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UNITED STATES NATIONAL MUSEUM

BULLETIN 217

SMITE SONLAN ENS TETUTEION

WASHINGTON, D. GC,

1960

VT RO BON

Oaet.”

Birds of
Anaktuvuk Pass,

Kobuk, and Old Crow

A Study in Arctic Adaptation

By LAURENCE IRVING

Research Associate, Smithsonian Institution

Publications of the United States National Museum

The scientific publications of the United States National Museum include two
series, Proceedings of the United States National Museum and United States National
Museum Bulletin.

In these series are published original articles and monographs dealing with the
collections and work of the Museum and setting forth newly acquired facts in
the fields of Anthropology, Biology, Geology, History, and Technology. Copies
of each publication are distributed to libraries and scientific organizations and
to specialists and others interested in the different subjects.

The Proceedings, begun in 1878, are intended for the publication, in separate
form, of shorter papers. These are gathered in volumes, octavo in size, with the
publication date of each paper recorded in the table of contents of the volume.

In the Bulletin series, the first of which was issued in 1875, appear longer, sep-
arate publications consisting of monographs (occasionally in several parts) and
volumes in which are collected works on related subjects. Bulletins are either
octavo or quarto in size, depending on the needs of the presentation. Since
1902 papers relating to the botanical collections of the Museum have been pub-
lished in the Bulletin series under the heading Contributions from the United States
National Herbarium.

‘Untrep Srates Government Patntine Orrice, WasHincron, 1960

For sale by the Superintendent of Documents, U. S. Government Printing Office
Washington 25, D. C. - Price $2 (paper)

Contents

Chapter
Acknowledgments
1. The Background .
2. Anaktuvuk Pass .
3s Kobuk,s.. ..
4. Old Crow .
5. Status and Diveribution -
6. Migration and Origins
7. Residence in the Arctic. . .
8

. Biological Aspects of Miers, esta Wesings: MG Se tie

9. Arctic Metabolic Economy of Warm-blooded Animals. . .
1

ECE EOIDED gfe ot as 5 4a iy Sb) gL g, to ee 2a
Literature Cited .
Appendix . .
Fee ish hse

ole Nd Je dhe

Acknowledgments

The observations on which I have based these studies were made
during expeditions for physiological investigations of adaptation
to the arctic Alaskan climate. These were supported by the Office of
Naval Research, through the Arctic Research Laboratory, from 1947
to 1949 and by the United States Public Health Service, through the
Arctic Health Research Center, after 1949.

I am also grateful to the Arctic Institute of North America for a
grant, from funds provided by the Office of Naval Research, to assist
our biological reconnaissance at Old Crow in 1957, and for grants from
the Explorers Club in aid of the reconnaissance on the Ahlasuruk in
1953 and at Old Crow in 1957.

Sig Wien, President of Wien Alaska Airways, whose many years
of arctic flying has brought him a thorough understanding of the fea-
tures of the country, and Thomas Brower of Barrow, whose keen un-
derstanding of life on the arctic coast had led him to suspect that
many birds reached there by migration through the mountains of the
Alaskan interior, were instrumental in leading me to choose the central
Brooks range as the key to the faunal situation in arctic Alaska.

Thomas P. Brower of Barrow, son of Charles Brower, also pointed
out to me the probable significance of the mountain passes as migra-
tory routes of the birds which he knows so well on the Arctic Coast
and related to me many pieces of information, sustaining that view,
derived from his own observations on the coast and accounts of Es-
kimo travelers in the interior. In 1949 he made extensive collections
and observations for me at Anaktuvuk which early presented an outline
of the migration. Since that time his friendly advice has repeatedly
helped to sustain my studies of arctic Alaskan animals.

Since 1950 my colleague John Krog has joined me in physiological
studies of the effects of arctic temperature. I owe to him many long-
sustained observations, and I have gained from him many clear views
of arctic life which were supported by his vigor and skill in combining
experimentation in the field with studies in the laboratory.

My colleague at Point Barrow, and later in Anchorage, Raymond
Hock, has given me much assistance from his observations at Tuluak
Lake during a few days in the spring of 1948, and from his observa-
tions on land and from the air in the lower Colville Valley during
that summer.

VI

ACKNOWLEDGMENTS VII

Robert Rausch first went with one of our expeditions from Point
Barrow to Tuluak in 1948. He has since then intensively studied the
mammals and parasites of the Anaktuvuk region (Rausch, 1951, 1953)
and he has generously given me many keen observations and numerous
specimens of birds. Neal Weber joined one of our expeditions to
Anaktuvuk in 1948, and he has subsequently made important con-
tributions to our knowledge of arctic insects (1948, 1949, 1950).

Carl Henkelman also accompanied one of our expeditions in 1948,
in order to make a survey of the dentition of the mountain Eskimo,
finding dental conditions excellent among these people subsisting al-
most exclusively on meat. Vladimir Walters in 1949, studied the fishes
in the region of Tuluak Lake (Walters, 1955).

Lloyd Spetzman in 1949 made collections of the flowering plants
at Anaktuvuk Pass, and George A. Llano, a former colleague in nu-
merous undertakings, in the summer of 1949 collected lichens (Llano,
1950). Attracted by accounts of the mountain people, my son William
Irving accompanied me in 1950 and 1951 in the Anaktuvuk and Kil-
lik Valleys to examine the people anthropologically, and has found
many artifacts of flint and bone in sites occupied by the former human
inhabitants of the mountains (William Irving, 1951, 1953).

It is with great pleasure that I acknowledge the helpful and pleas-
ant association with these good colleagues in an area where our com-
mon interests in natural history could be well rewarded. Our sep-
arate observations have built up a good view of the background of
life in the arctic mountain valleys, and have affirmed the early pre-
dictions of Tom Brower and Sig Wien as to the strategic importance
of the locality for the advancement of biological knowledge in arctic
Alaska.

I am sure that I express the views of all these scientific visitors in
acknowledging with gratitude the hospitality and assistance of the
mountain Eskimos. Without their help, existence for us would have
been too difficult to permit methodical scientific investigation; thanks
to their careful observations, accurate knowledge of country, and
ability to travel, our studies proceeded rapidly. In these people, who
live in primitive fashion, I have found a proud regard for accurate
and penetrating observation. In their complete knowledge of their
country they find the basis for a philosophical appreciation of nature
which has added admiration to my feeling of grateful friendship.

From Alexander Wetmore, I have had kind assistance in naming
several forms of birds, along with a continuing personal interest which
I gratefully acknowledge. Remington Kellogg, who has always been
a stimulating friend to field research, has cordially facilitated my
working at the U. S. National Museum. Herbert Friedmann has
given me much critical help and encouragement particularly in prob-

Vill U. S. NATIONAL MUSEUM BULLETIN 217

lems of distribution and systematics. Ira N. Gabrielson has assisted
me with determinations and information from his wide experience
with Alaskan birds. While I take responsibility for the determina-
tions, except as noted, I could not have undertaken them without
the guidance of Herbert Friedmann, nor would I have compiled the
studies without his help and encouragement. Herbert Deignan was
most ready in extending to me the use of the collections and working
facilities of the Museum, and Bernard Feinstein aided the identifica-
tion of the specimens from Old Crow by making many comparisons
and references, including a complete review of my arctic specimens.

The collections made include skins and sets of eggs with nests, which
are deposited in the U. S. National Museum, where I was kindly
afforded opportunity to compare the skins with those in the collections.
Most of the series of birds were quite uniform and conformed with the
average characteristics of their kind, so that identifications were, in
general, not difficult.

Tn his editorial capacity John Lea has suggested many arrangements
of the text, tables, and figures which would clarify the presentation of
the information.

Whatever is interesting and worthwhile in this work I wish to
dedicate to my wife and to our son Alan, whose short life was marked
by its great promise.

. Lavrencre Irvine
Anchorage, Alaska
February 1959

Birds of Anaktuvuk Pass,
Kobuk, and Old Crow

PLATE 1.
ANAKTUVUK Pass
(U. S, Air Ferce Photo)

Lonc-TaILeD JAEGER Stercorarius longicaudus (see pp. 78, 200, 247).

I. The Background

HEN I ENTERED the field of physiology after World War I,

scientific research was beginning a period of rapid expansion.
Accompanying this expansion, to enlarge the scope of scientific
observation, was a corresponding development of increasingly com-
plex instrumentation. For the analysis of physiological processes
this instrumentation is essential, but it is most easily applied in the
laboratory to the reactions of domesticated animals.

A consequence of this need to observe animals in laboratories is that
the physiological reactions are seldom viewed in context with the
natural situations in which the variety of responses evolved. In the
various natural situations in which each species finds itself, animals
have only the power of their intrinsic physiological and social proc-
esses with which to meet the environment and maintain the orderly
distribution of life. Under domestication the artificial selection that
produces animal varieties is exercised by processes extrinsic to natural
biological systems. This extrinsic process is arbitrarily applied by
human choice to decide the forms and associations in which life will

1

2 U. S. NATIONAL MUSEUM BULLETIN 2:17

exist In proximity to man, whereas natural selection proceeds accord-
ing to natural law.

Thus, in spite of the progress in biology resulting from the applica-
tion of new physiological instrumentation in the laboratory, we
usually have had to speculate about the mode of life of animals that
survive under those particular natural conditions which the physio-
logical processes we were able to study in the laboratory seemed quite
inadequate to cope with.

For example, the prolonged dives of whales and seals are impossible
for land animals, which are characteristically intolerant of inter-
ference with breathing. But as soon as I began to observe and measure
characteristics of respiration in diving animals, I found that only a
small range of the respiratory capability of mammals had come to
view because the laboratory study had been limited to a few domesti-
cated species whose rapid breathing concealed the basically discon-
tinuous process of respiration which supplies oxygen for continuous
metabolism. I found, for example, that when a seal dived the pro-
longed intervals banner its breathing magnified the appearance of
the respiratory reserves and lengthened the cycle of br eathing so that
its sequential processes could be separated.

Asa further consequence of their adaptation of respiration to long
periods of interrupted breathing a few species of mammals and
birds can enter waters which are well stocked with cold-blooded
animals. These warm-blooded birds and mammals, with their
air-breathing habit primarily suited to land, have a physiological
mechanism superior in its output of power te the cold-blooded mech-
anisms. This special adaptation of respiration has enabled them to
use the superiority of their warm-blooded metabolism for the exploita-
tion of an aquatic environment.

Plans fer Aretic Studies

A concern for these subjects led me, during World War II, to a tour
of duty in the Air Force, during which I was engaged in developing
techniques for human survival in the Arctic, and my interest in that
region thus become aroused. When, therefore, in 1946 I returned to
the Edward Martin Biological Laboratory at Swarthmore, I began,
with some colleagues who, like myself, had just been released from
the confinement of military service, to examine the pr ospects for
further investigations in the Arctic. We saw that the establishment
of routine inansdnctie air transport was opening the region to common
experience, and that a new phase of Arctic exploration, involving
comprehensive and intensive scientific research, was required to pro-
vide the information upon which would depend the establishment of
extensive human life there.

THE BACKGROUND 3

The present sparsity of human population in the Arctic and
Antarctic gives the impression of vastness, but these nevertheless are
the smallest of the earth’s climatic regions. Concentrated in the
ice of these limited areas are stored relics of ancient weather which
buffer the seasonal changes in temperature produced by current rates
of solar heating of the earth. Winter in the Arctic is so much colder
than the freezing temperatures at which vital processes slow down to
unproductive levels that it is, in a way, surprising to find there many
animals and plants which indicate their long arctic establishment by
having evolved special arctic forms. On the other hand, it is even
more surprising that some arctic species are scarcely distinguishable
from closly related populations living in warmer climates.

Only a small number of mammals, about 30 common species, and a
sunilar small variety of birds, are resident in the Arctic, owing to
the uniformly cold winter climate and the limited extent of the area,
so it seemed feasible to make a comprehensive survey of their essential
physiological adaptation to cold.

Research at Barrow

It was from these beginnings that plans grew for an expedition to
make physiological studies of adaptation to arctic cold at Barrow,
Alaska. This northermost point on the American Continent (latitude
71° N.) had the added advantage that an elaborate base was being
established there to support exploration in the Naval Petroleum
Reserve covering a large area in arctic Alaska. Means for support
of the expedition were generously provided by the Office of Naval
Research, stimulated by the interest shown by M. C. Shelesnyak, and
in August 1947 seven members of our expedition of physiologists
reached the Point Barrow navy station by air, a few days ahead of
the arrival of our heavy equipment by boat.

Per F. Scholander, who had been my colleague for eight years at
Swarthmore and in the Air Force, provided the scientific insight and
vigor, as a result of which our program of research soon showed sig-
nificant progress toward understanding physiological adaptation to
arctic cold. ‘These prospects encouraged us and the Office of Naval
Research to extend our stay for a second year.

We found, however, that the investigation of physiological adapta-
tion involves consideration of the many biological and physical con-
ditions which define the environment to which animals and plants are
suited by adaptation. Our need to consider these environmental con-
ditions served to outline the wide scope of scientific research which
could be usefully undertaken in arctic Alaska. As a result the Office
of Naval Research was led to name the small quonset hut occupied by
our party of physiologists the Arctic Research Laboratory, and during

4 U. S. NATIONAL MUSEUM BULLETIN 217

the winter I was asked to prepare a plan for a larger laboratory
structure. From the skillful designs of Ben Atkinson, and with
energetic construction by the Arctic Contractors, the new laboratory
was ready in the summer of 1948 to be occupied by about 40 scientists.
About 20 remained during the winter of 1948-49.

The summer of 1949 Scholander and I spent compiling the results
of our studies in Alaska. George MacGinitie had taken over direc-
tion of the Arctic Research Laboratory. I was more attracted to
research in the field than to the duties which tend to surround the
direction of scientific programs, and as Joseph C. Mountin and Jack C.
Haldeman were preparing to establish for the Public Health Service
at Anchorage a laboratory for studying conditions basic to the main-
tenance of health in arctic Alaska, in the autumn of 1949 I joimed
them in the Arctic Health Research Center, where I have since con-
tinued my studies on adaptation to arctic cold.

At Barrow in 1947 we had found the birds of the arctic coast
fairly well identified by the succession of naturalists (Bailey, 1948)
who followed the pioneer study of Murdoch (1885) during the First
International Polar Year. It is interesting that Ray (1885), the
young lieutenant of engineers in charge of making these two years of
geophysical recordings, also made the first significant anthropological
report on the arctic Alaskan Eskimo. The coastal mammals, too,
were known in 1947 but, as with the other fauna, their distribution
and relations to terrain and seasons could not be described from the
occasional collections which had been made. Only a few land
animals can live through the long cold winter of the arctic coast, with
its strong and often violent winds; and from June through August,
when overcast skies and fog are more common than sunshine and snow
flurries and frost occur frequently, warm days are too rare to produce
more than a scant growth of vegetation for their sustenance.

The Interior of Arctic Alaska

For our studies we needed to know a greater variety of animals than
we could find along the arctic coast and so we turned toward the inte-
rior of arctic Alaska. It is true that much had been earlier observed
in the interior of arctic Alaska, but the biological information was
discontinuous in time and space and restricted to identification of
forms in only a few systematic categories. Geologists had surveyed
the interior. During the winter of 1926-27 A. EK. and Robert Porsild
(1929) traveled by dog team from Buckland River along the Alaskan
coast to Aklavik, surveying the flora and conditions for the route of
the Canadian reindeer herd which was subsequently settled on the
eastern side of the Mackenzie Delta. In 1947 there had been no
reports on man or animals in the interior north of the ornithological

THE BACKGROUND 5

survey of Grinnell (1900) and the archeological exploration of Gid-
dings (1952) along the forested part of the Kobuk River. Olaus
Murie (1928) had reported on some of the birds which he found along
the Koyukuk and Alatna Rivers, but otherwise the arctic interior of
Alaska and Yukon was biologically unexplored.

It was the necessity to obtain more animals that first sent some of
us on short trips inland from Barrow. Gradually the scope of these
trips increased, but the distribution and habits of the animals were
unfamiliar, and without more information about their environment
we were in a poor state for estimating the adaptive significance of
their physiological processes. At first I could not imagine by what
means we could acquire the necessary comprehensive information about
biological conditions in the area of interior Alaska for 300 miles north
of the arctic circle and ranging 700 miles eastward from Cape Lisburne
to the Mackenzie Delta.

Fortunately Sig Wien, president of Wien Alaska Airways, was
then often at Barrow, and because he had been flying over arctic
Alaska for a number of years he thoroughly understood the features of
the country. Tom Brower, a native and merchant of Barrow was
also a good adviser, for Tom, his eminent father, Charles, and his
brothers knew the arctic coast and its adjacent interior. He could
view the country as a naturalist of experience, for his family was
credited by Bailey (1948) with making known 63 of the 153 species
of birds recorded from Barrow. To Wien and Brower the key to the
faunal situation in arctic Alaska appeared likely to be found in the
mountains of the Brooks Range. Bailey (1948, p. 177) expresses
Brower’s view: “He stated that there is a definite migration route
over the Endicott Mountains and down the Colville, and that many
water birds use this route to and from the eastern arctic coast.”

We decided upon the system of passes about Anaktuvuk as the
most favorable starting location for several reasons. The several
passes that interconnect in that region constitute the primary system
of easy northward passage of the Brooks Range overland or for low-
powered aircraft west of the Canadian border and the air transport
route to the arctic coast follows Anaktuvuk Pass (fig. 1), through
which low-powered aircraft in contact flight can guide upon a well
defined valley with ground elevations below 2,500 feet. In this com-
plex of passes, lives a small community of inland Eskimos, who were
said by Wien to know thoroughly the terrain and the wild life of
their native mountains. His estimate of these people gave us reason
to think that with their aid we might make a biological survey in an
area where to establish our own bases and to travel unguided would
require impossible expeditionary effort in time and expense.

I decided to utilize my interest in birds for developing a picture

U. S. NATIONAL MUSEUM BULLETIN 217

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U. §. NATIONAL MUSEUM BULLETIN 217

THE BACKGROUND

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8 U. S. NATIONAL MUSEUM BULLETIN 217

of a faunal group which would well indicate the relations between
the animals and their environment. There are a sufficient number
and variety of birds, and their taxonomic relations are well worked
out. Like ourselves, birds are principally directed by sight and sound
in activities by daylight, and so they demonstrate themselves to our
senses as to their own. Consideration of the relations of animals to a
country brings to mind their connection with populations of other
regions, and the visible flight of birds indicates how populations are
distributed. In particular the migrations of birds had impressed upon
my interest that their courses annually exhibit the operation of distri-
butional inclinations which are invisible in sedentary animals,

I did not anticipate in 1947 that for the ensuing 12 years I would
be studying adaptations to life in arctic Alaska, nor did I then realize
how many biologists would become associated in their attention to
that area. But after several years it was apparent that the bird life of
arctic Alaska indicated it to be a well characterized faunal region,
and I was led to look for the geographic features by which the environ-
ment might be defined as a habitat for its somewhat special fauna.

Physiography of Arctic Alaska

The principal physiographic feature of arctic Alaska (fig. 1) is the
Brooks Range, which extends eastward near the 68th parallel from
Cape Lisburne for about 700 miles to the Mackenzie Delta. In east-
ern Alaska the peaks of the Brooks Range rise above 9,000 feet.
They diminish in the center to about 7,000 feet, and to about 4,000
feet within 100 miles of the western coast. In the northern watershed
are many short swift rivers, the central ones draining into the Colville,
which flows eastward through the rolling tundra of the arctic slope.
East of the Colville mouth the coast approaches close to the moun-
tains, and north of the Colville the arctic slope gradually lowers to
a flat coastal country of many lakes.

Along the southern watershed the westerly flowing Koyukuk River
receives the rivers from the central mountains of the Brooks Range
while those in the east drain into the Yukon and Porcupine. The
Noatak River runs west from the western Brooks Range and enters
Kotzebue Sound just north of the Kobuk River; together they drain
the southwestern base of the Range. Although the Kobuk is not con-
nected with the Yukon River, its southern watershed is open at low
elevations to the Koyukuk Valley and so to the Yukon Valley. The
Koyukuk River flows into the Yukon and from its upper valley a
broad low pass leads to the upper Yukon Valley. In Yukon Terri-
tory the Porcupine River, draining the southeastern watershed of the
Brooks Range, forms a continuation of the Yukon Valley. The south-
ern watershed of arctic Alaska thus is the northern part of the great

THE BACKGROUND 9

Yukon Valley system, which provides a continuous valley across
northern Alaska and Yukon Territory extending from the western
coast to within 50 miles of the Mackenzie River without significant
obstruction to the distribution of animals. McDougall Pass, at an
elevation of 1,200 feet through the Richardson Mountains, is the
lowest and shortest pass through the western mountains of North
America.

Arctic Climates Present and Past

The northernmost treeline in America is in arctic Yukon. East
of the Mackenzie River the forest retreats from the arctic circle and
its distribution roughly follows the June isothermal line of 50° F.
mean temperature, which reaches the Arctic in the interior of Alaska
(fig. 2). In winter, interior northern Alaska and Yukon include the
coldest part of North America, but in summer warm air from the
north Pacific tempers the northwestern interior climate as far north
as the Brooks Range. North of the Range the cold summers of
arctic shores prevail. Many birds and mammals can endure any
degree of arctic cold, but their existence depends upon the provision
of food which is ultimately the product of vegetation. Many plants
also endure cold but their growth occurs in summer warmth. The
vegetation that results from these climatic conditions in the Yukon
Valley enables the woodland animals there to reach their northern-
most limit in America.

Since summer temperatures are cold north of the mountains and
warm on the southern slope, in passing northward across the Brooks
Range a rapidly cooling gradient of summer temperature is experi-
enced. This steep northward gradient is peculiar to western arctic
America, for east of the Mackenzie Valley summer warmth equaling
that in the northern Yukon Valley reaches only as far north as James
Bay. The northward extension of summer warmth into western arctic
America is the result of conditions in the earth’s circulation of heat
which have been long in operation. This is shown by the southern
limit of extensive permanently frozen ground in North America, which
in Alaska lies north of Anchorage (lat. 61° N.), while in the middle
of the continent around James Bay it extends farther southward by
about 8 degrees of latitude. Near the arctic coast of Alaska explora-
tory drilling for petroleum shows that permanently frozen conditions
extend for 1,000 feet below the surface. (Black, 1951.)

Relics of the great American ice fields of the past now exist only in
southeastern Alaska although evidence for recent, more extensive
glaciation is shown in the gradual invasion of vegetation into valleys
from which it is apparent that glaciers have recently melted. The
western limit of Alaskan glaciation extended only as a shelf into the

10 U. S. NATIONAL MUSEUM BULLETIN 217.

deep and warm Pacific Ocean. The southern foot of the ice cover
of the Wisconsin age was at about latitude 60° N. in central Alaska,
while the midcontinental ice fields extended 15° farther southward.
In the late Wisconsin age continuous ice fields discharged glaciers
north of the Alaska Range to form moraines along the southern sides
of the Kuskokwim and Tanana Valleys (Péwé, 1953). North of the
Brooks Range, the Wisconsin age glaciers also extended just beyond
the mountains. In general it appears that in Alaska north of about
latitude 64° extensive lowland areas were not then covered by ice and
that the continuous ice cap in central Alaska extended farther south-
ward across only about 5° of latitude (see map, fig. 3). McConnell
(1890) first remarked that the northern part of Yukon Territory
appeared to have been free from the continuous ice which covered
the country far to the southward and which probably extended over

4 ANS
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110 =
SEA-LEVEL ISOTHERMS : JANUARY (°C)

Ficure 2.—Sea-level isotherms for January (left) and May (right). (From C. F. Brooks,
Univ. Press, 1936,

THE BACKGROUND 11

the arctic coast east of the Mackenzie (Flint, 1947). So in western
arctic America the glacial regime appears to have left large areas in the
Yukon-Porcupine Valleys and on the arctic slope of Alaska free from
ice fields during the late Wisconsin age. The conditions northwest
of the American ice cap were quite different from those on the massive
ice sheets which continuously covered the rest of Alaska, Canada,
and some northern States.

The records of glacial geology lead Flint (1947) to propose that
Bering and Arctic seas were frozen during the last great Wisconsin
glaciation, and that the precipitation forming the arctic ice caps
came from the tracks of storms like those of the present. This moisture
would then have originated in the temperate oceans. However, re-
cently acquired oceanographic evidence of the history of temperatures
in the Atlantic Ocean has led Ewing and Donn (1956) to a novel

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SEA-LEVEL ISOTHERMS : MAY (°C)

A. J. Connor, and others, “‘Climatic Maps of North America,’ Cambridge, Harvard
maps 1 and 3.)

12 U. S. NATIONAL MUSEUM BULLETIN 217

hypothesis that the moisture for the latest American ice cap came
from the Arctic seas, which they postulate as unfrozen while the
maximum glaciation developed. Geological and geophysical methods
are rapidly developing evidence of climatic history, which is probably
closer to elucidation than is indicated by these contrasting views on
the climatic mechanisms of glaciation.

For the present we must be cautious in postulating what the climate
was like on the arctic coast of Alaska adjacent to a sea which might
have been frozen or unfrozen but in the interior the climate could not
have been warm and was probably cold in the proximity of the great
ice cap over the Alaska Range and the extensive glaciation in the
Brooks Range. It appears that, in most of the ice-free northern
American area which was connected with Siberia, the climate in the
area that served as a “refuge for survival” was at least cold and could
suit only animals like those now hardy in the north.

Climate and the Animal Populations

The masses of water frozen in polar and arctic ice near the end, and
apparent maximum, of Wisconsin glaciation so reduced the oceans
as to connect Alaska and Siberia over Bering Strait. Fossils of ex-
tinct animals as well as the systematic relations of many animals now
found, indicate that some taxonomic groups in America derive from
Asiatic predecessors and some in Asia from American predecessors.
It is popular to speculate that the intercontinental exchange implied
in the relations of the present and fossil animals of America and Asia
was effected by traffic over the land bridge.

On the basis of systematic relations and the geographical and chron-
ological sequences in which they are found, it is reasonable to propose
that between Asia and America migrations have occurred that have
altered the previous geographical distribution of animals. Walters
(1955), for example, has shown that the land bridge of the late Wis-
consin age could have lead to the present distribution of fresh water
and coastal fishes in western arctic America and eastern Siberia. In
the aquatic environment of fishes in the north, the temperature is al-
ways above freezing, but not much warmer, and suitable climatic con-
ditions for fishes can be reasonably inferred for the waters about the
_ recent land bridge.

Most of the species now living can be traced through fossil records
to Pleistocene times but their evolutionary modification has been too
slow to produce variation in form which would indicate an exchange
between Alaska and Siberia in Recent time. A number of land
mammals, many conspicuous by their size, have become extinct since
the Pleistocene, some of them only a few thousand years ago (Flint,
1947). Unfortunately the situation of their remains in Alaskan loess

THE BACKGROUND 13

Ficure 3.—Glaciation of the Wisconsin age compared with that of the present. Black
areas are existing glaciation, hachured outline marks limits of Wisconsin glaciation,
isotherms are present July means in degrees F, (From L. S. Dillon, “Wisconsin Climate
and Life Zones in North America,” Science, vol. 123, fig. 1, 1956.)

deposits do not well preserve the record of the surroundings in which
they lived, and we cannot yet list by their remains the land animals
which existed in Alaska during the last great glaciation about 10,000
years ago or say whether any then changed their range by migration
over the land bridge.

If, as it appears, new species have not evolved during the Pleistocene,
and these animals have passed through the climatic variations of
recent ice ages without much evolutionary change in form, then the
mammals and birds resident in ice-free arctic America and Siberia
during the Wisconsin age could have been like those now resident
there. Some of the large mammals have recently become extinct and
some have moved in from the south since the ice sheet melted. By
that time, however, the land bridge probably no longer existed, and in

14. U. S. NATIONAL MUSEUM BULLETIN 217

any event it does not seem to have been a suitable intercontinental
route for animals from a warm climate.

It is not to be suspected that during the glacial periods birds could
make annual migrations across the American ice cap. Whether mari-
time and oceanic birds could migrate to Bering and Arctic sea coasts
from the north Pacific would have depended, for one condition, upon
the amount of ice-free coastal and inland waters. It does not seem
likely that extensive nesting grounds were open to them on the coast
or in the interior. Ice free corridors through the east Siberian
glaciers probably existed (Flint, 1947), but for long distances the
routes to warm areas for wintering in Asia were in proximity to ice
fields. The routes to Alaska and Siberia were either impassible or
difficult, and the limited unglaciated arctic area and its conditions
would seem to have provided insufficient attraction for the annual
migration of birds to the arctic.

When the ice caps melted, the American lands suitable for animals
were increased by the opening of most of Alaska, Canada, and the
northernmost states. Populations have extended to occupy these new
lands but new species have not evolved. The source of the present
avian populations may have been in survivors in the ice-free refuge
and in those which have come in from the south as migratory routes
developed. While it seems possible that some species of birds and
mammals might have survived the last maximum glaciation in the
unglaciated parts of Alaska and Yukon, the annual migrations of
birds to nest in the Arctic have certainly developed since the ice
caps melted. In each spring the migrants now pass northward over
lands which have only gradually become occupied during the last
10,000 years. Many of the populations that pass annually from
a warm wintering place to nest in the cold arctic spring encounter
cooling climatic gradients by their own movements, thus reversing
the seasonal transitions in temperature encountered by northern
sedentary populations. All the expanding populations of Canada
and most of Alaska have had to adjust to the new and changing condi-
tions of the environments in which they have settled. They have also
had to adapt social habits to the requirements of moving and develop-
ing populations. The animals which live near the arctic limits of life
must have possessed a great range of adaptability toward environ-
mental and social conditions to have succeeded during the changes of
the Jast 10,000 years.

na ee
E. PALL POSIT ee ayn ae ca
aa : eae SS ay =

a x i Pi oe

=i a,

1,

U. S. Post Orrice at Summit, ANAKTUVUK Pass, 1951 (see p. 25).

2, Anaktuvuk Pass

hie THE BoRDERS of Yukon Territory westward to the sea the
broad valley of the Yukon River offers no barrier to the flight of
birds. Along the northern part of the central Yukon watershed the
Koyukuk is its principal tributary, running approximately west near
the southern border of the mountains. Near Bettles the John River
enters the Koyukuk, after flowing about 90 air miles south from the
divide locally called Summit (lat. 68°10’ N., long. 151°40’ W., eleva-
tion about 2,400 feet). Just north of the divide at Summit the
Anaktiktoak River comes in from the eastward to join the head of
the Anaktuvuk, which then runs about 100 miles nearly north to the
Colville River.

A forest of spruce with some birch lines the John River and extends
up its tributary valleys to maximum elevations of 3,500 feet on south-
ward slopes as far north as Publatuk Creek, about 30 miles south of
Summit. In the main valley, tree line is passed below 2,000 feet,
where the forest terminates abruptly. Northward, willows are re-
stricted to certain areas in the valley. In some places they may reach

15

16 U. S. NATIONAL MUSEUM BULLETIN 217

a height of 15 or 20 feet, in others they are only a few feet high. A
few small cottonwoods occur in occasional and restricted areas and
a few alders are found at Akvalutak Creek.

Northward in the John River Valley the traces of glaciation be-
come more apparent, and the upper John and Anaktuvuk Rivers lie in
an obvious glacial cut trough about 3 or 4 miles wide between steep
mountain walls of much dissected metamorphic rocks. No superficial
ice now remains permanent in the Pass, but a small mass of ice near
tree line in the upper John Valley, not now evident, was reported
by Schrader (1904) as of glacial origin.

Along Anaktuvuk Pass the mountains rise steep and sharp to 5,000
or 6,000 feet. Through their walls enter occasional narrow and pre-
cipitous valleys much eroded by water since their initial formation by
tributary glaciers from former high snow fields. Now the sparse snow-
fall outlasts the hot June sun in only a few places. With its meager
precipitation this country has some resemblance to the desert region
in the southwestern States. The wide fans of boulders and gravel at
the foot of the tributary streams attest the occasional power of ancient
slides and occasional flood flow, and the appearance of lateral moraines
on the mountain walls west of Kalutak Creek suggest that the ice
levels of the glacier may have been 1,000 feet above the present floor
of the John River Valley. The northern margin of the mountains
rather sharply forms a mountain line, north of which only an occa-
sional elevation rises to 2,000 feet among generally long-sloping
smooth hills.

At the base of the steep side walls of the valley the talus slopes end
at a level a few hundred feet above the valley floor. Below that level
the gently sloping terrain is often dry and sparsely covered with
short vegetation over the occasional, well drained areas of gravel.
More often the surface is wet and irregularly spotted with
hummocks of sedge clumps called niggerheads. The small tributary
stream beds are often elevated upon fans of their own detritus, into
which they cut annually varying channels. These are frequently lined
with willows that are a few feet high near the mountain walls and up
to 12 or 15 feet high in occasional stands along the river flats, where
the basal trunks may be 5 or even 6 inches in diameter.

From the junction with the larger east fork (Anaktiktoak) just be-
low Kangomavik, the Anaktuvuk meanders north in the mountains
for about 15 miles through a flat valley floor of sand and gravel among
small moraines, leaving several cutoff, or oxbow, lakes (see map, fig.
4). On the tundra adjacent to the river are many marshes and small
lakes, some of which are of the strange but not unusual arctic type
resembling limestone sinks, with steep sides cut deeply into the per-

ANAKTUVUK PASS

17

152°00" 40’ 20° 151°00' 40'
68°30 oma
SM Yo Narivukararuk Lake ‘— Snow *
) Fed 7 ralukpug
Ji A,
t S
os
20° —| 20’
10° 10’
oP ANING@r5,
fi Da onn
ae “
<p,
By ee
“oto, £e4.
"low,
Me
,
Uy Ney
Ny
50’
4S ANAKTUVUK PASS
if Con Statute Miles
\e. SUMMIT = 0 2 4 6 8 10
S fav Foot Corty, \ COMPILED BY WM.‘IRVING
\
ro) $ DATA FROM SIMON PANEAK
nig S
l) (ESS
Bo Ja OEY,
x a"
{J Contact Creek <7 "
67°40 : o 67°40'
152°00’ 40’ 20 151°00' 40° 4

Ficure 4.—Sketch map of Anaktuvuk Pass.

ey U. S. NATIONAL MUSEUM BULLETIN 217

10 \TOLUGAK L. CHANDLER L. KUNGOMOVIK TOLUGAK L. KUNGOMOVIK
+ CONTACT CR. KALUTAK CR. | | 4
-10 © 900 ° ooo
° o
-20

-30

DEGREES CENTIGRADE

SNow So”
RIVERICE 12°

-40

10 20 30 10 20 3| 10 20 30 10
SEPTEMBER OCTOBER NOVEMBER DECEM

Ficure 5.—Temperatures in Anaktuvuk Pass. Top: Winter temperature 1948-1949,
taken daily at 8 a.m. (from the journal of Simon Paneak). Bottom: High and low
temperatures, spring 1951 (from the journals of John Krog and Simon Paneak).

manently frozen foundation of the tundra. Few of the lakes in the
valley are a mile in length.

Tuluak Lake on the east side of the valley (Lat. 60°19’ N., Long.
151°26’ W.), is formed principally by the clear streams from Nakag-
nik Springs, rising from about a quarter-mile east of the lake, and in
places is open throughout the winter. Tuluak is a deep lake, nearly
a mile in length, well stocked with grayling, a large and a small char,
small Jake trout, and whitefish. It is a pleasant spot, much frequented
by migrating birds, and is still a favorite Eskimo campsite. Local
stories and traces of old habitations nearby indicate its ancient im-
portance to the mountain Eskimo, who formerly called that portion
of the valley Tuluak, meaning “raven.” The Eskimos say that they
called the valley Tuluak and applied Anaktuvuk to another valley east
of there. Schrader, who named the localities (1904), was apparently
not guided there by the mountain Eskimos and he applied some names
varying from historic Eskimo usage. Subsequent maps have often
failed to use established aboriginal names and have applied trite,
meaningless English names.

Weather and Climate

The record of temperature in figure 5, prepared from observations
by Simon Paneak and John Krog, shows no extreme cold, but
sudden fluctuations in winter temperature often occur. These changes
are frequently accompanied by winds so violent and turbulent that,
particularly in the Killik Valley, Eskimo boys were taught that winter
travelers must camp only at sites known from experience to be shel-

ANAKTUVUK PASS 19

KALUTAK GR. AHYANITOK CR.- JOHN R. 10

BER JANUARY FEBRUARY MARCH APRIL

DEGREES CENTIGRADE

1314 15 16 17 1819 2021 22 23242526 27 2829 3031 | 2345 6 7 8 910 II 1213 1415

MAY JUNE

tered from the dangerous force of the winds. The power of the winds
is shown by the wind erosion of the high rocky pinnacles above Kan-
gomavik, by many large old dunes along the river, and by the furrow-
like depressions on the surface of old dunes now covered with vegeta-
tion. In the Killik Valley these old and at present fast-building dunes
are spectacular. From early June to mid-August storms are rare and
in July the weather has often been the most delightful I have expe-
rienced.

North of the timbered southern slopes snowfall is sparse, probably
not over two feet in winter, and it is usually drifted and compacted
nearly but sometimes not quite hard enough to bear a man’s weight,
so that small snowshoes are in common use. Many exposed areas
are blown free from snow, thus clearing the vegetation for the caribou

20 U. S. NATIONAL MUSEUM BULLETIN 217

and rock ptarmigan to feed upon. The branches and twigs of the
willows protruding through the snow are exposed during most of
the winter. Since the larger arctic animals and birds are not bothered
by winter cold (Scholander, Hock, Walters, Johnson, L. Irving, 1950;
L. Irving, 1951), the exposure of their sources of food gives a certain
advantage to tundra residents over those in the windless forest areas
where snow and frost cover the branches as well as the ground.

The lakes are not usually iced over until late in September, but by
early October the ice may be a foot thick. On any extensive ice sur-
faces the snow may be blown into hard drifts with extensive clear
areas, so that the frequent winds facilitate winter travel for men and
animals. Thus, if properly considered and utilized, the tempestuous
winter weather in the mountains facilitates the feeding of animals
and the travel of man. And, in fact, the variety of human artifacts
found widely distributed in Anaktuvuk and Killik Valleys (W.
Irving, 1951, 1953), shows that people using Mesolithic implements
once inhabited these valleys.

In late May the winter snow has generally melted or evaporated
and the streams and rivers have broken through to flood over the
ice which still remains fixed to the bottom. In some lakes the ice
may remain sufficiently firm for landing light aircraft until after
June 10, although the margins of many lakes are open sufficiently
for the feeding of shore birds and ducks after the middle of May.
Fresh snowfalls and cold freezing weather occurred in the first week
of June in 1948 and 1949, and in 1949 another period of several days
followed, around June 20, when the ground was covered with 6 inches
of fresh snow. These June snow storms obviously harassed the shore
birds, for they then had no place to feed or land except upon a few
rocks protuding above the stream beds. The distress of the expressive
yellowlegs and tattlers was pathetic to watch. Many nests were cov-
ered and the eggs destroyed by the snow; and yet a few days after the
return of warm weather the exuberance of bird life in Arctic spring-
time reappeared. In spite of these recurring hazards, the bird popu-
lations flourish as a whole, and the occasional hardships are appar-
ently offset by generally favorable conditions. It seems that these
populations migrating to the arctic encounter an average condition
of abundant food and favorable weather and only individuals suffer
from the vicissitudes of climate.

After mid-June, freezing temperatures are rare, rain is uncom-
mon, and long sequences follow of brilliantly clear days with most
agreeable temperatures. Daylight is continuous and during warm
parts of the long days children play as noisily in the river as they do
in any climate. The well drained areas of the tundra and the mountain
slopes rapidly dry to a desert appearance.

ANAKTUVUK PASS 21

By the first of June willows are flowering and large active bumble-
bees cover themselves with reddish pollen, often working in freezing
temperatures when the sun is obscured. By mid-June mosquitoes are
active and soon form an abundant cloud about each person and dog,
even in the wind. Later they are supplemented locally by much
more poisonous black flies.

Ranunculus (anemone) and Pedicularis (lousewort) flower early
in June, followed by Potentilla and Dryas, while in July Pinguicola,
a beautiful small insectivorous plant, H’pilobiwm (fireweed), and a
number of lupines and vetches brighten the tundra. The willow
leaves are well out in July, when the dwarf Arctic Rhododendron
flowers fall, and the early part of that month is a colorful period,
particularly along the small streams. Grasses, sedges, and leafy
plants develop so slowly that their green does not conceal the old
vegetation until mid-July. Many lichens show little color in the dry
summer, but in the moist weather of August their green begins to
freshen and it remains bright long after being covered with snow in
the fall. Altogether, the summer scene is delightful and in no place
have I seen people, animals, and birds, appearing so well adjusted
to their environment as in these arctic mountain valleys.

The Nunamiut

The 70 Eskimo people of the mountains trace their origin to
Nunamiut ancestors. For them, the term designates an ethnic group
as well as a community, and they consider themselves different from
other Eskimo people. Their ancestors have lived in the mountains
north of the forests, along the southern watershed of the Colville
River, where they formed the flourishing interior population which
Stoney and Howard met in 1885 and 1886 (Stoney, 1900; W. Irving,
1953; L. Irving, 1953; and L. Irving and Paneak, 1954). In historic
times, and in the relatively recent prehistoric times which their stories
penetrate, they ranged westward to meet for trade the Eskimo of the
Kobuk, Noatak, and Utokok Rivers. But they did not themselves
regularly traverse the mountains west of Howard Pass and they
have little knowledge of the western arctic coast. Southward they
sometimes encountered the Indians, with whom there have been
alternating periods of conflict and rather indifferent accord. A
tundra culture requires considerable modification in order to suit
forest life, and the southern experience of the Nunamiut apparently
just penetrates the edge of the forests. Their territory has not
extended north of the Colville River in recent times. As a result of
occasional conflicts at the border they have carefully left that country
to its inhabitants, the coastal Eskimos, except for traffic along regular
routes of travel.

23 U. S. NATIONAL MUSEUM BULLETIN 217

Eastward, the Nunamiut used to descend the Colville River in
spring to obtain seal oi] and sealskin for boots in exchange for caribou
skin, wood, and flint products. Although the present Nunamiut are
mountain people, they well understand sea travel and hunting for
seals along the summer arctic coast. In fact, the eastward travels of
some of the present group have extended to the Coppermine River, —
and they regularly correspond with friends and relatives all along
that section of the Alaskan and Canadian arctic coast.

In keeping with these habits, they know accurately the birds of the
arctic coast east of the Colville, but their stories show no familiarity
with the conspicuous and distinctive avifauna of the Bering Sea coast
and the western Arctic (L. Irving, 1953). In physical characteristics
they appear to be distinguishable from Eskimos of the northern arctic
coast, and their own view of their distinction from the present dwellers
of the coast regions should be seriously considered.

Progress of the Investigation

On October 17, 1947, Per Scholander, Clay Kaigelak of Barrow,
and I flew with Sig Wien from Point Barrow into the camp of the
inland Eskimo people. The camp, consisting of 4 families and about
25 people, was situated on the west side of Chandler Lake Valley near
the north end of the lake (lat. 68° 19’ N., long. 152° 35’ W.). We
were courteously received by the people and were soon comfortably
settled. After studying the country by day, we discussed during the
long evenings in their domed tents of caribou skin the natural history
of the mountain valleys. Simon Paneak best presented the descrip-
tions in his accurate and expressive English, and his observations were
supplemented by well considered comments from Jesse Ahgook and
Elijah Kakena in Eskimo and by Frank Rulland in both languages.
Their wives and older children were often consulted for confirmation
and sometimes served as referees in the discussions. While the aurora
flashed in the winter nights outside, the people described many birds
with the intimate detail in which I had known them as a boy in the
northeastern states. They pictured the springtime arrival of migrants
as a wave, passing through or stopping to nest in the valleys. My
curiosity about the birds of their country aroused their interest, and
we soon came to discussing places and dates which would be suitable
for making decisive observations. It was apparent at this first meeting
that accurate reports and intelligent appreciation of birds could be
obtained from the keen observations of these Eskimo residents of the
mountains.

When we left them, the people of the village were preparing to
travel south to winter within the spruce timber on the upper John
River near Hunt Fork, for the low willows about Chandler Lake

U.S. NATIONAL MUSEUM BULLETIN 217 PLATE 2

EGGS

Top: Eastern American golden plover, near Kiminiaktuk, June 14, 1951. Bottom: Short-
billed gull, near Summit, June 24, 1951.
469496—60 3

U.S. NATIONAL MUSEUM BULLETIN 217 PLATE 3

EGGS

Top: Parasitic jaeger, with 16 gauge shell, near Kiminiaktuk, June 14, 1951. Bottom
phoebe, at Napaktoaloitch Cave, June 17, 1951.

U.S. NATIONAL MUSEUM BULLETIN 217 PLATE 4

Top: Kennicott’s willow warbler, at Contact Creek, June 23, 1951. Bottom: Eastern robin,
at Nacharach, June 10, 1952.

U.S. NATIONAL MUSEUM BULLETIN 217 PLATE 5

Top: Common redpoll, at Imaiginiek, June 27, 1951. Bottom: Gambel’s white-crowned
sparrow, at Nacharach, June 10, 1952.

U.S. NATIONAL MUSEUM BULLETIN 217 PLATE 6

EGGS

Top: Smith’s longspur near Nacharach, June 10, 1952. Bottom: Western tree sparrow at

Nacharach, June 11, 1952.

U.S. NATIONAL MUSEUM BULLETIN 217 PLATE 7

E

THE PANEAKS

Top: Simon Paneak at Chandler Lake, November 1947. Bottom: Mrs. Susie and Franklin
D. Roosevelt Paneak (still called Mister) at Chandler Lake, November 1947.

U.S. NATIONAL MUSEUM BULLETIN 217 PLATE 8

ANAKTUVUK VALLEY

Top: Simon and George Paneak before a rack of 37 wolf skins, February 1952. Bottom:
Homer Mekiana’s tent at Contact Creek, the new U.S. Post Office at Anaktuvuk Pass,
August 1951.

U.S. NATIONAL MUSEUM BULLETIN 217 PLATE 9

ANAKTUVUK VALLEY

Top: While a Nunamiut hunting party stops to cook sheep meat, one of the boys surveys
the valley through his telescope, near Kungomavik, July 1950. Bottom: Jesse Ahgook
among remains of old dwellings at Kiminiaktuk, looking eastward down the valley by which
the creek emerges from the mountains, July 1950.

U.S. NATIONAL MUSEUM BULLETIN 217 PLATE 10

«* 7 *
. ay
. 2 aia: ee Ax

KILLIK AND ANAKTUVUK VALLEYS

Top: Clumps of alder near Akmalik Creek in the Killik Valley, August 1950. Bottom: Cotton
sedge in Itikmalukpuk Creek about 10 miles north of the mountain line, July 1950.

U.S. NATIONAL MUSEUM BULLETIN 217 PLATE 11

ANAKTUVUK PASS AND CHANDLER LAKE

Top: Terminus of boating on Anaktuvuk River about 10 miles north of the mountains, look-
ing southward, July 1950. Bottom: Camp near Chandler Lake, looking northeast across
Chandler Valley, November 1947.

U.S. NATIONAL MUSEUM BULLETIN 217 PLATE 12

ANAKTUVUK PASS

Top: Camp village at Nacharach (Summit) near Contact Creek, looking westward at the
wall of the Valley, June 10, 1952. Bottom: Looking southeast across Tuluak Lake at the
camp then located there, June 1, 1949.

U.S. NATIONAL MUSEUM BULLETIN 217 PLATE 13

IMAIGNIEK

Top: Looking southwest at the camp at Imaigniek, August 1950. Bottom: Looking south-
ward along the Valley from camp at Imaigniek, August 1950.

ANAKTUVUK PASS 23

afforded meager fuel for winter use. In March 1948, they returned
north into Anaktuvuk Valley to camp on the east side of Tuluak Lake.
This lake (lat. 68°19’ N., long. 151°26’ W.; see fig. 4) is situated on
the eastern side of the valley near the northern edge of the mountains
which they call the mountain line. The lake, almost a mile long, is
fed on its eastern side by streams from Nakagnik Springs which re-
main open through the winter. Remains of ancient habitations and
stone corrals for caribou drives, and the stories told by old Eskimos,
indicate that this region has been the site of many old camps. Flint
artifacts found near there at Imaiginik are assigned to the Cape
Denbigh type of the earliest Alaskan artifacts yet described (William
Irving, 1951). The modern residents of the mountains, who still sub-
sist mainly by hunting caribou, thus follow the ancients in recognizing
the suitability of this mountain valley for their way of life.

On June 1, 1948, with my younger son, Laurence H. Irving, I again
visited the Tuluak Lake camp for a week. The winter snow had melted
from the valley floor and lower mountains. The first flowers and
pussywillows were showing. Wandering tatlers and yellowlegs were
numerous and several of them, with some long-tailed jaegers, were held
captive for me in the tents of the Eskimo people. Plovers and sand-
pipers, ducks, loons, Alaska and Smith’s longspurs and, in fact, all
birds were either still moving north or settling down to nest in the
valley. This lively spring scene was changed after a few days with
the fall of 6 inches of snow and the return of cold weather. Shore birds
perched disconsolately on the few emerging rocks or willows along
the shore of Tuluak Lake, but in a day or two the weather again
warmed and migration and nesting were resumed.

Many records, specimens, and accounts were presented tome. Every
report was clear as to location, time, and circumstances. These reports
covered observations made during the preceding winter at the camp
in the spruce timber along the John River, during the gradual travel
of the Eskimo people north through Anaktuvuk Pass, and during
their stay since the end of March at Tuluak Lake. The terse and
precise written or spoken narrative style of Simon Paneak impressed
me then with high regard for his powers of observation and his re-
spect for accuracy. The outline picture of the resident, migrant, and
nesting birds presented to me during my winter visit at Chandler
Lake began to gain supporting data. With my own eyes I could ob-
serve the intensity of the migratory and nesting activity. It was
plain that the arriving birds were robust and in fine shape. They
were not groping their way into the arctic regions exhausted by
forced flights nor were they harried by competitive search for a
meager food supply. On the contrary, the arctic mountain valley
provided so well for their sustenance that the long days were filled

469496—60—_3

24 U. 8. NATIONAL MUSEUM BULLETIN 217

with bird songs and the varied activities of flocks migrating and sepa-
rating for courtship and nesting. Feeding, in their daily program,
appeared to require only incidental attention.

With Neal Weber and Carl Henkelman I next visited the Valley
at the end of August 1948, and found that the people had moved to
Pitaich, or “Hole,” Lakes, a group of small lakes on the west side of the
valley just north of the mountain line. Asis common, the caribou had
been absent from the valley during the summer and only a few had
returned recently to relieve the meager diet of fish. Living was em-
bellished by supplies which we brought in, for I have made a practice
of carrying in rations more than adequate for our parties so that we
might in no way drain the natural resources upon which the resident
people depend.

Old squaws, white-winged scoters, and scaup ducks were numerous
in the many small Pitaich Lakes set deep, like limestone sinks, in
the high tundra about the camp site. I was pleased to obtain several
greater scaup ducks, to see over the tundra a marsh hawk flying as it
does in any climate, and to examine a sparrow hawk shot while hover-
ing curiously over a group of equally curious and excited children.
Longspurs, redpolls, and tree sparrows were numerous and restlessly
moving south; by the first snowfall in September they seemed to be
gone. We left early in September to escape the freezing weather
which would close the Lakes to float planes.

During the winter at Barrow Tom Brower and I discussed plans
for a careful survey during the nesting season of 1949, an undertaking
for which his keen interest and thorough knowledge of the birds of the
arctic coast eminently fitted him. Robert H. Stapleton visited the
mountain Eskimo people for me at their winter camp near the north-
ern edge of the spruce in the John River to take in some hardware,
tools, clothing, and food which, by easing their life somewhat, would
give them more time and better means of observing and recording for
our survey parties. Simon Paneak now understood the projected
survey so well that he and Bob Stapleton could settle upon the details
of our plans for the next summer.

Tom Brower landed at Tuluak Lake on April 26, 1949, and found
the Eskimo’s camp established. His arrival preceded that of the first
migrating birds, and he recorded them meticulously, collecting 215
specimens and 338 nests and eggs. He so carefully selected, studied,
and described the birds that his notes give a comprehensive picture of
migration and nesting. These notes were supplemented by Simon
Paneak’s records, which included the observations of the other people
in the village. When I visited Tuluak on June 1 with my colleague
Robert Rausch, it was evident that our information about the birds
was rapidly developing and that a complete record would soon be

ANAKTUVUK PASS 25

obtainable. In this summer, about 40 people from the Killik Valley
community of mountain Eskimos joined the people at Tuluak Lake
and the two communities have remained since that time at Anaktuvuk.

My departure from Barrow in midsummer prevented the review of
our observations until January 20, 1950, when, with Rausch, I visited
Simon Paneak again in the Eskimo winter camp at Kalutak Creek
on the upper John River (lat. 68°6’ N., long. 151°54’ W.). There I
was able to see some of the winter birds resident in the Pass, to discuss
the observation of the last year, and to make plans for the extension
of our survey. We also met Helge Ingstad, who was spending the
winter with Simon Paneak’s family while collecting material for
his book “Nunamiut” (Ingstad, 1954).

Traveling from Kalutak by air to Barrow I discussed Paneak’s
records and views with Tom Brower. We could now see that we
needed more information on certain birds and particularly on the
arrival of the birds and their nesting. This I obtained in July and
August 1950, while at the Imaiginik Camp with Paneak and my older
son, William Irving, who was investigating the numerous and varied
archeological remains in the mountain valleys. For about 15 days in
early August the three of us were able to collect on the Killik River
(see map, fig. 6) between Odrivik Lake (lat. 68°30’ N., long. 154°9’
W.), and the mountain line at Akmalik Creek (lat. 68°25’ N., long.
154°12’ W.).

In February 1951, I was able to consult again with Brower while
carrying on physiological experimentation on sled dogs at Barrow,
and with the reindeer herd at Topagoruk, about 70 miles south.
Review of our data on the Anaktuvuk birds showed that it could be
best improved by further observations of dates of arrival and by
information about weight of arriving birds, nesting, and the develop-
ment of young birds.

On May 11, 1951, my colleague John Krog landed at Contact Creek
and transported our supplies to the site on the east bank of the River,
opposite Imaiginik, where Simon Paneak, Elijah Kakena, and Frank
Rulland were camped with their families. Krog and Paneak care-
fully observed the arrivals and obtained weights of over 500 birds
during a spring season slightly earlier, drier, and more variable in
weather than usual.

The weather of May ran the gamut of temperature variation from
about —20° C. to 15° C., but had less than usual precipitation. Con-
sequently, when I arrived in June with William Irving the tundra
was drier than I had ever seen it, mosquitoes were a smaller plague
than usual, and walking was a pleasure. After Krog left, I continued
my observations of birds through the nesting period. Because of
the easy walking on the dry tundra our radius extended for about

26 U. S. NATIONAL MUSEUM BULLETIN 217

20 miles north and south along the Valley and 5 miles east and west,
but without going above 4,000 feet. Nests and downy young of many
species were found.

Early in July, after I left Tuluak, Simon Paneak and William
Irving for 18 days conducted a combined ornithological and archeo-
logical reconnaissauce in the Killik Valley from the forks at Easter
Creek in the south to Odrivik Lake north of the mountain line, Wil-
liam Irving then returned to Tuluak, where he remained until Sep-
tember 1.

While passing through Bettles, on the Koyukuk River, I fre-
quently had short opportunities to examine the birds there, but never
for long or at a time favorable for obtaining decisive data on migration.
In August 1951, however, John Krog and I were fortunate in being
able to make a journey of reconnaissance for three weeks by boat down
the Koyukuk to Alatna Village and thence for about 100 miles up
the Alatna River to Oscar Creek, just north of the southern mountain
line and well within the mountains.

In late February and March 1952, John Krog and I studied
the temperature of birds and mammals with Simon Paneak in the
vicinity of Tuluak Lake. In this year the willow ptarmigan were
very numerous, caribou moved south and west through the Valley
in large numbers, and wolves were frequently in sight. By trap-
ping and by his near phenomenal open-sight rifle shooting Simon
Paneak took 37 wolves, and the community took 160.

On our way back from Tuluak, we spent a week on the upper.
Alatna Malemute in Ernie Johnson’s hospitable cabin, located in a
spruce-forested valley high in the southern watershed of the moun-
tains, where the snowfall is heavy. That spring saw a heavy accumu-
lation of late snow in the mountains, particularly on their southern
slopes and the rivers on the northern arctic slope were nearly two
weeks late in breaking up, so that pilot J. L. Anderson was still
able to land a Cessna 170 with wheels on the ice of the lake at Sum-
mit on June 12. At Fairbanks, the emergence of buds and leaves was
nearly two weeks later than usual.

Terris Moore, President of the University of Alaska, became inter-
ested in seeing the passes through the Arctic mountains, and shortly
after commencement, on May 26, we set out in his plane to the head
of the Noatak River, where we spent a few days in Howard Pass.
From there we flew eastward through the mountain passes, crossing
the spectacular trails of a great westward migration of caribou, of
which we saw several thousand laggards still moving. We landed in
Anaktuvuk Pass, at Summit (called Nakrak by the Eskimo), where
for about three weeks I continued to observe the migrant and settling
birds in this uncommonly late season.

ANAKTUVUK PASS oy

154°30' 25° Pes 154°15"

68°31)’ 68°31
Odrivik Lake
KILLIK PASS
Statute Miles
a Jaane HS
COMPILED BY WM. IRVING Imaiknikpuk HS
1951 Lake Hs
\/3 Ikusik
DATA FROM SIMON PANEAK :
25 = 25"
20' ut a0"
Cr.
Kaiksuk
8 Pacn
Anvarvik
; aes
Toguyuk c
ac § 15
ve Nigaktutoy
Enikaklik f] ra
5 |
J/0 |
tl NE; 410
ri KS iksook a |
Soguke! 5 Xo, Kone |
: |
River |
zs —— ee
3B
€o,, Mayugiokluk
Cr
A.
re 68°05’
20° 154°15'

Ficure 6.—Sketch map of Killik Pass.

In early February 1953, I was detained in Bettles for 8 days by a
spell of weather below —40° C., too cold for flying to Anaktuvuk. The
time was used for making observations on the winter dessication of
leaf and flower buds. The snow cover was unusually thin and the
residents feared the effects of the consequent thick formation of ice
in the marshy ponds in which muskrats and beavers had to live in the

28 U. S. NATIONAL MUSEUM BULLETIN 2:17

small remaining supply of unfrozen water. When J reached Anak-
tuvuk in early March I found that both willow and rock ptarmigan
had been even more numerous than during the preceding winter, and
from Bettles northward along the John Valley their winter progress to-
ward the tundra was shown by their tracks in the snow. With Simon
Paneak I discussed his winter observations of birds and his collection
of data on the feeding and night shelter of the willow ptarmigan.

We met again in Bettles in mid-June to fly to Howard Pass, where
Terris Moore and I had briefly surveyed the snow-covered country
in the preceding spring (L. Irving and Paneak, 1954), and landed
on Itivlik Lake 20 miles east of the Pass. For 6 weeks we searched the
valley of the Ahlasuruk, finding 46 species of nesting birds. This is
14 fewer than in Anaktuvuk, and the absence at Ahlasuruk of some
migrants from the American continent that are very familiar in Anak-
tuvuk suggests that on the Ahlasuruk, 140 miles westward, the strength
of their westward migratory movement diminishes.

Since that time I have visited with Paneak and his family oceasion-
ally at several camps in Anaktuvuk Valley, at Savioyuk River, and at
Anivik Creek, the last two being winter camps in the northern border
of the forest. Simon and his wife Susie have continued to record bird
migrations, and in 1955 he visited me at Anchorage for a month.

In March 1957, while he was in Fairbanks to join our expedition to
Old Crow, Paneak developed an illness which was diagnosed as pul-
monary tuberculosis and he went to the Alaska Native Hospital in
Anchorage for treatment. His illness, when I saw him during the
summer confined to the hospital, was in sad contrast to his former
vigor. In October surgery promptly changed his outlook and pros-
pects and late in the winter we enjoyed his frequent visits in the
laboratory and for dinner at home. During March and April he
kept watch from the fifth floor of the hospital with a telescope, record-
ing the arrival of the earliest migratory birds. On April 22, 1958,
he was discharged, and left eager to see his family and home after
a year’s absence.

The Birds of Anaktuvuk Pass

The accounts that follow here and in subsequent chapters are pre-
ceded by a 5-column tabulation of the number and sex of all specimens
collected; the date on which they were collected; weight in grams;
fatness on a scale ranging from fat (F), medium fat (MF), little fat
(LF), to very little fat (VLF); and, where important, certain
measurements.

ANAKTUVUK PASS 29

Family GAVIIDAE: Loons

Gavia immer (Briinnich)

2 males June 4, 1948 weight (1), —— wing 379, 342 mm.
June 21, 1951 3600 g. culmen 78, 73 mm.
tarsus 79, 83 mm.

The measurements of these two examples of black-billed loons fall
among the small-sized group which Bishop (1921) designated Gavia
immer elasson. Inasmuch as I have not found a clear demonstration
that a small race is discrete in size or separate in distribution, I would
prefer to call these two specimens Gavia tmmer, as is done by Willett
(1933, p. 11) and by Rand (1947), who did not think that races
were distinguishable.

This loon is called Tasingik, meaning “black-billed,” in Nunamiut.
It is considered not common but of regular occurrence in the moun-
tains. Two black-billed loons were recorded seen May 27, 1949. A
pair was reported seen by Elijah Kakena at the lake near Summit
July 23, 1950. In June 1951, several were seen and heard in Tuluak
Valley so often that they were probably resident. One was observed
calling on Napaktualoitch Lake during two days in June while we
camped nearby. Calls and occasional subsequent views in flight near-
by suggested that lake as the center of a nesting pair.

In Anaktuvuk Valley no nests have been found, but on a lake in Ok-
milaga Valley, about 50 miles west, a pair was known by Simon Pa-
neak to have nested several years ago. During July 1951, several black-
billed loons were observed and heard by Simon Paneak and William
Irving in the Killik River Valley.

They are only seen on large lakes where fishing for human food is
likely to be good. Without wind they cannot take flight, whereas
yellow-billed loons take off somewhat more readily. Loons of both
species frequent large bodies of water where they are clearly visible,
and the black bill distinguishes them from yellow-billed loons at great
distances in the bright light of arctic summer.

The lakes and country are daily searched by the Nunamiut with tele-
scopes in their surveys for game, and large loons and even small birds
are carefully examined until identified. All birds seen are noted and
with other natural phenomena are the subject of careful discussion
when people get together. Every such observation which a mountain
Eskimo makes or hears seems to become recorded in his memory, for
I have often asked them to recapitulate records and never found de-
viation in a statement of factual observation. It is therefore unlikely
that there was another pair of black-billed loons in the Valley between
Summit and the mountain line in 1950, but there may have been more
than a pair present in 1951. The black-billed loons acted like steady

30 U. S. NATIONAL MUSEUM BULLETIN 217

residents in a lake, whereas the yellow-billed loons which were seen
more often, appeared to fly more readily from lake to lake.

In the windy weather of late summer, black-billed loons are often
heard calling while in flight. Simon Paneak thought that they fre-
quently called while flying through light rain, but he doubted if the
flight call predicted the onset of rainy weather.

The first records of the season are June 4, 1948, May 27, 1949, May
21,1951 and June 1, 1954.

On the arctic coast Bailey (1948) mentions the report of a large
black-billed loon which had been killed east of the Colville mouth.
Anderson (1921) had been told by Leffingwell of one killed near Flax-
man Island, and Leffingwell (1919, p. 651) reported that only one
had been seen there and that the Eskimos only knew of one being
seen on the Colville. I found among the Nunamiut the recollection
of circumstances about the loon which Leffingwell mentioned and
which is apparently the basis for all these published comments. ‘They
consider it a rarity on the coast, but I have found several coast Es-
kimos who know the black-billed loon from one observation or through
a report from other Eskimos, and I do not question that black-billed
loons occasionally migrate to the arctic slope and coast.

The specimens and these accounts show that émmer is regularly pres-
ent in and sometimes nests in the mountains. In addition a few prob-
ably migrate northward for nesting, for a pair was known by Simon
Paneak to have nested some years ago north of Anaktuvuk on the lower
Colville River.

Gavia adamsii (G. R. Gray)

1 female - Sept. 28, 1949 — — wing 375 mm.
culmen 93 mm,
1 female, not June 9, 1951 weight 4050 g. medium fat —
preserved

Yellow-billed loons, by the reflection from their bills, are easily
distinguished from common loons at surprisingly great distance in
arctic light.

The earliest recorded arrivals are May 27, 1949, May 20, 1950, May
28, 1951, May 30, 1952, May 23, 1953 and June 1, 1954. Two were
seen at Odrivik Lake in the Killik Valley daily August 13-15, 1950.
On July 6, 1951, Simon Paneak and William Irving saw a pair with
one young ona lake near Amorgoayat in the Killik Valley, where they
also reported seeing frequent pairs and on July 10, observed six
together in Akmalik Lake.

To the Nunamiut the yellow-bill is well known as Tootlik. Like
black-billed loons they frequent the larger lakes, where they are sure
to be observed by the frequent surveys of the countryside which the

ANAKTUVUK PASS 31

Eskimos make with their telescopes. Their call is unlike that of the
common loon, and for another contrast, yellow-billed loons take
wing more readily. No nests have been found in the mountain valleys,
but these loons are seen and heard frequently during the summer.
The Nunamiut consider black-billed loons more common in the
mountains, although I was inclined to regard yellow-billed loons more
numerous. I now think that the Nunamiut view is correct and that
mine was formed because yellow-billed loons fly more frequently from
lake to lake and in late summer occasionally indulge in aerial
maneuvers.

The regular presence of these loons and the observed young are
assurance of their nesting in large lakes near the mountains. There
are few lakes large enough for them or for black-billed loons within
the mountains in Anaktuvuk Valley, and so it is thought that most
are visitors from larger lakes north of the mountain line, whereas
the black-billed loons which are seen appear to remain steadily within
a more restricted summer territory.

I consider yellow-billed loons to be migrants that visit and rarely
nest in the mountains.

Gavia arctica pacifica (Lawrence)

1 male June 23, 1949 weight 2326 g. — —_
1 female June 14, 1949 weight 1850 g, — —_
1 downy young July 31, 1950 weight 1242 g. — ——
male, probably
of this species

The earliest recorded appearance of Pacific arctic loons is June 1,
1949. One or two weeks later, when the ice breaks up completely in
the lakes, many of those a half mile across or larger, are occupied by a
pair of Pacific loons. At the end of July, one parent may be seen
closely accompanied by a large downy young or occasionally two.
A parent and young were observed on the landing lake near Imaiginik
and two days later a large young downy loon was brought in from
there. It is listed with question under this species, to which it probably
belongs.

The Pacific loons are at least 10 times as numerous as black-billed
loons. Like the yellow-billed loons, they fly swiftly and frequently,
and these two species seem to fish over a considerable area. They
frequent larger bodies of water than the more sedentary red-throated
loons, and are only about a tenth as numerous.

As Malirgik they are well known by the Nunamiut to be regular
nesting residents in the lakes of the mountain valleys. The greater
numbers seen in spring and late summer are thought to be migrating.

32 U. S. NATIONAL MUSEUM BULLETIN 217

Gavia stellata (Pontoppidan)

1 male June 11, 1949 weight 1900 g. — —

1 female June 11, 1949 weight 1522 g. —— —

1 male, not pre- Jume 28, 1951 weight 1711 g. — ===
Served |

1 female, not pre- June 2, 1951 weight 1613 g. — eggs 30 mm. and 20
served. mm,

The earliest recorded appearances of red-throated loons are May 23,
1950, June 2, 1951, May 18, 1953, and June 1, 1954. The size of the
developing eggs found in the bird on June 2, 1951, shows that the
nesting date was close at hand. During summer they are common,
particularly in smaller lakes, even those high up among the mountains,
entering some which may be less than 100 yards across, although
when these are narrow they are usually elongate. In these restricted
waters they are much less shy than the other loons, and late in summer
they may be seen there with their young. They are the most numerous
of the loons and, because of the number of small bodies of water suit-
able to them, may be ten times as numerous as Pacific loons.

A late observation, September 8, 1950, at Contact Creek suggests
that these loons traverse the mountains southward in fall migration,
for at that date the lakes north of the mountains are likely to be ice
covered.

In early June, they hold closely to the small lakes and are not as
often seen flying as are the other loons. But at the end of June, their
call in flight begins to be heard. The call is the basis for their Nuna-
miut name, Aaksrauk. During July and August, this fast repeated
call is much heard overhead, and it is difficult to connect it with the
bird, which flies so swiftly and so high that the sound seems to trail
behind its origin. To search for the source of the call high in the over-
cast above the arctic tundra is a dizzying effort, but the Nunamiut like
the trial of locating Kaksrauk in flight.

Red-throated loons nest in the valley and migrate northward and
southward in numbers.

Family PODICIPEDIDAE: Grebes

Podiceps grisegena holbollii Reinhardt

Red-necked grebes have not been reported in the mountains, but
Sidney B. Peyton told me that he found one floating dead in a lake
near Bettles Village late in June 1954. Charles Sheldon of Kobuk
reported them common there, with the Eskimo name Shoolishookruk.
Since red-necked as well as horned grebes occur along the Koyukuk
and Kobuk they may likewise be expected to venture north into the
mountains.

ANAKTUVUK PASS 33

Podiceps auritus cornutus Gmelin

2 males June 25, 1951, weight 481, 459 g. — —
Tuluak Valley

I had repeatedly described grebes to Simon Paneak, who said that
his father knew them as Malikak, meaning “small loon,” and that he
remembered his father shooting several in 1907 with his .44-caliber
rifle when the family was about 40 miles above the mouth of the Col-
ville River. Jesse Ahgook also remembered seeing dfalikak, along the
Colville, but in the scale of his experience with 80 years of life the
date was only recalled as “long ago.” Elijah Kakena and Frank
Rulland did not think they had seen a grebe.

On June 25, 1951, a horned grebe pushed its weird looking head out
of the horseshoe lake just west of Akvalutak Creek within 30 feet of
me and promptly drew it under water again. During the next two
hours, Frank Rulland and I ran around the lake, locating three male
horned grebes and obtaining the two male specimens, both with small
testes and little fat.

In camp, Simon at once recognized the birds as the ones which his
father showed him in 1907 and called A/alikak. As Frank Rulland de-
scribed the appearance of grebes and their methods of diving, along
with a graphic and humorous commentary upon our hunt, it was evi-
dent that no one of the people had seen them before in the mountains.

Six weeks later on the Alatna River just below Helpmejack Creek,
which is just south of the southern mountain line, John Krog and I
saw four horned grebes fishing in the fast current of a sharp bend in
the river. This location was about 100 miles southwest from
Anaktuvuk.

I believe that these grebes only rarely come into the mountains.
I class them as visitors because they are not far from their common
range in the Yukon and Koyukuk Valleys.

Family ANATIDAE: Swans, Geese, Ducks

Olor columbianus (Ord)

The earliest northward flights of whistling swans is recorded only
in one year, May 20, 1949, when a flight passed over Tuluak Lake.
To the Nunamiut, they are known as Hogruk. Every year some are
seen and heard passing over, often flying higher than the mountain
tops. It is thought that, unlike the birds of weaker flight, they do not
confine their course through the mountains to the passes. A few occa-
sionally land and the flights and landings through the Killik Valley
are said to be more numerous than in Anaktuvuk. On June 10 and
16, 1951, we saw a single swan in Tuluak Valley in grayish plumage,
but the flights have passed by early June. I have also seen one in

34 U. S. NATIONAL MUSEUM BULLETIN 2:17

grayish plumage near Anivik Lake late in June. In fact a lone swan
of grayish color has now (in 1954) been seen a few times in each
summer. The summer occurrence and its appearance are so unusual
that it is spoken of as if it were one individual.

Swans are not known to have nested recently in the mountains, but
Simon Paneak recalls that a swan’s nest was found by his father
about 50 years ago on a lake near Akmalik Creek in the Killik Valley.
He also recalls finding swans nesting along the Colville River from
the junction of the Anaktuvuk to its mouth. Swans are so conspicuous
on the tundra that their nesting would be observed. They are re-
garded as regular migrants which do not nest in the mountains.

Branta canadensis taverneri Delacour
3 males May 18-20 weight (1) 2551 g. aaa ae
3 females May 18-23 weight (1) 2150 g. — —

Until clarification of the systematics of the Alaskan forms of Branta
canadensis, by Delacour and Zimmer (1951), a single specimen from
Anaktuvuk was regarded as close to but not identical with specimens
marked leucopareia in the U.S. National Museum.

With the aid of Herbert Friedmann I compared the single specimen
then available with a specimen obtained by John Krog at Amchitka
(J. Krog, 1953). By Delacour’s definitions the Amchitka goose was
leucopareia and the Tuluak bird was taverneri.

In August 1951, John Krog and I collected 4 males and 2 females
along the Koyukuk and Alatna Rivers within about 100 miles of
Anaktuvuk. These were identified as tavernert. They were among
family groups containing well grown young which were just achiev-
ing flight in early August. By mid-August families were beginning
to associate and a little later well formed flocks were assembling the
geese of the vicinity.

In 1952 geese were collected by Tom Cade from among families of
that season along the Colville River and identified by Delacour as
taverneri (Kessel and Cade, 1958).

In 1953 five specimens were obtained from the flights which alighted
more frequently than in other years at Anaktuvuk. These are all
taverneri. ‘The testes of three males measured May 18-20, 1953, were
26 mm. long. Two of the birds were noted to have little and one much
fat. In a female examined May 18 the largest egg measured 22 mm.

Charles Sheldon obtained a specimen for me from Kobuk village
on May 19, 1954. It was the male in breeding condition of a pair
and the specimen was identified by Dr. Herbert Friedmann.

It is thus established that ¢averneri nests along the rivers of the
wooded southern watershed of the central Brooks Range. They mi-
grate through Anaktuvuk in spring, where they have been earliest
reported May 22, 1949, May 11, 1950, May 22, 1951, May 18, 1953, and

ANAKTUVUK PASS 35

may 15, 1954. Southbound flights are seen in late summer but they
land less often than in spring. One fiock passed over Kangomavik
on August 26. None are known to nest in the mountains, and the
migrating flocks stop for only a brief time. During nesting season
they are also on the Colville River near Umiat. These two nesting
areas are different and separate, one being on the rivers in thickly
wooded country and the other 250 miles north, on the treeless arctic
slope beyond the intervening mountains.

Geese of the Canada type are known to the Nunamiut in the moun-
tains and on the coast as Eksrahgotiliz:, “light-colored cheek.” The
occurrence of differences in size has been observed but has not been
regarded systematically.

Many of the geese of this as well as of other kinds are regularly
seen in migration, and judging from those which are seen and heard
a great many traverse the mountains. Often they. fly higher than the
mountain peaks and it is thought that they need not follow the valleys
except occasionally, when they wish to land.

Branta nigricans (Lawrence)
1 male May 30, 1952 weight 1494 gr. — a

The single specimen is normal to the characteristics of Pacific black
brant (Delacour, 1952). Its testes at 20 and 25 mm. in length were
probably not mature.

The earliest recorded observations of brant are May 26, 1950, May
96, 1951, and May 29, 1952. Usually they are the most commonly
seen geese in spring, and land most frequently. But in 1953, when
other geese were numerous, no brant were seen. Often they fly low
through the valley, in their swerving and circling flight calling as if
seeking a landing place. At other times they fly direct at the eleva-
tion of the high mountain peaks.

Brant have not been observed in the mountains during the late
summer on the southbound course. Along the arctic coast near Bar-
row, I did not see migrating flocks of brant as often in the spring as
in the late summer when the flocks commonly flew westward low in
the air above the coastal tundra or over the sea close to the coast.
Their migratory course in spring probably crosses the interior of arctic
Alaska at many points, for they are known there by the Nunamiut in
Chandler Lake, Okomilaga, and Killik Valleys. They are also
common spring migrants at Kobuk and Shungnak, whence they ap-
pear to head northward in the direction of Howard Pass.

The Nunamiut call brant Viklinagak, meaning “almost like (white-
fronted) geese.” It is well known to them that many brant nest along
the arctic coast about the mouth of the Colville River, but none are
known to remain for nesting in the interior.

36 U. S. NATIONAL MUSEUM BULLETIN 2:17

Anser albifrons frontalis Baird

female May 23 weight 2844 g. — —

White-fronted geese are familiar migrants, making frequent spring
and fall flights through the mountains. The earliest observations are
May 16, 1949, May 21, 1950, May 9, 1951, May 18, 1952, May 18, 1953
and May 15, 1954. In Nunamiut they are called Vklivzk, which means
“eoose.” Most of the numerous flights are heard passing high among
the mountains during the short nights of late May. Landings have
occasionally been observed in spring but not from the southbound au-
tumn flights. The nearest nesting of these geese known to the Nuna-
miut is along the lower Colville River. It was my impression that
white-fronted geese flew higher and more directly on their course
than any but snow geese.

Chen hyperborea hyperborea (Pallas)
1 female May 20, 1954 — very fat egg 30 mm.

Snow geese were first reported seen on May 20, 1949, May 15, 1950,
May 19, 1951, May 14, 1953, and May 15, 1954, and many landed in
Anaktuvuk Valley just north of the mountain line in 1948. In recent
years none had been killed until a few landed in 1954. ‘These geese,
called Aangok in Nunamiut, are known as regular spring migrants
which sometimes land in the Valley. They often fly high among the
mountains, being the highest flying of the migrating birds. Occa-
sionally they are seen in the fall, but they are not known to land at
that season. A few are known by the Nunamiut to nest about the large
river mouths on the eastern Alaskan arctic coast. Snow geese are
more rarely seen than other geese and usually only in small flocks.
Occasionally one or a few of them join in the flights of white-fronted
geese.

Anas platyrhynchos platyrhynchos Linnaeus

2 males May 24, 1949, June weight (3), 1099, — —
26, 1951 1151, 1210 g.
1 female May 28, 1949 weight 800 g. == ==

The earliest records of common mallards are May 20, 1949, May
16, 1950, May 13, 1953, and May 15, 1954. To the Nunamiut they
are well known and are called Ogiuguk, a word representing their
call, and a better representation of the sound than our “quack.”
Six mallards were reported May 24, 1949, several groups of 8 to 10
were seen in 1953 and a few are seen in each spring. A male was
seen in the Killik Valley near Odrivik Lake August 10, 1950, and a
moulting male unable to fly was taken June 26, 1951, in Pitaich Lake.
None are known to nest in the mountain valleys, but from the fact that

ANAKTUVUK PASS BY /

they are seen in pairs in spring or singly in summer, it is believed
that nesting occurs nearby.

Anas acuta Linnaeus

5 males May 21-June 25 weight (18) (May — —_—
10-June 25) 754 to
1030, average 817
1 unlabeled, in as ious eas nee
male plumage
5 females May 21, Aug. 7 weight (7) without — —
eggs (May 17-
June 11) 679-835,
average 731 g.
lnest, 6slightly June 10, 1949 — — —
incubated eggs,
female parent

Pintails were earliest recorded May 16, 1949, May 11, 1950, May 6,
1951, May 18, 1952, May 13, 1953, and May 15, 1954. The last date
was probably several days after the first arrival. On May 17, 1949,
12 were recorded by Tom Brower and thereafter he reported the num-
bers seen as increasing to between 60 and 100 daily from May 21 to 30.
The northbound migration goes on steadily during at least two weeks
and many flocks comprise a dozen or more and are well formed. Some
of these migrants stop to nest in the valleys, where in the extensive
marshy regions they are the most numerous of the summer resident
ducks. The number which remains to nest in the Valley is however,
only a small percent of the great number of migrating pintails.

In 1951 developing eggs were found to be 6 mm. in length on May
18, one was ready to be laid on May 21, 12 mm. in length on May 22,
and 12 mm. in length on May 28.

A nest found in a marshy area by Tom Brower June 10, 1949, with
6 slightly incubated eggs, was made of small grasses mingled with
down. Another set of 6 eggs was discovered on the same date but
not taken. It appears likely that eggs are laid between about May 20
and June 10. These dates are earlier than for the old squaw and
merganser. A female with 8 young was observed in Akmalik Lake
in the Killik Valley by Simon Paneak on July 8, 1951, and on August
28, 1950, young pintails were reported able to fly for some distance.

On June 16, 1951, six males were observed together at Napaktua-
loitch Lake, and thereafter the males were commonly grouped in small
bands along the streams and in the lakes. One male taken on June
16, 1951, was moulting and just able to fly, while three others took wing
with difficulty, unlike their usual ability to spring from the water
into flight.

The Nunamiut name for the pintail is Korugak. 'They are the most
numerous migrating and nesting ducks in the Valley.

38 U. S. NATIONAL MUSEUM BULLETIN 2:17

Anas carolinensis Gmelin

6 males May 24-June 26 weight (6) 293-379, — —
average 332 g.
3 females May 20, 29 weight (3) 265, 297, — ——

334, average 343 g.

The first green-winged teal were reported seen at Tuluak Lake on
May 20, 1949, May 22, 1950, May 11, 1951, and May 18, 1953. As
Korualorgosik, meaning “smaller than pintail,” they are well known
in migration and as summer residents. The nests had not been found
but on May 20, 1951, a female contained an egg 26 mm. long. The
bird then weighing 410 grams. In the cold spring of 1952, teal were
not seen until June 2 when the testes of a drake were 35 mm. in length
and a duck contained a 12-mm. egg. Two days later a small duck
contained a 85-mm. egg. About this time numerous pairs and single
teal were present in the Valley. On June 5, 1950, Raymond Paneak
found a nest with 6 eggs. On July 11, 1950, a female with 9 downy
young, the wing feathers just showing, swam from Nakagnik Creek
to Tuluak Lake. Young birds are seen each summer. In 1950 they
were lastr eported seen August 27.

The teal nest in the Valley regularly and some migrate northward.
Because they keep much in cover and do not congregate in flocks,
they are probably a larger population than appearances indicate.

Mareca americana (Gmelin)

2 males May 24, 1949 weight (1), 714 g. — —

The first American widgeon were seen May 24, 1949, May 20, 1950,
and May 18,1953. The Nunamiut name is Horuaknak, meaning “like
pintail.” Small groups arrive in spring and they are occasionally seen
in summer, but no nests have been found in the mountains. In 1948,
an adult and six young were seen on the west side of Tuluak Valley.
A female and about eight large downy young were seen August 6,
1950, on a lake about four miles north of Akmalik Creek in the Killik
Valley. They are less numerous residents than green-winged teal,
and they are estimated to be fewer in migrating numbers, but they
are usual summer residents and nest in the mountains.

Spatula clypeata (Linnaeus)
1 male June 1, 1953 weight 578 g. — —
Two shovelers were reported by Thomas Brower on May 238, 1949,

and one on May 26. Since he has obtained specimens of shovelers at
Barrow his identification is correct. Simon Paneak recalled seeing

ANAKTUVUK PASS 39

a shoveler shot at Humphrey Point in 1926 and although they are con-
sidered rare in the arctic I find that they are well known to many
Eskimos. A rare bird may be well known to Eskimos because each one
which appears in any way unusual is studied by all who are near
and the circumstances are discussed for years among a widening
circle. I mentioned that three reports in the literature of the arctic
coast appeared to stem from one black-billed loon which Leffingwell
(1919) and some Eskimos saw sometime before that date. I have
also heard of this same loon from several Eskimos and in different
villages. So the well known shoveler is familiar from a few well re-
membered observations but it is rare in central and eastern arctic
Alaska from the mountains northward.

On June 1, 1958, Elijah Kakena saw a pair and obtained the male.
Its testes at 5 mm. were not much developed. These reports warrant
designating the shoveler as migrating through Anaktuvuk, although
I have no report that they nest north of the mountains.

Aythya marila nearctica (Stejneger)

3 males May 24-June 4 weight (17), 844—- — ee
1046, average 932
g.

5 females May 27-July 10 weight (9), 856-1117, —_ —
average 957 g. ‘

1 downy male July 10, 1951 weight 91 g. ae ——

ldowny female July 10, 1951 weight 89 g. — —~

Throughout the north, ducks are important for human food and
are consequently so shy that I could rarely come close enough to distin-
guish by sight the greater from the lesser scaups. Among those
taken for food, when closely observed, the extent of white on the wing,
the larger bill, and the heavier weight of the greater scaup distinguish
the latter from lesser scaup. No intergradation of these characters
was noticed.

Judging from the number of birds taken by the Eskimos the greater
scaup are probably at least ten times as numerous as the lesser. The
earliest recorded observations of scaup are May 24, 1949, May 19, 1950,
May 24, 1951, and May 14, 1953, and as specimens of both forms were
taken early, they apparently arrive at about the same time, coming
in numerous small flocks of both sexes which soon break up for mating.
Many continue their flight northward. No nests have been found, but
there is no doubt that the greater scaups, which among ducks are next
in summer numbers to old squaws and pintails, nest about many of
the larger lakes.

469496—60-——4

40 U. S. NATIONAL MUSEUM BULLETIN 2:17

The progress of egg development in the female greater scaup, as
shown in the tabulation below, indicates that in 1951, eggs were ready
to be laid June 5 and that one female had not completed laying on
June 16.

Date (1951) Weight (g.) Length of eggs (mm.) Date (1951) Weight (g.) Length of eggs (mm.)
25 May 856 4, 2 15 June 1067 8
28 May 918 15 15 June 985 6
28 May 1117 +15 16 June 1034 31, 26, 17, 12, 10,
28 May 978 1,2 8, 5, 3
28 May 964 4,2 10 July 740 (with downy
31 May 992 young)

5 June 1069 (1 larger egg

broken)

Young birds are often seen during the late summer accompanied by
females. A female greater scaup, remarked upon as containing very
little fat was collected at Kidgavik Lake, near the northern moun-
tain line in the Killik Valley on July 10, 1951 with two downy young.
These young birds, weighing 91 and 89 grams were then heavier than
I would expect the eggs to weigh. I would guess that they were not a
week old, and that the eggs from which they came had been laid early
in June. Young scaup, not distinguishable as to the species were
recorded as unable to fly on August 28, 1950, in Tuluak Lake.

Until about mid-June, male and female birds were observed to-
gether, and a female, collected with a male on June 16, 1951, contained
eggs, the largest of which measured 31 mm. and was nearly ready to
be laid. Two days later two small flocks of four and six males were
seen on Napaktualoitch Lake and on June 28 several flocks composed
exclusively of from 20 to 30 males were assembled near the center of
Margaktuk Lake.

The Nunamiut call the scaup Aaklutuk, meaning “big-billed.”
Although they recognized that variations in size occurred they did not
distinguish between the two scaup until the decisive characters were
pointed out. They were not impressed by the difference until after
several years of examining and weighing the birds. Now they accept
the difference in size and bill as distinguishing the greater from the
lesser scaup.

Aythya affinis (Eyton)

4 males May 27-June 4 weight (3), 769, 681, —_— —_
694 g.

4 females May 29-Sept. 10 weight (2), 601, 651 —_ —
(thought to be g.
of this form)

1 large downy Aug. 7, 1950 weight 350 g. — —
male

1 large downy Aug. 7, 1950 weight 331 g. —_— —

female

ANAKTUVUK PASS 41

The fact that similar numbers of specimens of adult greater and
lesser scaup were taken does not indicate populations of equal size.
Out of about 100 dead scaup which I have examined only about 10
were lesser scaups, and since I looked particularly for them, I suspect
that the greater scaups are more than 10 times as numerous.

Lesser scaup were taken about as early as the greater, so that their
time of arrival is similar. Flocks of ducks are very difficult to
approach, so that most specimens are singles. A male collected on
May 31, 1951, at Imaiginik by John Krog was one of nine scaup shot
on that day in the same lake, all the others being greater scaup. A
female which I shot on June 28, 1951, was alone in the outlet stream
from Margaktuk Lake, but in the lake nearby were over 80 males.
Two females were collected September 10, 1950, about the latest time
when ducks could find extensive open water, but I have also examined
greater scaup shot at about that time. I cannot report any difference
in the habitat of the two forms, but they are certainly frequently
associated,

The female which was shot on June 28, 1951, had already laid eggs
and from its actions might have had a nest nearby. The two downy
young birds taken August 7, 1950, in a small lake near Akmalik Creek
in the Killik Valley were among six young ducks with an adult female
scaup, a pintail, and a white-winged scoter. The scaup attended the
young most closely, and the young were not scoters and look like
downy scaup ducks. AsI watched them, for some time I thought that
the attendant bird was a lesser scaup, but I was unable to confirm my
view by collection.

From these observations and their presence throughout the summer
I consider that the lesser scaup nest in the mountain valleys, but not
having seen them further north I cannot say that they migrate beyond
the mountains.

As I remarked in describing the greater scaup, the Nunamiut have
only one name, Aaklutuk, for scaup, and they did not distinguish
between them until several years of observations showed that the
size characters made two discontinuous groups. While they know
that scaup nest very commonly in the mountain valleys, they cannot
add evidence for the nesting of the lesser form.

Clangula hyemalis (Linnaeus)

1 male June 1, 1949 weight (6) 621-868, —_ =
average 778 g.

1 female, nest, and June 30, 1949 weight (6) 510-713, — ===
7 fresh eggs average 606 g.

A young female oldsquaw on August 25, 1948, weighed 380 grams.
The earliest records are May 21, 1950, May 17, 1951, and May 23, 1953.

42 U. S. NATIONAL MUSEUM BULLETIN 217

Few dates have been recorded, perhaps because the arrival of the
oldsquaws is such a familiar event.

The largest unlaid eggs were 10 mm. in length on May 31, 1951, and
34 mm. on June 5, 1951.

The nest was situated on a dry slope near a small stream, and it was
composed of dark-colored down entangling some small dry leaves. If
the date of fresh eggs obtained on June 21 is representative, nesting
is late and growth must be rapid to reach the weight of 380 grams
observed on August 25.

Oldsquaws and pintails are the most numerous ducks in the Valley.
They frequent open water, where in spring they are conspicuous and
bring attention to their presence by their pleasant notes. In Nuna-
miut they are called Ahalik, which represents their call. Eskimo chil-
dren try to toll any ducks with this call and frequently the ducks
seem to be attracted. The large lakes usually contain several families
of oldsquaw and even the small lakes may provide nesting sites, al-
though islands seem to be preferred.

Coast and inland Eskimos like and admire these hardy arctic birds.
As they fly through the haze on the arctic coast or in the mountains,
ascending and descending as if to get ground bearings, it is most de-
lightful to hear their musical calis, which give the impression that
the oldsquaws are signaling to each other. The individuals of a
group diving in the lakes or ocean seem to watch each other and to
seek food in a group, like a fleet of fishing boats. On the coast when
a boat goes among them they continue their fishing, opening up the
fiock to allow it to pass. They do not often take precipitous simul-
taneous flight as do the nervous scaup ducks or wary geese.

For all their abundance and lack of anxiety, they are not easy birds
to shoot, and their loose system of organization never concentrates
them like eider ducks so as to allow numbers to be taken at one shot.
As food for man they are no delicacy, although superior to loons and
scoters, and while they are good for hungry people, it requires more
ammunition and hunting to obtain nutrition from oldsquaws than
from any other common arctic duck.

I find them the most interesting birds of the duck family. I am
sure they associate in well organized societies and they are, as far as
I know, the only arctic ducks, which are not subject to flock alarms
and retain their presence of mind and flock organization when an
enemy has penetrated among them.

Histrionicus histrionicus (Linnaeus)
3 males June 2-4 weight (2), 639, 618 — —

Ze
1 female June 2 weight 562 g. —— ae

ANAKTUVUK PASS 43

Western harlequin ducks were first reported June 2, 1949, May 31,
1950, and May 29, 1952. After that time they are occasionally seen,
usually along the swift streams. In 1951, however, we did not record
a single harlequin, although they are well known and conspicuous in
habit and profile. Often they are called “current ducks” by the Nuna-
miut from their frequenting fast water, but the regular Nunamiut
name is Ahaliknak, “like old squaw,” which well indicates their adept
diving. Sheldon (1911) reports that harlequin ducks occupied most
of the swift streams along which he passed to the sheep country high
in the mountains of western Yukon Territory. In August 1951, Wil-
liam Williams of Alatna Village showed me a pair of harlequin ducks
which his wife had mounted very nicely. These were shot in 1950 on
the Koyukuk from a group of about six, and he considered them to
be of common occurrence in that region. During three weeks on the
Koyukuk and Alatna Rivers in August 1951, however, we saw none,
and Williams had no positive observations in that year.

They usually remain evident during summer, and several families
of young have been seen by the Nunamiut, although nests have not
been discovered. Resident harlequins may be nearly as numerous as
baldpates but they are fewer than green-winged teal. The Nunamiut
knew them along the Colville River, and I do not hesitate to say that
some seen in spring at Anaktuvuk are migrating farther north.

Polysticta stelleri (Pallas)

Steller’s eider was well known to the Nunamiut, when they were
on the arctic coast, as Lknikauktok. Simon Paneak reported to me
his great surprise at seeing a male and a female in late June standing
on the gravel along Contact Creek. He watched them in amazement
through his telescope to verify their marks but they flew before he
could approach within range of shot. Although we are confident of
the identification we agreed that the observation of an eider duck in
the mountains was too anomalous for its inclusion in the list of Anak-
tuvuk birds. It was Simon’s suggestion that these eiders may have
ascended the Colville looking for another ocean.

Melanitta deglandi (Bonaparte)

3 males June 11-14, 1949 weight (6), 1413- _ a=
1907, average 1650
g.
2 females June 17, Aug. 6 weight (2), 1548, —_ —
1700 g.
2 downy young August 6, 1950 _ — —
females with
female adult.

The first white-winged scoters were recorded May 31, 1950, May 24,
1951, May 20, 1953 and May 28, 1954. No large flocks are reported,

44 U. S. NATIONAL MUSEUM BULLETIN 217

and it appears that they come in principally to nest. At any rate,
there are white-winged scoters on many of the larger lakes in the
Valley and upon some of the smaller ones. A female with nine small
downy young was seen July 26, 1950, on the landing lake near Imaigi-
nik, On a small lake near Akmalik Creek in the Killik Valley the
female and two downy young were collected from a group of five on
August 6,1950. This was one of the deep round lakes about 200 yards
across, sunk like a limestone sink in the terrain of wind blown sand
on river gravel. The scoters swam always near the opposite shore,
but by running around the lake it was possible to approach them, for
the young, which would weigh about 500 grams, swam very slowly.
The young could dive well and swim some 50 yards under water, but
the devoted female did not leave her escort duty even when the young
dived and scattered.

These ducks, fewer in numbers than scaup, are well known as
Tongargagruk, meaning “devil” in Nunamiut. Asa few are known to
range far north to the arctic coast, some of these observed in spring
are undoubtedly migrating northward. Eggs contained in one female
bird measured 30 and 20 mm. in length on June 5, 1951.

I could not distinguish my specimens from specimens of white-
winged scoters from the east and west coast of America. It was also
apparent that the bills of my scoters, which were taken while soft and
perhaps distended in early summer, had warped into shapes influenced
by conditions of drying. Accordingly, I could not assign them to a
western form on the basis of small variations in size and shape of bill
and I could see no other distinction warranting their assignment
among specimens of eastern or western origin.

Melanitta perspicillata (Linnaeus)

1 male May 20, 1951 weight (5), 964-1006, — —
average 987 g.
1 female May 20, 1951 weight 980 g. — Ss

Until 1951, my knowledge of surf scoters in the mountains was
limited to Nunamiut accounts, which gave clear descriptions of their
appearance and habits, and the Nunamiut name Avilyuktok. It was
reported that a number of these scoters regularly spent the summer on
Chandler Lake and that young birds were seen there. None were re-
ported in Tuluak or adjacent valleys during 1948, 1949, and 1950.
From their conspicuous appearance and common choice of the open
and deeper lakes, I believe that very few could have been present if
they escaped the keen observation applied to the country often with
the aid of 15-power binoculars and strong telescopes.

In 1951, four were observed on Tuluak Lake on May 20. ‘These were
collected and proved to be three males and one female, the latter with

ANAKTUVUK PASS 45

its largest egg 6 mm. in length. Two of the males had large testes
and were moderately fat. The female was very fat. The third male,
which was wounded, was not recovered until two days later, at which
time it had little fat remaining. Another male which I shot near
Pitaich Lake June 15, 1951, had little fat.

Along the Killik River near Togoyuk Creek in the mountains, one
was observed on July 5, 1951, by Simon Paneak and William Irving.
They reported that on June 18 a flock of about 100 surf scoters landed
on Odrivik Lake, about four miles north of the mountain line, and
that 5 landed there on July 15 and 10 on July 16.

I have no observations upon nesting in the Anaktuvuk or Killik
Valleys, but I consider that the report of young birds at Chandler
Lake and the numbers and conditions of the birds observed during
1951 indicate that surf scoters nest among the Endicott Mountains.
The irregular records indicate that the numbers of surf scoters which
reside or migrate fluctuate widely from year to year, and they are the
only birds I know to vary conspicuously in annual numbers at Anak-
tuvuk.

The surf scoters I have eaten were the nearest to being objectionable
meat that I have encountered in a country where any meat is acceptable.

Mergus serrator serrator Linnaeus

3 males May 26, 30-June1 weight (5), 992- aaa —_—
077 g.
1 young partly Aug. 29, 1950 _—_ _— nest with 9 eggs
feathered
June 24, 1949
2 females May 30, 1951 weight (3), 808-975, — -——

average 918 g.

The first recorded red-breasted mergansers were seen June 2, 1948,
May 23, 1949, May 21, 1951, May 29, 1952, and May 18, 1953. In the
latter part of May, many of the lakes are occupied by a pair of
mergansers, which in spring regularly travel in pairs, as do many
mallards, teal, and harlequins. I did see one group of six mergansers
flying north on May 29, 1952.

A nest with very large eggs was found by Thomas Brower among
low willows about ten feet from the shore of a small island. Only
one of the eggs showed signs of incubation. The nest was made of a
number of small dry leaves with down. In a hole nearby were many
remains of fish. The parent bird was seen, but it escaped.

The young bird obtained August 29, 1950, was partly feathered,
but its primaries were still only an inch long. It seems late in the
season for so small a bird to be able to complete its growth, for the
lakes would be frozen over in a few weeks. Among the eggs found
on June 24, 1949, only one showed signs of incubation. Among three

46 U. S. NATIONAL MUSEUM BULLETIN 217

females examined on May 380, 1951, the largest undeveloped eggs
recorded were respectively 2,4, and 27 mm. in length. It appears that
eggs are laid during the first half of June.

In Nunamiut the red-breasted merganser is called Akpaksruayook,
which means “runs (like a man) on top of the water.” Because of
their utilization of even small lakes and the river, a wide area of
suitable habitat is open to them and the population is rather large,
probably, because of their wide distribution, exceeding the number of
white-winged scoters.

Family ACCIPITRIDAE: Hawks, Harriers

Accipiter gentilis atricapillus (Wilson)

1 male November 1952, weight 1280 g. _ —
Ikiakpuk Creek
1 female. Mar. 8, 1950, John weight 1290 ¢. — —_

River

The first specimen of goshawk was taken by John Morry in the
spruce timber along the John River not far from Hunt Fork. It is
known to the Nunamiut as Hidigavitch kiringit, and the old folks
were familiar with the regular nesting of a pair in that area. Gos-
hawks are not conspicuous in the forest and they were considered un-
common, or at least they were infrequently seen. I did not hear of
their being found on the tundra until Mory Maptigak brought me a
specimen he had shot along Ikiakpuk Creek, some 30 miles north of
tree line on the southeastern approach to the low divide between
Tkiakpuk Valley and the watershed of Chandler Lake.

In the period 1954-1956, goshawks were more frequently seen than
in earlier years near the limit of timber and a few were seen several
miles farther north. Iam inclined to think that the increased observa-
tions of goshawks resulted from their following the great numbers
of willow ptarmigan which were migrating during those years.

I still regard the goshawk as a bird of the forest which, as in the
case of our winter specimen, may venture a few miles over the tundra.
It is therefore designated a visitor.

Buteo lagopus (Pontoppidan )

No specimens of rough-legged hawks have been collected in the
Endicott Mountains, so that the race cannot be designated, although it
probably is B. l. s.johannis, which Tom Cade informs me he regards as
the form which he found along the Colville River. They are well
known under the name /lyirgik, which means “basket sled,” a designa-
tion which might refer to their feather clad legs. In 1949 the first
record was on May 5, in 1950 on May 19, in 1951 on May 3, in 1952

ANAKTUVUK PASS 47

on May 9, and in 1953 on May 3. In 1949, early observations were
made more continuously than in other years by Thomas Brower, who
reported one or two daily early in May and six on May 138. All these
birds were flying northward in evident migration. One nest was seen
by Simon Paneak in 1938 at Okpikyuak River near the mountain line
in Tuluak Valley and another just east of the Killik Valley, but they
are known to nest commonly on the cliffs which extend from Umiat
northeast along the Colville River and this site is presumed to be the
destination of these migrants through the Pass. A few are seen during
the southward migration, but they are then never as conspicuous as
when northward bound. They are probably rare as nesting summer
residents in the mountains but regular as migrants in the Pass.

Aquilae chrysaéios canadensis (Linnaeus)

No American golden eagles were brought into the collection, but
several were examined for parasites by Robert Rausch, and numerous
others have been brought in for my examination. The earliest
sight records are April 7-10, 1948, when four were caught in wolf
traps; March 29, 1949, April 3, 1950, April 10, 1951, March 28,
1952, April 4, 1953, and March 23, 1954. During April, May, and
June, more eagles were seen than during the summer, the largest
number reported by Thomas Brower being 14 on June 20, 1949. Dur-
ing the summer one or two eagles are usually in sight soaring high over
the mountains or along the precipitous slopes of the upper sides of
the Valley. In spite of their impressive size and capable flight, they
are without expression, not social, and, in comparison with the ravens,
uninteresting. In their individual and varied social activities, ravens
are always expressive of purposive play or shrewd hunting.

In Nunamiut, the eagle is called Tikmiakpuk, meaning “largest
bird.” They nest in the mountains and occur in larger numbers dur-
ing the fall migration. The latest record was September 14, 1950,
at Contact Creek.

Many eagles migrate through the Pass in spring, but as there are
few cliffs upon which they could nest north of the mountain line, and
as eagles are seldom seen on the arctic coast, their only known north-
ern destination on the arctic slope is the cliffs along the Colville River
northeast of Umiat.

A single example for Contact Creek weighed 5900 grams.

Haliaeetus leucocephalus alascanus Townsend

Northern bald eagles have been occasionally seen in the mountains,
but only as visitors, giving no appearance of seeking residence. Simon
Paneak recalled seeing several in the Valley in 1948, and had only
rarely seen one along the eastern Alaskan arctic coast. I saw one on

48 U. S. NATIONAL MUSEUM BULLETIN 2:17

June 1, 1952, over Inukpasugaruk Creek. The Nunamiut name was
not recalled by the people, but they described the bald eagle accurately
and recognized it. Since these eagles are regular and common resi-
dents along the rivers in the wooded country to the south, their occa-
sional visits are not unexpected.

Circus cyaneus hudsonius (Linnaeus)

lunsexed butin Aug. 28, 1950 weight 478 g. — ——
female plumage

I saw an American marsh hawk flying as if hunting over the tundra
August 29, 1948. One was reported on May 380, 1951, May 20, 1952,
and May 19, 1953. Simon Paneak saw one several times in early June
1951, around Tuluak Lake. Susie Paneak and May Kakena saw one
“with reddish-brown breast” on June 11, 1952. The Nunamiut know
them as Papiktook, which means “long parka tail,” and Simon Paneak
had described them to me accurately before the specimen was taken
by Jesse Ahgook. No nests have been found and the occasional
individuals seen appeared to have been on a hunting or exploring
trip and not resident in the mountains. On August 9 and 14, 1951,
I saw two marsh hawks while traveling up the Alatna River, the
first about 10 miles and the second about 90 miles north by river from
the Koyukuk and within 60 miles of Anaktuvuk. As their common
range extends through the Yukon Valley, occasional visits to the
tundra can be expected.

Family PANDIONIDAE: Ospreys
Pandion haliaetus carolinensis (Gmelin)

Several of the Nunamiut saw an osprey’s nest before 1947 in a tree
near Island River in the forested midpart of the John River, but I
have not seen one along the upper Koyukuk and Alatna Rivers. On
May 28, 1954, May Kakena saw one at the outlet from Tuluak Lake.
Charles Sheldon pointed out to me a pair of ospreys over a nest in
a high tree which had been occupied for many years on the Kobuk
River about 10 miles above the village. On June 1, 1952, I saw an
osprey flying over Contact Creek. The osprey is known to the Nuna-
miut from their occasional journeys to the forest country as Kallok-
stoyuk, meaning “goes after fish.” They are occasionally seen flying,
but not fishing in the mountains. It is not surprising that occasional
visitors should venture to the Valley since it is only an hour’s flight
from their known nesting range in the wooded valleys. It is inter-
esting that the Nunamiut remarked that the visiting ospreys had not
been seen fishing in the tundra lakes,

ANAKTUVUK PASS . 49

Family FALCONIDAE: Falcons

Falco rusticolus obsoletus Gmelin

1 female, Nov. 17, 1952 weight 1431 g. —= <=
Itikmalikpuk

1 young mele, July 2, 1951 weight 1330 g. == +
Trivik Creek,
Killik Valley

Gyrfalcons have been explicitly reported seen in the mountains
in every month of the year and I can see no indication that the num-
bers vary in a way to suggest migratory habits. As I came to know
some of the well established locations of the nesting pairs it appeared
that the birds observed were usually within some miles of a nesting
place and it was our opinion that a pair remained within its territory
throughout the year.

During the late summer and autumn of 1952, gyrfalcons were occa-
sionally seen around the heights of Soakpuk mountain. Birds seen
a year earlier near the head of Kangomavik Creek were thought to
belong to this family, and individuals were occasionally seen in the
nearby valley during the months of February and March 1953. The
location of the Soakpuk nest in the high cliffs was impossible to
determine.

About the end of March 1952, Simon Paneak frequently observed
2 pair of gyrfalcons near Naniksruk, an open fishing place on the
Anaktuvuk River some 60 miles north of the mountains. He thought
they were then preparing to nest in the place which he knew as a
historic nesting location. The young specimen was one of two full-
sized nestlings shot by William Irving on a nest in a cliff near Irivik
Cave in the Killik Valley. The other bird lodged in the nest, which
was inaccessible, and which had long been a known but unattainable
nesting site. It was remarked that another nest also inaccessible was
located within a hundred yards.

The Nunamiut say that none of the hawks can overtake a sound
ptarmigan or duck in flight at the same level. They also believe that
a gyrfalcon strikes its prey with its breast and only seizes it with
claws when it is struck to the ground. While this is not the common
opinion of naturalists it was expressed by Nunamiut for whom ob-
servation of animals is an inherited career.

There are several Nunamiut names for gyrfalcon: Okiotak, Kit-
gavikroak, and for young birds in autumn, Atkaruak, meaning “like
caribou mittens.” The Nunamiut interest for gyrfalcons arises from
their spectacular habits and their usefulness to a hunting people.
Wing and tail feathers were used to guide arrows and spears, as were
those of the duck hawk, pigeon hawk and rough-leg. For this purpose
the first four primary feathers were preferred. Two feathers were

50 U. S. NATIONAL MUSEUM BULLETIN 2:17

common on each arrow, but sometimes three or even four were used.
Paired feathers were preferably matched from corresponding feathers
of the right and left wings. If tail feathers were used, matching parts
were used by preference in pairs. I did not learn the principle or prac-
tical reason on which this practice was based. I also asked whether left
wing feathers were preferred for convenience of attachment by right-
handed arrow makers, as Bridges (1949) relates to have been the case
among the Ona Indians of Tierra del Fuego. I could find no com-
parable discrimination among the Nunamiut.

Along the low arctic coast hawks are rare and feathers were accord-
ingly one of the uniquely valuable articles of the old Nunamiut in
their trade with the coast. Some of the old Nunamiut people were
specialists in catching hawks with whalebone snares set near a perch
or nest, and the nesting locations were established in old Nunamiut
knowledge. I learned that many nests were known to have remained
in use during the lives of the narrators and during their father’s times,
and so far as it appeared, indefinitely. The present generation had
discovered a few nests which, so far as they knew were unknown to
their fathers, but none were thought to be at new sites. From ac-
counts of several older Nunamiut men has been compiled the follow-
ing list of 17 nesting sites of gyrfalcons which they know in the area
from the Killik on the west to the Anaktuvuk on the east, an area
near 100 miles on each side (no nests are known on the Colville below
the Anaktuvuk’s mouth) :

On Anaktuvuk River: 30 miles from mouth, near Naniksruk (specimen
obtained); 60 miles from mouth; and 90 miles from mouth, Soakpuk
Mountain.

On Colville River: 25 miles above Umiat; and 4 miles above mouth of Killik
River.

On Killik River: 15 miles from mouth; 45 miles from mouth; 20 miles above
Odrivik Lake, near Irivik Cave (two nests, both occupied in 1950, very
close together; specimens obtained); 2244 miles above mouth; and at
Akmalik Creek.

On Okpikyuak River, east of Killik.

On Okomilaga River: 8 miles east of preceding location; 3 or 4 miles inside
mountain line; and at summit.

Between Okomilaga and Chandler River: At mountain line; and 6 miles
north of preceding location. ;

It is thought that nests do not remain unused. If one bird of a
pair is killed a substitute promptly joins the bereaved mate and the
pair continues at the site. I could not press to learn the length of the
delay except that replacement is thought not to be deferred until the
next season but to occur right away. There seems to be a reserve, gen-
erally unobserved, of single birds in this population which otherwise
exists in firmly attached pairs.

ANAKTUVUK PASS 51

Seventeen pairs are known in occupation of about 10,000 square
miles. A gyrfalcon could cross the area in three hours flight and the
element of transportation would be a small obstacle to keeping the
pairs of the population in communication from the positions which the
Nunamiut views ascribe to them. It is interesting to speculate as to
what social bonds relate the few pairs of gyrfalcons in a population
which appears remarkably conservative in its total numbers and in the
location of its component pairs.

Falco peregrinus anatum Bonaparte

1 male, Anaktu- May 1, 1948 —_— — —-
vuk Pass, Sum-
mit

1 young male, June 11, 1951 weight 590 g. — —-
Anaktuvuk
Creek

The male American peregrine from Summit was taken from the
upper John River and north of the last spruce May 1, 1948, sent to
George Sutton and identified by him.

The young male was taken by Jesse Ahgook where Akvalutak
Creek comes out from the mountains into the Valley. While in pursuit
of a companion peregrine, Jesse fell and his shot gun discharged both
barrels so as to nearly sever the muscles of his leg anterior to the
femur. After two days of first-aid treatment in camp, he was taken
out by small plane to Bettles, and by the 10th Rescue Squadron to
Fairbanks, where under skilled treatment he made a recovery which
bids fair to extend his great usefulness as a bold and skillful hunter
and a gentle companion through his 9th decade.

In Nunamiut, the name Aidgavitch Kiriat, meaning “small hawk,”
refers to a medium-sized swift flying and striking hawk known to
nest along the cliffs of the Colville but not often seen in the Valley.
They are occasionally seen as migrants, and judging from Nunamiut
accounts it is uncertain that they nest in the mountains.

Simon Paneak related to me that along the cliffs of the Colville
River he had seen a peregrine dive at a rough-legged hawk. The latter
turned in the air and seized the attacker but was forced to release it
by the onslaught of the peregrine’s mate. Hawks frequently harry
other hawks and even eagies, and the maneuvers are evidently serious-
ly intended, for there seems to be no play in the relations among the
species of predatory birds. In level flight ptarmigan and ducks are
considered able to outstrip hawks, which are therefore dependent upon
surprising their prey at a disadvantage or on the speed of their dives.

52 U. S. NATIONAL MUSEUM BULLETIN 2:17

Falco columbarius bendirei Swann

1 male Aug. 12, 1950 weight 169 g. = —
2 females June 16, 1952 weight 142, 231 g. == ==
Aug. 25, 1950

The male western pigeon hawk had been seen on two days flying
South directly through the willows covering the sand dunes on the
south side of Odrivik Lake. On the last trip, its direction was fol-
lowed for about a half mile nearly to the bank of the Killik River, and
as it returned flying northward through a narrow marshy hollow be-
tween two willow covered dune ridges, it was shot. Its behavior so
well suggested that a nest or perch was in the vicinity that a careful
search was made in the area from which it had last appeared, but no
sign of nest or perch could be found. The conviction that a residence
site was near, led me to further observation of the area from the top
of a dune a little distant. On the next day a dark bird of the same
size left the area and flew swiftly across the Kallik River and dis-
appeared in a small patch of willows. The day was then dull and
identification of this bird could not be established in the poor light
and prolonged observation gave no other information. I think it
likely that the bird collected was a resident and not a migrant. On
two successive days in early June 1952, I saw a pigeon hawk fly
through the willows on Contact Creek. A female collected there on
June 16 had already laid eggs.

A pair was watched by Simon for several hours near the site of
the gyrfalcons’ perch on Soakpuk Mountain. The small hawks were
screaming in agitation at the gyrfalcons and probably at the human
intruders, but after watching for over an hour, he could find no nest
of either hawk. One of the pair of pigeon hawks was collected.

One female specimen was collected at Tuluak Lake on August 25.
That region had been under close observation through the summer,
and as no pigeon hawks had been reported, it is believed that this
example, recorded as having ample subcutaneous fat, was a returning
migrant. Another pigeon hawk was reported seen September 8, 1950,
near Contact Creek, and Simon Paneak and William Irving reported
one seen July 6, 1951, in the Killik Valley among the mountains.

In August 1951, several pigeon hawks were seen along the Koyukuk
and Alatna Rivers. A resident Indian, William Williams, said that
they regularly nested at certain locations along the river. One was
collected at the mouth of the Iniakuk River, at the southern mountain
line and about 70 miles southwest of Tuluak Lake. In mid-March
1951, Ernie Johnson pointed out a small hawk which had offended him
by harrying his tame jays and pursuing small birds about his cabin
near the northern limit of spruce on the Alatna Malemute. John

ANAKTUVUK PASS 53

Krog and I were convinced it was a pigeon hawk but in spite of
Ernie’s encouragement we were unable to eliminate the disturber of
his jays. Dall remarked that pigeon hawks were at Nulato all the
year round (Dall and Bannister, 1869).

The Nunamiut name is Kidgaviatchaurak, which means “smallest
hawk,” a name which might also apply to the sparrow hawk. The
latter is an unusual visitor, while the pigeon hawk is considered well
known among the Nunamiut and nests among the cliffs of the Colville.
It seems certain that pigeon hawks also nest in the mountain valleys.

These three specimens, together with a female collected on the
Alatna River, conform well with the common appearance of speci-
mens marked dendirei in the U. S. National Museum, and not with
those marked columbarius.

Falco sparverius sparverius Linnaeus

I saw a small hawk repeatedly hovering over the camp at Pitaich on
August 25,1948. Elijah Kakena shot the bird, which I identified as
a female American sparrow hawk. The skin was prepared but among
the difficulties of travel in those days it has been lost. The Nunamiut
had not seen a sparrow hawk before, and it is certainly unusual in
the Valley. Since sparrow hawks in Alaska are known to nest as
far north as tree line, this bird can be called a visitor.

Family TETRAONIDAE: Grouse, Ptarmigan

Canachites canadensis osgoodi Bishop

3 males (sex of 2 Sept. 13-Oct. 9 weight (2), 611, 590g. —_ —=
determined
only by
plumage)

1 male (sent to Feb. 23, 1948 noe — —-
G. M. Sutton)

2 females Oct 3, 1950 weight 649, 734 g. — —

Four of these Hudsonian spruce grouse were obtained from the head
of the Savioyuk River and one from the upper John River. All
were from forested country. The male taken February 23, 1948, at
Hunt Fork, was sent to Dr. G. M. Sutton and kindly identified by
him, A female taken May 13 near Anchorage weighed 577 grams
and possessed little fat, while the autumn specimens in the mountains
were fat.

This bird, which does not venture north of the forest, inhabits the
northern tree limits, often so abundantly that it is important as food
for hunting parties. When the Nunamiut sometimes go in winter to
the timber for trapping, they know the spruce grouse as Napaktom
Kadgia, which means grouse of the spruce.

54 U. & NATIONAL MUSEUM BULLETIN 217

Lagopus lagopus alascensis Swarth

3 males Dee. 13, 1948- mean weight (25), — —
May 31, 1952 641 g., std. var.
75
4 females November 1948- mean weight (88), — —
May 30, 1952 602 g., std. var.

78

These Alaska willow ptarmigan inhabit a territory ascribed to
Lagopus lagopus alascensis, and compare well with specimens of that
species in the U.S. National Museum.

The willow ptarmigan are Kadgivik in Nunamiut, which is ex-
plained simply to signify “real ptarmigan.” They are so familiar to
the Eskimo life and economy that for them the name has no known
descriptive meaning. The pebbles which are regularly found in the
gizzards of ptarmigan are called “heaters” by the Nunamiut, using
the designation of the stones which in former times were heated in
the fire outside and then brought in to warm the people in their skin
tents. The “heater” stones which the rounded pebbles in a ptarmigan’s
gizzard resemble were specially selected fire-resistant stones, well
rounded by water. Many stones, even well rounded ones, crack and
even burst dangerously in the fire, but the kind selected for heating
were probably free from strain and thus less likely to burst. ‘There
is no implication that the function of the pebbles in the gizzard is
for heating.

In summer a few willow ptarmigan remain in the mountain valleys
and occasionally nests are found. In July 1951, William Irving and
Simon Paneak saw several adults with young birds in the Kilhk
Valley. On Contact Creek I obtained a female with developing eggs
25 mm. in length on May 30, and several pairs were courting on the
open valley and among the willows.

By the first of October many ptarmigan come from the north into
the valley, and through the autumn their movement is southward,
but I have not found the southbound movement of any birds to be
as evident as their northward migration, so the southward movement
of ptarmigan is ill defined in our records. In January the traffic
appears to be least, and in February a northward movement becomes
apparent and continues through May, but with relaxed intensity
during the middle of the migration.

The Nunamiut say that the ptarmigan migrate in two waves sepa-
rated, around the end of March, by an interval of less frequent appear-
ance. This pause in migration is evidently regular, for the Nuna-
miut sometimes date events as occurring during the first or second
migration of ptarmigan. It seems to be an established schedule in
the habits of ptarmigan but I do not know of any distinguishing form

ANAKTUVUK PASS 55

or condition of the birds. I suggested that they might come from dif-
ferent areas, but Eskimos are not much interested in speculation upon
determinable but unknown facts. By the end of May the last mi-
grating ptarmigan disappear and comparatively few remain to nest
in the mountains.

In February and March 1952, willow ptarmigan were common in
scattered flocks of 20 to 40. In the winter of 1952-1953 they were
even more common, and this great abundance continued through the
winters of 1954 and 1955. In 1957 they were still very numerous.
Few of the willows lacked signs of removal of buds and twig tips.
In the dark twilight before the brief day the ptarmigan could be
heard calling. Before noon, feeding stopped and the birds rested on
the snow or perched in the willows for one or two hours before re-
suming the feeding which filled their crops for night. Crops weighed
in the morning and at noon were lighter than in the afternoon and,
according to numerous weights obtained by Simon Paneak and John
Krog, willow ptarmigan retire in winter at 4 or 5 o’clock with well
filled crops weighing from 50 to 100 grams.

This store suffices for the 14 hours or so which they spend in indi-
vidual burrows in the snow. The accumulated feces show that each
bird remains in one spot. These burrows are a foot or so beneath
the snow and extend from 18 inches to 2 feet. In the morning the
ptarmigan leave their snow burrows usually by flying directly up
through the soft snow. There is no indication that any weather causes
them to remain longer than one night in their burrows. Reports of
willow ptarmigan burrows in the snow come from other parts of
Alaska, Yukon Territory and Newfoundland (Wetmore, 1945).

Although at times the tracks, noise, and sight of ptarmigan are
reminiscent of the density of population of a farm yard, and although
the willows showed signs of their intensive feeding, the ptarmigan
were in good condition in the time of their greatest abundance. Occa-
sionally, when the frost thickly covered the willows the ptarmigan
sought sedges exposed by the wind on which to feed, but in winter
willows alone afford probably 90 percent of their food. And yet as
far as can be seen neither ptarmigan nor the restricted willows of the
tundra suffer from the intensity of cropping which occurs when the
ptarmigan are most numerous.

Ptarmigan meat never has enough fat to make it a satisfactory
arctic food. Nevertheless, despite the arctic cold, and the competition
on their crowded range from successions of migrating flocks, the
ptarmigan in winter have flesh that is better eating and the birds are
heavier than in summer, when they have access to a flourishing
vegetation.

469496605

56 U. 8. NATIONAL MUSEUM BULLETIN 217

TaBLe 1.— Weights of willow ptarmigan from several parts of Alaska

Males Females
Date Weight (g.) Weight (g.)
No. No. 2
Extremes | Averages Extremes | Averages
Anaktuvuk Pass
November—December 1948 4 504-627 600 2 500-625 563
April 1949 1 706 4 504-627. 560
February-March 1950 Gays 461-804 601 7 430-704 544
March 1952 13 580-743 633 20 527-706 617
25 (621) 33 (591)
Arctic Village
December 1951—February 1952 14 506-730 615 15 495-749 64)
ARCTIC 39 (619) 48 (607)
Chugach Mountains
December 1951—February 1952 12 450-617 530 8 446-525 495
Talkeetna Mountains
December 1950, January 1951, and Feb- ;
ruary 1951 3 473-519 539 5 493-627 573
SUBARCTIC ; 15 (535) 13 (525)

In the winters of 1948, 1949, 1950, and 1952 the mean of the weights
of 128 arctic ptarmigan was 619 grams. Among willow ptarmigan
taken in winter in the Chugach and Talkeetna Mountains about 600
miles south from Anaktuvuk the mean weights of 28 birds were 531
erams. In Mount McKinley National Park, Alaska, Dixon (1927)
reported the average weight of male willow ptarmigan (of this race)
in the breeding season to be 507 grams. My records (11 males from
Anaktuvuk during April and May weighed 538 grams.) indicate that
arctic willow ptarmigan are lighter in summer than in winter. While
he was on the eastern arctic coast of Alaska in winter Anderson (1921)
reported that 15 willow ptarmigan weighed 22 pounds which gives an
average weight of 652 grams.

The distribution of weight about average in arctic willow ptarmigan
is shown in the following tabulation:

Percentage distribution about average

weight
Number Sex +10% +20% +30%
39 Male 54: 96 100
49 Female 52 89 100

The difference between the winter ptarmigan of arctic and subarctic
Alaska points to the arctic ptarmigan being significantly heavier than
those from the subarctic locality. On the basis of weight differences
of similar proportions (about 1 kg. in 5 kg.) von Zedlitz (1924)

ANAKTUVUK PASS 57

thought that geographical races of Zetrao urogallus living in lands
about the Baltic Sea were distinguishable, although not recognizable
by superficial taxonomic distinctions. Near the northern limits of
spruce in the interior of Keewatin 6 adult males of Lagopus lagopus
albus taken between May 31 and November 7 averaged 631 grams in
weight (Harper, 1953).

Considering the variations in weight of domesticated animals which
can be produced by feeding one would conclude that differences in the
natural nourishment provided in parts of the range of a species could
easily account for the differences in weight of grouse found in several
distinct localities. Among the wild birds of Anaktuvuk, however, I
find that the differences in weight among individuals of specific popu-
lations are small and they are evidently homogeneous in respect. to
weight. I am inclined to favor von Zedlitz’s suggestion that genetic
differences are likely to be at the basis of the distinction of weight
in certain parts of the range of grouse. Nasimovitch (1936) reported
an increase in the weight of Lagopus lagopus progressing eastward
into that part of their range lying in the central regions of western
Siberia.

In the years under observation (1947-1957) the abundance of willow
ptarmigan in winter at Anaktuvuk has varied from their being com-
mon to very common. I have not found evidence for regular cycles of
abundance. The difference in actual numbers seemed to be as much as
ten fold, and I suspect that some orderly conjunction of events under-
lies the phenomenon, As no annual differences have been observed in
their weight and condition of flesh the good nutritional condition of
willow ptarmigan in winter is a characteristic of the season and locality
in spite of the fluctuations in numbers which at times appeared to
crowd the country with ptarmigan. It also seems that in the sub-
arctic locality a distinction in weight occurs, although we have seen
no superficial character to distinguish them from birds of other
localities.

Lagopus mutus nelsoni Stejneger

9 males May 8-July 14 weight (20), 411-559, — —
average 466 g.
8 females May 8-June 5 weight (7), 416-480, — —-
average 427 g.
1marked female, July 31 weight 412 g. — oo
but in male
plumage
2 downy males July 3 weight 15.2, 16.3 g. — ——

These Nelson’s rock ptarmigan correspond in beak size and in the
distinctive summer pattern of males to those marked nelsoni in the
U.S. National Museum.

As Niksaktongik, rock ptarmigan are nearly as well known, and
form a more interesting part of Nunamiut life than are the larger wil-

58 U. S. NATIONAL MUSEUM BULLETIN 217

low ptarmigan. In an old Nunamiut story Willow Ptarmigan chal-
lenged Rock Ptarmigan. The latter said that although smaller he was
more agile and had better staying power. In the ensuing combat he
demonstrated his superiority and finally killed Willow Ptarmigan.
Then his wife tattooed the black marks with soot on either side of his
mouth as the sign used among Eskimos of one who has killed a man,
or, along the coast, of one to be recognized for killing a whale. Rock
ptarmigan appear bolder and more energetic than willow ptarmigan.
In the rock ptarmigan the mothers seem to be more attentive in the
care of the young chicks and male birds often share with them in the
early care. The disposition for individual independence and effective
care of family make the rock ptarmigan also seem like substantial
mountain residents in the opinion of the Eskimos.

Rock ptarmigan do not appear to migrate as do willow ptarmigan
but they do change their common feeding range during the year.
They are familiarly seen in summer on high ground or occasionally
on the gravel bars among the willows in the creeks, while in winter
they often keep to higher rounded slopes where the low vegetation is
exposed by the strong winds. Not as numerous as willow ptarmigan
during their period of maximum number, they have a more restricted
habitat which appears less productive of food than the willows, with
their good supply of buds, on which the latter feed.

The largest eggs developing in females were measured in 1951 on
May 17 at 14 mm.; May 18,2 mm.; May 20, 12 mm.; May 21, 18 mm.;
May 23, 45 mm.; and in 1952 on May 29 at 14 mm.; June 3, 44 mm.;
June 5, large. The last was the size of eggs in nests. The two downy
young taken on July 3 were, judging from their size, recently hatched.
In country along the Itivlik similar to Anaktuvuk chicks were just
able torun on June 25.

In early May the red about the eye of the male expands and be-
comes brilliant, and by mid-May a few dark feathers appear on the
neck. On May 27, 1952, in Howard Pass the white males were con-
spicuously posturing, fighting, and occasionally pursuing the sly
females, which, their backs and heads changed to summer plumage,
remained well concealed on the alternating grassy and snow covered
spots. A week later at Anaktuvuk the courtship appeared to have
been accomplished. Through early June the male birds, still white
except for a few dark feathers showing on their necks, stood in
statuesque posture on the knolls where they often permitted a close
approach before flying.

So seriously do male ptarmigan take exhibition as their function
while the female is sitting on the nest that they stand for hours in
full view, often with empty crops. During this period, their preoc-
cupation with exhibitionism hinders feeding and makes them lean.

ANAKTUVUK PASS 59

The females feed busily and frequently while courting and nesting.
By June 5, the females in their beautifully patterned summer plumage
are difficult to detect.

During the whole period of mating and nesting, the females are
as secret and retiring as the males are conspicuous. The secrecy of
the females conceals them and their nests. The display of the males
looks provocative to predators and it would be interesting to think
that the males in this way distracted predation from the brooding
females. As a predator myself I have often been attracted to the
vicinity of nests by the displays made by the nonbrooding parent, and
I conclude that the occasional danger of predation must be less than
the constant need for advertising the occupation of a territory by a
family of nesting birds. For successful breeding each family must be
kept as a distinct unit within the society of rock ptarmigan. Thus
both the isolation of families and their cohesion in an orderly society
is effected by the display of the male. The inclination for display
dominates the behavior of breeding male birds to degrees varying
among the species. For the rock ptarmigan this concern for its family
and society seems to suppress the attention for individual protection
which is apparent in its behavior at other seasons.

Pedioecetes phasianellus (Linnaeus)

In addition to spruce grouse another timber grouse is called
Odgillyim kadgia, “birch grouse,” by the Nunamiut. At Bettles a
sharp-tailed grouse was clearly described by David Tobuk, an Eskimo
long resident there, who said that he had not seen one in recent years.
Simon Paneak recalled taking one at Hunt Fork about 1939. Older
Eskimos at Kobuk related to me the occurrence during one period in
that region of “birch grouse.” I have not yet been able to develop a
chronology which would enable me to estimate the years when they
were well known, but I suspect that sharp-tailed grouse have been
rather common near the northern limit of timber during a period
which ended some 20 years ago.

Cade and Buckley (1953) relate various accounts of sharp-tailed
grouse which indicate that in 1934 the population became dense in the
Tanana Valley. On one occasion in that year a large flock was seen
to fly away. This movement seemed to mark their disappearance from
that part of the Tanana Valley for thereafter there were no reports
of their occurrence near Fairbanks. Since that date these grouse have
not been widely common throughout interior Alaska. It seems pos-
sible that the reports of sharp-tailed grouse which I obtained in arctic
Alaska may be related to a date when they were abundant in much of
the subarctic interior.

60 U. S. NATIONAL MUSEUM BULLETIN 217

Family GRUIDAE: Cranes

Grus canadensis canadensis (Linnaeus)

Lesser sandhill cranes are seen and heard regularly during spring
migration. They were reported May 30, 1949, and May 25, 1954. As
Tattidgak, their characteristic mode of flight and calls are well known
to the Nunamiut. It is said that flocks greater than six in number are
not seen, and that they do not alight in the Valley. A few are known
to nest along the lower Colville.

Family CHARADRIIDAE: Plovers, Turnstones, Surfbirds

Charadrius semipalmatus Bonaparte

16 males May 19-June 22 weight (11), 39-48, — —
. average 42 g.
2 females June 22-Aug. 6 weight (8), 41, 40, 39 — —
g.
1 young male Aug. 16 — = ===

2nests with3and June 22, 1949 == == —
4 fresh eggs; one
male, one female
parent.

The attractive semipalmated plover was recorded in Tuluak Valley
May 20, 1949, June 2, 1950, May 6, 1951, May 23, 1953, and June 1,
1954, One which had been collected at Tuluak June 3, 1948, was
examined but could not be saved. A considerable number in migrant
flight pass through late in May, and many settle to disperse along the
sand and gravel bars of the river, where they may often be seen
running with surprising swiftness for such small birds. The first set
of eggs was found by Thomas Brower; it contained three eggs on
June 12, 1949, and since no addition was made, it was collected partly
incubated June 22. The other set listed was fresh.

The bird in young plumage was collected at Akmalik Creek, Kallik
Valley, on August 6. It could then fly well and was near full length
and wing size. It was with an adult female, and both appeared to be
in residence. The last record for 1950 was September 1 at Contact
Creek. .

One of the nests collected by Tom Brower was found on the fine sand
near a little stream and was made of coarse and finer grasses with a
few leaves and fragments of willow and other dry tundra material.
The other was found on a ridge above the river and contained moss,
fragments of grass, and small leaves from the adjacent dry tundra.

These handsome and alert little plover are well liked. The
Nunamiut name A odrakoruk is given them in resemblance of their call.

There is a considerable northbound flight of semipalmated plover.
Those which remain in the Valley are conspicuous by their calls and

ANAKTUVUK PASS 61

for the swiftness with which they run over the gravel bars of the river,
but because of their restricted habitat, the resident population in the
valley is not large.

Charadrius vociferus vociferus Linnaeus

On the basis of recognition of pictures and from description of its
appearance and habits, the killdeer was called Z’alikvak, by the Nuna-
miut. Simon Paneak says that a few may be seen as spring migrants,
and reported one sight record May 29, 1950, at Tuluak Lake. On
May 29, 1951, one was wounded by Elijah Kakena but it escaped. The
Nunamiut think that they do not nest in the mountains, but that they
nest on the lower Colville. These Nunamiut accounts undoubtedly
refer to a plover or sandpiper differing from any which we obtained
and resembling the illustrations and descriptions of killdeer. But
there have been only two specimens of killdeer taken on the arctic
coast of Alaska (Bailey 1948) and there is only one sight record from
Yukon Territory (Rand 1946). Considering it possible that I could
have been confused in our exchange of descriptions, I cannot now
identify conclusively the killdeer in the list of birds of the mountains.

Pluvialis dominica dominica (Miiller)

20 males May 19-July 25 weight (36), 127-169, — —
average 144 g.

18 females May 21-July 25 weight (24), 126-169, —— ——
average 146 zg.

1 young male July 25 weight 105 g. — —_—

2young females July 25 weight 111, 121 ¢g. — —_

5nests, each with June 9 od FS

female and 4

fresh or slightly
_ incubated eggs.

In addition to the specimens of eastern American golden plover re-
corded, seven birds were collected May 12-23, 1948, and about 100
others were examined. None were questioned as deviating from the
form dominica. The earliest records are May 12, 1948, May 16, 1949,
May 10, 1950, May 8, 1951, May 19, 1952, May 18, 1953, and May 17,
1954. Thereafter increasing numbers were seen daily feeding and
whistling on the tundra, where the drier and more elevated part of
the valley floor was their preferred habitat. Because the flocks were
few and loosely formed, it is difficult to distinguish northbound mi-
grants from the many birds which take residence and impossible to
estimate the proportions of the two categories.

In early June pairs are established in their territory, where both
often stand, conspicuous and statuesque, as the male whistles the cheer-
ful call for which it is called 7’odlék by the Nunamiut. The first
nests were recorded on June 9, 1949, two sets then being slightly in-
cubated, one on June 8, 1951 and a nest with three eggs on June 10,

62 U. S. NATIONAL MUSEUM BULLETIN 217

1952. Some nesting occurs earlier, for eggs were used for food when
a snowstorm on June 3, 1948, drove many birds from their nests. Fe-
males taken May 28, May 30, and June 1 contained eggs ready to be
laid.

Nests were located on dry ridges near the river and over the valley
in dry spots as far as the slopes leading to the steep valley walls.
They were simply composed of fragments of lichen, a little grass, and
a few small leaves. With the five nests female birds were taken in
each case, but the males remained nearby.

The first young were recorded June 25. On July 8, 1951, a young
bird taken at Amorgoayat in the Killik Valley weighed 77 grams.
Thereafter, the young birds were inconspicuous until about July 20,
when they began to reveal themselves as they ran over the tundra un-
der watchful supervision of the parents. On July 25, 1950, of four
young in one group, one escaped by flying and the largest of the three
taken could fly a little. On July 24, 1950, seven young flew so well
that they could not be run down and were taken with shot. They
weighed from 121-135 grams, average 127 grams. Plover were nu-
merous and usually still in pairs until August 2, but none were seen
August 3-15, in the Killik Valley and none have been recorded later
than August 15, 1950,in Tuluak Valley. About three months of arctic
residence appears to satisfy them for their long flight from southeast-
ern South America.

With their extensive range in the Valley, golden plover are ex-
tremely numerous and they probably outnumber the common Baird’s
and semipalmated sandpipers and northern phalaropes, all of which
have a more restricted range than plover. Their numbers cannot com-
pare with those of redpolls, tree sparrows, or Alaska longspurs. ‘These
observations will indicate the large size of the population of plover
in the Valley, which is representative of great areas which I have seen
on the arctic slope.

Their large numbers and calm, unhurried attitudes demonstrate that
the satisfaction of their food requirements in the arctic environment
is easily accomplished. The ability to obtain basic sustenance with-
out strain suggests how these beautiful birds support their tremendous
migratory flight. The charm of their appearance matches the splen-
did vigor of these birds, which can travel so far yet have the time and
inclination to be lively but unexcited and alert but confiding, and
which always present to an observer the appearance of being satisfied
with and interested in their arctic environment. .

Squatarola squaetarola (Linnaeus)

4 males May 31-June 3 weight (6), 206-234, — —
average 214 g. :
1 female May 27, 1951 weight 192 ¢. == ===

ANAKTUVUK PASS 63

Black-bellied plover were earliest recorded on June 3, 1948, May
14, 1949, May 30, 1950, May 29, 1952, May 23, 1953 and June 1, 1954.
They are called Z'odlivak by the Nunamiut and being conspicuous
by habit and appearance their presence is well known. A few are
seen each spring but they do not remain in summer, and they are not
seen in the fall.

A light-colored female specimen was one of four birds reported
by John Krog to be similarly light in plumage, and it is evidently
a year-old bird. It contained eggs 12, 6, 4, and 2 mm. in length and
was very fat. As the largest egg was about 14 of full length, the
bird was ready to lay soon. The Nunamiut know their nesting places
along the arctic coast.

In 1952, black-bellied plover were seen more commonly than in
other years. Eskimos on the arctic coast have told me that the num-
bers of these plover seen varied greatly from year to year. They
qualified their comments by remarking that their own chances to
observe birds varied with the requirements of their families, the
nature of their hunting, and the localization of their interests. They
believed, nevertheless, that the numbers of black-bellied plover
varied.

Where I have seen these plover, which is not on their nesting
grounds, they have been shy and mute compared with golden plover.
When watched through binoculars, their alert and confident posture
is most impressive, and for their statuesque poise I admire them beyond
even golden plover.

Arenaria interpres interpres (Linnaeus)

1 male May 30, 1949 weight 105 g. — —_—
3 females May 24, 31 weight 90, 97 g. === —

The earliest records for turnstones are June 1, 1948, May 18, 1949,
May 30, 1951 and May 27, 1954. ‘Two shot June 1, 1948, were examined
but could not be saved. The female collected May 30, 1951, contained
three eggs about 4 mm. in length. Well known as Talkvikeak in
Nunamiut, they usually pass hastily through Anaktuvuk Valley in
spring, but none are known to nest there and they have not been seen
later in summer.

T have followed Herbert Friedmann’s identification of these speci-
mens. Bailey (1948) named Alaskan specimens of turnstones
interpres but remarked upon the difficulty of distinguishing them
from A. 2. morinella. The nesting areas assigned to interpres and
morinella overlap east of Alaska and it would seem as if either the
identifications or the taxonomic separation of the two as races is open
to question.

64 U. S. NATIONAL MUSEUM BULLETIN 217

Family SCOLOPACIDAE: Woodcock, Snipe, Sanderlings

The list of shorebirds is extensive for an inland mountain region,
and it is evident that there is much migratory traffic in addition to
some nesting. I have no evidence to suggest that additional forms
may be found, although I have frequently been asked by ornitholo-
gists for information about bristle-thighed curlew in the mountains.
There appears little doubt as to the birds which are nesting or
transient.

The weights of ten of the shorebirds shown in table 2 can be used
for comparison of the size of the two sexes. In two species the average
weight of males is heavier by more than 10 percent, in three the
females are heavier, and there is no significant difference in the other
five (inclusion of specimens carrying a temporary excess of weight
because of being in laying condition can, of course, account for a
higher average weight). I have heard it said that female sandpipers
are larger than males, but that generalization does not apply to the
dimension of weight. —

TABLE 2.—Average weight of the two sexes of Scolopacidae of ten species

Male Female
Name
Number | Weight | Number | Weight
(g.) (g.)

Capella gallinago delicata 6 100 3 102
Numenius phaeopus hudsoni-

cus 2 366 4 456
Heteroscelus incanum 13 101 16 116
Totanus flavipes 14 81 6 81
Erolia melanotos 25 86 10 60
Erolia bairdii 28 39 12 39
Erolia minutilla 16 20 14 22
Limnodromus scolopaceus 28 100 ll 109
Ereunetes pusillus 32 24 10 26
Tryngites subruficollis 4 71 6 53

Capella gallinago delicata (Ord)

4 males May 21-June 20 weight (6), 95-109, — —
average 100 g.
1 female May 22, June 2 weight 92, 113, 105 — —_—

g.

The earliest recorded Wilson’s snipe were seen at Tuluak Lake May
15, 1950, May 16, 1951, heard at Summit May 19, 1952, and seen on May
14, 1953, and May 25, 1954. Not many are seen in the spring, but
the flight sound of snipe is heard almost every day in the Killik and
Anaktuvuk Valleys until about mid-July. Between June 19 and
June 24, 1951, I saw two Wilson snipe at Contact Creek and one at

ANAKTUVUK PASS 65

Kangomavik. Simon Paneak saw one at Contact Creek and John
Morry, three. In late May and early June 1952, I saw numerous
snipe, mostly in pairs, along Contact Creek.

The Nunamiut know them well as Avzkiak, which means “like the
walrus, Aivik,” referring to the similarity of the far-carrying blowing
sound made by walrus and the winnowing of snipe. According to an
old story, killing a snipe would bring on bad weather, and the snipe
is often called “weather maker.”

Occasionally young birds have been seen by the Nunamiut in sum-
mer, and although nests have not been found, I believe that they nest
in the Valley. None have been seen at the end of summer. The
numbers seen in spring and the frequency with which snipe are heard
in summer suggests that they may approach the yellowlegs in number,
and that the retiring disposition of nesting snipe contrasts with that
of the demonstrative yellowlegs sufficiently to give a false impression
of their relative abundance.

Numenius phaeopus hudsonicus Latham

2 males June 1, 1949, 1950 weight 382, 350 g. a oe
3 females May 23, 26 weight (4), 407-493, —— —
average 456 g.

The earliest records of whimbrels are May 20, 1949, May. 23, 1950,
May 19, 1951, May 26, 1952, May 14, 1953, and May 31, 1954. They
appear briefly in spring, often in pairs or in groups of 6 to 10, but
they act like transients and give no appearance of lingering in the
Valley.

Two females taken on May 23 and 25, 1951, contained eggs re-
spectively 10 and 12 mm. in length. As this is about one-fifth of the
length of the eggs when laid (Bent, 1929, p. 117), the birds might
easily with their swift flight reach the arctic coast before laying.

The Nunamiut call them Sgoktovak, meaning “long-billed,”’ and
they have not seen them in the mountains in summer. They know of
nesting places near the mouth of the Colville River. One was reported
seen at Tuluak on August 19, 1950, but fall appearances are rare. One
shot on May 29, 1952, contained many crowberries in its crop.

Bartramia longicauda (Bechstein)
1 female May 24, 1949 — es eee

The single example of an upland plover is in excellent condition
of plumage although it was beyond the range usually ascribed to it.
The Nunamiut designated it a bird which they had not previously
noticed.

On June 1, 1952, Simon Paneak, Elijah Kakena, Frank Rulland,
and Old Hugo saw three sandpipers which they thought resembled

66 U. S. NATIONAL MUSEUM BULLETIN 217

the upland plover taken in 1949 and which differed from any other
sandpiper they knew. A week later Paneak and I saw two birds
which by appearance we could not refer to any other kind of sand-
piper than upland plover. Susie Paneak thought she saw one at
Pitaich on June 18, 1954.

There are a few Alaskan records of upland plover. Grinnell (1900,
p- 75) did not see one on the Kobuk but mentioned Townsend’s (1887,
p. 12) specimen from there. It has been reported in west-central
Yukon (Rand, 1946) and nesting settlements have been reported in
central western Yukon (Buss, 1951). It has also been reported nest-
ing in Mount McKinley National Park (Dixon, 1938, p. 77). Charles
Sheldon of Kobuk recognized the illustration and description of this
bird and gave me as its Kobukmiut name Vanum kanockdoroagna,
“land longlegger.” I suspect that its occurrence in the mountains
is not unique and that it is an occasional visitor from not distant ranges
in the ornithologically little known interior of Alaska.

Actitis macularia (Linnaeus)

3 males, not pre- 31 May, 14 June weight 30.2, 34.1, — —
served. 35.2 £.
2 females 28 May, 6 June weight 37.7, 50.9 g. — —

IT have frequently seen spotted sandpipers along the Koyukuk River
from Bettles to Allakaket and on the lower Alatna River. The Nuna-
miut recognized a specimen which I had brought from near Bettles,
named it Oklaktak, and said it was familiar to them as far north as
the arctic coast. In 1952 I obtained a specimen on Contact Creek near
an old caribou skin to which it may have been attracted by the nu-
merous blow fly larvae. The bird was fat and its eggs, measuring 3
mm. in length, had started development. Another individual was re-
ported seen soon afterward. In 1954 a second female was obtained
by Simon Paneak and 3 males were weighed. The heaviness of the
female indicates it was in laying condition.

In view of the Nunamiut knowledge of these birds, and with several
specimens and records, I consider these spotted sandpipers to have
been migrating birds northbound for nesting. They are reported by
Kessel and Cade (1958) to be rather common in the Colville Valley.

Tringa solitaria cinnamomea (Brewster)

2 males May 17, May 20 weights 55, 54 g. == ar a

These western solitary sandpipers are patterned like a specimen
in the U. S. National Museum collected by Dall at Nulato (Dall and
Bannister, 1869). The Nunamiut know the bird as Xipilugoksiyook,
which means “looks for insects.” They describe it as a lonely sand-
piper found near lakes and in wet places, and seen occasionally in

ANAKTUVUK PASS 67

the mountains, and they recalled it as of regular occurrence near the
arctic coast. Sidney B. Peyton informed me that in June 1954 he
collected a set of eggs from the common nesting place of this sand-
piper. I believe that solitary sandpipers are rather rare migrants
through Anaktuvuk and not stray birds.

Heteroscelus incanum (Gmelin)

12 males May 28-July 9 weight (13), 87-114, os ——
average 101 g.

12 females May 20-July 9 weight (16), 98-130, —_— —
average 116 g.

1 young female July 31, 1950 weight 94 g. ee ==

2 young downy July 9, 1951 weight 43, 44 g. — —

males
1 young downy July 9, 1951 weight 45 g. — —

female

First records of wandering tattlers are May 25, 1949, May 19, 1950,
May 26, 1951, May 29, 1952, May 22, 1953, and May 18, 1954. At this
season they became common among the boulders on the gravel bars
of the rivers and on the gravel fans where the swiftly running streams
come down from the mountains. They are then familiar and friendly.
Their calls, the Nunamiut describe in their name, Stlyirisoktok, “it
sounds like sharpening with a stone.” Captured tattlers emitted this
call in their cages, or, since they soon became adjusted to human
company, as they ran about among the people in the tents and picked
bits of food from the floor or from dishes. In their ready psychological
adjustment to captivity tattlers are like lesser yellowlegs.

Soon after arrival the tattlers become very inconspicuous. Some
doubtless fly north, but an occasional one or pair may be found among
the small boulders of the rocky fans, often closely covered by willows,
where the small streams rush down to the valley from the mountain
sides. After the event, I was informed that my son Laurence and I
ate tattler’s eggs along with those of golden plover, on June 2, 1948,
when many of the birds had abandoned their nests after a snow fall
in early June. The tattler’s nest had been found by Susie Paneak in
the gravel of Nakagnik stream near Tuluak Lake, and Simon Paneak
remarked that the nesting place was at lower elevation than usual.

Nesting is known to the Nunamiut to occur in the boulder-strewn
stream beds, and I have evidence to confirm their story. A male bird
taken June 21, 1949, had bare incubating spots on its breast. It was
the smallest adult male recorded. The young female bird, taken at
Kangomavik Creek July 31, 1950, weighing 94 grams, was sufficiently
young to bea local fledgling.

In 1951, we examined eggs in the tattlers collected. The largest
measured 6 mm. on May 26; 6 mm. on May 28; 2 mm. on May 28; 7
mm. on June 1. These figures do not set the exact date of nesting.

68 U. S. NATIONAL MUSEUM BULLETIN 217

We particularly searched for nests. Susie Paneak reported that on
June 10 she had seen a tattler that acted as if nesting in Akvalutak
Creek near its emergence from the mountain wall, but repeated search-
ing failed to discover a nest. On June 24, a female tattler was col-
lected on Kangomavik Creek about halfway from the river to the
mountain wall of the Valley. Its behavior strongly suggested the
proximity of a nest or young, but Simon Paneak and I were unable
to find anything after long search in the grass-grown gravel. On
July 9, Simon Paneak and William Irving collected a pair of tattlers
with three downy young on Akmalik Creek about 114 miles west of
the Killik River. They were located on a grassy bar among the wil-
lows. Since these young tattlers were probably near twice the weight
of the eggs from which they had hatched they had probably left the
nest several days earlier. Simon and William had, in July, seen an-
other pair of tattlers acting as if near a nest at Togoyuk Creek several
miles higher in the Killik Valley.

In 1956 Simon Paneak’s son Roosevelt found four fresh tattler’s
eggs ona gravel bar in Tuluak Creek. These were collected for Sidney
Peyton.

We have no record of the late summer or return flight, for the
birds remain inconspicuous all summer in the valley stream beds which
they frequent and which are lined and filled with dense willow brush.
During migration, tattlers are much less numerous than golden plover
but more common. than black-bellied plover. Some probably go on
northward, although their summer habitat in the valley would only
be found in a few places north of the mountain line. Among the
Nunamiut, tattlers are considered to be a rather common nesting
bird, but their concealing habitat and shyness in this period is the
reason for uncommon summer observations.

It is interesting to compare the tattlers, yellowlegs, and snipe. The
last are always of secretive disposition on the ground, particularly dur-
ing nesting, revealing themselves in summer only by their flight sounds
and in occasional flights. Yellowlegs show off their presence con-
spicuously all during the summer, when wandering tattlers mostly
disappear up the small creeks. Although only yellowlegs are com-
monly seen, I suspect that all three birds are present in considerable
and perhaps similar numbers nesting in the mountain valleys.

Totanus flavipes (Gmelin)

10 males May 26-July 3 weight (14), 69-94, — ——
average 81 g.
4 females June 13-July 3 weight (6), 77-92, == =

average 81 ¢g.
6, sex unknown May-Aug. 6 — === ——
1 nest, 4 slightly June 13, 1949 = => awe
incubated eggs,
with female

ANAKTUVUK PASS 69

The earliest records of lesser yellowlegs are May 15, 1949, May 16,
1950, May 18, 1951, May 16, 1952 and May 23, 1953. Thereafter they
are conspicuous summer residents. No large numbers were noted in
spring. They proceed rapidly to nesting in the wet marshy areas near
willows and the nest collected by Tom Brower with the female bird
contained 4 eggs already developed by several days incubation. This
nest was on a dry ridge under short willows near the river and made
of short fragments of grass and small leaves. It contained numerous
pieces of the equisetum which grows abundantly in the marshy places
which they frequent. This nest was visible from 10 feet away.

The Nunamiut name Ovingoayook means “whistling.” Yellowlegs
are not a numerous population, but they are conspicuous in habit and
appearance. As one approaches a wet marshy area the yellowlegs
may rise close at hand and fly fluttering about to land and balance on
a slender willow branch, there continuing its sharp whistling call.
Soon it may leave to flutter and repeat the balancing and calling.
Even at a distance too great to explain the disturbance by intrusion
of the observer, yellowlegs may rise, fly about, and balance upon the
willows calling so that their restless behavior reveals the location of
the population.

Although very attentive to human interference, they are apparently
not deeply agitated by contact with man because captured yellowlegs,
like alee readily accept a prisoner’s life and soon learn when
released within a tent, to run about among and over the children and
people. They feed readily upon bits of meat or crumbs and some have
been kept for several weeks in this close association with the families.
Their gentle behavior in captivity is like that of well-mannered guests.
This and their demonstrative attention to human company when free
makes them well liked by the children and adults, who appreciate their
dainty friendliness. Their behavior contrasts with that of the sullen
jaegers, which can also be kept for a time, but only as inexpressive
captives showing no sentiment but greed.

The population of yellowlegs is conspicuous but not numerically
large in comparison with that of the northern phalaropes. Judging
from the frequency of summer calls of the Wilson snipe, they are as
numerous as yellowlegs, but a snipe is seldom seen in summer. The
spring numbers of yellowlegs are somewhat greater than in summer, so
that there is a migration northward, where there are many marshy
areas among willows resembling those inhabited in the mountains.

Erolia melanotos (Vieillot)

18 males May 18-June 3 weight (25), 64-105, — —_—
average 86 g.
9 females May 18-July 29 weight (10), 58-69, — _

average 60 g.

70 U. S. NATIONAL MUSEUM BULLETIN 217

The earliest arriving pectoral sandpipers were recorded May 18,
1949, May 24, 1950, May 20, 1951, May 26, 1952, May 18, 1953, and
May 27, 1954. In the next week, considerable flights passed through,
obviously on the way northward. By early June, migrating flights
had ceased, but birds were frequently seen and occasionally heard in
the grassy marshes in which, however, like snipe, they lay rather close
and remained inconspicuous.

The eggs in several females were measured in 1951, and their length
was found to be 1 mm. on May 24, 2 mm. on May 26, 4 mm. on May
27, 4 mm. on May 28, 4 mm. on May 29, and 4 mm. on June 1. From
these figures, I derive the belief that nesting occurred after June 1.

No nests were found but the Nunamiut consider them to be nesting
birds in many grassy places, where the soft booming sound of the
males is often heard through June. The source of the sound is indi-
cated in the name Poviaktook, which means “inflating the chest,” and
nesting is considered to occur rather commonly, but not to be easily
discoverable. I am disappointed by the vagueness of my summer
information upon the pectoral sandpiper, the nesting of which is re-
ported upon so commonly in other arctic regions.

A female taken July 29 was in fresh plumage, with adult character-
istics. It was one of a group of three found near the north end of
Margaktuk Lake and is considered to be a resident bird, for no flight
movements had yet been seen. In fact, southward flights have not
been reported, and as in most species, they apparently do not form into
flocks in the valleys.

Erolia fuscicollis (Vieillot)

1 male June 1, 1954 weight (2), 40.2, — —
31.7 g.

The first white-rumped sandpiper to be recorded was collected by
Simon Paneak and so named by him in the field from Peterson’s
(1941) descriptions. On the same date he weighed another male,
found alone. Both birds were fat and the testes, which were 8 and 9
mm., respectively, were progressing toward breeding size.

Bailey (1948) reported that 8 specimens and 2 sets of eggs had
been collected along the arctic coast between Wainwright and Demar-
cation Point. It is possible that the two specimens were migrating to
nest further north but the record of two males only warrants their
designation as visiting birds.

Erolia bairdii (Coues)

34 males May 15-July 26 weight (28), 32-48, — —_—
average 39 g.
8 females May 16-July 20 welght (12), 34-45, — ==

average 39 g.
1 young male July 26 weight 26.4 g. — ==

ANAKTUVUK PASS Tt

The first arrivals of Baird’s sandpiper were recorded May 18, 1949,
May 29, 1950, May 18, 1951, May 26, 1952, May 138, 1953, and May 12,
1954, In 1949, 200 were reported seen May 16 and again on May 17,
but by May 23 only 8 or 10 were reported daily. The early large
numbers seen along the lakes and streams probably included many
northbound migrants as the resident birds often resort to the higher
and drier parts of the tundra, even to elevation 1,400 feet above the
valley floor. The rush of migration passed in late May 1949. ‘These
sandpipers, which are only known to nest north of the arctic circle,
make their way northward in large numbers through the Pass.

On June 6, 1951, John Krog found a nest about 1,400 feet above the
valley floor in the mountains west of Tuluak Valley. Robert Paneak
found the young just breaking through the shell in a nest on the val-
ley floor June 24,1951. A female contained an egg 20 mm. long May
28,1951. These observations set the date of nesting as about June 1.

Often they stand, like plovers, rather alert and suspicious, taking
wing to rise swiftly and soar about singly or in pairs. They are more
shy than semipalmated and least-sandpipers, but more conspicuous on
the dry ground which they often occupy. During migration they are
more conspicuous than the semipalmated and least sandpipers as is
also the case of the resident birds insummer. They appear to be about
as numerous as any migrant shore birds in their short season of flight.
In summer they are probably not as numerous as semipalmated sand-
pipers.

The Nunamiut cal] Baird’s sandpipers Vwuksruk, which means
that “they sound like a man with a bad cold.” They can be confused
in the field by the Nunamiut, as by us, with semipalmated sandpipers,
but the name, like the birds’ habits, was specifically distinct and was
correctly applied in about 20 cases when skins were examined for a
test.

Erolia minutilla (Vieillot)

4 males May 16-June 8 weight (16), 16-23, — —_
average 20 g.
5 females June 5-June 11 weight (14), 17-27, — =

average 22 g.

Least sandpipers arrive in the latter part of May, and when first
seen migrating are difficult to distinguish from semipalmated and even
Baird’s sandpipers. First arrivals, often in pairs, have been recorded
May 30, 1950, May 16, 1951, May 29, 1952, May 22, 1953 and May 15,
1954. As one becomes accustomed to their habits and as they separate
into their habitat, which is usually small streams and pools, their be-
havior is quite distinctive. They stand more quietly and look more
delicate than the larger sandpipers. While capable of swift flight,
they often flutter their wings like a moth in making turns, whereas

469496—60——-6

72 U. S. NATIONAL MUSEUM BULLETIN 217

semipalmated sandpipers bank and glide swiftly through their turns.

The faint sweet song of these two sandpipers has brought their
similar Nunamiut names. Liva livawrak means “smaller Liva liva”
and designates the least sandpiper. Its diva liva song is repeated more
faintly and more rapidly than that of the semipalmated sandpiper.
Like its mothlike behavior in flight, Liva livaurak’s song reminds me
of an insect call, while that of the semipalmated sandpiper has the
qualities of a frog’s call. The diva liva song may be given from the
ground or from high in the air, but I have never with certainty seen
more than one Liva livaurak at a time calling in flight.

A female examined June 5, 1949, contained an egg about ready to
lay, but I have no other observation related to nesting.

Least sandpipers are more numerous in spring than later and a con-
siderable number fly on northward. They are less conspicuous and
more retiring birds than the larger sandpipers, less common in sum-
mer than semipalmated sandpipers and probably fewer than Baird’s
sandpipers. Nevertheless they are common and familiar birds which
the Nunamiut like for their song.

Erolia alpina pacifica (Coues)

1 female June 10, 1954 weight 58 g. — ==

On May 14 and May 16, 1949, Thomas Brower reported seeing a
red-backed dunlin. Because these distinctive sandpipers were familiar
to us at Barrow his sight record is considered certain. On June 2,
1952, Simon Paneak saw one at Summit and on June 5 I thought I saw
one in flight, but these were both short views and they are uncertain
evidence. On June 10, 1954, Simon Paneak obtained the specimen.

The example had little fat and its eggs, at 4 mm. in length, were
just starting to develop. These sandpipers are so distinctive in ap-
pearance and so familiar to our view on the arctic coast that the few
records in the mountains show that dunlins rarely alight in Anaktu-
vuk. The evidence does not show the use of Anaktuvuk as a regular
migratory path and at present I regard the birds recorded as visitors.

Limnodromus scolopaceus (Say)

21 males May 23-June 3 weight (28), 90-114, — —
average 100 g.
10 females May 22-June 3 weight (11), 93-119, — —

average 109 g.

Pitelka (1950) gives as mean measurements for the long-billed
dowitchers, which he designated Zimnodromus scolopaceus, the fol-
lowing: males—wing 139, culmen 62; females—wing 144, culmen 72.
Our dowitchers agree closely with these measurements. _

The mean weight of birds of this race taken in spring and fall in

ANAKTUVUK PASS ek

British Columbia and the United States is given by Pitelka (1950,
p- 17) as males (11), 99 grams, females (7), 115 grams. In view of
the known capability of some species of birds to deposit large amounts
of fat and as small warm-blooded animals necessarily consume their
intrinsic resources rapidly, weights recorded at different seasons and
localities might not be of much use as indices of race. The agreement
of our weights in the arctic with those of Pitelka shows that these
dowitchers do not vary significantly in weight with season and place.
Apparently they neither store nor exhaust much substance during
migration.

In 1954 ten examples of dowitchers were weighed at Anaktuvuk on
May 25 and 26 by Simon Paneak. This sampling, taken in a spring
season reputedly uncommonly cold, did not deviate significantly from
the mean weights of the preceding 5 years.

By comparison with specimens in the U. S. National Museum our
dowitchers agree in characters with those from Alaska marked
scolopaceus.

The flights of dowitchers are numerous but pass rather swiftly.
The first records are June 3, 1948, May 20, 1949, May 26, 1950, May 18,
1951, May 29, 1952, May 16, 1953, and May 25, 1954. Many birds
are seen in pairs but the largest developing egg found was 9mm. In
1949 the maximum number recorded by Thomas Brower was 200 on
May 30. Soon thereafter only a few were seen and it is believed that
they pass rapidly to nesting grounds further north.

The Nunamiut name, Azlyaktalik, means “like a bundle when seen
from behind,” a rather apt description which should not, however,
disparage the excellent ability of dowitchers in flight. They are
said to be seen very rarely in the valleys during summer, but 140 miles
west, along the Ahlasuruk, we have seen dowitchers in summer which
acted as if nesting (L. Irving and Paneak, 1954). While it is thought
that they may nest at Anaktuvuk, no indications of nesting are
explicitly recalled, so I do not list them as nesting birds.

Micropalama himantopus (Bonaparte)

9 males June 1-4 weight (10), 53-66, — =
average 58 g.
3 females, not June 8-12 weight 54, 56, 55 g. SS a Tay
preserved.

Stilt sandpipers were also recorded from two examples which Ray
Hock and I examined after they had been taken on May 30 and
June 3, 1948, and in observations by Thomas Brower between June 1
and 6, 1949. Since that time they were not again reported until
between May 31 and June 12, 1954, when two singles, a group of 8,
and a group of 3 were reported by Simon Paneak.

74 U. S. NATIONAL MUSEUM BULLETIN 217

The Nunamiut did not give me a name for stilt sandpipers. Our
failure to notice them more often may arise from our lack of
familiarity with these birds. I cannot believe, however, that many
have been present without attracting our attention. Even A. C.
Bent (1929) remarked that he could not give a general pattern for
their appearance in various localities, and like some other not un-
common birds, records of stilt sandpipers seem to provide no clear
track of their movements. I consider those which we have seen to
have been normal migrants, but they are not common and may not
appear in some years.

Ereunetes pusillus (Linnaeus)

27 males May 21-July 20 weight (32), 20-29, — —
average 24 g.
14 females May 20-July 24 weight (10), 21-27, — —

average 26 g.
5 downy young June 26-27 == == ==

3 young males July 20-26 weight 20.5, 19.1, — —
20.5 g.
1 nest with 4 June 12, 1949 === Sess —
fresh eggs and
male bird.
1 nest with 4 June 18, 1949 aoe — —

largely incubat-
ed eggs and male
bird.

Semipalmated sandpipers are the most common sandpipers. Large
numbers arrive in early May, but because of easy confusion with
least sandpipers, the earliest records may not be first arrivals. They
have been reported May 20, 1949, May 30, 1950, May 14, 1951, May
24, 1952, and May 27, 1954. The earliest five males to be collected,
between May 21 and May 23, were heavier than any males subsequently
taken, weighing from 26.0 to 29.4 grams, and were then certainly not
exhausted from their northward flight.

Soon after arrival they spread over the marshy tundra, feeding
much by pools and lake edges and small streams, while the numbers
diminished as some of the migrants moved north. During early June
they are conspicuous as one, two, or several birds ascend rapidly some
200 feet in the air to circle about singing, and carrying out aerial
evolutions in alternate fluttering and soaring flight. The sweet diva
liva song, faint in the distance, was indefinitely repeated without
variation or interruption, but because of the numbers and their
distance I could not tell how long each bird sustained its calling. As
I watched a pair on a sandy bank on Tuluak Creek, the one which
was more forward in its attentions called repeatedly in 15-second
runs, while the companion or mate called only occasionally in short
runs of a softer sound. Frequently I have heard them on the ground
calling only the two sounds, liva Kiva.

ANAKTUVUK PASS 75

A female bird examined on May 24, 1951, contained eggs 10 mm.
long and one on May 28 had eggs 12 mm. long. Nests were found June
12 and June 18 with respectively fresh and incubated eggs. On June
26 a single downy female weighed 9.1 grams and was just able to walk
slowly with an adult female in close attendance. Near Itikmalikpuk
Creek on June 27 we found 4 small downy young huddled in a nest,
still unable to stand. Their weights were 1 male, 4.3 grams; 3 fe-
males, 3.4, 3.5, 3.6 grams. At this date semipalmated sandpipers were
feeding among the small waves on the sandy lea shore of one of the
small Pitaich lakes. About a dozen were associated but not formed
as a flock. From that time on, they are seen in larger numbers feed-
ing along the sandy beaches of the windward shores of the lakes if the
waves are small. Their association in these meeting places becomes
closer as the young birds grow up, and they become more concentrated
along lake and river shores than in their earlier wide distribution
among tundra pools and streams.

The Nunamiut call all small sandpipers Ziva liva, but this name
really designates the song of the semipalmated and least sandpipers.
These two are called Liva livakpaurak by the Nunamiut meaning
“larger Liva liva” and Liva livaurak meaning “smaller Liva liva.”
At first view these and Baird’s sandpipers are not readily distin-
guished, but upon careful observation one sees that they look, walk,
fly, and sing so differently that it is not surprising that many, but
not all, Nunamiut adults and children name each with specific pre-
cision. The arrival of Ziva liva is a welcome event in spring and
people may stand alone or in groups to search with delighted interest
to locate the obscure source of the sweet liva liva song in the small
birds flying high overhead.

Tryngites subruficollis (Vieillot)
4 males May 29-June 3 weight (4), 64.2- _—— —-
80.5, average 71

g.
6 females May 29-June 5 weight (6), 50-58, —— —
average 53 g

Buff-breasted sandpipers are often seen in spring migration. The
earliest observations recorded are May 24, 1949, and May 31, 1950,
and the brief but fairly numerous flight is soon over. Probably for
this reason it is not always observed.

They were well known on the arctic coast to the Nunamiut as
Aklaktak, meaning “spotted,” but they are not known to remain in
the mountains.

76 U. S. NATIONAL. MUSEUM BULLETIN 217

Limosa lapponica (Linnacus)

A small flock of godwits was reported on May 28, 1950, at Tuluak
Lake and a few were heard on May 18, 1953. During each spring a
few are said to appear in small groups, but none are known to remain,
although their nesting at the mouth of the Colville in 1911 was recalled
by Simon Paneak. Although I have not seen them, the clear descrip-
tion and Nunamiut name, Torhatoruk, convince me that the occasional
view of migrating godwits represents normal migration through
Anaktuvuk.

Crocethia alba (Pallas)

2 males June 4, 1949 weight 52, 51 g. —_ ——
1 female June 4, 1949 weight 52 g. cod =
Sanderlings were recorded in 1949 between June 1 and June 6,
when they were not uncommon. I saw a single one on May 29, 1952.
It seems that their migration through Anaktuvuk Pass is brief and
not numerous but of frequent occurrence, and that they do not linger
to be observed. The Nunamiut name, Kimithkoilyak, meaning “having
no heel,” is appropriate and would not be applied without close ex-
amination. .

Family PHALAROPODIDAE: Phalaropes

Phalaropus fulicarius (Linnaeus)

3 males May 30, 1950 weight 42, 41, 37 ¢. ee ——
2 females May 30, 1950 weight 52, 51 g. == Sa

Some red phalaropes were seen late in May in 1948 and 1950. The
early records are May 18, 1951, May 18, 1953 and June 9, 1954. The
red phalaropes examined by Simon Paneak in 1954 had little fat.
Eggs in the females had developed to about 2 mm. and the testes in
the males were from 9 to 12 mm. in length.

Red phalaropes are usually seen in small groups or alone. Often
they are so much engaged in their feeding or courting activities as
to be quite undisturbed by human approach. Their Nunamiut name
is Auksruak, which means “colored like blood.” They are familiar
spring migrants which act as if visiting for a short time or even con-
sidering residence, but they are not known to remain in the Anaktuvuk
Valley in summer. Judging from their behavior in the Valley, some
are close to their nesting ground. The Nunamiut know that many
nest along the arctic coast. I have no record of their southbound
flight at Anaktuvuk, but in late summer at Barrow large flocks are
seen flying westward along the shore.

ANAKTUVUK PASS Ti

Lobipes lobatus (Linnaeus)

9 males May 23, July 24 — weight (14), 29-35, = iE
average 32 g.
12 females May 18, June 24 weight (7), 29-33, — —_

average 35 g.
2 nests, each with June 22, 1949 —_— -— —_—
4 fresh eggs and
1 parent

The earliest records of northern phalarope are May 18, 1951, May
24, 1950, May 30, 1949 (when 40 were seen), May 18, 1952, May 19,
1953, and June 1, 1954. In late May, many migrants are seen, but
within a few days the migrants hastily pass and the remainder of
these nervous but friendly little birds settle on most of the pools and
lakes of the Valley. The Nunamiut name Kaiyiorgon, meaning “float
like an Eskimo in a kayak,” aptly describes the appearance of phala-
ropes floating lightly on the water with head erect and dark eyes alert
like a watchful Eskimo in a kayak.

Examination of developing eggs in 15 birds between May 18 and
June 2, 1951, showed none longer than6mm. The first nest reported
that year was found by John Krog at Imaiginik June 8.

These phalaropes are very numerous, probably equaling in their
numbers the Baird’s and semipalmated sandpipers,

Family STERCORARIIDAE: Jaegers, Skuas

Stercorarius pomarinus (Temminck)

Pomarine jaegers are uncommon in the mountains. They are well
known to the Nunamiut, from their experience on the arctic coast, as
Isongngakluk. Accordingly, I credit Simon Paneak’s sight records
on May 11, 1950, and two dark and one light bird on June 10, 1950,
and Elijah Kakena’s report of three on May 31, 1952.

In the mountain valleys, they are said to be occasionally seen in
small groups in spring, but are thereafter much less frequently ob-
served than other jaegers and usually as single individuals. It is not
known that they nest in the Valley, and they are so uncommon that I
call them visitors. I would be surprised if these jaegers migrated
northward through Anaktuvuk, for they are so seldom reported from
interior Alaska that I suspect that those which come in to the
mountains come from the north.

Stercorarius parasiticus (Linnaeus)

2 males, 1 dark, May 27, June 9 weight 513, 428 g. —— =
1 light

3females,1 dark June 11, July 29 weight 540, 520, 412 —_ a
2 light g.

The earliest recorded sight of 6 parasitic jaegers in 1949 was May
22, at Tuluak Lake. One was reported May 21, 1951, one on May 30,

78 U. S. NATIONAL MUSEUM BULLETIN 217

1952, one on May 23, 1954, and one on June 1, 1954. In the 1950
camp near Imaiginik it was the most commonly seen jaeger, whereas
usually long-tailed jaegers are more numerous.

As a dark pair were approached while feeding at the carcass of a
dog on the river bank July 14, 1950, two small gray birds the size of
large sandpipers ran into the willows nearby while the old birds
waited about and then flew only a short distance as if they had young
to look after. We could not find them.

On June 14, 1951, Raymond Paneak showed me two eggs which his
mother and he had found the day before. They were situated in a
slight depression of the vegetation among small, marshy pools on the
tundra near Kiminiaktuk. A single bird left the nest at some distance
from us, but it could be clearly distinguished.

Their Nunamiut name, Wirgiaksyook, means “go after vomit.” It
is much better than our name, jaeger, for it refers to the common
habit of these jaegers along the coast which is to harry gulls until they
disgorge their food, which the jaegers then seize. They also feed
greedily upon carrion. They regularly frequent Anaktuvuk Valley,
commonly searching the tundra in twos and threes. Young birds are
occasionally seen, and nesting is considered to occur regularly.

There appeared to be two dark and one or two in light plumage
searching the tundra regularly in the vicinity of Tuluak Lake in 1950
during the summer. In spring a few more are usually seen, either
in pairs or small groups. On June 5, 1952, a group of 17 was seen
flying northward. They are seen at Point Barrow as early as our
records in the Pass, and from the dates they might as well have come
south to Anaktuvuk from the Colville Valley as from the southern
interior.

Stercorarius longicaudus Vieillot

1 male, in 2d year June 30 weight 216 g. — —
1 male —— weight 305 g. — —
2 females May 26, June 20, weight 347, 353 g. — —

1949

Long-tailed jaegers have been earliest recorded as four live cap-
tives received June 1, 1948, and caught a few days earlier. Other
early dates are May 20, 1949, May 11, 1950, May 27, 1951, May 24,
1952, May 22, 1953, and May 28, 1954.

The new tail feathers of the second-year male are only the length
of those of a parasitic jaeger. The bill is small and not distinguish-
ing. The gray feathers of the ventral area are barred as in immature
birds, although the throat is faintly yellowish, indicating transition
from immature to adult state. The identification is based upon the
light coloration of the tarsi as compared with the black feet. The
other birds are clearly distinguished.

ANAKTUVUK PASS 79

They are called Jsongnatcheak, meaning “young or new jaeger” in
Nunamiut. In spring they are frequently seen in pairs or small groups.
Apparently they are then in migration, for they are less numerous dur-
ing the summer. On June 5, 1951, a female contained eggs 18 and 12
mm.long. The female taken June 20, had a bare spot, as if incubating.
Young birds have been reported, although I have not seen them. Long-
tailed jaegers are usually more numerous than parasitic jaegers both
in spring and summer.

Our early records in Anaktuvuk Valley are later than the time for
arrival of jaegers at Barrow, as Tom Brower first pointed out to me.
It may be that jaegers and some other birds migrate southward from
the arctic coast in spring rather than into the mountains by the route
through interior Alaska. Either route would be consistent with our
date records for jaegers.

A number of captive long-tailed jaegers lived well for several weeks,
but they became extremely dirty. They ate voraciously, and showed
neither fear nor curiosity. They did not evince any interest in their
surroundings nor did they appear depressed by them. Their stolid
behavior resembled that of captive gulls, but was quite unlike the
lively interest and adaptability to domestication which is shown by
some sandpipers, ravens, and jays and by some of the Fringillidae.

Family LARIDAE: Gulls, Terns

Larus hyperboreus barrovianus Ridgway

1 male, adult, not May 18 weight 1605 g. —— SI
preserved

1 male, immature, June 11 weight 1824 g. == ==
in 2d year, wing 435 mm.
white culmen 60 mm.

The white-plumaged gull with some faint dirty brown on the breast
appears to be a second-year bird and these measurements conform
with those given by Bailey (1948) for western glaucous gulls from
Barrow. The other male, with matured testes, was fat.

These gulls are called Nauyavak by the Nunamiut, meaning “larger
gulls.” The earliest recorded arrivals are May 14, 1949, May 10, 1950,
May 6, 1951, May 29, 1952, May 4, 1953, and May 6, 1954. Between
May 19 and 22, 1949, from 5 to 8 were seen daily. Between May 6
and 14, 1951, a few were seen each day, generally flying northward.
It would be possible for these wide-ranging gulls to have come to
Anaktuvuk either by northern or southern routes, for they arrive
earlier at Barrow than at Anaktuvuk. During the summer of 1950
an occasional solitary bird patrolled the river daily. Frequently it
was seen standing alone on the river bars. When disturbed it would
fly only a few hundred yards as if reluctant to leave its territory.

80 U. S. NATIONAL MUSEUM BULLETIN 217

The Nunamiut say that nests are occasionally found in the Valley
and I do not hesitate to designate it a regular but uncommon nesting
bird. In spring many more are seen than in summer. I have not
seen them along the upper Koyukuk and Alatna Rivers, where the
herring gull seems to be the regular large nesting gull of the interior.
Further west, on the upper Kobuk and Ahlasuruk Rivers, about 125
miles from the coast, the glaucous gull is very common, while the
herring gull is rare.

Larus argeniatus smithsonianus Coues

2 males July 12, 1949, July weight 1189, 1293 g. ——= =
27, 1952

The first specimen was obtained by Robert Rausch. One was re-
ported seen on July 2, 1951, by Simon Paneak and William Irving
on the Killik River within the mountains, but they are only rarely
seen in the mountains. I have seen herring gulls with their young
of the year along the Koyukuk and Alatna Rivers within 100 miles of
Anaktuvuk. An adult male collected on the Alatna was identified as
smithsonianus. . ,

These and the few glaucous gulls of arctic interior Alaska are wary
and difficult to shoot or to approach within range for distinct recogni-
tion. While this conduct embarrasses the naturalist, it is a blessing
for the resident people that gulls do not rob and foul the meat caches
as they do the environs of every boat and village along populated
coasts. It appears that large aggregations of gulls and people induce
reactions in behavior quite different from that which is natural to
individuals or small groups.

The Nunamiut know the large gull with black wing tips which occa-
sionally appears in the mountains but they give it no explicit name.
It is probably an occasional but not infrequent visitor from the for-
ested valleys of the central interior.. There it seems to be common
and the glaucous gull rare.

Larus canus. brachyrhynchus Richardson

3 males May 17-20, 1949 weight (5), 394-452, —_ —
average 428 g.

1 female June 20 weight 408 g. = =

1 female, in one June 20, 1949 weight 300 g. == at
year plumage,
badly worn

1 female, of this Aug. 7, 1950 weight 422 g. === =
year

Mew gulls were first recorded on May 13, 1949, May 16, 1950, May
3, 1951, May 29, 1952, and May 18, 1953. Small flocks of at most 15
are sometimes seen during the last two weeks of May., Thereafter,
during the summer, a few pairs are about and one or two may be seen

ANAKTUVUK PASS 81

almost any time flying over Anaktuvuk Valley, feeding on the river
banks or lakes. The Nunamiut call them Nauyatcheak. They say
that some are seen even at the mouth of the Colville.

A single egg was found on the wet mosses of a little hummock in
a small tundra lake near Summit, June 24, 1951. One bird was sit-
ting on the nest, and as I approached, it took off and was joined by
another which sailed about over me as I took pictures.

The young first-year female bird was taken August 7, 1950, on the
Kilhik River near Akmalik Creek. It had attained adult weight
but not length. It flew strongly and might have come from a distance,
but it was in the company of adult birds which acted as if in residence.

The latest record in 1950 was September 8 at Contact Creek. Mew
gulls are common summer residents which nest in the Valley. Greater
numbers pass through in spring than remain as residents. These are
probably northbound migrants.

Occasionally these gulls fly high among the mountains. I watched
one soaring with more grace and speed than an eagle as it rose over
a 5,000-foot mountain. Early in summer, small groups carry on aerial
evolutions, calling as they fly above the mountains and often pass
vertically beyond range of glasses. I estimate that they reach alti-
tudes of 10,000 feet.

Rhodostethia rosea (Macgillivray)

Ross’s gull is well known from earlier annual visits of the Nuna-
miut to the eastern arctic coast, and it is called by them Kakmakloak.
At Barrow I saw these beautiful gulls first as if coming from the east
early in September. Only one has been reported to me in the moun-
tains, when in April 1949, it circled and hovered about the camp at
Kalutak Creek for some minutes watching and watched by the chil-
dren, who were joined by several adult witnesses before it flew away.
The event made a distinct impression upon the people. The reported
occurrence of this coastal gull in the mountains and in spring is so
far beyond the scope of its reasonable wandering that this report does
not entitle it to inclusion among the known avifauna of the Pass. The
Nunamiut do not know of its nesting place.

Xema sabini (Sabine)

One Sabine’s gull was reported seen by Thomas Brower at Tuluak
Lake on June 24, 1949, and the Nunamiut say that they occasionally
see it in the mountains. This beautiful gull is well known as Kadga-
giak to the Nunamiut from their former annual visits to the eastern
arctic coast. The name is said to refer to the sound made by the gull.
The name also resembles that of the willow ptarmigan and there is
something about the gull which reminds me of a male ptarmigan as it

82 U. S. NATIONAL MUSEUM BULLETIN 217

begins to gain specks of dark color in summer. I do not hesitate to
designate Sabine’s gull a normal visitor.

Sterna paradisaea Pontoppidan

3 males May 25-July 20 weight (4), 90 to == ==
122, average 101
g.

4 females June 1—July 20 weight (4), 84 to == ===
101, average 96 g.

1 young male Aug. 8, 1950 weight 102 g. —— ——s

1 young female Aug. 8, 1950 weight 101 g. — =

Arctic terns were recorded as seen June 1, 1948, June 1, 1949, May
25, 1951, May 29, 1952, May 23, 1953 and June 1, 1954. Migrant and
resident behavior of terns is not distinguishable until the regular
flights over their fishing grounds become established at nesting time,
but inasmuch as the early population exceeds that in summer, some
must be migrant birds. In May, terns could scarcely subsist on the
arctic slope, and it is quite certain that they arrive from the south.
They are well known as Mitkotailyak to the Nunamiut, and in some
of the old stories, terns are important, usually interesting, and often
spicy characters.

In the summer of 1950, about a dozen terns daily patrolled the river
near Imaiginik occasionally cutting out to circle, or where it seemed
favorable, to fish a lake. Their nesting place was described as among
the gravel and rock bars some 10 miles down river, from which they
flew in the morning and toward which they returned in the evening.
This colony appeared to be the only one along about 20 miles of valley
under observation. A like number was found present in 1951. A
female examined in Tuluak Valley June 2, 1951, had laid an egg and
still contained another ready to lay.

No young birds were seen until a colony of a dozen was found on
the Killik River, roosting on the steep cut earth bank about 15 feet
high opposite the mouth of Akmalik Creek. The two young birds
obtained flew well; they were above the mean weight of adults and
equal to them in wing length, but had not the adult length of tail.

Family STRIGIDAE: Typical Owls

Bubo virginianus lagophonus (Oberholser)

1 female Nov. 27, 1947, Hunt = sr a
Fork
The single specimen of a northwestern horned owl was obtained
from within the forested region at Hunt Fork and sent to George
Sutton, who kindly gave me this identification. Horned owls are well
known in the spruce forest and called Nukisirgak by the Nunamiut.
In a discussion among four Nunamiut hunters, they recalled from their

ANAKTUVUK PASS 83

experience of some 40 years, several instances of the capture of horned
owls in wolf traps north of tree line. The valley is close to their nor-
mal range, and, while they are not common visitors there, it is there-
fore not surprising that these powerful owls venture over the tundra
for exploration.

Nyctea scandiaca (Linnaeus)

1 female Feb. 27, 1951, weight 2267 g. — —
Contact Creek

Snowy owls are frequently seen in the Anaktuvuk Valley during
the winter, and they are well known by the Nunamiut name Oxpik.
Simon Paneak saw individual owls almost daily north of the moun-
tain line in late March and early April 1951, and from the reports of
other Nunamiut hunters then taking advantage of the lengthening
days for extended hunting excursions, Simon remarked that there
were quite a few owls in the Valley. The latest recent date for one
to have been reported came from John Krog, who saw one May 25,
1951, at Tuluak Lake. They have occasionally been seen in summer
at Chandler Lake, but no nests have been found in the mountains. I
have heard no comment indicating in what circumstances the small
numbers of owls seen vary systematically from year to year, nor is there
any comment to indicate that there are cyclic changes in the small
population of lemmings in the mountains. Any considerable change
in either population would have been observed and remarked upon.

Along the arctic coast of Alaska the cyclic variation in numbers of
lemmings is attended by variation in the numbers of all predators. I
do not have enough information about the variation in numbers of
snowy owls in the central parts of Alaska for it to serve as evidence
for a cyclic migration. Neither do I have specific evidence that the
winter movement of snowy owls to Anaktuvuk has the consistency
which appears in the regular annual migrations of birds. For the
present it seems best to designate snowy owls as visitors from the north
coming to Anaktuvuk in winter.

Surnia ulula caparoch (Miller)

1 male Mar.1, Hunt Fork, weight 350 g. —= ——
1 female Feb.10, JohnRiver weight 350 g. — ——
American hawk-owls are well known as Neakoktoakruk among the
Nunamiut to be dwellers in the timbered country, where they have
been seen as far north as the last spruce at Publatuk Creek and on
the upper Savioyuk River. They are not reported as venturing north
of timber and are not included among the avifauna of Anaktuvuk
Pass. Our records show that some hawk-owls of the northernmost
population in the forests are winter residents.

84 U. S. NATIONAL MUSEUM BULLETIN 2:17

Strix nebulosa nebulosa Forster

The American great gray owl, Vattak in Nunamiut, is well known
in the spruce forests and is occasionally seen at the northern limits
of spruce between Hunt Fork and Publatuk Creek. It is not con-
sidered likely to pass north of timber and, like the hawk owl, is not in-
cluded in the avifauna of Anaktuvuk Pass. Itis my opinion that this
owl and the hawk owl are closely attached to the forest environment.

Asio flammeus flammeus (Pontopiddan)

1 female May 29, 1949 weight 400 g. —— ==

Short-eared owls, known as Vipailyutak to the Nunamiut, are regu-
larly seen in Anaktuvuk Valley in summer. The earliest records are
May 18, 1949, May 10, 1950, May 8, 1951, May 18, 1952, May 4, 1953,
and May 15, 1954, at Contact Creek. In 1950, the last was observed
September 11 at Inakpasugaruk Creek. During the summers of 1950
and 1951, one or two were seen frequently as if they were residents,
and they undoubtedly nest in the Valley.

On one occasion, I shot at and missed a short-eared owl. As it flew
away, some small bird the size of a redpoll rose to attack it. Ata
distance of about a mile, a golden plover harassed it severely and
when last seen, two terns were pursuing the owl out of sight. The lives
of these predators seem to be much disturbed by those whom they are :
represented to oppress.

In spring, and particularly as evening twilight develops in the late
summer, several of these owls are often seen together and they are
apparently then in migration to or from the arctic slope.

Something about short-eared owls seems to favor their use in Nuna-
miut stories to exemplify human traits and characters. In one a young
woman who was unbearably influenced by unsatisfied sexual impulses
was transformed into the erratically flying owl.

In another Nunamiut story, a short-eared owl and a Pacific loon
were at one time arguing as to which could first fly from the moun-
tains to the arctic coast. The loon asserted that he could fly much too
fast for the owl’s company and set out in its swift linear flight, while
the owl floated upward on light wings until it was only a little ball
in the sky. Then it dove far down with the swiftness shown in its
short swoops over the tundra until it came to rest on the coast where
it was quietly waiting when the loon arrived all out of breath from
the labor of its direct flight.

The Eskimos say that the short-eared owls sometimes hang their
small prey in the bushes as do the shrikes.

ANAKTUVUK PASS 85

Aegolius funereus richardsoni (Bonaparte)

1 male Oct. 3, 1950, weight 115.8 g. — —
Savioyuk

These small boreal owls were described as residents of the spruce
forests and were named Zakpilyakruk by the Nunamiut before speci-
mens were at hand for verification. Normally residents of the timber,
they do, however, occasionally venture north. The specimen recorded
is from within the most northern spruce forest at the head of the
Savioyuk River. But one was recorded observed May 18, 1949, at
Tuluak Lake and another September 27, 1950, at the head of Inak-
pasugaruk Creek. From these records they are considered to be
occasional summer visitors to the treeless country of Anaktuvuk
Valley.

Family PICIDAE: Woodpeckers, Wrynecks
Colaptes auratus borealis Ridgway

On May 31, 1952, I saw a yellow-shafted flicker 3 times at the edge
of the willows near the mouth of Inakpasugaruk Creek, and I saw
it again twice the next day. I could find no opinion that one had
previously been seen in the mountains. However, from my descrip-
tion, Simon Paneak recognized it as like the skin of one which David
Tobuk had shot while it was hammering on a pole by his house at
Bettles Village. The people at Kobuk know the flicker well and say
that to have its skin brings good luck. The bird observed at Inak-
pasugaruk is considered a rare visitor from the forested country.

Dendrocopos pubescens nelsoni (Oberholser)

1 female Sept. 7, 1948, —— = Ta.
Tuluak Lake

Another specimen of Nelson’s downy woodpecker was obtained No-
vember 27, 1947, within the spruce forest at Hunt Fork and sent to
George Sutton, who identified it. _

The Nunamiut distinguish two woodpeckers, Zoyuk, and a larger
bird Toyukpuk. The name for the smaller bird refers to the downy
woodpecker, which is well known in the forest and occasionally seen
in the willows north of the mountain line, but only in winter. The
3-toed woodpecker has never been seen north of the forest.

The willows of the northern valleys occasionally reach a diameter
of six inches at a height of from one to three feet above ground, and
there are very rare sparse northern stands of small cottonwoods.
Dead, stunted willows do not look like good residential material even
for a small woodpecker, and I could find no evidence that nest holes
have been seen in the tundra willows. It may be that the downy wood-
pecker nests in timber and sometimes ranges north after the nesting

86 U. S. NATIONAL MUSEUM BULLETIN 217

season, a behavior in keeping with its independent, hardy nature and
one for which its habits of feeding in small brush would well adapt it.
The often associated Yukon chickadee may also nest in the timber and
move north in winter.

I am inclined to think of the downy woodpecker as a visitor after
nesting season.

Along the Yukon and on its Delta Turner (1886) reported these
woodpeckers as common and present in winter.

Picoides arcticus (Swainson)

Charles Sheldon of Kobuk informed me that both a ladder-back and
a black-backed woodpecker with yellow head were well known in the
spruce timber along the Kobuk. To demonstrate his point, he later
sent me the skin of a male of arcticus, obtained April 22, 1954, weigh-
ing 71 grams. Grinnell (1900) had found fasciatus nesting on the
Kobuk at Hunt River. It is apparent that both these forms of Picoides
occur in arctic Alaska near the limit of spruce timber, but it is unlikely
that either would pass north of the forests.

Picoides tridactylus fasciatus Baird

Three examples of northern 3-toed woodpeckers have been brought
to me from the spruce forest in the vicinity of Hunt Fork. These
are a male, January 16, 1948; a female, December 22, 1947, and
another male November 27, 1947. The skins of the two former were
sent to George Sutton and identified by him. The third, obtained in
the spring of 1948, was too badly decomposed for preservation.

This is the woodpecker which the Nunamiut know as Toyukpuk, but
as it means only “larger Z’oyuk,” the name is hardly specific. There is
no indication that these woodpeckers ever abandon the wooded areas
nor does it seem likely that their feeding habits among the large spruce
trees would enable them to do so. But they do dwell in the most
northern timber. They are not included in the avifauna of the Valley.

Family TYRANNIDAE: Tyrant Flycatchers

Sayornis saya yukonensis Bishop

1male,1 female June 17, 1951, — —_ eae
Anaktiktok Val-
ley, Napaktua-
loitch Cave

1male,lfemale June 23, 1952, weight 25, 24 g. —— =
Anaktuvuk Soak-
puk Mountain

1male,1female July 2, 1951, Killik weight 22, 19.1 ¢. — =
Valley, Irivik

Creek

1 male July 29, 1950, weight 26.0 ¢. = TT
Kangomavik

1 female Aug. 7, 1950, weight 20.5 g. Ss a

Akmalik Creek

ANAKTUVUK PASS 87

I had been looking for the Yukon Say’s phoebe although I could
find no recollection among the Nunamiut of their having seen one.
The male specimen collected in 1950, was with another and smaller
phoebe which acted like a young bird among the low willows in a creek
bottom near the western valley wall and about two miles north of
Kangomavik.

The female taken in 1950 was one of three phoebes among the wil-
lows on the sand dunes just south of the mouth of Akmalik Creek
in the Kalhk Valley. It was provided with abundant, soft, yellow
subcutaneous fat.

The latter bird certainly, and the former apparently, were members
of family groups. In 1951, two phoebes were seen near Napaktualoitch
Cave on the north side of Anaktiktoak Valley where we were looking
for archeological remains. Close to them was a characteristic
phoebe’s nest on an overhung ledge above the entrance to the cave.
It contained five rather well-incubated eggs. Although there was an
old robin’s nest nearby there were no other signs of earlier nesting by
phoebes.

The two birds collected in the Killik Valley were likewise near a
eave which William Irving and Simon Paneak were searching for
signs of its ancient use by man. Two other phoebes were present, but
could not be obtained and no nest was found. One phoebe was seen
June 4, 1952, high in a branch of Inakpasugaruk Creek. The two
taken June 23, 1952, were a pair, the female having a bare brooding
area, They were obtained at the base of high cliffs on Soakpuk
Mountain in difficult terrain where the nest could not be found.

I am surprised and the Nunamiut were embarrassed that they did
not know these phoebes, for the bird’s habits and appearance are
conspicuous and distinctive. Their nesting on rock ledges interested
the Nunamiut for the caves and overhanging ledges have been used
in the past for storage. The caves are frequently visited, and it is
hard to see how persistent and regular nesting phoebes could escape
the keen interest of the Nunamiut in natural events of this sort. And
yet in 1950, one family of phoebes was seen in Anaktuvuk and one
in the Killik Valley, and in 1951, one family was seen at a different
location from the preceding year in each valley. In 1950, our observa-
tion was too late to show that the birds nested near where they were
seen, and in 1951, only one nest was found. Simon Paneak was in-
clined to believe that they had been missed in observation rather than
to suggest that phoebes were newcomers in the mountains. I believe
that some migrate further north.

469496—60——_7

88 U. 8S. NATIONAL MUSEUM BULLETIN 217

Family ALAUDIDAE: Larks

Eremophila alpestris arcticola (Oberholser)

15 males May 14-July 22 weight (15), 33-41, — —
average 37 g.

7females May 19-June 23 weight (11), 30-40, — —
average 37 g.

1 young male July 13 weight 31 g. —_— —

1 young female July 22 weight 32 g. —_— —

Seven pallid-horned larks from Tuluak Lake were brought to me
for identification in 1948. They had been taken on May 15, 16, and
23. The dates of earliest record are May 15, 1948, May 18, 1949, May
28, 1950, May 3, 1951, April 28, 1953, and May 3, 1954. ‘The irregu-
larity in the recorded annual arrivals of horned larks result because
the high ground which they frequent is off the common paths of travel
in early spring. No flocks are recorded nor do I know the extent of
the migration except that larks are common in many parts of the
arctic slope to the northward.

The larks apparently nest earlier than any of the Fringillidae. A
large egg was found in a female on May 22. Developing eggs 10 mm.
in length were found in a female examined on May 31, 1951, and an-
other female on that date was noted as having already laid its eggs.
On May 29, 1951, a female was collected near its nest of four eggs.
Four eggs found by Tom Brower on June 18, 1949, were so far
incubated that they could not be saved. A nest with four eggs was
reported to me near Imaiginik on May 28, 1951. Two young specimens
taken on July 13 and 22 were flying and near adult size.

Although they are among the earliest migrant birds to arrive, some
of the larks were plainly moving southward in preliminary migratory
movement as early as August 5. Their arrival in the cold weather of
the arctic spring conforms with the apparent suitability of their
feathers for effective insulation. But in spite of their ability to live
normally in cold weather they begin to leave soon after the warmest
part of the arctic summer and all are gone before the onset of cold
weather.

Larks kept pretty well to the elevated dry portions of Anaktuvuk
Valley and its sides, and in July were found occasionally along the
drier parts of the slopes. They ranged to 300 or 400 feet above the —
valley floor on the edge of the terrain occupied by the wheatears at
the foot of the talus slopes, but they were commonly at lower levels.
None have been recorded seen at high elevations. Larks were never
numerous at one place, but they spread over a large area and therefore
outnumber the wheatears and gray-crowned rosy finches. In 1950
at Summit larks were less frequently seen than Smith’s longspurs.

ANAKTUVUK PASS 89

They are, nevertheless, rather common birds, and generally more
numerous than the latter. Their Nunamiut name NVakrulik means
“horned.”

Family HIRUNDINIDAE: Swallows

Tachycineta thalassina lepida Mearns

In summer I have regularly seen violet-green swallows at Bettles,
but they are the least common swallows there. Only in 1954 did
Simon Paneak report to me that he had killed one on May 29 at
Anaktuvuk. He had seen them at Bettles with me, but as we have
no other report of the occurrence of this swallow in the mountains
I consider it to be too unusual to designate it as a part of the avifauna
of Anaktuvuk.

Tridoprocne bicolor (Vicillot)

1 female June 7, 1949 Weight 19.5 g. — =
1 young male Aug. 15, 1949 —- — ——
1 young female Aug. 15, 1949 — ——— —_

The two young tree swallows were identified by Alexander Wetmore.
They were large enough to fly with adult ability. One was recorded
seen at Tuluak Lake May 20, 1950, by Robert Rausch, and I saw two
at Summit June 5,1952. On the same day some boys clearly described
tree swallows which they had seen. Swallows are said to come
regularly each year into the Valley, but none are known to nest or
reside. The Nunamiut name 7’olugaknek, which means “like a raven,”
is applied to all swallows without specific distinction, nor with much
accuracy as to color. But I must observe that the flight of ravens in
the winds high over the mountain cliffs and peaks, where they may look
almost as small as swallows, has a similar lightness of aerial
maneuver.

Tree swallows are common on the Koyukuk River at Bettles, and in
1951, I have seen them along the Koyukuk and lower Alatna Rivers,
the nearest places of occurrence yet reported. As these places are
less than 100 miles distant from Tuluak, two hours’ flight could
bring in a visiting swallow, and that is probably the status of all
of the swallows seen in the Valley.

Riparia riparia riparia (Linnaeus)
1 male June 6, 1949 weight 16.8 g. —_— —

Occasionally an individual bank swallow is reported in Anaktuvuk
Valley, but none appear to remain. Knowing that Anderson (1921)
had reported seeing a cut bank on the Hula Hula River “thickly per-
forated with holes,” I have looked for their nesting holes in the river
banks among the mountains. I have not found a sign of their nesting

90 U. S. NATIONAL MUSEUM BULLETIN 217

north of the Koyukuk and lower Alatna Rivers, nor did any of the
Nunamiut recall seeing nests north of those localities. The con-
spicuous method of nesting would have brought notice of bank swal-
lows which remained to breed in the mountains. Accordingly, I con-
clude that the bank swallows occasionally seen in the mountains are
visitors from nearby colonies along the Koyukuk and Alatna Rivers,
where I have found them to be numerous and regular nesting birds.
Some of these colonies are within 80 miles, which would be an easy
flight.

The single specimen collected by Tom Brower, two which I obtained
from a regular nesting colony at Bettles, and nine adult and ten young
bank swallows collected for me by Tom Cade from along the Yukon
River in the vicinity of Eagle and Forty Mile all fit among specimens
referred to Riparia riparia maximiliani by Arny (1952), but con-
sidered uncertainly separable from typical riparia. °

The bank swallows are only known as visitors to the Valley.

Hirundo rustica erythrogaster Boddaert
1 male June 6, 1949 weight 17.7 g. — —

The barn swallow is known by the Nunamiut to be an occasional
visitor to their country, but they do not know its nesting place. I
have not seen one in the Koyukuk or Alatna Valleys. Charles Sheldon
of Kobuk clearly described their appearance and nests and told me the
Kobukmiut name for barn swallow. He said that nests were very
uncommon near Kobuk. Tom Brower told me that he collected the
young barn swallows at Barrow which are remarked upon by Bailey
(1948) and that they were too young to have flown very far. Unusual
nesting sites appear so often attributed to swallows that they may be
currently venturing to extend their range. In Alaska, violet-green,
tree, bank and cliff swallows now nest intensively about human
habitations which only a few years ago did not exist. These species
have recently taken new sites for nesting and the adaptability which
they exhibit may also indicate a current tendency to extend the terri-
tory over which they breed.

As far as concerns the occurrence of barn swallows at Anaktuvuk
there is no indication that their rare visits there represent other than
the usual explorations of strong fiying birds.

Petrochelidon pyrrhonota (Vicillot)

In the winter of 1908 or 1909, Simon Paneak recalled, he and some
other boys, who were then near the head of the west branch of the
Kuparuk River (lat. 68° 35’ N., long. 149° 20’ W.) found over a hun-
dred mud nests built against the rock cliffs. When these were knocked
down for examination they were found to contain frozen young

ANAKTUVUK PASS 91

birds which were well feathered and nearly ready to fly when they had
died. Paneak and I discussed this story again as we watched the cliff
swallows building their nests against the houses at Bettles. We agreed
that he had certainly observed a colony of cliff swallows nesting on the
Kuparuk River cliffs some 40 miles northeast of Tuluak. He has
not seen nor heard of such nests anywhere else north of the timber.

The nesting of cliff swallows in natural arctic sites is not unknown.
“On May 28, 1949, Mr. Rae found this swallow on the banks of the
Coppermine, having constructed its clustered nest against the cliffs
at the mouth of the Kendall River, latitude 67° N., but not yet laid
its eggs” (Richardson, 1852, p. 395).

Cliff swallows are common nesting on the houses at Bettles Village
and Airfield, but no other report than Paneak’s has shown them to
nest north of there in Alaska. The invasion of the arctic tundra for
nesting turned out to be an unproductive effort for that particular
colony of cliff swallows.

I do not have any records of their occurrence at Anaktuvuk.

Family CORVIDAE: Jays, Magpies, Crows

Perisoreus canadensis pacificus (Gmelin)

4 males Feb. 17, Apr. 2, weight (1), 79 g. — ——
May 15, Nov. 17

6 females Jan. 26, Feb. 1, 21, —_— — —
July 31, Oct. 24

3 not sexed Mar. 2, Oct. 2 weight (1), 76 g. —_— —

Alaska jays, called Hirik by the Nunamiut, are commonly seen north
of timber in winter, where 11 specimens have been taken along the
headwaters of the John River. During summer these birds are not
seen. The two summer specimens are a full-sized young female found
at Kangomavik July 31, 1950, and a male at Tuluak Lake taken May
15, 1951. Three were reported seen by Tom Brower May 8, 1949, at
Tuluak Lake. A young jay shot in the Killik Valley July 6, 1951, by
Simon Paneak and William Irving was a young enough bird to settle
all doubt that it was raised in the mountains. The Nunamiut remark
that they have never found a jay’s nest is a good comment on the bird’s
extreme secretiveness in nesting season, but a few evidences of jays in
summer and the young bird found on the Killik assure us that jays
nest in the mountains.

Their sociable winter behavior makes the jays well known even on
the tundra in that season in contrast with the mystery surrounding
their nesting. No cold restricts their activity or range, but on cold
winter days they erect their feathers until they appear spherical when
at rest. They are seen so much more frequently in winter that I be-
lieve some jays travel north from their common haunts in the timber

92 U. S. NATIONAL MUSEUM BULLETIN 217

to winter among the small willows of the treeless mountain valleys,
but apparently only a few remain to nest on the tundra.

Corvus corax principalis Ridgway

5 males — weight (5), 1100- — —
1400, average 1240
£.

4 females — weight (3), 1125, — —
1050, 1800 g.

Northern ravens may appear alone in the mountains or in groups
sometimes as large as 25 on almost any day and in any weather. They
may fly singly and direct, or a dozen may soar into view above the
highest mountain to maneuver aloft before diving down in complex
flight patterns, calling as they disappear among the cliffs.

The Nunamiut name the raven Z'’uwluak, which as they sound it re-
sembles the raven’s common call. In the coldest weather, ravens are
active. ‘Their adroitness in flight and keenness in finding caribou
killed by hunters or wolves are much admired. Many stories attest
the Nunamiut’s appreciation for the raven’s aerial skill, hardihood,
and undoubted ability to communicate while hunting and in the course
of their social flight maneuvers. The application of the raven’s name
to a man is a compliment to his searching ability and discretion, or
glossy dark clothes, but it has the same connotation as the name of the
fox in our usage.

Circling ravens often call the attention of Nunamiut hunters to
dead animals upon which bears or wolves may be feeding. Against
this service, the circling ravens may see the hunters and by their
action and calls warn his prey.. Thus the behavior of ravens is signifi-
cant to the other residents of the north. To the Nunamiut ravens
are also interesting, for they recognize in them observant, curious
personalities which communicate with each other and so reveal their
social methods of play and work.

No temperature or weather limits the raven’s appearance, but they
are seldom seen far from the rocky mountains about which they like
to fly and where they nest. Observations in March, April, and May
suggest that there is more raven traffic in those months than in others,
but it is not obvious that there is any seasonal change in territory or
numbers in the mountains.

As to the extent of the raven’s travels in the mountains, I have
a story from Simon Paneak that in his grandfather’s time a man on
the Colville River saw a raven suddenly drop dead from the cliffs.
Examination showed that it had been killed by a whalebone spring
such as in the old days was coiled and embedded in meat or fat and
then frozen, serving by expansion on thawing in the stomach to kill
a fox or wolf. The mark on the whalebone identified its owner.

ANAKTUVUK PASS 93

Later, when the observer of the raven’s death met with the owner
of the whalebone trap, it developed that it had been set near Publatuk
Creek along the John River, about 150 miles from the cliffs of the
Colville. Traces in the snow had shown that a raven had made off with
the bait from Publatuk on the morning of the same day during which .
it had died 150 miles away on the Colville. This incident is considered
by the Nunamiut to substantiate their impression that the common
daily travel of ravens extends over wide areas.

Simon Paneak reported ravens nesting in March among cliffs about
20 miles north of the mountain line. A group of about five, which
appeared like a family, were heard and seen high among the pinnacles
of Napaktualoitch mountain during several days when we camped
near there about June 15, 1951. On June 24, I surprised this group
on a hillside as I walked over a moraine slope just south of Kango-
mavik. Two ravens flew at once, but the other three, which were full
sized, waited until I walked within 40 yards of them, and we decided
that they behaved like young birds.

I consider ravens to be established residents. I believe that some
ravens come into the Anaktuvuk Valley from distant resorts, and that
as the old Nunamiut believed, ravens often make long daily flights.
In the mountains, there is no indication of seasonal migration.

Although displaying a certain cleverness, ravens have not always
been successful characters in old Nunamiut stories. At one time a
young raven was much admired for his bold manner and glossy dark
feathers. He was singled out for attention by the richest man in the
village and eventually Raven won the affection and hand of the richest
man’s daughter. But the affair turned out badly, for Raven was
unable to procure any meat but carrion and the young lady’s proper
disparagement of that fare made Raven’s life miserable.

Before the time of this story and doubtless long ago, Raven and
Pacific Loon both had coats of light-colored feathers about like those
of the glaucous gulls. One day Raven and Loon were discussing the
handicap of their light feathers, which were so conspicuous on the
summer tundra. They decided to darken each others coats with
charcoal.

Raven first worked on Loon and applied the symmetrical black and
white pattern which we now see on Loon’s back. Since Loon could
not hold his belly off the ground it retained the original lightness,
but both of them were rightly pleased with the result. As Loon started
work on Raven’s back the curious bird could not restrain his impatience
to see the progress. After standing a certain amount of Raven’s
fidgeting, Loon said the constant inspection unnerved him and had
already caused him to mark the two sides unevenly.

Then Raven agreed to close his eyes while Loon completed the work

94 U. S. NATIONAL MUSEUM BULLETIN 217

in hand. Smiling quietly to himself Loon marked Raven all over
with black. When Raven opened his eyes and looked curiously at his
feet, back, and tail he croaked with anger at the unhappy sight of his
uniform black appearance. Loon burst out in the crazy laughter for
which he is renowned. Maddened by the thought of how he looked,
Raven seized a handful of ashes from the fire and threw them at Loon.
Loon was smart enough to dive into the lake, but not so quick but that
he got the ash-gray patch on his head as it went below the water.

Now when Loon recalls what he did to Raven he cannot restrain
his long bursts of crazy laughter as he sits safely on the lake. Raven,
on the other hand, when he thinks of the affair expresses his feelings
in his sad hoarse voice.

F amily PARIDAE: Titmice, Verdins, Bushtits

Parus atricapillus turneri Ridgway

1 male Jan. 26, 1950, — — peewe,
north of timber
1 male Oct. 7, 1950, weight 11.1 ¢. = =

Savioyuk River,
just south of
timber
1 unsexed Jan. 26, 1950, —_— = ==
Summit

A number of other specimens of Yukon black-capped chickadee
could not be preserved. Four from Summit were brought into camp at
Kalutak Creek and examined on January 26, 1950. A specimen from
the upper John River, 1950, otherwise without data, was kindly iden-
tified by W. Earl Godfrey. Numbers were observed in late September
and early October 1950 just within the northern limit of spruce forest
on the Savioyuk River.

The Nunamiut have a name for only one chickadee, Misikakak.
They recognize the brownish form and say that Misikakak applies
strictly only to the black-capped kind. It is known and liked for
the way it occupies the willows north of tree line in winter with the
attitude of being the proprietor. The Nunamiut are accustomed to
seeing it north of tree line in winter but not in summer. It is possible
that this chickadee, downy woodpeckers, pine grosbeaks, and possibly
jays, may be required by nesting habits to remain in the wooded area
in summer, but that upon completion of duties for reproduction, they
then wander north.

Although it is difficult to see how extreme arctic cold and wind can be
endured by such a tiny bird, it appears oblivious to both. Its feathers,
like those of the jay and grosbeak, provide effective but thin insulation
and its habit of searching for food on the willow branches and twigs,

ANAKTUVUK PASS 95

which are blown clear of snow in the windy mountains, may make the
wind-cleared twigs of the arctic willows a more suitable winter range
than the still, snow-covered forest.

Parus hudsonicus hudsonicus Forster

1 male Oct. 13, 1950 weight 12.3 g. — —
1 female Oct. 13, 1950 weight 10.8 g. — —

These two Alaska chickadees were obtained by Simon Paneak near
the head of the Savioyuk River and within the spruce timber. The
Nunamiut consider the brownish chickadees to be confined to the
forest and it is not explicitly designated by their name for the black-
capped chickadee. I have no observations to suggest that the Alaska
chickadee ventures north of the forest and I include it as a bird of the
nearby forest but not of the treeless country.

A female was collected at Hunt Fork on February 26, 1950, by
Robert Rausch, an unsexed bird was obtained in 1950 from the wooded
part of the upper John River, and I collected another unsexed bird
on August 15, 1950, near Bettles. These were examined by W. Earl
Godfrey and are deposited in the National Museum of Canada. He
proposed including our Alaska specimens with those which he would
designate P. h. evura.

Family CINCLIDAE: Dippers

Cinclus mexicanus unicolor Bonaparte

1 female Apr. 29, 1949 weight 49.4 gr. _ —

This specimen along with a female dipper from Hunt Fork, Febru-
ary 3, 1948, sent to George Sutton, establish the winter residence of
these birds north of the Arctic Circle, a fact which has been frequently
described in narratives but not, so far as I can tell, earlier confirmed
by specimens or the observation of ornithologists. We have several
additional sight records of these birds with strange aquatic habits
which fit so well into the tiny and uncommon niches of the arctic en-
vironment in which water remains unfrozen throughout the year.

The Nunamiut name for dipper is Anaruk kiviruk, which was trans-
lated as “old woman sunk.”

At Nakagnik Springs, running into Tuluak Lake, Raymond Hock
saw two dippers in May 1948. In attempting to secure them, both
were unfortunately washed under the ice below the open water. At
the same place the following year two dippers were shot by Tom
Brower April 29, 1949. One was recovered and preserved, the other
washed under the ice and was only recovered later when too badly
damaged to be saved. Observations were made at this spring daily

96 U. S. NATIONAL MUSEUM BULLETIN 217

in 1950 and 1951, but no dipper was reported. Raymond Paneak saw
two dippers there in April 1953. At the head of the Savioyuk River
near open water and just within the northern limit of spruce forest,
Rausch recorded an observation of a dipper on October 10, 1950.
Four were reported from Savioyuk on February 10, 1951, by Simon
Paneak.

Two dippers were seen by Robert Paneak at Naniksruk Springs,
a winter fishing place, some 40 miles north of Tuluak Lake, on March
31,1951. One was seen on Kanayat Creek by Elijah Kakena on April
9, 1951.

Where the waters are unfrozen a dull cloud of ice fog called “water
smoke” may be seen for a long distance rising in the still winter air.
These places of open water are called springs by the Nunamiut. They
provide fishermen with access to water in winter, and they are there-
fore carefully examined and well known. In a stretch of about 100
miles south to north on the upper John and Anaktuvuk Rivers,
dippers have been reported in four places during winter. In the
country within 15 miles of that line there are three or four
other places which might be suitable, making places for about 6 or 8
pairs of these small birds in about 3,000 square miles of arctic moun-
tains which are otherwise completely snow and ice bound for eight
months of the year.

Some of these open water locations are only a few yards in extent,
and there are so few of them that individuals or pairs are often sepa-
rated from each other during more than 6 months by 30 or 40 miles
of frozen country completely unsuited for a dipper’s sustenance.
Territories vacated by death are nevertheless filled, so that in spite
‘of the winter isolation of its members and their usual close restriction
to a minute feeding territory, the population tenaciously f90EE the
integrity necessary to preserve its small numbers.

The population density of dippers in northern Alaska is probably
the least of any small bird in the north, and it may be suggested that
the individuals and pairs live in isolation from their kind unique
among animal populations.

Our scant physiological knowledge indicates hypothetical diffi-
culties for so small a bird feeding under water in arctic winter condi-
tions. But the individuals show no signs of strain and are undoubtedly
adapted so that their lives are passed without continuing physiological
stress. Their social scheme must also present a fine adaptation to
their isolation in order to preserve the coherence of this remarkable
population of widely separated families.

ANAKTUVUK PASS 97

Family TURDIDAE: Thrushes, Solitaires, Bluebirds

Turdus migratorius migratorius Linnaeus

9 males May 15-Sept. 18 weight (14), 70-91, — —
average 79 g.
4 females May 24-Sept. 18 weight (6), 72-94, mee we

average 82 g.
5 young males June 21-Aug. 7 =e — ==
6 young females Aug. 7-Sept. 18 —_ — —

The earliest records for eastern robins are May 138, 1948, May 18,
1949, May 11, 1950, May 10, 1951, May 18, 1952, May 19, 1953, and
May 25, 1954. Robins appear to go directly upon arrival in spring
to nesting places which are mostly in rather large areas of fair-sized
willow brush, as around Contact Creek and Kangomavik. I have seen
nests at many places in the narrow valleys where a few high willows
grow and on a rock ledge at the mouth of Napaktualoitch Cave. On
the open tundra around Tuluak Lake they are not often seen.

Around Contact Creek in the spring of 1952, the first arriving rob-
ins appeared to act in pairs at once to establish nesting territory, and
females were taken within two days of the first male specimens in
1951. However, I saw groups of six robins and numerous single birds
on the upper Noatak River and at Howard Pass on May 25, 1952, so
that I believe that separation of the small migrant bands into pairs
occurs about at the time of arrival at the arctic nesting grounds.

The tabulation of social and family activities observed among robins,
below, presents a good chronicle of their summer activities because
in the north, as elsewhere, robins are of conspicuous and demonstra-
tive behavior:

May 10-18, 1948-52 First arrival Anaktuvuk

May 25, 1952

May 28, 1951
May 29, 1952
May 30, 1952
June 5, 1952

June 10, 1952

June 17, 1951

June 21, 1951
June 22, 1950

July 31, 1950
August
September
Sept. 12, 1950
Sept. 18, 1950

Groups of six and single
birds

Pair nesting

Nest building

All pairs settled

First egg

First set of 4 eggs com-
pleted

2 well-feathered nest-
lings

Young in first flight

Young male flying
weighed 63 g.

Young—76 and 79 g.

Family groups

Flocks frequent

Flock of 50

Last flock of 12

Upper Noatak River and
Howard Pass

Akvalutak

Contact Creek

Contact Creek

Contact Creek

Contact Creek

Napaktualoitch Cave

Contact Creek
Kangomavik

Kangomavik
Summit

Summit
Summit

98 U. S. NATIONAL MUSEUM BULLETIN 217

Robins have been observed nesting at Umiat on the Colville River
(Karplus, 1952). We have found them common in the Killik Valley
in summer, and I have seen them in migration in the upper Noatak
Valley and at Howard Pass. The conspicuous flocks seen about Sum-
mit in early September best indicate the large numbers of robins
which have summered in the arctic mountains and among the larger
willows of some of the more northern valleys. Because their range
of terrain is limited, the total arctic tundra population of robins is
locally conspicuous but not comparable with the numbers of the com-
monest birds.

While searching for the races of Dall’s sheep high among the moun-
tains along the Macmillan River in Yukon Territory, Sheldon (1911,
p. 127) remarked upon the numerous passing flocks of migrating
robins and noted that the last was seen on September 19.

In early summer, robins take poses nearly as conspicuously statu-
esque as are those of the golden plover. They sing cheerily, but to my
mind much more weakly in the northeastern States, where, however, I
heard them with younger ears. They appear vigorous and interested
in their arctic environment but unhurried by anxiety for the short-
ness of the summer and the remoteness of their winter quarters, con-
cerns which are in the usual thoughts of human strangers although not
apparent in normal arctic residents.

Combative against other robins intruding upon their nesting terri-
tory in the early part of the season they later become increasingly
noisy toward any potentially dangerous enemy to their young. Asthe
young grow larger, other pairs from adjacent regions join in these
disturbances with as much singleness of purpose as if nothing but the
commotion among robins were important. Their excited calling and
generally conspicuous parental] solicitude draws attention to the pres-
ence of the young birds, whose answering youthful calls precisely lo-
cate their position. Whether these parental demonstrations are, on
the whole, effective in protection of the young I cannot say. To me
they illustrate such an exaggeration of parental solicitude as would
attract a predator. Sometimes the social demonstrations of roebins
are joined by birds of other species, but a pair of robins can seriously
trouble a marauding shrike so that his only safe tactic is to seize a
nestling victim in a stealthy dash and hope to get out of range intact.
In judging the interaction of animals we must recall that feeble birds
sometimes pursue and obviously discomfit large predators, and that
unexpected occurrences often disturb large mammals to flight. A hu-
man hunter can be much alarmed by strange sounds heard in dark-
ness when there is insufficient sensory information available for rea-
soned calmness. Animals must often be mentally unstable in the dark-
ness of their unreasoning minds.

ANAKTUVUK PASS 99

The noisy exhibitions of the robins indicate how their early summer
concern for family affairs merges into a communal interest which
widens and strengthens in early September to join them in well
organized flocks ready for departure southward.

The aggressive, swift-flying robins, with their great concern for
their own pattern of social behavior, apparently find the arctic nest-
ing grounds very satisfactory. Perhaps it is because of this aggressive
and demonstrative concern for their own social pattern that we white
people find familiar sympathy for robins. The Nunamiut name for
robin is Koyapigaktoruk, described to me as referring to their “noise”
rather than to their song. Although the robins live about the Eskimo
camps I did not detect as much Nunamiut interest in them as they
have in many birds whose more gentle behavior better conforms to my
impression of Eskimo manners.

Hylocichla minima minima (Lafresnaye)

7 males May 29-June 14 weight (7), 26.1-33.6, = —=
average 30.6 g.
lfemale, 1 nest; June 22 weight (2), 27.8-35.5 — =
4 fresh eggs, g.
with female

A northern gray-cheeked thrush was brought to me May 30, 1948,
and I saw many along Contact Creek after May 29, 1952. The first
birds seen appeared to be settled, and Grinnell (1900) reported that
they were already paired when first seen on the Kobuk on May 24.
The Nunamiut know the bird well, calling it Niviokruksioyuk, mean-
ing “goes after flies.” It is one of the regular and common summer
inhabitants among the dense willows of the Valley.

The nest collected by Tom Brower was typically situated in the fork
of high willows about two feet off the ground. It was bulky, com-
pact, and rather well formed of flexed willow twigs and coarser
round grass stems. It was lined with fine grass to form a deep
depression.

They are not conspicuous birds after nesting begins in the moun-
tains, but males sometimes come to small willows in the open and to
the top of the willows in thick brush to sing. Nesting sites are among
dense willows. In 1951, I found many gray-cheeked thrushes in the
thick willows around Contact Creek. In 1952, they were more common
there than robins and we saw many more than in preceding years.
Karly in June that year, the male thrushes were often engaged in
driving away intruding males, which they did more decisively and
in swifter flight than even the robins. I rank these thrushes and
Alaska yellow wagtails ahead of robins for the sudden violence of
their reaction to preserve their domestic territory for their own ex-
clusive use.

100 U. S. NATIONAL MUSEUM BULLETIN 217

While many pairs were established in the dense willows along Con-
tact Creek by May 31 they were elusive in the thick brush and I have
only one explicit record of nest building on June 11 in addition to the
set of eggs which was collected on June 22. Nesting thrushes are
about as numerous as robins but nest somewhat later. At Umiat a
family of nestling thrushes left the nest on July 6, four days later
than a family of nestling robins (Karplus, 1952).

Oenanthe oenanthe oenanthe (Linnaeus)

14 males May 21-July 31 weight (15), 21.0- —- —
28.1, average 23.9
g.

5 females June 27-July 31 weight (6), 19.9- — —
26.3, average 23.5
g.

4 young males July 13-31 weight 29, 24, 24, 25 — —
g.

2young females July 22-29 weight 24.6, 24.2 g. — ; —

6 downy nestlings June 28, 1951 weight 6.4-7.9 zg. _—— —

These birds correspond to the specimens of European wheatear in
the U. S. National Museum. The earliest record is of the specimen
taken May 21, 1949, May 23, 1951, May 23, 19538, and June 1, 1954,
but in their mountainside haunts they cannot be closely observed.
They occupy in summer a narrow band of dry ground along the wall
of Anaktuvuk Valley starting just above the level of the Valley floor.
There they may sit quiet but alert on a short willow or rock and when
approached, rise nervously high into the air like the Alaska yellow
wagtail, but usually they fly more directly away and for a greater
distance. When one of a family group thus darts into the air, the
others are likely to follow. Their rapid, periodically swooping flight
along the Valley walls makes the Nunamiut name 7%kmiakpaurak,
“little eagle,” easily appreciated by one who has pursued them over
this steep, rough terrain.

Because of their location above the Valley floor, our information
upon their arrival is uncertain. We also failed to discover nests until
1951, although the Nunamiut said that they were there but well hidden
beneath rocks. Three males and three females taken June 25-29 had
bare brooding areas and behaved as if their nests were nearby. A male
watched on July 9, 1950, made two trips to the low willows along the
depression of a small creek. It perched for some minutes on the same
sharp stone before and after each trip. It returned one time carry-
ing a moth and appeared ready to watch me indefinitely before moving
again. Its earlier, although repeated visits to the willows gave me no
indication of a nest or young family. No signs of a nest could be
found, nor did the place look favorable.

On June 28, 1951, I saw a wheatear on the west side of the Valley
and about 800 feet above its floor. After about a half hour it went to

ANAKTUVUK PASS 101

a small opening under a flat rock, where it was joined by its mate
and the two made repeated visits with food, apparently insects, during
the next hour. The birds were extremely alert and nervous. Lifting
aside the covering rock I found a nice nest of fine grass lined with
ptarmigan feathers and including 6 very small young weighing from
6 to 8 grams each. They must have been newly hatched. I estimate
that the last egg of the clutch had been laid about 14 days earlier.
The first egg would then have been laid 20 days earlier, or about
June 8.

Young birds were flying well and had reached adult weight by
mid-July, but through the rest of that month they remained asso-
ciated in family groups as well organized as those of robins.

The fat of wheatears in late summer is white and fluid, similar in
consistency but not in color to the yellow fat observed in the insectiv-
orous Yukon phoebe of similar haunts. I suspect both birds of de-
riving their fat from a diet of insects and that the different pigmenta-
tion might offer a clue as to its origin.

Late summer observations were not regular, but two wheatears
came into camp on the Valley floor at Imaiginik, August 15, 1950.
Like the Alaska yellow wagtails in late summer, their shyness was
gone, and they had discarded the temperamental behavior observed
during the period of family life.

In their regular summer haunts wheatears are associated with oc-
casional Alaskan longspurs, pallid horned larks, and American pipits.
Along the narrow bank below the talus slopes, wheatears are numerous
but the area occupied is small and their population is less than that
of pallid horned larks. Since the extent of their habitat much exceeds
that occupied by robins, they are probably the most numerous
of the thrushes.

Luscinia svecica svecica (Linnaeus)

I have been on the lookout for the red-spotted bluethroat but have
failed to find one about Anaktuvuk. In June 1953 I obtained a female
near Itivlik Lake, about 130 miles west of Anaktuvuk (L. Irving and
Paneak, 1954). The conspicuous foxy color of the tail and the wren-
like attitude of the bird as I watched it convinced me that I had not
previously seen one like it. When Simon Paneak saw it he was sure
that he had not seen one like it nor had he heard it described among
the Nunamiut until I had made known my interest in finding one.
When my colleague Robert Rausch saw the specimen he was sure
that he had not seen one during his extensive travels in the mountains
and on the arctic slope.

The bluethroat has been found near Wales, Wainwright, and Barrow
(Bailey, 1948). With the recent simplification of transportation to

102 U. S. NATIONAL MUSEUM BULLETIN 217

the arctic interior, several ornithologists have observed it during brief
visits to Umiat. In 1952 Cade and Schaller recorded bluethroats on
thirteen occasions while traveling along the Colville River (Kessel,
Cade, and Schaller, 1953).

I am puzzled by the fact that I could find no acquaintance with a
bluethroat among the Nunamiut, for had it been seen along the Col-
ville, Simon Paneak is sure that he would have heard remarks about
its distinctive appearance. To be sure, the regular summer experi-
ence of Nunamiut along the Colville is now more than 20 years past,
but for the other birds of that region their recollections remain clear.

The recent observations of scientists have not sufficient sequence to
warrant a suggestion that this migrant from Asia is significantly
changing its range or abundance in arctic Alaska. It may be that the
bluethroat is a late arrival in spring and that it occupies restricted
habitats in certain localities.

Family SYLVIIDAE: Old World Warblers, Gnatcatchers, Kinglets

Phylloscopus borealis kennicotti (Baird)

5 males June 19-June 24 weight (7), 8.8-10.2, —_— =
average 9.4 g.

3 females June 19-July 23 weight (4), 8.5-9.9, — ——
average 9.1 g.

1 sex unknown June 24 weight 8.1 g. =— a

2 young males Aug. 14 weight 9.7, 10.2 g. = re

3 young females July 23-Aug. 14 weight 9.3, 10.7, 9.8 == east

g.

I had hoped to see one of these birds because of the reference by
Baird (1869) of Dall and Bannister’s (1869) specimen from Nulato
to the name of Kennicott, the brilliant leader of the early scientific
exploration of interior Alaska. Frequently I had discussed their
appearance with Simon Paneak, and he had tentatively connected them
with birds which he had known asa boy in the Killik Valley and which
were named Songakpalutunygik, meaning “small bird the color of
bile.”

In August 1950, we were delayed for several days at Odrivik Lake
in the Killik Valley by malfunction in the lubrication of the airplane
in which John Cross had come to take us out. While I was hunting
among the willows covering the sand dunes along the river, I saw
several very small warblers actively feeding among the willow tops,
apparently taking small insects. I failed to get them in view of my
glasses, and they disappeared. As they were moving southward along
the river when seen, I ran to make intercepts of their apparent course.
At the third try and near a mile to the south I picked up a band of
five, from which I collected 3 specimens. Simon recognized them as
the Songakpalutunygik which he had known as a boy.

ANAKTUVUK PASS 103

On June 19, 1951, among the willows on Contact Creek, I saw two
which acted as if in residence, watched them, and heard the male
singing, although my unfortunate insensitivity to bird songs prevents
me from definition of their faint but agreeable sounds. In the next
4 days I hunted daily along Contact Creek and formed the estimate
that about 20 pairs were resident along one mile of the thick high
willows along the stream.

On June 23, John Morry’s son, Riley, brought me a willow warbler
with its nest and two eggs. With his slingshot, the boy had wounded
the bird, and had found its nest on the ground with one egg. As boys
and men will often do with animals that they have just failed to
kill, he compassionately brought bird and nest back to his parents’
tent. There the bird had laid one additional egg before expiring,
and in the morning Riley brought me the nest with its two eggs and
the parent bird.

On June 24, 1951, as we returned northward to our camp at Imai-
ginik, Simon Paneak and I found two pairs of these little warblers
among the willows of Kangomavik Creek, and I suspect there were
other pairs there. In 1950 we had hunted this area carefully on several
days wthout seeing willow warblers.

When I reached Contact Creek on May 29, 1952, I heard reports
from Simon Paneak that a few willow warblers had already been seen.
In the next 10 days, I saw two birds like willow warblers, and there
were several observations reported by Eskimo adults and children.
Unless one can see a willow warbler stationary and from nearby it is
difficult to identify, because of its insignificant size and faint mark-
ings. When it is moving among the wind-shaken willow leaves and
viewed against a light sky background its color and even shape are
hard to distinguish. Several instances of singing willow warblers were
reported to me, but in that year I saw none I could identify. By this
time the Nunamiut shared my interest in these little birds and they
gave me several reports indicating that willow warblers were not
uncommon. But it was not until July 23 that Paneak collected an
adult female and two young females of adult weight at Kangomavik.

On occasions the willow warblers have been easy for me to see clearly
as they sang or moved about in the edge of the still willows. I have
had so many views of them and received so many reports that I am
sure there is a considerable population of them usually nesting among
the willows along Contact Creek and at Kangomavik. The earliest
date when one has been reported to me is May 30, 1954.

Simon Paneak and I saw many willow warblers among the willows
around Lake Itivlik and along the Ahlasuruk River. A female that
we collected had a bare area as for incubation (L. Irving and Paneak,
1954).

469496—60——_8

104 U. S. NATIONAL MUSEUM BULLETIN 217

The Nunamiut identification of willow warblers with a name re-
called from their childhood is a good indication that the birds are
historic members of the avifauna. That they had not recently seen
them until I obtained a specimen in 1950 can be explained by the fact
that the people with whom I most associated have resided north of the
thick willows since 1936, whereas from 1920 to 1936 they had lived
principally on the arctic coast.

I believe that my fragments of evidence combine to indicate the
regular arrival of willow warblers at certain dense willow patches in
the valleys at the end of May, where they promptly establish nesting
areas and lay eggs in mid or late June. The young birds appear to
be near adult size at the end of July and by mid-August they are in
migratory movement.

It is quite understandable to me that the records of willow warblers
are few and variable, and that no nesting in arctic Alaska has hitherto
been confirmed in reports. From the vague literature on the subject,
doubts of their firm establishment in Alaska were reasonably ex-
pressed (Bent, 1949). But my observations give no evidence that the
population of willow warblers in Anaktuvuk is small, unstable, or
irregular, and I believe that their existence is historic because of their
designation with a specific Nunamiut name by Eskimos whom I have
found to be singularly precise in the recognition and nomenclature of
birds (L. Irving, 1953).

Regulus calendula calendula (Linnaeus)

I have had several reports of ruby-crowned kinglets at Anaktuvuk
and, surprisingly, they point to its winter residence there. In the
spring of 1948 Jesse Ahgook brought me an example with the bright
red crown feathers of a male, reporting that he had collected it in
early winter at Hunt Fork. The bird had deteriorated beyond possi-
bility of saving. Simon Paneak said that the kinglet was familiar to
them, that a few were usually seen in the northern part of the forest,
and that some remained in winter. He thought that kinglets were
occasionally seen north of the timber in winter.

We discussed kinglets frequently and several people thought that
they had seen them among the willows north of timber in winter,
but realizing the critical evidence required they did not at first wish
to present their opinions as records. John Krog and I looked for
them in winter and summer without success at Anaktuvuk. We
obtained one specimen, identified as Regulus calendula calendula
(Linnaeus) at Bettles on August 25, 1951.

Simon Paneak reported that he saw a kinglet among the thick
willows at Nakrak just after Christmas 1952. He watched it from
within a few feet during several minutes, but it was too close to

ANAKTUVUK PASS 105

shoot. A few days later his wife, Susie Paneak, watched several in
the same locality and heard their faint calls. In early March 1954,
while we were in camp at Kivik Creek on Hunt Fork, Simon described
three kinglets which he had recently seen feeding among the alders
in the company of a large flock of redpolls.

Bailey (1948) reports that Brower obtained a specimen from Cape
Halkett in September and that one was collected near Barrow on
April 30. Anderson obtained one found dead aboard ship off Cape
Halkett on September 24. On the arctic coast winter conditions are in
effect on these dates.

Our evidence warrants inclusion of the ruby crowned kinglet among
the birds of Anaktuvuk. But its recorded winter range is southward
from Nebraska to West Virginia. Without a winter specimen I do
not now feel warranted in saying that it is a winter bird above the
arctic circle in Alaska. Until further evidence is at hand I list the
kinglet asa visitor at Anaktuvuk.

Family MOTACILLIDAE: Wagtails and Pipits

Motacilla flava tschutschensis Gmelin

12 males June 4-Aug. 3 weight (10), 15.1- -—— _—
17.7, average 16.3

g.
9 females June 6-Aug. 3 weight (9), 14.0- =—— -_
17.6, average 16.1

; g.
4nests, 5fresh eggs June 16, 17, 19, 20 — -— —

My first specimen of Alaska yellow wagtail was collected at Umiat
on the Colville River June 4, 1949, while the ground was still covered
with deep winter snow. This bird flew violently to expel another
wagtail and so seemed definitely located and committed to the protec-
tion of his own territory. At this date in Tuluak valley, 100 miles
south, the winter snow has disappeared from the open tundra and
the season is near a week ahead of Umiat. Wagtails were not re-
ported at Anaktuvuk until June 6, 1949, June 4, 1951, June 7, 1953,
and June 5, 1954. Bailey (1948) reports the collection of two males
near Wainwright May 30, 1939, and June 17, 1941. These dates and
my record at Umiat suggest that the wagtails when coming from Asia
migrate eastward in Alaska through or north of the mountains. At
St. Michaels they were reported by Turner (1886) to arrive about
June 12 and to proceed at once to nest.

Nests were found with complete sets of fresh eggs from June 16 to
20. Even for the energetic wagtails to prepare their nests complete
with eggs in 10 days after arrival seems rather rapid progress and it
is possible that they arrive earlier than we have recorded.

106 U. 8. NATIONAL MUSEUM BULLETIN 217

During early July on the valley bottom many willow patches a
few hundred yards across contained a wagtail. The males perch
briefly on the tops of high willows and while observed even at a dis-
tance beyond 50 yards, every half minute or so may rise rapidly high
into the air, darting about in quite unpredictable directions, often
to settle soon with equal suddenness near the original perch. It did
not appear that these maneuvers were executed in alarm or in pursuit
of insects, but as if they were expressions of the restlessness of the
birds. I could see no capture of insects, but it may be that my exasper-
ation in trying to follow these restless birds caused me to mistake some
gainful activity for behavior which I attributed to their nervous
disposition.

Wagtails are common and well known to the Nunamiut by the
name Piorgak, referring to their call. They are more numerous in
summer than Savannah sparrows and occupy the upper part of the
same habitats in the fair-sized and more or less dispersed willows.

The nests are commonly under willows on a niggerhead, covered
and difficult to find. Usually they are near wet ground. They are
constructed loosely of grass, willow roots and bark, sometimes with
considerable quantities of moss, and lined with fine grass, caribou hair
and a few feathers.

At the end of July near Imaiginik, wagtails were evidently free
from nesting cares for they had ceased their aerial demonstrations
and started wandering calmly among the willows along the sandy
river bars. In the Killik Valley by Odrivik Lake after August 3,
1950, in the mornings after the brief shadow of the first late summer
arctic nights, numbers of wagtails were feeding among the willows.
At this season they somewhat resembled the pipits in coloration, but
they kept more in the branches of willows than do the ground-feeding
pipits. During the next ten days the numbers were far greater than
were seen in July and the trend of the groups was distinctly southward
toward the mountain line about five miles away. This movement was
evidently the early withdrawal of great numbers coming from the
valley of the Colville. Judging from the habitats in which I have seen
them, they might prefer routes north of the forests such as could be
found in the willow-filled valleys in the mountains west of the head
of the Killik.

In the wooded part of the Kobuk River valley Grinnell (1900) did
not report wagtails, although he found them abundant and nesting in
the treeless areas about the Kobuk delta. We found them numerous
along the Ahlasuruk, 130 miles west of Anaktuvuk and about the same
distance north of Grinnell’s winter camp. (L. Irving and Paneak,
1954). In Yukon they had not been reported (Rand, 1946), but
Munro informed me that he had seen them near the mouth of the Firth

ANAKTUVUK PASS 107

River. Anderson (1921) found nestlings near Niklik at the mouth of
the Colville. Only a few records from the northern arctic coast are
given by Bailey (1948). From information provided by my col-
league Raymond Hock, they are common in the Colville Valley near
Umiat. It seems that data are near at hand to establish accurately
the distribution and abundance of these small birds which migrate
far eastward from the interior of Asia to energetically occupy in
summer the treeless section of arctic Alaska.

Anthus spinoletta rubescens (Tunstall)

26 males May 4-Aug. 7 weight (28), 19.2- — —-
25.5, average 21.6 g.
7 females May 11-July 7 weight (8), 18.6- —_ -——

23.2, average 20.1 g.

The earliest recorded arrivals of American water pipits were May
10, 1949, May 10, 1950, May 4, 1951, May 14, 1952, May 13, 19538 and
May 25, 1954. In 1949 the peak of migration was reached on May
19 when 500 were reported seen by Tom Brower. Thereafter the num-
bers dropped off sharply, but on May 30, 40 pipits were recorded.
Since this number is larger than the number of residents seen on later
days, it probably represents the late northward flight in that year. On
May 19, 1951, the pipits were reported by John Krog to have suddenly
disappeared from Nakagnik Springs where they had previously been
numerous.

Males collected May 4 and 10, 1951, had small testes, but all ex-
amined later were large. Eggs in females examined in 1951 were 3
mm. in length on May 21, 2 mm. on May 23, and 2 mm. on May 26.
On June 2 some females under observation had laid and one large egg
was found broken; on June 19 all had laid.

One nest, found by Tom Brower on the ground under some low wil-
lows upon which dead grass was hanging, was large enough to accom-
modate the 6 eggs it contained (these appeared large for a bird of
this size). It was loosely constructed of flat grass, lined with fine
round grass, and contained no other material but grass.

Pipits are uncommon on the floor of Anaktuvuk Valley in summer,
but in some high wet or dry grassy places and on the dry and rocky
ground as high as 4,000 feet, the limit of our regular surveys, they
are numerous, although inconspicuous. They ranged higher than the
wheatears, which favor the foot of the talus slopes, but a common
redpoll (Acanthis flammea) and the pipits were the highest ranging
small birds to be collected.

In midsummer, these high mountain regions are extremely dry, bear-
ing only a sparse low vegetation, and the pipits are inconspicuous on
the ground among the rocks and grasses, but they frequently stand ex-
posed on hummocks or rocks, revealing themselves only by twitching

108 U. S. NATIONAL MUSEUM BULLETIN 217

their tails. Like wagtails they may suddenly take wing for no ob-
vious alarm, fly high in the air, swooping about unpredictably, and
sometimes return close to their start, or having gained 100 or 200 feet
altitude they may fly directly out of sight.

In August, they begin to move more deliberately along and near the
ground on the Valley floor. In the Killik Valley in early August,
they were generally moving southward in considerable numbers, but
they were fewer than the wagtails. Some were seen at Chandler
Lake, August 16, 1950, and the last recorded was seen at Contact
Creek, September 2, 1950.

Many pipits remain for nesting in the Valley but as many more were
seen during the spring and fall migration. A large population tray-
erses the Pass toward the arctic slope.

The Nunamiut name Piorgavik refers “to the song.” ‘The name re-
sembles the designation Piorgak given to the Alaska yellow wagtail.

Family BOMBYCILLIDAE: Waxwings

Bombycilla garrula pallidiceps Reichenow

Robert Rausch and Homer Mekeana saw a flock of Bohemian wax-
wings near the northern limit of spruce on the Savioyuk River, Oc-
tober 15,1950. In August 1951 John Krog and I saw a flock of some
50 feeding among the bushes and tree tops along the Alatna River
north of the stream from Iniakuk Lake and just about at the southern
mountain line. I have obtained several specimens from Kobuk,
where the Eskimos name them Shooloktachailak, meaning “wearing
feathers like an Indian.”

There is no evidence that they pass to the open country north of
the forests, but in their wandering occupation of the wooded interior
of Alaska they occasionally reach its northern border. They are not
included in the avifauna of Anaktuvuk.

Family LANIIDAE: Shrikes

Lanius excubitor invictus Grinnell

4 males, adult Apr. 29-Sept. 5_ weight (8), 72, 73, —_— : ——
plumage g.

3 males, Ist year Aug. 6,7 — ad CTLET
plumage

2females, adult May 8, June 24 weight 71, 64, g. == sts
plumage

3 females, Ist year May 26-Aug. 7 weight (1), 69.5 g. — ee
plumage ‘

3 males, in first June 24, July 31 weight 68, 67, 67 g. — —
summer !

1,sexunknown, September f — — —
1st year

plumage

ANAKTUVUK PASS 109

The earliest observations of Northern shrikes were May 8, 1949 at
Tuluak Lake, May 9, 1950 at Contact Creek, April 29, 1951, at Itik-
malikpuk Creek, and May 9, 1952 at Contact Creek. The latest record
was September 28, 1950, just within the limits of the spruce forest
at the head of the Savioyuk River. Shrikes were observed almost
daily during early September as the Nunamiut moved from Tuluak
southward past Kangomavik to Kalutak Creek.

The Nunamiut name is Jrirgik, “eye extractor,” indicating the dis-
agreeable activity attributed to shrikes of picking out the eyes of
small birds and mice, after which, according to the story, the shrike
watches its blinded victim blunder about until ready to eat it. The
name and story are identical with the characterization of the shrike
as given by Eskimo people on the lower Kobuk River (Grinnell, 1900),
and as given me by Charles Sheldon of Kobuk.

A nest with one egg and a dead shrike was brought to me on June 1,
1948, but unfortunately it could not be preserved. On June 11, 1951,
I found an empty nest 9 feet up in a willow where Itikmalikpuk Creek
emerges from the mountains. It looked like a new shrike’s nest and a
shrike was seen nearby. One young bird, clearly hatched that season,
was near adult weight and although its tail was still short, it was
flying with other young among the willows at Kangomavik July 31,
1950. Two young males on June 24, 1951 were as heavy as the young
male just mentioned. Another bird judged from its plumage to be
of this summer was taken near Contact Creek in early September
1950, but the exact date and sex were not recorded.

Shrikes were observed commonly in 1950 and 1951 in the Killik
Valley near Odrivik Lake, along the river 4 miles north of Akmalik
Creek, and at least three family groups of three to five were observed
in 1950 around Akmalik Creek. Mice and small birds hung by them
er by short-eared owls in the bushes were occasionally found. The
plentiful numbers of shrikes in the mountains was a surprise to me,
but none were observed away from the rather large willows about 15
feet in height.

It appears that the summer produces a large increase in the popu-
lation of shrikes which feed upon small birds like the redpoll and
upon an occasional mouse. No evidence for migration further north
can be presented, but since the favored habitat of high willows along
the streams and the abundance of mice and redpolls are repeated
through the Colville Valley, and through the upper parts of the
valleys of northward flowing rivers like the Meade and Ikpikpuk it
is assumed that a number of shrikes move through the Pass to nest
on the arctic slope. Judging from occasional reports we will find
that some shrikes winter in arctic Alaska.

110 U. S. NATIONAL MUSEUM BULLETIN 217

Family PARULIDAE: Wood Warblers

Dendroica petechia amnicola Batchelder

2 males May 31, 1954 weight 9.4, 8.9 g. -_— —

These are the first records of yellow warblers obtained by Simon
Paneak, who collected them in the mountains. It may be that some
others have been mistaken for Kennicott’s willow warblers. Several
times warblers which appeared too yellow for willow warblers have
flown swiftly past me. On the other hand I have observed several
hundred willow warblers until sure of their identification, and I am
therefore certain that other yellowish or greenish warblers are uncom-
mon. At present the information only suggests that the two yellow
warblers were visitors.

Dendroica corenata hooveri McGregor

1 in male May 26, 1953 weight 13.8 g. —_— ———
plumage
The specimen listed above is the only record of the myrtle warbler
from Anaktuvuk. Since the male birds are conspicuous as they sing
and feed in the tops of poplar trees in the wooded portions of Alaska,
I believe that they would be noticed if they were often present at
Anaktuvuk. I consider this example to be a visitor.

Wilsonia pusilla pileolata (Pallas)

2 males June 3, 7 weight 7.2, 6.9 g. = —

These pileolated Wilson’s warblers are the lightest adult birds that
I have examined in the Arctic. The first one, with a rapidity em-
phasized by its small size, flew to the top of a willow to sing, revealing
to me its clear yellow underparts and black crown. For 15 minutes
I watched its restless behavior and listened to its singing before it
withdrew into the thick low brush. I was sure of its attitude as
proprietor of the area. The next morning, after I had waited for
about an hour, it returned at about three o’clock to sing in the same
willow. In 1954 Simon Paneak obtained another example.

From time to time in earlier years I had reports of small yellow
birds in the valley and I saw a warbler which I thought was brighter
yellow than Kennicott’s willow warbler or the yellow warbler among
the willows along Contact Creek. Although the Nunamiut did not
know the pileolated warbler explicitly I suspect that some of these
elusive little birds will be found nesting in the mountains.

After we had watched through the night for wolves to return to
their den, at about 3 a. m. I saw a male pileolated warbler sing twice
on a willow near the Ahlasuruk River about 130 miles west of Anak-

ANAKTUVUK PASS Li

tuvuk (L. Irving and Paneak, 1954). Cade found them at Umiat
(Kessel and Cade, 1958).

I consider that the indications warrant designating the pileolated
warblers as birds nesting at Anaktuvuk.

Family ICTERIDAE: Blackbirds

Euphagus carolinus carolinus (Miiller)

6 males May 15-Aug. 31 weight (4), 60-66, — =
average 63 g.

4 females May 17-July 8 weight (3), 48.0-60.0, == ==
average 63 g.

1 young male July 8 weight 55.4 g. — ——

1 young female July 8 weight 50.4 g. —— =e

1 young male Aug. 31 weight 55.5 g. —— —

Individual rusty blackbirds have been reported May 15, 1949, May
16, 1950, May 22, 1951, and May 18, 1952, and a few have been seen
each year in each summer month. Mating and nesting or summer
resident blackbirds were not remarked until 1951, and if they nested
commonly their revealing habits would scarcely have allowed them
to escape observation. But in 1951, Simon Paneak and William
Irving collected an adult pair and a well-grown young male and
female at Amorgoayat in the Killik Valley on July 8. I had formerly
thought of them as visitors from the Koyukuk and Alatna Valleys,
where I have frequently seen numerous family groups, but since a
family group has been seen in the Killik Valley, I consider it certain
that an occasional pair nests in the mountain valleys in addition to
numerous visitors which come in from the wooded valleys to the
south.

The Nunamiut name is Tolungiksyaurak, meaning “little raven.”

Family FRINGILLIDAE: Grosbeaks, Finches, Sparrows, Buntings

Pinicola enucleator alascensis Ridgway

1 male Nov. 11, 1948 _ — —
2 females Jan. 31-Nov. 11 == == ==

I have seen pine grosbeaks frequently in the timbered regions of
the Koyukuk and Alatna Valleys, and I have numerous records and
specimens from the northernmost spruce forest of the John and Savio-
yuk Rivers. They are also seen occasionally in winter among the
willows of the valleys as far north as the northern mountain line.
According to Simon Paneak none have been seen further north. The
Nunamiut name is Kayatavak.

I have only one observation recorded in summer north of the tim-
ber; Jesse Ahgook reported a pair to be nesting among the willows
of Contact Creek, June 7, 1951. His observations are usually ac-

112 U. S. NATIONAL MUSEUM BULLETIN 217

curate, but later we could not find the nest or birds. Jesse and Simon
Paneak agreed that since this was the only report of one north of
timber in summer that they could recall, it did not establish normality
of summer occurrence.

Small groups of pine grosbeaks appear occasionally to move about
in winter over the open tundra north of the spruce forests as they
do in the wooded regions. In winter the northern willows blown
bare of snow may afford easier feeding than the snow covered
branches of the windless forests. The grosbeaks’ feathers are the fluf-
fiest of any of the Fringillidae, and they and the redpolls (Acanthis
hornemanni exilipes) are the only members of that family which regu-
larly winter in the mountains. Their winter travel northward is a
frequent occurrence, and it seems correct to designate the pine gros-
beaks as winter visitors which at that season move from the forests
100 miles or so north of tree line. I also suggest their inclusion with
the Yukon chickadee, downy woodpecker, and Alaska jay in a prob-
ably more or less regular northward movement in winter from the
forest to the tundra.

Leucosticte tephrocotis irvingi Feinstein

7 males May 22-June 29 weight (8), 26.8, 29.1, — —_—
average 27.7 g.

5 females June 7-29 weight (5), 25.8, 30.3, — —_
average 28.6 g.

Gray-crowned rosy finches were found along the west side of the
valley at Tuluak as it slopes near and above the talus slopes 4 to 500
feet above the valley floor. They are well known and their rosy colors
bring them the name kaviksruak among the Nunamiut. The
earliest reports in spring are May 19, 1950, June 2, 1951, and June 1,
1954. The fall records show old birds with young of the year moving
south on August 29, 1950, at Kangomavik, and the last were seen
September 2, 1950, at Contact Creek, also moving southward.

No nests were found, but a male examined by Robert Rausch on
May 22, 1950, had testes measuring 10 mm. and was in obvious breeding
condition. Three male specimens taken on the mountain side between
June 18 and June 29 had bare brooding patches. The behavior of the
male and female observed by Thomas Brower on June 23, 1949, showed
the alert and baffling indirection of a bird near its nest. Its efforts for
diversion were successful, and no nest was found. John Krog observed
several pairs about 1,400 feet above the floor of Anaktuvuk Valley
several miles west of Tuluak and reported their behavior like that of
nesting birds on June 2, 1951. The Nunamiut believe that they nest
on the mountain sides fal from the evidence there is no doubt that
nests are made and young birds are reared pitas the Valley’s moun-
tain side, probably near the talus slope.

ANAKTUVUK PASS 113

It is unlikely that these finches move much further north, for beyond
the mountain line there are only a few isolated elevations with the type
of mountainside terrain to which they are usually restricted.

After repeated examinations these rosy finches appeared to differ
from L. t. tephrocotis of more southern origin in being more exten-
sively tinged with rosy color on upper and lower posterior parts of
the body. The brown of the Anaktuvuk specimens is more red. Ber-
nard Feinstein (1958), who has described them as a new race, remarked
a more distinctive difference from tephrocotis in the gray pattern
about the head, which approaches the pattern of /éttoralis. We con-
sider the Anaktuvuk birds to be distinguished as a separate race.

Brina Kessel kindly sent three specimens of rosy finches obtained
in the mountains near the head of Sheenjek River in the eastern part
of the Brooks Range, in Yukon Territory. These, a specimen from
Bettles, and two from the Cheena River also conform with our Anak-
tuvuk birds. A few specimens from Eagle are the northernmost
examples of tephrocotis that we could find.

Acanthis hornemanni exilipes (Coues)

12 males Feb. 14-July 19 weight (30), 10.7= —— a
16.1, average 12.7
g.
8 females Mar. 4-June 26 weight (24), 10.4- -—— =
14.8, average 12.8
g.
2 young males July 19, 24 weight 12.5, 12.2 g. === Ee
2 nests, each with June 16, 17 —— arr ao
5 fresh eggs and
female
2 nests, each with — —_ == ——
4 fresh eggs, no

female speci-
men, and there-
fore uncertain
as to whether
flammea or ezi-
lipes

Ira Gabrielson kindly reviewed my estimate of the redpolls col-
lected up to 1950 and Herbert Friedmann reviewed my distinction
of the whole collection of redpolls.

A few hoary redpolls are resident at Anaktuvuk in winter. In
addition to four specimens taken in February and March, four mum-
mified birds taken in February and March were identified by com-
parison in the U. S. National Museum. I have seen redpolls in late
winter at Umiat under the alders, consuming seeds on the snow where
they had fallen from the cones. They also feed on the bushes, usually
moving rather rapidly and coherently as a scattered flock, the indi-
viduals acting nervously and the flock often flying together. I have
not obtained a winter specimen of flammea. In winter plumage

114 U. 8S. NATIONAL MUSEUM BULLETIN 217

the two redpolls are more easily distinguished and I have then been
certain in observations of eailipes, but I have not identified any
flammea on the arctic tundra in winter.

During his winter on the forested part of the Kobuk, Grinnell (1900)
found redpolls the only birds usually to be seen. His specimens of
flammea were usually taken from flocks of exilipes. In his collection
there were 104 specimens of exilipes and only 7 of flammea (p. 48).
In his checklist (p. 77) Grinnell remarked that fammea was “common,
chiefly along the coast,” a comment which, I believe must result from
his observations around Kotzebue Sound in summer.

In mid-May 1949 small flocks began to appear near Tuluak and by
May 17 over 100 redpolls were reported daily by Thomas Brower.
The numbers then diminished and not more than six were seen
together after the end of May. Groups of five or six were occa-
sionally seen moving together after the others had begun nesting.

A nest was reported building June 8, 1949, and fresh eggs were
collected between June 16 and 20. With the first two nests were female
birds identified as eawilipes. Females were also collected and identified
from two nests in which the eggs on June 13 were too far incubated to
save. ‘The others were named by sight, so that we have identification
for only four individual parents of exilépes found nesting. Nests with
4 eggs have been found on June 4, but incomplete clutches are common
until June 15. Nestling birds were found June 25, and on July 4
a family had left the nest. Young birds were flying July 19. The
nests were bulky constructions low in the willows and made of small
roots, twigs, grass, occasional moss, and lined principally with caribou
hair and a number of ptarmigan feathers.

The following comments apply to exilipes, but I cannot determine
whether they apply to fammea as well. The Nunamiut said that red-
polls frequently have a second brood so late in the season that the young
birds are abandoned, starve and freeze. It was thought that parental
interest for their young was replaced in late summer by the desire
for flocking together and migration, so that the young were first aban-
doned and then froze after starving. On June 26 I have seen male
exilipes with testes of breeding size. At the same date I found a fresh
set of eggs of fammea which would probably have had time for normal
development of the young before cold weather. But a nest was then
near completion by redpolls which I called eazlipes, and I question
whether birds could mature from that set of eggs before cold weather
or the formation of flocks distracted the parents from family to
flock activities.

On July 27, 1950, family groups were joining and numerous flocks
of 8 or 10 birds were seen. Thereafter at Anaktuvuk and from Au-
gust 3 to 15 in the Killik Valley loosely associated flocks were building

ANAKTUVUK PASS 115

up in the willows. Some apparently came from farther north and
generally they were heading slowly southward as they fed. By Sep-
tember large flocks in fairly steady movement were working their
way along the sides of the valley and its floor near Contact Creek.
By the end of September the main migration had apparently passed,
carrying great numbers of summer residents and migrants from the
north toward the south.

In summer, resident hoary redpolls are the most numerous birds
near the willows. Because they spread over more territory they prob-
ably outnumber the tree sparrows. Redpolls were more often seen in
higher branches in the willows than tree sparrows, and they also dwelt
as high in the Anaktuvuk Valley as the willows grew up along the
narrow canyons of the tributary creeks to at least 1,000 feet above the
Valley floor. Many redpolls and snow buntings were seen by Simon
Paneak on August 29, 1950, above the head of Kangomavik Creek,
an estimated 3,000 feet above the valley floor. He could not be sure
of the kind, but they were apparently migrating.

In Nunamiut they are called Suksangik, a name which has no mean-
ing except redpoll. It does not distinguish exilipes from flammea.

Acanthis flammea flammea (Linnaeus)

17 males May 15-July 24 weight (21), 10.2- —_ —
15.0, average 12.9

g.

6 females May 19-July 7 weight (9), 10.1-14.2, — —
average 13.1 g.

1 young female Aug. 3 weight 12.5 g. — —

In view of the difficulty of making accurate distinction in the field
while the birds are in summer plumage, I cannot say to what extent
common and hoary redpolls mingle.

None of flammea have been identified in the mountains earlier than
May 15, and I do not think that they remain there during winter.

In Anaktuvuk Valley, I have found more of flammea near the dense
willows along Contact Creek than elsewhere, but I have specimens of
each redpoll from every region in which I frequently hunted. A
single flammea was collected among barren rocks at about 4,000 feet,
the highest elevation from which a redpoll was obtained, but I cannot
separate nor generalize upon the elevation of their ranges. It is my
impression that fammea is more often in the denser and larger willow
patches.

On June 27, 1951, a nest with five eggs was found about 4 feet above
ground in one of the small willows along the river near Tuluak Lake,
situated about 50 feet from a nest of ewilipes which I had found on
June 10. I could not be certain of the identity of the female, but a
male well colored with rose red occasionally came to feed her as she
sat on the nest. This bird was collected and identified in the U. S.

116 U. S. NATIONAL MUSEUM BULLETIN 217

National Museum as fammea. On June 11, 1952, I identified by sight
as flammea birds which were working on two unfinished nests at Con-
tact Creek, A third female allowed me to watch her from a distance
of 2 feet while sitting on a single egg. So I believe that both species
nest numerously in adjacent areas upon which I cannot now place any
distinction.

Until 1952 I believed that exilipes were ten times more numerous
than flammea. But I saw about equal numbers of each species as I
hunted principally around Contact Creek in 1952, where I had not
previously concentrated so much attention. I believe, however, that
over the whole valley the numbers of exilipes in summer greatly ex-
ceed those of flammea, and that they are distributed in more varied
situations.

In early and late summer, the migrating flocks of hoary redpolls are
very much more numerous, and I have not positively identified flocks
of flammea nor do I know whether they intermingle in migration with
flocks of ewilipes.

In weight the two forms are not distinguishable. In reports of
fatness prior to nesting both redpolls in early summer are noticeably
less fat than many common migrant species at Anaktuvuk. This ob-
servation can be compared with Barbara Oakeson’s (1953) view that
races of white-crowned sparrows of the Pacific Coast region making
short migratory flights are not as fat on their wintering grounds be-
fore migration as are those sparrows contemplating a long migration.
Furthermore, at the terminus of only a short migration, the Gambel’s
sparrows were not fat (Blanchard, 1942). Many redpolls remain dur-
ing winter in Alaska, but I have seen more of fammea and but few of
exilipes near Fairbanks and Anchorage.

The arrival of the redpolls is vaguely defined in my records, but the
rarity of fat redpolls at migration time presents a contrast with the
uniform fatness of arriving tree sparrows and Alaska longspurs, and
in fact with the fatness of many arriving migrant birds which are
known to winter in regions remote from Anaktuvuk. It may be that
the meager fatness of early summer redpolls at Anaktuvuk is a symp-
tom of a short migration, as Blanchard (1942) showed to be the case
among the white-crowned sparrows of the Pacific-coast States.

Of the two species of redpoll, eadlipes was found to be less fat than
flammea, as the tabulation below shows:

Very
Fat Medium fat Little fat little fat Lean

Acanthis hornemanni exilipes

Males 1 3 4 7 2

Females 2 2 2 1
Acanthis flammea flammea

Males 2 2 7 6 0

Females 1 4 2 0 0

ANAKTUVUK PASS 117

On this basis alone, I would not suggest that ewilipes make a shorter
migration to Anaktuvuk than flammea, but a difference in fatness of
the two birds reflects a difference in nutritional state which could well
be significant of different migratory habits. It is among such dif-
ferences in the physiology and habits of birds that we must look to
find expression of the influences which combine those of like race in
the patterns of similar flight activity, and which enable them to co-
here in their migratory route.

Loxia leucoptera leucoptera (Gmelin)

1 male July 10, 1954 weight 21 g. — —

Five white-winged crossbills were collected at Hunt Fork in the
winter of 1947-1948 and brought to me in spring when the skins could
no longer be saved. I saw a flock of about 30 feeding and flying about
in their careful method of association in the spruce tops along the
Alatna, just north of Helpmejack Creek, early in August 1951. Ihave
also seen them at Bettles and, in March, among the most northern
spruce around Ernie Johnson’s cabin on the Alatna Malemute.

Susie Paneak collected one of two crossbills which she found at
Itikmalikpuk Creek on July 10,1954. Simon Paneak recalled seeing
one west of Chandler Lake in 1938. The rare records and strangeness
of a crossbill’s occurrence north of timber cause me to omit the species
from the normal avifauna of Anaktuvuk.

Their Nunamiut name is Pakagik, and they are so familiar at the
northern limit of the spruce forests as to deserve mention in relation
to the northern area.

Passerculus sandwichensis anthinus Bonaparte

6 males May 16-July 17 weight (9), 16.1-19.8, — —
average 17.8 g.
1 female, nest June 13 weight 21.8 g. — —
with eggs
1 female July 2 weight 14.9 g. — —
1 young female July 19 weight 14.6 g, —. ——

Called Okpisoyuk by the Nunamiut, which means “staying mostly
near willows,” Savannah sparrows are regularly present in summer
and I have found them usually near marshy areas. Few are identified
because of their unobtrusive behavior, but they are not uncommon.
Our first recorded observations are May 13, 1949, May 16, 1951, May 25,
1952, and May 27, 1954. They are less numerous than Gambel’s spar-
rows but more common than fox sparrows. In some of the other arctic
valleys where the marshy areas are more extensive than at Anaktuvuk
the Savannah sparrows are more numerous.

The nest is neatly and compactly made of fine round grass that di-
minished uniformly in diameter toward the lining, where it is inter-

118 U. S. NATIONAL MUSEUM BULLETIN 217

woven with caribou hair. It was located under dead grass near the
stream from Nakagnik Spring.

Junco hyemalis hyemalis (Linnaeus)

2 males May 22, 1949, weight 19.5, 15.4 g. — —
May 15, 1952

Northern slate-colored juncos are not often seen in the mountains.
My only records are two taken by Thomas Brower at Tuluak Lake
May 22, 1949, and one seen at Nakrak May 9, 1952, in the same place
where the specimen was collected on May 15. The Nunamiut know
juncos and had described them for me before specimens were col-
lected. They call the junco Kayatavaurak. This name is like that of
the pine grosbeak Kayatavak, but with a diminutive ending added.
T could find no information about the nesting of juncos in the moun-
tains. Their appearance there is unusual, but the two specimens were
in good condition. I have seen numerous juncos raised in that season
in the wooded country along the Koyukuk and Alatna to the south and
a few have been reported along the Arctic coast (Bailey, 1948, p. 296).
It is my impression that juncos may nest in the mountains and that
their occurrence represents migration. But from present evidence,
they can only be called visitors to Anaktuvuk.

Spizella arborea ochracea Brewster

26 males May 15-Sept. 10 weight (43), 16.1- — —
22.0, average 18.4
g.
6 females May 21-Sept. 5 weight (11), 16.1- — ——
19.8, average 17.7
g.
8 young males July 24-Aug. 14 weight (8), 16-20 g. — —
2young females July 13, 29 weight (2), 16-17 g. — —

5nests with 5eggs June 9, 16 —_—
in 4, 6 eggs in 1

The first western tree sparrows were recorded May 14, 1948, May
18, 1949, May 11, 1950, May 8, 1951, May 20, 1952, and May 14, 1953.
The latest date recorded is September 12, 1950, at Contact Creek.
During May 1949, the numbers gradually increased until 40 were ob-
served on May 30. This observation does not indicate the actual
abundance, for they were throughout the summer second in numbers
only to the redpolls, with which they occupied the same habitat of
the low willow patches, particularly in the flat bottom of the valleys.
They did not appear to venture to high elevations up the willow-lined
streams, nor away from the willows among the dry rocky places, as
frequently as did the redpolls and longspurs. They were usually seen
near the ground among the lower willow bushes. Specimens are from
Tuluak Lake, Kangomavik, and Contact Creek in Anaktuvuk Pass,

ANAKTUVUK PASS 119

and from Odrivik Lake in the Kiliik Valley. Their Nunamiut name
is Misapsak, which refers to their song.

Among the series of seven young males collected between July 24
and August 14, all had attained adult weight but were deficient in tail
length. Numerous nests were found rather clearly visible on nigger-
heads among short willows or in nearby long grass. The nests were
rather bulky, predominantly of grass, occasionally of moss, were
lined with fine grass and caribou hair, and always contained white
ptarmigan feathers.

The tree sparrows practically overrun the short willows of the
Valley in spring without, however, appearing in organized flocks
as do the redpolls. A majority of the incoming birds act as if about
to take up residence, but a comparison between numbers seen earlier
and later indicates that many move north.

The Nunamiut regard them as common all the way to the arctic
coast, and I have seen them over that range. My estimate is that a
smaller proportion of arriving tree sparrows moves north than is
the case among the redpolls. A great number of both species migrate
northward.

As a natural accompaniment to the hardy behavior of tree sparrows
in their winter range, their body feathers are as thick and fluffy as
are those of arctic residents, and are of the same type. By that com-
parison, their feathers might well afford protection adequate for
winter residence in the arctic. I have no indications to suggest that
they winter in arctic Alaska, and the fatness of the arriving migrant
tree sparrows is appropriate to a long migration route; in this respect
they can be contrasted with the lean redpolls, which may be suspected
of making a shorter migration (see p. 116).

Zonotrichia leucophrys gambelii (Nuttall)

15 males May 16-Aug. 8 weight (29), 22.2- —_— —
29.5, average 25.6
g.

9 females May 28-July 31 weight (9), 21.7-27.4, — —
average 23.3 g.

4 young males July 29-Sept. 12 —_ — —

1 young female July 31 weight 26.7 g. — —

2nests, each with June 9, 12 = = —
5 fresh eggs

Earliest arrivals of Gambel’s white-crowned sparrows were recorded
May 138, 1949, May 19, 1950, May 16, 1951, May 20, 1952 May 14,
19538, and May 18, 1954.

On May 24, 1952, as Terris Moore and I were preparing for a flight
from Bettles to Kobuk and Howard Pass the Gambel’s sparrows were
apparently already settled in nesting places. This was a spring when
the late winter snow and ice persisted uncommonly late in northern

469496—60-——_9

120 U. S. NATIONAL MUSEUM BULLETIN 217

Alaska. The opening of the birch leaves was about two weeks later
than usual at Fairbanks. On the Kobuk River at Kobuk Village the
snow was still deep on May 25 and the streams, just beginning to flow,
had yet to break open the ice on the river. Here the white-crowned
Sparrows were in small groups, but a few appeared to be paired.
Seventy miles north, in the upper Noatak Valley I saw these sparrows
in small groups, with some apparent pairs.

When we reached Anaktuvuk several days later, the first arrivals
of white-crowned sparrows had been reported on May 20, three or
four days later than the mid-date for the preceding 3 years. If this
small delay in observation represented delayed migration, it was not
apparent in the behavior of the birds, which took territory, sang and
nested according to their regular calendar schedule.

Barbara Oakeson (1954) found that male white-crowned sparrows
became numerous at Mountain Village, Alaska (on the lower Yukon
River, lat. 62°07’ N.), before the females did. Of the specimens taken
at Anaktuvuk in May there were 20 males and 5 females. Eleven of
the males in which the greatest length of testes was measured ranged
in this character between 6 and 9mm. Six males measured in early
June ranged in length of testes from 8to10 mm. It appears that at
Anaktuvuk, as Mrs. Oakeson found at Mountain Village, male white-
crowned sparrows arrived on the breeding grounds in numbers before
the females. One female was taken at Mountain Village earlier than
any male, so the precedence of males is not exclusive but rather a
majority. Further, in the arriving males at Anaktuvuk the testes
have not reached full size. The development of eggs also had not
progressed beyond 5 mm. diameter in the latest female taken during
May, and it is apparent that both male and female birds complete
their physiological preparation for breeding after arrival on the nest-
ing grounds.

In connection with these comments upon gonadal] development it is
important to note that Mrs. Oakeson studied only white-crowned
sparrows which showed by their behavior that they had settled at
Mountain Village to nest. The locality is in the peninsula of the
Yukon Delta only 40 miles from the coast of Bering Sea. At Anak-
tuvuk we have not so carefully distinguished that the white-crowned
sparrows examined had settled for nesting. But these sparrows are
rare on the arctic coast of Alaska and there was no noticeable con-
centration of incoming birds in numbers nor were flocks larger than
small groups observed. Since only about 100 miles of possible nesting
territory for white-crowned sparrows exists north of Anaktuvuk the
population examined is at or close to its breeding ground and in this
respect comparable to the birds studied by Mrs. Oakeson.

In early June, many Gambel’s sparrows were singing, having al-

ANAKTUVUK PASS 121

ready settled their territorial claims. They nested on the ground,
usually well concealed under short willows or in long grass at the
edge of willow patches, with five or occasionally six eggs. Like the
birds, the nests were inconspicuous. Four early nests were found:
with two eggs on June 1, 1954, by Simon Paneak; with a complete set
of eggs, on June 9, 1949, by Tom Brower; on June 9, 1951, with eggs,
by Susie Paneak; and with five eggs, at Nakrak, on June 10, 1952.
Hatched nestlings were found by Tom Brower on June 25, 1949. The
nests were rather bulky, and as a result the bottom was raised 50 mm.
above the ground, they were made of fine grass with a few bits of
moss. Among birds resident in Tuluak Valley, Gambel’s sparrows
were common, perhaps 1/20 as numerous as the redpolls and tree
sparrows occupying the willow habitat. They were about 10 times
as frequently seen as fox sparrows.

Seldom conspicuous, and in midsummer keeping to the cover of
the willows, they were nevertheless easily discoverable about many
of the willow patches in Anaktuvuk Valley or in the small alder
thickets as well as in the Killik Valley. They were usually in willows
exceeding 10 feet in height but an occasional pair was found among
the low willows near damp places on the mountain sides. In the
thick willows along Contact Creek in mid-June, I found that when '
I remained still for a few minutes a pair of Gambel’s sparrows would
frequently approach within 5 feet and examine me carefully. But
outside the brush they were commonly shy. Their Nunamiut name
is Vungaktuakruk, which refers to their white-striped crown.

At the end of July, the young were heavier than the adults. Their
departure, like their arrival, is not conspicuous, and the latest record
noted two birds (one young) on September 12, 1950, at Contact Creek.
Although they spend a long season at Anaktuvuk their feathers are
not as fluffy as those of tree sparrows and they do not appear so well
insulated against cold weather.

The white-crowned sparrows were first seen at Mountain Village,
Alaska (on the lower Yukon River, Jat. 62°07’ N.), on May 9, 1950,
10 days ahead of the earliest record for that year at Anaktuvuk. On
arrival at their breeding grounds at Mountain Village the average
weight of 7 males was 25.8 grams and the average weight of 3 females
was 25.5 grams (Oakeson, 1958). The records of weight and fatness of
white-crowned sparrows at Anaktuvuk permit comparison to be made
subsequently (see p. 320) with Mrs. Oakeson’s information upon their
nutritional condition in relation to migration.

Among 25 adult birds examined with the aid of Herbert Friedmann,
1 had only black feathers between the eye and mandible, 16 had the
light superciliary line continuing to gray between eye and mandible,
and 8 had the superciliary line continuing in light or white feathers

b22 U. S. NATIONAL MUSEUM BULLETIN 217

between eye and mandible. Of the 2 additional birds from that
part of Alaska, 1 from Bettles agreed with the gray birds and 1 from
Ahlasuruk agreed with the lighter mark between eye and mandible.

The exceptional bird bears the pattern of the eastern white-crowned
sparrow, as Alexander Wetmore first pointed out tome. It was a male
taken on July 11, weighing 27 grams, with testes measuring 1.5 mm.
It could be interpreted as an accidental wanderer of the eastern race,
far from the usual range of its subspecies, or as an aberrant pattern of
plumage in gambeli that happens to resemble typical leucophrys.
The latter of these two choices is the one followed here, as it seems
more logical than the other.

Passerella iliaca zaboria Oberholser

5 males May 21, Aug. 6 weight (4), 34.3- — —
42.7, average 37.4
g:

2 females June, July 11 aes 30.2 g. == =
2 young males July 11, 1950 weight 34, 34g. == ae

These fox sparrows compare well with those kindly shown me by
Ira Gabrielson as representatives of this race from Alaska.

The earliest records are of a specimen brought to me May 19, 1948,
which unfortunately could not be preserved, June 8, 1950 (an un-
doubtedly late observation), May 17, 1951, May 29, 1952, and May 28,
1954. Because of their shy withdrawal among the willow thickets
early fox sparrows are unlikely to be seen until the males reveal them-
selves by singing from the tops of bushes.

The Nunamiut name for fox sparrows, /élikvik, means “tool bag.”
This name is very appropriate, for it refers to the quick, rattling
sound which the fox sparrow makes in the Arctic, as in temperate
regions, while scratching in dry leaves. It resembles the rattling of
the ivory bone and flint instruments formerly carried in an Eskimo’s
tool bag (the older men used to carry their tools in these bags as they
went to work together on their fish nets and other gear in the men’s
social house of the village, where they would not be disturbed by
women and children, who in turn also had liberty for their occupa-
tions and society). Barbara Oakeson informs me that at Mountain
Village on the lower Yukon the Eskimos also base their name for fox
sparrows on their scratching, but they are there called Shachtayolik,
meaning “he scratches.” As a small boy I first recognized fox spar-
rows when the odd rattling sound of their scratching among the
autumn dry leaves in a swampy thicket called them to my attention.

Almost as early as they have been seen, the males were singing their
fine, sweet song. Among early specimens, the testes varied consider-
ably in size, so that I suspect that they, like Alaska longspurs, are not
all ready for reproduction when they first arrive. Nevertheless a

ANAKTUVUK PASS 123

female bird contained an egg 17 mm. in length on May 29, 1952, and
their nesting must be among the earliest of the sparrows.

A nest was reported on Contact Creek June 7,1951. Through June,
I saw and heard many fox sparrows singing in the willows along Con-
tact Creek, where I suspected that they were already nesting.

There is no indication of extensive migration, but some must pass
northward, for the Nunamiut know that fox sparrows are common
in the willows along the Colville River.

Calcarius lapponicus alascensis Ridgway

30 males May 10-July 20 weight (27), 24.6- —_— ——
33.1, average 28.6
g.

18 females May 10-July 24 weight (22), 22.3- —— —
30.6, average 25.4
g.

9 young males July 5-Aug. 11 weight (8), 22.3- — —
24.7, average 23.6
g.

2youngfemales July 20, 24 weight (2), 24.4- —— —_—
27.4 g.

The earliest recorded arrivals of Alaska longspurs at Tuluak Lake
are May 15, 1948, May 1, 1949, April 29, 1951, May 9, 1952, May 4, 1953,
and May 138, 1954. About the first of May is the usual date of their
arrival. During the first two weeks of May a few were seen each day,
but during the third week in May, flocks increased until May 19, 1949,
when a thousand were reported by Thomas Brower. The popula-
tion remained at migrant size for about a week further and suddenly
declined to resident numbers and behavior before the end of the
month, Their Nunamiut name is Potokioluk.

Early in May and in some years to the end of that month, winter
snow persists in Anaktuvuk Valley. Then the longspurs may be seen
in great numbers on the higher levels of the Valley and particularly
where hummocks of sedge protrude above the snow. In the latter part
of the month, the males frequently sing in downward hovering flights
which terminate in swift, soaring glides to earth. In this time mating
competition goes on, before nesting areas are exposed, among flocks
which are generally moving northward on the ground and in short
flights. I suspect that some mating occurs before the final nesting
grounds are reached.

On May 10 and 11, 1951, five males examined were recorded to have
small testes. On May 14 and 15, the records show that the testes of
four were getting larger and in four, the testes had reached breeding
size. In 11 birds examined during the rest of May, the testes were in
all cases of breeding size.

All males (15) which were examined between May 10 and 16 were
fat. On May 21 and later in that month among 10 males examined

124 U. S. NATIONAL MUSEUM BULLETIN 217

only two were reported fat, three were designated medium fat and
five had little fat. These figures indicate less fat in the males in
breeding condition and during the time of mating, but they were
nevertheless in good nutritional state.

The progress of egg development may also be observed from records
in 1951. In 13 females examined between May 10 and 21, the eggs
were recorded as 1 mm. in length. Eggs were recorded as 2 mm. on
May 21 and 28, 10 mm. May 28, with only small eggs found on June
8 and 9.

On May 10 and 15, seven females were recorded to be fat. Of nine
females examined in the remainder of May, two were recorded to be
fat, three were considered medium fat and four had little fat. These
figures indieate that females, like males, are fat until about the time
of active mating and nest preparation and that thereafter they are
not commonly so well supplied with fat, although their state of nutri-
tion is generally very good.

Two nests found on June 6, 1949, and one on June 6, 1953, con-
tained two eggs each. On June 7, 1949, another nest contained four
eggs and on June 8, one set of two eggs had increased to four, the
other nest having been destroyed. On June 12, several nests were
found with four eggs too far incubated to be saved, and because of
this condition and destruction no specimens were preserved.

The first nest recorded in 1951 contained four eggs on June 4 and
10 days later it contained two nestlings and one egg. On this same
date, June 14, another nest contained four newly hatched birds.

The young were first seen flying inexpertly June 29, 1951, and the
young were all fiying over the grassy tundra in early July. In the -
series of nine young males taken between July 5 and August 11, 1950, |
of eight weighing from 22.3 to 24.7 grams, none had reached adult
weight or length except one which on July 2, weighed 28.5 grams while
still in juvenal plumage. Its testes measured 1.5 x 1 mm., large for a
young bird, and their condition may indicate the reason for enlarge-
ment of the bird. It was omitted from the series of weights as an
abnormal bird.

These longspurs are the most numerous birds of the wet grassy
tundra and in open grassy places among the willows. Commonly,
they remain near water, but they sometimes venture far up onto dry
areas and onto the talus slopes on the Valley sides. Around the tents
of Nunamiut, they are common visitors and in late summer the young
birds venture familiarly close among people and dogs

In the summer of 1951, I saw so many more males than females that
I tried to estimate whether this apparent predominance of males
observed represented the state of affairs in the population. About
mid-May, John Krog remarked that he saw about ten times as many

ANAKTUVUK PASS 125

males as females. During June, I noticed that there were frequently
groups of four or five males amiably feeding together on the dry parts
of the tundra, occasionally to be joined by a female. So frequently
were these small groups of males seen together at the same place that
it was my impression that they were regularly associated companions.
They walked about in their awkward, stiff-legged gait as deferential
to each other as a group of old gentlemen at their club.

When the young birds scattered from the nests at the end of June,
the females came out of retirement and were usually seen in the
proportion of one female to three or four males, but thereafter the
grouping of males was not noticed. The clubs were now broken up.
A collection of young birds which had left their nests was made in
1950. These birds were not selected and the sexes were not distinguish-
able in the field either by appearance or habit and yet the results gave
nine males and two females. When I consider the revealing habits
of these longspurs in May and June, I wonder if the males in the
population in 1951 considerably outnumbered the females.

In the Killik Valley in August 1950, Alaska longspurs were as
abundant as at Anaktuvuk and about twenty times as numerous as
Smith’s longspur. Two young birds from Odrivik Lake and Akmalik
Creek correspond in size with birds from Tuluak. In late July at
Tuluak and more noticeably in early August in the Killik Valley, long-
spurs frequently were grouped loosely in larger than family numbers
of a dozen or so, and the direction of flight was more often south than
north. They were observed August 15, 1950, at Chandler Lake. On
August 25, 1948, there were not many around Tuluak Lake and our
latest record is September 2, 1950, at Contact Creek.

Alaska longspurs arrive earlier and remain later than Smith’s long-
spurs. Alaska longspurs go far north on the arctic tundra in summer
and are common along the arctic coast, but they have not the feathers
typical of northern birds, and they retreat from the far north before
cold weather sets in.

The view of large but loosely organized flocks moving northward in
mid-May shows that Anaktuvuk Pass is an important thoroughfare
for these longspurs in their movement to the arctic slope.

Calcarius pictus (Swainson)

13 males June 4-July 10 weight (22), 25.5- —— —_—
31.8, average 28.5
g.
11 females June 4—July 20 weight (7), 22.0—26, — =
average 24.3 g.
1 young male July 27 weight 27.6 g. — cs
2young females July 24, 27 weight 28.1, 25.3 g. — —_—
Eggs, 4fresh with June 18, 1949 —— mae ——
female

Eggs, 4fresh with June 19, 1949 — — —
female

126 U. S. NATIONAL MUSEUM BULLETIN 217

The earliest records of Smith’s longspur are May 28, 1948, June 4,
1949, June 8, 1950, May 27, 1951, May 26, 1952, May 22, 1953, and
June 1, 1954. We have not noted flocks of these longspurs nor any
grouping of them that might be related to migration, and so I think
that most of those in Anaktuvuk have reached the northern terminus
of their migration. The latest to be reported was seen on August 24,
1950.

The Nunamiut consider that Kallorgosiksook, so named because of
the ability of Smith’s longspurs to sing like several kinds of birds,
are common in some years and scarce in others. During 1950, it was
estimated that they were about a twentieth as numerous as Alaskan
longspurs which would rank them as about the second species in abun-
dance over the open wet grassy part of the tundra at the level of the
floor of Anaktuvuk Valley. In 1951 and 1952, the proportion observed
in summer was about the same.

They did not frequent the dry places, but were often seen in wet
grassy places. Although they stand and fly like Alaska longspurs,
Smith’s longspurs when feeding move through the grass with the swift
purposiveness of a hunting weasel, whereas the Alaska longspur has
a slow, stiff gait, like that of an old man.

Examination of the testes of 10 of these longspurs between May 27
and June 9, 1951, showed all to be at breeding size at the time of
arrival. In this they were unlike the Alaska longspurs which had been
reported two weeks before the time all males were at breeding size.
Three male birds examined May 27 and 28, 1951, were fat. Between
June 1 and 9, out of seven males examined, two were designated
medium in respect to fatness, while five had but little fat. It seems
that like the Alaska longspurs the early male Smith’s longspurs are
fat, and that during the mating and nesting season their fatness
diminishes.

In four females examined between May 27 and June 1, eges were
recorded at 2 mm. in length (that is, to have undergone a little
growth), but on June 5, a bird contained a fully formed egg.

The nests collected by Thomas Brower in 1949 were located on hum-
mocks in the grassy tundra, slightly raised above the wet or damp
surroundings and not concealed. They were rather bulky and were
constructed of grass lined with fine round grass, some caribou hair,
and a few ptarmigan feathers. The sets of eggs were observed for two
or three days before being taken and were complete in number by
about June 15. This places the nesting date about 10 days later than
that of Alaskan longspurs. A set of three eggs was first observed on
June 23 and on June 26, having received no addition in number, it
was found that the eggs were too far incubated to be preserved.

ANAKTUVUK PASS 12%

In 1951, a nest found with two eggs on June 10 contained three eggs
on June 11 and four on June 12. Another nest with two eggs was
found June 11, one with four June 12, and one with five eggs June 21.

In 1952, a nest found on June 3 contained no eggs. Another nest,
empty on June 3, contained three eggs on June 7, four on June 8, and
five when next examined on June 10.

The young birds collected had reached adult weight on July 27 and
the latest recorded observation of Smith’s longspurs was on August 24,
1950. The evidence points to their being briefer summer residents as
well as less numerous than are Alaskan longspurs. Among the
Fringillidae, Smith’s longspur appears to be least suited for cold by
the style and coherence of their feathers. The observations made so
far do not show any formed flights of Smith’s longspurs heading
north through Tuluak Valley. In mid-August, however, more indi-
vidual birds and family sized groups were seen flying south through
the Killik Valley than had been earlier apparent in the local popula-
tion. It is concluded that the main northward movement extends to
the mountain line, but that some proceed further north.

Plectrophenax nivalis nivalis (Linnaeus)

11 males Apr. 2-May 26 weight (2), 38.9, — —
32.6 g.
3 females May 2-May 26 weight (1), 30.6 g. —_ ==

In their northward flight the first migrating snow buntings were
recorded March 28, 1949, April 3, 1950, April 3, 1951, May 9, 1952,
April 3, 1953, and April 16, 1954. The late 1952 observation may indi-
cate that the buntings were retarded by the late heavy snow cover
which persisted in interior Alaska that spring. But I am inclined to
think that they passed unobserved. They commonly reach the arctic
coast at Barrow in mid-April (Bailey, 1948). Sir John Ross (1835,
pp. 322, 511) recorded that the first snow buntings reached their
winter quarters at Boothia (lat. 69° 58’ N.) on April 17 in 1830 and
again in 1831.

At Bettles Hannah Anderson told me that she had seen the first
few snow buntings of 1953 on about March 10. She remembered from
her childhood there that snow buntings returned early in March and
remained numerous for several weeks. It appears that the most north-
ern part of the migration is prolonged on the Koyukuk and then
moves only slowly from Anaktuvuk to Barrow.

Snow buntings are known to winter in southern Alaska from the
Peninsula to Sitka (Ridgway, 1901). Brandt (1948) was informed by
local people that snow buntings were occasionally seen during winter
in the villages along the lower Yukon River. At Kobuk Village,

128 U. S. NATIONAL MUSEUM BULLETIN 217

Charles Sheldon has informed me, snow buntings were seen in Feb-
ruary and March 1954, and the local people expected to see them oc-
casionally every winter. Dall (1869) saw them in winter at Nulato
on the Yukon near the mouth of the Koyukuk. The wintering popu-
lation of snow buntings nearest to those of northern Alaska is in
British Columbia. There they have been reported at Atlin (Swarth,
1936), and I saw a flock near Fort St. John in mid-December 1949 and
again in January 1954. Between the areas where snow buntings are
known to winter in Alaska and Canada the mountains appear like a
barrier which would be formidable in winter and which would deter
east-west migration of snow buntings in spring. Of this we cannot
be certain, but Salomonsen (1947, 1950) thought the Alaskan buntings
might be of a race intermediate between the darker American nivalis
and the lighter Siberian pallidus. At present there seems to be no way
to establish whether snow buntings wintering in Canada join the large
numbers which proceed northward in spring to nest along the arctic
coasts of Alaska. .

By April 29, 1949, snow buntings had become numerous at Anak-
tuvuk. The peak of their abundance was reached on May 9, when over
400 were recorded by Tom Brower. The numbers diminished from
the hundreds seen daily near the peak of migration until the last flock
was recorded on May 18, but a final pair was observed on May 26 and
May 27.

~The Nunamiut name for snow bunting is Amauligak. Henry Col-
lins informed me that many Eskimo groups give that name to snow
buntings because it is related to the distinguishing black-and-white
pattern of plumage of male king eiders, which shows especially clearly
in flight. Simon Paneak remarked upon the resemblance of the two
patterns. Several other birds are named for their resemblance to a
single character of another bird, although the resemblance holds in no
other respect. Swallows are named for the resemblance of their acro-
batic flight to the aerial evolutions of ravens. Wheatears are named
little eagles because of their free flight over the rocky mountainsides.

During summer no snow buntings were seen on the floor of Anak-
tuvuk Valley nor at elevations up to 4,000 feet, which were frequently
visited. At the head of Kangomavik Creek; at an elevation estimated
to be about 6,000 feet Paneak saw snow buntings on August 29, 1950.
We thought these might have been early migrants returning from the
coast or possibly had nested in the mountains, for Ray Hock trapped
nine snow buntings at Barrow between September 11 and 29, 1947.
Mary Lobban saw them at Barrow and Wainwright until the end of
September 1957, and Anderson (1921) last saw them on the Hula Hula

ANAKTUVUK PASS 129

River on September 20. It appears that buntings do not leave the
coast until September.

While at Chandler Lake during the summer of 1955 Paneak fre-
quently observed young buntings with their parents in the high moun-
tains and Jack Campbell repeated these observations in 1956. It is
thus certain that some nest in the high mountains of the interior of
arctic Alaska.

Three snow buntings were seen high in the mountains above Kang-
omavik on October 2, 1950, by Robert Rausch and Simon Paneak, and
Paneak again saw several near Summit on October 20. It is scarcely
possible for the large northbound flights to return southward through
the lower levels of the Pass without being noticed, and I believe that
the southbound flights proceed through the high mountains.

Among the northbound birds the gonadal development was deter-
mined in only three males, which had testes of breeding size on May
15, May 26, and May 29. In two females the eggs were 2 mm. in
length on May 26.

Captive snow buntings lived well through the winter of 1947-1948
at Point Barrow in an outside cage exposed to the full severity of
arctic winter (Scholander, Hock, Walters, Johnson, Irving, 1950).
One was seen among the willows near the winter camp at Kalutak
Creek on January 30, 1950. The buntings possess the feathers char-
acteristic of northern birds, although they are not as fluffy as those
of the regular winter residents, jays and chickadees.

Anaktuvuk Pass is an important thoroughfare for snow buntings
in migration to the arctic coast. The peak of their migration comes
at least a week earlier than that of Alaska longspurs. At elevations
up to 4,000 feet at least there are no regular summer residents.

A nest was brought to me in the late summer of 1947 at Point Barrow
which resembled others of snow buntings in composition but which
was uniquely located, for it was ensconced in the well bleached skull
of an old Eskimo. One day about a year later I noticed that Chester
Lampe, a Barrow Eskimo who most capably cared for the cleanliness
of the laboratory, was looking at the skull and smiling as Eskimos
do in appreciation for humorous observations. Expecting some witty
comment I asked him if he did not think that an odd nesting place.
He said that it was unusual but that it was the bird’s nature to enclose
its nest in shelter. He then asked me very seriously if I thought the
nest had been built in the skull before or after the demise of its mortal
owner. Suspecting some fantasy of Eskimo wit, I cautiously suggested
that birds did not commonly nest in living human skulls. Thereupon
he gravely remarked that he had known several men who certainly
acted as if their skulls contained bird nests. I agreed that from the

130 U. §. NATIONAL MUSEUM BULLETIN 217

evidence his supposition that the skull was tenanted during the life
of its owner by a bird’s nest was as good as mine that the birds had only
sought shelter in the shell of a former intellect. So perhaps if Hamlet
had been as observant as an Eskimo he would not have felt warranted
in basing a soliloquy upon the probability that intellectual processes
had occurred in a single human skull selected at random.

TE ay Sees GP are —S
ak Xe, 3 SABE ©

Gray-Crownep Rosy Finca Leucosticte tephrocotis irvingi (see pp. 112, 148).

3. Kobuk

E WAS IN MAY 1952 that I was introduced to a new portion of the

northwestern interior of Alaska in the pleasant company of Terris
Moore, president of the University of Alaska, when he piloted me to
Kobuk village, the upper Noatak River, and Howard Pass (see
map, fig. 1).

The climate of the Kobuk Valley is tempered by the protection of
the mountains along its northern margin, so that subarctic plants
flourish and cottonwood, birch, and spruce reach good size. Asa result
of irregular glacial deposition and the meandering of the river through
its gravel plain the wooded areas vary in extent and pure stands of
timber are rare. Water collects on level areas because it cannot drain
through the underlying permanently frozen ground. Some of the fre-
quent marshy flats and lakes give way to areas of dry tundra over
the outwash from tributaries and moraines in the upper valley. Wil-
low and alder thickets are extensive. Because of the terrain and prox-
imity to the coast there are some tundra and maritime species of birds
in addition to the fauna of the northwestern forest.

131

132 U. S. NATIONAL MUSEUM BULLETIN 217

Grinnell (1900) is the only naturalist who has observed the inte-
rior of the Kobuk Valley during all seasons of the year. Earlier
McLenegan (1889) and Townsend (1887) had reported upon the birds
they found during arduous summer expeditions for exploration of
the Kobuk River, and Stoney (1900) left only a few comments on
natural history based on his magnificent explorational travels in the
region in 1885-1886. Grinnell described the birds which he found
around Kotzebue Sound and around his winter camp on the Kobuk
opposite Hunt River about 70 air miles from Kotzebue. I have selected
those species which the three authors remarked as occurring along the
wooded interior part of the Kobuk River. I find that McLenegan
recorded 60 species, Townsend 50, and Grinnell 85. Their combined
records list 99 species on the wooded Kobuk.

Since 1952 I have been at Kobuk Village on five occasions, during
which I have observed the birds there and obtained a few specimens.
The major part of my information, however, has been gained from
discussion with some of the 60 resident Eskimo, and especially from
migration records and lists prepared for me in 1954 by Charles Shel-
don, who has long been a local leader in school and church affairs.
(William Irving and I were led by Sheldon to a cursory examination
of a number of old depressions which probably represented sites of
ancient winter houses; these were at the mouth of Kugaluktuk Creek
and are now overgrown with sizable trees.) Harry Brown, the long
respected trader at Kobuk and member of the Alaska Game Com-
mission, introduced me to the people who best knew the country and
Simon Paneak, from his long acquaintance with the country and his
knowledge of the people helped me to evaluate the information
obtained.

As an aid for our discussion of birds we had a list of Nunamiut
names for about 90 species of birds (L. Irving, 1953). Sheldon named
103 species in the Kobuk manner of speech and recognized explicitly
110 species of the list of 122 birds of Kobuk which I have compiled
from published records and our information. Nunamiut and Kobuk
names differ to some extent (lL. Irving, 1958a), but Paneak and
Sheldon were familiar with each other’s nomenclature through the long
and friendly relations of their people. Sheldon prepared several
specimens and kept a journal of his observations during April, May,
and early June 1954.

The combination of ancestral Eskimo knowledge of natural history
with the current observations of individuals and communities affords
information gathered by lifelong observers in all weather and situa-
tions. Novisitor can by himself gain such understanding of the fauna
as is possessed by some of the older members of these Eskimo com-
munities. Itis not easy to prepare one’s self to receive the information

KOBUK 133

of these Eskimo experts in natural history, for the people are extremely
critical about accuracy in recognition of species and they cultivate
precision in observation. However, Eskimo appreciation for com-
petent interest in birds is so great that evidence of a visitor’s serious
scientific interest serves as an introduction which in due time opens for
him the store of their knowledge, and they then eagerly join in making
observations and records in order to provide for the transfer of their
knowledge into the terms of scientific ornithology.

The Birds of Kobuk

On the following pages is presented the new information gained
from our investigations at Kobuk, together with comments upon earlier
reports. From these sources, too, has been compiled the list given in
table 4 (p. 246) of species now known at Kobuk. Because the named
races give important indications of geographical distribution in arctic
Alaska, I have designated them in every case where it was possible
to do so on the basis of specimens found in the U. 8. National Museum,
on identifications by Townsend and Grinnell (made consistent with
modern nomenclature), and on specimens which Sheldon and I have
obtained. The status of most species in respect to residence, migration
and nesting appears clear. . The designation of a bird as a visitor is
not intended to suggest that the bird’s occurrence was accidental. In
most cases these birds were known in nesting areas or on migratory
paths nearby and their appearance at Kobuk, although not known to
be for nesting, migration, or residence, is not to be considered abnormal.
In order to reduce the categories used a few birds are designated
visitors because the scant information now available does not indicate
their true status.

Family PODICIPEDIDAE: Grebes

Podiceps grisegena holbollii Reinhardt
Podiceps auritus cornutus Gmelin

Red-necked and horned grebes were clearly described in appear-
ance, behavior, and call by Sheldon. Both had been found at the
mouth of the Kobuk River to the west (Grinnell, 1900). I have seen
red-necked grebes at Kobuk, horned grebes on the Alatna River
about 80 miles east of Kobuk, and I have collected and designated
the latter a rare visitor north of the forests at Anaktuvuk. Sidney
Peyton saw red-necked grebes at Bettles on the Koyukuk, 140 miles
east of Kobuk. Both species accordingly occur on the forested south-
ern watershed of the Brooks Range across Alaska. Since they also

134 U. S. NATIONAL MUSEUM BULLETIN 217

nest at Old Crow where the Porcupine River extends the Yukon
Valley almost to the Mackenzie River the normal range of both
reaches just north of the arctic circle over the full extent of the
Brooks Range.

Family ANATIDAE: Swans, Geese, Ducks
Branta canadensis taverneri Delacour

Sheldon obtained a male specimen from a pair of Canada geese on
May 19, 1954, at Kobuk. It conforms with six specimens obtained in
migration at Anaktuvuk and with six specimens of these geese from
families raised on the Koyukuk and Alatna Rivers for which Herbert
Friedmann confirmed my identification as tavernert according to a
description by Delacour and Zimmer (1951). The testes of the Kobuk
specimen were 3/ mm. long, greater than in three geese measured dur-
ing migration at Anaktuvuk. It was near reeds condition and may
represent the numerous Canada geese which nest along the wooded
Kebuk in the western interior of arctic Alaska. Grinnell (1900) re-
ported hutchinsi, the common nesting goose on the wooded Kobuk.
Townsend (1887) also reported minima and McLenegan (1889)
leucopacia. Since their specimens are not available these earlier iden-
tifications cannot be reliably related to the current nomenclature and
we cannot say whether other races than taverneri occur there.

Branta nigricans (Lawrence)

As Grinnell (1900) observed, numerous large flocks of brant migrate
through the Kobuk Valley late in May but are not seen returning in
autumn. Often the migrants fly over the Valley in a northeasterly
direction at 4,000 or 5,000 feet, which would clear the adjacent moun-
tains, but farther east I have watched and heard them clearing some
of the highest peaks in northbound flights at elevations estimated to
be over 8,000 feet. No specimens have yet been obtained from Kobuk
but at Anaktuvuk nzgricans has been identified.

Philacte canagica (Sewastianov)

Sheldon found an emaciated emperor goose near Kobuk in 1954. It
is known at Kobuk to be a coastal species which rarely straggles inland
and it is therefore not numbered in the regular avifauna of Kobuk.

Anser albifrons (Scopoli)

White-fronted geese appear about mid-May at Kobuk. Sheldon
said that some flocks disappear northward and I have seen them in
northward migration over Howard Pass. A large number remain in
the Kobuk Valley in summer but move out in August before the Can-
ada geese, some of which remain into September.

KOBUK 185

Spatula clypeata (Linnaeus)

Sheldon obtained and named the skin of a shoveler, which I have
seen and identified, and described it as one of an apparently settled
pair on May 23, 1954. It is known by the people at Kobuk that a
few shovelers usually nest in the vicinity, but not enough are seen
to show that migration proceeds further.

Aythya marila nearctica (Stejneger)

The Eskimos at Kobuk reported scaup tobe common. Some distin-
guished greater and lesser scaup by Eskimo names which they said
specified two kinds. Sheldon’s description seemed to distinguish two
sizes. In support of the description, he reported the weight of a male
scaup at 870 grams. This he named Kak/odook and said that it corre-
sponded to our “lesser scaup.” This is just within the smaller size
range of A. marila and greater than three weights recorded at
Anaktuvuk for male A. afinis (769, 681,694 grams). By reference to
these weights the bird was probably A. marila. The Kobuk names for
the two scaups differ by only a suffix, “pw,” meaning big. Although
lesser scaup may occur and are perhaps distinguished at Kobuk, the
evidence is not yet certain enough to include A. affinis in the established
list of avifauna.

Bucephala albeola (Linnaeus)

Sheldon’s appropriate description and an explicit Eskimo name
caused me to include the bufflehead as a regular occurrence at Kobuk,
although it had not been earlier reported.

The four Alaskan species of eider ducks are well known to Kobuk
people from their visits to the coast, but individual eider ducks which
occasionally stray inland should not be listed as part of the regular
avifauna.

Melanitta deglandi (Bonaparte)

The white-winged scoter, which had been earlier reported only by
McLenegan (1889), was clearly described by Sheldon and given an ex-
plicit Eskimo name. It is well known among Nunamiut and is also
known as a regularly nesting bird at Kobuk.

Oidemia nigra (Linnaeus)

The common scoter, previously reported at Kobuk by Townsend
(1887) and Grinnell (1900), was named by Sheldon. As it has not
been recognized at Anaktuvuk it probably does not range into the
arctic interior beyond the Kobuk River.

469496—60-———10

136 U. 8S. NATIONAL MUSEUM BULLETIN 217

Family ACCIPITRIDAE: Hawks, Harriers

Accipiter gentilis (Linnaeus)

McLenegan had earlier reported goshawks as seen “once or twice,”
and he reported a specimen. Goshawks are well known to Eskimos at
Anaktuvuk as woodland birds and they are designated with the same
name at, Kobuk, where they are known to these woodland Eskimos as
residents of the forest.

Accipiter striatus velox (Wilson)

There are a number of names for hawks in Eskimo. In some con-
versations the various kinds of hawks appear to be as vaguely desig-
nated as by English vernacular names. It has therefore been difficult
for me to relate the information of Eskimos to the species of this
family. Sharp-shinned hawks are not known to occur at Anaktuvuk,
and I could not be sure that they were. explicitly known by the Es-
kimos at Kobuk. Grinnell’s (1900) report shows that sharp-shinned
hawks are found at Kobuk.

Aquila chrysaétos canadensis (Linnaeus)

Although golden eagles had not been earlier reported on the upper
Kobuk, they are named there, as at Anaktuvuk, and are so well known
among Daeains that they are listed as regularly nesting 1 in the moun-
tainous areas along the upper Kobuk.

Haliaeetus leucocephalus alascanus Townsend

The bald eagle, earlier reported seen only by McLenegan (1889) was
clearly described and named at Kobuk but not known to nest there.
It isa rare visitor beyond tree line at Anaktuvuk.

Family PANDIONIDAE: Ospreys
Pandion haliaetus carolinensis (Gmelin)

Sheldon showed me an osprey’s nest in 1954 on a tall spruce near
Kogaluktuk Creek. He said that the nest had been used for many
years. In 1957 I saw an osprey carrying a fish.toward this nest and
was informed that the nest was occupied as usual. We have occa-
sionally seen ospreys north of the timber in the central Brooks Range,
ut Simon Paneak remarked that he had never seen one fishing in
those waters.

Family FALCONIDAE: Falcons

I am still uncertain that the Kobuk Eskimo name and description
refer explicitly to duck hawks (Falco peregrinus). It is likely that

KOBUK 137

they occur and are known on the wooded part of the Kobuk but since
no other report names them, their presence is not taken to be established.

The variety of names given to small hawks in Eskimo can be as
confusing as their vague designation in English. I do not think that
the sparrow hawk (/f. sparverius) is known in Kobuk. Only Mc-
Lenegan (1889) had named them on the upper river. In agreement
with Grinnell (1900) I do not consider their regular occurrence at
Kobuk to be established.

Family TETRAONIDAE: Grouse and Ptarmigan

McLenegan (1889) heard ruffed grouse (Bonasa wmbellus) drum-
ming in the woods along the upper Kobuk, but Townsend (1887) did
not mention them and Grinnell (1900) reported that he could find no
valid indication of their presence. The Eskimos at Kobuk would
know if ruffed grouse were now present, but they did not appear to
recognize them from descriptions. They do mention stories of “wil-
low grouse,” which had been seen about 1928, and which they
described like sharp-tailed grouse, Pediocetes phasianellus. Stories
among the Nunamiut, and information related to me by David Tobuk
and Big Joe Sousik, two older Eskimos at Bettles, indicate that
sharp-tailed grouse (often called “willow grouse” in Alaska and
Yukon) had occurred some years ago in the forests along the southern
slopes of the central Brooks Range. Nunamiut and Kobuk people
alike name these sharp-tailed grouse Odgillimakadga, the name mean-
ing “birch ptarmigan.” Dall and Bannister (1869) found ruffed
grouse at Nulato and generally along the Yukon. It is possible that
its range, and likely that of the sharp-tailed grouse, extended to
Kobuk at some former time. The northwestern border of their range
apparently does not now reach the forests of the Kobuk or the central-
southern watershed of the Brooks Range.

Family CHARADRIIDAE: Plovers, Turnstones, Surfbirds

Charadrius vociferus vociferus Linnaeus

Among Kobuk people and Nunamiut the killdeer was apparently
recognized in illustrations and its distinctive behavior and appear-
ance were suitably described. Sheldon gave it a different Eskimo
name from that of the Nunamiut, but both Eskimo names are akin
to those for the other plovers and unrelated to any of the names for
sandpipers. The appearance of killdeers at Kobuk would so far ex-
tend their reported range that the present evidence cannot be said
to establish their occurrence there.

138 U. S. NATIONAL MUSEUM BULLETIN 217

Arenaria interpres (Linnaeus)

The turnstone is described and named at Kobuk, as at Anaktuvuk.
At Anaktuvuk specimens have occasionally been taken during migra-
tion. With that inland range established and because of their common
occurrence at Kotzebue, I accept the Eskimo information as evidence
for the migration of some turnstones through Kobuk.

Aphriza virgata (Gmelin)

Three specimens were collected by Grinnell (1900) from a flock of
six. I did not find that descriptions and pictures of surfbirds were
recognized at Kobuk. Grinnell’s identification lists the surf bird and
his observations suggested to him that it nested there, but for lack
of evidence as to the status of the species I designate it a visitor.

Family SCOLOPACIDAE: Woodcock, Snipe, Sandpipers

Bartramia longicauda (Bechstein)

Sheldon’s description, recognition, and name, Vanwm kanockdo-
roagna which he described as meaning “inland plover” indicated his
familiarity with the upland plover. Townsend’s specimen from
Kobuk has provided the westernmost and until recently, the only re-
port of upland plover in northern Alaska. Tom Brower obtained
one specimen for me at Anaktuvuk and I have thought that several
Eskimos and I saw them there a few times during the last 10 years.
Although I think that the upland plover is known to Kobuk Eskimos,
it is included in the avifauna because of Townsend’s specimen.

Tringa solitaria cinnamomea (Brewster)

The absence of a name for solitary sandpiper at Kobuk probably
results from some misunderstanding. Since Grinnell (1900) found
these sandpipers common they should be known to the observant
Eskimos. Solitary sandpipers are occasionally found in spring mi-
gration at Anaktuvuk, where they are well known to the Nunamiut.
Sidney Peyton found their eggs near Bettles and the birds undoubt-
edly nest on the Kobuk and probably along the whole southern water-
shed of the Brooks Range, for they are common at Old Crow, Yukon
Territory.

Heteroscelus incanum (Gmelin)

Wandering tattlers are well known among the Nunamiut, who find
many of them and an occasional nest in the mountains, and we have
found them north of Kobuk on the Ahlasuruk (L. Irving and Paneak,
1954). Among Alaskan sandpipers they are so distinctive in appear-
ance and habits that Sheldon’s name for tattlers, Adlakuk, meaning
“stranger” is appropriate, although it differs from the onomatopeic

KOBUK 139

Nunamiut name. His report and a specimen denote the tattler’s oc-
currence at Kobuk. Because tattlers proceed quickly, after arrival
from migration, to nest along gravelly streams in the mountains, they
would have been seldom in the path of the ornithologists on the Kobuk.

Erolia bairdii (Coues)

Grinnell (1900) reported that few Baird’s sandpipers were seen on
the Kobuk, and in summer on the Ahlasuruk, 80 miles north, we saw
only one (L. Irving and Paneak, 1954), although numbers migrate
through Anaktuvuk and many remain there to nest. Sheldon recog-
nized and named Baird’s sandpiper as occurring at Kobuk but as
they seem to be uncommon, the Valley is probably off the main routes
of their migratory traffic. I have no evidence that they nest near

Kobuk.

Erolia alpina pacifica (Coues)

The appearance of red-backed dunlins is familiar to inland Eskimos
because the birds are well known on the coast. At Anaktuvuk only a
few have been collected during the time of northward migration.
From Sheldon’s account it seems that a few are seen at Kobuk, al-
though previous reports had not recorded their occurrence. Not being
sure that there is a regular migration through Kobuk, I designate them
as visitors.

Limnodromus scolopaceus (Say)

Only McLenegan (1889) had earlier reported dowitchers on the
middle Kobuk. They were found in summer north of there, on the
Ahblasuruk (L, Irving and Paneak, 1954). At Anaktuvuk dowitchers
are common for a few weeks during spring migration. They are well
known among Alaskan Eskimos, and Sheldon’s description and a
Kobuk name, although differing from that of the Nunamiut, indicate
that dowitchers are known to occur there in their proper habitat
during migration.

Micropalama himantopus (Bonaparte)

I could not discover that stilt sandpipers were known at Kobuk.
At Anaktuvuk they were recognized and their nesting on the arctic
coast was described. Several specimens have been taken there. The
numbers seen are few, however, and in some years they are not no-
ticed. If they are as irregular at Kobuk they could easily escape
recognition in the greater concealment of the woodland.

Tryngites subruficollis (Vieillot)

Buff-breasted sandpipers are well known to Eskimos on the eastern
arctic coast of Alaska. The duration of their migration through

140 U. S. NATIONAL MUSEUM BULLETIN 217

Anaktuvuk is brief and they have not been reported every year.
An explicit Eskimo name and apt description of their distinctive
plumage by Sheldon indicates their migration through Kobuk, al-
though no previous observer had reported them.

Limosa lapponica (Linnaeus)

McLenegan (1889) reported seeing godwits on the Kobuk away
from the coast. He listed his observations of godwits in the interior
under L. hemastica. As Bailey (1948) indicates, Japponica is well
known by specimens from the arctic coast and interior of Alaska, but
hemastica is known there by only two specimens from Barrow and one
attributed to Stoney on the Kobuk. The godwits seen by McLenegan
on the interior Kobuk and described and named by Sheldon were
probably /apponica. McLenegan’s description suggests that they nest
there, but I have only found evidence for their Eskimo recognition
and name. Until evidence for nesting is available the godwits seen
at Kobuk should be considered as migrating lapponica.

Crocethia alba (Pallas)

Only McLenegan (1889) had earlier reported sanderlings at Kobuk.
Sheldon recognized the appropriateness of their Nunamiut name
which means “having no heel,” but gave a different Kobuk name,
Akpukshookti, descriptive of their running on the shore. Since
sanderlings can easily be distinguished among sandpipers by descrip-
tion, I consider them known to migrate through Kobuk. Few sander-
lings have been seen at Anaktuvuk and they are apparently also
uncommon at Kobuk.

Family PHALAROPODIDAE: Phalaropes

Both phalaropes are well known at Kobuk and Anaktuvuk. Red
phalaropes are so conspicuous in appearance at the time of migration
that Sheldon’s explicit name and description indicate that in some
spring migrations they are seen for a short time at Kobuk in various
but not large numbers.

Family STERCORARIIDAE: Jaegers, Skuas

Pomarine jaegers had not been earlier reported at Kobuk, but
Sheldon knew their odd-shaped tail feathers well and reported them
as occasionally seen. Eskimos usually know the three jaegers and
their various habits. Sheldon’s description gave no evidence for
migratory traffic, but suggested that as at Anaktuvuk, the pomarine
jaegers which are occasionally seen should be called visitors at Kobuk
until some regularity in their appearance is established.

KOBUK 141

Family LARIDAE: Gulls, Terns
Larus hyperboreus Gunnerus

Glaucous-winged gulls are not known to the Nunamiut and I could
not find that they were known to occur at Kobuk. Nelson (1887)
found glaucous-winged gulls rare north of Bering Sea and remarked
that, no specimens had been taken from the interior. Bailey (1948)
reported that Grinnell’s (1900) Kobuk specimen was not a glaucous-
winged gull and that various other arctic coast specimens reported
to be glaucous-winged gulls should be otherwise identified. In August
1957, while occasionally traveling along the Kobuk between Ambler
River and Manneeluk I did not distinguish a glaucous-winged gull
among some 200 glaucous gulls which I saw. Adding this evidence
together convinces me that Townsend’s report of glaucous-winged
gulls along the Kobuk River is not characteristic of their distribution
and that they have not been identified in the interior arctic forested
country.

Larus argentatus Pontoppidan

Herring gulls have not been earlier reported at Kobuk, but it was
evident that they were well known to Sheldon for their wing pattern
and difference from short-billed gulls in size and color of feet and bill.
Occasionally a herring gull visits Anaktuvuk, and I collected a speci-
men from a family group on the Alatna River in its wooded section
about 80 miles east of Kobuk village. They appear to be the common
large gull nesting in the northern wooded interior.

Xema sabini (Sabine)

Grinnell (1900) saw only one group of six Sabine’s gulls near Hunt.
River. Sheldon recognized the descriptions of them as common gulls
on the coast and said that they were occasionally seen at Kobuk. They
are also seen in the interior of arctic Alaska as far south as Anaktu-
vuk, and the Nunamiut and Kobuk names are similar. There is no
indication that the Sabine’s gulls seen in the interior are other than
temporary visitors.

Family STRIGIDAE: Typical Owls
Bubo virginianus lagophonus (Oberholser)

Early in July 1954 Sheldon and I saw an adult and three young
great horned owls just able to fly clumsily in the trees along Kugaluk-
tuk Creek, the next large tributary to the Kobuk above the village.
I also saw one in August 1957 about 10 miles above Kobuk. They
are familiar to the residents of Kobuk and evidently nest in the
western arctic forest.

142 U. S. NATIONAL MUSEUM BULLETIN 217

Strix nebulosa nebulosa Forster

The great gray owl is known by both Nunamiut and Kobuk people
as a woodland resident with the same name and clearly described
appearance. Indians at Old Crow, Yukon Territory, named an ex-
ample, which one of them shot for us, and remarked upon its appear-
ance, habits, and even its merit as food which distinguish it from the
great-horned owls. The Old Crow Indian name, nastok, is one of the
three Indian specific names among 91 which resemble Eskimo naming.
(L.. Irving, 1958b). Arctic people seem to have a special interest in
collecting and recalling information about owls. The great gray
owl had been mentioned earlier on the Kobuk only in rather vague
comments by McLenegan (1889).

Aegolius funereus (Linnaeus)

The boreal ow] is also well described by Eskimos at Kobuk, although
it had not been earlier reported there. This little owl is named and
described alike by Nunamiut (L. Irving, 1953) and at Kobuk. Like
the great gray owl, it usually keeps in thick timber, where it is not
commonly seen, but I find that many people living in the arctic forests
are likely to give clear details as to localities and circumstances in
which owls have been seen and that they are usually precise in the
descriptive comments by which they distinguish the species.

Family ALCEDINIDAE: Kingfishers
Megaceryle alcyon caurina (Grinnell)

Sheldon gave me three names for kingfishers and showed his special
liking for them by saying that he would not care to shoot one. King-
fishers have been noted by all ornithologists on the Kobuk. I have
seen them there and on the Alatna and upper Koyukuk Rivers. The
Nunamiut have heard of them from their forest-dwelling neighbors,
but I could find no sign that kingfishers had been seen north of the
large forested rivers.

Family PICIDAE: Woodpeckers, Wrynecks
Colaptes auratus borealis Ridgway

The yellow-shafted flicker, reported only by McLenegan (1889) on
the upper Kobuk, was unmistakably described and named by Shel-
don who reported it frequently heard in the spring of 1954. When
I reported seeing a flicker at Anaktuvuk to the Eskimos there, they
were surprised at its appearance on the tundra for they had only
heard it described by neighbors living on the wooded Koyukuk.

KOBUK 143

Kobuk must be about the northwestern limit of the flicker’s range, but
apparently it regularly nests there.

Dendrocopos villosos (Linnaeus)

From illustrations and descriptions, Sheldon considered the large,
northern hairy woodpecker to be a resident bird at Kobuk, but with-
out giving it an explicit Kobuk name. Since I have no other evidence
for the hairy woodpecker’s occurrence in arctic Alaska it is not listed
among the birds of the Kobuk.

Dendrocopos pubescens nelsoni (Oberholser)

Although downy woodpeckers were only designated by the general
Eskimo name for woodpecker at Kobuk, they were well described by
Sheldon. There are a few specimens from Anaktuvuk, and they evi-
dently sparsely inhabit the northwestern forested regions of Alaska.

Picoides arcticus (Swainson)

Because it had not been earlier reported from arctic Alaskan forests,
Sheldon’s description of the black-backed three-toed woodpecker
caused me to question the correctness of our understanding until, to
conclude the discussion, he sent me a correctly named skin for which
Herbert Friedmann confirmed the identification.

Picoides tridactylus fasciatus (Baird)

With P. arcticus distinguished, it was clear that the ladder-backed
three-toed woodpecker also was known at Kobuk. It had been identi-
fied from the Kobuk near Hunt River by Grinnell (1900) and from
the northernmost forest just south of Anaktuvuk on the John River.

The ladder-backed three-toed woodpecker was the only one reported
by Grinnell (1900) on the upper Kobuk, and McLenegan (1889) had
reported seeing only a flicker and a downy woodpecker. Sheldon’s
accurate descriptions and specimen convince us that four species of
woodpeckers are known by the Kobuk Eskimos, with uncertainty about
the fifth, D. villosus. All are called toyuk by the interior Alaskan
Eskimos. The Nunamiut distinguish a toyuk larger than D. pube-
scens as toyukpuk, and have provided specimens of P. t. fasciatus from
the adjacent woodland for thisname. At Kobuk, toywk is applied to
D. pubescens, toyukpuk to P. arcticus and P. t. fasciatus, which are
nevertheless known as distinct forms, and the same name is given for
another which they describe suitably for D. villosus. C. auratus is
designated toyushookruk, meaning “beautiful” toyuk. Although Shel-
don recognized four, possibly five, woodpeckers he gave only three
modifications of a common name for them. It is surprising to find that

144 U. S. NATIONAL MUSEUM BULLETIN 217

people who have 18 explicit Eskimo names for their species of ducks
and 15 Eskimo names for their 16 species of sandpipers (Irving, 1958a)
are undiscriminating in nomenclature of woodpeckers, although they
distinguish the species.

Family TYRANNIDAE: Tyrant Flycatchers

Say’s phoebe (Sayornis saya) has not been reported in arctic Alaska
west of the Killik Valley. It was not found on the Ahlasuruk (L.
Irving and Paneak, 1954), nor has any flycatcher been reported in any
account of the Kobuk.

Family ALAUDIDAE: Larks
Eremophila alpestris (Linnaeus)

The horned lark had not been reported earlier on the Kobuk, but it
is familiar to the Eskimos and is named by Sheldon as by the Nuna-
miut, Vakrulik, meaning “horned.” Horned larks are not woodland
birds. In the Brooks Range they nest on open and usually dry eleva-
tions, away from the paths of observers on the main river. £. a. arti-
cola has been reported 80 miles north of Kobuk on the Ahlasuruk (L.
Irving and Paneak, 1954) and larks nest in suitable mountain habitats
near Kobuk.

Family HTIRUNDINIDAE: Swallows

Iridoprocne bicolor (Vieillot)

Tree swallows now commonly occupy nest boxes and occasionally
holes in the low buildings at Kobuk and villagers say they have become
more common, or at least more familiar, since the practice of erect-
ing nesting boxes was started. TZachycineta thalassina, violet-green
swallows, which nest less commonly in the same places as tree swal-
lows at Bettles, are not known at Kobuk.

Riparia riparia riparia (Linnaeus)

Bank swallows nest in many colonies along the river banks, and the
people in the village remarked on the bird’s skill in selecting suitable
sandy banks unlikely to fall during the terrific erosion caused by the
sudden and violent summer floods. Harly in August 1957 a few were
still on the Kobuk, the only swallows seen at that time.

Mirundo rustica Linnaeus

Barn swallows are known but are rare at Kobuk and are not now
known to nest there.

KOBUK 145

Petrochelidon pyrrhonota (Vieillot)

Cliff swallows are not now familiar near the village, where the houses
seem to be too low to be attractive for nesting. Cliff swallows at Bet-
tles nest under the eaves of buildings with a second story. Eskimos
at Kobuk know about their nests on cliffs and name them “mud swal-
lows.” Cantwell (1887) alone of earlier travelers reported seeing a
great many nesting on the cliffs by the falls near the head of the Ko-
buk. The location of nests and probably the numbers of all but bank
swallows have undoubtedly been much affected by the recent increase
in numbers and size of buildings in northern Alaska.

For species of swallows which occur near the Nunamiut country
and Kobuk there is an indistinction in the Eskimo nomenclature simi-
lar to that in their nomenclature for woodpeckers. Tree swallows (J.
bicolor), bank swallows (2. riparia), and barn swallows (Z. rustica)
are known at Anaktuvuk by their occasional appearance. On visits
to the Koyukuk, Simon Paneak promptly recognized cliff swallows
(Petrochelidon pyrrhonota) and violet-green swallows (7Z'achycineta
thalassina) astwo forms newtohim. But the Nunamiut have only one
name, meaning “like a raven” for all swallows. The Kobuk people
recognize four species of swallows but also give them only descriptive
modifications of the single name.

It might be suggested that Eskimo names for the woodland wood-
peckers and swallows are lacking because Eskimo nomenclature has
developed during residence on the tundra. Their language seems not
to have been expanded to fit such objects as the woodpeckers and
swallows which are part of a woodland environment which few Eski-
mos know well. Nunamiut and Kobukmiut individuals are prompt
to recognize distinct species of the woodpeckers and swallows which
they see when they are away from their own environment, for in gen-
eral new knowledge comes readily to the observant Eskimos, but new
words evidently come slowly into their language.

Family PARIDAE: Titmice, Verdins, Bushtits

Sheldon was only sure in describing the black-capped chickadee
(Parus atricapillus). Although the Nunamiut had collected speci-
mens of Brown chickadees (P. h. hudsonicus) for me from the spruce
timber along the John River and recognized them as differing from
black-capped chickadees, they named the latter alone as “the chicka-
dee.” I have not yet seen P. cinctus lathami, but McLenegan (1889)
and Grinnell (1900) have identified it at Kobuk. The absence of
information about the forms of chickadees in both Kobukmiut and
Nunamiut may result because only the black-capped chickadee is
known as a tundra bird, where it is apparently part of the classical
knowledge in the conservative Eskimo culture of the tundra.

146 U. S. NATIONAL MUSEUM BULLETIN 217

Family CINCLIDAE: Dippers

Cinclus mexicanus unicolor Bonaparte

The dipper was reported on the Kobuk by Grinnell (1900) from
the unmistakable account of white trappers, for it is one of the few
species of birds which white trappers and prospectors can describe
well enough to demonstrate their explicit recognition. The Kobuk-
miut and Nunamiut know the dippers and their habits because in
winter dippers frequent the open water of streams, places which the
Eskimos observe as dangerous for travelers and which they watch
for signs of fish. The two slightly differing Kobukmiut and Nuna-
miut names for dippers were translated for me as meaning “old
woman sunk.”

Family TURDIDAE: Thrushes, Solitaires, Bluebirds

Sheldon’s report to me that olive-backed thrushes (Hylocichla ustu-
lata) were distinguished from gray-cheeked thrushes at Kobuk, and
nested there, is new. His description seemed to differentiate the
bird and its nest from H. minima, but he had no explicit name, using
that (Beeauk) which is also applied to H. minima. It would not be
surprising to find H. ustulata incana at Kobuk for I have collected it
at Bettles, 140 miles east in the same latitude, and at Old Crow. It
seems probable to me that Sheldon’s report from Kobuk will be con-
firmed, but I consider it as yet uncertain evidence for inclusion of
olive-backed thrushes in the regular avifauna.

I have found the bluethroat (Luscinia svecica) once, at Ahlasuruk
(L. Irving and Paneak, 1954), although I have searched for it in the
Brooks Range. It has recently been found to be rather common in the
Colville Valley (Kessel, Cade, Schaller, 1953). The interesting range
of this small migrant to Alaska from Asia, thus far only found outside
the tree line, prompts curiosity as to whether it enters the forested
areas of Alaska. I have found no trace of Eskimo acquaintance with
the very distinctive blue-throat, although it has now been reported
to be rather common along the northern border of the country of the
Kobuk people and probably occurs through much of the country
formerly occupied by the Nunamiut.

Oenanthe oenanthe oenanthe (Linnaeus)

The wheatear is well known among Nunamiut and is apparently also
a part of Kobuk natural history, with the same name in both places.
While nesting, it is a shy bird of the rocky mountain sides in the
central part of the Brooks Range, where I have found it from Anak-
tuvuk to the Ahlasuruk and in intervening valleys. Kobuk hunters
for caribou and sheep know these areas, but they are not easy for a
stranger to examine.

KOBUK 147

Family SYLVIIDAE: Old World Warblers, Gnatcatchers, Kinglets

Phylloscopus borealis kennicotti (Baird)

Kennicott’s willow warbler was designated by Paneak with an
Eskimo name Songakpalutungtk, meaning small bird the color of bile,
and he recognized it when I first showed him a specimen. Since Shel-
don applied the same name to a specimen identified as Dendroica pete-
chia amnicola, his recognition of both species is questionable. Paneak
considered the name Songakpalutungik explicit for P. borealis, which
greatly outnumbered the not so common PD. petechia and W. pusilla in
our experience at Anaktuvuk. We found willow warblers common
on the Ahlasuruk (L. Irving and Paneak, 1954), and Townsend
(1887) and Grinnell (1900) identified them along the wooded part of
the Kobuk. Although they are now widely known from the arctic
tundra, Grinnell’s report from Kobuk seems to give the only identifica-
tion of willow warblers in the arctic Alaskan woodland.

Regulus calendula calendula (Linnaeus)

When naming ruby-crowned kinglets, Sheldon spoke of their pres-
ence in winter at Kobuk. Eskimos at Anaktuvuk have reported king-
lets there in winter, and an Indian at Old Crow, Yukon Territory,
told me it was a winter bird there. The few dates when specimens
have been taken on the arctic coast were in the early and late winter
(Bailey, 1948). Although I think that the information of the native
people is correct, confirmation is needed before designating kinglets
as resident during winter in the arctic forest.

Family MOTACILLIDAE: Wagtails and Pipits

Motacilia flava Linnaeus

Although yellow wagtails have now been frequently collected north
of timber in the Brooks Range and on the arctic slope, no previous re-
ports have shown them on the wooded part of the Kobuk River. On
the arctic tundra wagtails are found about patches of willow brush
on marshy ground. Sheldon reported that wagtails were found in
such situations near Kobuk. The bird is so distinct and well known
to Alaskan Eskimos that Sheldon’s statement is evidence for its oc-
currence near Kobuk.

Family PARULIDAE: Wood Warblers

Among the six species of Parulidae reported by previous observers
on the Kobuk, only the myrtle warbler was explicitly named by Shel-
don in Kobukmiut. The yellow warbler and pileolated warbler are
common near Kobuk, but in our discussion I did not establish their

148 U. 8. NATIONAL MUSEUM BULLETIN 217

recognition as explicit kinds. At Anaktuvuk, where specimens of
D. petechia amnicola, D. coronata hooveri, and W. pusillus pileolata
have been obtained, no explicit Nunamiut names were found for
them.

An ornithologist may realize that because of his interest or expe-
rience, he is alert in the field to recognize species in some taxonomic
groups and slow in others. Personal and cultural experience develops
and may also suppress acuity of observation, which is much influenced
by the preparedness of the observing mind. The small warblers are,
to be sure, unobtrusive, but they are easily distinguishable in arctic
Alaska. In one year, Grinnell identified six species near Kobuk, and
during one week in August I saw five, while Sheldon, a native resi-
dent, scarcely seemed to distinguish three of these birds which had
surrounded him during his lifetime, and he knew a Kobukmiut name
which was explicit for only one of the six species. By way of con-
trast, Sheldon described and named 15 species of sandpipers, many
of them not obvious; whereas, so observant an ornithologist as Grin-
nell listed only 11 sandpipers.

Family FRINGILLIDAE: Grosbeaks, Finches, Sparrows, Buntings

Among the Fringillidae, Sheldon, like Simon Paneak at Anaktuvuk,
had a name for only one redpoll. Distinguishing the species of red-
polls requires criteria developed in the museum rather than in the
field. Grinnell (1900) did not report. gray-crowned rosy finches (LZ.
tephrocotis) which Sheldon described and named like the Nunamiut,
probably because even more than Oenanthe it keeps above tree line
in the mountain country that is occasionally searched for caribou and
sheep by the Kobuk and Nunamiut hunters. Grinnell identified
golden-crowned sparrows (Z. atricapilia), which Sheldon named, but
which apparently do not range north of the forests in arctic Alaska.
Except for this form and the crossbill, the arctic Alaskan Fringillidae
are equally birds of the tundra and of the bordering forest country.

Plectrophenax nivalis nivalis (Linnaeus)

A few snow buntings are seen in winter at Kobuk, where small
groups occasionally remain for several days about the village. Snow
buntings wintering in southern and western Alaska are geographi-
cally far separated from those wintering in the Mackenzie Valley,
but their differentiation in form has, so far as I know, only been
proposed by Salomonsen (1947, 1950).

EMAGARA Ate 9 tear mene a ete inne Seen caaiaiae AM etn SO.

“i eyepranich hemi A Ce 1 Pte Masta.

es :

Arctic Loon Gavia arctica pacifica (see pp. 31, 158).

4. Old Crow

oF HE GAP IN OUR INFORMATION about the geographical distribution of

life in the northern interior of Yukon Territory has been especially
significant because there we could expect to find a transition from the
specialized fauna of Alaska to the more generalized American fauna
of the Mackenzie Valley, representing the central part of the American
continent.

The Terrain

The Porcupine River extends the Alaskan Yukon Valley eastward
across Yukon Territory to within 50 miles of the Mackenzie River
and the arctic coast of Canada (see maps, figs. 1 and 7). Passes
through the British and Richardson Mountains from Old Crow River
Valley to the arctic coast are numerous and shorter and lower than
the passes through the Alaskan part of the Brooks Range. Macdougall
Pass through the Richardson Mountains, leading from the eastern
tributaries of the Porcupine to the Mackenzie watershed, is shorter
and, with an elevation of 1,200 feet, lower than any pass through the

149

150 U. 8. NATIONAL MUSEUM BULLETIN 217

western mountains of America. The Porcupine Valley thus forms
the eastern 250 miles of a corridor extending over thirty degrees of
longitude from the Bering and western arctic coasts nearly to the
central arctic coast and the great interior valley of North America.

The Porcupine River provided the first route for Canadian com-
merce between the Mackenzie Valley and interior Alaska. After
having established Fort McPherson in 1840 John Bell crossed the
Richardson Mountains in 1844 with Indian guides and traveled down
Bell River to the Porcupine and southwestward to its junction with
the Yukon (Osgood, 1936). In 1847 Alexander Murray traveled
the same course to establish Fort Yukon as a trading post for the
Hudson’s Bay Company, and in his journal (1910) vividly described
the Kutchin people, the animals, and the country. Related Kutchin
tribes lived on the Peel River, where they traded with Fort MacPher-
son, and on the Porcupine, upper Yukon, and Chandalar Rivers,
where Murray sought their furs in exchange for the scarce and costly
trade goods which his company had transported across the continent.
Sir John Richardson, one of the earliest biologists (1829) to evidence
his understanding of the geographical distribution of life over all
northern North America, quickly recognized the importance of
Murray’s location by soliciting from him information about the mi-
grations of birds. Murray’s reply, which Sir John quotes (1852),
provides in brief and vivid expressions the first recorded information
of the great spring migration of birds passing the upper Yukon and
lower Porcupine Valleys.

Travel along the Porcupine was then easier than through the moun-
tains by way of the Liard to the headwaters of the Yukon. In the
early times the Kutchin people were better disposed toward the white
fur traders than were the Indians near the head of the Yukon River,
where the Chilkoot Tlingit, with the intention of reserving the area
for their own trade, destroyed the early Fort Selkirk. Above the
junction of the Bell River with the Porcupine, Lapierre House was es-
tablished on the west side of MacDougall Pass as an intermediate sta-
tion between Fort Yukon and Fort MacPherson. There Robert Ken-
nicott visited briefly in 1860, 1861, and 1862 (Preble, 1908). In
September Kennicott traveled down the Porcupine to stay at Fort
Yukon until the summer of 1861. From his inspiring influence upon
the educated employees of the Hudson’s Bay Company there developed
that interest for natural history which produced collections upon which
most of our early knowledge of northern Mackenzie depends. One of
these, B. R. Ross (1861 and 1862), then in charge of the Mackenzie
district, was first to report observations on birds in northern interior
Yukon from Lapierre House, and he also gathered an extensive col-
lection. By 1900 steam transportation on the Yukon and traffic over

OLD CROW 151

White Pass had concluded the usefulness of the Porcupine route for
transport and it has remained unused except by the resident Indians
and an occasional traveler. In 1926 Olaus Murie visited Old Crow
in summer and banded some birds, and Otto Geist traveled along the
Old Crow and Porcupine Rivers in 1954 and 1955 surveying the
region for fossils. Otherwise there is no record of biological attention
to the Porcupine, although the river provides an easy course for
travel.

Physiography and Climate

At Old Crow the Porcupine River is about 350 yards wide between
cut banks about 15 feet high. Its current is swift but there are no
falls along its length, and for a light boat it is an excellent waterway.
At the ramparts along the Alaska Yukon border the Porcupine has cut
a gorge through the highlands between the Ogilvie Mountains and the
Brooks Range. Above the ramparts the river meanders through a
valley between bluffs about 100 feet high and from two to five miles
apart. At Old Crow the river has commonly cut from ten to twenty
feet below the level of its valley plain through layers of fine dark
sandy soil on top underlain with gravel, but sand and gravel layers
have been much intermixed by the floods and meandering of the river.
On this more or less flat valley white spruce is common, often mixed
with birch and some poplar. In wetter parts willow and alder often
form dense thickets. The valley between the bluffs looks like a flood
plain of the present river, which in fact now floods much of its area
in spring.

The bluffs north of the village are of a fine grained soil like loess
into which the present flood plain has been cut. In places the bluffs
are overgrown and in others still bare and eroding. A fragrant sage
is one of the plants which settles upon the slopes even before they
cease to slide. The bluffs are cut by steep ravines. Half a mile north
of the bluffs the land rises to about 800 feet above the village, which
is at an elevation of about 900 feet. Beyond this point a gentle rise,
with black spruce gradually replacing the white, continues for about
two miles to an altitude of about 1,600 feet, where the spruce forest
gives way to a half-mile-wide ascending band of willow and alder, be-
yond which is reached the tundra at the foot of the Old Crow Moun-
tains. These rise abruptly to about 3,300 feet on their southern side,
with some steep slopes and slides of the sedimentary rocks of which
they are formed.

To the northwestward the Old Crow Mountains are detached from
the Brooks Range by several low, wooded valleys. Forests also ex-
tend along the Old Crow River to the northern margin of Crow Flats,

469496—60——11

152 U. S. NATIONAL MUSEUM BULLETIN 217

but the tree line is low (1,600 feet) and the forest rather marginal
along the Old Crow.

At the relatively low elevations of the headwater valleys north of
Old Crow occasional spruce extends to the headwaters of the Firth
River, flowing toward the arctic sea. This is about the northernmost
occurrence of timber in America. The broad valley east of Old Crow
River and north of the Porcupine is occupied by many lakes in Crow
Flats, which occupy a depression among the mountains in arctic
Yukon east of the high peaks of the Brooks Range in northeastern
Alaska. From Crow Flats, however, only little drainage passes
through Old Crow River after the spring flood, for the annual pre-
cipitation is evidently small. Between the Yukon arctic coast and
Crow Flats the narrow Range of the Richardson Mountains extends
the Brooks Range eastward along the arctic coast and continues south-
ward along the Yukon-Mackenzie border.

The Porcupine River which rises in the northern watershed of the
Ogilvie Mountains and runs 150 miles northeasterly before changing
its course westward near the mouth of Bell River, has its southeastern
headwaters not far from those of Peel River. On the map, the
passage of the Peel through the southern Richardson Mountains ap-
pears to provide a valley route at low elevation by which birds might
enter the upper Porcupine Valley from the Mackenzie, while south-
ward and southwesterly the Ogilvie Range interposes what looks like
a barrier of high and extensive mountains separating the upper Por-
eupine Valley from the upper sources of the Yukon. The route to
northern Yukon Territory which seems to be open for birds migrating
at low elevations from the Alaskan part of the Yukon Valley extends
northeasterly along the Porcupine. The Porcupine Valley in Yukon
lies north of the Rocky Mountains in some isolation by surrounding
ranges through which, however, it provides lowland connection be-
tween the interior valley of Alaska and the Mackenzie Valley.

Warm weather comes to Old Crow a few weeks later than to Fort
Yukon and summer is colder. About June 25, 8 inches of snow brought
toboggans into use at Old Crow for two days. Since frost usually
occurs in every summer month there is little chance for gardens. The
summer weather at Old Crow is more severe than at Bettles, but is
probably not quite so cool as Anaktuvuk. Old Crow is evidently
just within the western arctic region where the close spacing of iso-
thermal lines (see fig. 2) shows the rapid northward cooling of summer
temperatures.

Of special interest in regard to the present geographical distribution
of life in Alaska and Yukon is the history of the recent glaciation,
which has left many clear marks, and the related climatic history,
which is less obvious. In Wisconsin times Cordilleran ice (Flint,

OLD CROW 153

1947) diminished in thickness north of about latitude 54° and probably
did not reach the Porcupine Valley. Massive though probably not
continuous glaciers occupied valleys of the Brooks Range, but much
of the Yukon Porcupine Valley system was free from continuous ice,
as was also the arctic slope of Alaska. Thus northern Alaska and
Yukon, while not covered with ice were cut off from habitable America
by ice which apparently reached the sea west of the mouth of the
Mackenzie and along the Pacific shores of Alaska.

We can speculate that hardy birds like those now resident in the
Arctic might have survived the last glacial maximum in arctic Yukon.
There is no doubt that the invasion of migratory birds has occurred
since their paths from the south have become free from ice. The
10,000 years since the last maximum of glaciation have had fluctua-
tions in climate and in the attendant condition of the land surfaces.
If migrations have been long in operation the birds must have had a
remarkably varying experience in their migratory routes. Not only
have the climatic conditions changed but the changes in populations
accompanying the opening and closing of great areas must have
exerted powerful and changing social influences. Through this period
of change, however, the essential features of the Rocky Mountains
and Brooks Range have continuously maintained a northern Yukon
and Alaska geographically and climatically distinct from the adjacent
regions. At present we have then the maximal duration of time of
development of modern migrations, which passed through areas of
great climatic and biotic change and which are now established in a
geographically distinguishable area.

Modern Indians seldom pass into the barren coastal mountains of
Yukon Territory, the traditional barrier and boundary between them
and the Eskimo people with whom they have long had hostile, or at
best unfriendly, relations. ‘There has probably been some exchange of
material culture but the distinction between Indian and Eskimo
ways is striking for neighboring people in similar climates. Among
names of birds, for example, Indian and Eskimo names for about 90
species known by both people show resemblance in only three cases
(Irving, 1958b), and I think that this instance about represents
their general resemblance in respect to intellectual culture. Even in
material culture it is surprising to see that their few natural resources
have been used in many cases with such differing implement and fash-
ions, and judging from what I have heard from them the blood of
the two races has seldom mixed (Lewis, Chown, and Hildes in 1959
reported that blood samples taken in 1958 from most of the Indian
population of Old Crow showed no serological evidence for inter-
mixture with Eskimos).

154 U. S. NATIONAL MUSEUM BULLETIN 217

The Indians at Old Crow

At Old Crow about 170 Indians form the only village now in the
territory of the Vanta Kutchin (Osgood, 1936; Leechman, 1954).
The nearest settlements are Fort MacPherson, Herschel Island, and
Fort Yukon, about 140 miles east, north, and southwest, respectively.
Southward about 300 miles are the settlements on the upper tributaries
of the Yukon. Old Crow had only three cabins and a store when
Neil McDonald came there in 1913 but many of the residents are
probably derived from the Vanta Kutchin of the Porcupine Valley,
who were once more numerous, Long geographically isolated, their
old people were nevertheless great travelers, the Porcupine being
an ancient trade route between the Yukon and the Mackenzie, In
winter now some of them travel great distances to hunt or trap, and
to work or visit at MacPherson and Fort Yukon. Recently the prices
of fur have been so low that little beside muskrats is taken, but
although their price is cheap the numbers taken in spring on Crow
Flats provide the principal cash support of the village.

The present chief, Charlie Abel, was elected in spring of 1958 to
succeed Charlie Peter Charlie, who had been elected in 1953. The
two former chiefs, each served for more than ten years, Peter Moses
from about 1940-1953 and Joe Kay from about 1925-1940. All have
demonstrated the high quality of strong leadership which seems to
have been common among these Indian people. In addition a detach-
ment of two Royal Canadian Mounted Police Constables administers
law with the skill and wise concern for the people for which that force
is noted. An Anglican preacher operates the church to which prac-
tically every villager belongs. A recently established Roman Catholic
Mission has not acquired an Indian following.

Disturbed only by a monthly mail plane from Fort Yukon, and
under the wise and strong guidance of their chief, the police, and the
preacher, the community seems to be a well ordered and a more co-
herent body than most Eskimo villages, in which the regulatory power
of the council is small and where families join to make a village on a
very informal basis of sociable inclination. For sustenance, some fish
are obtained in spring, but the caribou which cross the Porcupine in
spring and fall migrations provide the principal source of natural
food. For many years the caribou had usually been available, but
in the autumn of 1957 their migration passed so far west of Old Crow
that shortage of dog food reduced the range of hunting and prevented
preparation for trapping in the next spring season. The people rose
to this serious situation with increasing exertions at hunting but they
were severely handicapped by the reduction of their radius of opera-
tion through the impairment of canine transportation.

OLD CROW 155

The people are strong travelers, good hunters, and lively com-
panions. As they learned of our interest in birds they told us their
observations and brought us many specimens. Without their help
and advice we could have obtained little information. With their aid
our survey shows the status of the birds in their country, Living and
working with the Old Crow people has been pleasant and profitable
for biologists.

From former chief Joe Kay I obtained Indian names for 99 of the
107 species which we now identify in the avifauna of Old Crow
(Irving, 1958b). He wrote them for me in the orthography devised
by Archdeacon McDonald (1911) for an Indian version of the Book
of Common Prayer and Bible. Most of Joe Kay’s identifications were
from specimens, but some were made by reference to illustrations or
descriptions of appearance and habits. I did not meet another Indian
with such command of nomenclature. Furthermore, Joe Kay in any
company was an exceptional man for his remarkable strength and
ability. At about 75 years of age his wisdom and social understanding
were impressive to us and respected by his own people.

He remarked that even among the older people few could recognize
and name all birds. I believe that the names he gave me were true
designations in his language, and yet I marvel that an unwritten
language could persist in so small a community where only a few
individuals can have such intelligence and learning as is required to
preserve the comprehensive and accurate Indian knowledge of their
natural environment. It is also revealing to find among the Indians
and Eskimos who socially dominate their fellows by physical strength
and skill the possession of such intellectual attainment as is illus-
trated by perfect recognition and naming of birds. Here, in a small
community living among primitive conditions, we see how refined an
intelligence can be developed, for the cultivation of such knowledge
as deals with small birds demonstrates mental exertion for the pure
love of knowledge.

Progress of the Investigation

Leonard Peyton and I reached Old Crow on April 2, 1957. We
were kindly received and lodged for our first night by Constable
P. A. Robin. The next day we obtained from Albert Abel the use of
his cabin, To store our gear would require more shelves than he had.
When I asked if we might build some shelves, I remarked that per-
haps their absence from his cabin showed some special reason prevent-
ing their construction and observed that perhaps he would not want
them. He simply replied that he had not needed shelves because he
had nothing to put on them.

156 U. S. NATIONAL MUSEUM BULLETIN 217

Within a few days we engaged Robert Bruce to guide and help us
in hunting and he proved to be a strong and clever hunter and a pleas-
ant companion. His decision to work with us required some study,
for he had planned to leave with his family for the spring period of
hunting muskrats on Crow Flats. He did not enjoy the prospect of
separation from his wife and children and the busy life in the musk-
rat-hunting camp which all the people so much enjoy. Within a few
weeks every family in the village had left for their camps except a
few who were too aged or ill, and in spite of his cheerful nature Bruce
was often unhappy for his absent family. Edith Josie who helped
us at skinning birds was usually smiling or laughing and soon acquired
skill in the preparation of skins.

Leonard Peyton took over the arduous care of our collection of
specimens so that I might be more free for hunting and observing.
Usually I went with Robert Bruce, but Peyton also seized many occa-
sions for hunting. His careful preparation of our supplies and equip-
ment, diligent administration of their use, and attention to the col-
lections and records were a basis for the success of our expedition.

During April the weather was generally cold, with prevalent north-
easterly winds. On the river the snow was firm enough for travel
on skis or snowshoes or with dogs and toboggan. Where the trails led
through woods and up hills the slewing of the toboggans on the curves
made the snow rough for travel. Occasional westerly winds brought
warm chinook weather and melting conditions.

In May the sun was warmer and melting periods became more fre-
quent. We were able to travel by dog team on the river until May 6,
when we met elderly Paul Joseph returning from Fort MacPherson
as weary as his dogs. Thereafter the overflow on the river increased,
although until May 16 we could still cross by wading through the
overflow, or during a temporary freeze. Then the ice began to erode
rapidly, and after showers in the night of May 22 the water began
to rise, lifting the ice from its attachment to the shores, and the floes
began to move steadily during the day. During the next few days
the water rose about 15 feet, nearly to the top of the cut banks, and
the ice floes varied in concentration as they came in from various tribu-
taries. In June some days were hot and mosquitos too common, but
manageable by the use of spray and repellent. Brooks and swamps
filled, making it difficult to walk.

On May 10, Francis Williamson joined us for a month collecting
mammals. His keenness in recognizing birds by their calls enabled
him greatly to extend our information. On May 28 Sidney Peyton,
Leonard’s father and an experienced collector of birds’ eggs, and Webb
McElvey joined us. The latter was a taxidermist collecting for W. J.
Sheffler of Los Angeles, whose specimens we were very kindly per-

OLD CROW 157

mitted to study and report upon as our own. On June 4 Robert Whaley
flew in from Anchorage bringing Mary C. Lobban, a physiologist
from Cambridge, England, to make studies of the daily cycle of uri-
nary excretion among the resident people during the long arctic days.
For sleeping quarters and a laboratory, we utilized the old church, now
deconsecrated.

Under Leonard Peyton’s careful supervision 361 bird skins were
prepared and have been deposited in the U. S. National Museum.
Webb McElvey prepared 90 bird skins for the collection of W. J.
Sheffler. Sidney Peyton, recorded the nesting of about 25 species
of birds and collected 35 examples of nests and eggs, which are now
in his collection. The nesting data are used in this report. Frank
Williamson obtained 66 specimens of 16 species of mammals upon
which he will report. Leonard Peyton and Frank Williamson collected
a number of fishes which they have sent for identification. Mary Lob-
ban’s report on the cycles of urinary excretion and the daily rhythms
will appear in physiological publications. A number of hearts of
newly arrived and settled migratory birds and of resident birds were
preserved and sent to George Burch, professor of medicine at Tulane
University, for a determination of whether the migration to the arctic
and the sudden transition there to sedentary habits has caused signs
of stress to appear in their coronary structures.

The collaboration of these colleagues, with their wide range of ob-
servations and variety of viewpoints, has been most productive,
and I am deeply grateful for the companionship in which we enjoyed
together, as spring progressed, the view of the country and its rapidly
changing biological phenomena.

The Birds of Old Crow

Family GAVIIDAE: Loons
Gavia immer (Briinnich)

Common loons were identified by sight and occasionally by call.
The first certain identifications were made on May 28, when several
single birds, pairs, and groups of four were seen in the evening flying
directly up the Porcupine. After two groups had been noticed over
the village a few others were observed to arrive from the west and
pass beyond sight eastward. Other common loons were occasionally
noticed at the end of May, apparently in migratory flight from west
passing eastward. Judging from the direction of apparent migratory
flight of the common loons, they come from a population wintering
on the west coast.

158 U. S. NATIONAL MUSEUM BULLETIN 217

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Ficure 7.—Porcupine Valley in the vicinity of Old Crow. (From Canadian Dept. Mines and

OLD CROW 159

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158 U. §. NATIONAL MUSEUM BULLETIN 217

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OLD CROW

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159

160 U. S. NATIONAL MUSEUM BULLETIN 217

On June 18, five common loons were recorded, and about ten times
as many Pacific loons identified by sight along 15 miles of river just
east of Old Crow. John Buckley informed me that during a canoe
trip from Old Crow to Fort Yukon in August 1954 he had identified
common loons among much more numerous Pacific loons along the
river.

Although Bailey (1948) found few records of common loons along
the Arctic coast of Alaska, they are well known in the Arctic interior
and are more often seen in the central Brooks Range than yellow-
billed loons.

The Indians call them Ttretetere.

Gavia arctica pacifica (Lawrence)

1 female June 15 weight 1751 g. little fat ege 6mm,

1 male June 25 weight 1382 g. medium fat testes 10x19, 6x11
mm,

1 slightly incubated June 24 —_— —_ =

ege

The first Arctic loon was seen circling in flight over the village on
May 22, at the time when the ice in the river was breaking up. There-
after they were frequently seen on the river and pairs occupied the
small portions of open water as early as they appeared in many of the
lakes in the valley. Here it was apparent that nesting was in prepara-
tion. On the lake behind the village where a pair had been observed a
nest was found abandoned because the birds had been shot or driven
away.

On June 24 Irwin Linklater found a nest on a moss platform at
the tip of a point of land extending into a small lake. It contained
one slightly incubated egg which would have been laid about June 15.

As Robert Bruce and I were pursuing an Arctic loon on a small lake,
it emerged from a dive and, seeing the canoe nearby, dove suddenly
with a loud scream. Soswift was its dive when fired at that it actually
seemed to have submerged before the shot splashed harmlessly about
the place of its disappearance. On June 15, this loon was about ready
to lay.

In the latter part of May the majority of Pacific loons as single
birds or pairs appeared from the west and passed eastward, seemingly
migrating up the river and presumably coming from a population
wintering along western coasts.

During June many Pacific loons were seen along the river, where
they were occasionally calling, often associated in pairs. When
alarmed they quickly took wing but sometimes flew only short dis-
tances as if attached to the area. It was not apparent whether they
nested along the river, but they occupied many lakes.

The Indians name the Arctic loon Thulvit.

OLD CROW 161

Gavia stellata (Pontoppidan)

Red-throated loons were not recorded as satisfactorily identified by
sight or call in flight until June 2, which was probably later than the
first arrivals. These loons were less frequently identified than common
loons on the river and none were noticed on the small lakes which were
visited. Speciments were obtained in 1958.

Family PODICIPEDIDAE: Grebes
Podiceps grisegena holbollii Reinhardt

5 fresh eggs June 15 — — aes
4 slightly incubated June 23 os ad aa
eggs

The first evidence of red-necked grebes, which the Indians call
Tekkin, was the distant sight of the head and neck of one among the
Equisetum at the margin of a small lake on June 13. The bird had
disappeared when the spot was reached, but a nest was found on the
floating clods of mud from a muskrat push up, with three eggs lying
partially in the water. The eggs felt warm. Two days later two addi-
tional eggs were found in the nest, so that the first eggs must have been
laid about June 11. The other parent was seen, but neither could be
approached within a quarter of a mile. A second nest was found by
Irwin Linklater at a similar site in another lake. Red-necked grebes
were rarely seen on the river and those which were seen were shy.

Red-necked and horned grebes have now been reported near the
northern border of the forest at Old Crow and in central and western
arctic Alaska.

Podiceps auritus cornutus Gmelin

1 female May 19 365 g. very fat egg 4mm,

Two horned grebes were watched on the slough near the village
on May 19 and on the same day one was collected for us by Philippe
Dicquemare. The size of the egg which it contained suggested that the
female was approaching readiness to lay at a date earlier than eggs
of red-necked grebes were found. Horned grebes, known by the In-
dians as Wotsik, were occasionally seen along the Porcupine River,
where they were less shy than red-necked grebes and more frequently
noticed.

Family ANATIDAE: Swans, Geese, Ducks
Olor columbianus (Ord)

At Old Crow village Father Mouchet reported that he had seen
a swan flying up the river on June 3. Peter Lord and other local resi-
dents knew swans well and said that they had not seen many near
the village and only a few on Crow Flats, where no groups larger

162 U. S. NATIONAL MUSEUM BULLETIN 217

than six were recalled. Ben Charlie and Freddie Frost reported that
they saw two swan’s nests containing five eggs each at Crow Flats
this season. Joe Kay gave the Indian name of the swan as 7'arwi, and
the stories related to me showed it to have a regular place in Indian
tales concerning natural history.

Richardson (1852, p. 304) cites a letter written to him by Alexander
Murray, who established Fort Yukon, in which occurs an interesting
reference to swans in the course of a vivid description of the migrations
of birds in spring at Fort Yukon: “Of the two kinds of swan only
the largest sort are seen here; they pass on to the northward of the
Porcupine River to breed in the lakes there.” After Murray’s ref-
erence to swans, Richardson wrote in “Cygnus buccinator.” 'The ob-
server and his reporter command such attention for the accuracy of
their observations that we must believe that in about 1847 trumpeter
swans migrated to the Porcupine. Dall and Bannister (1869) re-
ported that Lockhart had found a set of trumpeter swan’s eggs near
Fort Yukon. It is to be hoped that it may turn out that trumpeter
swans still migrate to the Porcupine. This hope is encouraged by the
recent discovery of trumpeter swans nesting on the Copper River
(Monson, 1956) and accumulating evidence that a fair number nest
on the Kenai Peninsula.

Branta canadensis taverneri Delacour

1 female June 6 weight 1741 g. little fat 6 ruptured follicles
(brood patch)
1 female June 19 weight 2219 g. little fat egg, 5 mm.

Canada geese were first seen May 6. Until May 11 a few groups
of two or four birds each were seen. During this period about 30
geese were recorded. Most of those noticed on the wing were look-
ing about as if interested in landing. A number of them alighted on
the ice of the river, where their sleeping postures and the accumula-
tion of droppings showed that they rested for some hours before
leaving. Some of the Canada geese seen in this first period were
flying from the east but the direction was not certainly established.
By their inclination to rest they seemed to be arriving from a long
flight. There was rarely a sound from them.

From May 11 to May 18 single birds, twos, and groups of four and
six occasionally passed, sometimes honking. They now often appeared
flying from the west along the Porcupine River, but their flight was
less decisive of direction than in the first few days, and they often
circled the village and swerved to the howls of dogs and calls of men.
The river was only melted in superficial pools which froze at night, so
that it could be regularly crossed afoot until May 16. Although
brooks were now running, no lakes were open, the snow cover was con-

OLD CROW 163

tinuous, and only a few mud and gravel banks were exposed. We
could not see a source of food available to the Canada geese which
passed during the first ten days nor did we get any indication that
they fed nearby.

On May 18, three days before the ice began to run in the Porcupine,
the first large flock of 20 Canada geese was noticed flying eastward
in a V and honking. From that date until June 1 occasional flocks
formed in V’s were seen, with considerable unorganized traffic. The
formed flocks were only noticed flying eastward. Two Canada geese
were seen flying in formation with a V of nine snow geese, and this
mixture was seen again. This association seems to indicate the west-
ern origin of these Canada geese.

Two additional specimens were taken in 1958.

On June 3 and 4 two pairs of Canada geese appeared as if settled
on the river bars near Kenneth Nukon’s cabin. Francis Williamson
reported that Canada geese were settled along the river in numbers
increasing eastward from Nukon’s cabin to Driftwood River and
Indians reported that in summer geese are most concentrated along
the eastern Porcupine and lower Bell River. Williamson collected
the first female specimen near Driftwood River, where it had evi-
dently begun to lay about June 1.

On June 13 a few separate pairs were seen near Kenneth Nukon’s
cabin. A flock of nine which included several birds without the
adult sharpness of the black-and-white pattern on the head were
feeding in a flock which included some mature appearing birds. As
they flew away honking, two others joined them.

On June 17 a bird was wounded out of a flock of 12 which had
landed on the muddy bank of the slough near the village. Two days
later Robert Bruce collected it alone in a lake across the river. It
was a female with eggs 5 mm. long but showing no indication of
having laid.

Peter Lord reported the arrival of geese at Crow Flats on May 7,
one day after the first sight of them at the village, a schedule sug-
gesting that they came in the same migratory movement which first
brought geese to the village. But at Johnson Creek, a tributary en-
tering the upper Porcupine about 80 miles south by east from Old
Crow, Lazarus Charlie reported the arrival of geese on April 30,
eight days before they were seen at Old Crow. This discontinuity
of schedule suggests that either migration is markedly retarded in
that area, or that Johnson Creek is on a different migratory route.
The arrival at Johnson Creek was slightly later than the dates reported
in earlier years at Frances Lake and Sheldon Lake, about 300 miles
further south by east from Old Crow (Rand, 1946). Murray’s old
report (Richardson, 1852, p. 304) places the arrival of geese at Fort

164 U. S. NATIONAL MUSEUM BULLETIN 217

Yukon near the time of arrival at Johnson Creek and suggests that
different populations provide the first arrivals at Old Crow and
Fort Yukon.

We have assembled four specimens of Canada geese from Anaktuvuk
with Delacour and Zimmer (1951) ascribed to the range of B. c.
taverneri, and also two specimens collected on Colville River by Cade
and Schaller. The latter were on their breeding ground and the former
were migrants collected while 150 miles south of Colville. We have
also a specimen taken by Hughes and Chamberlain at College, Alaska,
May 7 with largest ova 22 mm. The two specimens from Old Crow
conform with the seven from central northern Alaska and differ
from four specimens regarded as B. c. parvipes, taken in May near
Lake Athabasca, Alberta, and one from the site of Old Fort Good
Hope on Mackenzie River. Their measurements are shown in table 3.

TaBLE 3.—Dimensions of Branta canadensis parvipes and Branta canadensis

taverneri
Measurements of culmen (mm.) Testis
Date | Sex Locality or egg Weight
; (mm.) (g)
Wing Nail Length | Depth
PARVIPES
May 26| o | Lake Athabasca 411 12 40 20
May 26} @ Lake Athabasca 409 12.5 36 18
Jun 28] co | Old Fort Good Hope 432 13 42 22
Jul 3] o& | Athabasca Delta 420 13 45 21.5
Oct 5{ co | Lake Athabasca E 410 12.5 43 21
TAVERNERI
May 7 9 College 389 12 37 20 22 2688
May 18 fof Anaktuvuk 422 11.5 33 22 6
May 18 2 Anaktuvuk 400 13 35 19 22
May 19 fof Anaktuvuk 405 12 35 20 22 2551
May 20 2 Anaktuvuk 373 11 36. 5 19
Jun 6 fe) Old Crow 390 il 39(?) 19(?) 16 1741
Jun 10 2 Colville 394 9 31 19.5 9 2250
Jun 19 9 Old Crow 412 11 39 21.5 5 2219
Jun 20 2 Colville 387 10 30 18.5 6 2250
1 Laid.

The geese listed as taverneri have the small nail and short bill
characterized by Delacour. Since the bill is as deep at the base as that
of the longer billed geese, the angle formed by the top and bottom
of the mandible is relatively steep. The breast of taverneri is generally
darker, forming a sharper contrast with the white belly. The back
of tavernert is darker. It is especially dark at the transition from

OLD CROW 165

neck to back, where the darkness of taverneri is a well marked distinc-
tion from the four birds called parvipes.

Several specimens in the U. S. National Museum, collected by Olaus
Murie in 1926 along the Old Crow River, are regarded as J'averneri.

It appears that the specimens of geese of north central Alaska and
Old Crow can be included and distinguished as taverner. Except for
the specimen from College, Alaska, which on May 7 was probably in
migration, these specimens are in the breeding range ascribed to them
by Delacour. One of the geese from Old Crow was a nesting bird, and
Kessel and Cade (1958) obtained downy young on the Colville.
In addition we have obtained six immature specimens from the
Koyukuk and Alatna Rivers, where family groups were observed,
and one male in breeding condition from Kobuk. These specimens
appear to have the characters of taverneri and occupy the breeding
range ascribed to tavernert by Delacour.

Bailey (1948) distinguished under lewcopareia a dusky goose nesting
in Arctic Alaska west of Barrow, inland from the coastal range of
the very small minima, and approaching the coast east of Barrow.
The measurements resemble those later ascribed to taverneri and are
less than those of the northern interior geese which would be in the
range ascribed to parvipes (A. O. U., Check-list, 1957). Bailey also
pointed out the dusky character of the arctic Alaskan geese, which
is matched by our specimens.

Our two specimens from Old Crow do not establish that the small-
billed dusky goose is the only form nesting and migrating along the
Porcupine. Mary Lobban reported that she saw a small goose calling
in a manner only ascribed to minima.

During migration we gained the impression that Canada geese were
traveling eastward and westward although a greater number were
headed eastward.

The Indian name for the goose is Kyha.

Branta nigricans (Lawrence)

Only one flock of about 60 brant was noticed flying swiftly east-
ward close to the Porcupine River on May 25. They are, however,
well known to the local Indians. As late may migrants they are seen
passing eastward along the Porcupine at Old Crow, and more often on
a northerly and northeasterly course over Crow Flats. Cade (1955)
collected reports, mostly from older records, showing that numerous
and large flights of brant late in May pass through the interior of
Alaska. In part the brant pass Old Crow after coming up the Porcu-
pine, apparently from Alaska. None are known to summer on the
Porcupine and Old Crow Rivers (Murray, in Richardson, 1852, p. 304),
but there are numerous summer settlements of black brant on the

166 U. S. NATIONAL MUSEUM BULLETIN 217

islands off the Mackenzie Delta (Porsild, 1943), and for a short dis-
tance eastward along the Arctic coast (Snyder, 1957).

At about the time when black brant were named from a specimen
taken at Egg Harbor, N. J. (Lawrence, 1846), Alexander Murray,
the venturesome and observant founder of the Hudson’s Bay Com-
pany Post at Fort Yukon wrote to Sir John Richardson (1852, p.
304), “there are also black geese which I presume you have never
seen,” and gave a description which by Sir John’s comment “applies
pretty well to the brent goose.” Murray added that the “black geese”
breed only on the shores of the arctic sea.

Preble (1908, p. 308) wrote, “though keeping strictly to the seacoast
east of the Mackenzie during migration, many of the fiocks (probably
all of the eastward breeding birds) strike across Alaska from near
the mouth of the Mackenzie to the North Pacific.” The Indians re-
marked that brant were not seen near Old Crow on their return, and,
in agreement with Cade’s (1955) survey, I have not been able to obtain
reports on the return of brant across interior Alaska. Since Murray
observed that white and black geese returned in September and Octo-
ber flying high, Preble’s view of an overland return from the Mac-
kenzie is confirmed. Murray also observed that a few black brant
“pass down Peel’s River,” and the Porcupine may not be the only
overland course by which the western black brant pass from the Pacific
coast to the Mackenzie Delta and eastward.

The Indian name for the brant is Ttsun tratesit.

Anser albifrons frontalis (Baird)

We did not distinguish many or large flocks of white-fronted geese
migrating through Old Crow, but those we saw were flying eastward.
The first flock of seven was seen on May 16. Reports of the Indians,
who call this bird 7echyo, indicate that some white-fronted geese nest
around the lakes on Crow Flats, but we did not get certain informa-
tion as to whether they nest along the Porcupine.

In 1867-8 Dall (Dall and Bannister, 1869) found white-fronted
geese arriving at Nulato about May 6-10. In 1849 Murray wrote to
Sir John Richardson (1852, p. 305) that they arrived at Fort Yukon
from April 27 to 29 and nested on the lakes “north of the Porcupine”
where they are “more numerous than any other kind; and the number
that pass northward there are perhaps equal to that of all the other
species together.” White-fronted geese are still numerous in migra-
tion at Anaktuvuk and Kobuk about May 15. Rand (1946) reports
the banding of one by Olaus Murie at Old Crow as the only record in
Yukon Territory south of the arctic coast. From these reports and
our rather few observations it seems that the spring migration of
white-fronted geese along the Porcupine River represents the east-

OLD CROW 167

ward extension of the population which migrates northward along
the coast of British Columbia and “much less commonly in the in-
terior” (Munro and Cowan, 1947).

Chen hyperborea hyperborea (Pallas)

The first flock of nine snow geese, seen on May 20 included in its V
formation two Canada geese. This association of two species of geese
is often seen in arctic Alaska. Only a few flocks of snow geese were
recorded at Old Crow, all in V formation and flying steadily eastward.
It is reported by Indians that snow geese, which they call Hookeh,
occasionally land on Crow Flats, where they are more often noticed
than at Old Crow. Peter Lord reported that he first saw snow geese
on the Flats on May 16, but that they always pass northward and do
not remain in summer. The size of the flocks of snow geese and the
numbers sighted vary from year to year at Anaktuvuk and from
reports of the Indians, a similar variation appears to occur at Old
Crow.

Dall (Dall and Bannister, 1869) found snow geese common in spring
at Nulato, where they arrived on May 9 flying up the river. Rand
(1946) gives no reports on snow geese in southern Yukon, and Swarth
(1936) saw only three at Atlin—two on May 5, 1932, and one on
May 5, 1934. It seems likely that the eastward flight at Old Crow
brought snow geese from the Pacific Coast. Those which we saw
were flying rather low and their numbers would not make a substantial
contribution to the numbers nesting from the Mackenzie eastward.
But since these geese often fly high over mountains we may have
missed seeing most of the migration.

At Fort Yukon, wrote Murray to Richardson (1852, p. 305), the
snow geese arrive about May 15 or 16 and “breed only on the shores
of the Arctic Sea. They return in September and early October, flying
high and seldom halting.”

Subfamilies ANATINAE and AyTHINAE: Ducks

Of the 14 species of ducks at Old Crow, two have holarctic nesting
areas, two are not quite so widely distributed in the Arctic, and 10
are restricted in arctic nesting to Alaska, Yukon and Mackenzie.
East of central Mackenzie the northern border of nesting by these
species extends southeasterly to the southwestern shore of Hudson
Bay about parallel to but usually somewhat north of the forest. The
northern limit also runs about parallel to the steeply declining iso-
thermal gradient of summer temperatures which marks the extension
of summer warmth far northward in western North America. East
of the Mackenzie River the physiographic features also change, for

469496—60——12

168 U. S. NATIONAL MUSEUM BULLETIN 217

eastern Mackenzie and Keewatin, which extend over the Barren
Grounds and the pre-Cambrian shield, are not mountainous, in contrast
with the rugged terrain of Yukon and Alaska. There are thus changes
in the general climatic, geological, and vegetational conditions to
correspond with the northern limit of nesting.

Among 28 migratory species of North American ducks only three
are now divided into races (A. O. U., Check-list, 1957): the blue-
winged teal, with an Atlantic race of separate range; the common
elder, in which the maritime nesting range of the species is easily
separable into separate ranges for each race; and the white-winged
scoter, the races of which winter in widely separated maritime ranges
on either side of the continent.

White-winged scoters and the five other species of ducks most
common in migration at Old Crow showed the trend of their flights
to be eastward along the Porcupine in spring. On account of their-
western wintering location, goldeneyes (Bucephala) are also suspected
of migrating eastward. About 100 miles east of Old Crow the Rich-
ardson Mountains bound Mackenzie and Yukon, but they are low
and provided with short passes between the easternmost marshes of
the Porcupine Valley and the western marshes of the Mackenzie
Valley. The eastward trend of migratory ducks on the Porcupine
appears strong enough to lead them to contact with or even to inter-
mingling with, the populations migrating and nesting in the northern
Mackenzie Valley.

In addition to the six species observed and one estimated to migrate
eastward to make contact with the ducks nesting in Mackenzie, lesser
scaups nest so numerously along Porcupine Valley that their breed-
ing area is continuous from Alaska to Mackenzie. Greater scaups were
not recorded from southern Yukon (Rand, 1946), and we are not sure
of their abundance along the Porcupine Valley. But some are there
in summer and the Alaskan breeding birds are connected with the
Mackenzie nesting ground by at least a scattered nesting population
in a narrow band through northern interior Yukon. Barrow’s golden-
eyes are now known from all over Yukon south of the Arctic coast,
but American goldeneyes, like greater scaups, appear to form the
connection between Alaska and Mackenzie only in the north along
Porcupine Valley.

All the observed trends in migration of ducks at Old Crow were
eastward, and on grounds of propinquity the Alaska and northern
Yukon nesting ducks are suspected of arriving in spring from the
western parts of their wintering grounds. Spring migration and
nesting appear to separate, although not to isolate the far northwestern
population genetically so as to form geographic races. Only two
species of ducks are acceptably differentiated as races and therefore,

OLD CROW 169

presumably, genetically distinct, the others apparently containing
inheritance mixed from many localities by the wandering of indi-
viduals during the time before and after nesting.

Anas platyrhynchos platyrhynchos Linnaeus

1 male May 19 weight 1063 g. fat testes 18 x 48 ,23 x 50
mm.

1 male summer —_— medium fat —_—

1 female summer —_— medium fat -—

Mallards were first reported at Old Crow by Stephen Frost, who
saw a flock of ten land early in the morning of May 8 just across
the river in one of the few grassy sloughs of water then open. A small
female mallard was shot that morning on the ice of the river, where
it had alighted with two pintails. During the next week singles, pairs,
and small flocks were often seen. Some flew intently eastward, but
more circled as they flew up the river and searched for any of the few
small pools of water in which to land. Occasionally landing on ice,
they chose water more often than the pintails, which seemed to have
no aversion forice. At first the mallards were often with pintails, and
later both mingled freely with widgeons.

In the first week of migration practically all of what little water
lay on the ground was underlain with ice, and on several days of cold
there were no unfrozen pools. During this week we saw no signs that
food was available for ducks. Occasionally mallards, like pintails,
settled to stand resting on the ice or to float on pools over the ice.
The birds which alighted seemed ready to rest, but several that were
examined were fat.

The trend of migratory flights of mallards at Old Crow was up the
Porcupine River. During the first two or three days mallards were
almost half as numerous as pintails. Since they were not in well or-
ganized flocks the arriving mallards gave an impression of being nearer
to the end of their migration than the better organized flocks of pin-
tails. After the first week the numbers of other migrating ducks con-
tinued to increase, but the main migration of mallards was over.

Evidently the mallards at Old Crow are not so near the end of their
migration as at Anaktuvuk, where scarcely any but singles, and pairs
are seen. The numbers of mallards observed passing Old Crow did
not seem sufficient to populate the eastern and southern areas of Por-
cupine Valley.

On May 19 singles and pairs were flying locally and during the next
week males were often seen flying in pursuit of loudly quacking fe-
males. On June 7 a nest was found with three eggs, the first estimated
to have been laid on June 5. At that date a female mallard might
occasionally be seen swimming on the lakes .

The Indian name for the mallard is Vatakcho.

170 U. S. NATIONAL MUSEUM BULLETIN 217

Anas acuta Linnaeus

Pintails were the earliest of the ducks to be seen, arriving on May
7. They were first noticed in the morning flying up the Porcupine
River in small bands. Most of the early arrivals that morning seemed
to be searching for a landing place. They landed readily on the ice
or near shallow pools on the ice and sat quietly resting or sleeping.
There was apparently no place attractive for feeding and so the early
flights only stopped for a rest before continuing on in a generally
eastward movement. The next day the largest flock of pintails no-
ticed during the early season contained 30 birds, but for the first week
most flocks appeared rather casually assembled and often included mal-
lards. The number of individual ducks and small detached groups
gave the impression that they were looking for a nearby location in
which to settle, but the places open were too few and too rapidly chang-
ing in the alternate thawing and freezing of the overflow on the river.
In these groups of pintails the proportions of adult males, females,
and young varied. Often an adult male was the most wary and the
first to lead the group on the wing.

On May 17 the flights appeared to increase in size and orderliness
of formation. One well organized flock of 80 contained only pintails
and flew directly eastward. After a few days this more intensive
migration seemed to subside and there was more association with
groups of mallards and widgeons. Pairs were occasionally observed
about May 30, but migrating flocks of 20 or 30 pintails continued to be
seen after the mallards and most flocks of widgeons had separated.

The flights of pintails past Old Crow seemed to be numerous and
well organized enough to account for more than the summer popula-
tion, which is rather scattered along Porcupine River. This suggests
that the Porcupine River is an important route for the entry from
Alaska of many pintails, some of which pass on to the arctic coast and
Mackenzie Delta. The early arrivals (May 10, 1934, at Kittigazuit)
reported by Porsild (1948), could be these early pintails passing
through Old Crow and bent upon an easterly course.

The Indians call this duck Chinchityo and Nakostikyi.

Anas carolinensis Gmelin

1 male May 17 weight 307 g. fat testes 12x29, 12x27
mm.

1 male June 15 weight 286 g. little fat testes 12x31, 16x29
mm.

The first green-winged teal noticed were a pair seen on May 17.
For a week singles and pairs were common and males were soon fol-
lowing in flight after calling females and in small ponds teal were

OLD CROW L741

persistently splashing as they quarreled. After May 24 an occasional
female was seen swimming alone in marshy pools as if settled there
for nesting, but on June 1 two pursuit flights were seen to indicate
that courting continued. On June 15 about a dozen males were peace-
fully scattered on a small lake, no longer disturbed to jealousy by
impulses of sex, although the testes of a specimen had not regressed
in size. There were no females in view and the males flew away as if
unattached to the locality.

The combative nature of male teal is illustrated in an Indian story.
Once Swan declared himself, as the largest of the birds, their chief.
In order to make his position understood he called all birds to appear
before him and defeated each species in combat. At last only Teal
had not entered the contest, or demonstration, and when he was sum-
moned he appeared with a bandage stained with old blood over his
head and said that he was not in fit condition to fight. As Swan in
sisted that Teal, like every other bird, must take his turn, with ap-
parent reluctance Teal said that if he must fight he would do his best.
In a short struggle Teal knocked Swan to the ground. Swan claimed
he had not been honestly beaten and attacked again. This time Teal
threw Swan down and only released him after he had admitted defeat.

Teal nest in Arctic Alaska, Yukon, and Mackenzie. In Alaska
they nest north of tree line at Anaktuvuk, but they are rare on the
Arctic coast (Bailey, 1948).

The Indian name is Z'arui kaka.

Mareca americana (Gmelin)

American widgeons were first identified by one of a pair that
was shot on May 17. Some probably arrived earlier, but the
early migratory flight contained mainly pintails and mallards. From
May 17 to 19 some flocks contained 20 or 80 widgeons; other flocks
mingled widgeons, mallards, and pintails in various proportions.
Many of these flocks of mingled species appeared from the west,
settled temporarily and flew eastward as if organized for migrations
together. Other groupings of mingled species were formed when
scattered ducks accumulated by alighting at a favored or already oc-
cupied resting place. Individuals or groups seemed to detach them-
selves readily from these associations. All the mixed flocks seen after
about May 20 were apparently loose and temporary associations. As
with the pintails, flocks containing only widgeons were better syn-
chronized in action and usually held to decided easterly course.

The first established pair was noticed on May 30, and many were
seen after that date along the river, but there was no time during
June when small flocks might not be seen. The number estimated to

172 U. §. NATIONAL MUSEUM BULLETIN 217

be settled in the Valley, mostly seen along the river, was slightly less
than the number of scaup. The obviously migrating widgeons were
estimated as much fewer than the migrating flights of pintails and
white-winged scoters.

Porsild (1943) reported widgeons as “perhaps the most common
duck in the Mackenzie Delta.” They nest commonly across Arctic
Yukon, but not on the Arctic coast (Rand, 1946), and across interior
Arctic Alaska. The Indians know them as Xaloree.

Spatula clypeata (Linnaeus)

Shovelers were recognized by the Indians from illustrations, and
Joe Kay described the duck and named it Tetrik. Its description was
also recognized by several other Indians, who usually see shovelers in
summer on the lakes of Crow Flats. They are known to Eskimos and
Indians across arctic Alaska and Yukon, but rather by their unusual
appearance than for commonness. According to Porsild (1943) the
Eskimos said that shovelers nested in the Mackenzie Delta, and
Indians reported them nesting on Crow Flats.

Aythya marila nearctica (Stejneger)
1 male June 13 weight 917g. medium fat testes 11x31, 12x29
mm,

Although a number of the scaup seen were thought to be A. marila
only the specimen taken was identified satisfactorily by sight. Since
its testes were only about as large as those of teal, they were probably
less than breeding size, but the date was late enough for some regres-
sion to have occurred. Joe Kay recognized the specimen and dis-
tinguished it by name as Jani cho; We believe that greater scaup
are regularly present in summer and probably nest along Porcupine
River, but that they are uncommon in comparison with lesser scaup.

Porsild (1943) reported the greater scaup an “infrequent summer
resident and no doubt breeding” in the Mackenzie Delta. Rand (1946)
reported only one specimen from Yukon taken by Nelson at Fort
Reliance, and referred to Swarth’s (1927) opinion that a few sight
records from southern Yukon should not be accepted as distinguishing
marila, from affinis. Greater scaup are probably not one tenth as
frequent at Old Crow as lesser scaup, but the proportion at Anaktu-
vuk is about reversed. Judging from the few reports of greater
scaup in southern Yukon and on the arctic coast, the Porcupine is
the important arctic connection between the large Alaskan and
Mackenzie Valley nesting populations.

During September 1958, of about 25 scaup shot and examined at Old
Crow only one was A. maréla, the others were affinis.

OLD CROW 173

Aythya afjinis (Eyton)

1 male May 22 weight 640 g. medium fat testes 8x21, 9x18 mm,
1 female May 22 weight 613 g. little fat ——
8 slightly incu- June 2 —_ -_ —

bated eggs

On May 18 the male of a pair of scaup near the village was identified
as A. affinis by its purple head, but some seen a few days earlier were
thought also to be lesser scaup. Occasionally scaup alighted in
pools with mingled species of ducks, but they did not join with other
species in organized flights. In the few days just before breakup
occasional small flocks and many individuals and pairs of scaup
were seen. Only a few individuals could be positively identified as
lesser scaup and the directional trend of their flights was not deter-
mined. By the time of breakup several pairs of lesser scaup had been
identified by sight among the considerable number of pairs seen in
the partially open water of small lakes. The two specimens were
flying together when they were obtained for us by Father Mouchet.
The testes of the male specimen were only the size of a teal’s and
were probably immature.

Scaup were seen paired about as early as were mallards and teal,
but the pairs of scaup remained longer evident as they swam on the
lakes, while pairs of mallards and teal did not long appear together.
On June 13 the last demonstration of pairing appeared when a male
lesser scaup rushed away from a female to attack a surf scoter which
had been shot nearby. On June 15 there were about ten male lesser
scaup scattered about on a lake with no females in view and the males
flew about singly as if not attached to a particular locality. The re-
formation of flocks of male scaup was not noticed during June.

On June 28, 1957, Irwin Linklater found eggs in a nest on top of
a moss and willow hummock 50 feet from the shore of a small lake.
Estimating about three days of incubation, the first egg was laid on
June 18.

As evidence for late nesting of scaups, of a group of 4 flightless
young scaup 2 which were shot on September 2, 1958 had only pin-
feather primaries and were about half the adult weight.

Although Rand (1946) had found reports on lesser scaup only
from southern Yukon it is evident that the Mackenzie and Alaskan
birds are connected by continuous nesting across Yukon as far north
as Porcupine.

The Indian name is Vityitin.

174 U. S. NATIONAL MUSEUM BULLETIN 217

Bucephala clangula americana (Bonaparte)

1 male June 13 weight 1024 g. fat testes 11x18, 9x13 mm.

The single specimen of American goldeneye was shot from a flock
of 25 flying over the river. The testes appeared to be less than breed-
ing size. During the hour about the time when the specimen was
obtained a flock of this same size was observed three times as we
traveled along the river and we suspected that these were sights of
the same flock. In it we distinguished male birds to form the majority,
but we could not determine that there were no females. In our rec-
ords a pair of goldeneyes was reported on the lake back of the village
and three were reported seen on the river on June 14.

Because of our unfamiliarity with goldeneyes and their reported
rarity in Yukon (Rand, 1946) we were at first reluctant to accept
reports by Indians that they had seen goldeneyes. Robert Bruce
recognized the specimen when it was taken and Joe Kay named it
with an explicit Indian name, Zovi. Indians had spoken of seeing
goldeneyes at least as early as May 21 and we had been informed that
goldeneyes were rather common on Crow Flats.

We were also reluctant to accept reports of the presence of golden-
eyes because we rarely found in the extensive forest the dead mature
trees which could provide holes for nesting. This was in part because
dead spruce are harvested for fuel along traveled routes, but even away
from lines of travel mature trees do not appear to stand long after
they die.

Only one other report of an American goldeneye comes from Yukon,
a specimen from Forty Mile described by Grinnell (1909), but Swarth
(1936) found them regularly in small numbers at Atlin, B.C. Nelson
(1887) remarked upon Dall’s (Dall and Bannister, 1869) finding
American goldeneyes wintering on the Aleutians and passing along
the middle Yukon among the earliest migrants. It was the first
duck killed at Nulato, on May 8, 1868. According to these migra-
tory schedules the goldeneyes on Yukon came from the western winter-
ing grounds. Those on the Porcupine River might also have come
from the west. American goldeneyes are common in summer on the
Mackenzie Delta and have been reported first arriving on the Peel
Branch on May 20, 1934 (Porsild, 1948). Preble (1908) reported
American goldeneyes first seen at Fort Simpson on April 26, and
there are many records of them in that part of the Mackenzie Valley,
from which also migration to Porcupine Valley appears an open pos-
sibility. Past failure of our observations to distinguish the abun-
dance and migratory course of the two species of goldeneyes caused us
to miss valuable evidence bearing upon geographical distribution.

In September 1958 we identified several American goldeneyes which
had been shot, and thought that some of the numerous migrating

OLD CROW 175

flocks were also of this species, but examples of Barrow’s goldeneye
were more numerous in hunters’ bags and in flocks identified by sight.

Bucephala islandica (Gmelin)
1 male June 14 weight 914 g. medium fat ——

Philippe Dicquemare kindly provided us with the specimen of Bar-
row’s goldeneye, listed above. Joe Kay distinguished it from the
specimen of American goldeneye, and named it differently, 7’esttet ky.
The testes had been destroyed by shot, but it was in breeding plumage.
Kay’s familiarity with the two species indicates that both are known
on the Porcupine. Rand (1946) reported Barrow’s goldeneye as a
fairly common summer resident in southern Yukon. Kessel, Murie,
and Schaller, in an unpublished survey of the birds of the Sheenjek
River, reported a sighting of Barrow’s goldeneye on the Sheenjek
River in 1956, probably the northernmost record in western America.

Clangula hyemalis (Linnaeus)

Oldsquaws were not seen until just as the ice began to move in the
river on May 22. On May 23, Father Mouchet and Philippe Dicque-
mare provided us with two males and a female which had been shot
while together. During the next week small flocks were frequently
seen flying eastward over the river. Usually the flocks of oldsquaws
were not high and flew purposively, like the migrating scoters, and
unlike the frequently deviating courses of migrating flights of pin-
tails, mallards, and widgeons. Individuals and pairs of oldsquaws
were often seen during late May on the lakes and river.

On June 4 there was much crying and maneuvering among several
male oldsquaws attending females on a small lake. Males which
intruded upon the pairs were vigorously repelled. During their noisy
courtship the males lost their caution and allowed close approach by
land or canoe. The preoccupation of both sexes with mating made
them the least wary of the ducks. Until June 15 oldsquaws on the
lakes demonstrated little concern for anything but breeding. During
the following week several small flocks of oldsquaws were noticed
along the river.

A comparison of the density of the local summer population of old-
squaws near Old Crow with the intensity of eastward migration sug-
gests that the migratory flights would not much more than provide the
Porcupine and Old Crow Valleys with their summer population.
Rand (1946) mentions no observation of oldsquaws in Yukon south
of the Arctic coast, so the migration through Old Crow cannot have
traversed southern Yukon and probably came from the Alaskan Yukon
Valley. Since oldsquaws are said by Dall and Bannister (1869) to be

176 U. S. NATIONAL MUSEUM BULLETIN 217

rare along the Yukon, it is to be suspected that they cross the Alaska
Range from the Gulf toward the upper Yukon and Porcupine.

The onomatopeic Eskimo name for oldsquaw is close to the Indian
name, Ahaluk. It and the name for great gray owl are the only two
in which I can find resemblances between Old Crow Indian and
Kobuk (forest-dwelling) Eskimo names for 90 species of birds known
in both areas.

Histrionicus histrionicus (Linnaeus)

1 male May 29 weight 582 g. very fat testes 11x21.5, 9x14.5
mm,

We are indebted to Philippe Dicquemare for bringing us the harle-
quin he shot on the river near the village. It was our only sight of this
duck, but Joe Kay, who named it 7'sz tut kwilik, said it was familiar to
him in summer on the mountain streams. The testes were approaching
breeding size.

In length of wing, depth of bill, and length of tarsus we could see
no distinction between western and wintering harlequins. The four
specimens from Anaktuvuk and one from Old Crow cannot be assigned
to a racial division.

Melanitta deglandi (Bonaparte)

1 male June 1 weight 1564 g. very fat testes 8x17, 8x18 mm.
1 male June 14 weight 1649 g. medium fat testes 11x26, 12x30 mm.
1 male =— weight 1663 g. medium fat testes 10x20, 9x25 mm.

The earliest white-winged scoters were seen on May 20. In the next
10 days more flocks were seen each day. During this period none
were recorded as containing more than 30 scoters and they often in-
cluded a minority of surf scoters. These flocks regularly appeared
from the west and flew steadily eastward, usually 30 or 40 yards above
the river, in rather compact and little changing order. The main
migration of scoters was later than those of pintails, mallards, and
widgeon. Migrating flights of white-winged scoters were even
steadier on course than flights of oldsquaws.

Until June 15 flocks of from 10 to 30 white-winged scoters were not
uncommon, but no trend in the direction of their flights could be seen
after June 5. On June 3 I watched several groups of six or eight
scoters, mostly white-winged as they floated down river for about a
mile with the current and then flew upstream to repeat the course.
Pacific loons and an occasional red-throated loon floated nearby with-
out apparent interspecific interest.

On June 15 males and females appeared associated in pairs. The

OLD CROW 177

female usually dove on the splashing approach of the male, who fol-
lowed her under water. Each female remained in one part of the lake
and we did not observe them to go on shore, but the males occasionally
left the area. Among the three male specimens, the late increase in
size of testes was consistent with the late manifestation of breeding
activities.

The Indians, who call this duck Vya, say that male white-winged
scoters thrash about noisily in furious combat on the brushy shores, at

- which time they are heedless of the approach of a man or canoe. In
this state the males seem to be more preoccupied by combativeness than
male oldsquaws. Although male teal are regarded as redoubtable
fighters during the mating period they and most ducks only reduce
but do not abandon their alertness for other dangers.

White-winged scoters are conspicuous because of their size and be-
cause they remain in the open waters until their late nesting date.
There may be little difference in the numbers of white-winged scoters,
widgeon, and scaup nesting along the Porcupine, but the scoters are
more easily seen, and during their prolonged migratory passage the
steady succession of flights must account for the passage of an enor-
mous number toward the east and northeast. It would be difficult to
speculate upon whether these flights past Old Crow importantly con-
tribute to the summer populations of Mackenzie, but they appear to
be at least sufficient to occupy the Porcupine Valley as far east as
Richardson Mountains.

Swarth (1936) remarked upon the sudden arrival from the coast
of large numbers of white-winged scoters at Carcross May 24-26.
If that date is typical of their first arrival, those reaching Old Crow
before that date did not migrate through Carcross. I have seen
scoters during migration through the upper part of the Copper River
Valley, which is still south of the Alaska Range, but have found no
reports confirming the overland course from the Gulf of Alaska which
would bring them into the upper Yukon Valley. The regularity of
their flights up Porcupine River indicates that scoters reach arctic
Yukon from the population which winters along the Alaskan coast.

White-winged scoters seem to be the only species of migratory ducks
distinguished into eastern and western races (A. O. U. Check-list,
1957). Brooks (1915) named the western race dtxoni because he
found them to have shorter and broader bills. During the breeding
season the bills of scoters become more highly colored and the softer
parts, in the males, at least, seem to swell. When dry the bills of
scoters warp, and it is then difficult to compare their shape. We

178 U. S. NATIONAL MUSEUM BULLETIN 217

have made the following measurement of length of bills with some
reservation as to their significance as dimensions:

Length Length
Eastern race (mm.) Western race (mm.)
Mar. 7 Woods Hole 44 May 8 St. George Island 43. 5
Mar. 7 Woods Hole 40 May 22 Nushagak 38
July 3 Athabaska Delta 36 June 1 Old Crow 42
Aug. 3 Rupert House 36 June ll Tuluak 36
Dec. 9 New York 40 June 14 Tuluak 37
June 14 Old Crow 40
(aver.) 39 June 14 Old Crow 41.5
June 14. Old Crow Al
July 20 Alloknagik Lake 41
(aver.) 40

The length of bill does not distinguish these eastern and western
specimens.

It may be remarked that the type specimen was taken at Humphrey
Point, Alaska, where scoters are rare. As far as I know, scoters do
not nest along the arctic coast in Alaska (Bailey, 1948), in Yukon
(Rand, 1946), or in the Mackenzie Delta (Porsild, 1943). The type
specimen was taken from the northern limit of its range and not from
a nesting area. Distinction of an eastern and western race would
imply some isolating process in the habits of white-winged scoters
in contrast with the apparent intermingling within other species of
ducks at some time of year, a condition which prevents localization
of genetic forms.

These scoters cannot be distinguished on their breeding grounds as a
western race, although I believe that they have migrated from a west-
ern wintering population. If this view is correct, and the two winter-
ing populations are genetically alike they must mingle after breeding
and before nesting. Since they preserve their geographical separa-
tion in winter and apparently during their spring migration to arctic
Yukon, it is to be suspected that in summer, perhaps after nesting,
enough scoters exchange their western or eastern wintering habits to
maintain a genetically mixed population.

Melanitta perspicillata (Linnaeus)

1 male June 13 weight 983 g. fat testes 11x24, 9x25
mm,

Surf scoters were first remarked mingled in a flock of white-winged
scoters on May 20. Although they were usually in the minority of
mixed flocks of scoters, one flock contained only one white-winged
scoter among nine surf scoters. Scoters were not noticed in associa-
tion with ducks of other species, and flocks containing only one species

OLD CROW 179

of scoter became more frequent after the first few days. In a week
the two species appeared quite segregated.

The subsequent segregation of two species which seemed to be pro-
miscuously associated during early migration may be related to the
earlier appearance of mating behavior among surf scoters, for the pur-
suit behavior of males on June 5 was intense, but the mating pursuits
of white-winged scoters were not so intense until a week later. As
sexual ripeness develops, the different synchronization of its processes
in each species intensifies intraspecific bonds and in the Arctic must
practically preclude miscegenation.

The surf scoters in summer near Old Crow were estimated to be
not half so numerous as white-winged scoters, but their total seemed
to be only about a quarter that of the migrating scoters. It may be
presumed, as suspected for white-winged scoters, that the eastward
migration of surf scoters past Old Crow about suffices to occupy the
Porcupine and Old Crow Valley, and that the two species of scoters
come by a common route from a western wintering place.

Indians at Old Crow esteem white-winged scoters as food but dis-
like surf scoters, which they call Tetre la.

Dall (Dall and Bannister, 1869) did not remark upon surf scoters
at Nulato, but Swarth (1936) described them as “fairly common”
migrants at Atlin, B. C., where they were first seen May 8, 1931, and
May 11-24, 1934. From this we suspect that their spring flight passes
through the mountains in that region from the Gulf of Alaska to the
upper Yukon, but that a coastal route extending further westward
may serve western Alaska.

The two maritime wintering areas of surf scoters are separated by
the continent, although stragglers have been found in many interior
points. Since eastern and western winter specimens are not distin-
guished it may be presumed that there is no isolation to effect genetic
distinction. The spring arrival at Old Crow appeared to be so con-
sistently from the west that the mingling of the birds raised in various
localities evidently occurs later in summer.

Mergus serrator serrator Linnaeus
1 male May 22 weight 1157 g. very fat testes 19x42, 19x50
mm,

Red-breasted mergansers, which reach Anaktuvuk in pairs, nest
northward to the arctic coast of Alaska and Yukon, as far north as
pintails in those regions. Unlike the pintails, which approach their
nesting ground with flocks just disintegrating, the mergansers, highly
gregarious in winter, complete the separation into pairs before they
approach their nesting grounds.

180 U. S. NATIONAL MUSEUM BULLETIN 217

The first red-breasted mergansers were seen May 20 at Old Crow.
Some were single birds but most traveled by twos, male and female.
As a result of this relation of breeding to migration, the spring
course of migration could not be estimated nor could we compare the
relation between migrating and nesting numbers.

The testes of the merganser and mallard had reached the largest size
of any observed among ducks, indicating early maturity for breeding,
and a common basis for their early pairing.

In September Francis Williamson saw large flocks of mergansers on
the Porcupine. The trend of their flights was westward. Because
of this observation it is to be suspected that mergansers migrate to
Old Crow from the large population wintering on the western coast.

The Indian name is Z'trah.

Family ACCIPITRIDAE: Hawks, Harriers

A view of the dates of arrival of migrating hawks which prey upon
birds show that they precede the arrival of any appreciable migration
of their prey. This suggests that they are not attracted to the Arctic
solely by the abundance of migrating birds, whose schedule they
anticipate.

Accipiter gentilis (Linnaeus)

On two occasions we thought a goshawk was recognized in a brief
view while flying among the tops of the spruces. Joe Kay provided
the name, 72 choh, and an appropriate description after studying
illustrations of hawks. Since Olaus Murie in 1926 (Rand, 1946)
banded young goshawks on the Old Crow River, their nesting in the
area, is established.

Accipiter striatus Vieillot

A report of a sharp-shinned hawk on May 8 is regarded as uncertain,
but several times later one was recorded as seen flying through the
tree tops near the edge of the bluff back of the village. On June 3 one
was clearly viewed as it was slowed down in carrying a mouse across
the slough near Kenneth Nukon’s cabin. A specimen was obtained in
September 1958.

The Indian name is Chul rut tstt.

Buteo lagopus (Pontopiddan)

The earliest rough-legged hawk was seen circling high over Old
Crow Point on April 18. On April 29 two were seen several times
along Old Crow River near the cliff about 8 miles north of the village,
and an old nest there on June 10 was found occupied by a young
rough-leg about a week old. A few duck wings were in the nest and

OLD CROW 181

at a nearby perch were feathers and rabbit fur. During the last few
years Robert Bruce had seen the adults, but not the young perched on
this nest when he was returning down river from Crow Flats at the
end of the trapping season, just after the middle of June. Many
people in the village were familiar with this nest.

Between May 2 and 5, six rough-legs were recorded flying over
Porcupine River as we traveled with dog team to Dave Lord Creek
and back. We thought that several were migrating and that their
common direction was northward, but Ben Charlie, son of the former
chief, was of the opinion that most rough-legged hawks arrived in
April and were already settled before May.

The Indian name is Chut rhui chun tsik.

Aquila chrysaétos canadensis (Linnaeus)

Joe Bryant of the Canadian Wildlife Service at Aklavik reported
seeing a golden eagle about April 5 during a survey of muskrats at
Crow Flats. While at Old Crow on April 20, Ben Charlie said that
he had already seen a golden eagle on the Flats. The observations of
our party record only one sight of a golden eagle, on May 16 flying
eastward over Porcupine River. The mountainous areas we visited
were restricted in extent and contained few ground squirrels, Con-
sequently they were unfit for golden eagles. More suitable areas
would be found in the high mountains, where Kessel, Murie, and
Schaller (Unpubl. Ms.) reported golden eagles along the upper part
of Sheenjek River.

The Indian name is Chittese.

Haliaeetus leucocephalus alascanus Townsend

On May 18 a bald eagle was soaring high above Porcupine River
east of the village. <A flock of 20 Canada geese flew under at half the
eagle’s altitude without changing their V formation or course, and
the eagle continued its soaring flight eastward. On June 6, Francis
Williamson saw two bald eagles near Driftwood River. A few bald
eagles summer and probably nest along the Porcupine River, and
from occasional views of them along the river they are familiar to
the Indians, who call them Chizin.

Circus cyaneus hudsonius (Linnaeus)

1 male May 14 weight 367 g. little fat testes 5.5x15.5, 7x13
mm,
1 female May 19 weight 663 g. fat egg 16 mm.

On May 5 a marsh hawk with brownish flanks flew northward along
Dave Lord Creek and continued its flight north by west across Porcu-
pine River. Another seen earlier over Dave Lord Creek also was
migrating northward. On May 7 two marsh hawks, later distin-

182 U. S. NATIONAL MUSEUM BULLETIN 217

guished as male and female, were pursuing separate courses parallel
to the River hunting near the village. These birds repeated these
hunting flights many times daily, but were not seen working to-
gether. This is the pair which I collected. During early May a
marsh hawk was occasionally seen on the opposite side of the river
and appeared to occupy an adjacent territory. Marsh hawks were
seen several times along Porcupine Valley and a few times over the
tundra above tree line on Old Crow Mountains.

The male specimen contained a Microtus in its gizzard. The en-
larged egg in the female showed that it was near condition for laying.

The wide distribution of marsh hawks in winter from British
Columbia to Massachusetts does not suggest the probable wintering
area of the Yukon birds. The direction of a few migratory flights
suggests their arrival on the Porcupine from the southeast.

This bird is known by the Indians as Tzecho.

Family PANDIONIDAE: Ospreys

Pandion haliaetus carolinensis (Gmelin)

On May 17 two ospreys flew up Porcupine River and circled several
times close to the small channel of open water cut through the ice at
the mouth of Old Crow River. They then proceeded up Old Crow
River as if migrating northward. Later in the day an osprey was
seen searching over Porcupine River although only a few channels
of water were running on top of the ice. On May 20, still before ice
started running in the Porcupine an osprey was seen carrying a fish
westward past the village. A mile downstream it soared with its
load to a great altitude before disappearing. Subsequently trans-
portation of fish in this direction was observed several times, and
ospreys were occasionally seen along the Porcupine.

The Indian name for the osprey is Thuk.

Family FALCONIDAE: Falcons

Falco rusticolus Linnaeus

We did not recognize a gyrfalcon, although numerous peregrines
were seen. From an illustration and by comparison with peregrines
Joe Kay named the gyrfalcon Hwi tst chi, and said that it nested in
spruce trees. Gyrfalcons are not mentioned by Rand (1946) as having
been reported among birds of Yukon territory, but they were seen by
Kessel, Murie, and Schaller (Unpubl. Ms.) on upper Sheenjek River
in the Brooks Range. We consider this nearby observation to confirm
reports by the Indians that gyrfalcons occurred near Old Crow. In
the arctic interior of Alaska gyrfalcons are settled throughout the

OLD CROW 183

year in certain areas, where they also nest, and this is assumed to be
their status near Old Crow.

Falco peregrinus anatum Bonaparte
4 fresh eggs June 8 — pe ene =

On May 5 a male and female peregrine were calling and appar-
ently settled on the cliffs just below Dave Lord Creek. Guano near
perches there indicated long use. After May 16 a male and female
peregrine were occasionally seen and heard near the mouth of Old
Crow River, a vicinity where Robert Bruce said that they often nested.
At the end of the first portage across the long bend of the river, about
6 miles due east of Old Crow, a pair of peregrines were seen about
a cavity in the cliff, and there Sidney and Leonard Peyton obtained
four fresh eggs on June 8. Since peregrines are reported to lay their
eggs every other day (Bent, 1938) we can estimate that the first was
laid about June 1.

On June 13 eight peregrines, six of them associated as if in pairs,
were seen at separate cliffs during 40 miles travel along Porcupine
River. Because almost every suitable cliff was occupied, it is doubt-
ful if many of the nesting peregrines remained unseen.

The Indians know this bird as Chinechun.

We had noticed in April that over a hundred old nests of cliff swal-
lows were located along about a quarter of a mile of these cliffs where
the peregrines’ nest had evidently been situated for many years.
Stalagmites of guano showed the ancient occupation by cliff swallows
of some of the positions, and bulkier accumulations of guano near
perches of the peregrines showed their long use of the area.

At the time the peregrines’ eggs were found the nearest completed
nest of cliff swallows was 100 feet distant, and within 200 yards were
over a hundred nests in various stages of completion. It seems as if
cliff swallows would be especially susceptible to attack during their
social swarming, mating, and nesting. Accounts of peregrines’ feed-
ing indicate that in some places they capture and devour rough-winged
swallows (Bent, 1938). To pluck nestling swallows from their nests
would require no exertion of the strength of the peregrine’s claws.

In his description of cliff swallows, Alfred Gross (Bent, 1942) re-
ported that Coues, Taverner, and Forbush had each commented upon
swallows nesting on cliffs where prairie falcons also nested, and said
that Taverner also found them about a duck hawk’s nest. Moreau
(1942) has compiled numerous reports of African birds nesting close
to insects which repel the approach of other animals by stinging, and
several instances of small bird’s nesting on the nest structure of larger
birds which are usually considered to be predatory.

We did not observe the peregrines bothering the swallows, and it is

469496—60—18

184 U. S. NATIONAL MUSEUM BULLETIN 217

apparent that predation upon them has not been sufficient to destroy
that community of several hundred swallows, as would surely result
if the peregrines obtained from them as much as a tenth of the sub-
sistence needed during their Arctic residence, for it would take 100
swallows to weigh 3000 grams and provide enough to feed two adult
peregrines for about one out of the approximately ten weeks during
which they and the swallows are neighbors.

Perhaps the exertion for capturing a 30-gram swallow, about 149th
of a peregrine’s daily food, is not worth the effort when a peregrine
can take a duck equivalent to 30 swallows as easily as a. human can
remove an apple from his refrigerator. Whatever condition guides
the habits of peregrines it must permit the existence of a neighboring
population of cliff swallows.

Falco columbarius Linnaeus

The first pigeon hawk was reported seen on April 380. The swiftness
of its flight left identification uncertain, but this early date did not
surprise us, for a pigeon hawk was seen harrying jays in mid-March
on the Alatna Malemute, just south of tree line in the central Brooks
Range, and Dall (Dall and Bannister, 1869) found them in winter
at Nulato. Another pigeon hawk was reported at Old Crow on May 8,
but these small falcons are difficult to identify by sight because of the
swiftness of their flight when close to brush and timber. Pigeon
hawks have been reported on the lower Porcupine (Williams, 1925),
at Lapierre House (Ross, 1862), and on the upper Sheenjek River
(Kessel, Murie, and Schaller, Unpubl. Ms.).

The Indian name is Chin tettree.

Family TETRAONIDAE: Grouse and Ptarmigan

Sharp-tailed grouse (Pediocetes) were known by several Indians
who had seen them along the Yukon River, where they were called
“willow” grouse, a name also used in Alaska. They said that there
were none near Old Crow. No information was obtained concerning
white-tailed ptarmigan, but they have been recorded at Lapierre House
(Ross, 1862).

The numbers of willow ptarmigan near Old eae were locally re-
garded as showing a great decline in 1957 from their usual abundance.
Varying hares were also few. We did not hear of a living hare being
seen in spring of 1957 and ourselves saw only three fragments which
appeared to be relics from winter, but the tracks of hares were not
unusual and some were fresh. Periodic decreases of willow ptarmigan
and hare are known locally to occur, but we did not inquire about the
relation of their cycles.

OLD CROW 185

Some of the Indians suggested that the rarity of spruce grouse in
1957 might be associated with the reduced number of ptarmigan, but
the number of spruce grouse near Old Crow is not usually sufficient to
make their absence impressive.

Ruffed grouse, reported at Lapierre House (Ross, 1862) and Forty
Mile (Grinnell, 1909), are so little known at Old Crow that failure to
remark them in 1957 is not significant of a change in the population.

While rock ptarmigan are well known at Old Crow, their habitat is
not regularly observed and our inquiries did not evoke opinions upon
change in their numbers. I saw as many in the Old Crow Mountains
as I have seen at various times in some Alaskan localities.

The Alaskan races of willow and rock ptarmigan extend to Old
Crow in the arctic Yukon.

Canachites canadensis osgoodi Bishop

Wesaw no signs of spruce grouse in 1957, and residents of Old Crow
said that they had seen none that spring. In most years spruce grouse
are often seen in August and 9 specimens were obtained in September
and October 1958, at which time individuals and flocks were being
occasionally reported. It seems that they are never abundant, but
that a few are usually noticed in all seasons. The species is normally
resident in Alaska, Yukon, and Mackenzie to the northern edge of
spruce forests (A. O. U., Check-list, 1957).

Bonasa umbelius (Linnaeus)

Joe Kay, elected chief at Old Crow from about 1920 to 1984 and
a man possessing great wisdom about his country and its animals,
said that he knew and used to hear the “thumping” of ruffed grouse,
although for two years he had not heard or seen them. On the basis
of his report, ruffed grouse are listed among the resident birds of
Old Crow. The Indian name is Chut tul.

The species had been reported at Lapierre House (Ross, 1862),
Forty Mile (Grinnell, 1909) and Fort Yukon (Bishop, 1900) and
from Porcupine to Old Crow and Lapierre (Aldrich and Duvall,
1955). Residents in the Alaskan Arctic forest along the Koyukuk and
Kobuk Rivers do not seem to be familiar with ruffed SOAS PHi but Bie
seem to extend just across the Arctic circle in Yukon.

Lagopus Pgous alascensis Swarth

4 females Apr. 5-12 weight 532-627, little fat (2), no eggs 0.75, 1.5 mm.
average 584 g. fat (2)
6 sex unknown Apr. 5 weight 546-636, — —

average 591 g.
The average weight of willow ptarmigan in the small sample from
Old Crow was less than the 619-gram average for 128 L. 7. alascensis

186 U. S. NATIONAL MUSEUM BULLETIN 217

from the central part of the Brooks Range in winter. In summer,
arctic Alaskan willow ptarmigan are lighter, and their weight then
does not differ significantly from that of birds from Old Crow. They
are, however, significantly heavier than the average weight (522
grams), found in 32 examples of willow ptarmigan from the Chugach
and Talkeetna Mountains in the southern Alaskan range of alascensis.

Although not fat, none of the numerous willow ptarmigan brought
into the village appeared ill nourished. They fed on willow buds and
tips of twigs, reaching them from the ground or often from branches
as high as 20 feet, where they appeared clumsy and insecure while
reaching for the slender tips.

Some ptarmigan burrows in the snow showed by the quantity of
droppings in them that they had been occupied throughout cold nights,
but in moderate weather some of the open roosting depressions in the
snow were shown by their content of droppings likewise to have been
occupied during the night. It is not correct, however, to conclude that
burrowing is for protection from the cold, because it certainly affords
concealment and it is doubtful if the insulation of snow is needed in
addition to the physiological adaptations which protect ptarmigan
from cold. In mild weather the snow is heavier and since it may form
a crust during the night, it seems to be generally unsuitable for
burrowing.

At Old Crow, as in the Brooks Range, some but not all willow ptar-
migan in white winter plumage have their feathers suffused with a
delicate shell pink like a faint but beautiful fluorescent dye. This
color appears in white ptarmigan and short-billed gulls in spring,
and is most striking in Ross’s gulls when they appear at Barrow in
September. It disappears from specimens after a few days.

The nine specimens listed were all obtained within a few miles of
Old Crow Village among the willows along the river. I had not no-
ticed that Arctic willow ptarmigan regularly concentrated according
to sex during migration. At Anaktuvuk, the sexes have been found
mixed in the usual flocks which migrate near the level of the river, but
Simon Paneak told me that in spring small] flocks found occasionally
on the hillsides might contain a majority of either sex. There is evi-
dently some segregation by sex at times during migration.

Near Old Crow during a walk of 10 miles along the river in early
April, several flocks of from 10 to 50 birds might be seen, but I did
not record seeing more than 100 birds on any day. The people of Old
Crow used the expression “no” ptarmigan and “no” rabbits in 1957 to
compare the scarcity in 1957 with the numbers common in other years.
The numbers of willow ptarmigan seen at Old Crow were few com-
pared with those seen on occasions during the winters 1951 to 1954 at

OLD CROW 187

Anaktuvuk, where Simon Paneak reported they were still at the peak
of exceptional abundance during the spring migration of 1957.

Moses Tizya, who returned April 14 from winter trapping on the
Porcupine about 85 miles east of Old Crow, remarked that there were
“no” ptarmigan there during the past winter, although the valley had
been “white” with them in other winters. Paul Joseph, who returned
May 2 by dog team from Fort McPherson, also reported that there
were “no” ptarmigan along that route, covering about 140 air miles
eastward, so the scarcity of willow ptarmigan in 1957 extended east-
ward at least to the Richardson Mountains, Several Indians remarked
that no spruce grouse had been seen this winter although usually there
were a few. Ruffed grouse, which are considered rare, had not been
seen at all for several years. It seems that the period of scarcity
among ptarmigan is not synchronous in the race which extends across
arctic Yukon and Alaska.

The last flock of willow ptarmigan was recorded on April 9, but a
single bird was collected on April 12. On April 8 fresh tracks of wil-
low ptarmigan were found in the scrub birch and spruce near tree
line at about 1,600 feet elevation on Old Crow Mountain, and Ben
Charlie reported about 1,000 in one flock on April 18 above timber on
the trail passing over the eastern shoulder of the mountain. He said
that on April 26 willow ptarmigan were still on the mountain and at
Crow Flats. While on patrol from Old Crow to Herschel Island dur-
ing the week before April 26, Constable Robin saw ptarmigan, some
with dark spots, as far north as the edge of timber, but he saw none
northward to the coast.

The Indians informed us that in summer willow ptarmigan moved
north of the timber or above tree line in to some of the extensive moun-
tain tundra. The spring migrations probably occur in a direction
which produces the requisite summer vegetation or climate either by
change of latitude or elevation. The land along the Old Crow River,
normally well occupied by willow ptarmigan in winter, extends with
some timber northward for about 75 miles and the valley is practically
cleared of ptarmigan during the summer. The Koyukuk Valley about
Bettles is occupied in winter, but in summer it and the southern John
River Valley are pretty well cleared for 50 miles by the northward
migratory movement. On the Alaskan arctic slope willow ptarmigan
are few in winter and usually rather numerous in summer. At Anak-
tuvuk, the center of density of ptarmigan seems to move from south
of the divide in the Brooks Range in winter to north of the divide in
summer, perhaps a distance of 200 miles. In the Old Crow Valley,
_ where the forested wintering ground approaches to within 50 miles
of the arctic coast, the mean migratory displacements appear to be
shorter.

188 U. S. NATIONAL MUSEUM BULLETIN 2/17

Willow ptarmigan are usually near Old Crow in winter, but as
the flocks rove about they cannot be called settled residents. Their
annual schedules of movement appear to recur in each locality and
so there is a true if relatively short distance migration.

The race L. 7. alascensis in Alaska, is distinguishable from L. 7. albus
by the size and shape of the beak, and our specimens from Old Crow
were typical of the Alaskan race. The range of albus is reported to
extend to northern Yukon (A. O. U., Check-list, 1957), but our speci-
mens show that the Alaskan race extends eastward on the Porcupine at
least to Old Crow.

The Indian name for this ptarmigan is Take.

Lagopus mutus nelsoni Stejneger

9 males Apr. 11-June 9 Weight 389-435, fat (1), medium fat testes 4-18 mm.
average 413 g., (2), little fat (1),
coeff. of var. very little fat (8),
4.3% no fat (2)

3 females Apr. 21-May 25 weight 405-446, medium fat (1), eggs 1.5-23 mm.
average 425 g. little fat (1), very

little fat (1)

The average weight of male rock ptarmigan at Old Crow was found
less than the mean weight (466 grams) in spring of male Z. m. nelsont
at Anaktuvuk, but the three females did not differ in weight from
those at Anaktuvuk.

The beautiful faint pink of the white winter feathers appeared often
in rock ptarmigan of both sexes at Old Crow as it often appears in
willow, rock, and whitetailed ptarmigan of Alaska.

In the vicinity of Old Crow, rock ptarmigan were seen only on the
mountains above tree line at elevations from 2,000 to 4,000 feet, the
highest examined. The small tracks and open night-roosting places
in the snow were identified by droppings at the edge of the spruce,
dwarf birch, and alder at about 1,600 feet elevation on April 8. On
April 11, a wild flock of about 50 birds found at 2,400 feet scattered
into several groups when pursued. One group which contained about
15 birds was approached while feeding at about 2,700 feet. On April
29, eight birds were seen in groups of two or three and no flocks were
seen then or subsequently. The males were calling and mating was in
progress.

In the breeding season females can often be distinguished by their
compressed feathers, which give them a slender appearance as they
stand or walk, and by their practice of carrying their head and tail
extended nearly horizontally. The males stand straighter and with
feathers erected, giving the impression of larger size, although the
weight of the two sexes is not different. A trace of dark feathers was
appearing on the head of the females and the red eye combs of the

OLD CROW 189

males were larger than on April 11. The testes had not reached half
their maximum length when these initial steps toward pairing were
in progress.

Specimens taken on April 17 and 20 were stained with crowberry
(Empetrum) and these berries were found in the crop of one bird.
At this time the snow had cleared from about 40 percent of the steep
southeastern slope, but was continuous on the less steep and less sunny
areas.

On May 10, a windy day, only one male rock ptarmigan was seen
at about 2,500 feet. On May 25 three collapsed ovarian follicles and
a brood patch in a female specimen showed that it had laid three
eggs, the first on May 23. Only 10 percent of her white contour
feathers and white primaries remained. A male companion had a
few speckles on the crown and one on the back replacing not more than
5 percent of the white feathers, and its eye comb and testes were the
largest noted. It is interesting that the female, although in the course
of an intensive program of egg laying, was nevertheless marked as
“medium” fat, which is about as fat as ptarmigan are found. The
very considerable extra metabolic expenditure of laying was well
covered by the food supply, leaving no sign of nutritional strain to
suggest that the bird was under stress. This is in accordance with the
appearance of unhurried actions of the female rock ptarmigan during
the breeding season in arctic Alaska.

On May 26 two rock ptarmigan were seen as they stood on ex-
posed rocky points at an elevation of 2,500 feet, in the statuesque pose
which males take during nesting season. At this time and on June 9,
other male rock ptarmigan stood on occasional eminences high in the
first and second mountains at the conspicuous posts from which they
signal their nesting territory. The testes of specimens taken at these
times were smaller than the maximum size observed about May 25
but dark feathers had only covered the head and a few spots on the
neck and body.

Measurements (Mm.) Plumage
Change (%)
Date Social Behavior Testes Ova Male Female

Apr. 11 flocks disintegrating 0 0
Apr. 17 3x4, 3x4 1.5 0

6x8, 5x7 0
Apr. 20 pairing 6x5, 7x5 2 0 Trace
May 22 o' signals, 2 laying
May 25 o' signals, 9 laying 11x18, 11x17 123 i 95

11x16, 9x13
May 26 o' signals, ° laying 11x18, 9x14 1
June 4 o signals, 9 incubating 11x13, 9x12 5
June 9 oO signals, 9 incubating 11x14, 5, 10x13 5

1 Three collapsed ovarian follicles.

11x13, 9x13

190 U. S. NATIONAL MUSEUM BULLETIN 217

The flocks in winter occasionally flew out of sight several miles
away. In summer male birds signaling or nesting were not seen
closer together than about a mile. In the barren Old Crow Moun-
tains that much range might be required for a brood of chicks, and
the winter flock of 50 would accordingly need some 25 square miles
of mountain for nesting. The first Old Crow Mountain scarcely con-
tains 10 square miles of nesting territory. The others of that iso-
lated mountain group may provide 100 square miles for nesting.
These rough estimates suggest that winter flocks convene from a nest-
ing area with peripheral points 30 miles separated. Within this area
some may be resident through a year. Tracks showed that early in
April rock ptarmigan had been among the dwarf birch, willow, and
alder along the margin of the spruce forest, at about 1,500 feet eleva-
tion. We saw them in April, May, and June above 2,000 feet on the
bare rocky slopes and mountain tops. Winter and summer move-
ments of rock ptarmigan over Old Crow did not show regularity worth
the designation of migrations, and we have not had certain evidence
for any regular migration of rock ptarmigan in Alaska. As they are
one of the swiftest among birds and have large hearts for their weight,
rock ptarmigan are physically equipped for long flights.

The race L. m. nelsoni, which is easily distinguishable by the pat-
tern of even a few feathers of summer plumage in both sexes, occupies
all Alaska except the southeastern part. In arctic Yukon this race
extends eastward at least to Old Crow without deviation in character-
istics. Somewhere between there and northern Mackenzie occurs the
border between nelsonz and rupestris. Porsild (1943) reports rupes-
tris a rare visitor to the Mackenzie Delta where the forested lowland
may separate the races of these mountain and tundra birds. In south-
ern Yukon, however, Rand (1946) assigns 15 specimens to rupestris.
As is the case among several species of migratory birds, the race found
in Alaska extends eastward only across northern Yukon.

The Indian name for the rock ptarmigan is Z’ako.

Family GRUIDAE: Cranes

Grus canadensis canadensis (Linnaeus)

The first crane that we saw on May 14 was alone circling and call-
ing high in the air. Again we heard one on May 18 and saw two on
May 22. Cranes are well known to the Indians, who said that they
nest on Crow Flats, but evidently not many come to Porcupine Valley.
Their Indian name is Chya.

Rand (1946) quoted reports of great migrating flocks passing over
points marking courses from the Liard River to the upper tributaries

OLD CROW 191

of the Yukon River. Evidently this route 600 miles south of Old
Crow is followed by the bands of cranes which traverse southern
Yukon and nest throughout Alaska.

Family CHARADRIIDAE: Plovers, Turnstones, Surfbirds

At Old Crow the species and numbers of plover were inconspicuous.
It seems to be off the route of the migrations which take so many
plover to nest in western arctic America.

Charadrius semipalmatus Bonaparte

2 females 20 May, 10 June weight 38.3, 54.1 g. little fat 4 broken follicles
brood patch

Three semipalmated plover were the first of the species seen at Old
Crow on May 19. They flew away eastward. During the time of
breakup and until June, while the river remained high, a few semi-
palmated plover were seen about marshy places near the village. As
the water fell pairs of these plover were occasionally seen on the
gravel bars and banks of the river.

The female specimen taken on June 10 had evidently begun to lay
by June 7. Although she had completed laying, she weighed 14 grams
more than the average of the other females and three females from
Anaktuvuk, an overweight condition sometimes noticed in females
of other species at the time of laying.

The breeding range of the semipalmated plover extends across
Alaska and Yukon without separation from the Mackenzie area.
We have no evidence as to how the Porcupine is reached from the
widespread migration through North America, and their wintering is
widespread but apparently continuous across southern United States.

Its Indian name is Shishenetyi.

Charadrius vociferus vociferus Linnaeus

On May 20 Francis Williamson and Leonard Peyton saw a killdeer
standing and calling by a grassy pool behind the village. On June 1
Leonard and Sidney Peyton also saw one which might have been the
same individual.

Rand (1946) mentions only one sight record of a killdeer in Yukon
territory. Its occasional presence in arctic Alaska is confirmed by
specimens from Barrow (Bailey, 1948) and on the basis of Eskimo
reports is suspected at Anaktuvuk and Kobuk.

There is no report of killdeers nesting north of British Columbia
but it is apparent that individuals frequently wander in the western
Arctic.

192 U. S. NATIONAL MUSEUM BULLETIN 217

Pluvialis dominica (Miiller)

It was thought that several golden plover were seen in flight over.
the village but not confirmed in 1957. Father Mouchet described sev-
eral single golden plover and pairs which he had seen about the Mis-
sion in spring, 1956. Golden plover have been reported in Yukon
north to the arctic coast (Rand, 1946). Apparently Old Crow does
not lie on a route significant for the migration which brings so many
golden plover to nest in northern Alaska. These could only belong
to the race which winters in Argentina and migrates northward
through the Mississippi Valley. They nest on the arctic coast of
Yukon and some migrate across southern Yukon (Rand, 1946) but
the available reports do not show where the bulk of the great popula-
tion of golden plover nesting in Alaska traverses Yukon in its westerly
course from the northbound migration in the upper Mackenzie Valley.
Except near the arctic Yukon coast the golden plover nesting in
Alaska seem to be separated from those which nest over such a wide
area between Mackenzie and Hudson Bay.

Family SCOLOPACIDAE: Woodcock, Snipe, Sandpipers

Only 9 species of sandpipers were seen at Old Crow, a small list
compared with the 16 known at Kobuk and 18 at Anaktuvuk. The
terrain along Porcupine River suits snipe, spotted and solitary sand-
pipers, and lesser yellowlegs, but among these we could only establish
that snipe and solitary sandpipers appeared in sufficient numbers to
demonstrate numerically significant migration through Old Crow.
As seen at Old Crow the Porcupine Valley is an unimportant migra-
tory path for sandpipers as well as for plovers. .

Six of the species of sandpipers at Old Crow are restricted in their
Arctic nesting to Mackenzie, Yukon, and Alaska.

The fact that six of the nine species of sandpipers found at Old
Crow are not divided into races suggests that these northwestern
breeding birds are not genetically differentiated and that, as among
ducks, the populations which are localized for breeding exchange
members with the rest of the population of the species during migra-
tion or on their wintering grounds.

Among the three races of sandpipers there is no local diftérentia-
tion among North American snipe (Capella). The whimbrel (Vw-
menius) and solitary sandpiper (7ringa) are western races that mi-
grate farther south in winter than the eastern races and they migrate
farther north in summer. In both form and geographical distribu-
tion the western and eastern populations are distinct.

In southern Yukon snipe are uncommon (Rand, 1946) and whimbrel
rare. Probably arctic Yukon is a geographically significant connec-
tion for the Alaskan and Mackenzie populations of these species.

OLD CROW 193

Hypothetical migration routes of Scolopacidae are shown in the fol-
lowing tabulation :

Species Arctic nesting in Northern wintering Route through Coming
U. S. and Canada toward
Old Crow
from
Capella gallinago Mackenzie, Yukon, British Columbia to (?) East or West
delicata Alaska Alabama
Numenius phaeo- Mackenzie, Yukon, Central California Pacific Coast East
pus hudsonicus Alaska
Actitis macularia Mackenzie, Yukon, British Columbia (?) (?)
Alaska
Tringa solitaria Mackenzie, Yukon, Ecuador Mountain (2)
cinnamomea Alaska
Totanus flavipes Mackenzie, Yukon, Mexico Prairie Northwest
Alaska
Erolia melanotos Southampton to Siberia Peru Interior Northwest
Erolia bairdii Greenland to Siberia Ecuador Prairie Northwest
Erolia minutilla Mackenzie, Yukon, Oregon to North (?) (2)
Alaska Carolina
Ereunetes pusillus Baffin Island to Alaska Gulfcoast Interior Northwest

Capella gallinago delicata (Ord)

1 male May 22 weight 96.3 g. medium fat testes 7x15, 5x7 mm.
1 male June 5 weight 93.0 g. very little fat testes 7x15, 8x12 mm.
4 slightly in- June 7 = == —

cubated eggs

Wilson’s snipe were reported by Neil McDonald during the evening
of May 7. On the next day their presence was apparent in the per-
sistent winnowing over the lake and marshes back of the village.
Until June the winnowing was often carried on with several snipe in
the air steadily for a few hours, but some individual birds were
noticed descending after only five minutes of winnowing flight. After
May the winnowing was heard less often.

Joe Kay found a nest with four eggs on June 7. The first egg
would have been laid about May 31. Peter Lord reported hearing the
first snipe at his camp in the northern part of Crow Flats on May
16. Barbara Oakeson (1954) reported snipe on May 9 at Mountain
Village, Alaska. At Anaktuvuk the mean first date from 1950 through
1953 was May 16, but in 1957 the surprisingly early date of May 8 was
reported there. The number of snipe heard and seen during the
early season at Old Crow appeared to be considerably in excess of the
nesting territories about the village, suggesting that there was some
migratory passage.

The Indian name is Paeaale meaning “calling from the sky.”

Snipe reach the Arctic in Mackenzie, Yukon, and Alaska. Since the
wintering range extends from British Columbia across the northern
States there is no basis in recorded migrations for suspecting how the
Arctic nesting population travels. Rand (1946) remarked that “
have no records: of snipe: having been even moderately common in

194. U. S. NATIONAL MUSEUM BULLETIN 217

Yukon.” They are common in the Mackenzie Delta (Porsild, 1943)
and through Alaska. The plentiful nesting population on the Porcu-
pine, apparently a main connection between abundant Mackenzie
and Alaskan nesting populations, implies that the migration to the
Porcupine is either from the west or from the east since central and
southern Yukon seem to be but little visited by snipe.

Numenius phaeopus hudsonicus Latham

1 male June 4 weight 364 g. little fat testes 8x12.5, 4x9 mm.
1 male i June 9 weight 312 g. little fat testes 8.5x15, 6x10
mm,
2sets of 4 slightly June 4 — — —
incubated eggs

A whimbrel was heard on May 24 on the sedgy saddle extending
for about a mile between the southern and next northern of the Old
Crow Mountains. Hummocks of sedge interspersed with lichen cover
extended over this gently sloping saddle at elevations from about
1,600 to 2,000 feet. On May 25 two pairs were watched and four
other whimbrels were flying about calling the trill which resembles
the loud call of a tree frog. ‘These birds often landed and all appeared
to be attached to the locality.

Two whimbrels were seen at the same place on May 27 and Father
Mouchet saw several on May 29. On June 4 Sidney Peyton collected
a male bird and two sets of four slightly incubated eggs which must
have been first laid about May 27. On June 9 three pairs were again
seen on the same saddle. The larger appearing bird of one pair was
collected and turned out to be a male with a brood patch. Both of
these birds were observed calling, were unwilling to leave the vicinity
in spite of an hour of pursuit and several shots fired at long range.

Whimbrels, which the Indians call Zetnjyo, nest on tundra loca-
tions in arctic Mackenzie, Yukon, and Alaska. They are rare in the
remainder of Yukon (Rand, 1946), and are common migrants only
along the coast of British Columbia (Munro and Cowan, 1947). It is
indicated that the arctic nesting population comes from the migra-
tion along the Pacific coast, and presumably reaches arctic Yukon and
Mackenzie from the Yukon Valley after crossing the mountains from
the Gulf of Alaska.

Actitis macularia (Linnaeus)

Imale June 8 weight 45.5 g. medium fat testes 7x12, 10x9
mm.

1 male June 13 weight 39.7 g. no fat testes 5.5x13, 7x9
mm.

1 female June 13 weight 41.7 g. little fat egg 5mm.

4 fresh eggs June 24 — — —

A spotted sandpiper was first seen flying near the bank of the Por-
cupine River at the time of the break up, May 22. No other was

OLD CROW 195

noticed until May 29, when one was seen on a mudbank just beginning
to emerge as the river subsided; after that time, single spotted sand-
pipers and pairs were frequently seen along the banks. The two
specimens collected on June 18 were together and apparently a pair.
The female contained one 5-mm. egg but had not yet laid. A nest was
found on June 24 with four fresh eggs, of which it was estimated the
first was laid about June 18, the latest start of laying found among
sandpipers.

Spotted sandpipers, known by the Indians as 7’raruk, are common
along the Porcupine River but there was no evidence of migratory
traffic in excess of the summer residents.

These sandpipers seem to nest north to the tree line, and
so extend into the Arctic from Mackenzie, across forested Yukon and
Alaska. In winter some remain as far north as British Columbia
(Munro and Cowan, 1947) but are spread over more southerly States
in the east. Their migratory course is too widely distributed to sug-
gest from which direction Old Crow is reached.

Tringa solitaria cinnamomea (Brewster)

Males Females
Weight Fat Testes Weight Fat Eggs
Date (g.) (mm.) (g.) mm.)
May 16 53. 6 F 5x7, 5x6.5
May 17 52.2 F 5x10, 4x6
48, 2 F 5.5X7, 4x5
May 18 53. 7 F 5.5x8.4, 5x7 54.6 F 6
May 20 59.6 VF 5x7.5, 5x6.5
52.2 VF 5x7, 4x6
May 23 72.2 LF 12
June 1 55.4 MF 6x10, 4x5 brood patch
June 5 55. 4 MF 5x8.5, 3x4.5
June 8 54.7 LF 5x8, 3x4 65. 2 LF 16
(aver.) 53.8

4 broken follicles.

Solitary sandpipers, which bear the Indian name 7Zwe, had been
heard on May 14 and several were seen on May 16 and 17. On May
18 about 12 were remarked in groups of 2 or 3 around marshy pools.
From then until May 31 they were seen more often than they were
later on, and their numbers seemed to exceed the old nests which
could provide their eventual housing in the vicinity. Many of these
birds appeared to be pausing in migration, and it was our impression
that most of them took off in an easterly direction. Francis William-
son found solitary sandpipers at Fort Yukon when he arrived there
on May 10.

A number of solitary sandpipers were settled around marshy pools
and streams in June. Although we had not earlier observed many old
nests there were enough new ones of robins and thrushes which could

196 U. S. NATIONAL MUSEUM BULLETIN 217

subsequently provide nesting places for a great many solitary sand-
pipers.

No eggs were found, but the female specimens taken on May 18
and 23 were evidently near to laying, for they had eggs measuring
6and 12mm. A male on June 1 was remarked with a brood patch and
a female on June 8 contained four ruptured ovarian follicles. It ap-
pears that eggs were first laid about May 25.

The females taken on May 23 and June 8 were much heavier than
the female taken on May 18. Such variation is not seen among adult
birds except in the transient condition of females being overweight
while ovulating.

Male specimens taken between May 16 and 20 were fat or very fat,
but those taken subsequently were medium and little fat.

Solitary sandpipers appeared the most common species of the family
to settle around Old Crow, and it is estimated that the migrating
numbers could provide for their extension over much of Porcupine
Valley.

All were distinguishable from 7’. s. solitaria in length of wing and
in the heavier marking of the throat and breast. Of 12 Old Crow
specimens, 11 had clear marbled patterns on the inner side of the first
primary. The examples are typical of the western race, and in most
the back pattern contained more buff and was rather more brownish
than in solitaria.

The western race nests in arctic Mackenzie, Yukon, and Alaska,
perhaps confined to the forested areas. In spring they migrate from
Keuador (and southward) principally through the intermountain
region, reaching Atlin, B. C., on May 2, 1931, May 12, 1933, and May
10, 1934 (Swarth, 1986). It is possible that their arrival at Old Crow
is either from the west or from the southeast.

Totanus flavipes (Gmelin)

1 male June 11 weight 66.5 g. == testes 3x5, 2x4 mm.

1 female May 31 weight 100.0 g. medium fat 4 ruptured follicles, 1
formed egg, brood
patch

The first report of a lesser yellowlegs on May 7 seems early in view
of the weather and the condition of the snow and ice two weeks before
breakup (Williamson found them present at Fort Yukon when he ar-
rived there on May 10, and they have not been reported at Anaktuvuk
before May 16). Additional yellowlegs were not recorded until May
23, but on May 31 the condition of a female specimen showed that it
had begun to lay on May 28. This bird weighed 100 grams or about 25
percent more than the average of six females in spring at Anaktuvuk.
This is another example of the transient overweight of laying females.

OLD CROW 197

The male specimen was abnormally light and its testes measured only
5 mm., which is smaller than usual among breeding sandpipers.

The lesser yellowlegs did not at any time seem to exceed the con-
siderable numbers which were found in the marshy area during nest-
ing time, and so there was no evidence for their migratory passage
through Old Crow.

Yellowlegs migrate from their Mexican wintering places northward
chiefly through the interior, (A. O. U., Check-list, 1957). Since they
are abundant in autumn and scarce in spring transients in British
Columbia (Munro and Cowan, 1947) we may suspect that spring
migration to the northwestern Arctic is from the southeast. They had
reached Atlin on May 3, 1980, May 2, 1983, May 5, 1934 and May 3,
1935 (Swarth, 1936); Ross Post, May 11, 1930 (Rand, 1946); and
Anaktuvuk about May 17 in five years. It seems as if their spread to
arctic Alaska is either slow or reached by a different contingent of the
migratory population.

The Indian name is Z’echuh.

Erolia melanotos (Vieillot)

Males Females
Weight Fat Testes Weight Fat Eggs
Date (g.) (mm.) g.) (mm.,)
May 23 76. 2 VL 6x13, 7x12
May 24 74.0 N 6.5x11.5, 7.5x11
Junel . 96. 7 F 8.5x16, 10x13 53.6 L 4
54.8 L 3
57.7 L 4
53.8 L 3
61.1 L 4.5
53.2 F 3
52.7 F 3
58.5 F 3
57.4 F 4
61.5 F 4
57.0 F 2
(aver.) 82.2 56. 5

A flock of eight pectoral sandpipers was seen on May 23. During
the next week from one to six were occasionally seen about the marshy
pools behind the village. The latest record was of a female shot on
June 11 but not preserved. Eggs were from three to five mm., so that
the females were approaching breeding condition. The testes of two
earlier male specimens were smaller than mature size but on June 1
they were full size. The pectoral sandpipers seen were evidently in
migration and there was no indication that they nested near Old Crow.

The large variations in weight among male pectoral sandpipers are
not unusual. In a series of 25 weighed in spring at Anaktuvuk were
examples lighter and heavier than those weighed at Old Crow. In
both sexes the visible fatness was also conspicuously variable. Among

198 U. S&S. NATIONAL MUSEUM BULLETIN 217

the species of Arctic migratory birds, the pectoral sandpipers are no-
tably variable in weight.

The arrival of pectoral sandpipers at Old Crow was later than the
usual date for the beginning of the large migration through Anaktu-
vuk. Judging from the few birds seen in comparison with their num-
bers migrating through Anaktuvuk, Old Crow is not an important sta-
tion on the migratory route of pectoral sandpipers, and there was no
evidence for their nesting. From their wintering places pectoral sand-
pipers migrate in spring northward from Peru mainly across interior
States and Provinces east of the mountains (A. O. U., Check-list,
1957), so their arrival in Arctic Yukon and Alaska is probably from
the east and southeast.

The Indian name is Teggetesel.

Erolia bairdit (Coues)
2 males May 29 weight 38.4, 39.5 g. fat (2) testes 5x7, 5x5 and
5x8.5, 4x6.5 mum.
From their wintering place south of northern Ecuador, Baird’s
sandpipers migrate northward through the prairie States and Prov-
inces (A. O. U., Check-list, 1957) ; and probably reach Old Crow from
the east or southeast, but only a few groups containing two or three
Baird’s sandpipers were seen there, in sharp contrast to the great
numbers migrating through and nesting near Anaktuvuk.

Erolia minutilla (Vieillot)

1 male May 20 weight 22.1 g. medium fat testes 3.5x6 mm.

1 male May 23 weight 22.1 g. little fat testes 2.5x3.5 mm.
1 female May 17 weight 20.1 g. fat eggs 1.75 mm.

1 female May 23 weight 24.9 g. fat eggs 2.75 mm.

It gives reassurance that spring is progressing to see these little
sandpipers arrive in the Arctic. ‘Two were seen feeding at the border
of a marshy pool on May 17 and later a few were seen each day during
May. On May 23 a dozen least sandpipers were running over the float-
ing debris among the flooded willows on the margin of the river. No
flocks were seen and their delightful song was not heard. On June 3
near Kenneth Nukon’s cabin a pair was seen fluttering about together
near the ground in courting manner with their wings rapidly beating
like those of a large moth. The migration of least sandpipers through
Old Crow was small and left only a few pairs nesting in the valley.

Since there is a wide wintering range from south of Oregon to
North Carolina (A. O. U., Check-list, 1957) the source of these tiny
migrants cannot be suggested.

The Indian name is Z'agatsil.

OLD CROW 199

Ereunetes pusillus (Linnaeus)

1 male June 1 weight 21.7 g. fat testes 4x7, 4x5.5 mmyg
1 male June 3 weight 22.7 g. medium fat: testes 5x8 mm.

1 female June 2 weight 23.3 g. fat eggs 2mm,

1 female June 3 weight 26.5 g. medium fat eggs 3mm.

A flock of seven semipalmated sandpipers was noticed on June 1,
a later date than their usual first arrival at Anaktuvuk. A few more
small groups were remarked, and on June 13 three were seen feeding
on the sandy shore of a slough, but the date of observation gave the
only suggestion that they were present for nesting. The migration of
semipalmated sandpipers through Old Crow was small and late.
Their nearest wintering range is in the gulf-coast States and the
migration northward passes through interior States and Provinces,
usually east of the mountains (A. O. U., Check-list, 1957).

The Indian name is Teggetselve.

Family PHALAROPODIDAE: Phalaropes

Lobipes lobatus (Linnaeus)

male May 24 weight 33.0 g. medium fat testes 7x13, 6x10 mm;
female May 23 weight 35.3 g. little fat egg 5.5 mm.
1 female May 24 weight 33.9 g. fat egg 4mm.

A few northern phalaropes were first seen on May 23, the day after
the ice moved in the river. For a few days some of the few phalaropes
seen appeared to be resting during migration, and courting pairs were
not recorded until May 29. Since the two female specimens had eggs
measuring 6 and 4 mm., courtship was probably in progress earlier
than it was recorded. During June individual phalaropes were often
seen floating on marshy pools in quiet contrast to their dashing speed
and remarkable aerial evolutions when in flight. A small migratory
passage through Old Crow was believed to occur, but a good many
phalaropes remain to nest in Porcupine Valley.

Believing that the other birds wintering in maritime situations come
to Old Crow from the Pacific we suspect that these phalaropes are
from the population wintering over the Pacific southward off Ecuador.
Swarth (1936) says they are common migrants through Atlin, which
they reached May 15, 1931, May 22, 1932, and May 12, 1934; but as
they are widespread in summer through Yukon (Rand, 1946) and
common all along the Yukon (Dall and Bannister, 1869), they may
pass from the Gulf of Alaska to the interior by numerous routes
through the mountains.

The Indian name is 7'revug.

469496—60——14

200 U. S. NATIONAL MUSEUM BULLETIN 217

Family STERCORARIIDAE: Jaegers, Skuas

Stercorarius parasiticus (Linnaeus)

1 male June 14 weight 579 g. little fat testes 12x20, 11x16
mm,

The only record of a parasitic jaeger was obtained from the specimen
brought to us. It seemed to be recognized by Joe Kay, who gave as its
Indian name 7tze kug. From the size of the testes it was in breeding
condition.

It is to be suspected that these jaegers winter with the Pacific popu-
lation which ranges off shore as far north as southern California.

Stercorarius longicaudus Vicillot

1 male May 26 weight 313 g. fat testes 8x17, 8x13
mm,

Long-tailed jaegers were seen only above the tree line on Old Crow
Mountains, where the specimen listed above was shot. Six were re-
corded on May 27 on or over the sedge hummocks of the saddle between
the southern and central mountain. Judging from their actions they
had already paired and several pairs appeared to have settled upon
nesting localities. The testes of the specimen were about mature in size.

It is to be suspected that this jaeger would have wintered with the
Pacific population off shore from South America south of latitude
10S:

Family LARIDAE: Gulls, Terns
Lorus hyperboreus barrovianus Ridgway

Larus argentatus smithsonianus Coues

1 male in adult June 11 weight 1435 g. very fat testes 10x12, 6x9
plumage mm,

1 female in sec- June 12 weight 1006 g. little fat egg 5mm.
ond nuptial i
plumage

It came as a surprise on May 15 to see overhead four adult glaucous
gulls flying swiftly eastward in compact formation close above the
trees at the edge of the steep bluff overlooking the mouth of Old Crow
River. Soon after another group of four appeared on the same
course, then five and again four, all within an hour. It became evi-
dent that their course along the bluff gained advantage from a local
wind condition by which their eastward flight was favored in the
prevailing northeasterly wind, for the gulls lost speed in the turbulent
air as they rounded the point. Then they found a northerly course
with favoring wind conditions, resumed speed, and disappeared up
Old Crow River.

OLD CROW 201

That afternoon and on May 16 these impressive flights continued
to be composed mostly of four, but sometimes of five, or three gulls.
Herring gulls, which the Indians call Tetyet kkya, were often among
the groups, in a minority of about one in four. The number four also
prevailed in the mixed flights and their formation and course were not
different from the flights composed exclusively of glaucous gulls. A
few of the flights passed eastward over the Porcupine River. On that
day about 100 large gulls were carefully watched from their first
appearance about two miles below the village. At the first sight of
them it appeared as if their course was set and there was no hesitation
at the mouth of Old Crow, where most of the flights turned north as
if prepared by anticipation for the change in course.

The flights of these large white birds were so conspicuous and so
obviously a purposive migration that it seemed warranted to estimate
that the 100 gulls carefully watched comprised a sample numbering
possibly ten percent of the two-days’ traffic and perhaps less than
one percent. This assumption leads to the estimate that between one
and ten thousand large gulls flew up the Porcupine to Old Crow on
May 15 and 16. Peter Lord first noticed large gulls flying north over
Crow Flats on May 16.

On May 17 most flights continued up the Old Crow, but an increas-
ing number went eastward over the Porcupine River. These two
migratory courses were followed in similar fashion until May 22, but
on some of the later days the route to the village passed over the river,
in which the ice was beginning to move. The orderliness of the flights
diminished as herring, and more rarely glaucous, gulls occasionally
stopped along the swift running river after the breakup. Only adult
gulls were noticed before the breakup.

Glaucous gulls are rare in the interior of Alaska south of the Brooks
Range in summer. One in second-year plumage was taken at Old
Crow November 28, 1957.

Since pairs of herring gulls in breeding season are seldom as close
as a mile apart on the large rivers in the arctic interior, one day’s
migratory flight past Old Crow could probably introduce the entire
nesting population of herring gulls found on the upper Porcupine.
No significant part of the more numerous migration of glaucous gulls
remains in the interior, It is likely that this impressive migration of
glaucous gulls and fewer herring gulls brings up the Porcupine River
from Alaska an important contribution to the summer population
of large gulls on the shores of Mackenzie Bay and the Delta. The
glaucous gulls were first sighted at Reindeer Station in the Delta on
May 15 (Porsild, 1943, p. 30), and the summer population of western
glaucous gulls extends along the American arctic coast about to An-
derson River (Snyder, 1957).

202 U. S. NATIONAL MUSEUM BULLETIN 217

Peter Lord said that large gulls did not remain on Crow Flats in
summer. In June we obtained two specimens of herring gulls on the
Porcupine River and saw a number of adults apparently settled.
The male specimen, in adult plumage on June 11, by the size of its
testes was judged to be not then in breeding condition. The female in
second-year plumage had eggs somewhat enlarged, but is supposed not
to lay for another year. We saw no glaucous gulls on the Porcupine
after migration.

The two specimens were identified as Larus argentatus smithson-
zanus and not L. a. thayert. Judging from the date they were sam-
ples of the summer resident birds and do not represent the migrating
gulls with dark wing tips, which might have included thayeri in the
great transarctic flight which carries them from their Pacific winter-
ing place as far east as Greenland. Thayer’s gull is abundant in winter
in southern British Columbia (Munro and Cowan, 1947, p. 117), and
it is remarked in migration along the southeastern coast of Alaska.
Apparently the next common records are from the Alaskan arctic
coast, where numerous specimens have come from the vicinity of
Barrow. There Thayer’s gull is reported as a regular fall migrant
by Bailey (1948), who remarks that there are few spring records.
Tt might be that the spring migration of 'Thayer’s gull crosses arctic
Yukon from the Gulf of Alaska. Such a course is apparently taken
by black brant.

If we believe that the association of glaucous and herring gulls
has been maintained since their departure from the sea, the migra-
tory course has probably not followed the Yukon River from Bering
Sea, because glaucous gulls seldom pass up the River (Dall and Ban-
nister, 1869), and such a migration as we saw at Old Crow would prob-
ably have been noticed. Glaucous gulls have not been earlier reported
south of the arctic coast in Yukon (Rand, 1946) nor have they been
remarked at Atlin (Swarth, 1936). However, Nelson (1887, p. 52)
reported that fur traders had taken glaucous gulls at Fort Reliance,
on the upper Yukon, on September 28 and October 18 when ice cov-
ered the river. He also remarked that Dall had obtained the young
from the famous arctic collector Lockhart, who found them at Fort
Yukon. The mystery of how these conspicuous gulls could pass unob-
served from the Gulf of Alaska or the Peninsula to Old Crow should
check speculation, were the answer less interesting.

Larus canus brachyrhynchus Richardson

1 male May 16 weight 435 g. very fat testes 12x20, 11x14
mm.
1 female May 16 weight 436 g. very fat egg 5mm.

Three mew gulls on May 3 were the earliest: gulls seen flying over
the frozen river. Until May 9 groups as large as 12 settled on the

OLD CROW 203

ice, and later, as small pools appeared, floated on them. Like the
early ducks and geese the early arriving mew gulls which landed
remained still, as if resting. There was certainly scant food available
to excite the gulls on the icy river and snow-covered land, and they
seemed to accept the scarcity without wasting efforts on search.
Unlike the direct flight of the large gulls the flight of mew gulls ap-
peared casual and easily diverted, although there was little difference
to be seen in the choice of the frozen landing places. It was clear that
they came up Porcupine River from the west and apparent that most
of them continued, or after a rest departed, eastward.

Between May 9 and 11 the eastbound traffic past the village became
more intense. In their wavering flight and irregularly organized
flocks these gulls nevertheless make swift progress. The flights com-
prised from 1 to 16 gulls. On May 13 few were seen and on May 14
none. It seemed that the early migration had left no settlers at Old
Crow.

On May 16 a compact flock of 30 settled on the ice in mid-river.
Four Bonaparte’s gulls were with this flock, keeping near each other,
but not otherwise discriminating as to position. Three times this flock
made a great crying as it was disturbed, but alit again nearby after
each disturbance. When two were wounded the flock scattered and
individuals flew close to the wounded birds on the ice until I ap-
proached. The flock then reassembled and after many circles disap-
peared on an erratic eastward course. This was the last day on which
we crossed the river on the ice. From May 15 until May 19 small
flocks and single birds occasionally alighted, but it seemed that most
of them continued eastward.

As the water began to run over the ice the migratory trend became
less evident and gulls often left the river as a course. When the ice
started moving on May 22 it appeared that many mew gulls were
scattering over the country nearby, although a continuing easterly
trend in the flights suggested that some migration persisted until
June.

Peter Lord did not remark any gulls on Crow Flats until May 16.
He said that only small gulls remained there in summer, and that they
were common. Lazarus Charlie reported the first sea gull at Johnson
Creek, about 80 miles south by east on the upper Porcupine on May 7.
Spring weather is said to come earlier there than at Old Crow, and
migration of a few species was reported in the upper valley of the
Porcupine before it reached Old Crow. It seems likely that many of
the earliest migrating mew gulls, passing Old Crow when the country
seemed destitute of opportunity for gulls to feed, were following
Porcupine River toward the more advanced spring in its southern
valley.

204 U. S. NATIONAL MUSEUM BULLETIN 217

These hardy little gulls, graceful and swift in flight, often wheel
and tower high in the air. The desultory appearance of their flights
made it appear as if they were not bent upon long migratory course.
They are likewise the first gulls to fly through Anaktuvuk in the same
manner early in May. There they settle the arctic interior more
densely than do glaucous gulls, but they are not common along the
coast. The desultory nature of the flights of mew gulls through Old
Crow and the large number which remain in Porcupine Valley make
it difficult to speculate upon their eventual destination. But the great
numbers which moved up the Porcupine suggest that mew gulls from
those flights overflow Porcupine Valley and contribute significantly
to the nesting population in the Mackenzie Delta.

These gulls winter on the Pacific coast from southeastern Alaska
southward. Since they arrive in migratory flights from the west it
can be suspected that they come from the Yukon Valley but we do not
know by what course they leave the Gulf of Alaska and traverse the
mountains. Swarth (1936) reported their arrival at Atlin, B. C., on
May 7, 1932, May 3, 1933, and April 22, 1934. These dates are early
enough to make the vicinity of Atlin a possible area for crossing the
mountains to reach the Yukon system.

The Indian name for these gulls is Vyouw.

Larus philadelphia (Ord)

Four Bonaparte’s gulls were seen May 16 with 30 mew gulls. An-
other group was later seen with a flock of mew gulls. They were
known and named Chit tryo by Joe Kay and other Indians, and are
evidently common enough in summer in the upper Porcupine Valley
to be familiar, but in small numbers compared with mew gulls. Their
occurrence in company with the mew gulls suggests that they also
came from the west along the Porcupine River and had wintered on
the Pacific coast south of Washington.

Sterna paradisaea Pontoppidan

Only a few arctic terns were seen near Old Crow. Mary Lobban
first reported them: on June 5 about ten miles down river from Old
Crow. - Robert Bruce saw one near the village on June 14. On
that same day it was remarked that no tern was seen in traveling’:
40. miles up the Porcupine River and back. To the Indians terns.
are familiar and not well liked because of their persistent diving
at and even striking intruders into their nesting areas on Crow
Flats. These terns are probably from the migratory population
which travels the Pacific to their wintering place on antarctic waters.
(Cooke, 1911). Probably they migrate to Yukon from the west.

The Indian name is Kkya notetuthga. .

OLD CROW 205

Family STRIGIDAE: Typical Owls

Indians, like Eskimos, are interested in owls, take pains to observe
them, and are clear in relating what they know about them. We
did not see short-eared owls at Old Crow and our illustration and
description did not evoke recognition of them which could be con-
sidered to be explicit. The resident owls and snowy owls were well
known.

Bubo virginianus lagophonus (Oberhoiser)

1 male May 3 weight 1445 g. fat testes 7x1i, 6x9 mm,

John Moses brought us the great horned owl listed above from
his camp 10 miles down river from the village. He reported that
two owls were about a nest, but his later examination showed the
nest to be old and the second owl could not be obtained. The testes
of the specimen were less than breeding size, and the date of nesting
would more probably have been earlier than later. On May 11
Francis Williamson observed a great horned ow] hunting over the
village at midnight. A feather thought to be from a great horned
owl was found in May near a slough a mile south of the river, and
in this area on June 5 Robert Bruce and I saw a large brownish
owl while it made several short flights through the woods during
which a rusty blackbird was able to engage the owl more closely
than we could accomplish.

Great horned owls, because of their conspicuous appearance and
calls rather than for their abundance, are well known to the Indians,
who call them Veesay. Judging from what we could learn the
residence areas of these owls are a number of miles apart.

From examples of northwestern horned owls in the U. S. National
Museum this specimen differed in the general light color, small amount
of brown, and the contrast of the black and white markings. When
two additional examples, obtained in central arctic Alaska, had been
borrowed from George Sutton of the University of Oklahoma who
has designated them B. v. lagophonus, it became apparent that the
specimen from Old Crow was a rather light example of lagophonus,
distinguishable from algistus, saturatus, and most readily from wap-
acuthu. It is the form so far recorded in arctic Alaska and Yukon,
whereas wapacuthu is the form to be expected in the Mackenzie
Delta.

Nyctea scandiaca (Linnaeus)
None of our party saw a snowy owl, but to the Indians at Old

Crow who call them Miseitivay, they are well known as occasional
winter visitors and are more frequently seen on Crow Flats.

206 U. S. NATIONAL MUSEUM BULLETIN 217

Surnia ulula caparoch\(Miiller)

1 male Apr. 17 weight 322 g. medium fat testes 7.5x11, 5.5x9
mm,

1 female Apr. 10 weight 384 g. medium fat egg 2.5 mm.

1 female May 2 weight 310 g. no fat egg 8mm.

1 female May 4 weight 336 g. no fat egg 8mm,

The first hawk-owl obtained made itself known by repeating a
low call like a robin’s note of alarm as it sat high in a willow on
the island east of the village. It had a layer of white abdominal
fat several millimeters thick and nearly as hard as beef tallow at
ordinary room temperature. Hawk owls are familiar to the Indians
for their boldness, since they often come into the village to perch
on poles. The eggs in the specimens taken May 2 and 4 were 8 mm.
in diameter. These birds appeared to be in condition for laying
early in May. The May 2 specimen was taken from the top of a
broken spruce near a cavity which looked as if it had been used last
year for a nest and which showed some signs of preparation for use
this year. About two feet lower was a flicker’s hole apparently con-
structed about a year ago but not in use this year.

Pairs of these small but demonstrative predators are apparently
distributed at intervals of only a few miles in the arctic forests from
Mackenzie across Yukon and Alaska. The Indian name is 7’chichitoo.

Strix nebulosa nebulosa Forster

1 female May 3 weight 1092 g, no fat egg 2.5 mm,

The specimen of great gray owl listed above was first seen flying
among the spruce trees which predominated in the area at the base of
the bluff a mile north of Kenneth Nukon’s cabin, and was pursued
for a mile of short flights. Its appearance of large size belied its
actual weight, for although the thick feathers of the great gray
owl make it look larger, great horned and snowy owls are much
heavier birds.

There are few specimens or even sight records of great gray
owls in western arctic America and the birds seem to be more widely
separated than great horned owls. They are well known to Eskimos
and Indians in the arctic forest, and from their reports I believe
that great gray owls sparsely occupy the arctic forest across Yukon
and Alaska. Joe Kay discussed owls and their habits in detail, even
remarking that great gray owls are not bad eating, while the slightly
more common great horned owls are not particularly good. As a
matter of fact owls are seldom eaten, but his people have accumulated
and preserved every observed detail about the birds of their country.

The importance of owls for food is calorifically small, but Indians,
Eskimos, and some arctic travelers have encountered situations in

OLD CROW 207

which a bird or animal usually disregarded as food nevertheless
served for a meal. Eskimos and Indians relate these experiences as
objective comparative characterizations of the flesh and structure
of animals rather than from gastronomic interest. The Indian name
is Nastok.

Aegolius funereus (Linnaeus)

Although we did not see a boreal ow] they were obviously familiar
to the Indians at Old Crow, who call them Wastotesul. Joe Bryant
of the Canadian Fish and Wildlife Service reported that he saw an
owl of this size about April 5 at Crow Flats, and Albert Abel gave
a good description of one which he saw there on April 16. Although
I have not seen one of these owls alive they are clearly described
by Indians and Eskimos as not unusual sights in the arctic forest
of Yukon and Alaska.

Family PICIDAE: Woodpeckers, Wrynecks

Colaptes auratus borealis Ridgway

1 male June 4 weight 154 g. no fat testes 6x15, 7x8
mm., brood path

1 female June 4 weight 152 g. little fat egg forming 3 rupc-
tured follicles,
brood patch

A flicker was heard calling north of the village on May 16 and
near Crow Point later on the same day. On May 19 Francis Wil-
liamson saw one near the village. On June 4 one was calling as it
moved about in the mixed spruce and birch near Kenneth Nukon’s
cabin. After a prolonged search Robert Bruce collected a pair when
they came to inspect an old flicker’s hole near the nest of a pair of
hawk owls (see p. 206). This hole had not been used this year by
the flickers, but the condition of the female showed that she had
first laid about June 1.

Flickers occur in the arctic forest from Mackenzie across Yukon
and Alaska. In migration they pass through British Columbia
(Munro and Cowan, 1947) as well as the Provinces to the east, but
the wintering area is too broad to assign a probable migration route
to the birds of the northwestern arctic forests. Their Indian name
is Chut lut.

Picoides tridactylus fasciatus Baird

male June 14 weight 61.3 g. little fat testes. 3.5x5, 4x5
mm,
John Moses sent word that a nest of this northern three-toed wood-
pecker was near his camp. Leonard Peyton found it about four feet
above ground in a stump and took the male, which was inside with sev-

208 U. S. NATIONAL MUSEUM BULLETIN 217

eral nestlings. The bird had a brood patch, but its testes had appar-
ently regressed from breeding size. The eggs would have been laid
about June 1. The resident range of this race extends in the arctic
forests from Mackenzie (Porsild, 1948) across Yukon and Alaska.
The Indian name is 7'utchun Tsya. 3

We looked carefully for signs of woodpeckers’ work. Dead birch
or poplar, even on burned areas, seemed to rot rapidly and seldom
bore visible marks of woodpeckers. Dead spruce stumps were rare.
Several were noticed with holes suitable in size for flickers and
about six had old holes suitable for three-toed woodpeckers, but none
were found that were small enough to be appropriate for downy
woodpeckers.

Family TYRANNIDAE: Tyrant Flycatchers

Seyornis saya yukonensis Bishop

1 male June 5 weight 23.7 g. medium fat testes 3x7, 3.5x4.5
Inm,.

Leonard Peyton collected the male phoebe listed above as it flew
in to light on a tent frame in the village. Its testes appeared to be
near but not at breeding size. On June 6, Williamson saw a phoebe
carrying nest-building material near the cliff opposite the mouth of
Dave Lord Creek, but search there on two later occasions did not dis-
cover the phoebe again. The Indian name is 2 kut ttst.

Yukon phoebes have not been found west of Anaktuvuk but evi-
dently extend across arctic Yukon and western MacKenzie. Since
they are usually east of the Cascade and Coast Ranges in British
Columbia (Munro and Cowan, 1947) they probably migrate through
that part of British Columbia.

Empidonax traillii (Audubon)

1 male June 12 weight 14.9 g. little fat testes 4x7, 4x5.5
mm.
1 female June 17 weight 12.4 g. little fat eggs 3mm.

Francis Williamson reported that Traill’s flycatchers were at Fort
Yukon when he arrived there on May 10, a date which seems early
for flycatchers to be in that northern locality. The first record near
Old Crow was of one heard on May 27 in the willow-alder brush
adjacent to the village. Thereafter one was occasionally heard or
seen. They were not easily discovered in the thick brush where they
were calling, but since Leonard Peyton located four by their singing
positions on June 12 they were apparently not uncommon. The
testes of the male specimen appeared to be at breeding size; the female
contained a small egg but had not yet laid. Although nesting appears
demonstrated by the behavior observed during nesting time its date
is not known.

OLD CROW 209

At Fort Yukon in 1958, Williamson and Peyton found Z. traillia
on May 7, a week after they arrived there.

The difference in date of arrival between Fort Yukon and Old
Crow suggests that migration does not proceed between those two
localities and suggests arrival of the migrants from the southeast.
Since they remain east of the Cascade and Coast Ranges in British
Columbia that is perhaps the western limit of their migratory course.
This flycatcher is known by the Indians as St tri gichi 2zeh.

Nuittallornis borealis (Swainson)

lmale May 24 weight 35.5 g. medium fat testes 4.5x12,
4.5x9.5 mm.

Francis Williamson recognized this olive-sided flycatcher by its
call on May 24. We followed the bird about among mixed birch,
poplar, and spruce at the top of the bluff for about an hour without
clearly sighting it, because it kept changing its calling position about
every five minutes as it moved around in an area about 34 mile across.
Robert Bruce, who was hunting separately in the same area, finally
shot it from the top of a poplar. Judging from the size of the
testes it was near condition for breeding.

This flycatcher nests in the forested areas of central arctic Alaska
(A.0.U., Check-list, 1957) and arctic Yukon, but has not been re-
ported in arctic Mackenzie. The more southern range from coast to
coast does not indicate a common migratory pathway for the species.

The Indians know it as 72¢v7.

Family ALAUDIDAE: Larks

Eremophila alpestris arcticola (Gberholser)

Males Females

Weight Fat Testes Weight Fat Eggs
Date (g.) (mm.) (g.)

May 25 36. 3 MF 5.5x9.5, 6x7

May 26 36.9 MF 5x8, 5x6

May 26 37.5 VLF 5x8, 5x6, brood patch 37.5 LF 4 broken follicles, formed
egg, brood patch

Horned larks were first seen as they were singing on the southern-
most of the Old Crow Mountains on May 25. Estimating from the
condition of the female on May 26 the first egg had been laid about
May 22, and the larks must have arrived at least a week earlier, as
is their schedule at Anaktuvuk.

Horned larks were seen only on the dry and barren rocky mountains
where a few pipits, wheatears, and rock ptarmigan somehow existed
in spite of the barrenness. Whenever the sun was warm many spiders
and some other insects could be seen, and these undoubtedly served
the small insectivorous birds. The occurrence of two medium-fat

210 U. S. NATIONAL MUSEUM BULLETIN 217

birds shows that even in the barren appearing mountains there is
enough food.

The arctic breeding range of this race extends across Alaska and
Yukon and to Mackenzie (A.O.U., Check-list, 1957) although Porsild
mentions that /. a. hoyti occurs on the Mackenzie Delta. The winter-
ing range in the northwestern States indicates that these larks migrate
through the western mountains.

The Indian name is Katu.

Family HIRUNDINIDAE: Swallows

Four species of swallows extend their nesting range beyond
the arctic circle in Yukon and Alaska and two nest also in arctic Mac-
kenzie. These western ranges are far north of their limits in the east,
and indicate northward migration through the mountain States and
Provinces. Inasmuch as the northwesterly nesting races of several
eastern species of birds extend farthest north, the tree swallows of
Alaska may come from eastern wintering situations. ;

The four species have quite separate wintering areas, but view-
ing their entire distribution only the violet-green swallows (7 achy-
cineta) are restricted to the west. The early presence of violet-green
and tree swallows at Fort Yukon is so far ahead of their arrival at
Old Crow that migration evidently does not enter the Porcupine Val-
ley from Alaska and probably does not pass westward from Old Crow
along the Porcupine River.

Tachycineta thalassina lepida Mearns

1 male May 21 weight 18.5 g. fat testes 4.5x6, 3.5x4
mm.

This violet-green swallow was brought to us by Joe Kay who
recognized it among the tree swallows then common about the village.
By comparison with breeding tree swallows the testes of this violet-
green swallow were not mature. Kay explained that violet-green
swallows are not often to be expected around the village, for as indi-
cated by its Indian name, 7ta shait sove, it is a mountain swallow.
Francis Williamson had seen them when he arrived at Fort Yukon
on May 10. On June 21 a pair occupied a nest box at Old Crow.

The nesting range of these swallows extends in Alaska northward
to Bettles but they are sometimes driven from their nesting places
by the more aggressive tree swallows, which are more numerous
where their common range just passes the arctic circle. Southward,
violet-green swallows nest in western Alberta and British Columbia
and migrate to southern California and Costa Rica in winter.

OLD CROW 211

Iridoprocne bicolor (Vieillot)

Male Female
Wei Fat Testes Weight Fat Eggs
Date (9. (mm.) Od
May 16 20. 8 F 7x10, 7x9 21.8 F 1
May 20 22.9 F 7x9.5, 8x10.5 24.2 F 1
May 22 22.7 F 7x10, 8.5x11
May 24 21.4 F 1
— 19.8 F 1
(aver.) 22.1 21.8

The first tree swallow was seen at about 2:30 p.m. on May 16, and
within a few hours a number of them were inspecting the bird houses
about the village. The numbers increased until on May 23 about 30
were circling in an area about a quarter mile long over the river bank.
A little later the group comprised about 40 which were coursing in
circuits about 300 yards in length over the meadow retrieving insects
from among the flooded grasses. At this date they were still flying
actively at 8:30 p.m. Toward the end of May the number of tree
swallows about the village diminished as every nest box became oc-
cupied by a pair. Some must have migrated further or dispersed
to natural nesting sites. Along the river tree swallows were occasion-
ally seen but their natural nesting places were not discovered.

The number of tree swallows, which the Indians call Sha sove, was
much exceeded by the cliff and even more numerous bank swallows.

It was noticed that some tree swallows with scarcely a trace of blue
on their slate-gray backs were mating as females and occupying nest
boxes. Two gray females taken for examination were indistinguish-
able in appearance of maturity from blue females.

Tree swallows are not reported from the Mackenzie Delta (Porsild,
1943) but extend across the wooded portion of arctic Yukon and
Alaska. Their arrival was surprisingly early for an insectivorous
swallow, preceding the breakup of river ice by six days, but all speci-
mens were fat. In view of their early commonness in British Colum-
bia (Munro and Cowan, 1947) and arrival April 10 at Craig, Alaska,
it is likely that the migration passes to the northwest through many
western valleys and is likely over a western flyway.

Riparia riparia riparia (Linnaeus)

On May 31 a few bank swallows were noticed, and by June 1 they
were as numerous about the village as tree swallows, whose feeding
areas they shared. Their number then increased until it exceeded by
many times that of the other swallows.

In groups of various sizes bank swallows nested in selected situ-
ations along the river. Their numbers are tremendous, but the greatest
concentration was noticed in the vicinity of the village.

242 U. S. NATIONAL MUSEUM BULLETIN 217

On June 3, holes were started by one group of about 50 swallows
along a section of bank, and in a few days there were 30 or 40 holes
under construction within a stretch of 50 feet. This particular group
seemed to swarm together as they flew in their unbelievably swift
banks and turns, giving the impression of social organization in the
pattern of flight. Many other holes were started in the bank in front
of the village and scattered or concentrated groups of holes were
formed in many banks exposed by the rapid fall of the river. ‘The
swallows are adept at selecting banks which do not disintegrate, as in
many places they do during the summer by thawing and drying. The
swallows also selected nest sites in firm but early-drying sandy loam.
The construction of holes was still in progress on June 15. It is
obviously a laborious operation for such small birds with their slight
equipment for digging.

A recorded first arrival (Bent, 1942) in Fairbanks, May 14, pre-
cedes the arrival at Old Crow by two weeks. Alfred Gross (Bent,
1942) refers to an account, written for him by Frank L. Farley, of
a large migratory flight of bank swallows passing northward over
the Athabaska River about 125 miles northwest of Edmonton on
May 11. Since they are reported to be rare west of the Casgade and
Coast Ranges (Munro and Cowan, 1947), it is presumed that the
northward migration travels through the Mackenzie Valley.

The Indians name this bird Shai tso ve.

Petrechelidon pyrrhonota hypopolia Oberholser

Males Females
Weight Fat Testes Weight Fat Eggs
Date (g.) (mm.) (g.)
June 10 30. 2 MF 7xi0, 10x10
June 14 27.0 MF = 9x11, 8x8 30.1 MF 1 in oviduct, 1 broken follicle, brood
patch
June 14 30. 0 F 1 in oviduct, 2 broken follicles
June 23 MF 5x9, 7x8
June 17 5 fresh eggs, 4 slightly incubated

eggs

A single cliff swallow was noticed on the evening of May 25. During
the next day their number increased, and on May 27 was twice that
of the approximately 50 tree swallows which had arrived in the village
10 days earlier. The arrivals at once set about examining the mud
nests remaining from.last year under the eaves of buildings. Many
old nests had been destroyed, but those most nearly intact were first
claimed and their reconditioning was begun. These did not suffice
for all the swallows, and work was gradually started on badly broken
nests and at new positions. The construction was a laborious op-
eration, and often the instinctive engineering was frustrated by the

OLD CROW 213

choice of unsuitable material or situation. Progress was slow, but
eventually about a hundred nests were completed on various buildings.

On June 14 a female was found to have laid an egg. On June 17
one nest contained four slightly incubated eggs, the first of which
would have been laid about June 9, and another five fresh eggs, the
laying of which would have started a few days later. At this date
many nests were still uncompleted, and laying probably extends
through June,

At several cliffs along the River, colonies of swallows were found
nesting. The nests were usually concentrated on firm rock faces.
In a few sites the droppings of swallows, accumulated in stalagmites,
showed the length of the nests’ occupation.

In the villages people often destroy the nests and sometimes block
the approach to the site with wire. Then the determination of the
birds is increased and for several days they may mill about trying to
replace the destruction or to evade the obstacle. The destruction of
nests is not wanton but because the droppings smear the windows,
assault the person and the birds are reputed to bring noxious vermin
unpleasant for man, a fair indictment since they are unnaturally using
our habitations.

On three cliffs extensively used by swallows, peregrines were seen
at nesting time. At one cliff where a peregrine was sitting on her
eggs a swallow’s nest was within 100 feet, and over 100 nests were
within 200 yards. A peregrine’s perch with abundant guano to show
its long use was within 6 feet of a cluster of nests, near which swallow
guano also showed their long use. We observed these in winter and
did not actually see the predator and swallow in such close proximity,
but during June, when we passed the cliffs, the swallows and their
nests showed no signs of disturbance.

Cliff swallows nest across the wooded part of arctic Alaska, for
IT have found them common along Koyukuk River and they are
known at Kobuk, but specimens are not available for identifying
the race in interior arctic Alaska.

In spring migrating cliff swallows move more rapidly northward
in the far west’ than in the east and at a much accelerated pace
through British Columbia, reaching far higher latitudes in Alaska
and Yukon than in the rest of Canada (Lincoln, 1952). Their
arrival at Atlin, B. C., was reported on May 21, 1934 (Swarth, 1936),
and at Peace River, Mackenzie, on May 20 (Bent, 1942). Our swal-
lows could have arrived by any course between these localities but
probably not from east of the Mackenzie River.

The Indian name for this bird is Shattso.

214 U. §. NATIONAL MUSEUM BULLETIN 217

Family CORVIDAE: Jays, Magpies, Crows

Perisoreus canadensis pacificus (Gmelin)

Males Females
Weight Fat Testes (mm.) Weight Fat Eggs
Date qg. q@.-) (mm.)
April 9 VLE 4.5x7, 4x6
April 15 85.9 VLE 6x9, 5x6 74.9 VLE 1
April 17 80.4 NF 5x6, 4x7.5
April 23 72.2 VLE 2x4, —
April 23 76.0 VLF 5.5x8, 5.5x6
May 1 73.5 NF 3x4.5, 2x3 61.5 NF 0.5
June 1 76.5 NF 2x4, 2x3
June 10 87.8 LE 2x3, 1.5x3 73.5 LE 0.5
June 17 75.4 LE
June 28 85.8 MF 2x2.5, 1.5x3 74.0 LF 0.3
Young Young
Male Female
May 30 66.5 NF
June 14 9 LF
June 28 73.1 LF

A few. Canada jays often came into the village looking for scraps
around the feeding places of the dogs. The Indians had some dis-
like for jays in addition to their prejudice against them for stealing
meat. Their name for them is 7%t¢mkotom.

Jays were encountered through the woods in situations indicating
pairs to be about a mile separated. On April 8 a pair appeared to
be courting, and during the latter part of April jays could often
be attracted by call and came in twos. Sometimes three appeared
together and on April 23 two males were called, and were collected
together.

Among the specimens collected the largest testes were found in
those taken April 15 and 17, and smaller testes were found in those
taken after April 23. The earliest young birds just flying were
seen on May 28 and recent fledglings were occasionally seen in early
June. Presumably the eggs were laid between middle and late April.

Old Crow and Anaktuvuk specimens of Canada jay had less brown
than P. c. canadensis, but matched satisfactorily with P. c. pacificus.

Corvus corax principalis Ridgway

Early in April one or two ravens were often seen searching for
scraps of dog food. At that time their flight was marked by calling
and maneuvering, but toward the end of the month linear flights
became more frequent, suggesting that nesting or feeding young
was in progress. No variation in the numbers of ravens was observed
which would suggest migration.

The Indian name for this bird is 7’atoo.

OLD CROW 215

Family PARIDAE: Titmice, Verdins, Bushtits
Parus cinctus lathami Stephens

We did not see any gray-headed chickadees. Olaus Murie (1928)
remarked that after nesting season they were rather conspicuous
along Old Crow River but that along Porcupine River he found only
boreal chickadees, an interesting distinction of range which he
thought to result from the preference of P. cinctus for the narrow
fringes of forest near the limit of trees.

Parus hudsonicus hudsonicus Forster

We occasionally saw boreal chickadees, usually in pairs, in a va-
riety of situations among thick willows and alders and in mixed
timber, but since they were rather quiet and secretive the popula-
tion may have been larger. The largest testes found were in
a specimen collected on May 21, but we have no other indication of
the time of laying. Olaus Murie (1928) found these chickadees
common along the Porcupine River but it is interesting that along
Old Crow he found none, although gray-headed chickadees (P.
cinctus) were common.

Our series of chickadees is uniformly a little paler brown on back
and crown than most examples of P. A. hudsonicus. Their tails are as
long as the longer examples among Alaskan birds. In these respects
they differ from the characters of P. h. farleyi, which Earl Godfrey
(1951) named as in the neighboring range of southern Mackenzie.
The specimens from Old Crow resemble two from the Savioyuk River,
at the northern limit of trees in central arctic Alaska. Our specimens
fit the vague characterization of P. h. evura by Coues (1884) and
Godfrey kindly informed us that he regarded three chickadees sent
to him from the John River, Alaska, as that race.

When Alaskan specimens were compared for degree of brownness
of back the palest and the darkest groups contained examples mingled
from eastern and western localities. Since we could not see that the
brown was distributed among Alaskan specimens according to a geo-
graphic pattern it seemed to serve no purpose to distinguish races on
this basis among Alaskan boreal chickadees. There is no advantage
apparent in naming the boreal chickadees of Old Crow differently
from the nominate race.

The Indians call this bird Tchichika.

469496—60——_15

216 U. S. NATIONAL MUSEUM BULLETIN 217

Family CINCLIDAE: Dippers

Cinclus mexicanus unicolor Bonaparte

We did not see in the ice-covered arctic rivers any of those infre-
quent areas where the water remained open and where dippers remain
in winter, usually in pairs. Stephen Frost carefully described two
dippers he had seen in April in the stretch of open water near the
mouth of Bluefish Creek and said that in February 1956 he saw similar
small birds at several of the open water stretches over 200 miles from
the mouth of the extraordinarily crooked Black River. Joe Kay con-
firmed these observations from his own much earlier observations and
named the dipper as soon as he saw its picture.

The Indian name for this bird is 722 rzuz.

Family TURDIDAE: Thrushes, Solitaires, Bluebirds
Turdus migratorius migratorius Linnaeus

Males Females

Weight Fat Testes (mm.) Weight Fat Eggs
Date (g.) Gg.)
May 15 71.5 LF 5x11, 8x12.5 75. 2 LF 2 mm.
86. 7 LF 6x14, 6x15
74.3 LF 7X12, 6.5x11
May 16 81.0 LE 7x11, 8x10 79.4 LF 1 mm.
76.2 LF 7x11, 7x10
June 6 80.0 F 9x14, 10x13
June 9 60. 2 LF 8x13, 9x15
59.1 9x18, 10x15
(aver.) 73. 5
June 2 4 fresh eggs, 4
slightly incu-
bated eggs

The first robin was seen on May 8. It was shy and quiet and a week
elapsed before more than one was noticed, although a few may have
been about but not observed. On May 15 a few more robins were
about and their numbers, noise and swift and often aggressive flights
rapidly became conspicuous until on May 18 the village appeared to
contain more than a suitable population for nesting. The robins
seemed to be of this opinion also for their quarreling continued until
the latter part of May. We then thought that some robins must have
left the area, for, in addition to the subsidence of their activity, their
numbers appeared to decrease.

Two nests were found on June 2. These we presume to have been
laid about May 25 and 27, 10 days after the conspicuous influx of
robins on May 15, but 17 days after the first was seen. In 1952 the
first robin was noted at Anaktuvuk on May 18 and the first egg was
reported on June 3, an interval of 16 days.

The maximum testes size was observed in a robin collected on June
9. Judging from the first eggs, some males were in breeding condi-
tion by May 25.

OLD CROW 217

Robins at Old Crow had little fat. Among 10 males at Anaktuvuk
in May one was fat, the others had little fat. They are one of the
arctic nesting species which are not fat during the time of arrival from
migration, but September arctic specimens may be fat and heavier by
about 10 grams.

Lazarus Charlie’s report of a robin on Johnson Creek on May 8 is
a date consistent with a migratory path through the southern part of
Porcupine Valley to Old Crow. Williamson found robins at Fort
Yukon when he arrived there on May 10 and they were reported arriv-
ing at Anaktuvuk on May 12, 1957. They were reported as first seen
at Frances Lake on April 22, 1943 (Rand, 1946). As Frances Lake
is 300 miles south by east from Old Crow there appears a discontinuity
in dates of arrival across the Ogilvie Mountains, which also appear to
offer a difficult migratory course. It can be suggested that robins may
enter the southern Valley of the Porcupine through Peel Valley from
Mackenzie Valley.

They probably travel the central flyway to the Mackenzie Valley
and turn westward to occupy central British Columbia, Yukon, and
Alaska.

The Indian name is Syo.

Ixoreus naevius meruloides (Swainson)

Males Females

Weight Fat Testes Weight Fat Eggs
Date (g.) (mm.) (g.)
May 4 71.5 LF 5x9, 4x7
May 20 66. 7 NF 7x12, 8x11
May 25 86. 5 F 4 broken follicles, brood
patch
June 1 2 fresh eggs
June 3 81.5 F 6x10, 6x10 5 half incubated eggs
June 4 98. 2 MF 3mm.
June 5 4 slightly incubated

eggs

(aver.) 73.2

The earliest varied thrush seen was flying from the top of a spruce
into the willow brush along Dave Lord Creek on May 4, when the
first signs of melting were seen. Its testes were not so large as were
those of specimens taken later in May. No more thrushes were noticed
at Old Crow until May 16, when they were heard singing among the
spruce. The interval between the first sighting and evidence of com-
mon establishment was longer than for robins which were conspicu-
ously exposed about the village. After May 16 varied thrushes were
often heard but not easily approached.

The female collected on May 25 had begun laying about May 21.
This early date of laying suggests that pairs probably started prepara-
tions for nesting some 10 days earlier and that the first arrivals near

218 U. S. NATIONAL MUSEUM BULLETIN 217

Old Crow were unobserved. On June 1, 3, and 5 nests were found
containing eggs which would have been first laid about May 23, 27,
and 81, respectively. Within 100 yards of one of these nests were
three old nests among small spruce, a situation conforming to the
reported return of varied thrushes each year to a location close to
their previous nesting sites. The other nest found this year was about
a third of a mile distant.

The weight of the two female varied thrushes was greater than that
of the males. Probably this represents the condition of overweight
which has been frequently observed among laying females of other
species.

The range of this thrush to Kobuk and Old Crow indicates its
extension north of the arctic circle to near the edge of the forest
across Mackenzie, Yukon, and Alaska. Since it winters from southern
British Columbia to Baja California, it is a western migrant, prob-
ably passing through the mountain valleys.

The Indian name is Sya.

Hylocichla ustulata incana Godfrey

Males Females
Weight Fat Testes Weight Fat Eggs
Date (g.) (mm.) (g.)
May 24 31.6 M 4 broken follicles, brood
patch
May 29 28.8 VLE 7x12, 7x12
June 3 29.4 LF 7x8.5, 6x9
June 3 28.9 LF -_— 6x11, 6.5x10.5
June 23 26. 0 LF 5x7, 7x8
(aver.) 28. 2

Francis Williamson heard a Swainson’s thrush sing, and obtained
the female specimen listed above on May 24. From its condition
it must have begun laying about May 20, so that these thrushes must
have been about the earliest to arrive in the vicinity of Old Crow.
They were the most retiring of the thrushes, but they were by no
means scarce because they were heard frequently and seen rather
often although usually only briefly.

Sidney Peyton found a nest on June 29 with three fresh eggs in
a slender spruce about 15 feet from the ground, in a thick growth
of spruce. The first egg in this nest was laid about June 26, 37 days
after the first female had begun laying. It does not seem possible
that such a late laying could have followed successful fledging of
an earlier brood. Before the date when this late clutch was started,
the gonads of one male had been found at less than full size, and
June 26 is an uncommonly late date of laying for this latitude.

OLD CROW 219

A specimen which I obtained from Bettles on July 1, 1951, estab-
lishes the occurrence of H. uw. incana there in a habitat typical for
this thrush. Reports from Eskimos at Kobuk described the nesting
and singing of another thrush very much like the gray-cheeked in
appearance, but differing in song and nest, and in the typical habitat.
The range attributed to zncana (A.0.U., Check-list, 1957) should be
extended from northern Yukon across arctic forested Alaska to Bet-
tles, and I suspect to Kobuk, near the western extremity of the forest.

The Indian name for this thrush is 7’z2 chi tho.

Hylocichla minima minima (Lafresnaye)

Males Females

Weight Fat Testes Weight Fat Figgs
Date g.) @.)
May 31 28.6 VLF 8x12.5, 10x13
June 1 28.1 LF 3 mm,
June 3 31.7 NF 6x10, 6x10
June 5 29.3 LF 2 mm,
June 7 31.7 LF 6x11, 7x10
June 10 30. 4 LF 7x13, 9x13
June 12 30. 4 LF 9x12, 8x12 32.5 LF 4 broken follicles,

brood patch

June 13 30. 6 LF 7.5x12, 8x11 34.3 LF 10 mm.
June 20 30.7 LF 8.5x10.5, 9x10
June 22 26.8 LF 2.5 mm.

Francis Williamson reported first hearing a gray-cheeked thrush
on May 22. On May 23 I saw a gray-cheeked thrush as it dashed
from the willows in the swift pursuit of another, an action charac-
teristic of male thrushes when attacking invaders of their territory.
After that date thrushes were frequently heard and occasionally seen
in the thick willow brush back of the village.

The testes of the first male specimen taken were as large as any
found later. Date of laying can be set by the condition of the female
specimen taken June 12, which indicated laying prior to June 9, This
female and the one taken June 13, which contained an egg 10 mm.
long, were above usual weight, a condition frequently remarked dur-
ing laying.

The first date of laying by gray-cheeked thrushes was later than
was observed among the three other species of thrushes which were
common near Old Crow. It was remarked at Anaktuvuk that gray-
cheeked thrushes were not found nesting early, and at Kobuk Grin-
nell (1900) recorded their first arrival on the rather late date of
May 24.

The specimens from Old Crow and Anaktuvuk were typical of
the nominate race which extends from arctic northwestern Alaska
through southwestern Alaska and northeastern British Columbia.
In migration they use the Mississippi flyway to Central and South

220 U. S. NATIONAL MUSEUM BULLETIN 217

America. Their northward flight in spring must proceed to about
latitude 60° N. before turning westward for over 1000 miles to traverse
Alaska and penetrate deeply into northeastern Siberia.

The Indian name for this bird is 7zintzzo.

Oenanthe oenanthe cenanthe (Linnaeus)

1 male June 9 weight 32.0 g. little fat testes 5x8, 6x6 mm.
1 female June 9 weight 35.5 g. fat egg 10 mm., 4 empty
follicles

Wheatears were not observed on the southernmost of the Old Crow
Mountains but Robert Bruce and I found a pair near the base of the
next northern and higher mountains on June 9. Altogether we saw
about ten in a few hours while climbing over the barren rocky mount-
tain at elevations from about 2,000 to 3,500 feet. They were as shy
and restless as usual during nesting time in the Alaskan Brooks
Range. Joe Kay identified the specimens at once, giving their Indian
name as 7'tha Tze, and was familiar with their shyness on the rocky
mountain slopes.

The condition of the female listed above established first laying
about June 6. The soft fat of the wheatears was creamy white, as
was remarked at Anaktuvuk. The female was fat and above usual
weight showing its adequate nutrition while living on the barren
mountain during the intensive production of eggs.

Wynne-Edwards (1952) reported the southeastern record of these
Asiatic birds in America in the eastern part of the Mackenzie Moun-
tains at latitude 64° 20’ N., longitude 128° 20’ W., a point which he
remarks upon as not much more than 1,000 miles west of the western
record of wheatears from eastern America. They are reported sum-
mering in mountain areas of southwestern Yukon and along the arctic
coast (Rand 1946). Their migration leads from Alaska to the
Chukchi Peninsula and southwestward along the Stanovoi Moun-
tains to Udski and thence farther through the interior of Asia
(Stejneger, 1901, p. 474). The nearest reported regular wintering
place of this race is far westward from Alaska in North China, and
their migration which almost reaches the Mackenzie, leads them
farther eastward into the American Continent than any of the other
land species wintering in Asia.

Because of the large weight of these two wheatears, Bernard
Feinstein kindly reexamined the measurements of these specimens
and those from Anaktuvuk. He found them all to conform to
O. 0. oenanthe, no distinction being signified by these weights.

OLD CROW 221

Family SYLVIIDAE: Old World Warblers, Gnatcatchers, Kinglets

Regulus calendula calendula (Linnaeus)
1 male June 3 weight 6.7 g. no fat testes 4x5 mm.

On May 17 a kinglet was observed. The male which was collected
was in company with another, apparently its mate. Joe Kay remarked
that kinglets were sometimes seen in winter at Old Crow, where they
are called by the Indians Khut traluk. We have also had reports from
Eskimos at Anaktuvuk and Kobuk, Alaska, that kinglets were oc-
casionally seen there in winter, but these reports are not confirmed by
winter specimens.

In the northwest the kinglets are all 2. c. calendula. The winter
range of this race is eastward and southward from Nebraska (A.0.U.,
Check-list, 1957), so it is likely that the northwestern population
migrates southeasterly along the eastern parts of the mountains.
Records are scarce in British Columbia (Munro and Cowan,
1947), whereas Rand (1946) found these kinglets “fairly common”
in southern Yukon.

Family MOTACILLIDAE: Wagtails and Pipits

Anthus spinoletta rubescens (Tunstall)

13 males May 6-June 9 weight 18.8-23.6, fat (8), medium fat —_
average 20.7 g. (1), little fat (3),
no fat (1)

Joe Kay on May 8, brought in the first two specimens of pipits,
which the Indians call Kwit kkyo zyo. For six days they occasionally
alighted in the meadow by the river near our cabin. Some of the
flocks seen contained 12 pipits, but because they were usually so rest-
less we could not form an estimate as to how many were migrating
through Old Crow. The first eight specimens obtained near the
village and during migration were fat males with testes less developed
(5 to 7 mm.) than when they were found later on the mountain.

After May 15 pipits were not seen near the river. In June they
were occasionally heard and seen in grassy or rocky places above
2500 feet in the mountains where three males were collected. Their
testes were then enlarged, (8 to 9 mm.) they had little or no fat, and
appeared to be occupants of breeding territories.

Our specimens of pipits from Old Crow (13), Anaktuvuk (33),
and Ahlasuruk (2) are identified as A. s. rubescens, the range of which
is thus extended across arctic Yukon and Alaska to within 200 miles
cf the western arctic coast. <A. s. pacijicus has been named by Bailey
(1948) as the race at Barrow, and northern Alaska and Northeastern

222 U. S. NATIONAL MUSEUM BULLETIN 217

Siberia are given as the range of pacificus in the A.O.U. Check-list
(1957), which does not extend rubescens west of arctic Yukon. Since
Ahlasuruk River is about 200 miles from the western arctic coast of
Alaska, rubescens and not pacijicus extends across arctic Alaska at
least in the interior.

As this race of pipits winters east of Texas their migratory flight
probably passes east and west across most of Yukon (Rand, 1946).
Since they are not recorded from British Columbia (Munro and
Cowan, 1947) they probably travel the Mackenzie Valley and the
central flyway.

Family BOMBYCILLIDAE: Waxwings

Bombycilla garrula pallidiceps Reichenow

3 males Apr. 18-June 1 weight 49.6, 50.3, fat (2), medium fat —
average 54.0 g. (1)
4 females Apr. 18-June 29 weight 59.1-58.7¢. fat (4) —

A flock of ten Bohemian waxwings was seen on April 10 in their
characteristically erect positions on the tops of some birches and
poplars. Later they were frequently heard and seen in small flocks
flying, feeding, or resting in the tops of the birches and poplars.
Six of the seven specimens taken were fat. The testes of the last male
specimen, taken on June 1 were enlarged but perhaps not to breeding
size, and on June 29 a female specimen had a brood patch. These
dates bracket egg laying.

There was no apparent change in the numbers of waxwings present,
but flocks were not remarked after June 1.

The Indians spoke of waxwings as winter birds near Old Crow,
where they are called hut tsa luk, and at Kobuk they are considered
to be winter birds by the Eskimos. Rand (1946) called the waxwings
migratory in Yukon and we have not found reports of them in arctic
forests earlier than April or later than October 15. At these seasons
winter conditions prevail. The birds of several species in the arctic
forests are migratory in the sense that in winter flocks concentrate
and move for some distance in search of favorable feeding places.
If they are true forest-dwelling birds the trend of their concentration
and movement in the northern margin of forest could only take them
southward. It has also been remarked however that in central arctic
Alaska some chickadees and pine grosbeaks move north of the forest
on to the tundra in winter. We have not now the evidence which
would show in these wanderings the precision of course that charac-
terizes the great spring migrations. The distinction of the marked
and long migrations is not only their distance, but the regularity

OLD CROW 223

which demonstrates that they are developed through long experience
of the population and repeated through the operation of memory.
These are quite different influences from the current impulses ap-
parently directing movements of waxwings without known directionai
regulation.

Family LANIIDAE: Shrikes
Lanius excubitor Linnaeus

On April 16 a shrike lit in the top of the willows along the river
and after a few short flights from our pursuit disappeared among
the willows on the island just above the village.

Joe Kay recognized the illustration of a shrike and gave the name
T si kwut go katshilyi, which he said referred to the shrike’s habit of
hanging its prey on bushes. No other shrike was seen. Above the
timber on the mountains we did not find the situations in extensive
willow-filled valleys where shrikes are rather common in spring and
summer in central arctic Alaska,

Family PARULIDAE: Wood Warblers

Four of the wood warblers at Old Crow are assigned to races which
extend far eastward from Yukon. Vermivora celata celata, Den-
droica petechia amnicola, and D. striata nest across northern
Yukon and Alaska, while Wilsonia pusilla pusilla is replaced by
W. p. pileolata at Anaktuvuk. V.c. celata, D. p. amnicola, and D. s.
lurida seem to be confined to the northern part of Yukon. This area
serves as an arctic bridge connecting the Canadian and Alaskan
populations, which extend to the northern limit of forest.

It is difficult to decide whether the water thrushes from Old Crow
are Seiurus noveboracensis notabilis or S. ns timnaeus. If they are
notabilis they should be counted as eastern warblers, if Z¢mnaeus they
are of a western race, as is clearly the Old Crow D. ¢. hooveri. The
western races of warblers extend farther south in Alaska and Yukon
than do the four eastern races.

It is not yet clear whether the eastern races of warblers migrate, to
Alaska along the Porcupine Valley, where they nest, or through
southern Yukon, where the migrating races have not been clearly
separated. In fact better identifications and range determinations
of warblers in Alaska and Yukon will give significance to what now
appear to be peculiar distributions.

I am particularly indebted to Bernard Feinstein of the U. S. Na-

tional Museum for his help in making the comparisons by means of
which the wood warblers are identified.

224 U. S. NATIONAL MUSEUM BULLETIN 217

The species at Old Crow are listed below with regard to their
habitat preference and nesting abundance:

Order of Order of
habitat nesting

Species wetness abundance
Vermivora celata celata 5 4
Dendroica petechia amnicola 3 1
Dendroica coronata hooveri 6 5
Dendroica striata 4 6

Seiurus noveboracensis notabilis or

limnaeus i 2
Wilsonia pusilla pusilla 3
Vermivora celata celata (Say)

Males Females
Weight Fat Testes Weight Fat Eggs
Date (g.) (mm.) (g.) (mm.)
May 21 8.7 LF 4x5, 3x4.5 8.8 MF 1, 25
May 24 9.6 LF 4x5, 4x4 8.9 LF 1
9.1 LE 3.5x6, 4x4
i 9.5 LF 4.5x6, 5x6
May 27 9.7 VLF 5x7, 5x5
May 28 10.0 LF 4x45, 3x4
May 31 8.8 LE 4x6, 4x5
June 1 9.5 LF 4x5, 4x4 10.1 LF 5
9.1 LF 4x5, 4x4
June 10 9.4 LF 3x6, 4x6
June 29 10.0 LF 1.5
(aver.) 9.3 (Coeff. of var. 4.4%)

Two orange-crowned warblers were first heard by Frank Wil-
liamson on May 20 singing among the poplars and scattered spruce
along the slope of the bluff. Thereafter several might be seen in
that area daily, where they came readily and swiftly to a call. Fre-
quently a pair of birds came and pairs were seen among the earliest
orange-crowned warblers. Until May 28 these were the most com-
mon of the warblers, but thereafter water thrushes and yellow
warblers became much more numerous. This was in part because
of the rather limited areas in which orange-crowned warblers were
found. The testes of the males taken during the month showed no
trend in changing size.

On June 1 a female contained an egg 5 mm. long, which must
have been nearly ready for laying.

The nominate and most northern race of orange-crowned warblers
nests from Alaska to Quebec and winters widely across the southern
states. In Yukon this race has only been identified from Old Crow,
not from further south. The range of this eastern wintering race
seems to extend for about 1000 miles in arctic western America near
the border of forests.

The Indians call this bird Tz? vit tich kwatlo.

OLD CROW 225

Dendroica petechia amnicola Batchelder

Males Females
Weight Fat Testes Weight Fat Eggs
Date (g.) (mm.) (g.)
May 27 10.2 LF 4.5x7, 5x5.5
May 28 10.0 VLE 4.5x7, 4.75x 5
May 31 9.0 LF 5x7, 5x5 10.4 LF 6mm.

9.3 LF 4x7, 5x5
8.7 LF 6x6, 5x5
9.3 LF 4x5.5, 3.5x5

June 1 9.5 LF 5x7, 5x5 9.6 LF 3mm.
June 2 9.6 LF 4.5x8, 5x6

10.3 LF 4x6, 4x5
June 6 9.4 NF 5x7, 5x5 9.8 MF 2mm.

9.5 LF 5x7, 5x5
10.3 LF 5x8, 6x6

9.6 LF —, 5x6.5
9.7 LF —.. 4x4.5
June 7 8.5 LF 6x8, —— 9.5 LF 2mm.

9.8 NF xs,
9.8 LF 6x7, 4.5x6

June & 8.7 NF 5x7, 5x6 9.7 LF 1mm.
8.6 NF 5x7, 5.5x7 8.8 VLF 2mm.
June 10 8.8 LF 3mm.
8.5 LF 2mm.
June 12 8.1 NF 2x3, 5x6 9. 4 LF 2mm.
1 broken follicle
9.3 LF 4x5.5, 4x4 10.8 LF 1 egg forming
8.1 LF 2x5, 5x5
June 16 3sets of 5 fresh eggs
June 18 1set of 5 fresh eggs
June 20 10.5 LF 2mm.
June 23 1 set of 5 fresh eggs
(aver.) 9.3 (Coeff. of var. 7.0%.) 9.6

The first male yellow warbler was seen May 23 in the willows
above the overflow from the river. No others were noticed until
May 27, when they were occasionally seen in the willow-alder brush
on the flat behind the village. The numbers kept increasing until
about June 8, when they were the commonest birds in the vicinity,
for a pair might be seen every 50 feet along a trail for a distance of
several hundred feet. Apparently some pairs occupied a territory
with the center only 25 feet distant from the edge of a neighbor’s
territory. In other willow-alder thickets along the river many yellow
warblers were seen. Farther up the Porcupine River the prevalent
spruce reduced the extent of their habitat.

About June 8 their territories and mating appeared to be in settled
good order. On June 12 the testes of three male specimens either
had regressed or were still undeveloped, and a female specimen con-
tained a formed egg and a collapsed ovarian follicle. On June 16
three nests, each with five eggs, were found by Sidney and Leonard
Peyton, and on June 23 one with five fresh eggs was found. Yellow
warblers were the latest of the warblers to arrive and nest. The
first eggs in these nests were laid about June 10, 11, and 16 respec-

226 U. S. NATIONAL MUSEUM BULLETIN 217

tively. The first egg was thus laid 18 days after the first male was
seen and 14 days after yellow warblers became common.

The excess of weight in females over males may be related to the
presence of numerous females with enlarged eggs. In several other
species it has been remarked that gravid females are heavier than
those not laying. At no time during our observation were any of
the species of warblers found to be fat. Of 91 specimens only three,
all females, were found to be medium fat.

These yellow warblers and those from northern Alaska were all
consistent with the race amnicola, which ranges across the continent
undifferentiated from Alaska to Labrador. The arctic range, how-
ever, is confined to the area west of the Mackenzie River.

The Indians call this bird 7’seéso.

Dendroica coronata hooveri McGregor

Males Females

Weight Fat Testes Weight Fat Eggs
Date (CD) (mm.) @g.) (mm.)
May 20 12.0 NF 4.5x7, 4x4.5 11.8 NF li
May 21 11.8 NEF —, 3x5
May 24 18.5 LF 5x8, 5x7
June 3 13.0 LF —, 5x5
June 4 13. 2 NF 6x10, 5x6
June 28 14.0 LF 5.5x9.5, 8x8

(Aver.) 12.9

Frank Williamson heard the first myrtle warbler singing on May
18. Between May 20 and 28 an individual or a pair could occasionally
be called in the mixed spruce and poplars along the slope of the
bluff, but thereafter they were seldom noticed. Williamson found
them rather common at Driftwood River, and near Kenneth Nukon’s
cabin, 15 miles east of Old Crow, there were a few about on June 3,
4, and 5. No eggs were found but because of the presence of pairs
during the breeding season it is concluded that myrtle warblers
were nesting. The Indians call this bird Ayekyszez.

This northwestern breeding race of myrtle warblers migrates to
winter from southern Oregon through the valleys of California (Wet-
more, 1926, p. 206).

Dendroica striata (Forster)

Males Females

Weight Fat Testes Weight Fat 8
Date Go (mm.) (.) i
May 29 12.4 LF 5x7.5, 6x6
June 3 12.1 NF 5x9, 5x7
June 5 13.7 LF 5x7, ——

12.6 LF 5x7, 4x5
June 6 12.5 LF 5x9, 5x6
June 10 11.4 NF 4x6, 4.5x5.5
June 23 12.8 MF 5 heavily in-

cubated eggs

OLD CROW 227

The first blackpoll warbler was heard singing by Frank Williamson
two days before the first one was seen and collected on May 29.
Since they are considered common in far northern forests we searched
for them but saw fewer than of any other warbler around Old
Crow. At Kenneth Nukon’s cabin, 15 miles east of Old Crow, black-
polls were common on June 3. The only nesting record was June 23,
when Sidney and Leonard Peyton collected the female specimen with
5 well incubated eggs. The first egg was estimated to have been
laid about June 12.

The Indians call this this bird 7'22 wit sitik kwarzui.

At the U. S. National Museum, Bernard Feinstein remarked that
our blackpoll warblers and specimens from Alaska differed from
eastern specimens in having less black on the back and generally duller
gray upper parts, as described by Burleigh and Peters (1948), for
Dendroica striata lurida, a form not recognized by the A.O.U. Check-
list (1957). The area in which intermediates between Jurida and the
more eastern form were found was around Fort Severn, Ontario, and
Churchill, Manitoba, so that our Old Crow specimens were far from
the eastern border of the range of their race.

Seiurus noveboracensis notabilis Ridgway

Males Females
Weight Fat Testes Weight Fat Eggs
Date (g.) (mm.) (g:) (mm.)

May 21 17.4 VLF 4.5x7, 4x6
May 28 17.9 VLF 4x6, 4x5

17.7 VLE 5x5, 4x5
May 29 16.6 NF 6x7.5, 5x7

15.7 NF 4x4.5

16.9 LF 4,5x6
May 31 17.8 LF 5x6, 6X7
June 3 17.4 LF 5x7
June 5 15.6 VLE 5x6.5, 5x6 18.5 LF 2
June 7 16.1 NF 3x7, 4x5.5

17.3 LF 5x9, 6X7

16.4 NF 4x7, 5x6

18.1 LF 5x6, 5x5 17.9 NF 6
June 12 16.4 LF 4.5x6, 5x6.5 22. 5 LF 17
June 13 16.6 LF 5x7, 5x6
June 20 16. 5 LF 5x8

(Aver.) | 17.0 (Coef. of var. 3.8%.)

11 broken follicle, 1 egg forming.

On May 20 the first northern waterthrush was heard singing in the
willows, above the noise of the river, and thereafter their loud clear
song could be heard, often at considerable distance from the swampy
willow brush. The male would fly to the top of a willow, sing his
series of notes about half a dozen times and then dive groundward
into the dense brush. Within five minutes he might be up again for
a period of song, this time on another willow, until after occupying

228 U. S. NATIONAL MUSEUM BULLETIN 217

four or five positions he returned to a point near the first perch. The
song and singing positions were conspicuous but the thickness of the
brushy surroundings and the brevity of the bird’s appearances made
it difficult to observe them near the ground.

After several males had been collected, a special effort was made
to find females and evidence of nesting. The sexes could not be dis-
tinguished in appearance and apparently were not distinctive in action
in their wet marshy terrain under the thick willows, for only three
females were obtained. One of these, taken on June 7, had an egg
ready to lay and one taken on June 12 contained a formed egg and a
collapsed ovarian follicle.

The Indians know these birds as Chootzi.

After examining our waterthrushes at the U. S. National Museum,
Herbert Friedmann wrote me, “McCabe and Miller (1933) indicate
that S. n. notabilés exists in two geographically separate populations.
These birds [from Old Crow], from the northwestern of the two
groups, are darker above, varying in the direction of lémnaeus, from
which race some of the individuals are difficult to separate. This
raises the possibility that the western section of notabilis may better
be grouped with 7imnaeus, and in this way we can do away with the
discontinuity in the distribution of notabilis which McCabe and Miller
indicated.”

The migratory course of these waterthrushes does not appear defin-
able from the scattered records of localities. Since they approach the
western race in appearance it may be presumed that their migration
passes west of the Mississippi.

Wilsonia pusilla pusilla (Wilson)

Males Females
Weight Fat Testes Weight Fat Eggs
Date @.) (mm.) (g.)
May 20 8.5 NF 4x7, 4%4.5
May 21 7.7 LF 3.5x5, 3.5x4
May 22 7.3 VLF 0.75 mm.
May 29 7.9 NF 3x4, ——
June 1 7.6 VLF 5X7, 4.5x5
7.3 VLF 4x6.5, 5x5
7.7 LF
June 7 8.8 LF 4 broken follicles,
1 formed egg,
brood patch
June 8 7.5 NF 5.5x8.5, 5x6.5 8.1 LF 2mm.
June 10 8.3 LF 5x8, 5x6

7.3 LF 4.5x6, 5x6
(Aver.) 7.8 (Coeff. of var. 5.2%.)
The first Wilson’s warblers seen were singing in low willow brush,

where they were subsequently often seen, usually near running water.
The first part of their Indian name, 7’setso khekuz, means water.

OLD CROW 229

They appear in lower brush than waterthrushes and less commonly
over standing marshy water. The female specimen taken on June 7
indicated first laying about June 4. A Wilson’s warbler was seen in
the 4-foot high willow brush along a small stream valley between the
second and third of the Old Crow Mountains, above tree line and at
an elevation of about 2,000 feet.

I have found W. p. pileolata the form occurring along the head-
waters of streams flowing northward through the arctic tundra in
Alaska. This, together with Bailey’s (1948) reports on their occur-
rence in northern Alaska, establishes this species, the smallest in the
family, as the most northerly of the wood warblers.

On June 20 Sidney Peyton found six newly hatched young in a nest
well hidden in the moss at the bottom of a small bank. The first egg
was probably laid about June 4. On June 21 snow began to fall, and,
on the next day it was 6 inches deep. On June 24 all the young were
found dead.

Three of the specimens trend a little in coloration from the darker
backs of pusilla toward the lighter pileolata but all match pusilla
in color of underparts. It appears that the specimens are variants
among pusilla rather than intergradations with pileolata. On this
basis they are extensions westward of an eastern population. Porsild
(1943) had reported pileolata from the Mackenzie Delta, and Rand
(1946) attributed Ross’ (1862), Lapierre House record to pileolata.
In southern Yukon pileolata has been commonly recorded, Our
specimens probably indicate that the eastern form extends westward
through the Porcupine Valley meeting the Alaskan range of pileo-
lata, which extends along the Yukon River. This extension of
pusilla westward is north of the range attributed to pileolata in
southern Yukon and northwestern British Columbia.

Family ICTERIDAE: Blackbirds

Euphagus carolinus carolinus (Miiller)

Males Females
Weight Fat Testes Weight Fat 8
Date 5 (mm.) 3 sh

May 10 52.1 MF 1.5 mm.
May 22 63.5 LF 10x15, 7x10
June 6 66. 2 LF 7x10, 9x8
June 26 60.9 MF 9x13.5, 12x12.5 52.9 MF 2mm.
June 7 5 half incu-

bated eggs

A female rusty blackbird was collected near the village on May
10 by Joe Kay. Until May 17 blackbirds were frequently seen as
they were apparently settling in the willow alder brush near by.
On May 19 a male was chasing a female and on May 20 some pairs

230 U. S. NATIONAL MUSEUM BULLETIN 217

appeared to be settled. The numbers increased and pairs became
more numerous, until by about May 24 the population of blackbirds
became about one fourth as numerous as the robins. Keeping near
the tops of the trees, they were as conspicuous as robins, but much
less active. On June 5 a blackbird was seen flying in hot pursuit of
a large owl.

On May 22 the testes of specimens taken had reached the largest
size measured, and on June 5 a nest with 5 half-incubated eggs was
found, the first evidently laid about May 25.

Lazarus Charlie reported that the first blackbird arrived on May
2 at Johnson Creek, a tributary to the southern part of the Porcupine
River. The air distance from Old Crow is only 80 miles south by
east, but it is three times as far along the river.

The winter range of this bird is so widespread that it provides no
suggestion as to the migratory course taken to Old Crow.

The Indians call this bird Chilly cho.

Family FRINGILLIDAE: Grosbeaks, Finches, Sparrows, Buntings

Three species of Fringillidae are found in winter at Old Crow.
Only pine grosbeaks (Pénicola) are differentiated as a race confined
to Alaska, Yukon, and Mackenzie. White-winged crossbills (Leu-
coptera) and hoary redpolls (Acanthis hornemanni ewilipes) have a
wide North American range. The arctic Yukon population of hoary
redpolls is significant, however, because they have not been reported
in southern Yukon, and so this northern population forms the nar-
row connection between the numerous hoary redpolls of Alaska and
Mackenzie. Snow buntings have not been reported wintering in
Yukon, which thus separates the populations wintering in Alaska
and Mackenzie. In their early spring migration the arrivals at
Old Crow seem to come from the Alaskan wintering population.
One other migratory species (Calcarius) forms a race which nests
principally from Alaska to Mackenzie. Northwestern America is
thus the special range of a race of resident pine grosbeaks, a race of
migratory Alaska longspurs, and a population of migratory snow
buntings.

Among the other migratory species of Fringillidae six races nest
in arctic America only from the Mackenzie westward. Only the
fox sparrow (Passerella) can be called an eastern American race,
the remainder migrating through western States and Provinces.

It is probable that snow buntings approach Old Crow from the
west and fox sparrows from the east or southeast. The arctic routes
of the others cannot now be indicated.

OLD CROW

231

This information is summarized in the following tabulation :

Species
Acanthis hornemanni
exilipes
Acanthis flammea flam-
mea

Passerculus sandwichen-
sis anthinus

Junco hyemalis hyemalis

Spizella arborea ochracea

Zonotrichia leucophrys
gambelii
Passerella iliaca zaboria

Melospiza lincolnii
lincolnii

Calcarius lapponicus

_ alascensis

Plectrophenax nivalis
nivalis

Arctic nesting range

Nearest wintering place

Lapland east to Old Crow

Mackenzie
Scandinavia
east to
Keewatin
Alaska to Mac-
kenzie

Alaska to Mac-
kenzie

Alaska to Mac-
kenzie

Alaska to Mac-
kenzie

Alaska to Mac-
kenzie

Alaska to Mac-
kenzie

Alaska to Mac-
kenzie

Alaska to
Greenland

Northern
Alaska

Southwestern
British
Columbia

Southwestern
British Co-
lumbia

Southern
British Co-
lumbia

California

Eastern Kansas

Northern
California to
northern
Georgia

Montana

Alaska

Migratory course

Western Moun-
tains

Western Moun-
tains

Western Moun-
tains

Western Moun-
tains

Mississippi,
Mackenzie

Western Moun-
tains
Porcupine River

The species present at Old Crow are numbered below in order of
their nesting abundance:

Species Order of nesting Species Order of nesting

abundance abundance
Pinicola enucleator alascensis iff Spizella arborea ochracea 3
Acanthis hornemanni exilipes 4 Zonotrichia leucophrys gambelii 1
Acanthis flammea flammea 5 Passerella iliaca zaboria 2
Loxia leucoptera leucoptera 8 Melospiza lincolnii lincolnii —

Passerculus sandwichensis

anthinus
Junco hyemalis hyemalis

a ©

Pinicola enucleator alascensis Ridgway

Weight
Date (g.)
May 6
May 13 59.3
May 19 60.1
May 30 59.2
June 4 53.5
June 20 55.6
(Aver.) 57.5

469496—60——16

Calecarius lapponicus alascensis -
Piectrophenax nivalis nivalis -

Males Females
Fat Testes Weight Fat Eggs
(mm.) (g.) (mm.)
LF 2.OX3, 2X2 VLE 1
VLF 7x10, 7x8 61.9 LF 2
VLF §.5x7.5, 5.5x6.5 59.0 VLF 2
VLF 11x13.5, 12x12 64. 6 VLF 2
NF 9x11, 9x10
LF 8x9, 8x9 54.5 LF 2
60. 0

232 U. S. NATIONAL MUSEUM BULLETIN 217

Early in April pine grosbeaks were commonly seen feeding in wil-
lows. The small groups included some pairs, for the bright rosy males
were conspicuous. Some males with very little pink could not be
distinguished in coloration from females. Both sexes were singing,
as some female specimens were collected while singing.

Of the specimens collected, those taken, at the end of April had
testes of the largest size and their plumage was the deepest red.
The males were then exhibiting themselves most conspicuously, so
that nesting was probably in progress. Even in April, the feathers
of specimens were loose and in June could scarcely be kept on the
skin. Apparently moulting followed soon after the early nesting
date.

Pine grosbeaks and willow-ptarmigan, both of which feed on wil-
lows in the Arctic spring, are commonly lean birds. The grosbeaks
were the only species of Pringillidae at Old Crow which commonly
had very little fat. At the same season the migrating snow buntings,
which feed upon seeds of annual plants, and the resident cross-bills,
which feed upon spruce cones, presumably eating their seeds, were
fat. The resident waxwings also were fat.

The Indian name for the pine grosbeak is T'eevay.

Acanthis hornemanni exilipes (Coues)

Males Females
Weight Fat Testes Weight Fat Eggs
Date (g.) (mim.) (9.) (mm.)
Apr. 19 14.6 F 3x4, 3x3
May 1 13.3 VLE 5x6, 5x5
May 16 13.3 ME 5x6.5 12.5 MF
May 18 14.3 LF 5x6.5, 4x6.5
13.3 NE 5x6.4, 4x4.5
May 22 13.4 LF 6.5x7, 4.5x6
May 24 13.4 VLE 6x8, 6x7 13.5 F 0. 75
June 6 13.9 MF 5.5x6, 6x6
June 7 12.6 LF 4.5x5, 5x5.5
June 12 13.1 LF 6x6, 5x5.5 12.0 NF 2
June 12 14.0 NEF 5x6, 5.5x5.5
(aver.) 13.6 12.7

During April small groups of redpolls were occasionally seen. They
moved so restlessly that few of them could be identified, but those
which could be seen were thought to be hoary redpolls. The Indians
consider redpolls winter birds, and no increase in numbers sug-
gesting migration was noticed until after May 20, when the first
common redpolls were remarked. Hoary redpolls as well then be-
came more conspicuous, but it appeared that the increase in num-
bers was temporary, as if a small migratory flight passed through.

No flocks of more than ten redpolls were seen, and the numbers
were few in comparison with the great numbers visibly migrating in
spring and nesting in summer on the tundra in the central part of

OLD CROW 233

the Brooks Range. There a few hoary redpolls are also winter
residents.

Hoary redpolls have been reported in southern Yukon only from
Forty Mile (Rand 1946). Since that country has been traveled by
a number of observant naturalists, the lack of records indicates that
the large hoary redpoll populations of Alaska and Mackenzie are
separated in the south but connected in the north.

The Indians call this bird Z'aloo.

Acanthis flammea flammea (Linnaeus)

Males Females
Weight Fat Testes Weight Fat Eggs
Date (g.) (mm.) (g.)

May 21 12.3 VLF 5x7, 4.5x5
May 24 12.8 NF 5x8.5, 5x6
May 31 13.0 MF 5x6, 5.5x5.5

12.2 LF 5.5x6, 5x6 1 egg, brood patch,

2 empty follicles

June 5 13.6 MF 6x7, 5x6 15.7 ¥
June 6 12.4 NF
June 9 11,2 LF 1.75 mm.
June 12 13.0 LF 5x6, 5.5x6
June 17 12.8 NF 5.5x7, 5.5x6 14.9 F 1.75 mm.
June 26 14.5 LF 7x8, 7X7

(aver.) 13s

The earliest common redpoll observed was collected on May 21.
Thereafter specimens were sought more eagerly than were hoary
redpolls, but in spite of our bias it seemed that after their migratory
arrival common redpolls were more numerous than hoary redpolls.
On June 1 a redpoll’s nest with two eggs was found and on June 17
a nest was found with five slightly incubated eggs, but neither parent
was identified. A female specimen of common redpoll had laid two
eggs on June 5.

The common redpolls at Old Crow were less fat than the hoary
redpolls and weighed slightly less. The average weight of common
redpolls at Old Crow, 18.1 grams, was not significantly different from
the average 12.9 grams at Anaktuvuk. Common redpolls are not
known to winter in either locality, but they are found near Nulato
(Dall, 1869) and at Forty Mile (Rand, 1946). This sight record ap-
pears to indicate winter residence, and one of Grinnell’s (1909) speci-
mens from Forty Mile was dated November 5. The wintering common
redpolls of southern Mackenzie or Alaska might be the source of the
nesting common redpolls. We have no indication of which direction
they travel to Old Crow, but it seems likely that it is either eastward
or westward, rather than northward over the Ogilvie Mountains.

Common redpolls are recorded from various parts of Yukon (Rand,
1946) and their distribution seems to be continuous in summer across
Yukon and is probably continuous across southern Yukon in winter.

234 U. S. NATIONAL MUSEUM BULLETIN 217

In the southern continuity of their range from Alaska to Mackenzie
the common and hoary redpolls seem to differ.

Loxia leucoptera leucoptera Gmelin

2 males Apr. 8 and 18 weight, 22.5, 28.0 g. _fat (1); medium testes 4x4, 8x8 mm.
1 fematie Apr. 18 weight, 24.6 g. ie fat —_—

On April 8 a loosely associated flock of twelve crossbills included
some which remained associated as pairs while they were feeding
among the black spruce near its upper limit. The testes of the male
specimen were then 4mm. in length. On April 18 a pair was collected
in the black spruce at about 1,200 feet, the male having well developed
testes, but the eggs in the female were little enlarged. Occasional
small flocks were seen in April and pairs were found thereafter,
always in the spruce. The Indians speak of crossbills, which they
name T%zinkee, as birds resident throughout the year.

Passerculus sandwichensis anthinus Bonaparte

1 male May 21 weight 18.4 g. very little fat testes 5x9, 7x8 mm.

Accustomed to the commonness of Savannah sparrows in much of
northern Alaska, we looked carefully for them at Old Crow, but
only about four were noticed, too few to allow us to seek information
about them from the Indians there. The areas searched did not have
much of the grassy marshland where these sparrows are so familiar
in northern Alaska. Porsild (1943) remarked that in some years
they were common, in others rare in the Mackenzie Delta.

Junco hyemalis hyemalis (Linnaeus)

Males Females
Weight Fat Testes Weight Fat Eggs

Date (g.) (mm.) (Cp) (mm.)
May 15 19.4 VF 1x125, 0.5x0.75
May 16 18.0 F 0. 5-1
June 10 20.0 LF 12
June 17 19. 4 NF 2
June 28 20.3 LF 6.5x8.5, —— (?)

June 1

1 Brood patch.
2 5 fresh eggs.

Two juncos were seen on May 15 and on the next day a few were
heard singing. Thereafter they were often heard singing and were
seen occasionally. Most often they were in the rather open mixtures
of spruce and deciduous trees or brush.

On June 1 a nest with five fresh eggs was found by Sidney Peyton.
The first of these must have been laid about May 25, ten days after
the earliest junco was seen. A female specimen on June 10 had a
brood patch.

The Indian name of this bird is T'chikikeekeejay.

OLD CROW 235

Spizella arborea ochracea Brewster

8 males May 16-June 20 weight 16.7-20.2, medium fat (2), little ——
average 17.8 g. fat (2), very little
fat (1), no fat (1)
6 females May 20-June 29 weight 14.1-19.4, medium fat (2), little —_—_
average 16.8 g. fat (3), very little

fat (1)

In this small series the male and female tree sparrows at Old Crow
were a little lighter in weight than the respective averages of 18.4
and 17.7 grams found during several years at Anaktuvuk.

The first male was collected on May 16. On the next day males
were singing extensively in the willow-alder brush near the village.
By May 18 many territories seemed to be settled but the numbers of
birds seen increased until May 24. Tree sparrows were then nearly
as numerous as Gambel’s sparrows but not quite so conspicuous as
fox sparrows. In fact their arrival, settlement, and general behavior
was quieter than it was for most of the migratory species. By June
tree sparrows were widely distributed in grassy places and deciduous
brush, but not in the continuous spruce forest. They were found at
elevations 500 feet above the upper limit of spruce growth.

On June 5 a nest with five eggs was found by Francis Williamson
at Driftwood Creek, where tree sparrows were uncommon in the
extensive spruce forests. Another nest with five half-incubated eggs
was found on June 29, one of the latest discovered. It might have
been a second attempt at nesting but was too early to follow a suc-
cessfully raised brood.

The Indians call this bird T’chinkee.

Zonotrichia leucophrys gambelii (Nuttall)

18 males May 15-June 22 weight 25.3 g., — —
average; coeff.
var. 6.5%.
10 females May 21-June 22 Weight 23.4 g. — —
June 1 — -_-— 5 fresh eggs
June 2 — — 5 fresh eggs
June 14 -— _— 6 half-incubated

eggs

In the grassy areas and willow-alder brush around Old Crow
Village white-crowned sparrows, which the Indians call WVatsik, were
about equally as numerous as tree and fox sparrows. Wherever spruce
was replaced by grass or deciduous brush they might be found. They
were about the commonest species seen in the whole area. They were
above tree line, and a nest with five eggs was found at about 3,000 feet
on the steep, barren, dry, rocky slope of the second Old Crow Moun-
tain. Because of the extent of their varied habitat white-crowned
sparrows are considered the most numerous among the species of
Fringillidae nesting about Old Crow.

236 U. S. NATIONAL MUSEUM BULLETIN 217

Gambel’s white-crowned sparrows were recorded as first seen at
Old Crow on May 14. By May 17 many were singing. On May 19
the numbers had further increased until they were the most commonly
seen bird in the vicinity of the village. At this date they appeared
to have settled most territorial limits, in which the males were sing-
ing, but there were still some territorial adjustments in progress
through invasion and judgment by combat.

In average weight of males and females white-crowned sparrows
at Old Crow did not differ significantly from males and females at
Anaktuvuk, or from males at Mountain Village, Alaska (Oakeson,
1953). The average weight of six males in the first 5 days after
arrival was greater than the average of nine males weighed during
the next 30 days, and the degree of fatness recorded indicated that
some fat which was present at the end of migration had been con-
sumed at the time when the young were fledged (see fig. 19, in the
Appendix).

During the time of laying eggs three females weighed 31, 33, and
31 grams, about 8 grams, or over 30 percent above the average weight
of nongravid females. The heaviest, which had laid five eggs, as
shown by collapsed ovarian follicles, and contained a formed egg, was
then nearly 10 grams heavier than the usual nonbreeding female. An
individual egg might weigh three or four grams, but this is not enough
to account for the excess in weight of the gravid females. One of these
females was marked as having little fat and two as medium fat. The
degree of fatness was not greater than in nongravid females, so that
not all of the tissues responsible for the increment in weight can be
indicated.

The efficiency of a hen in converting food to egg may be 77 percent
(Brody, 1945), so that in forming its own weight of eggs, as is con-
tained in a clutch of six, a sparrow must consume more than its own
equivalent of food for egg production during the six days of laying.
The rate of energy consumption in forming eggs probably requires
something like twice the usual consumption of food by the female
bird during laying. Evidently the resources in the sparrow’s meta-
bolic capability and its food supply in arctic spring are sufficient
to permit this sudden extra metabolic effort required during egg
laying. I have earlier remarked that while male birds of several
species appear to decline in weight and fatness after arrival in arctic
Alaska and during breeding, females do not so often show a decline
until after the young birds are being fed. The good condition of the
laying birds is evidence that these events impose no stress.

The reproductive schedule of Gambel’s sparrows at Old Crow can
be indicated from the ruptured ovarian follicles found in two female
specimens, each of which also contained a formed egg, from three

OLD CROW 2S.

sets of eggs collected, and from three broods of birds near fledging
which were banded (see fig. 19). The degree of incubation or progress
toward fledging was estimated by considering 9 days the duration
of nestling growth, following Barbara Oakeson’s (1954) studies on
Gambel’s sparrows at Mountain Village, Alaska, and our observa-
tions at Anatuvuk (L. Irving and Krog, 1956), and 12 days the
duration of incubation. In six nests the first egg was calculated
to have been laid between May 25 and May 28. The first egg was
laid 11 days after the first male was seen and the interval between
the arrival of the first male and the average date for the first egg
in six clutches was 13 days. The schedule appears reliable because
there is no error in calculating the date of the first egg from the dis-
covery of ruptured ovarian follicles and a well formed egg. As addi-
tional evidence for early laying, an egg 7 mm. long was found in the
first female collected on May 21. This egg could be expected to
be laid within a few days.

At Anaktuvuk, two eggs were recorded by Simon Paneak as found
on June 1, 1954, and two. females, taken on May 25 and May 27,
each contained an egg 8 mm. long. The first migrants are usually
seen there about May 15. This suggested an interval of about 15
days between the first arrival and the first egg, three days shorter
than was estimated from occasional earlier records at Anaktuvuk.

The careful records of the schedules of individual Gambel’s spar-
rows at Mountain Village presented by Barbara Oakeson (1954)
showed that in 1950 the first male arrival on May 10 was followed
by the first egg on May 25, an interval of 15 days. At Old Crow the
completion of all of the preparations for the first laying observed oc-
curred in the 11 days after the first male bird was seen. There the
country is evidently well suited for the large breeding population,
while at Mountain Village, Barbara Oakeson found Gambel’s spar-
rows not numerous, and at Anaktuvuk they were far less common than
at Old Crow. Mrs. Oakeson found that the duration of the intervals
between arrival and laying at Mountain Village was only about half
as long as for Z. 1. pugetensis at Friday Harbor. There is undoubtedly
acceleration of the breeding cycle at Old Crow and shortening of
its duration by synchronization of the schedule throughout the popu-
lation. The shortening of the schedule is of course a necessity in a
high latitude where the season suitable for breeding is so short. From
information which Mrs. Oakeson kindly provided on the breeding of
Gambel’s sparrows at College, Alaska, in 1957, the interval between
the average first arrival of males and first egg was 13 days and only
4 days after the average date of arrival of the females.

Her studies will indicate whether or not there is acceleration in the
physiological processes preparatory to breeding. We do not have that

238 U. 8. NATIONAL MUSEUM BULLETIN 217

evidence. I do not suspect that incubation can be environmentally
modified and I would be surprised if nestling growth could be signifi-
cantly modified by environment (L. Irving and Krog, 1956). The
conspicuous influence upon the breeding cycle appears in the synchro-
nization of events so that their succession is swift in the entire popu-
lation. The adaptation of breeding to the arctic environment thus
appears as a social phenomenon rather than as a modification of the
individual’s physiological processes, many of which are so deeply
rooted in the origins of the species that their adaptive modifiability
to local conditions is hardly to be suspected. (For a further discus-
sion of this subject see also chapter 7.)

In six of the nests where complete clutches were apparent the aver-
age of the numbers of eggs was 5.5, I think it proper to regard the
set of three nestlings as diminished by casualty. At Anaktuvuk sey-
eral complete clutches contained either five or six eggs. Mrs. Oakeson
kindly permits reference to her comparison of clutch size among white-
crowned sparrows in various localities:

Average
Race Location Clutches 8ize
nuttalli Central California 147 3.25
pugetensis Seattle, Friday Harbor, Victoria 44 4.09
gambelii Mountain Village 6 4.7
gambelii College 13 4.9
gambelii Old Crow 6 5.5

Our observations do not have the significance of her more careful
studies, but they show that the number of eggs laid near the arctic
limit of the range of white-crowned sparrows is largest. We would
not be inclined to relate this northward increase to latitude, for the
increase of eggs is not proportional to the differences in latitude. We
have earlier remarked that in many species, like plovers and sand-
pipers, the size of clutch is invariable, and the influence of environ-
ment upon size of clutch is not general. The large number of eggs
laid is, however, certain evidence of the adequacy of nutrition near
Old Crow in a dense population of white-crowned sparrows. The
rarity of casualties observed in nests is another indication of the suc-
cessful adaptation of breeding habits to the locality.

Passerella iliaca zaboria Oberholser

25 males May 14-June 20 Weight, average —— s=s=
(25), 35.9 g.,
coeff. of var. 5.8%

7 females May 16-June 20 Weight, average —_— —_
(3), 34.3 g.

2 young males June 20 Weight, average = =
(2), 26.8 g.

1 young female June 20 Weight 29.0 g. — ==

4 half-incubated Junel — —_—_ =

eggs

4 heavily incu- June 11 — aes ees
bated eggs

OLD CROW | 239

A few fox sparrows, known by the Indians as 7'cheekeekak, were
seen in the willow-alder brush near the village on May 14 and 15
and on May 16 numbers were singing and vigorously repelling inva-
sions of the territory they had assumed. On May 17 the numbers had
increased, there was much local flying as invasions were made and
repelled, and on May 18 it seemed that the area was saturated. The
quarreling persisted, and evidently extra males were seeking terri-
tories; an hour after one had been shot while singing, another was
singing on the same willow. By May 20 there was little disorder and
much placid singing.

On May 24, fox, Gambel’s, and tree sparrows were very common
in the willow-and-alder brush about the village and along the river.
Fox sparrows were found wherever the spruce was thin in swamps
or on slopes, usually in willow brush. They were seen on the moun-
tain as high as the brush was still continuous at about 1,700 feet.

Males were not far ahead of females in arriving, for the first male
was collected only two days before the earliest female specimen was
taken. Two sets of eggs with four eggs each and four sets of nestlings
were examined and two young birds were captured when just able to
fly and recently fledged. The degree of their development was esti-
mated on the basis of the scale observed by Oakeson (1954) in white-
crowned sparrows at Mountain Village, viz., eggs laid one day apart,
but with incubation twelve days and nestling stage ten days as we
found among tree sparrows at Anaktuvuk (Irving and Krog, 1956).
Using these figures for calculation implies that the most advanced
nesting birds developed from clutches for which the first egg had been
laid May 20 (see fig. 20, in the Appendix).

Two sets of eggs were started 6 days after the first male arrived
and the average interval between first arrival and first laying was
8 days in seven of the eight nests. For the first male observed on
May 14 to have taken a territory and a mate which could prepare
a nest and egg within 6 days is rapid progress in those complicated
social and marital processes. Other evidence points to the earliness
of fox sparrows’ laying at Old Crow. In four female specimens
taken eggs were measured as 3 mm. on May 16, 9.5 mm. on May 20,
7 mm. on May 22, and 3 mm. on May 22. A brood patch was remarked
on a female on June 1.

The arrival of fox sparrows is made conspicuous by their songs,
the flashing of their bright rusty color in flight through the gray
willows, and the vigorous demonstrations of the males in taking
nesting places. Four observers in the scientific party were daily
searching for birds in the habitat about the village where fox sparrows
were very numerous, and earlier reports of the observant Indians
coincided with our first sight of common species within a day. The

240 U. S. NATIONAL MUSEUM BULLETIN 217

6-day interval between first observed arrival of fox sparrows at Old
Crow and their first egg was even shorter than for Gambel’s sparrows.

It was earlier remarked that the community of fox sparrows was
in good order by May 20, for territorial transgressions were not then
frequent, although some nests were not a hundred feet apart. Within
about 8 days the entire society of fox sparrows had become changed
from an organization of individuals in which the influence of sex
was scarcely visible to a new social order in which sex impulses were
dominant in separating the society into pairs. The social reorgani-
zation was mainly effected by the demonstrative activities of male
birds.

The swift settlement of large numbers of fox sparrow in their re-
spective territories proceeded so rapidly and, in spite of the brief
period of combat, with so little disorder that provision for the social
processes which could organize them into a community of families
must have been already inherent in the individuals when they reached.
Old Crow. The expression of social organization appeared only in
the nesting environment, but the physiological preparations had been
completed and the inclination of every bird toward the correct se-
quence of social behavior must have been already perfected. Some
common bond, as well as temporal and spatial association, must have
joined the birds during their migration toward Old Crow, producing
a synchrony in their physiological and social states. Somewhere in
each bird’s “memory” there is brought about the synchronous prepa-
ration of breeding activity which eventually segregates and orders a
predetermined segment of the whole migrating population to proceed
to settlement on its nesting ground at Old Crow. The social “memory”
cannot be described by analytical reference to physiological processes
and mechanisms alone, for the organization of societies also involves
the environmental terms of time and space that are so evident in the
annual migrations of birds.

In specimens taken of arriving male fox sparrows the testes were
not found at their largest size until about May 27, but some must have
been ready for breeding before May 20. The early males were mostly
medium fat, and examples during early nesting had little fat, but
picked up fat again while nestling birds were being cared for. Their
generally good condition attests the adequacy of the food supply and
suggests that the revolutionary physiological and social transitions
while nesting were effected without nutritional strain.

As was remarked about female white-crowned sparrows, several
female fox sparrows preparing to lay were about 8 grams, or 20 per-
cent, heavier than the average of nongravid females. This increase is
near four times the probability of normal variation in weight of
males, and is about twice the weight of an egg. As those individuals
were fat, its accumulation and the ovarian enlargement are plausible

OLD CROW 241

explanations of the oversized weights. They show the expansive
capacity of avian metabolism for egg laying, the adequacy of the
food supply, and imply that the individuals were secure for an ex-
panded feeding program in a social system organized to preserve
individuals from stress during the period of reproduction. This view
goes with the appearance of leisurely behavior among the birds, at
least until the nestlings must be often fed.

The fact that no casualties were observed among the eggs or
nestling birds suggests that reproductive efficiency was high in this
rather dense population of fox sparrows, which went through its re-
productive cycle with such amazing speed.

Since some of the patterns of individual breeding behavior were
compressed within such short intervals, it seems quite likely that the
contributory development of some physiological changes was acceler-
ated. The duration of incubation and of nestling growth are not to
be suspected of being much influenced by the environment for they are
ancient characters of species and even higher systematic relations
(Irving and Krog, 1956). The conspicuous modification in arctic
breeding occurs in the synchrony of the stages in the entire population,
as Barbara Oakeson has so clearly shown in Alaskan Gambel’s spar-
rows (Oakeson, 1954). That this synchrony greatly reduces strife
and promotes social order is undoubtedly a factor of advantage for
the economy of arctic nesting, and the resultant short period devoted
to breeding in the Arctic must further reduce the social and physical
stress which birds encounter during the longer breeding seasons of
warmer climates. Because of its apparent advantages, the synchrony
of the progress of arctic birds through their arctic breeding cycle
can be regarded as an adaptation. This viewpoint is discussed in
detail in chapter 7.

Fox sparrows of the race zaboria winter mainly east of the Missis-
sippi. Their northern nesting range reaches the Arctic only in Mac-
kenzie, Yukon, and Alaska. They are the only species of the Fringil-
lidae nesting there which can be called eastern in reference to their
wintering range, and yet they, with gray-cheeked thrushes and some
wood warblers, also eastern wintering races, migrate from near ex-
treme southeastern to extreme northwestern parts of North America
to nest.

Melospiza lincolnii lincelnii (Audubon)

1 male May 31 weight 19.1 g. little fat testes 5x10, 8x10 mm.

Francis Williamson heard the male Lincoln’s sparrow singing in
the rank grassy clearing by the Roman Catholic Mission and collected
the only record of its occurrence at Old Crow. The bird acted as if
it had assumed a territory. It has been identified at Forty Mile
(Grinnell, 1909), found sparsely along the Yukon River west of the

242 U. S. NATIONAL MUSEUM BULLETIN 217

international boundary to Circle (Bishop, 1900) and is not uncom-
mon in southern Yukon (Rand, 1946).

Calcarius lapponicus alascensis Ridgway

é Males Females
Weight Fat Testes Weight Fat Eggs
Date (g.) (mm.) (g.) (mm.
May 4 24.6 ME
May 8 23. 0 F 3.5x4.5, 3.5x4
May 9 28,4 F 4x5, 3x4
May 13 27.1 LE 5x7.5, 4.5X7
May 14 27.7 MF 5x6.5, 6x6
May 17 een 24.3 F I
(aver.) 26. 2

In contrast with the great flocks of longspurs which migrate
through the Alaskan Brooks Range, only a few small groups were
seen at Old Crow. The largest count recorded was four. From May
4 to May 12 only males were seen, and from then until May 24 an oc-
casional female was seen with a few males. Those seen at Old Crow
do not represent the great migration of longspurs as it is seen at
Anaktuvuk. They were lighter in average weight and with less fat.
The earliest arrivals (May 4) were apparently expected at Old Crow
by Joe Kay, who promptly obtained specimens, named them Shinjee,
in the Indian language, and described their singing. Their arrival
there is apparently part of a regular schedule, but the vicinity of
Old Crow does not provide the grasslands and tundra where longspurs
congregate in large numbers.

Plectrophenax nivalis nivalis (Linnaeus)

16 males Apr. 5-May 21 weight 35.5-44.9, very fat (6), fat testes 1.5-4 mm.
average (7), 40.3 (10), little fat (2)
g., coeff. of var.
8.1%

At Old Crow in 1957 the residents reported that snow buntings
had first reached there about the middle of March. Their name for
the bird is Kukuzu. After we arrived on April 2, a flock of 20 or 30
was usually to be seen along the river bank in front of the village
feeding among the accumulated debris of the winter. This rubbish
contained little food which could please a small seed-eating bird, and
the interest of these earliest migrants to the Arctic was more likely in
weed seeds than in garbage.

Some flocks were followed back and forth for a mile along the river
bank, a few hundred yards at a move. The flocks often showed their
attraction to the village by returning after starting to fly away. It
was suspected that some flocks remained for a day or longer, but it
could not be established by observation and pursuit that a flock re-
mained for longer than several hours. Most of the records are of
course from the village, but occasional observations along the river
and reports of Indians show that while the flocks probably congre-

OLD CROW 243

gated at villages, they may stop at any clearing. After April 30, none
were reported. A single bird was noted on May 15, two on May 17,
five on May 19, and the final recording of a single bird was on May 21.

On Old Crow Montain four buntings were seen on April 11 and
two were collected by Joe Kay on April 17, but after the snow melted,
they were not seen, for the Old Crow Mountains are not high enough
to provide the sort of weather in which they nest along the arctic
coast and in the high mountains near Chandler Lake, Alaska, where
Simon Paneak reported them in 1955.

The fatness of these snow buntings at Old Crow was an impressive
indication of the adequacy of their food supply. In 1954 Susie Paneak
at Anaktuvuk recorded weights of 13 male snow buntings which av-
eraged 34.9 grams, and which were not as fat as at Old Crow, rating as
2 VF,1F,5 LF, and 1 VLF. Seven snow buntings in September
1947 at Point Barrow were, however, like the Old Crow birds in weight,
so there is no evidence in weight records to distinguish buntings at
Old Crow from those in central arctic Alaska. The variation in
weight among individual birds of each series of snow buntings is
larger than is found in most series of arctic migratory birds at Old
Crow, for the coefficient of variation was 8.1%. The mean weight of
the Old Crow and Anaktuvuk series differed according to the degree
of fatness reported in each series, but it appears that in this species
individual weight as a function of stature is a variable factor as well
as fatness. As observed visually, the fat of snow buntings was light
corn yellow and soft at room temperature. The consistent appearance
of the fat suggests its origin in a constant dietary source and implies a
single major component in the food.

Only male birds were found among the 16 specimens sexed in spring
at Old Crow. At Anaktuvuk, from 1949 to 1953 11 males and 3 fe-
males were collected. In 1954 there were 3 females in a series of 16
weighed at Anaktuvuk. No females have been collected at Anak-
tuvuk during April and there are accordingly indications that at
Anaktuvuk and Old Crow males are segregated in some flocks.

The testes of the specimens at Old Crow were but little enlarged
and the series did not show growth. When the latest flocks were
leaving Old Crow, at the end of April the birds had at least two weeks,
but only one hundred and forty miles before the earliest arrivals would
be due at their nesting grounds on the arctic coast.

In central arctic Alaska the stages in progress of migration of
snow buntings northward are indicated by first arrival dates at Bettles
about March 15, at Anaktuvuk April 1, and at Barrow April 15. It
appears that this schedule results from a movement the front of which
traverses the last 300 miles northward to their arctic Alaskan nest-
ing grounds in a month. Indians of Old Crow Village say that snow
buntings first reach Crow Flats later than they reach the village, and

244 U. S. NATIONAL MUSEUM BULLETIN 2.17

that they remain numerous on the Flats until later in spring. Ap-
parently the northern part of their migration is as leisurely and as
prolonged in Yukon asin Alaska. (A general discussion of migration
is to be found in chapter 6.)

Fourteen of our Old Crow specimens still possessed much brown
color on the head and rump, a color usually lost at breeding time,
leaving the white in sharp contrast with the black on wings and back.
It was apparent that the edges on the brown feather tips of our
Specimens were much weakened and about ready to be shed. The
change is not one of wear, as it is commonly called, because it occurs
at a predetermined position and leaves the edges of the feathers in
a sharp new outline quite in contrast with the irregularly worn con-
dition of tips of wing and tail feathers often apparent just before
the moult. The brown edging of contour feathers is retained for
about nine months and until the buntings are near their breeding
grounds.

We could not detect any distinction in size or plumage which would
distinguish between snow buntings from Alaska, Yukon, and
Mackenzie. In describing the snow buntings of Siberia as Ps; n.
pallidior, Salomonseh (1947) remarked that a few moulted speci-
mens from western Alaska showed the lighter backs by which he
distinguished pailidior from nivalis of “Arctic America, perhaps ex-
cept Alaska.” We did not have enough specimens in fresh plumage
to test Salomonsen’s suggestion of a distinct Alaskan color character.

At the U. S. National Museum Bernard Feinstein pointed out that
specimens from Alaska had stouter claws than a series of specimens
taken during migration at Fort Simpson on Mackenzie River. On
the basis of the average difference in stoutness of claws the Old Crow
birds go with those from Alaska rather than with those from
Mackenzie.

In the discussion of snow buntings at Anaktuvuk it was mentioned
that many occur in winter scattered through central and southern
Alaska. They are separated from those wintering in Mackenzie by
a mountainous region in Yukon which seems unsuited for wintering
buntings and from which none have been reported in winter. That
the geographical separation in winter may also isolate the two popu-
lations on their breeding grounds is further suggested by the fact
that it has been possible to sort Alaskan from Mackenzie buntings
on the basis of stoutness of claws. The resemblance in claw stout-
ness between Old Crow and Alaskan buntings suggests that both
belong to the Alaskan wintering population and that migration
passes eastward along the Porcupine at least as far as Old Crow. In
summer there is apparently no geographical gap in the extension of
breeding snow buntings along the arctic coasts of Alaska, Yukon,
and Mackenzie.

SSE

Loox

if

i
wget
} t

Mh
a

PINTAIL AND SPECTACLED EIDER, Anas acuta (see pp. 37, 170) and Lampronetta
fischeri (see pp. 35, 168).

5. Status and Distribution

| Sean arctic AtasKa has been comparatively little changed by

its human occupants, for until the last few years it has been subjected
only to aboriginal methods of exploitation. In modern times, how-
ever, in those parts of Alaska where the search for gold has been carried
out, the application of water and mechanical power has caused in-
tensive unnatural erosion. Forests near the mining operations have
been destroyed by lumbering, clearing, and fire, and the invading
population has added greatly to the aboriginal use of game. Along
the arctic coast whaling, the fur trade, and reindeer grazing have
‘intensively removed natural products that earlier had been sparingly
utilized by the aboriginal inhabitants. As a result these areas have
been visibly changed.

The tundra of the arctic interior has been less affected by civili-
zation than the coast or the forest, but we can nevertheless record the
total disappearance of the musk ox and Eskimo curlew as indications

245

246

of recent instability among the animals of the Alaskan tundra. There
are no records to show how the other animal populations have been
changed, but the lack of evidence of depletion suggests that the arctic
interior of Alaska is exceptional for the retention of conditions as
they were established by the natural progress of erosion and climate.

These considerations, together with the results of my own travels
and observations, and the reports of the few naturalists who carried
their observations through the seasons until they obtained compre-
hensive views of the avifauna, allow me to believe that I can present
a comparative view of the natural distribution of birds in arctic Alaska
and Yukon.

In the description of each species at Anaktuvuk, Kobuk, and Old
Crow (chapters 2-4), I have discussed the season when the birds
are present, their migration, and their nesting. In table 4 I have
summarized the information about their seasonal stay at these locali-
ties in terms descriptive of the seasonal extent and main biological
function served by their presence. I have used the characterization
“visitor,” which for us would imply a purpose, to designate only the
appearance of certain birds without evidence that they carry out
activities related to breeding. At Anaktuvuk, for example, all the
species called visitors are known to nest within a few hours flight,
and most have been seen frequently and in good condition, so their
appearance there gives no evidence of abnormality. Inasmuch as the
designation of birds as “visitors” defines an activity which appears
normal, however, it does include the implication of biological purpose
often suggested by a regular form of animal behavior.

U. S. NATIONAL MUSEUM BULLETIN 217

Taste 4.—Siatus of birds at Anaktuvuk, Kobuk, and Old Crow

[Explanation—(R) resident, (RM) resident and migrating, (M) migrating but not remaining to nest,
(MN) migrating to nest and also migrating, (N) migrating to nest there and not migrating farther, (V) vis-
itor, (WV) winter visitor, (O) not present.]

A- z
Species nak- | Ko- | Old Species fae Ko- | Old
tu- } buk | Crow tu- | buk | Crow
vuk vuk
Gavia immer MN | MN | MN Anas platyrhynchos platy-
Gavia adamsii MN] M Oo thynchos M N | MN
Gavia arctica pacifica MN | MN | MN Anas acuta MN | MN | MN
Gavia stellata MN | MN | MN Anas carolinensis MN | MN | MN
TOTAL GAVIDAE (4) (4) (4) (3) Mareca americana MN | MN | MN
Podiceps grisegena holbdllii oO N N Spatula clypeata M |MN|MN
Podiceps auritus cornutus V N N Aythya marila nearctica MN | MN | MN
Total PODICIPEDIDAE (2) | (1) (2) (2) Aythya affinis MN| O | MN
Olor columbianus M | MN|MN Bucephala clangula americana oO oO M
Branta canadensis taverneri M | MN | MN Bucephala islandica oO oO M
Branta nigricans M M M Bucephala albeola oO N 10)
Anser albifrons frontalis M |MN/}] MN Clangula hyemalis MN | MN | MN
Chen hyperborea hyperborea M M M Histrionicus histrionicus MN | MN | MN

STATUS AND DISTRIBUTION

247

TaBLE 4.—Status of birds at Anakiuvuk, Kobuk, and Old Crow—Continued

Species

Melanitta deglandi
Melanitta perspicillata
Oidemia nigra americana
Mergus serrator serrator
TOTAL ANATIDAE (21)
Accipiter gentilis atricapillus
Accipiter striatus
Buteo lagopus
Aquila chrysaetos canadensis
Haliaeetus leucocephalus alas-
canus
Circus cyaneus hudsonius
TOTAL ACCIPITRIDAE (6)
Pandion haliaetus carolinensis
TOTAL PANDIONIDAE (1)
Falco rusticolus obsoletus
Falco peregrinus anatum
Falco columbarius bendirei
Falco sparverius sparverius
TOTAL FALCONIDAE (4)
Canachites canadensis osgoodi
Bonasa umbellus
Lagopus lagopus alascensis
Lagopus mutus nelsoni
TOTAL TETRAONIDAE (4)
Grus canadensis canadensis
TOTAL GRUIDAE (1)
Charadrius semipalmatus
Charadrius vociferus
Pluvialis dominica dominica
Squatarola squatarola
Arenaria interpres interpres
Aphriza virgata
TOTAL CHARADRIIDAE (6)
Capella gallinago delicata
Numenius phaeopus hudson-
icus
Bartramia longicauda
Actitis macularia
Tringa solitaria cinnamomea
Heteroscelus incanum
Totanus flavipes
Erolia melanotos
Erolia fuscicollis
Erolia bairdii
Erolia minutilla
Erolia alpina pacifica
Limnodromus scolopaceus
Micropalama himantopus
Ereunetes pusillus
Tryngites subruficollis
Limosa lapponica
Crocethia alba
TOTAL SCOLOPACIDAE (18)
Phalaropus fulicarius
Lobipes lobatus

Species

TOTAL PHALAROPODIDAE (2)! (2)

469496—60——17

(2)

(9)
O
MN

(1)

Stercorarius pomarinus
Stercorarius parasiticus
Stercorarius longicaudus
TOTAL STERCORARIIDAE (8)
Larus hyperboreus barrovi-
anus
Larus argentatus smithsoni-
anus
Larus canus brachyrhynchus
Larus philadelphia
Xema sabini
Sterna paradisaea
TOTAL LARIDAE (6)
Bubo virginianus lagophonus
Nyctea scandiaca
Surnia ulula caparoch
Strix nebulosa nebulosa
Asio fiammeus flammeus
Aegolius funereus richardsoni
TOTAL STRIGIDAE (6)
Megaceryle aleyon caurina
TOTAL ALCEDINIDAE (1)
Colaptes auratus borealis
Dendrocopos pubescens nel-
soni
Picoides arcticus
Picoides tridactylus fasciatus
TOTAL PICIDAE (4)
Sayornis saya yukonensis
Empidonax traillii traillii
Nuttallcrnis borealis
TOTAL TYRANNIDAE (3)
Eremophila alpestris arcticola
ToTAL ALAUDIDAE (1)
Tachycineta thalassina lepida
Tridoprocne bicolor
Riparia riparia riparia
Petrochelidon pyrrhonota hy-
popolia
Hirundo rustica erythrogaster
TOTAL H1iRUNDINIDAE (5)
Perisoreus canadensis pacificus
Corvus corax principalis
TOTAL CORVIDAE (2)
Parus atricapillus turneri
Parus cinctus lathami
Parus hudsonicus hudsonicus
TOTAL PARIDAE (8)
Cinclus mexicanus unicolor
TOTAL CINCLIDAE (1)
Turdus migratorius migrator-
ius
Txoreus naevius meruloides
Hylocichla ustulata incana
Hylocichla minima minima
Oenanthe oenanthe oenanthe
TOTAL TURDIDAE (5)

can Ko- | Old
tu- | buk | Crow
vuk
Vv Vv oO
MN|MN] V
MN | MN | MN
(3) | @) } @)
MN|MN| M
Vv N | MN
MN |MN|MN
O V |MN
V V 10)
MN | MN | MN
(5) | () | ()
Vv R R
WV |Wwv| WV
O R R
O R R
MN |MN} O
Vv R R
(4) | (6) | (6)
oO N oO
(0) | @) } @
V N N
WV| R O
oO R oO
10) R R
(2) | 4) | @
MN| O N
O O N
O oO N
qd) | () | @)
MN | MN | MN
qa) |} @) } @
O oO N
V | MN| MN
V |MN|MN
oO N | MN
Vv Vv oO
(3) | 4 | ®
R R R
R R R
(ODE A CO
Wv| R oO
O R R
oO R R
qa) | @) }] @
R R R
Mel @)e ia)
MN | MN | MN
10) N N
oO O N
MN | MN | MN
MN/|MN] N
(3) | 4) 1 @)

248 U. S. NATIONAL MUSEUM BULLETIN 217

TaBLE 4.—Sitatus of birds at Anaktuvuk, Kobuk, and Old Crow—Continued

Phylloscopus borealis kenni- Junco hyemalis hyemalis V |MN| N
cotti MN|MN} O Spizella arborea ochracea MN | MN} MN
Regulus calendula calendula V ijMN;] N Zonotrichia leucophrys gam-

TOTAL SYLVIIDAE (2) (2) | (2) | @) belii MN | MN | MN
Motacilla flava tschutschensis | MN | MN} O Zonotrichis atricapilla oO N oO
Anthus spinoletta rubescens MN | MN |} MN Passerella iliaca zaboria MN | MN | MN

TOTAL MOTACILLIDAE (2) (2) (2) (1) Melospiza lincolnii lincolnii O oO V
Bombycilla garrula pallidiceps | O R R Calcarius lapponicus alascensis | MN | MN | M

TOTAL BOMBYCILLIDAE (1) |} (0) | (1) | GQ) Calcarius pictus MN} O oO
Lanius excubitor invietus MN|MN] V Plectrophenax nivalis nivalis M M M

TOTAL LANIDAE (1) qd) |} @d)] @ TOTAL FRINGILLIDAE (16) (12)} (14)| (42)
Vermivora celata celata Oo N N SS aa
Dendroica petechia amnicola V N N TOTAL, ALL FAMILIES (139) | (106)} (122)} (107)
Dendroica coronata hooveri Vv N N
Dendroica striata (0) N N SuMMARY or STATUS
Seiurus noveboracensis nota-

bilis oO N N (R) resident 5 20 18
Wilsonia pusilla pusilla O Oo N (RM) resident and migrating 2 3 2
Wilsonia pusilla pileolata N N oO (M) migrating but not remain-

TOTAL PARULIDAE (7) (3) | (6) } ©) ing to nest 21 12 9
Euphagus carolinus earolinus N | MN|MN (MN) migrating to nest and

TOTAL ICTERIDAE (1) qa) | @)} @ also migrating 49 57 48
Pinicola enucleator alascensis | WV | R R (N) migrating to nest but not
Leucosticte tephrocotisirvingi | N | MN] O migrating farther 3 19 24
Acanthis hornemanni exilipes | RM | RM | RM (V) visitor 22 10 6
Acanthis flammea flammea MN | RM | MN (WV) winter visitor 4 1 1
Acanthis flammea holboellii 0) Vv (0) —]—_—|-——_
Loxia Jeucoptera leucoptera oO R R TOTAL 106 | 122 107
Passerculus sandwichensis an-

thinus MN|MN]| N

The Birds of Anaktuvuk Pass

Anaktuvuk Pass has been so little disturbed by human activities
that the birds still live there in an environment as stable as the natural
terrain and climate. Their comparative stability greatly simplifies
description of how their activities relate them to the populations of
birds elsewhere. Among the 11 species of birds regularly present
there in winter, 7 also remain to nest in summer, as shown in table 5.
But the nesting birds cannot be called residents simply because indi-
viduals are seen there at all seasons of the year. For example, the
number of nesting willow ptarmigan (Lagopus lagopus alascensis)
is very small in comparison with the great numbers which migrate
northward through the valley in late winter. Only occasionally are
hoary redpolls (Acanthis hornemanni exilipes) seen in winter but
great numbers migrate through in May and September. At present
there is no indication whether the nesting individuals of these two
species had been present in winter or arrived during migration. The
migrant ptarmigan and redpolls at Anaktuvuk are probably a thousand

STATUS AND DISTRIBUTION 249

times more numerous than those which remain to nest. The disparity
in numbers of rock ptarmigan (Lagopus mutus nelsoni) present in
winter and summer is not as great, but the migrants are perhaps ten
times as numerous as the nesting individuals. Accordingly, in case
a species is differently represented during the year at Anaktuvuk we
cannot conclude whether a specialized sedentary habit distinguishes
a resident from a migratory section of the populations.

TaBLE 5.— Winter birds in the valleys at Anaktuvuk Pass, north of the forests

Nesting | Migra- | Winter
tory i

itor

Falco rusticolus obsoletus x

Lagopus lagopus alascensis x x

Lagopus rupestris nelsoni x x

Nyctea scandiaca x
Perisoreus canadensis pacificus x

Corvus corax principalis x

Dendrocopos pubescens nelsoni x
Parus atricapillus turneri x
Cinclus mexicanus unicolor x

Pinicola enucleator alascensis x
Acanthis hornemanni exilipes x x

Because of their secretive nesting at a time when softening snow
impedes travel we are also uncertain as to whether the few jays
(Perisoreus canadensis pacificus) which are seen in winter remain to
nest on the tundra. It would be only a guess to suggest that jays
circulate back and forth to some extent between forest and tundra.
But although they are only occasionally seen they have always been
familiar to the Nunamiut and there is no indication that in recent
years the numbers or activities of jays on the tundra have changed.

Gyrfalcons (Falco rusticolus obsoletus) appear to occupy the coun-
try around their nesting sites throughout the year as resident birds.
Although they are in a degree isolated from the rest of the population
of gyrfalcons, they can reach the territory of adjacent pairs quickly.

At certain arctic sites where open water persists in winter, dippers
(Cinclus mexicanus unicolor) have been seen at various times through-
out the year. I believe that they are residents in these small areas
which for more than 8 months in winter are surrounded by such dis-
tances of frozen country as would seem to form barriers completely
isolating each pair of dippers from the rest of their population. Yet
their numbers seem to remain about the same and their dwelling sites
are known to have been occupied for many years. In spite of their
apparent isolation it is evident that they are coherent members of a
population of dippers.

250 U. S. NATIONAL MUSEUM BULLETIN 217

Ravens (Corvus corax principalis) are often seen throughout the
year in the vicinity of the high cliffs where they nest. They are con-
sidered to be birds of established residence. It is also known that
they often make long flights which are considered to imply that
their hunting extends over wide ranges. The frequent association
of various numbers of ravens shows that they are related by their
sociability to the ravens of a greater area.

Among those species which are seen throughout the year at Anaktu-
vuk, gyrfalcons and dippers appear to be birds of settled residence and
the ravens are probably mainly fixed residents. In the case of the
other four species which are represented throughout the year at
Anaktuvuk I have no opinion as to whether the settled residence is
a habit fixed in certain individuals. If so, it might be connected
with traits individually inherited.

Four birds not found nesting there in summer are seen in winter at
Anaktuvuk and are listed as winter visitors in table 5, although their
appearance is too frequent to be considered casual. The snowy owls
(Nyctea scandiaca) come from near the arctic coast, where they nest
and where some spend the winter. The numbers seen vary with the
years. More are usually seen in late winter, but individuals do not
remain long in one place. It is not apparent whether they are in mi-
gration southward through Anaktuvuk. Their visits, while usual,
do not show any regularity that I can detect. Downy woodpeckers
(Dendrocopus pubescens nelsont), black-capped chickadees (Parus
atricapillus turnert), and pine grosbeaks (Pinicola enucleator alascen-
sis) also do not seem to have fixed winter locations on the tundra,
where they move about among patches of willows. ‘Their winter
foraging on the tundra appears to be well established. All are
hardy birds, well able to live among low willows, where they may
find the branches bared by the winter winds of the tundra better
suited for feeding than the branches in the still forests, which are
often thickly covered with snow. In summer these three species have
not been seen on the tundra. They reverse the usual seasonal direction
of movement by coming at least a limited distance northward in
winter. It seems likely that they return southward to the forests
for nesting in areas where the others of their kind are regularly
present. :

In spite of several reports on its winter occurrence on the tundra
I list the ruby crowned kinglet (Regulus calendula) as a visitor to
mark my uncertainty rather than to describe the bird’s status. With-
out a winter specimen I do not like to say that it occurs in winter
3,000 miles north of the range currently ascribed to it.

In addition to the birds listed in table 5 as present in winter at
Anaktuvuk, one snow bunting (Plectrophenax nivalis), one goshawk

STATUS AND DISTRIBUTION 251

(Accipiter gentilis atricapillus) , and several great horned owls (Bubo
virginianus lagophonus) have been found on the tundra in winter.
There is nothing remarkable about finding these birds on the tundra
in winter except that it has been too uncommon to be called a habit.

The group of birds listed below include some of those familiar
as residents in the northern borders of the forests but not reported on
the tundra:

Bird Locality of observation

Canachites canadensis osgoodi Savioyuk River
Pedioecetes phasianellus Hunt Fork

Surnia ulula caparoch Hunt Fork

Strix nebulosa nebulosa Hunt Fork
Picoides tridactylus fasciatus Hunt Fork
Picoides arcticus Kobuk Village
Parus hudsonicus hudsonicus Savioyuk River
Bombyeilla garrula pallidiceps Savioyuk River
Loxia leucoptera leucoptera Hunt Fork

In the forest they occasionally encounter weather colder than any
that occurs on the tundra, so cold is not the barrier to their northern
dispersal. Among them the crossbills (Lowia leucoptera leucoptera)
are associated with spruce for feeding, and the three-toed wood-
peckers (Picoides) feed on tree trunks but not on branches. It is
likely that the exercise of certain essential habits requires trees and
thus these hardy, and in some cases wide-ranging, birds are held within
the border of the forests.

Among the 73 species of birds shown in table 4 to migrate to or
through Anaktuvuk, 39 have been established as nesting by specimens
or photographs of eggs, downy young birds, or eggs ready to be laid,
found in collected female specimens. The behavior of birds demon-
strates their intentions to each other through sight and sound, senses
which we can appreciate, so that behavior gives reliable circumstantial
evidence for the breeding of birds. In 20 additional species, such
circumstantial evidence for breeding was regarded as conclusive indi-
cation of nesting, giving a total of 52 migratory and 7 resident species
nesting at Anaktuvuk. Evidence for the nesting of most of this
group of birds was obtained repeatedly and conformed with informa-
tion which Eskimo people gave me from former years.

Some species are seen in different numbers each year, but the varia-
tion is not as noticeable as is the impression of regularity in the
annually repeated occupation of the Valley by each common species.
For example, the number of surf scoters observed varies greatly from
year to year and I know of only one locality in the mountains
(Chandler Lake) where the sight of young surf scoters has provided
evidence that they nest in the area. Their long establishment in the
mountains is shown by old Nunamiut accounts and the application

252 U. 8. NATIONAL MUSEUM BULLETIN 217

of an explicit name. In spite of the history of long occurrence, the
variation in numbers seen from year to year shows that surf scoters
occupy the mountains irregularly. In this irregularity the surf
scoters stand in contrast with most nesting species.

Where so many sharp-sighted people were engaged in observation,
the reported absence of birds at nesting time is significant. The
designation of birds as migrating individuals implies that they are
traveling to nesting grounds and not wandering. Of the 21 migrants
that do not remain at Anaktuvuk in summer, as listed in table 5, we
know the northern nesting areas of most. Taverner’s goose (Branta
canadensis tavernerz) isan interesting example. I have obtained speci-
mens from family groups of these geese along the Koyukuk and
Alatna Rivers about 100 miles south of Anaktuvuk and a specimen
from Kobuk, about 140 miles west of the Alatna and also on the
southern watershed. ‘Two specimens were obtained at Old Crow.
Cade and Schaller found families of these geese along the Colville
River, about 100 miles north of Anaktuvuk (Kessel and Cade, 1958).
The nesting area of these geese in the forested and in the tundra areas
of central arctic Alaska and Yukon is separated by an unoccupied
area, of mountains about 200 miles across. Although there is a recol-
lection of a pair of swans’ nesting in the Killik Valley about 50 years
ago the failure of swans (Olor columbianus), geese (Branta, Anser,
Chen), and cranes (Grus) to nest in the mountains is historic
Nunamiut knowledge.

I am somewhat doubtful about the correctness of designating
shovelers (Spatula clypeata) as migrants because I do not know that
they nest north of Anaktuvuk. Although they have been reported in
only two years of my observations, they are well known to Nunamiut
and other Eskimos, but Eskimo acquaintance with shovelers shows
that rarity rather than frequency of observation distinguishes these
unique-appearing ducks. From questioning I am sure that very few
shovelers have been seen in the Arctic and that the birds are not
common, but since the arctic appearance of shovelers is not a new event
the birds probably should be considered as migrants.

Godwits (Limosa lapponica) are not often seen at Anaktuvuk but
the few observations indicate that they move swiftly through the
mountains. ‘They are well known and named explicitly by the
Nunamiut, who have seen them nesting farther north, and I do not
hesitate to call the godwits regular migrants through Anaktuvuk.

Stilt sandpipers (Aficropalama himantopus) and _ sanderlings
(Crocethia alba) have not often been reported at Anaktuvuk and the
numbers observed have been so few that I am not certain that they
were on a normal migratory path which would lead them successfully
to a nesting ground. Sanderlings are well known along the arctic

STATUS AND DISTRIBUTION 253

coast insummer. Nunamiut have reported to me that stilt sandpipers
nest near the Colville mouth, and Tom Brower knew of a nest that
was found near Barter Island. I include these sandpipers as migrants
thinking that their rapid progress in part accounts for the few
observations.

Under the designation of visitors I list the following 22 species
which come to Anaktuvuk in summer but not to nest there or farther
north:

Nearest known

Visitor nesting place
Podiceps auritus cornutus Bettles
Accipiter gentilis atricapillus Hunt Fork
Haliaeetus leucocephalus alascanus Koyukuk River
Circus cyaneus hudsonius Alatna River
Pandion haliaetus carolinensis John River
Falco sparverius Sparverius. Yukon River
Bartramia longicauda Eastern Alaska
Erolia fuscicollis. Arctic coast
Erolia alpina pacifica Arctie coast
Stercorarius pomarinus Arctic coast
Larus argentatus smithsonianus Alatna River
Xema sabini. Arctic coast
Bubo virginianus lagophonus. Hunt Fork
Aegolius funereus richardsoni. Hunt Fork
Colaptes auratus borealis Bettles
Hirundo rustica erythrogaster. Kobuk Village
Iridoprocne bicolor. Bettles
Riparia riparia riparia Bettles
Regulus calendula calendula Hunt Fork
Dendroica petechia amnicola. Kobuk Village
Dendroica coronata hooveri Kobuk Village
Juneo hyemalis hyemalis Alatna River

I regard their attachment to Anaktuvuk as more casual] than that of
the three species that usually come northward from the forests to
feed in winter among the willows on the tundra, or of the snowy owls
(Nyctea scandiaca) that usually appear in late winter. I suspect,
however, that some of the species now called visitors will subsequently
be found nesting near or north of Anaktuvuk. For example, marsh
hawks (Circus cyaneus hudsonius) have been seen a number of times
and are so familiar to the Nunamiut that an explicit name is applied
to them.

Two reports indicate that swallows have nested north of the Brooks
Range. Anderson (1921) reported seeing the characteristic burrows
of bank swallows (Riparia riparia riparia) along the Hula Hula
River. I have not seen their burrows north of the woodlands and
the Nunamiut, who study as problems the absence of birds well known
elsewhere, do not know of any burrows north of the Koyukuk. We

254 U. S. NATIONAL MUSEUM BULLETIN 217

occasionally saw bank swallows in spring at Anaktuvuk and they
nest commonly 100 miles south, along the Koyukuk and lower Alatna
Rivers.

Simon Paneak recalled (see p. 90) seeing mud nests with frozen
young cliff swallows (Petrochelidon pyrrhonota) while hunting with
some other young men along the west branch of the Kuparuk River,
about 40 miles northeast from Anaktuvuk. His report indicates that
a colony once ventured to settle in an Alaskan location 200 miles north
of any place where they are now known to nest. Their attempt was
unproductive and it cannot have been often repeated or the con-
spicuous evidence of their nesting would have been remarked.

In 1849 Rae reported seeing the nests of cliff swallows on cliffs along
the Coppermine River (Richardson, 1852), but I do not know whether
that was a single venture or a sustained habit. I have never had a
report of a cliff swallow at Anaktuvuk, although they are the most
numerous nesting swallows around Bettles. They are more colonial
than the other swallows and perhaps do not wander so much indi-
vidually.

It was only in 1952 that I obtained evidence that a rusty blackbird
(Huphagus carolinus) had raised young in the mountains. Indi-
viduals are seen each year and as their common nesting grounds are
nearby this event cannot be taken to represent a new trend. Except
for the two reports on swallows the evidence obtained so far indicates
no tendency of the birds designated as visitors to expand from
transient to more lasting occupation of Anaktuvuk. In the stories
from past years I can find no sign which shows that expansion or con-
traction of the range of birds at Anaktuvuk is visibly in progress.

The common range of these visitors is not far from Anaktuvuk,
17 being regularly seen along the Koyukuk and Alatna Rivers, and 4
being known to nest along the arctic coast. The frequency of their
brief appearances varies, but only a few hours’ flight would be re-
quired for them to travel from their known regular ranges.

As I have noted in numerous instances in the descriptions of the
species, the occasional appearance of these visitors has been long
remarked by the Nunamiut. Some of the intervals between observa-
tions were long (about 40 years separates two occasions when horned
grebes were seen) but the birds were well remembered, explicitly
named, and regarded as rare but not strange appearances.

While we must regard the evidence of bird occurrence at Anaktuvuk
as indicative of a well established fauna, it is reasonable to suggest,
however, that the transient visits of certain species represent a tendency
among birds from both north and south of Anaktuvuk to make occa-
sional sallies from their usual range. That is, the visiting habit is
thus a normal form of behavior in those populations, even though it is

STATUS AND DISTRIBUTION 255

infrequently apparent. What it shows is that some birds are curious,
restless, or otherwise unconstrained by the influences which hold the
rest of their populations to their present nesting grounds. <A very
elaborate and unlikely conjunction of physiological and environmental
conditions would be necessary to expand or contract a well established
nesting area. The persistence of visiting nevertheless suggests that
inherent inclinations exist which could, under special conditions, ef-
fectuate an increase in range. The long persistence of the occasional
expression of the tendency shows how seldom it changes the range of a
population and indicates the essential conservatism of bird distribu-
tion at Anaktuvuk.

The Birds of Kobuk

At first view the Kobuk avifauna as summarized in table 4, seems not
to differ as much from that of Anaktuvuk as do the environmental con-
ditions. Kobuk is in a wooded valley, Anaktuvuk is in mountain
tundra. Climatically they differ, in that at Kobuk frost-hardy vege-
tables can be grown in the usual summer warmth, whereas at Anak-
tuvuk warm summer days are too infrequent for growing vegetables
and hard freezes occur there as often as mild summer frosts at Kobuk.

A rather large difference appears in the status of the birds of the
two areas, for whereas at Anaktuvuk 22 are summer visitors from their
nearby ranges, at Kobuk only 10 species, because they are not known
to nest or normally migrate there, are considered to be summer visitors.
The winter best demonstrates the large difference between biotic con-
ditions on the tundra and in the forest, for only 7 birds are regular
winter residents at Anaktuvuk in contrast with 23 at Kobuk.

Numerous other differences are apparent to the observer who has
lived in these places. Swallows are rare visitors to Anaktuvuk whereas
three species nest at Kobuk. Six species of warblers nest in large
numbers at Kobuk, but only one has been occasionally found nesting
at Anaktuvuk. Several species of sandpipers, common along the
tundra streams, are scarce in the wooded valleys, and at Kobuk long-
spurs (Calcarius), pipits (Anthinus), and larks (Hremophila) are to
be found only in the tundralike elevations above the valley floor.

Large migratory flights of geese and ducks and lesser flights of gulls
pass along the Kobuk River in spring, the gulls appearing to come
from the coast. It is not apparent whether the geese and ducks come
from the interior or from the coast, but flights of geese can be often
seen to leave the river for the north. The direction of migration of
small birds is especially obscure in wooded country. Since most of
the land birds and many of the shore birds migrate northward through
western Canada, these must have traveled some 500 miles westward
across Alaska in order to reach Kobuk. The few records of arrivals

256 U. S. NATIONAL MUSEUM BULLETIN 217

indicate that the migratory species reach Kobuk a few days before
their arrival at Anaktuvuk. Several of these species also appear to
reach Old Crow earlier than Kobuk in conformity with their suspected

Birds of Old Crow

Longer observation at Old Crow might add to the small number
of visitors now recorded there, but it seems that the nesting birds are
established firmly in their migration to the Porcupine Valley, which
is also the northern limit for migration of more species than either
Kobuk or Anaktuvuk. Also the smallest number of species migrate
through Old Crow without leaving any nesting pairs. Summer at
Old Crow is less warm and is shorter than at Kobuk, but is sufficient
for the growth of spruce to fair size. Warblers, thrushes, and spar-
rows form dense populations in some of the brushy areas along the
Porcupine River, showing little attenuation in density of population
at the northern limit of their ranges. Surface-feeding ducks are not
especially numerous but for old squaws and scoters it is an important
nesting ground. The marked discrepancy at Old Crow appears in the
small number of species and individuals of sandpipers and plovers,
both nesting and migrating. Except for a few species using the in-
land marshes the Porcupine Valley is an insignificant nesting area and
unimportant as a migratory route for members of these groups.

Birds of the Arctic Coast

For records from the Yukon arctic coast I have selected from Rand
(1946) the reports from Herschel Island westward to the Yukon-
Alaska boundary. From Anderson’s (1921) journal account and from
Brooks (1915) and Dixon (1943) I have taken records pertaining
to the arctic coast of Alaska between the mouth of the Colville River
and Demarcation Point.

Bailey’s (1948) extensive report on the birds of arctic Alaska
covers a coast line and certain interior localities very diverse as habi-
tats of birds. I have chosen Barrow as the best studied locality.
From Bailey’s explicit references to the locations where observations
were made I have been able to select a list of the birds found within
about 40 miles distance from the village of Barrow. Even with that
restriction, the list of birds found at Barrow presents a strange
assortment difficult to resolve into a pattern related to our knowledge
of the geographical distribution of birds.

Bishop (1944) described the situation at Barrow well by saying
that “of about 140 forms recorded from the northern coast of Alaska
about 48 have been taken not over four times and 34 only twice. This
would indicate that most of them were lost birds, wandering until

STATUS AND DISTRIBUTION 257

they came to the sea.” Since that time new observations have been
made, but Barrow is still a locality where unusual birds appear so
often that its exceptional avifauna warrants further comment.

Out of the 126 species counted as from near Barrow in table 7, about
17 are still known from only one specimen and 8 from only two speci-
mens. For many years expert naturalists have studied at Barrow.
In addition to the published accounts of these distinguished natural-
ists there are the observations, extending over 60 years, of Charles
Brower and his sons, a family which has provided the first specimens
of 63 species and subspecies of the birds now known from arctic
Alaska. From his vivid, pictorial memory and with precise recollec-
tions of time and circumstances Thomas Brower has related to me that
these rare appearances of unusual birds clearly implied to him that
they had come as strangers to Barrow.

TABLE 6.—Distances from their nearest reported nesting grounds Of some
birds recorded at Barrow

from Speci-
nearest Birds mens
nesting

place

2400 | Piranga olivacea
1800 | Tyrannus tyrannus
1800 | Euphagus cyanocephalus
1200 | Troglodytes troglodytes alascensis
1100 | Agelaius phoenicus arctolegus
1100 | Piranga ludoviciana
1000 | Charadrius vociferus vociferus
1000 | Chordeiles minor minor
900 | Spizella passerina boreophila
800 | Hylocichla guttata guttata
800 | Passerella iliaca unalaschcensis
800 | Oporornis tolmiei
800 | Contopus sordidulus
700 | Falco sparverius
500 | Sialia currucoides
300 | Tachycineta thalassina
300 | Iridoprocne bicolor
300 | Hirundo erythrogaster
300 | Petrochelidon pyrrhonota
300 | Riparia riparia
300 | Dendroica coronata
200 | Wilsonia pusilla

me O68 Om mh he BOD om ee DO |

Some land birds found at Barrow were far from their known nest-
ing grounds and on a terrain and in a climate very strange to them.
Some of these strange birds from Bailey’s report with my estimate of
the distance to their usual nesting grounds are listed in table 6. The
location of their nearest nesting ground was obtained from the A.O.U.
Check-list (1957) or from my own experience with birds nesting in
Alaska. Although better information may modify the estimated

258 U. S. NATIONAL MUSEUM BULLETIN 217

distances, the fact will remain that many birds have reached Barrow
after long travel over country in which their ability to exist comes
to us as a surprise.

By comparison most of the visitors to Anaktuvuk (see p. 253) are
known to nest within 100 miles of that locality and none were more
than 300 miles from their normal nesting range. In distance from
regular nesting grounds they resemble only the end of the list of oc-
casional birds at Barrow, and no bird was as far from its proper
nesting place as were the birds of 15 species which have been recorded
at Barrow.

The presence of strange birds at Barrow and the lack of them at
Anaktuvuk is apparently characteristic of each locality, for it does
not seem possible that such a great disparity in phenomena can be
ascribed to difference in intensity of observation. Point Barrow is
unique because it is the northernmost continental locality in the
western arctic. The promontory is worked by conflicting ocean cur-
rents that deposit along the coast to the west of Barrow, driftwood
originating in the Yukon Valley and to the east driftwood from the
valley of the Mackenzie (Giddings 1952) and often the coasts on
each side of Barrow have such differing ice conditions that ships have
to wait for a change before rounding the point. It is near the
eastern end of the annual migration of Pacific walrus. Many older
villages have occupied the locality, which in the past has evidently
attracted people to settle there during several cultural periods. These
characteristics indicate that Barrow has been an important point in
the distribution of animals and man.

Land birds may concentrate at a promontory and they are likely
to be attracted to the dwellings of people in a landscape otherwise
little marked. These circumstances concentrate stray birds and lead
to their observation, but they do not explain why the birds went so
far from their regular haunts, traversing a country far different from
the normal experience of their populations. It is not possible to in-
voke storms or weather as causes which could divert them for such
distances and through such a diversity of terrain and local weather
as they must have passed. At present we must take the appearance of
humerous strange birds at Barrow as an interesting fact we cannot
yet connect with the rest of our knowledge of the geographical dis-
tribution of birds.

It is these strange birds which make the Barrow avifauna more
numerous than that in the other localities. Among the birds normally
found at Barrow are several of the maritime birds so numerous on
Bering Sea. These birds are absent from the interior, and relatively
few are found along the coast eastward from Barrow. Few land

STATUS AND DISTRIBUTION 259

birds except snow buntings (Plectrophenawx nivalis) and Alaska long-
spurs (Calcarius lapponicus alascensis) are found common at Barrow,
and many that are common at Anaktuvuk occur there rarely.

Distribution in Western Arctic America

In table 7 is presented a comparison of distribution in six localities
(see maps, figs. 1 and 8): (1) Anaktuvuk; (2) Kobuk Village, on the
forested middle and upper part of Kobuk River about 150 miles
west by south from Anaktuvuk; (3) the arctic part of interior Yukon
about Old Crow, 350 miles east of Anaktuvuk; (4) the eastern arctic

Taste 7.—Species of birds in the families reported from six localities
of western arctic America

Arctic coast
Families Anak- Kobuk Old
tuvuk Crow Eastern
Yukon | Alaska | Barrow

Gaviidae
Podicipedidae
Procellariidae
Phalacrocoracidae
Anatidae
Accipitridae
Pandionidae
Falconidae
Tetraonidae
Gruidae
Charadriidae
Scolopacidae
Phalaropodidae
Stercorariidae
Laridae
Alcidae
Strigidae
Caprimulgidae
Alcedinidae
Picidae
Tyrannidae
Alaudidae
Hirundinidae
Corvidae
Paridae
Cinclidae
Troglodytidae
Turdidae
Sylviidae
Motacillidae
Bombycillidae
Laniidae
Parulidae
Icteridae
Thraupidae
Fringillidae

_

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OoccoccononoococroroocoocorF ORF KF OH YE NF ON KR OOF &
_

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NOK WRFONNWOF FN WHERE NOCOCROUTUWN OAKEY NRK ATO OF
ROR ORF RF NNR OrF AON RF ORF ODOSOAWNATRFWNH DBWOON

WONWWOOCONKNRFN FOF REF NF WOCrKH WN OCW NH RP NE NWO NW

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ToTaL 106 122 107 62 58 126

260 U. S. NATIONAL MUSEUM BULLETIN 217

coast of Alaska from Flaxman Island to Demarcation Point, some
250 miles northeast of Anaktuvuk; (5) the arctic coast of Yukon
Territory, about 350 miles east by north; and (6) Barrow, about
250 miles north.

I believe that the small number of species reported on the eastern
Alaskan (58 species) and Yukon coasts (52 species) is significant
of the true situations there. While observers and collections have
been few, Anderson (1921), Dixon (1943), and Brooks (1915) each
spent the cycle of the seasons there. They were skillful and energetic
naturalists and I think it unlikely that they missed much. Reading
their records impresses me with their reliability as indices of an
avifauna which is meager in species and probably in numbers. Their
records agree with information I have derived from Nunamiut and
other Eskimo accounts.

While birds of strict maritime habits are absent from the arctic
interior, some loons, ducks, certain sandpipers, jaegers, and gulls that
in winter are maritime birds on the Pacific coast and adjoining islands
migrate in summer to nest in the interior of arctic Alaska and Yukon.

Of the four species which migrate from the continent of Asia to
nest in Alaska only wheatears (Oenanthe oenanthe oenanthe) were
found at Old Crow. We did not find Kennicott’s willow warbler
(Phylloscopus borealis kennicotti), Alaska yellow wagtails (Motacil- .
la flava tschutschensis) and red-spotted bluethroats (Luscinia svecica
svecica), nor do they appear on Rand’s (1946) list of Yukon birds.
On the eastern arctic coast of Yukon only wheatears and wagtails
among the Asiatic birds have been reported. Thus the migration of
these land birds from Asia hardly carries them beyond Alaska on
the American continent.

There is a difference in the ducks of arctic Yukon and Alaska in
that the proportion of lesser scaup (Aythya affinis) to greater scaup
(Aythya marila nearctica) at Old Crow was about like the propor-
tion of greater to lesser scaup at Anaktuvuk. Golden-eyes (Buce-
phala clangula and islandica), common at Old Crow, do not appear
to reach arctic Alaska.

The species and individuals of plovers and sandpipers at Old Crow
were fewer than at Anaktuvuk and Kobuk and except for snipe
(Capella gallinago), yellowlegs (Totanus flavipes) and solitary sand-
pipers (Z'ringa solitaria cinnamomea) none of their populations were
numerous. In Rand’s (1946) list only 16 species of sandpipers are
reported from all Yukon. The common coastal forms seem seldom
to migrate through interior Yukon, although many traverse Alaska
at Anaktuvuk and Kobuk. In the Mackenzie Delta Porsild (1943)
reported only 11 species of sandpipers.

STATUS AND DISTRIBUTION 261

I had earlier shared Bishop’s opinion, formed as he traveled west-
ward along the Yukon River that the number of species of American
land birds gradually diminished westward in Alaska. But compari-
son of the avifauna of Old Crow, Anaktuvuk, and Kobuk shows the
decrease to be marked only in the family of flycatchers (7'yrannidae),
none of which have been reported at Kobuk. Swainson’s thrush
(Hylocichla ustulata ineana) has not yet been identified at Kobuk, but
I think that it occurs there. Otherwise the swallows (Hirundinidae),
thrushes (Z'urdidae), and even the small wood warblers (Parulidae)
populate the arctic woodland thickly from Old Crow to Kobuk and
some species of thrushes and wood warblers commonly nest on the
tundra.

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ic teat Sty igh

Pireaupae tae Pariede: hm Kyte Fk a4

DY Mesos: WRG Sop aee

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4
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i

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eet Or fe ee BGs ¢ ea 5 ere his 4
Ra; hae ea Payee vata At Ae: ia

; i i

hy e é. 4, a i . » oe
, er ft es i Se ee eo a ae

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vv

ne nga le

M iy 4 pnveny hpl
i Poa e. Sk PN
“f at, Ni Liat papel ‘ in , ie a ahd Rete 3 , wea Ben 6, " re Ms i they

ry NEE it Pt ie Mg r: enn

‘ho aoe Rad soda! bar sala oe meats 40 ri agape ab ;
Bs) bawer” Sain eh! Ab oe

| oe cm aed Me —
ay Cat vallowlnan (Patan

ch, onli tf hy ;
Hck nodule hy rineeinientaim sihe ii “ein ps

PRI YES ah M ists
h

Beem
oan, * : ;
koartad ri a)
tf obagarate ‘Chroust |
at Samad anc Tope nt
TOeh shee any. ad Ag ben alee pit anil iat

Canapa Goose Branta canadensis (see pp. 34, 134, 162).

6. Migration and Origins

fi ICE OF THE LAST and maximal Wisconsin glaciation, it has been
earlier pointed out, lay over Canada and southern Alaska for
many thousand years (Flint, 1947) and separated an unglaciated area
in northern Alaska and Yukon from the southern parts of North
America, which during that time remained in habitable condition
for birds. It is doubtful that the climate and ice of northeastern
Siberia then permitted migration to Alaska from temperate parts of
Asia. The maximum stage of this ice period was probably about
10,000 years ago and current developments in glacial geology tend
to shorten the estimates of the time during which the migrations we
now see could have established their present routes to arctic North
America. It is evident that the migration of birds to Alaska is a com-
paratively modern development and the distances many species have
traveled to occupy new areas point out the extent of this recent re-
distribution. There must also have been a great multiplication of
numbers as new populations developed to occupy the extensive lands
in most of Alaska, all of Canada, and some northern States as they
263
469496—60——-18

264 U. S. NATIONAL MUSEUM BULLETIN 217

were released from the cover of ice. This expansion in distribution
and numbers of birds appears favorable for the development of new
populations and their subsequent variation from the species type.

It is tempting to hope that in their geographical courses the annual
migrations we now observe may show traces of their development
which, in relation to climatic history, will recapitulate the kinetics
of avian processes of distribution and reveal the geographical in-
fluences which assist the differentiation of species into races. It is
obvious that I think that the species now migrating to Alaska and
Yukon did not remain there during the last ice age, and that they
then occupied milder southern territories. The habits and tolerance
of cold of most of the migratory species make it unlikely that they
had traffic with the Arctic over an ice cap. I do not feel equal assur-
ance in speculating upon whether the species now resident in arctic
Alaska and Yukon remained in the unglaciated regions while the land
to the south was covered with ice, although the hardiness of the
present resident land birds makes it plausible that they could have
remained if the conditions provided them with food.

TasLeE 8.—Ranges of birds present in winter
[Explanation—(P) presence established. American range: (A) Alaska, (Y) Yukon, (M) northwest

Mackenzie.]}
Range
Species Anak- | Kobuk Old
tuvuk Crow

World American

Accipiter gentilis atricapillus P P Eu As Am

Falco rusticolus obsoletus P P P Eu As Am
Canachites canadensis osgoodi P Pp Am AYM
Bonasa umbellus yukonensis P Am AYM
Lagopus lagopus alascensis P P P Eu As Am AYM
Lagopus mutus nelsoni P P iP. Eu As Am AYM
Bubo virginianus lagophonus P P P Am AY
Nyctea scandiaca P P P Eu As Am

Surnia ulula caparoch P P Eu As Am

Strix nebulosa nebulosa P P Eu As Am

Acgolius funereus richardsoni P P P Eu As Am
Dendrocopos pubescens nelsoni P P Am

Picoides arcticus P Am

Picoides tridactylus fasciatus P P P Eu As Am

Corvus corax principalis P P P Eu As Am

Perisoreus canadensis pacificus P P P Am AYM
Parus atricapillus turneri P P Am A
Parus cinctus lathami P P Eu As Am AYM
Parus hudsonicus hudsonicus P P Am

Cinclus mexicanus unicolor P P P Am

Bombpycilla garrula pallidiceps P P Eu As Am

Pinicola enucleator alascensis P P P Eu As Am AYM
Aeanthis hornemanni exilipes P P P Eu As Am

Acanthis flammea flammea P Eu As Am

Loxia leucoptera loucoptera P P Eu As Am

Total (25) (13) (24) (21)

oe ———

MIGRATION AND ORIGINS 265

The winter-resident birds of the forested parts of arctic Alaska
and Yukon (table 8) are well represented at Kobuk and Old
Crow, but only 4 of the 13 at Anaktuvuk are strictly birds of
the tundra, showing the restriction imposed by lack of trees. These
species are now adapted to life on arctic tundra and might have
survived under similar conditions during an ice age. Inasmuch as
all 25 species also now live in arctic forest, that condition would
have been more suitable. As for their present world distribution, 9 of
the species are now confined to America and 16 also extend widely
over Asia and Europe. Of the 10 American species 5 are dif-
ferentiated as races restricted to Alaska, Yukon and northwestern
Mackenzie. Among the 16 holarctic species are 4 races restricted to
the area. The differentiation of 9 populations resident in the area
implies that they have been long isolated from interbreeding with the
remainder of their species. It also indicates a certain degree of faunal
homogeneity of the arctic interior of western North America from
the western border of forest to some line near the Mackenzie Delta
where contact is made with other resident races.

Species and their Migration Routes

Distinguishable populations of 19 migratory species of land birds
nest mainly in Alaska, Yukon, and western Mackenzie, as shown in
table 9. Of these, 3 (in Aphriza, Heteroscelus, and Limnodromus)
have no nesting range outside the area except on the adjacent
coast of Siberia. The American populations of Oenanthe and Plec-
trophenax are not recognizable by taxonomic distinction; but I have
indicated that the Alaskan snow buntings appear distinguishable in
the thickness of toes and claws. The wheatears are not taxonomically
distinguishable from those of Europe and Asia, although their migra-
tion to nest in northwestern America takes them far east of their
Asiatic range. Among the 14 populations assigned to taxonomic
races nesting only in Alaska and Yukon, I find local differentiation
visible in 8 racial designations that are not considered as established
by the A.O.U. Check-list (1957): Branta canadensis taverneri,
Leucosticte tephrocotis irvingi, and Arenaria interpres interpres. The
separation of the latter from A. 7. morinella rests upon distinctions
which are difficult to make and accordingly their ranges are uncertain.
In spite of these reservations as to the distinction of the populations,
3 species and 11 races nesting mainly in northern Alaska and Yukon
are commonly recognized and the area is characterized by the nesting
of a number of distinguishable populations of migratory species.

266 U. S. NATIONAL MUSEUM BULLETIN 217

TABLE 9.—Land birds migrating ‘to nest mainly in Alaska, Yukon, and western
Mackenzie

[ Explanation—(P) present, GD migrant. Migration routes: (W. Am) western America, (EH Am) eastern
America, (Pac) Pacific, (Asia) Asia, (Al) Alaska.]

Birds Migration | Anak- | Kobuk |Old Crow| Range of
route tuvuk the species
Branta canadensis taverneri W Am M Pp 12 Am
Arenaria interpres interpres Pac M M Eu As Am
Aphriza virgata W Am Pp Am
Tringa solitaria cinnamomea W Am Pp P P Am
Heteroscelus incanum Pac Pp P Am
Limnodromus scolopaceus W Am Pp 12 Am
Limosa lapponica baueri Asia P Pp Eu As Am
Sayornis saya yukonensis W Am P P Am
Eremopbila alpestris articola W Am Pp P P Eu As Am
Phylloscopus borealis kennicotti Asia P P Eu As Am
Motacilla flava tschutschensis Asia P Pp Eu As Am
Txoreus naevius meruloides W Am 12 P Am
Hylocichla ustulata incana E Am iP Am
Luscinia svecica svecica Asia P Eu As Am
Oenanthe oenanthe oenanthe Asia P P P Eu As Am
Dendroica coronata hooveri W Am P P Am
Leucosticte tephrocotis irvingi W Am P ie Eu As Am
Calearius lapponicus alascensis W Am P P M Eu As Am
Plectrophenax nivalis nivalis Al M M M Eu As Am

TOTAL (19)

The migratory routes of these populations are diverse: 10 pass
through western America, 1 through eastern America, 2 over the
Pacific Ocean, 5 through Asia, and 1 lies within Alaska; while 9
of the species listed nest only in America, and 10 have a widespread
distribution in the three northern continents. It is unlikely that
any of these species migrated to Alaska and Yukon during the maxi-
mum of Wisconsin glaciation and their present migrations have
developed since the American ice cap melted. Their differentiation
in form has accordingly been brought about by special migratory
habits which have isolated these populations within Recent geological
time.

Of the 35 species, shown in table 10, that nest in Alaska and Yukon
and winter or migrate as maritime birds along Pacific American
coasts, 11 also nest over variously wide extents of North America and
the remainder are holarctic in distribution. A number of these species
also migrate along Atlantic coasts, but we have observed birds of
several of these species migrating to Old Crow from the west, and
there is evidently some segregation of the Pacific populations. Only
3 of these species (in the genera Vumenius, Erolia, and Larus) are
differentiated as western American races, but these also nest east of
Mackenzie. Although neither restricted in range nor differentiated
as local races the use by these winter maritime species of long over-
land flight routes through complex terrain shows that special migra-

MIGRATION AND ORIGINS 267

tory habits are established among these maritime species which nest
in Alaska and Yukon. In none of these maritime species, however,
have the specialized migratory habits resulted in the visible differ-
entiation which distinguishes the resident and most of the migratory
populations of land birds which are restricted in nesting to Alaska
and Yukon. Judging from their powers of flight and adaptability
to arctic conditions these species could have been among the earliest
to come to Alaska after the last glacial maximum. It appears that
the isolation produced by special migratory paths and special nesting
areas is broken down by behavior after nesting which leads to mixing
with the remainder of the species. Here the specific behavior has
prevented physical differentiation.

TaBLE 10.—Birds wintering or migrating along American Pacific coasts

[Explanation—(P) present.]

Birds Anak- | Kobuk |Old Crow| Range ef
tuvuk the species
Gavia immer iz iP 12 Am
Gavia adamsi P P Eu As Am
Gavia arctica pacifica P 1p qe Eu As Am
Gavia stellata Pp ie le Eu As Am
Podiceps grisegena holbdéllii P P Eu As Am
Podiceps auritus cornutus P ie P Eu As Am
Olor columbianus ie if 1 Am
Branta nigricans P P P Am
Chen hyperborea hyperborea 12 ip 1 Am
Aythya marila nearctica P if P Eu As Am
Bucephala clangula americana P Eu As Am
Bucephala islandica P Am
Bucephala albeola P Am
Histrionicus histrionicus P 12 TE Am
Clangula hyemalis P P P Eu As Am
Melanitta deglandi 12 12) lz Am
Melanitta perspicillata P P 1 Am
Oidemia nigra americana ie Eu As Am
Mergus serrator serrator ip Ie iE Eu As Am
Aquila chrysaetos canadensis Pp P ip Eu As Am
Haliaeetus leucocephalus alascanus P P ie Am
Squatarola squatarola ie P Eu As Am
Numenius phaeopus hudsonicus P P AE Eu As Am
Erolia alpina pacifica P ie Eu As Am
Crocethia alba P P Eu As Am
Phalaropus fulicarius P P Eu As Am
Lobipes lobatus P P P Eu As Am
Stercorarius pomarinus le ig Eu As Am
Stercorarius parasiticus iz iP P Eu As Am
Stercorarius longicaudus Le: P te Eu As Am
Larus hyperboreus barrovianus le iz Pp Eu As Am
Larus canus brachyrhynchus P Pp LE Eu As Am
Larus philadelphia if P Am
Xema sabini P P Eu As Am
Sterna paradisaea Pp le Le Eu As Am

Torat (36) (30) (34) (26)

268 U. § NATIONAL MUSEUM BULLETIN 217

Birds of 25 species that migrate overland in northwestern Canada
and probably through the western mountains to nest in northern
Alaska and Yukon, are listed in table 11a. Although swallows
(Hirundinidae) are widely distributed across North America during
migration I have included all 5 species as probably flying northward
along a western course to Alaska because the early northward progress
of the spring migration of swallows in the west fits their time of
arrival in Alaska and Yukon. Two of the five swallows (Zachycineta
and Petrochelidon) are differentiated as western races. The difficulty
in proposing the migratory paths of some wide-ranging species 1s
shown in the cases of the Savannah sparrow (Passerculus) and Junco
(Junco), for both migrate through most of the northern states. In
these cases one may suspect that these migrants to Alaska come
from the western fringe of the widespread populations. The majority
of these birds can be regarded as western American and 14 of them
are taxonomically distinguished as western races. Of the species
listed in table 11a, 5 nest in the Arctic far east of Yukon, 9 have
already been referred to (table 9) as providing races mainly restricted
to nesting in Alaska and Yukon, and 11 are assigned to races which
nest over a considerable American area south of the Arctic. But
among these 25 western migrants, 20 reach the Arctic only in
Alaska, Yukon, and western Mackenzie, so that their most northern
nesting range in the west is apparently related to the favorable
environmental conditions of Arctic Alaska and Yukon.

The 17 species that migrate principally east of the Rocky Mountains,
as shown in table 11b, can be assigned to the central flyway, which is
routed over the watershed east of the Rocky Mountains. Among
these species the 2 thrushes (7urdus and Hylocichla), the kinglet
(Regulus), pipit (Anthus) and 5 wood warblers (Vermivora, Den-
droica petechia and striata, Seiurus, and Wilsonia) also migrate
through paths east of the central States and Provinces. Alaska and
Yukon are only part of the extended nesting areas of these birds, but
9 of them reach the Arctic only in Alaska, Yukon and Mackenzie.
Again the favorable nesting environment in the western arctic is
apparent.

Only 5 species nesting in Alaska and Yukon (table 11c) can be
regarded as migrating mainly east of the Mississippi River. There is
some migration of each of these species west of the Mississippi but
they are commonly regarded as eastern birds. The arctic nesting races
of two of these species are restricted to Alaska and Yukon. All of
them also nest south of the arctic circle.

MIGRATION AND ORIGINS

TaB LE 11.—Status of various migrating birds
(Explanation—Arctic nesting range: (A) Alaska, (Y) Yukon, (M) Mackenzie,

(Eu) Europe, (As) Asia, (Am) America.]

Arctic Anak- Old
Species nesting tuvuk | Kobuk Crow
range
A. MIGRATING THROUGH BRITISH COLUMBIA
Branta canadensis taverneri AY ie) 1 P
Anser albifrons iz P 1S
Falco columbarius bendirei AY iP Le iy
Grus canadensis canadensis P iP 12
Tringa solitaria cinnamomea AYM pe Pp P
Limnodromus scolopaceus A P 1
Megaceryle aleyon caurina A P
Sayornis saya yukonensis AY 1 P
Eremophila alpestris arcticola AY P P 12
Tachycineta thalassina lepida AY 15) P
Tridoprocne bicolor AYM 12) P
Riparia riparia riparia AY P ie
Hirundo rustica erythrogaster AY P
Petrochelidon pyrrhonota hypopolia AY P P
Ixoreus naevius meruloides AY Pp iy)
Lanius excubitor invictus P P 1s
Dendroica coronata hooveri AY P P P
Wilsonia pusilla pileolata A Ie P
Leucosticte tephrocotis irvingi AY 1e Es
Passerculus sandwichensis anthinus P P Ve
Junco hyemalis hyemalis AY P P
Spizella arborea ochracea P Te P
Zonotrichia leucophrys gambelii AYM je iv P
Zonotrichia atricapilla A Pp
Calcarius lapponicus alascensis AYM 12 P P
TOTAL (25) (16) (24) (19)
B. MIGRATING BETWEEN THE ROCKY MOUNTAINS AND THE MISSISSIPPI RIVER
Bartramia longicauda 1p i)
Totanus flavipes 12 ie de
Erolia melanotos P 12 P
Erolia bairdii P 1s) P
Tryngites subruficollis ge 1
Empidonax traillii traillii ne P
Nuttallornis borealis ¥ 12
Turdus migratorius migratorius P 1y jE
Hylocichla ustulata incana AY P
Regulus calendula calendula P ye) Te
Anthus spinoletta rubescens 12) 1E P
Vermivora cclata celata AY 12) P
Dendroica petechia amnicola AY P P Pp
Dendroica striata AY ae P
Seiurus noveboracensis notabilis AY P P
Wilsonia pusilla pusilla AYM P
Calcarius pictus AYM P
Torta. (17) (10) (12) (14)
C. MIGRATING EAST OF THE MIssISsIPPI RIVER
Pluvialls dominica dominica P P bod
Micropalama himantopus P
Hylocichla minima minima P 12 P
Dendroica striata AY P ie
Passerella iliaca zaboria AY P P P
Torat (5) (4) (4) (4)

269

270 U. S. NATIONAL MUSEUM BULLETIN 217

TABLE 11.—Status of various migrating birds—Continued

Arctic Anak- Old
Species nesting tuvuk Kobuk Crow
range

D. MIGRATING TO ASIA FOR THE WINTER

Limosa lapponiea baueri A P P
Oenanthe oenanthe oenanthe Eu As Am P P Pp
Tuscinia svecica svecica A P
Phylloscopus borealis kennicotti A P P
Motacilla flava tschutschensis AY P P
TorTat (5) (5) (4) (1)
E. MIGRATING OVER THE PACIFIC OCEAN
Arenaria interpres interpres AY (?) P P
Heteroscelus incanum A P P
TorTAL (5) (2) (2) (0)
F. WINTERING IN ALASKA
Acanthis hornemanni exilipes Eu As Am P P P
Acanthis flammea flammea Eu As Am P P P
Plectrophenax nivalis nivalis A (?) P P P
TOTAL (8) (8) (3) (3)

A remarkable 5 species (table 11d) migrate after nesting in north-
ern Alaska to winter in Asia. Among these species the godwit
(Limosa) winters on Asiatic coasts, while the others seek some as yet
undefined wintering areas on the continent of Asia. The wheatears
(Oenanthe) are not differentiated from a widespread Asiatic and
European race. The four others are ascribed to races which are re-
stricted to nesting in northern Alaska and Yukon. Long establish-
ment of their remarkable migratory paths to a restricted nesting area
in western arctic and subarctic America is indicated in the racial
designations which distinguish them from close relatives in Asia.

There are 2 species (table 1le) which nest mainly in northern
Alaska and Yukon as far north as the arctic coast and which migrate
widely over the Pacific Ocean to winter on its coasts and islands.
Among the three migratory species which both winter and nest in
Alaska (table 11f) I can only distinguish the Alaskan snow buntings
(Plectrophenax) by a small difference in the thickness of their claws
from the widespread North American race. The winter distribution
of snow buntings suggests isolation of these in Alaska, I have no
evidence to suggest whether the nesting redpolls of Alaska and Yukon
come from the populations wintering in Alaska or from the western
Provinces of Canada.

MIGRATION AND ORIGINS 271

Main Paths of Migration

By utilizing as indicators of migratory paths the differentiation
of races, direction of observed migration and a few significant re-
corded dates of migration on the approaches to the northern nesting
grounds, I have assigned 93 of the species nesting in northern Alaska
and Yukon to migratory paths in North America, over the Pacific,
and in Asia. The majority of these migrations pass through western
Canada and along American Pacific coasts, and only 22 of these
species appear to migrate east of the Rocky Mountains. The totals
from tables 10 and 11 are summarized below:

Along American Pacific coasts 36
Through western Canada 25
Rocky Mountains to the Mississippi 17
East of the Mississippi 5
To Asia 5
Over Pacific Ocean 2
In Alaska 3

Total 93

Without this analysis I would have subscribed to Dixon’s (1988, p. 5)
comment on the birds of Mount McKinley Park, Alaska, a fauna
similar to that of northern Alaska, “a large proportion of the breeding
birds of the McKinley regions consist of eastern forms.” As it is,
I do not.

The designation of birds as western or eastern American depends
on the location assigned to the geographical boundary between east
and west, and to whether the reference is to species or race. It is
further influenced by whether current distribution or the origin of
the form is meant. The present development of taxonomy, geograph-
ical distribution, and recent climatic history are leading to the assign-
ment of each taxonomic form to a meaningful position in time and
space.

The migration of 22 species of birds through North America east
of the Rocky Mountains into northwestern Alaska and Yukon is im-
pressive. In his explorational travels in the Yukon Valley, W. H.
Dall (1870, p. 288) detected the eastern character in the fauna north
of the Alaska Range, and remarked upon the common distribution
of North American pike (#soa# lucius) in the Yukon River as far
west as tidewater as an indication of a nearly continuous connection
by water between northern Alaska and the interior of North America.
By recognizing an extensive fauna in Alaska related to that of the
interior of Canada, Dall concluded that the northern mountains
were readily traversed by birds migrating to Alaska. In his great
study of the fauna of northern North America, Sir John Richardson

272 U. S. NATIONAL MUSEUM BULLETIN 217

(1852, p. 124) saw the geographical relations of the Mackenzie Valley
in a manner different from Dall, for he remarked, “On the Mac-
kenzie there is an intermingling of the flora of both coasts, as
well as of the migratory feathered tribes, the Rocky Mountains not
proving a barrier to either.” Actually Dall’s view, limited to Alaska
and overemphasizing the eastern character of the fauna properly
accentuated the remarkable trend of bird migration in northern North
America to the westernmost parts of northern Alaska and even into
Siberia. This westerly trend in migrations is undoubtedly related to
conditions produced by the east-west direction of mountains and
valleys in Yukon and northern Alaska.

The very complexity of the terrain of southern Yukon and Alaska
leads me to believe that, although we now have few significant obser-
vations of northern routes of migration, we can by reasonable specu-
lation predict the situations in which these routes may be discovered.
It is my belief, based upon observation in northern Alaska, that the
high mountains of the coastal and Alaska ranges must funnel much
of the migratory bird traffic through the few low passes by which the
cold and barren high elevations can be avoided.

Some of the maritime birds wintering on Pacific American coasts
(table 10) must leave the Gulf of Alaska to pass overland to the
Porcupine Valley, where a number of these species were seen arriving
at Old Crow from the west (see p. 157). Their crossing of the Coast
Range may be suspected to start its overland passage east of the high
St. Elias Mountains. Westward along the Gulf of Alaska the next
break in the lofty Coast Range is through the Copper River Valley, .
which in its lower part is still raw from the recent retreat of great
glaciers. Further west the Susitna Valley opens from Cook Inlet
and leads northward, as does the Copper Valley, to passes through
the Alaska Range into the Tanana Valley. From Cook Inlet, also,
Rainey Pass leads through the Alaska Range west of Mount McKinley
to the Kuskokwim Valley. Some of the maritime birds wintering
along the Alaska Peninsula and Aleutians probably cross and round
the Peninsula. These routes are postulated from a few records of
migration but mainly from geographical appearances and as sug-
gestions for a strategy of further observation.

A few of the maritime species (table 10) which winter on south-
western Alaskan coasts probably pass from Bering Sea eastward
along the Yukon, for Dall (Dall and Bannister, 1869) remarked upon
the early arrival of goldeneyes (Bucephala clangula) at Nulato from
the west. According to Dall they winter extensively in the Aleutians.
Mary Lobban informed me that late in October 1957 goldeneyes were
numerous about Cold Bay, near the western end of the Alaska Penin-
sula. Since much of their nesting area lies east of Alaska the west-

MIGRATION AND ORIGINS 273

ern wintering population probably traverses its entire breadth east-
ward in spring. Many maritime birds of the western coasts in winter
come to nest and migrate farther northward in the passes through the
Brooks Range.

Many of the land birds of the Pacific-coast States (table 1la) have
been reported in migration northward between the Coast Range and
the Rocky Mountains in northern British Columbia (Swarth, 1936).
I suspect that those which are headed for Alaska pass through south-
western Yukon into the upper Tanana and Yukon Valleys. Those of
them which reach Old Crow may be suspected of avoiding the cross-
ing of the Ogilvie Range to travel along the Yukon until they can
enter Porcupine Valley. In this way they would be following the
eastward routes of inland travel observed among loons, geese, ducks,
and gulls in their visible flights past Old Crow.

The species which migrate northward between the Mackenzie and
Rocky Mountains (table 11b) are seen in considerable numbers during
migration in southern Yukon. It is likely that they pass over the
valley of the upper Liard River into the headwater valleys of the
Yukon and proceed from there westward into Alaska. With ex-
ception of the sandpipers, 13 of these species are very common nest-
ing birds at Old Crow. If we consider the Ogilvie Mountains likely
to bar them from a direct approach to the Porcupine, they could pass
along the Yukon to the Porcupine and enter its valley from the west,
as do the maritime wintering birds. There is an objection to con-
sidering that these birds would enter Porcupine Valley from the
west because some of them have been observed in the Yukon Valley in
Alaska a week or more earlier than they appear only 140 miles
northeastward at Old Crow. But not enough dates of northward
progress are yet available from the right locations for us to use the
calendar to plot the probable routes of these migratory flights. Some
eastern birds may cross the mountains in northern British Columbia
to reach their nesting grounds around Cook Inlet, but the majority
of these populations nest north of the Alaska Range and seem likely
to enter Alaska along the Yukon.

The five species which migrate through the United States east of
the Mississippi (table 11c) pass northward in the Mackenzie Valley,
probably holding to its warmer western watershed. In its northern
part they bear west to enter Yukon. Since golden plover (Pluvialis)
are rare at Old Crow they probably traverse northern British Co-
lumbia and southern Yukon to occupy their extensive Alaskan nest-
ing areas. The gray-cheeked thrush (Hylocichla minima), blackpoll
warbler (Dendroica striata), and fox sparrow (Passerella) migrate
extensively through southern Yukon with other birds from east of
the mountains.

274. U. S. NATIONAL MUSEUM BULLETIN 217

Near Old Crow we saw a few hawks and geese flying as if in migra-
tion from the southeast. Lazarus Charlie recorded some arrivals of
land birds at Johnson Creek, on the Porcupine 80 miles southeast of
Old Crow, before we saw them. These observations suggest that
some eastern birds reached the Porcupine by flights northwesterly
from the Mackenzie Valley. We have no clear evidence whether
the Porcupine is an important migratory route for small eastern birds
on their flights toward Alaska, but the Peel River breaks through
the southern Richardson Mountains in low passes which, from their
appearance on the map, would be favorable routes from the Mac-
kenzie to the Porcupine Valley, so I suspect that eastern birds, and
possibly some eastern components of the western migrations enter
by this course.

For birds from the westernmost United States to reach western
arctic Alaska requires a westerly flight over at least 40 degrees of
longitude. Birds arriving in arctic Alaska from the Mississippi
Valley must have traveled 70 degrees westerly and some golden
plover (Pluvialis dominica) when southbound from Alaska to Nova
Scotia must traverse 100 degrees of longitude easterly in order to
ciear the Atlantic coast on their overwater flight to South America.

The 45 species of birds which migrate over land through North
America must pass westerly from at least 30 to more than 70 degrees
on their flights to arctic Alaskan breeding grounds. The amount
of westing involved may be seen by comparison with that of about
100 degrees from San Francisco to Manila. The changes in local
time at solar noon encountered by the migrating golden plover
(Pluvialis) during a few weeks of late summer is comparable in
rate and hours to that during a voyage by ship across the Pacific Ocean.

The Asiatic wintering birds (table 11d) reach western Alaska by an
as yet undefined oversea route from Siberia. Since in eastern Alaska
they are concentrated in its northern and arctic parts, I suspect that
they cross Bering Strait.

It may seem that these Asiatic wintering species make only a feeble
penetration into the American continent, but they are not always
easy to see, and they occupy secluded haunts within untraveled areas.
In my experience, with the exception of the bluethroat (Luscinia),
they are numerous and widely distributed. Although their hold upon
the American continent has been regarded as tenuous, actually they are
established in Alaska by easterly migrations as impressive as those of
any American wintering birds. If the wheatears (Oenanthe) which
nest in eastern Alaska come from the central wintering area for the
race in Arabia (Mackworth-Praed and Grant, 1951) their spring
migration passes halfway around the world. Ascribing to them the
most easterly winter residence possible in China, they would still have

MIGRATION AND ORIGINS 275

to migrate across 100 degrees of longitude to reach Yukon. This is a
remarkable flight for a bird weighing 24 grams. And from its nearest
known wintering place in the Philippines (Parkes and Amadon, 1948)
Kennicott’s willow warbler (Phylloscopus) also travels eastward about
100 degrees of longitude to nest in central arctic Alaska. It weighs
less than 10 grams.

The two species which fly over the Pacific (table 11e) also fly along
its shores. Present records give no clue as to their Alaskan landfall.

Along the arctic coast of Alaska a powerful trend of bird migra-
tion moves eastward in spring, in contrast to the predominant west-
ward trend of land birds in the Yukon Valley. These eastbound
migrants along the arctic coast may be assigned to three groups,
according to the length of their arctic flight courses. One includes
the birds wintering on north Pacific coasts (table 10). At Barrow
flights of brant, some shore birds, gulls, and terns are often seen
migrating eastward in spring and westward at the end of summer.
On the eastern arctic coast of Alaska, Dixon (1948) remarked east-
bound flights of loons, brant, and other geese in June and westbound
flights of brant in August. It has been my impression that along
the coast near Barrow the flights of these North Pacific wintering
birds break up in spring and assemble in late summer. East of
Barrow the flights of these birds appear to diminish, while west-
ward they are more often reported, so that coastwise migration of
North Pacific wintering birds tapers off between Barrow and eastern
Alaska.

At Barrow some of the migrating loons, geese, ducks, and sand-
pipers appear in spring as if they were arriving from the interior.
Before we began our studies at Anaktuvuk, Thomas Brower believed
there were indications on the coast that many water birds, and prob-
ably all land birds, coming to the arctic slope and coast had passed
through the Brooks Range. My occasional observations in the coun-
try between the coast and mountains showed that many birds were
coming from the south in spring and forming fiocks for southward
migration inlatesummer. At Anaktuvuk, in the Killik Valley, and at
Howard Pass the migratory movements I have seen were suflicient
to bring large numbers of birds to the arctic slope and coast.

A second group of species, which migrates northward over
Bering Sea and then east along the arctic Alaskan coast, contains
members of the great bird associations characteristic of the Bering
Sea coast in summer. The distinctive birds are cormorants, auks,
murres, guillemots, and puffins. These species reach Barrow and
individuals occasionally pass eastward, but their abundance rapidly
diminishes east of Cape Lisburne, where the last good nesting cliffs
are found.

276 U. S. NATIONAL MUSEUM BULLETIN 217

A spectacular phenomenon of arctic life is seen in the eastbound
flights of four species of eider ducks along the coast in spring at
Barrow. King eiders outnumber all other species and move in such
close sequence of large flocks as to exceed any migration I have
seen. Sir Hubert Wilkins remarked to me that they constitute the
most impressive sight of migratory movement in the world. At the
same time gulls (Zarus hyperboreus and occasionally Rissa tridactyla
and Xema sabinz) move eastward, but in much smaller numbers. The
flights of loons, geese, ducks, and shorebirds, while sometimes numer-
ous, are irregular, so that they do not compare with the flights of
eiders as orderly migratory phenomena.

From reports originating on the eastern arctic coast of Alaska I
gain the impression that the orderly movement of king eiders con-
tinues an easterly migration far beyond Alaska. For other birds the
eastern arctic coast of Alaska does not seem to be an important
migratory route.

There are no reports known to me which indicate that spring
migration passes westward from Mackenzie to the arctic coast of
Alaska. The short list of the avifauna on the eastern Arctic coast
of Alaska suggests that the region is a terminus for the migration
of only a few land birds. The evidence available suggests that some
of the land birds arrive there after crossing near the center of the
Brooks Range and flying eastward down the Colville Valley.

The migratory movements of birds in northwestern America can be
represented by a diagram (fig. 8) showing the main trend of the sev-
eral migratory routes which have been described. For the movements
pursued in arctic Alaska and Yukon the evidence is rather clear, but
the other migratory routes referred to are at present hypothetical.
We thus have the courses of migration delineated with greatest cer-
tainty near their arctic terminus and through a country commonly
regarded as little known. There an orderly pattern appears which
can be easily distinguished because the migratory populations are
homogeneous at the time of their proximity to their nesting grounds.
These orderly currents of migration are related to such major topo-
graphic features as the Rocky Mountains, the Yukon Valley, the
Brooks Range, the Bering and Arctic coasts along with their asso-
ciated influences upon weather and climate.

Migration, the Climate, and Taxonomic Differentiation

I have pointed out a number of distinctions in the races of birds in
northern Alaska and Yukon: all resident species restricted to the
area form taxonomic races (table 8), and of the 19 migratory species
principally nesting in the area, 14 form American taxonomic races

MIGRATION AND ORIGINS 277

BEAUFORT

_——— —
pS ~

—

CHUKCH! SEA

GULF OF ALASKA

bs
oo

Ky
4
t
t

Paairic 2

“%

Ficure 8.—Main migration routes in northwestern America. Large circles (1-3) mark lo-
calities where comprehensive observations have been made by the author. Numbers 4-10
mark localities where observations have been made several times for brief periods, or
once over a period of at least 3 weeks.

Explanation:

1—Anaktuvuk 5—Ahlasuruk River 8—Alatna River
2—Kobuk 6—Killik River 9—Bettles
3—Old Crow 7—Chandler Lake 10—Big Lake
4—Barrow

(table 9). Thus, 23 local populations are distinguished taxonomically
out of 28 resident and migratory species which nest mainly in northern
Alaska and Yukon.

The migratory species come from wintering grounds in America and
Asia distributed over at least 170 degrees of longitude, or nearly half
of the circumference of the earth. The diversity of their winter ex-
perience stands in contrast to the similarity of their summer experi-
ence on the breeding ground which they occupy for only about three
months of the year. If we assume that the races nesting in northern
Alaska and Yukon became differentiated as a result of their segrega-
tion there as isolated breeding populations, the formation of 28 dis-
tinguishable races out of 28 species is a significant step in the separa-
tion of species into races. The characters by which the local races are
distinguished are matters of color, pattern, or dimensions to which
no adaptive value can be assigned and so, as far as can be seen, it is

278 U. S. NATIONAL MUSEUM BULLETIN 217

isolation of the breeding populations which has brought about the
differentiation. Kor the differentiation into races to have affected so
many species implies that the isolation of the northwestern breeding
populations has been in operation for some time. This view leads
to an inquiry as to the nature of the special conditions which are re-
flected in the distinction of the avifauna of northern Alaska and
Yukon.

The geography of the country was discussed in chapter 1
(see p. 6). A. E. Porsild (1951) has remarked that the flora of
arctic Alaska and Yukon is very old and rich in isolated and endemic
species. At the Mackenzie River a sharp boundary separates the
Yukon flora from that of Arctic America in the east. In contrast
there appears an affinity of relation to Asiatic plants in one third of 604
species of Alaska and Yukon. The North American isothermal lines
move rapidly northward in spring west of the Mackenzie River, al-
though in winter only the western coast is markedly warmer than the
interior. Spring advances rapidly northward in the west, or more
correctly, warm Pacific air moves inland over the Rocky Mountains
and over the Alaska Range to the Yukon Valley and extends about as
far eastward as the Mackenzie Valley. Many species of migratory
birds reach arctic Alaska and Yukon by May 15, or at about the date
when they come to northern Ontario in the east. At the time of spring
migration the isothermal lines above freezing extend nearly 20 de-
grees of latitude farther north in western than in eastern America.

The distinctive flora and fauna of northern Alaska and Yukon
about correspond with the area in which the lowlands were not cov-
ered with ice during the last stage of Wisconsin glaciation (see figs.
3and9). Inthinking about the effect of this area as a refuge in which
life might survive while the rest of the northern part of the continent
was covered with ice it must be recalled that some climatic conditions
which left the country free from continuous ice in Wisconsin times
still exist. The general formation of the mountains has not varied,
and the resulting climate and low preciptation may well have been a
persistent distinction of the extreme northwest. Until we can discover
their remains and assign them to dates in glacial history, it is pure
speculation to view the wintering birds now present as derived from
stock which survived in an unglaciated arctic Alaskan refuge. It
seems more plausible to suspect that the warmth of the Pacific Ocean
early melted the coastal ice and made the coasts habitable for maritime
birds and a fit path for coastwise migration before the interior was
cleared. But even now the glaciers around the St. Elias Mountains
leave only a narrow path for land birds along the coast. Although
the majority of the taxonomically differentiated migratory races in
the Alaskan breeding avifauna are western American birds, the fact

MIGRATION AND ORIGINS 279

Ficure 9,—Hypothetical life zones of the Wisconsin age: (A) arctic glaciation, (B) tundra,
(C) Hudsonian, (D) Canadian, (E) Transitional, (F) Upper Austral, (G) Lower Austral.
Heavy broken line marks the limit of 20-inch mean annual rainfall. (From L. S. Dillon,
‘Wisconsin Climate and Life Zones in North America,” Science, vol. 123, fig. 11, 1956.)

that some are of eastern and Asiatic species show that birds from all
sources became differentiated by their relation to the Alaskan nesting
grounds.

Some hardy species of birds, like those which winter, in the Arctic
may have survived in ice-free lowlands of northern Alaska and Yukon,
during maximum glaciation in the Mankato substage of Wisconsin
time but land birds could not migrate over continental ice caps. The
migratory populations now nesting in the northern part of America
must, therefore, have become established within the last 10,000 years.

During the earlier glacial maxima of Wisconsin time the existence
469496—60-—19

280 U. S. NATIONAL MUSEUM BULLETIN 217

of life on most of the northern part of America must have been
similarly reduced, but in the interglacial periods climatic conditions
may have been suitable for repopulation.

The assumption that differentiation of the migratory species of birds
has taken place since the ice cover melted sets a time when populations
migrating to nest in western arctic America could have started their
isolation from more southern members of the species. But although
melting of the ice removed one barrier against the migration of birds
imto the Arctic, it did not at once make a path fit for their travel
and nesting, for the changes were gradual and did not progress
steadily toward the conditions we now see. Relics of vegetation under
sediments and in Alaskan bogs provide evidence that marked climatic
fluctuations have occurred there as elsewhere during the last few
thousand years. The warmest period since the Wisconsin ice, for
example, is indicated to have been widespread in North America
about 6,000 years ago (Karlstrom, 1957), and many species could
then have extended populations far north of their present limits, for
a warmer climate seems likely to favor increase of the variety of living
forms. Only a few hoary redpolls (Acanthis hornemanni) now
winter in the Arctic, and snow buntings (Plectrophenax) and com-
mon redpolls (Acanthis flammea) sparsely winter along the margin
of the Arctic, but a little moderation of climate would allow large
populations of these species to remain as residents and the extent of
many migrations would be reduced.

The larger species of northern mammals are visibly adapted to
cold by their insulation. Although the physiological adaptability of
small mammalian species is limited, large populations of lemmings,
mice, and shrews live in the Arctic sheltered from the cold by their
adaptive behavior. Arctic human populations are adapted to their
cold environment mainly by each individual’s learning of the culti-
vated practices of an arctic society: Birds seem to have devised few
methods of behavior which are obviously protective against cold;
the larger species are adapted by their insulation, but the obviously
successful adjustment of small birds to arctic cold is not now satis-
factorily explainable in terms of physiology and behavior.

Following Bergmann’s (1847) important discussion of animals’
economy of heat in relation to size, it has become popular to say that
the northward increase in dimensions of clines within some species
demonstrates natural selection of a climatically adaptive trend in
variation. P. F. Scholander (1955) has shown that such small di-
mensional changes as are reported in clines are insignificant for the
economy of heat. Therefore they could not be selected for adaptive
value in relation to climate.

MIGRATION AND ORIGINS 281

The physical characters of some races may be adaptive, but it is
a difficult task to demonstrate the adaptive nature of most physiologi-
cal characters even in full species. In a few important studies, local
populations have been shown to have adaptive distinctions. In his
classical demonstration of physiological adaptation of local popu-
lations R. Goldschmidt (1948) found populations of Lymantria dis-
pars at high elevations which were more hardy in cold than popu-
lations living at lower and warmer elevations. Some of these
physiologically distinguishable neighboring populations were shown
to be genetically differentiated by incompatibility for reproduction.
J. A. Moore (1949) has shown that examples of Rana sylvatica from
northern ranges produce eggs which are viable in colder water than
can be endured by the eggs of southern populations. Some incom-
patibility in fertilization by parents from remote ranges also indi-
cates their genetic distinction. The differentiation of Rana sylvatica
has occurred in northern areas and is probably of recent origin. These
obviously adaptive and genetic variations distinguishing populations
show that adaptively significant variations sometimes become
established within species.

I consider it inescapable that natural selection will favor any adap-
tive variation according to its value to the possessor. If the variation
is an inheritable character it will proceed through the genetic mecha-
nism with a bias for its preservation in proportion to its adaptive
value. However, if we consider how rarely the small variations
within a species have been proved to be inheritable adaptations, we
must admit that there are few demonstrations of opportunities for
natural selection, per se, to operate in the differentiation of discrete
populations within a species. Some other factor or agency must then
be sought for the differentiation of the numerous taxonomic races
which are found.

The variations shown in the northern races of Alaskan birds have
no obvious adaptive value. Nevertheless, they clearly mark off geo-
graphic populations readily distinguishable from other populations
of the same species elsewhere. The distinctions, whether of large or
small degree of difference, appear to be maintainable only by prev-
alent inbreeding among the members of each race. Segregation of
a species into recognizably distinct populations cannot take place
without a sorting process, by means of which the homogeneous species
becomes fragmented into races. Rather than surmise that some in-
visible adaptation in form or in function favors the orderly segre-
gation of populations within a species, it seems preferable to seek
out the conspicuous behavioral activities of birds that can keep these
groups apart and intact as reproductive segments of the whole. This

982 U. S. NATIONAL MUSEUM BULLETIN 217

leads us to consider the possibility that the species involved, by virtue
of their behavior, are their own active isolating agent rather than
that they are the more or less passive recipients of molding influences
of external factors operating on undiscernible variational values.

We have seen that in neither resident nor migratory species of arctic
birds are the taxonomic distinctions of their races adaptive. Natural
selection cannot operate directly upon nonadaptive characters, but
yet in 9 resident species and 14 migratory species, the populations nest-
ing mainly in northern Alaska and Yukon have become differentiated
to the point where they are taxonomically distinguishable as races.
It would appear that some influence other than natural selection must
operate to bring about the segregation of their special although non-
adaptive variations from the common attributes of the species.

Small inheritable variations mark individuals of a species, and any
sample of individuals contains a genetic assortment which is some-
what different from the whole species. The smaller the sample of a
population, the less complete is its representation of the genetic com-
plement of the species and the greater is the probability that the genes
will vary during their hereditary transmission. If a population is re-
productively isolated it may well be different in its total genetic
range from the remainder of the species and that difference may well
increase with reproduction in isolation. These conditions for varia-
tion of a population are fulfilled by the isolation of the breeding
populations which are restricted to northern Alaska and Yukon.
The breeding isolation conforms with geographic limits, but for strong
and mobile birds like ptarmigan the restriction by inert geographic
obstacles appears to be not so much a factor as their active coherence
in a population that selects its own range and associations.

Birds of many species return to the nesting places from which they
have been experimentally removed, and they recognize their own nests
and their young even after only brief acquaintance. Birds of some
species persistently return in spring to nest in the locality where they
were raised, evidently attracted by some pattern imprinted on their
memories by early experience in the nesting locality. This particular
impression upon memory appears not to be inheritable and is acquired
by each individual.

Some races breeding mainly in Alaska and Yukon migrate to remote
wintering grounds different from those of other races of the same
species. Western solitary sandpipers (Tringa solitaria cinnamomea)
winter mainly south of the equator in South America, while the east-
ern race (7'. s. solitaria) winters from Texas and Louisiana south to
the equator. Near the time of autumn departure and spring return
the appearance of organized flocks of some species on their nesting

MIGRATION AND ORIGINS 283

grounds shows that there is some association of individuals of the
breeding populations apart from the time of nesting. Common ex-
perience and some association of members within each arctic popula-
tion while away from nesting grounds is also indicated by their
synchronous return in spring with all individuals in the same phase
of the brief arctic nesting cycle. Since the schedule of a bird’s physio-
logical preparation for breeding and migration can be experimentally
modified by changing the duration of daylight, the physiological
synchronization is not a genetically fixable character and it can arise
only from similarity in the physical and social environment en-
countered while away from the nesting grounds.

Birds migrating to the Arctic are similarly oriented in time and
space. It would be difficult to imagine the whole mechanism of in-
dividual behavior, physiology, and social activity which operates to
keep these migratory populations on the same coordinates of time and
space, but an acquired recollection of locality seems to be the essential
instrument of their isolation. Memory is the guide which associates
individual birds in populations along their long migratory paths
throughout the year. The strength of its influence for return to the
nesting grounds at the right time is especially impressive in relation
to the small size of some species. Kennicott’s willow warblers, for
example, weighing only 10 grams, remain a distinct population within
their wide ranging species while annually moving swiftly over a large
and geographically complex part of America and Asia.

The results of this behavior permit the migratory races to exploit
an arctic area which is only suitable for a brief season. The establish-
ment of such migratory races shows that their migratory behavior has
met the test of natural selection for value. The essential instrument
for their segregation is the memory pattern which acts as if to direct
energy of avian activity toward isolating racial characters from the
random mixture of the whole species.

Many maritime species (table 10) nest in arctic Alaska and Yukon
without having formed recognizable races. Although the area has
long been occupied by these species, they have not become differ-
entiated into taxonomically discernible entities. Occupation of the
arctic nesting grounds must be useful to the species, or birds would
not have become established as migrants to that area, However,
either the area is not an isolated one as far as they are concerned, or
each individual bird’s association with one locality is not maintained
consistently enough to segregate the requisite proportion of breeders
to allow for their eventual differentiation. It cannot be assumed that
those birds lack sufficient homing ability to maintain group solidarity,
as this ability is known from experiments in which several species

284. U. S. NATIONAL MUSEUM BULLETIN 217

of gulls and terns returned to their nests after being transported long
distances away. It may be suggested that these maritime, locally
undifferentiated birds are not persistently affected by companionship
with nesting associates during the nonbreeding season away from their
breeding grounds, and that they may be diverted to join individuals
bound for other nesting areas sufficiently often to keep the character-
istics of the entire species in a mixed state.

Outside of Alaska and Yukon, examples of long and specialized
migration routes can be seen to be associated with taxonomically dis-
tinguished populations. Although maritime species have not formed
races confined to nesting in the northern interior of Alaska and
Yukon, R. C. Murphy (1936) has described many examples of popula-
tions of oceanic birds which regularly return to nest on small South
American Islands after remarkable long oceanic migrations. Thayer’s
gull (Larus argentatus thayert) winters with other herring gulls on
the coast of British Columbia and migrates across arctic America to
nest in its eastern part. The physical characters of the race are only
significant for taxonomic purposes. It shares the strength and hardi-
ness of other herring gulls but behavioristic association with a locality
isolates the race on an unusual migratory path.

There are many variations in form among races of some species
which are not visibly adaptive and which cannot reasonably be as-
sumed to have been established by natural selection. And yet with-
out some segregating influence these characters could not be placed
in separate order from the common characters of the species. Isola-
tion, which in birds is often visibly governed by an acquired pattern
of geographic memory, can segregate breeding populations and thus
provide conditions suitable for variation. Under the protection of
isolating behavior small and nonadaptive differentiation could pro-
gress until it became significant for natural selection to perpetuate or
destroy. Isolating behavior could protect differentiation until even
completely nonadaptive characters became so distinct that the variant
population was genetically separated from its former associates in
the species. Small variations are common and nonadaptive racial
distinctions arenumerous. Isolation by behavior seems to be a process
which preserves these adaptively insignificant variations until they
distinguish populations.

During great biotic changes in northern lands, and in fact since
Pleistocene times, the surviving species of birds have remained essen-
tially unaltered (Wetmore, 1931, p.5). Although they have probably
moved their ranges northward and southward to escape some drastic
climatic changes, the northern species have met the fiuctuating con-
ditions of recent times without modification of their specific char-

MIGRATION AND ORIGINS 285

acters. The versatility of specific physiological processes and be-
havior is made impressively apparent by this stability. Thus, al-
though it is often proposed that the occurrence of taxonomic races
shows an evolutionary process by which species have been formed, there
is no convincing evidence that the races of birds form steps toward the
evolutionary divergence of species. We can say that natural species
formation has not been evident during the enormous environmental
fluctuations of the last 100,000 years. It is amazing that the species
were physiologically so versatile as to meet these changes without es-
sential modification.

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Lesser Scaup AND WuHITE-WINcED Scoter, Aythya affinis (see pp. 40, 135, 173) and
Melanitta deglandi deglandi (see pp. 43, 135, 176).

7. Residence in the Arctic

we OUTSIDER ENGAGED in arctic exploration and exploitation usual-
ly follows a program quite different from the native way of life.
Travel and construction by nonindigenous methods take place mainly
during the few months of open water and unfrozen ground. A
year’s labor must often be completed within a few months. Haste
drives the white man through the arctic summer and he is inclined
to think that natural processes move by his schedule. For the natural
arctic resident, however, life proceeds at a normal] rate in each season.

Annual changes in the arctic temperature are large and the progress
of related natural phenomena is pronounced because of the con-
spicuous character of seasonal transitions. It would be easy to believe
that the intensity of seasonal change requires acceleration of those
biological processes which must run through their cycle in the arctic
summer, for in three months or less birds must terminate their migra-
tory flight, mate, nest, rear young, and prepare their departure south-
ward. How these important periods are compressed within the short
duration of the birds’ summer residence in the Arctic is the subject
ofthischapter. _

287

288 U. S. NATIONAL MUSEUM BULLETIN 217

APRIL 30 MAY 10 2,0 31JUNE 10 20
Gavia immer { 1 Co)
Gavia adamsii 3
Gavia arctica pacifica 9
Gavia stellata 3 Ce)
Olor columbianus 4
Branta canadensis taverneri 03 19
Branta nigricans O12
Anser albifrons frontalis
Chen hyperborea hyperborea 30 9
Anas platyrhynchos platyrhynchos
Anas acuta ! (2)
Anas carolinensis 1

t]
Mareca americana 3 9 9

—
°
7]
)
i)

Spatula clypeata
Aythya marila nearctica 6| 391 | eS SS
Aythya affinis 9

Clangula hyemalis 1 |o3 31 fae Ore ay oS
Histrionicus histrionicus 2098

Melanitta deglandi 4 ‘| N
Melanitta perspicillata 1
Mergus serrator serrator 3 \19 2 yo ee Ee See
Buteo lagopus 9 862 0 |

Aquila chrysaetos canadensis APR 7,48 WAR 29,'49 APR 3.60 APR 10,51 MAR 23,62 af 4,53

Circus cyaneus hudsonius 3 8 nl

Falco peregrinus anatum 6 I
Falco columbarius bendirei W2
Grus canadensis canadensis 3
Charadrius semipalmatus 1 q 3 N==2 >
Pluvialis dominica dominica 1qe 9 32 4 SSS=
Squatarola squatarola ; 9
Arenaria interpres interpres 9 os
Capella gallinago delicata 30! 2
Numenius phacopus hudsonicus 3 199 0 2
Tringa solitaria cinnamomea 9 |
Heteroscelus incanum o|/3 91 2
Totanus flavipes o219) 3 aS —
Erolia melanotos BI 02
Erolia bairdii 913 2 ON I ==

N — —

Erolia minutilla =~ 1 3 20
Erolia alpina pacifica 9
Limnodromus scolopaceus 31 ‘ 20
Ereunetes pusillus 1

Tryngites subruficollis 9 co)
Limosa lapponica 3 oO
Crocethia alba 2 9
Phalaropus fulicarius 13 08
Lobipes lobatus 123 O/+,_-9 Nee ei
Stercorarius pomarinus fC)
Stercorarius pacar 1S3 2 = |

Stercorarius longicaudus () $3261
Larus hyperboreus barrovianus 31 Qa 9
Larus canus brachyrhynchus i 9 03 |
Sterna paradisaca 3 10 298-
Asio flammeus flammeus ee ) 29
Sayornis saya yukonénsis Nr
Eremophila alpestris arcticola 3 ! 8 9 Ne esas
Lanius excubitor invictus 31 290 hs =
Wilsonia pusilla pileolata ie 2

Euphagus carolinus carolinus 9021
Leucosticte tephrocotis irvingi 0
Acanthis hornemanni exilipes N---|-----|-
Acanihis flammea flammea Na----4-
Passerculus sandwichensis anthinus 9 1 N+--—

Spizella arborea-ochracea . 1 |o398 2 N---T-

Zonotrichia leucophrys gambelii 26 | 02 N--—

Passerella iliaca zaboria 16 9.2
Calcarius lapponicus alascensis 1|9 3 2 8 ===

Calearius pictus 3 218 ‘ee=re5

Plectrophenax nivalis nivalis MAR 28,48 APR 3,50 APR3,'51 MAYD,'52 APRB,53

Ficure 10.—First arrivals of migrants at Anaktuvuk, 1948-1953. Number is the last
figure of the year, N marks the date when the first egg was found, and dashes mark the
period during which eggs were laid.

RESIDENCE IN THE ARCTIC 289

Arrival

The arrival of migrant birds in spring makes a great impression
upon arctic people. First a few early comers appear. Shortly
thereafter a wave of individuals, pairs, and flocks sweeps in to popu-
late the formerly lifeless tundra. Not only do the birds suddenly
enliven a formerly quiet scene but each species at once displays its
special pattern of living on its nesting ground. Separating from
social groups the birds mate and establish territories in which to pursue
domestic and family life. Their program is as orderly as the settle-
ment of a traveling nomadic community of people in a new village
site. The community separates into groups of individuals for whom
the family is the dominant association, whereas in traveling the
direction of the community prevailed.

The conspicuous similarity of basic social concerns in the lives of
birds and men is clearly an attraction for the interest of people who
live a simple life. The birds are graceful, alert, neat of plumage,
and competent in managing their own affairs. Even the jealousy
shown in claims upon mate and territory is redeemed by the faithful
support which bird parents give to their families. Such staunch
support of their families is appreciated by Eskimos whose lives are
bound up in the preservation of family associations.

The birds arrive in an arctic season changing at a rate which can-
not be imagined from experience confined to temperate regions.
Daylight becomes continuous in May. In that month, from levels
lower than those of the coldest part of winter in the temperate
regions, the arctic temperature rises to the warmth of summer. The
water in the environment changes its physical state from snow and
ice to liquid. But in any one year these day to day changes are not
orderly, for until early June pleasant warmth, snowfall, and bitter
cold can succeed each other within a few hours (see fig. 5). In these
vagaries of the arctic weather in spring the birds arrive, settle, and
proceed to nest with such regularity that their behavior sets the
steadiest schedule among the natural phenomena.

I have recorded in figure 10 the earliest birds seen in spring at
Anaktuvuk from 1948 through 1953. These records come from
Nunamiut observers, from specimens obtained in the course of their
hunting, and from my own observations. Tom Brower carefully
studied the birds arriving in migration in 1949, and John Krog re-
ported on the condition of migrant arrivals in May 1951. During
each of the 6 annual migrations under consideration I have been
present at least some of the time. In the open valley so favorable
for observation, and with the interested assistance of the keen and
understanding Nunamiut, I obtained records of migration that I be-
lieve are unusually complete. Adding to the effectiveness of Eskimo
observation was their pleasant social custom of spending long periods

290 U. S. NATIONAL MUSEUM BULLETIN 217

in conversation and discussion upon the events of natura] history.
In this way all observations made in the village were brought into
a, pool from which Simon Paneak and I could study in careful, critical
discussion the details and circumstances by which the migration was
marked.

Often numerous individuals of the commoner species are seen on
the same day in spring when they are first reported. Within a few
days the presence of the common species is familiar to all the
people. Thereafter new arrivals of the species may be distinguishable.
The early arriving snow buntings are followed by successive flocks
which continue to move northward for as long as 6 or 7 weeks. The
early eagles all arrive within a period of 2 weeks. Golden plover
continue to gain new migrant arrivals for about 2 weeks, but Baird’s
sandpipers seem to arrive during a shorter time, as do robins,
Gambel’s sparrows, and Smith’s longspurs. The arrival of migrants
appeared to be concentrated in the latter part of the season.

If the appearance of migration is compared with a moving wave,
its crest occurs several days after its initial appearance. The length
of the waves certainly differs among the species by as much as from
1 to 6 weeks. But if the analogy of migration to a wave is pursued,
the initial appearance is a characteristic presaging the birds which
follow. The first birds which arrive are seen as individuals or as
flocks which are fewer than the entire migration. Since the birds of
a population are coherently related, their average behavior and con-
dition indicate the normal characteristics of their coherent action.
On the other hand certain occasional behavior is seen which is incon-
sistently related to migration. These observations of aberrant be-
havior cannot be used in describing the progress of migration.

Migratory birds arriving at Anaktuvuk either settle there to nest
or they proceed northward as far as the arctic coast. There is no
likelihood that significant numbers pass eastward or westward beyond
arctic Alaska. No Alaskan land lies more than 200 miles north of
Anaktuvuk, and the geographical center of the arctic slope north
of the Brooks Range is less than 100 miles distant. The location of
Anaktuvuk is accordingly within a few hours of the limit of north-
ward flight overland in this sector and thus is near or at the terminus
of migration. We may say that, practically, the locality is the northern
terminus and that the condition of the migrant birds represents their
state upon arrival from a long migration to an arctic region. Con-
sidering the rapidity of change in behavioral and physiological proc-
esses in birds about to nest, there will be some loss of physiological
detail in considering all these migrants through Anaktuvuk alike as
birds that have reached their breeding grounds, yet they indicate
- better than do those at lower latitudes the general condition at the
terminus of migration.

RESIDENCE IN THE ARCTIC 291

Distance Traveled and Arrival Date

In studying the arriving populations it must be considered that they
come from such widely distributed wintering grounds as are shown
in table 12. Among the migratory birds, only hoary redpolls are
known also to be resident in winter at Anaktuvuk, and few are seen
then in comparison with the large numbers present late in May. Dur-
ing that month the numbers progressively increase, but there is no
conspicuous climax in numbers which accurately designates a mean
date of migration. Nevertheless, the numbers seen in late May exceed
those seen in winter many hundred times. The many redpolls seen in
summer are still not a hundredth of those seen in May or again in
late August. In winter hoary redpolls are sporadically common or
occasional in interior Alaska. There are not enough indications to
suggest whether the arctic nesting hoary redpolls come from winter
resorts in nearby Alaska or from their winter range, which extends to
northern United States.

Common redpolls have not been found at Anaktuvuk in winter and
their nearest reported wintering area is in the Tanana Valley (Cade

TABLE 12.—Least migratory path of birds arriving on central Alaskan arctic
nesting grounds.

[Anaktuvuk (location, lat. 68°10’ N., long. 151°46’ W.) is taken as representative of the northern destination.
The reference localities of the wintering range areas represent the nearest well known points now assignable
to the species designated.]

Ce see eS Se eee

Least migration distance

Genus or migration group and Nearest winter Winter residence
number of species range reference locality Lati- | Longi-
tude | tude | Miles
(de- (de-
grees) | grees)
Acanthis 2| Alaska
Plectrophenax 1 | Alaska Anchorage, Alaska 7 2W 450
American Pacific coast (table| 19 | Gulf of Alaska Seward, Alaska 8 2W 550
10)
West and Northwest American} 20 | Northwestern Seattle, Wash- 2| 29W 1800
(table 11a) States ington
East of Rocky Mts. (table1ib) | 12
(Hylocichla) Tastern base of Great Falls, Mon- 22 | 42W 2100
Rocky Mts. tana
(Passerella) Southeastern States | Atlanta, Georgia 35 | 67 W 3800
(Pluvialis) Argentina Buenos Aires, 104 | 93 W 8500
Argentina
North American (table 11c-f) 31 | Mountain States Great Falls, Mon- 22| 42W 2100
tana
Pacific 2 | Northwestern Seattle, Wash- 22 | 29W 1800
(Heteroscelus) States ington
(Arenaria) Aleutians Cold Bay, Alaska 14 12E 1100
Asiatic 3 | China Shanghai, China 38 95 E 5200
(Oenanthe)
(Phylloscopus)
(Motacilla)

TOTAL SPECIES 90

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292

U. S. NATIONAL MUSEUM BULLETIN 217

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RESIDENCE IN THE ARCTIC 293

1953) and in the Kobuk Valley (Grinnell 1900). They may winter a
little farther from their arctic nesting ground than the hoary redpolls.

Snow buntings winter in southern and western Alaska and are also
seen in winter in northeast British Columbia, about 1,200 miles from
Anaktuvuk. Inasmuch as Salomonsen (1950) considered the Alaskan
buntings to be distinguishable in color from the Canadian birds, I
regard the migrants to arctic Alaska as coming from wintering
grounds within that territory. The front of the northbound move-
ment of snow buntings appears to pause in the Koyukuk Valley and
then to proceed slowly northward. It reaches Anaktuvuk early in
April but does not usually reach Barrow until mid-April, apparently
covering the northernmost 200 miles in about 2 weeks. This rate is
slow in comparison with the progress of the later season migrants.
The slow progress in the Arctic of snow buntings is nevertheless regu-
lar from year to year at Anaktuvuk and Barrow. It is also interesting
to note that James Ross (Ross, 1835) reported sighting a snow bunt-
ing in upper Boothia on April 17 in 1831 and 1832. The position cor-
responds in latitude with Barrow and the date of arrival of snow bunt-
ings agrees in both places.

The remainder of the migrants (89 species) come from wintering
grounds which are from 500 to 8,500 miles distant. In view of the
remoteness of most of the wintering grounds, the use of Anaktuvuk
rather than Barrow or Umiat as a northern reference point cannot
much affect estimates of the distance traversed in migratory flights.
On the other hand, wintering grounds of many species are extensive.
For example, wandering tatlers and turnstones are reported in winter
on many Pacific islands and continental coasts. There is no present
way of knowing whether the migration of these species to Anaktuvuk
comes from the areas of their nearest reported occurrence in winter,
which for the tatlers are coasts of northwestern States and for the
turnstones the Aleutians.

It is not intended to imply that arctic Alaska is a unique focus of
migration. It is, however, a point where observations show the arrival
of migrant birds at their breeding grounds after traversing many
different routes. Some of these routes cross many degrees of latitude
and longitude with the related changes in climate and the variation
in the direction of migrating flight relative to the sun’s position.
Most of these flights cover more than 600 miles of land before reach-
ing their destination at Anaktuvuk. Even for bird migration this is
a long distance, and with few reservations the Anaktuvuk birds can
be regarded as long distance migrants.

By plotting the dates of first arrivals observed in each species
during 1948 to 1953 the most common arrival dates, as shown in
figure 11, is about May 20. Also about half the May records fall

294. U. S. NATIONAL MUSEUM BULLETIN 217

before and half after that date. Accordingly about May 20 is the
mean and median date for recorded first arrivals of all species. The
arrivals about this date are not arranged in a normal distribution
curve, but form one negatively skewed. For early season progress
the curve is long and concave, and after May 20 it is short and convex.
Since the peak of arrivals precedes the maximum of summer heat by
about two months the relations to maximum heat is not explanatory
for migration.

Influence of Seasonal Phenomena

The accelerating progress of environmental conditions in spring
and a factor resulting from contingencies of observation may both be
concerned in the skewed distribution of arrival dates. No bird can
be recorded before it arrives, and even with intensive search some de-
layed observations inevitably result, tending to skew the curve. The
separate plotting of the records for the years 1949-1953 in figure 12
shows the component years of the record. The median dates of
distribution of arrivals in those years occurred on May 20 in 1948,
1949, 1950, 1951; on May 24 in 1952; and on May 18 in 19538. Observa-
tions of arrival were deliberately relaxed in 1952 for the interval
from May 21 until May 26 in order to give attention to other studies.
On May 29 they were resumed intensively, and it may be suspected
that the large block of records in the last of the month resulted from
delayed observation. The low temperature and late snow melting
in May of that year will be discussed later. It was also the year
between 1949 and 1953 when fewest observations were made, and the
abridged observation makes it doubtful whether the resulting records
well represents the dates of first arrival. With these reservations
about 1952, the close correspondence of the median of dates of recorded
first arrivals in each year suggests that the composite curve presents
a summation of substantially similar annual migration records.

It might be suspected that the records for those species most
consistently observed would contain fewer observational errors than
for birds which could be less regularly observed. In order to examine
the data by this criterion I have separated in figure 11 the first re-
- corded arrival dates of species for which 8 or more annual records
were obtained. The median date for this group falls on May 19.
The small advance is consistent with the view that the most evident
birds are reported a little more promptly than those which are less
conspicuous. Altogether these views of composite and selected records
suggest that the homogeneity of the data results from regularity in
bird behavior and has been little modified by erroneous observation.
Because of the skewed characteristic of the curves it is likely that

295

THE ARCTIC

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469496—60——20

296 U. S. NATIONAL MUSEUM BULLETIN 217

delayed observation adds a little lateness and consequently a slight
increase to the appearance of regularity in the plotted median dates.

Inspection of the arrival records indicates that species arrive at
various dates. In many cases the dates of first record of a species are
close together in succeeding years. I have shown in figure 13 the
number of days separating first recorded arrivals of a species in 3
years. It did not seem wise to assess commonness of occurrence or
our familiarity with a bird as a basis for selecting the most reliable

io)

SPEGIES
a

DAYS 5
SPAN OF FIRST RECORDS IN
3 YEARS. 47 SPEGIES

SPECIES
a

DAYS
SPAN OF FIRST RECORDS IN
4 YEARS. 37 SPECIES

Ficure 13.—Top: Span of first arrivals in the three closest annual arrivals among 4/ species.
Bottom: Span of first arrivals in the four closest annual arrivals among 37 species.

RESIDENCE IN THE ARCTIC 297

observations. All records of the shortest interval covering the ar-
rival reports of each species in any 8 years are therefore plotted to-
gether. They comprise 51 species and the range of first recorded ar-
rival dates extends from one to 11 days. The median range is between
3 and 4 days. Among 380 species the range of arrival dates did not
differ more than +2 days. Since there are few records from 1948, the
period comprised is principally from 1949-53.

The range of first-arrival records in 4 years, shown in figure 13
for 37 species, extends from 4 to 12 days. In 23 of the species 4 annual
arrivals were recorded within a period of 7 days. In a longer series
of years the arrival of the birds can vary more than in a shorter
period, and errors of observation likely to prolong the span of arrival
records can also enter more frequently. It is my belief that in the
4-year series, owing to the influence of erroneous observation, the
variations in date of arrival are more apparent than real, and that
the first birds of any one species will be found to arrive at close to the
same date each year. For example, during 3 years out of 5, with
1948 eliminated, 30 species were first seen within 2 days of the mean
date of the species and 45 species arrived within 5 days of the mean
date. As shown by the records of 37 species during 4 of the 5 years,
27 were first seen within 314 days of the mean date characteristic
of the species and all were reported within 6 days of their mean date
of arrival.

As a seasonal event, migration must be related to annual changes
in the sun’s heat. In weather records this seasonal variation is in-
dictated by measurements of air temperature. These are, by present
convention of the U. S. Weather Bureau, measured as daily maximum
and minimum, and their mean is designated daily mean temperature.
For a longer period of time, the daily mean temperatures are averaged
as the mean temperature of the period. C. EK. Watson and Robert F.
Dale, U. S. Weather Bureau climatologists, kindly made available to
me the mean temperature records for 3 Alaskan localities and for 3
localities in northern states representative of western, Rocky Mountain
and east-central areas through which the northbound migration of
American birds must pass. I have plotted in figure 14 winter to sum-
mer progress of the monthly mean temperatures in these 8 localities.
It shows that in the Arctic the change of air temperature in spring
is pronounced, and that in general both rate and amount of rise of
spring temperature increases in higher latitudes, while the seasonal
change diminishes southward and is imperceptible in the low tropics.

A sequence of official records of mean temperatures is not yet avail-
able at Anaktuvuk, for only in July 1953 was an observaiton station
established there by the Weather Bureau. The temperatures shown
in figure 5, from Simon Paneak’s journal records and from our oc-

298 U. S. NATIONAL MUSEUM BULLETIN 217

casional comparisons, indicate that winter temperatures in the nar-
row mountain valley at Anaktuvuk do not reach the extreme lows
measured in the broad valleys at lower elevations at Umiat, Bettles,
and Fairbanks. It appears that the mean monthly temperatures in
spring at Anaktuvuk lie between those of Umiat and Bettles. Be-
tween March and June the mean temperature at Anaktuvuk rises
from about —20° C. to about +10° C.

204

TLE
5s

| GREAT FALLS
; ©

| MINNEAPOLIS

uso FAIRBANKS wage
: sea

{ BETTLES

“JAN FEB. - MAR. APR MAY JUNE

Ficure 14.—Winter to summer progress of monthly mean temperatures in three Alaskan
localities and in three northern States.

During the days before and after May 20, when most migrant bird
species first reach Anaktuvuk, the mean temperature of the air is
just above freezing. Migratory birds can be regarded as land birds
at Anaktuvuk in the sense that they feed on snow-free ground or in
shallow pools and waters free of ice. This requirement of unfrozen
feeding ground is probably common for most of the birds migrating

RESIDENCE IN THE ARCTIC 299

into arctic and boreal regions. It does not limit the advent of birds
predatory upon mammals or feeding upon carrion, seeds, and insects
in exposed vegetation. Eagles, rough-legged hawks, snow buntings,
and longspurs precede the appearance of water and snow-free ground.
But for most species of migrants water and unfrozen ground seem
to be the main feeding places.

Although brought about by solar heat, the breakup of ice on large
rivers is also infiuenced by the thickness of winter ice and snow and
by the amount of water impounded at the time of the autumn freeze-
up. The date of the breakup of the Koyukuk River near Bettles
has been observed at various dates from early to late May. In 1949
and 1952 small airplanes could land on wheels on the ice of lakes
at Anaktuvuk until mid-June. In 1951 the ice was insecure in late
May and Pilot J. L. Anderson landed with us on floats before the
end of May. The complete breakup of ice is too variable and ill-
defined to use in dating a phenomenon which appears with such
regularity as bird migration. These events are all related to the in-
creasing heat of the sun, but not by obvious, simple parameters.

Heavy arctic snow cover in May would appear to be a handicap
to birds migrating from the south. In 1952 snow was unusually deep
over the mountain valleys and along the Koyukuk until the end of
May. In 1950 and 1953 most of the ground was bare early in that
month. The dates when the ground is mostly free from snow may
differ by 3 or 4 weeks in successive years.

In May the arctic land vegetation shows little change that could
be significant for birds. The first pollen appears on willows at the
end of the month, when a few bumblebees are seen. Early flowers
of sedge serve ptarmigan and caribou but seem to afford no sustenance
for most migratory birds. In short, the spring outburst of land
vegetation is considerably preceded by the advent of the birds.
Farther south at Bettles and Fairbanks the emergence of birch leaves
is a conspicuous and rather sudden event. I know of no published
records, but my memory of the events of spring and examination of
the recollections of old observers indicates that the emergence of
birch leaves in interior Alaska may differ by as much as two weeks
from one year to the next. It commonly follows the arrival of most
migratory birds.

The spring progress of arctic and subarctic vegetation is scarcely
visible when the main migration occurs. I have not noticed many
seeds of flowering plants at any time in the Arctic and I suspect that
they are not abundant in spring. During May flying insects are
rare and few can be discovered moving on the ground. Terrestrial
arthropods could have been little influenced to new growth in the
arctic spring when the birds arrive and those serving as food for birds

300 U. S. NATIONAL MUSEUM BULLETIN 217

must have survived the winter. But an important change in the
environment of arctic aquatic life has been remarked by John Krog
(1954) to occur some time before the waters are ice free. Ice is
impermeable to air and in the underlying pools of water which remain
unfrozen during the northern winter the oxygen supply becomes
much depleted because of the thickness of the covering ice. In late
winter the added pressure of water trickling under the ice from
melting snow begins to crack the ice on streams and cause the “over-
flows” which are so dangerous to travelers. As the season progresses
more of this circulating water is returned below the ice by the pressure
changes resulting from heat and barometric variations. As this
circulation increases, aerated water is mixed with the oxygen-deficient
sluggish streams and pools beneath the ice. The restoration of the
aquatic oxygen supply begins some time before the breakup of ice
occurs and before the temperature of the water changes appreciably.
It appears that the metabolism of arctic aquatic plants and animals
starts a spring revival as soon as the first localized melting starts the
circulation and reoxygenation of the waters. Thus, although the
formal breakup of ice is often later than the principal arrival time
of the birds, the beginning aeration of the water is earlier. I suspect
that this process initiates an efflorescence of aquatic growth some time
before the ice breaks and starts production of a new crop of aquatic
food in time for the arriving birds, for while watching beaver trap-
pers cut holes through several feet of ice in April, I have found the
buds of lilies pressed against the bottom of the ice. A few days after
the ice melts from Alaskan lakes the lily buds rise above the water,
having grown four or five feet from the bottom before an appreciable
change in water temperature takes place.

That the curves for mean temperatures during the time of migra-
tion vary greatly from year to year can be seen in figure 15. At
both Umiat and at Bettles the mean temperatures of May differed
by as much as 6° to 8° C. between 1948 and 1953. The cold mean
of —9.1° C. at Umiat in May 1952 was but little warmer than the
mean of —12.5° C. there in April 1948. In a longer series of arctic
records the mean temperature of May can vary +5° C. If birds
guided their arctic migration upon a fixed mean temperature, in some
years they would have to vary the date of their arrival by several
weeks, whereas the annual variation in arrival of a species at the
arctic terminus of their flight is not more than a few days.

While daily temperatures in May are not so variable as in April,
between May 15 and 20 temperatures as low as —16° and as high as
18° C. were recorded at Anaktuvuk in 1951 (fig. 5). Wide spring-
time fluctuations in temperature are so frequent in the Arctic that
birds must commonly encounter days of more severe cold there than

RESIDENCE IN THB ARCTIC 301

1948 1949 1950 1951 1952 1953

ce)

FAIRBANKS ce

BETTLESO

UMIAT

Ficure 15.—Mean temperatures in March, April, and May at Fairbanks, Bettles, and
Umiat, 1948-1953. (Compiled from climatological data.)

302 U. S. NATIONAL MUSEUM BULLETIN 217

most of the species have met at any time on their wintering grounds.
We have no measure of the casualties which result from this exposure
to cold, but by their long existence it is evident that the birds regularly
endure these wide variations in temperature. The tolerance for cold
exhibited by such small birds as sandpipers is remarkable but it
must nevertheless be regarded as a normal ability, for it is exercised
every year.

Nesting

The social behavior and family relations of birds are elaborately
developed and their importance for the existence of races is more con-
Spicuous than is generally the case among other animals, which can
detect others of their kind through scent and in darkness. Such ani-
mals may thus remain strange to us because of our lack of suitable
perception, whereas among birds social and family coherence is main-
tained by visible and audible actions. These are as apparent to us as
they are to the birds for which these signals are produced, and in many
cases their significance is in a general way understandable to us,
Without claiming full comprehension of the demonstrations by which
birds maintain coherent flocks and families we can often recognize the
general purport of their activity ata given stage near breeding,

Duration of Migratory Behavior

The latest date when the birds of a species nesting in the area have
arrived can be determined only by circumstantial evidence, but because
of their expressive behavior it is generally easy to distinguish in even
a single bird whether it is migrating or has settled down to nest. Mi-
gratory flocks disintegrate before the pairs nest. In any single season
the record of the first migrating bird observed may be a day or so after
the first arrival of the species on migration. To decide from journal
notes when the migration has ended is not soeasy. While there is thus
uncertainty in setting the end of migration as a normal date, it-appears
to be fairly regular, and in most years probably few birds of a species
arrive after the dates registered for the last migrants in table 13. I
believe, however, that favorable or unfavorable weather may shorten
or prolong migration in any year, whereas the first arrival does not
appear to vary in date.

The duration_of the migratory arrival-period among 28 species
varies from 2 to 60 days.- The 2 days attributed to the-wagtails is
probably shorter than the actual period, which is nevertheless brief.
The long period of arrival shown for snow buntings is substantially
correct, for none remain to nest at Anaktuvuk. Since these early mi-
grants first appear at Barrow about April 15 they also have tima there

RESIDENCE IN THE ARCTIC 303

TABLE 13.—Duration of phases of migratory and reproductive cycles in 28 species
of birds at Anaktuvuk

Arrival date Duration Days Duration
of arrival | First egg |from mid-| Latest | of depar-
Species period date arrival |departure} ture
First Last (days) to mid- date period
migrant | migrant laying (days)
Plectrophenax nivalis nivalis Apr. 1] June 1 60 j9June 1 34 |¢Sept. 10
Aquils chrysaetoes canadensis | Apr. 3] Apr. 18 15
Lanius excubitor invictus May 4] May 16 12 | May 27 23 | Oct. 15
Calcarius lapponicus alascensis | May 6] June 7 32 | May 31 14 | Sept. 2 20
Buteo lagopus May 8 | May 23 15
Eremophila alpestris arcticola | May 10| June 1 20 | May 29 13 | Aug. 24 19
Anthus spinoletta rubescens May 11] June 6 26} June 1 12 | Sept. 3 30
Anas acuta May 13] June 6 24 | May 23 4 |>*Sept. 10 10
Spizella arborea ochracea May 13} May 30 17 | June 3 17 | Sept. 12
Pluvialis dominica dominica May 14] May 31 17 | May 28 10 | Aug. 15 15
Turdus migratorius migrato- | May 15 | May 30 15 | May 30 12 | Sept. 18 13
rius.
Zonotrichia leucophrys gam- | May 16 | May 29 13 | June 3 16 | Sept. 12 12
belii
Charadrius semipalmatus May 16] June 5 20 | June 12? 21 | Aug. 16 15
Anas carolinensis May 18] June 6 14 | May 27 6 }bAug. 30 10
Totanus flavipes May 19] May 27 8 | June 1 13 | Aug. 6
Erolia bairdii May 19 | May 31 12 | May 29 8 |>Aug. 10
Aythya marila nearctica May 20] May 26 6| June 5 17 |+Sept. 10
Clangula hyemalis May 20 | May 28 8| June 4 15 |*Sept. 10 10
Lobipes lobatus May 22] May 29 7| June 6 16 |bAug. 25 10
Ereunetes pusillus May 23} June 1 9
Mergus serrator serrator May 23} June 3 ji | June 9 17 |*Sept. 10 ll
Oenanthe oenanthe oenanthe May 23] May 27 4} June 5 14 |>Sept. 1 20
Erolia minutilla May 24] June 5 12 | June 3 8
Stercorarius parasiticus May 24] June 5 12} June 8 13 |>Sept. 10
Melanitta deglandi May 25] June 14 21 | June 6 9 |bAug. 30 10
Calcarius pictus May 26] June 3 8| June 4 12 | Aug. 24 10
Sterna paradisaea May 28} June 6 9 | May 81 4 |bAug. 30 10
Motacilla flava tschutschensis | June 3] June 5 2 | June 10 10
TOTAL (28) (aver.) (14)

« Estimated from observations at Barrow.
> Estimated from growth of young.

sufficient for a prolonged arrival from migration before their nesting
begins in early June. At Anaktuvuk the equally early eagles (Aquzla
chrysaetos canadensis) and somewhat later shrikes and rough-legged
hawks arrive in a wave lasting only a quarter as long. Such early and
plentiful birds as pintails, Alaska longspurs (Calcarius lapponicus
alascensis), horned larks, and pipits have also a rather prolonged
migration. Species which are numerous are not necessarily diverse
in behavior, but the populations of those species which are numerous
at Anaktuvuk are not so closely synchronized in migration as those
less numerous.

Snow buntings winter in much of Alaska south of the Alaska range
from Unalaska and Nushagak to Sitka, along the lower Yukon and
north of the arctic circle at Kobuk, and also east of the Canadian

304 U. S. NATIONAL MUSEUM BULLETIN 217

Rocky Mountains in an area separated from Alaska by extensive and
high mountains. I suspect that those nesting in Alaska winter there,
but since its winter climate is very diverse and its terrain is separated
by massive mountains into regions with distinct conditions, the origin
of migrants in such diverse regions can be a cause of variation in their
migratory schedules.

Urner’s (Urner and Storer, 1949) long records of the migratory
flights of plovers and sandpipers along the coast of New Jersey show
the duration of migration in that locality. From their reported rec-
ords between 1928 and 1938, I have extracted the dates during which
were observed principal migratory flights of some species also familiar
at Anaktuvuk.

Duration in days

Inclusive dates of flights in New Anak-

New Jersey Jersey tuvuk
Charadrius semipalmata May 4-June 4 31 20
Totanus flavipes April 24-June 13 50 8
Erolia minutilla April 29-May 30 31 12
Ereunetes pusillus May 7—June 7 31 9

It is apparent that in New Jersey the birds of these species are
migrating toward nesting in localities variously distant northward,
and that the samples are from populations known to be heterogeneous
in regard to nesting area. The shorter migration waves at Anaktuvuk
are produced by populations which are alike in nesting within Arctic
Alaska. Heterogeneity of migratory populations in regard to either
nesting or wintering areas may prolong the migratory passage over
any locality. In fact it seems likely that diversity of either back-
ground or future events might reduce the synchrony of migratory
action.

Regardless of the explanation for the diversity in the duration of
their migratory flights, it is apparent that the birds of no single
species arrive synchronously at Anaktuvuk. Let us consider the im-
plications in the lack of synchrony of Baird’s sandpipers which con-
tinue to arrive for about 12 days. If they had traveled at the rate
of 60 miles a day during the 12 days preceding their arrival the popu-
lation would have been spread out along a course 720 miles long.
It is immaterial how closely this assumption fits the progress of migra-
tion in its northern course, for in relation to the range of sight of birds
the migrating elements of either flocks or individuals would be scat-
tered beyond range of communication with the remainder of the
population most of the time. I cannot imagine by what sense small
migrant birds could communicate in order to bring about coherent
action when spread over the time and distance in which each species
appears to be distributed during migration. In terms of proximity
of individuals relative to the range of sensory communication, the

RESIDENCE IN THE ARCTIC 305

migratory process is not concerted but widely distributed. It seems
as if individuals and groups, but not the populations as units, guide
the migration.

Progress Toward Laying

Judging from the size of testes, males of most species are ready for
breeding at the end of migration, although Alaska longspurs with
small testes were taken during the first 10 days in which migrants
were arriving. However, readiness for reproduction does not termi-
nate migration and start settled residence on the breeding ground,
for immature individuals of some species migrate to summer near
the breeding grounds.

Mallards, mergansers, golden plover, and birds of several other
species commonly appear on the nesting grounds in pairs. On the
other hand, some flights of arriving sandpipers are composed prin-
cipally of males, and the males of some passeriform species take up
nesting territories before females are seen. The variety of physio-
logical phases and behavior of arriving migrants is interesting, but
no common sequence can be derived from it to mark the conditions
which transform a migrating bird into a settled bird with all its in-
terest focused upon mating and preparations for subsequent nesting.
From my general impression of their behavior, I would have said that
pintails ducks and all earlier arriving migrants took the transition
from the migratory to breeding state in more leisurely fashion than
did later migrants, yet an examination of my records gives no sug-
gestion that the interval between arrival and laying is related to
earliness of arrival.

The recorded arrival period of migrants of nesting species lasted
from 2 to 32 days. I think that more migrants arrived to settle during
the first half of the period of each species than arrived later. Some
later migrants appeared in haste to move northward and those which
settled found less free territory. The period from midmigration to
the middle of the week after nesting begins is a fair measure, for
most of the species at Anaktuvuk, of the time elapsed between ar-
rival and laying. This elapsed time is estimated for 24 species in
table 18. The interval is 4 days for 2 species, 5 to 9 days for 4, 10 to
19 days for 16, and 20 to 23 days for 2 species. These estimates in-
dicate that the interval between arrival and laying differs among
species and that for a few species it is less than a week. In temperate
climates some early nesting pairs of many species lay as soon after
arrival as do the arctic birds. Mrs. Oakeson (1954) shows that at
Mountain Village, Alaska, for Gambel’s sparrow the interval between
the average arrival date and first laying was about 8 days. The earli-
est nesting birds of the race of white-crowned sparrows (Zonotrichia

306 U. S. NATIONAL MUSEUM BULLETIN 217

leucophrys pugetensis) at Friday Harbor, Washington, on the average
went through the same phases of the reproductive cycle about as
rapidly as the Alaskan birds. The individuals of the Puget Sound
race, however, were not so well concerted in phase of the reproductive
cycle as were the Alaskan birds.

At Barrow, Pitelka (1954) found that 80 to 90 percent of all
breeding Alaska longspurs and snow buntings began laying within
a single week in the first half of June. These birds pass Anaktuvuk
in a relatively prolonged migration extending, respectively, over
about 30 and 60 days. Both longspurs and snow buntings winter in
quite diverse areas, but at Barrow their reproductive activity is
synchronized to occur within a remarkably short time.

The simultaneous breeding of arctic populations is based upon
synchronization of the internal reproductive condition of individual
birds. As Mrs. Oakeson (1954) observed, individual arctic birds may
take a nesting area, mate, and copulate a little more quickly than in
a warmer climate. The sequence is a little expedited, but I suspect
that the shortening is produced by reduction in the time required for
transition from the end of one physiological eae to the beginning
of another.

At Old Crow, in Yukon, our observations were carried on through
only one season and the records in table 14 show the date between the
first individual seen of the species and the first egg observed. This
information is empirically determined, while the interval between
middle of arrival of the species and the middle of laying at Anaktuvuk
includes a factor of judgement estimated from a number of years’
observations. The species reported upon at Old Crow also differed,
including only 5 of those at Anaktuvuk. The average of the intervals
observed among 27 species at Old Crow was 17 days between first
arrival and first egg, whereas among 25 species at Anaktuvuk it was
14 days, indicating that the progress from migratory to laying con-
dition was at similar rates in these two arctic localities.

In each locality the progress toward laying varied among the
species. The longest interval at Old Crow was 31 days and at Anak-
tuvuk 34 days. The shortest observed interval at Old Crow was 7
days for the fox sparrow, and at Anaktuvuk 4 days for the pintail
and arctic tern, a period which seems likely to be too short. At both
places the progress of some species from migration to laying was
remarkably rapid and of others comparatively leisurely. The species
likewise differed in the rapidity of their transition from migration
to reproduction.

It is apparent that the speed of the transition varies among species
of ducks at Anaktuvuk and within the family of sandpipers in both
localities. Only in the family Fringillidae are the species much

RESIDENCE IN THE ARCTIC 307

TasieE 14.—Progress from migratory to laying condition in 27 species of birds

at Old Crow
Species First First | Interval
arrival egg (days)

Gavia arctica May 22 | June 15 24
Anas platyrhynchos May 8 June 5 28
Aythya affinis May 18 | June 18 31
Buteo lagopus Apr. 18 | May 5 17
Circus cyaneus May 5 May 20 15
Falco peregrinus May5 | Junel 27
Charadrius semipalmatus May 19 | June 7 19
Capella gallinago May 7 May 31 24
Actitis macularia May 22 | June 18 27
Tringa solitaria May 14 | May 25 11
Totanus flavipes May 7 May 28 21
Colaptes auratus May 16 | June 1 15
Petrochelidon pyrrhonota May 25 | June 9 15
Turdus migratorius May 8 May 25 17
Txoreus naevius May 4 May 21 17
Hylocichla minima May 22 | June 9 18
Vermivora celata May 20 | June3 12
Dendroica petechia May 23 | June ll 20
Dendroica striata ; May 27 | June 12 16
Seiurus noveboracensis May 20 | June 8 17
Wilsonia pusilla : May 20 | June 4 13
Euphagus carolinus May 10 | May 25 15
Acanthis flammea May 21 | May 30 9
Junco hyemalis May 15 | May 25 10
Spizella arborea May 16 | May 26 10
Zonotrichia leucophrys May 14 | May 26 12
Passerella iliaca May 14 | May 21 i
TOTAL (27) (Aver.) (17)

alike at Old Crow, but considering the long interval apparent in the
snow bunting at Barrow, there is variation in the rate of progress
toward breeding among the species in this family as well. It is
significant that the intervals in four of the five species common to
the two localities differ considerably, i. e., yellowlegs (Totanus
flavipes) 21 and 13 days, robin 17 and 12 days, tree sparrow 10 and
17 days, and Gambel’s sparrow 12 and 16 days. The best documented
very short interval of transition from migratory to laying conditions
is afforded by the 7 days recorded at Old Crow for fox sparrows.
There is undoubtedly a minimum time, probably differing among
species, in which the necessary physiological and behavioral proc-
esses can convert a migrating to a laying bird. Evidently external
circumstances can variously prolong the physiological process in an
individual, or at least prolong the physiological state of readiness
to lay. Accordingly, I conclude that the transition of a species from
migratory to breeding condition in two localities can be extended
beyond the minimum physiological requirement by amounts varying
according to local environmental conditions.

308 U. S. NATIONAL MUSEUM BULLETIN 217

The progression of physiological changes toward reproduction
seems to be punctuated by actions or stimulation which must come
from another bird (Tinbergen, 1953). Thus copulation, which may
occur at any time during a certain reproductive period, is necessary
to start the formation of fertile eggs. Thereafter, in the Arctic the
eggs of most birds are laid on successive days, and once the phase of
egg laying is initiated subsequent progress is invariable in all climates.
Such internal processes occur in homoiothermous birds under condi-
tions separate from external environmental influence and must proceed
at rates prescribed by ancient phylogeny. Certain types of behavior
which are the outward expressions of these internal processes must
be equally inevitable and unmodifiable by local and transitory
climates. But during the intervals between phases in the reproductive
cycle stimulation from another bird or even from the environment
may be quicker to set in motion the operations of the next phase.

Elaborate patterns of behavior concerned with possession of terri-
tory, courting, and mating are precisely followed by birds of each
species in sequences related to reproductive condition. Among indi-
viduals in the same phase of their reproductive cycle no scrutiny
is needed to determine that any bird of the same sex is physiologically
ready to compete for territory or mate. All birds of opposite sex are
disposed to the one phase of reproduction current in the local popula-
tion. In such a society no bird is likely to untimely action and one
of the common causes of social disorder is removed by the uniformity
of impulses in a population progressing synchronously through the
sequences of reproductive behavior. The duration of competition and
conflict in reproduction is shortened and actions decisive to the se-
quence of events in reproduction are prompt.

I do not imply, in this respect, that a natural society ordered by such
well synchronized impulses would be favored for existence in all
environments. But it appears to be by the ordering of society rather
than by acceleration of the individual’s physiological processes that,
adaptation of the breeding populations of birds conforms with the
shortness of the arctic season.

Incubation

Mrs. Nice (1954) has pointed out the frequency of erroneous reports
of the time during which eggs of wild birds are incubated. For a
clutch of eggs laid on successive days, it is difficult to set as a single
figure the duration of incubation. If incubation is considered to
begin only when the last egg has been laid its termination may differ
by as much as several days in the successive hatching of individual
eggs. The stage of embryonic development which is attained when

RESIDENCE IN THE ARCTIC 309

eggs of wild birds are hatched has not been well defined. Accordingly
the observed duration of incubation is not a factor in the velocity
or chronology of embryonic growth, but only in the behavioral process
of incubation.

Tf the conventional definition of the duration of incubation among
wild birds is taken to start when the last egg is laid and to extend
until the last egg hatches, rather consistent records are obtained for
each species. After examining the incubation of many species of
birds in equatorial Africa, the Moreaus (1940) remarked, “there is no
doubt that the incubation period is, within limits, specific... .” In
only a few species in tropical Africa did they find the incubation
period to be slightly longer than among their relatives in temperate
parts of Europe.

Lack (1948) commented upon the common similarity between the
duration of incubation and the time which nestling birds of altricial
habit spend in the nest before leaving. It is certainly remarkable
that these two periods are often numerically similar and appear as
time constants of species. If incubation and nestling periods are
constants of species, and probably of larger taxonomic groups, they
are not likely to be much influenced by climate. But to seek by
comparisons among species whether incubation or nestling periods
can adapt certain species to particular climates these periods should
be defined in terms of developmental processes or stages in growth.
Otherwise we may be comparing such different stages of growth as are
shown in extremes by the development of precocial and altricial birds.

- Once the egg is laid, incubation provides heat, so that the tempera-
ture maintained during incubation is the only visible parental in-
fluence upon embryonic development. Kendeigh and Baldwin (1928)
showed that the mean temperature of the eggs of house wrens
(Troglodytes aedon) in nests near Cleveland was about 35° C. In the
same summer climate Huggins (1941) found the mean temperature
during incubation of 37 species of 11 orders to be 34.0° C. Among
the passeriform species the mean temperature during incubation was
33.8° C. At Cleveland, Ohio, birds of diverse phylogeny adjusted
their incubating behavior to the weather so as to maintain a surpris-
ing similarity and constancy in the temperature of their eggs.

In summer at Anaktuvuk the temperature of the air is more variable
than at Cleveland. Freezing is occasional and temperatures are
mostly between 0° C. and 10° C. Among eggs in nests of birds of 7
species the median range of temperature was found to be between
33° C. and 35° C., with 76 percent of the records between 33° C. and
37° C. (L. Irving and J. Krog, 1956). We had thought that some
arctic nests appeared constructed for good insulation. But among
the eggs of a sandpiper (Zrewnetes pusillus) with no nest the tem-

310 U.S. NATIONAL MUSEUM BULLETIN 217

perature was like that in the well insulated nest of tree sparrows.
It is not by nest construction but by the amount of heat contributed
by the incubating parent that the temperature is kept in arctic nests
at the same level as in temperate regions. The extra heat dissipated
to cool arctic air could be described in physiological terms, but the
uniform temperature in the nest is regulated by the behavior of
parent birds.

This fact has long been known. In his famous essay Claude Ber-
nard (1876) referred to the even warm temperature maintained during
incubation as an example of regulation by behavior, and mentioned as
a remarkable illustration the Australian mound builder, which con-
structs a mound of earth and vegetation in which heat from fermenta-
tion supplements solar heat to maintain the correct temperature for
incubation. And a recently reported measurement by H. J. Frith
(1956) of the temperature around the eggs of the Australian mallee
fowl (Leipoa ocellata) shows it to be at the common level for avian
incubation. The birds work over the mounds daily to modify the heat
conductance of the material in accord with the varying supplies of
solar and fermentative heat.

Our temperature measurements among eggs of arctic birds were
not as well controlled as those made near laboratories in a temperate
climate, but the records appear no more variable than those observed
under better technical conditions by Kendeigh and Baldwm and by
Huggins. It thus seems that incubation probably does not in general
modify avian embryonic development through temperature, the only
course which we can see open. Having no measure of the progress of
development within the eggs I have not recorded the duration of
incubation. The progress of arctic reproduction suggests that-its
essential physiology is about the same as in temperate regions, and
on general grounds I doubt if embryonic development can be modi-
fied in race or local populations as a climatic adaptation.

Growth of Nestlings

After the eggs hatch the parent birds might influence growth of
the nestlings by the temperature of brooding and by the amount of
food provided. Karplus (1952), observing the short duration of
nestling life in a brood of robins at Umiat, related it to the prolon-
gation of feeding in the 20-hour daily active period of the arctic
parents. In Swedish Lapland (lat. 68° N.) Armstrong (1954)
remarked upon the prolonged daily parental feeding in connection
with the apparent shortening of the period of nestling growth.

There is some evidence for shortening of the duration of nestling
life from the tropics northward. Among 10 species of Central Amer-

RESIDENCE IN THE ARCTIC 311

ican Fringillidae, Skutch (1945) reported nestling life of from 12
to 15 days. Lack (1948) listed the duration of nestling life among
16 European species as being between 11 and 1414 days. Mrs. Oake-
son’s (1954) observations of three races of western white-crowned
sparrows nesting respectively at Berkeley (lat. 38° N.), Friday Har-
bor (lat. 49° N.), and Mountain Village (lat. 68° N.) are pertinent
indications that in the highest latitude the nestling period was shorter
in the relation 10.1, 9.6 and 9.0 days. In the duration of the nestling
stage at Anaktuvuk we found indications of about 10-percent shorten-
ing among several species of Fringillidae. This difference is not
related to the much greater prolongation of daily arctic feeding.
In fact, without some true measure of nestling development the dura-
tion of nestling life is only a time of parental and nestling behavior
and not a dimension of growth.

We measured the temperature of nestlings in seven nests of five
species at Anaktuvuk (L. Irving and J. Krog, 1956). Starting with
the incubation temperature between 33° C. and 35° C. at hatching
the temperature increased but did not quite reach the temperature of
adult birds. In general the temperature of the nestlings was rather
even, for 72 percent of the records fell between 36° C. and
42° C. and 86 percent between 34° C. and 42° C. While the mean
temperature of nestlings was cooler than that of adults, adult birds
may be 8° C. cooler at night than at rest by day and they may warm
8° more during activity (L. Irving, 1955). In arctic nestling
birds the variation in temperature little exceeds the changes in body
temperature which an adult usually passes through in each day.
The range of body temperature of the adult is regulated physiologi-
cally. The regulation of nestling body temperature is effected by
the parental behavior of brooding, although in older stages the nest-
lings contribute some metabolic heat and a certain amount of regula-
tion as they acquire insulation and control of heat production.

It has long been known that in hot climates parent birds shield their
young from intense heat of the sun (Wetmore, 1921). Bartholomew,
Dawson, and O’Neill (1953) showed that brooding behavior was
accurately regulated to protect nestlings in Baja California from
exceeding a normal body temperature.

The temperature requisite for growth of avian embryos or young
does not appear to be modifiable for adaptation to local climatic
conditions. Although the mean temperature during growth is lower
than that of the parents at rest, the variations to which the young
are exposed scarcely exceed the normal transition in adult body tem-
perature from sleep to intense activity; and although the young lack
control of their body temperature, it does not seem useful to regard

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312 U. S. NATIONAL MUSEUM BULLETIN 217

them as poikilothermous when they live with and probably require
about the same uniformity of temperature as their parents.

Since this sort of homoiothermism seems to be innate in the embryo,
which originates from cells In a warm parent, there appears little
opportunity for the environment to modify avian development through
the influence of temperature. At present this opinion is speculative
because it is based only upon observations of time and temperature
‘without support from measurements of differentiation or growth.

Among 28 species for which I can prepare schedules of migration
and nesting at Anaktuvuk the first eggs, as shown in table 14, were
found between May 23 and June 10. The latest fresh eggs found in
this latitude were in a robin’s nest on July 21 (L. Irving and Paneak,
1954). Late in June I have found a few pairs of redpolls beginning
to lay. The Nunamiut tell me that among the bare willows in winter
they occasionally find nests containing frozen emaciated nestling red-
polls. Starvation had preceded freezing. The Eskimo view is that
as August progresses redpoll parents become so anxious to join a flock
that they lose interest in family affairs. This good subjective descrip-
tion of the situation shows that it is not the cold weather which marks
the end of parental care but the onset of molt or other internal changes
which are represented in the social transition from family attachments
to the formation of flocks.

It seems clear that young and adult birds must prepare in summer
for the change of social regime which goes with the change of
season. In order to leave time enough for the late summer prepara-
tions for change of habit, residence, and season, arctic birds must pass
rapidly through the stages of the reproductive cycle occurring between
migration and nesting.

Departure

The departure of most species of birds from Anaktuvuk in late sum-
mer is not easy to record accurately. The termination of concern for
breeding first becomes obvious among some species of ducks, like
scaup and pintails, when the male birds begin to assemble in flocks in
mid-July. By the end of July the young of most of the passeriform
birds are flying. Family associations then relax and the birds of some
species begin to assemble in more or less well organized flocks. Other
species, like tree sparrows, withdraw from view and are infrequently
seen. Some conspicuous associations before migration southward can
be seen in the following examples:

RESIDENCE IN THE ARCTIC 313

Flocks compact Flocks coherent but not compact Southward movements loosely
associated but apparently coherent
Anas acuta (<7) Clangula hyemalis Eremophila alpestris
Aythya marila (07) Ereunetes pusillus Motacilla flava
Melanitta deglandi (7) Sterna paradisaea Anthus spinoletta
Turdus migratorius Acanthis hornemanni Calcarius lapponicus

Calcarius pictus

While they are molting the birds of several species become very
retiring and they are not seen again in their earlier numbers. Some
common species simply disappear without any visible preparatory as-
sociation. I know of no behavior by golden plover which signals
their departure, but the birds of most species give indications which
suggest that they are about to depart. By various signs as well as
from the last sight records fair estimates can be formed as to the end
of their migratory departure, as given in table 27. The duration of
the departure on migration can be estimated, but this is the most un-
certain observation in the birds’ program at Anaktuvuk.

The last sight records differ so much among the species that the
duration of summer occupation of the arctic tundra is clearly seen to
vary considerably. Plovers and sandpipers leave early. From my
recollection of migration near Barrow some of the sandpipers and
phalaropes appeared there as migrating flocks later than the last rec-
ords for those species at Anaktuvuk. The plover and sandpipers are
so inconspicuous in August at Anaktuvuk that near the ground level,
at least, no extensive southbound flights were seen to assemble there
or to pass through from northern arctic Alaska.

Numerous flocks of ducks, geese and a few flights of swans and
cranes are seen on their southbound course, but brant have not been
reported in the southbound flights. With that exception the south-
bound geese and ducks seem to be about as numerous as they are in
northbound flights. The flights seen moving southward do not alight
as often as they do in spring.

I have only one record of southward movement for the snow bunt-
ings. A flock was seen in the high mountains at the end of August.
A few others have been seen in the Valley in fall, but I do not know
the southbound route of the large numbers which fly northward in
spring. Our failure to record a date for the southbound eagles and
shrikes is not significant because there are not many of these birds.
A few rough-legged hawks are seen in late summer, which suggests
that their return occurs for the most part unnoticed along the spring
route.

314 U. S. NATIONAL MUSEUM BULLETIN 217

The variety of departure dates among the passeriform species is
conspicuous. Robins are the latest recorded, except for shrikes. The
latter, I suspect, occasionally winter in Alaska. My records fail to
show many late tree sparrows. Although they are occasionally seen
in September I have no idea when most of the large summer popula-
tion of these sparrows leaves Anaktuvuk. The early start of the
movement of wheatears and yellow wagtails is not surprising if we
regard the great distance of their migratory course to Asia, but a few
wheatears have been seen on the first of September. It cannot be said
that migrants to the nearer wintering grounds leave Anaktuvuk late
in the season, for the early arriving horned lark has its winter range
nearer than those of many species which depart at later date. There
is no clear general relation between duration of residence or date of
departure and remoteness of wintering ground.

The late birds remain in the arctic until ice and snow are likely
to cover their feeding places. At this season they encounter cold
weather, but it is not as severe as near the common date of arrival
in spring. The majority of the birds leave in August when the mean
temperature is above freezing and while even frost is of brief duration.
Evidently cold sets the latest termination of the stay of migratory
birds in the Arctic, but many birds leave just after the peak of sum-
mer’s heat and while the weather is still warm. As it appeared at
the time of arrival, the departure of migration is not in phase with
the temperature changes of the seasons.

Thus, it can be seen that the departure schedule of the various
species differs as to terminal date and as to approximate duration
of the process of departure. Although estimates of the duration of
departure are far from precise, there are evident large differences
among the species. In no case is the departure process short enough
to suggest that all individuals of a species act in concert. Considering
the homogeneity to be expected in populations breeding in the same
area, the duration of both arrival and departure seems rather long.
Neither part of the migratory process suggests that it is sufficiently
shortened at Anaktuvuk to indicate a strong, general, synchronizing
influence from the shortness of the arctic season.

: Gee

ny a aA

Guitemor Cepphus grylle mandtii (see p. 275).

8. Biological Aspects of Migration
and Nesting

W: WHO WALK 80 SLOWLY consider with amazement small birds such
as sandpipers, wheatears, Kennicott’s willow warblers, and yel-
low wagtails that fly half the distance between the poles and a nearly
equal distance eastward or westward to nest in the Arctic. Ac-
customed to warmer climates, people see in the short, cool arctic
summer a difficult time for the nesting of birds; and even the sym-
pathy of the Eskimos is evoked when the frequent cold weather of
May smites these small migratory birds, which appear so ill fitted to
encounter snow and cold. Yet, as we will find, the arctic season suits
their biological activities in a manner which dispels early impressions
that arctic life is passed in hardship.

I have discussed in chapters 6 and 7 the various courses of birds
migrating to the Alaskan arctic regions and the impressive magnitude
of the distances of their nesting from their wintering grounds. The

315

316 U. S. NATIONAL MUSEUM BULLETIN 217

19 migratory species nesting principally or exclusively in northern
Alaska (table 9, p. 266) are of many families and vary in weight from
10 to 2500 grams. Some winter below the equator in South Ameri-
ca, on the shores of the Pacific Ocean, or in the continent of Asia.
They have few common characters except their habitual convergence
on the same arctic nesting area. The family Scolopacidae is most
numerously represented among these arctic-nesting birds, but 12
other families are also represented at Anaktuvuk by one or more
species. Among the 80 migratory species arriving to nest at or north
of Anaktuvuk, the 34 listed below nest only in the Subarctic or in the
Arctic, and of these species, the 18 marked with an asterisk (*) breed
solely in the Arctic.

*Gavia adamsii *Micropalama himantopus

Olor columbianus
*Branta canadensis taverneri
*Branta nigricans
*Anser albifrons frontalis
*Chen hyperborea

Clangula hyemalis

Grus canadensis

Pluvialis dominica dominica
*Squatarola squatarola
*Numenius phaeopus hudsonicus
Heteroscelus incanum
*Hrolia melanotos
*Hrolia bairdii

Hrolia fuscicollis
*Hrolia alpina pacifica
Limnodromus scolopaceus

Size of Clutch

Hreunetes pusillus
*Tryngites subruficollis
*Limosa lapponica
*Crocethia alba

Phalaropus fulicarius
*Stercorarius pomarinus
*Stercorarius parasiticus
*Stercorarius longicaudus

*Larus hyperboreus barrovianus

Sterna paradisaea

Sayornis saya yukonensis
Oenanthe oenanthe oenanthe
Phylloscopus borealis kennicotti
Montacilla flava tschutsehensis
Calearius lapponicus alascensis
Plectrophenax nivalis nivalis

There are many records of the numbers of eggs in the sets of
birds’ eggs which have been assiduously collected, and if the collector
unobtrusively watched the nest until the Jast egg was laid and then
determined that no subsequent addition was made, the number taken
as a set represents the reproductive accomplishment of the female bird.
But if sustained observation of the nest is not recorded, there is a
chance that collection of the eggs preceded the completion of laying
or that accident or predation may have removed one or more of them.
This can introduce a, statistical bias in any calculation, based on col-
lected sets, of the normal number of eggs laid in a clutch.

In order to illustrate data available from collections by experienced
naturalists, I have listed below the number of eggs in sets taken by
Murdoch and his associates at Barrow during their expedition for
observation of the First International Polar Year 1882-1884, and in

BIOLOGICAL ASPECTS Sh?

some sets taken by A. C. Bent from several American localities.
(These sets were deposited in the U. S. National Museum and the
information is abstracted from the catalog in the Division of Birds) :

Eggs in set
1 2 8 = 5
Pluvialis dominica dominica, Barrow
(Murdoch) 3 5 11
Bartramia longicauda, Martha’s Vine-
yard to Saskatchewan (Bent) 1 9
Actitis macularia, Massachusetts to
Labrador (Bent) 6 15 i
Catoptrophorus semipalmatus semipal-
matus, South Carolina (Bent) 1 6
Catoptrophorus semipalmatus inornatus,
Saskatchewan to Utah (Bent) 4 4
Erolia bairdii, Barrow (Murdoch) 1 5
Tryngites subruficollis, Barrow (Murdoch) 2 6 1
Ereunetes pusillus, Barrow (Murdoch) 1 6 13
Phalaropus fulicarius, Barrow (Bent) 1 8
Lobipes lobatus, Barrow (Murdoch) 2 4 7 14
Lobipes lobatus, Unalakleet (Bent) 1 14
ToTAL 2 8 35 86105 2

Sets of the 7 species of sandpipers listed average a fraction over 3 but
less than 4. A. C. Bent’s (1927, 1929) published reports on the nest-
ing of 50 species of sandpipers (Scolopacidae) in North America, give
4 as the characteristic size of a clutch in all cases. He chose to gen-
eralize by the criterion of his vast experience with the nesting of
birds in concluding what their normal reproductive performance is.
Numerous published reports upon the number of eggs charac-
teristic of the clutch of each species in different parts of the world,
do not allow a determination of whether the bias often present in col-
lecting of eggs has been eliminated from the conclusions. It is there-
fore necessary to restrict comparison of the number of eggs laid in
the arctic and in other climates to cases in which rather large dif-
ferences in reproductive effort seem to distinguish arctic birds from
those in warmer climates.

Although the family Scolopacidae, discussed by Bent (1927, 1929) as
occurring in North America, includes many species which nest in the
north there are some (eg., Catoptrophorus semipalmatus) which do
not migrate away from warm temperate regions. Others, lke
spotted sandpipers (Actztis macularia), nest over a range extending
from warm temperate states to the Arctic. Some of the species which
nest farthest north regularly migrate many times as far as those which
nest in the south. This condition holds true among North Ameri-
can sandpipers; and the same sized clutch is also characteristic of the
sandpipers in Germany and England, for Lack (1947) reported that

318 U. S. NATIONAL MUSEUM BULLETIN 217

in 18 species the clutch contained 4 eggs. Since the number of eggs
in a clutch is characteristic of 63 widely distributed species of the
family Scolopacidae, this constant number is a reproductive charac-
teristic of the family and presumably is not modifiable for adaptation
of any of the species to any climate. —

In other species of birds, like the snowy owl, the reproductive pro-
cedure is not limited to a fixed annual production of eggs. The
number of eggs laid by snowy owls, however, has been related to the
great annual fluctuation in the population of lemmings (Salomonsen,
1950) and is not an example of climatic influence on egg production.

The birds of some species lay fewer eggs in a clutch in tropical
climates than in Europe, according to Lack (1947), who reported that
in most families the clutches of species nesting in equatorial Africa
were smaller than the clutches produced by species of the same families
nesting in mid-European latitudes. Over the nesting range of
western white-crowned sparrows the largest clutches, averaging 5.5,
were laid at Old Crow (see p. 238). Less extensive observations indi-
cated that 5 or 6 eggs were usual at Anaktuvuk. In this species two
arctic populations laid more eggs in a clutch than were found in
southern localities. The average number of eggs in a clutch of song
sparrows (Melospiza melodia) in Baja California was near 3 and
in southern and southwestern Alaska it was about 4 (Johnston, 1954).
In some species the difference of about one egg seems to represent
the distinction in clutch size between hot and temperate or between
temperate and arctic climates. However, it is doubtful that the in-
creased size of clutch in a few species can be usefully related with
latitude, for latitudinal influences are applied without exception to all
widely distributed birds and in only a few species is the size of clutch
affected. In a significant relation to a general condition there should
be no exceptions.

Repeated Nesting

Differences amounting to less than one egg in an average clutch
can affect the reproductive rate, but the effect is likely to be con-
siderably less than that produced by multiple clutches in areas with
longer breeding seasons. In the long summer of temperate climates
there is time for twice or even thrice the number to be added to the
populations of fast breeding birds in comparison with the number
from the single clutch possible from an arctic pair. Thus, I have
found only one robin’s nest late enough in the season to have allowed
for the successful rearing of an earlier brood yet leaving time enough
for the young to grow sufficiently to migrate before freezing weather
terminated the arctic season for robins (L. Irving and Paneak, 1954).
I have found a few fresh eggs of redpolls late in June (see page 114),

BIOLOGICAL ASPECTS 319

but I doubt if they were late enough to have permitted their parents
to rear an earlier brood. Simon Paneak has related to me that
occasionally they find late nestling redpolls which have perished before
cold weather because the social attraction of the preautumnal flocks
of redpolls distracted the inclination of the parents from further care
for their young. The Eskimos believe that seasonal change in parental
attitude rather than immediate cold weather dooms the late-hatched
birds to destruction. I agree with the philosophical conclusion based
on Eskimo experience, that in arctic life a family is unlikely to sur-
vive behavior at variance with the regular seasonal program of a
population. Pitelka (1954) remarked that near Barrow late broods
are doomed because premigrational molting terminates parental care
of young. In his study of the behavior of snow buntings in Green-
land, Tinbergen (1939) remarked that second broods were rare and
he did not allude to the possibility of their successful contribution to
the population.

Migration and the Reproduction Rate

Loons and some other species do not produce more than two eggs
from each pair. Since visible casualties destroy some eggs and young
birds, I would estimate that these migrant populations returning
from the Arctic in autumn do not include more than one young bird
for each pair of adults which had arrived in spring. These species
could not preserve stable numbers unless at least two-thirds of the de-
parting migrants returned in the next spring.

Among the broods of sandpipers, plovers, and robins, I have ob-
served two is a common number to survive into late summer and ap-
proach adult size while the families are still distinguishable.
Considering that some families are wiped out and that some adults do
not reproduce at all, I doubt if the arctic populations of these species
are twice as numerous at the start of the southbound migration as
they were when they arrived in spring. During the eight or nine
months of their absence from the Arctic and in migratory flights over
thousands of miles an average mortality of less than 50 percent is
provided for by the new birds raised in the arctic summer.

It might be possible for the species like robins, which nest over
extensive ranges, regularly or occasionally to recruit from the sections
of their populations raised in warm climates in order to maintain the
arctic sections. But there is no nesting reservoir in milder climates
from which to recruit bird populations nesting exclusively in arctic
regions, and yet there is no sign that they are less stable than the popu-
lations of species which nest over a wide range of latitudes.

Estimates of the young birds reared are, of course, conjectural,
but a population of birds reared in the Arctic seems unlikely to start

320 U. S. NATIONAL MUSEUM BULLETIN 217

south with numbers permitting on the average 50 percent loss before
it returns to the breeding ground in spring. The losses from some
populations probably cannot be as much as 30 percent during their
absence from the Arctic. These estimates show that a major annual
loss from an arctic population cannot be normal and that it must be
rare. If an estimated death rate of 50 percent were distributed uni-
_ formly over 8 months of absence from the arctic the allowance would
be only 6 percent in each month. Some deaths must occur rather regu-
larly, because aging is not spasmodic and predators feed throughout
the year. ‘There does not appear to be any reproductive provision to
offset the possibility that arctic populations may lose heavily at any
single stage of migration.

Energy Resources for Migration

Hunters and ornithologists well know that storms cause mi-
grating birds to fly low and that bad weather may force them to land.
After severe weather during the time of migration some birds are
found which have perished from exposure or which have been scat-
tered to localities so far from their usual ranges that they can never
rejoin their populations. Many reports mention birds landing on
ships at sea exhausted during stormy weather and it is at these times
that birds are most often found injured by collision with lighthouses
or buildings. These reports indicate some of the hazards for birds in
migration. They do not give any idea of the normal proportion of
mischances because in fair weather the flights pass without presenting
unusual observations worthy of note. Often the normal flights are not
even visible in the darkness or occur on remote migratory routes.

There are numerous records attesting that birds arrive at the end
of long flights over water in exhausted condition. Wetmore (1939,
p. 178) vividly described the fatigue and emaciation which he saw
prevalent among the small migratory birds landing on the shores of
Venezuela from flight over the Caribbean Sea. In later studies at the
same landfall Voous (1953) reported that hundreds of blackpoll
warblers appeared tame because of fatigue, and remarks that black-
polls and redstarts weighed only half what Beebe had reported for
these warblers at their assembly points in northern Venezuela, before
their return northward. On Dry Tortugas, Florida, Bartsch (1919)
remarked that the migratory birds looked worn out in spring and
autumn. The condition of the overseas migrants may vary, for an-
other observer on Dry Tortugas reported that in autumn the migratory
land birds appeared in good shape (Sprunt, 1951).

The records of land birds seen over the western Atlantic Ocean
during cruises for oceanographic researches have been analyzed by
Susan Irving Scholander (1955). Some land birds were found at
sea further than 400 miles from nearest land and they must have cov-

BIOLOGICAL ASPECTS 321

ered these distances by continuous flight. While many of these birds
alighted exhausted on the ships there is evidently some traffic of cer-
tain species of land birds between America and Bermuda (600 miles)
which, if not regular, nevertheless occurs each year. Seasonal winds
had certainly influenced some of the flights, but recorded winds could
not be used to explain the frequent occurrence of these apparently
hazardous flights of lone land birds over the ocean. The repeated oc-
currence of these long flights demonstrates the long distance that a
single bird can cover without social guidance or landmarks.

For a hummingbird to fly across the Gulf of Mexico would exceed
the requirements of energy which Pearson (1950) could calculate to
be at its disposal. Weigold (1926) reported considerable variations
in the weights of migrant bird species arriving at Helgoland which
suggested that the lighter individuals had exhausted their reserves.
It is certain that long oversea flights tax and at times exceed the nor-
mal capability of birds, but a stationary observer is more likely to
remark upon migrating birds which are unsuccessful than to report
the swift flights of normal migrants.

Among Eskimos at Anaktuvuk it is commonly the impression that
migratory birds arriving in spring are in good condition and usually
fatter than later during the nesting season. It is also their view that
birds of some species become fatter again just before their departure
southward at the end of summer than they were at the time of arrival,
but the arctic people hesitate to generalize upon the condition of de-
parting migratory birds because the time of their departure is so
uncertain and the birds are then so often inconspicuous.

After examining many birds in 1948, 1949 and 1950 it was my im-
pression that in general they were in good condition and commonly
fat during the spring season of migration. The weights of birds taken
for specimens had been recorded along with occasional notes on their
apparent fatness. After 1950 the fatness of the birds was routinely
recorded. The designations were based upon the amount of fat visible
on the birds in terms fat, medium fat, little fat, and very little fat.
These terms mean whatever the observers associate with them in re-
lation to their value as food. Eskimos know that the fattest ptarmi-
gan has hardly as much fat as a rather lean loon or duck. The great
differences in the characteristic amounts of fat in the various species
required that the birds be compared according to the amount of fat
appropriate for each species. Finding good agreement in the designa-
tions of fatness by several Eskimos I followed their usage of the
terms, for their judgments were based upon their established code.

Among several thousand records of the fatness of birds which were
examined when taken for food or as specimens, I found fewer than
ten birds which were designated as lean or skinny. Wounded ducks
have been found after several days to be actually in emaciated con-

322 U. S. NATIONAL MUSEUM BULLETIN 217

dition, so that it is apparent that birds can survive to the point of
depletion of their visible reserves of fat. Even the birds which we
designated as very little fat still appeared active and in good con-
dition of plumage. They were not shrunken in other respects than
fatness. I believe that the birds in this category were not injured by
starvation.

TABLE 15.—Distribution of degrees of fatness among males of 21 species and
females of 9 species of common migrants to Anaktuvuk

Including redpolls | Not including red-
polls
Fatness

Number | Percent | Number | Percent

Fat 136 28 132 30

Medium fat 124 27 113 26

Little fat 165 34 150 35

Very little fat 55 11 37 9
Total (480) (431)

The records of fatness of common migratory birds at Anaktuvuk
are shown in the Appendix (figs, 21-86). For male birds of 21 com-
mon species and the female birds of 9 species the distribution in each
category of fatness is shown in table 15. The male and female hoary
redpolls (Acanthis hornemanni exilipes) and male common redpolls
(Acanthis flammea flammea) included in the first two columns of
figures, were less fat than the other migratory birds, having only 8
percent of fat birds. Since I suspect that redpolls winter not far
from Anaktuvuk, they are subtracted from the totals in the last two
columns to show the fatness of those species which migrate for long
distances. These records of fatness of migratory birds at Anaktuvuk ©
show that during the whole season only a small proportion were in
the lowest category of fatness.

The records in figures 21 to 36 were mostly concentrated during
the early season when the birds came in as arriving migrants, Among
the males of 9 species (Anas acuta, Aythya marila nearctica, Limno-
dromus scolopaceus, Erolia bairdii, Ereunetes pusillus, Hremophila
alpestris arcticola, Anthus spinoletta rubescens, Zonotrichia leuco-
phrys gambeli, and Calcarius lapponicus alascensis) the weight curve,
as Judged visually by the best fitting line, declined between 10 and
15 percent after the first arrival of migrants in spring. Among
males of 4 species (Pluvialis dominica dominica, Heteroscelus inca-
num, Eroka minutilla and Calcarius pictus) the decline was between
4 and 9 percent. The males of Spizella showed no change. The two
species of Acanthis, suspected of wintering nearby and probably
making only short migratory flights, showed no indication of chang-

BIOLOGICAL ASPECTS 323

ing weight during the early season. One well known migratory sand-
piper, Zrolia melanotos, showed an increase in weight during the
early season. There is wide variation in the weight of individuals of
this species (see p. 197).

In three species (Z'urdus migratorius migratorius, Oenanthe oenan-
the oenanthe, and Motacilla flava tschutschensis) the weights were
not distributed in a manner that would show seasonal change. The
weights of two species of Turdidae were affected by the heaviness of a
few individuals just before migration southward and by sparse early
season observations. The weights of J/otacilla are too few to indicate
a trend.

Less extensive data is available to show the condition of female
birds. In 5 species (Pluvialis dominica dominica, Heteroscelus in-
canum, E'rolia bairdii, FL. minutilla and Calcarius lapponicus alas-
censis) weights declined in the early season by between 10 and 15
percent. In the early part of the season the females of Lobipes
lobatus gained weight. The females of Acanthis hornemanni exilipes,
like the males, showed no change. Because of my belief that they are
not long-distance migrants they are not comparable with the other
birds. The number of weights of females of Zonotrichia leucophrys
gambelii is too small to show a trend, but it is reported for its relation
to Mrs. Oakeson’s interesting studies of this race.

Although the records of fatness are fewer than those of weight
and not numerical, they show a general correspondence with the pre-
dominating decline in weight after the first birds arrived. The males
of 13 species and the females of 2 species which declined in weight
during the early season also diminished in fatness. The males and
females of Acanthis did not show a significant change in fatness or
in weight. In comparison with many other species they were not
fat. Males of Erolia melanotos appeared to gain in fatness as they
did in weight. The species Z’urdus, Oenanthe, and Motacilla gave as
uncertain indications about fatness as they did in their weight.

These data are summarized in table 16.

TABLE 16.—Summary of differences in weight and fatness of common migratory
species after arrival at Anaktuvuk

Weight Fatness
Type of migrant
Lost No Gained Lost No Gained
change change
Long migration
Males ; 13 1 1 14 0 1
Females 5 0 1 2 0 1
Short migration
Males 2 2

Females il 1

324 U. S. NATIONAL MUSEUM BULLETIN 217

There is such a sudden change in the activity of migratory birds
when they reach their nesting grounds that it would be surprising
if visible signs failed to indicate a change in their physiological con-
dition. The northward progress of migration involves all migrants
to Anaktuvuk in flights which must proceed at a rate sometimes
greater than 60 miles a day in order to conform with the indications
of the northbound schedule. After reaching their destination many
species do not fly often or far and become sedentary birds in contrast
with their activity during migration. The mechanisms of flight
remain usable but they are little exercised.

There is a possibility that the musculature of flight may diminish
in migratory birds which become sedentary on their nesting grounds.
The evidence available is insufficient to suggest whether the birds like
gulls, jaegers and hawks, which continue to fly extensively while on
their breeding grounds, maintain their migratory weight while
breeding.

The average weight of specimens of birds in various parts of their
range is known to differ in some cases without other visible taxonomic
distinctions appearing. Von Zedlitz (1924) reported that the average
weight of Zetrao urogallus from Sweden was 3.65 kg. and from
Hinterpommern 5 kg. I have discussed the variations in the weight
of willow ptarmigan (Lagopus lagopus alascensis) from various
localities, which appear to be significant of populations differing in
weight although without taxonomic distinction. It is not known
whether these differences in weight are maintained through heredity
or whether they are simply induced in individuals by the two environ-
ments.

The birds nesting in the part of arctic Alaska we are considering
are geographically homogeneous populations, and this factor may be
important in the small “coefficients of variation” which were found in
the weights of the species. I have taken as “coefficient of variation”
the usual meaning, i. e., standard deviation as percent of the mean
weight. In the weights of adult males of 21 species the coefficients of
variation were between 5.3 and 12.6 percent. Among 9 species the
coefficients of variation in the weights of female birds were from 7.2 to
12.4 percent. The average of the coefficients of variation in male
examples of 21 species, each composed of from 10 to 40 individuals,
was 8 percent. Among samples of the females of 7 species the average
of the coefficients of variation was 9 percent. Considering the chances
for error in gathering figures for weight under the conditions of
arctic field study the birds of each species appear homogeneous in
their weight characteristic at Anaktuvuk.

The weight of a small bird may change considerably during the
course of a day, as is shown in the report by Linsdale and Sumner

BIOLOGICAL ASPECTS 325

(1934) of a golden-crowned sparrow weighing usually about 28 grams
which lost 20 percent of its weight between one afternoon and the next
morning. That this change was exceptional appears from their rec-
ords showing that several days elapsed before the bird regained its
usual weight. Many records show that a decline in weight of small
birds occurs over night and that their weight increases during the
day. Baldwin and Kendeigh (1938) reported that the average diurnal
curves for the weight of several passeriform species changed from
about 97.5 to 101 percent of the mean weight between 7 a.m. and 6
p.m. Since sparrows weighing 20 grams lost about 10 percent when
caged without food during a night, the rate of loss of the captives
exceeded the average nightly loss of free wild birds in summer. I
am inclined to think that the regularity of weight records for each
species at Anaktuvuk, although they were taken at random times
during the day, is influenced by the fact that with scarcely an hour
of darkness there in early May, feeding is never for long interrupted.
It is also my impression that the birds there feed frequently but not
for very long periods.

The daily food intake of small birds is undoubtedly a quite
appreciable part of their body weight. Captive English sparrows
weighing about 24 grams consumed from 4 to 9 grams of mixed feed
daily, according to my calculations from the metabolic heat production
described in Kendeigh’s detailed study (1949). The fact that average
weight characterized each of Kendeigh’s sparrows shows that their
food intake and its utilization were kept in close relation in captivity
when food was abundant. The same close accord between consumption
and utilization of food seems to be the rule under natura] conditions.

It has been implied in our discussion that variation in weight among
the birds of a homogeneous population represents changes in the
quantities of nutritive substances in their bodies disposable for met-
abolic purposes. The metabolizable substances include food which
has been recently consumed and parts of the tissues themselves.
Weights of Anaktuvuk species under consideration differed nearly
100 fold from redpolls (Acanthis) weighing only 12 grams to scaup
(Aythya) as heavy as 1000 grams. The basal metabolic rate of birds
has been found to vary with their size according to a formula given by
Brody (1945, p. 871) empirically describing the metabolism of do-
mesticated birds:

kilo-calories per day=89 X weight (in kilograms)?/*
A few wild birds which we have examined in Alaska (Scholander,
Hock, Walters, and L. Irving, 1950; L. Irving, H. Krog, and M.
Monson, 1955) conformed sufficiently well with this relation to war-
rant its use for calculations. A bird weighing 10 grams has a meta-
bolie rate at rest which would expend about 4.8 k.-cal. in 24 hours. A

326 U. S. NATIONAL MUSEUM BULLETIN 217

bird of 1000 grams would utilize 84 k.-cal. in 24 hours at the resting
rate. In case the two sizes of birds metabolize substances of equal
value for energy, the bird weighing 10 grams would consume a given
percentage of body weight about 6 times as fast as the bird weighing
1000 grams.

In order to relate the conversion of energy with the substances which
_ are actually metabolized it is necessary to define the energy provided
by the substances which are oxidized in the course of metabolism.
Fat has the highest value for energy production and its storage and
use involve little water. In the metabolic oxidation of fat, one gram
yields 9.4 k.-cal. (Brody, 1945, p. 385). A bird weighing 10 grams
would accordingly obtain 9.4 k.-cal. by the oxidation of 10 percent of
its substance in the form of fat. The oxidation of a given weight of
protein or carbohydrate produces only about half as much energy
as is derived from fat. These lower yielding substances are further
less suited than fat for metabolic reserves in that they are likely to
be stored and used along with a certain amount of water.

The comparison of metabolic rates just given was based upon the
condition of animals resting in air warm enough so that no special
metabolic provision would be needed for maintaining the warmth of
the body. These metabolic rates are accordingly at the minimum at-
tainable experimentally for brief periods and they do not make pro-
vision for the usual activities of life or for cold weather. Reasonable
estimates can be prepared to indicate the magnitude of the daily me-
tabolism of birds migrating during the cool season when they reach
the arctic. These are shown in table 17. Because of the limited
natural insulation of birds of small size, at just freezing temperature,
a bird weighing 10 grams would need at least double its basal metabolic
rate in order to preserve its warm body temperature (Scholander,

TaBLe 17.—Estimate of the daily budget of metabolism of three sizes of birds
approaching the arctic terminus of migration in O°C. average temperature

Relation to normal basal
metabolic rate

Item
Size 10 Size 100 Size 1000
2 g. :

Sleeping (10 hours) 20/24 10/24 10/24
Waking, but not very active (10 hours), 2

times basal rate 20/24 20/24 20/24
Flight (4 hours), 6 times basa] rate 24/24 24/24 24/24
Sum of metabolic periods 64/24 54/24 64/24
Relation to basic metabolic rate 2. 67 2.25 2. 25
Kg.-cal. per day 12.8 45.0 189
Equivalent grams of fat 1.36 4.8 20

Reserve (days) provided by metabolism of
10% of body weight as fat 0. 74 2.1 5

a aaa EEEEEE UTTER RIESE

BIOLOGICAL ASPECTS 327

Hock, Walters, Johnson, and L. Irving, 1950; L. Irving, H. Krog, and
M. Monson, 1955) but an arctic bird weighing 100 grams would prob-
ably be sufficiently insulated to get along at freezing temperature with
just the heat produced by basal metabolism. In table 17 I have al-
lowed 10 hours in the day for rest with metabolic rates on account of
cold, at twice the basal level in birds weighing 10 grams and at the basal
level in birds weighing 100 grams or more. I have then allowed that
during the 10 hours of the day when migratory birds would be awake
but not very active, they would require about double the basal rate
of energy expenditure in that period. I have allowed that during
4 hours of active movements of flying the metabolic rate would be six
times as great as the rate at rest, an expenditure of energy that cor-
responds with the rate of a man or dog walking rapidly. I have used
the figure, however, because Pearson’s (1950) measurements showed
that the hovering flight of a hummingbird elevated its metabolism
to about six times the resting rate.

The activity of a wild bird can hardly be regularly 50 percent
greater or less than I have allowed, for wild birds commonly pursue
their lives at a moderate pace. Even their swift movements in air,
which so far exceed the speed of man on the ground, are probably so
well within their capacity that they can probably fly as easily as a
trained man can walk. I have not noticed an arctic bird that appeared
to be exhausted by exertion, and it is my impression that they seldom
work at their highest rate.

These estimates of daily metabolic costs allow us to evaluate the
significance of the reserves for nutrition which exist in the body of a
bird. If these reserves are in the form of fat and amount to 10 percent
of the body’s weight they would provide the metabolic energy which
is expended during from 34 of a day to 5 days according to the size
of the common birds.

The accumulation of fat which we acquire on our bodies is not
readily removed by current metabolism. The fat of obesity for ex-
ample is sequestered from current use and can only gradually be with-
drawn. But fat in the amount of 10 percent of the body weight does
not constitute obesity. I have found ducks two or three days after
they had been wounded to bs in lean condition whereas ducks killed
instantly from the same flock were fat. It has been shown (p. 322)
that about a 10 percent decline in average weight is often visible among
certain species of arctic birds two or three weeks after their arrival
on their nesting grounds, and that (p. 824) there are numerous in-
stances of rapid decline in the weight of birds during starvation
which indicate that their substance can be quickly utilized for metabo-
lism when food is unavailable. Among the migratory birds after their
arrival at Anaktuvuk the amount of visible fat decreased as their

469496—60——22

328 U. S. NATIONAL MUSEUM BULLETIN 217

weight diminished (see p. 323). Asan example of extreme utilization
of reserves, the small weight of warblers which arrived exhausted
after flight over the Caribbean (see p. 320) shows that they had made
extensive use of their substance during a long flight (Voous, 1953).
It appears that the body substance of birds can be rapidly consumed
and that at times it is used to meet current metabolic demands or
emergencies during migration.

In comparing the weight of birds at the start of migration with
their weight at the terminus we are disturbed by the fact that a sample
of a migratory species taken on its wintering ground or along its
migratory route may be setting out for any part of its eventual nest-
ing grounds. There are many reports indicating that before depart-
ing for northbound migration birds wintering in temperate climates
acquire fat, but few reports show condition at the end of migration.
I will only refer to a few observers whose records of fatness appear
directly comparable with my observations at Anaktuvuk.

In the discerning view of von Zedlitz (1926, 1927) the terminus
of migration in Sweden was a favorable place for observation because
all the birds were alike in proximity to their nesting place. On his
estate in central Sweden, where he had watched the birds through
many years, he reported careful records of the weight and condition
of the migrant birds which for the most part came to settle and nest
there. The list of birds is interesting because their names were all
assigned by Linnaeus whose residence had been nearby. Corvus cornix
cornix, Sturnus vulgaris vulgaris, Fringilla coelebs coelebs, Columba
palumbus palumbus, and Scolopax rusticola rusticola arrived fat and
then lost weight at mating time. Von Zedlitz’ observations differ
from those of Weigold (1926) on Helgoland, a point along a migratory
route over water. There Weigold found variation in the weights of
examples from a number of species which he took to illustrate occa-
sional encounter with nutritional hardship during migration. Von
Zedlitz’ point of observation corresponds with mine in being at the
terminus of a normally completed migration. His conclusions agree
with mine that there is no evidence that successful migration leaves
signs of strain in the nutritional state of birds.

To T. T. McCabe (1948), a long-experienced collector of birds in
British Columbia, it was apparent that the northbound migrants were
commonly fat and frequently very fat but that they lost weight rap-
idly on their breeding grounds. As an interesting contrast Mrs. Oake-
son found that the Puget Sound race of white-crowned sparrows
(Zonotrichia leucophrys) on arrival at their nesting ground at Tilla-
mook were not fat (Blanchard 1942). Unlike the birds considered
by McCabe these white-crowned sparrows had reached Tillamook after
only a short migration from a nearby wintering ground. These in-

BIOLOGICAL ASPECTS 329

teresting studies developed the evidence for the proposition that races
of Pacific Coast white-crowned sparrows destined to make long mi-
gratory flights northward became fat in the period before departure
while races preparing for only a short migratory flight accumulated
little or no fat before the event.

Mrs. Oakeson has kindly sent me figures recording the weight and
fatness of examples of the race of white-crowned sparrows examined
by her just before leaving their California wintering ground and by
McCabe at a station in British Columbia more or less in midcourse
of migration. At both places the sparrows were generally heavy and
commonly fat. When members of this race were taken after arrival
on their breeding ground at Mountain Village, Alaska, they were
lighter and less fat than in the early stage of the migration through
Santa Barbara, California (Oakeson, 1953). The average weight of
7 males in May at Mountain Village was 25.8 grams. At Anaktuvuk
25.6 grams was their average weight during the summer, but during
the first week after arrival the males were about 5 percent heavier. I
conclude from Mrs. Oakeson’s records that in April when this race
started migration through California their average weight was as
much as 20 percent greater than after their settlement on their Alaskan
breeding grounds. A decline in weight of birds on the nesting ground
appears to occur soon after arrival, but I suspect that some migrants
start to lose weight during the last stages of their northern flight.

In several species at Anaktuvuk the females did not appear to lose
weight until after their eggs had hatched. This contrast between the
sexes conforms with the intense activity of male birds after they
arrive, while courting and maintaining their nesting territories. This
period does not appear to involve the female birds in especially in-
tense activity, for they take life easily during mating and incubation.
During the mating season the males may even be too busy to eat well.
Their fat, which is certainly not completely used up during migra-
tion, could serve them well on the breeding ground in the way that
reserves of fat enable male fur seals and bull caribou to neglect feeding
in order to devote their time exclusively to the jealous watchfulness
and demonstrations by which they seek to secure their paternity in
the offspring of their mates.

tie fo * as
fotos \
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‘ Vial? ; Sy) aa

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yy tai VY art shaw ba
evr creer TOIL
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bie yraen wilted Big BWOTE

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7
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high “anh ; iy thd 7 of

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A

aha Erie
i RW cit BS avi ieutey i Oiip
a aifinat ‘oe alge an Oa “HO¢ be ay Hall baw
atl SPN s hair ano TR wet

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hk tae ban
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Adak Gani urn dind
My

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TOES ak TES

Petacic Cormorant Phalacrocorax pelagicus (see p. 275).

9. Arctic Metabolic Economy of
Warm-blooded Animals

Wu SUPPLIES SHIPPED from outside, a foreigner to the Arctic can
afford to eat as much food for fuel as his metabolic process can
handle, but an indigenous people must maintain a culture, a society,
and families on food provided from local sources. Recognition of the
large portion of man’s schedule applied to other activities than feed-
ing should indicate to us the common necessity for animals also to
conserve their metabolic expenditures in order to have time in which
to maintain family and social systems, and we should therefore be
cautious about viewing the economy of animal heat in the arctic with
the thought that arctic animals can preserve their body temperature by
increasing their metabolic heat production much beyond the rates
normal for animals in other climates.

: 331

33y7 U. S. NATIONAL MUSEUM BULLETIN 217

The availability of food, the time applicable to obtaining food, and
the capacity of the metabolic processes set limits upon the extent to
which warm-blooded animals can, by increasing their metabolic pro-
duction of heat, adjust their lives to the cold arctic climate. These
prior considerations lead us to look particularly for the methods
by which metabolic heat may be conserved, as an essential to the
adaptation of warm-blooded creatures to arctic life, and to analyze
homoiothermism among arctic animals to see how the operation of
its component factors is suited to the arctic environment.

Body Temperature of Arctic Animals

The regular warmth of birds and mammals appears in sharp con-
trast with the temperature of arctic winter, which has been reported
as cold as —68° C. in eastern Siberia (Bartholomew and Herbertson,
1899; Court, Sissenwine, and Mitchell, 1949), and —65° C. in north-
western Canada (fig. 16). In experimentation at Barrow we found
that at temperatures frequently occurring in winter, arctic warm-
blooded animals could maintain their bodies some 80° warmer than

the air by the production of metabolic heat at their basal rate and by
its conservation with their natural insulation (Scholander, Walters,
Hock, and Irving, 1950).

We now have temperature measurements for many resident species
of Alaskan birds and most mammals except those of the smallest size
(L. Irving and J. Krog, 1954). In winter at —45° C. and in warm
summer the mean body temperature of 15 arctic and subarctic resident
species of birds averaged 41.3° C. The mean body temperatures
differed among the arctic species of birds by about as much as was
found by Wetmore (1921) among species from temperate regions,
and the means (fig. 17) for the arctic and temperate species exhibited
about the same differences.

Body temperatures of homoiothermic species are phylogenetically
differentiated, but the differentiation shows no common regard for
geographical range, and their temperatures are more stable than the
climates of the earth, which have changed appreciably, while the tem-
peratures of the birds that inhabit them apparently have remained
fixed. Body temperatures of arctic species show no indication of
modifiability by climate and the small differences in temperature
among the species show no adaptive qualifications for arctic life.

During our experimental study a number of arctic mammals were
observed to hold their body temperature with remarkable constancy
for several hours in air as warm as +30° C. or as cold as —50° C.
We have not as extensive a view of birds, but it appears that their
regulation of body temperature is about equally stable. Mammals or
birds which had been active showed temperatures elevated 2° or even

ARCTIC METABOLIC ECONOMY Sao

WORLD RANGE OF

CLIMATES ff ADAPTATION
-60

ARCTIC ALASKA
ARCTIC
30 MAMMALALS

200 BIRDS

TEMPERATE

Ficure 16.—Temperature ranges for which warm-blooded animals are physiologically
adapted compared with those of the world’s climates.

3° above the lower range of body temperatures at rest. It is my view
that the body temperature of arctic mammals and birds remains at
a miniumum value characteristic of the resting state except when
excitement or activity elevates it above this well regulated minimum
level.

During winter nights at Anchorage when the air was —10° C. to
—20° C. I found the body temperatures of six species of Alaskan wild

304 U. Ss. NATIONAL MUSEUM BULLETIN 217

e-
ae MEAN 50 FAMILIES
¢ © WETMORE

MEAN 29 SPEGIES
ALASKAN 4 =0.8

°C

4\|/- ——— — “ee — — — 6 ~~ @ARCTIC RESIDENT BIRDS
w @SUBARCTIC OR __
= MIGRATORY BIRDS
ae )
a
i 40
uJ
a
=
Lad MEAN
m 39 31 ALASKAN ®
é SPECIES 420.8 ida nit
5 | MEAN 56 SPECIES
= MORRISON ¢21.23

38F-OARCTIC MAMMALS
@SUBARCTIC MAMMALS

37

i §00 1000 10,000 100,000 1,00 0,000
LOG BODY WEIGHT IN GRAMS

Ficure 17.—Mean body temperatures of arctic and subarctic birds and mammals. (From
L. Irving and J. Krog, Journ. Appl. Physiol., vol. 9, fig. 6, p. 677.)

birds more variable than during the day (L. Irving, 1955). At night
the birds were often awake and active when we approached their cages.
On several occasions, however, the body temperatures of some of them
were 3° lower than the resting level by day, and I concluded that in cold
nights, probably during sleep, northern birds may cool below their
daytime body temperature by about as much as Dawson (1954) re-
ported for birds in temperate regions. This nocturnal relaxation
of either minimum temperature level or regulation among northern
birds, however, is not enough to have much significance for the con-
servation of bodily heat in the Arctic.

It is true that during hibernation at reduced body temperature
mammals save on the expenditure of metabolic heat (Hock, 1951).
At Barrow hibernating ground squirrels (Citellus undulatus) were
often near 0° C. (Erikson, 1956). At this low temperature their
heat production is so meager that it cannot protect them from freezing
in air only a few degrees colder. Accordingly their winter survival
depends upon obtaining such effective insulation from a nest and bur-

ARCTIC METABOLIC ECONOMY S50

row as will conserve their feeble production of metabolic heat. For
the very numerous arctic population of ground squirrels, hibernation
is a successful evasion of winter cold, but as far as I know the faculty
of hibernation at depressed body temperature is only exercised in the
Arctic by two mammalian species, Citellus and Arctomys.

It has been suggested that ptarmigan hibernate in their burrows
under the snow, but no indication was found that the body temperature
of willow ptarmigan shot early in winter mornings or late in the
afternoon differed from the midday level (L. Irving and J. Krog,
1954). Willow ptarmigan enter snow burrows at evening with crops
containing tips of willow twigs and buds amounting to as much as
one sixth their body weight. When they leave in the morning a large
mass of droppings shows that their digestion has been active during
the night.

Birds of the tundra lack the shelter of trees. Cliff dwelling gyr-
falcons and ravens on their roosts can find shelter from wind, but I
doubt if they find or require protection from arctic cold. Cade (1953)
has observed the common redpoll in winter near Fairbanks entering
apertures formed through the snow around weed stems, and it is
reasonable to suppose that such shelters afford some protection.
Simon Paneak remarked that hoary redpolls had been seen entering
the spaces in the snow through which willow bushes protrude. Inas-
much as these openings are sealed by each fresh snow and do not com-
monly form before the sun rises in winter, we did not think that the
redpolls would regularly be able to find shelter much warmer than in
the open air. I do not know where chickadees, jays, and grosbeaks
roost on the tundra, but I do not think that any small tundra birds use
shelters so well enclosed that at night the air in them would be warmed
much above the temperature outside. Under these conditions arctic
birds cannot reduce their body temperature by hibernation or in tor-
pidity, for because of the diminished heat production observed in
torpid birds they could not be expected to counteract the cold of even
just freezing weather (Pearson, 1953). Arctic tundra mammals
smaller than hares remain under the snow. Signs of pursuit by fox
and weasel reveal the extensive winter activity of the great numbers of
mice and lemmings otherwise concealed by snow. Nordenskidld (1882,
p. 114) described vividly how the melting of the snow disclosed the sur-
prising extent of the hidden winter activities of arctic small mammals
by showing the runways, nests, and dung. We have no measurements
of the body temperature of these small mammals in their natural winter
habitat, but the indications are that they are those usual for mammals.
Underneath the snow the small mammals are exposed only to modi-
fied arctic cold, but the small arctic birds seem to be unable to escape
the severe cold of arctic winter.

336 U. S. NATIONAL MUSEUM BULLETIN 217

Essentials for Maintaining Body Temperature

Before his sacrifice to the political tempers of the French revolu-
tion, Lavoisier (1777) had demonstrated that the production of animal
heat could be ascribed to the oxidation of carbon and hydrogen in the
animal’s body. Since that time physiological complications have often
been allowed to obscure the view that the heat produced by animal
combustions must pass to their environment in accordance with phys-
ical laws for the exchange of heat. The difficulty has been largely
methodological, and has arisen because man and the domestic and
laboratory animals commonly used for metabolic studies are so little
accustomed to cold that the factors of heat exchange are not large
enough for their relations to be readily apparent. The contrast be-
tween the conditions for heat exchange in arctic and tropical animals
presented their heat exchange in dimensions of such contrast that the
essential physical factors of the process could be related in a simple
form of Newton’s law of cooling (Scholander, Hock, Walters, John-
son, and Irving 1950).

Heat escapes from a warm body to cooler surroundings by conduc-
tion at a rate which is proportional to the difference in temperature
between the body and its surroundings. Warm-blooded animals, in or-
der to maintain a constant temperature, must produce as much heat
as escapes from their bodies. The loss of heat from animals proceeds
through surfaces which are insulated. Fur and feathers afford the
conspicuous insulation of animals but other natura] insulating sys-
tems also serve in the conservation of bodily heat. If all the devices
resisting the loss of heat are regarded as components which combine
to provide insulation in still air, the system preserving body tempera-
ture can be described by the use of Newton’s law of cooling (Scho-
lander, Hock, Walters, Johnson, and Irving, 1950),

T,—-T,=KI1H,

where 7’, and 7’, are the temperatures of the animal body and of the
surrounding air, Z is the overall insulation of the animal, H is the
production of heat by metabolism, and X is the factor appropriate to
relate the units of measurement used.

In order to compare heat exchange among animals of various sizes
the basal metabolic rate of each animal was represented as 100 and
the observed metabolic rates were plotted against air temperatures.
Figure 18, which shows these relations among arctic and tropical
species of mammals and birds, indicates that the larger arctic birds and
mammals maintained their basal metabolic rates in cold and could ap-
parently sustain any arctic winter weather without heat beyond that
produced at the level of basal metabolism. Tropical animals, on the

ARCTIC METABOLIC ECONOMY par

METABOLISM
MAMMALS BASAL 100

400

GR,
300 ”SaU/p
‘) pois tat
POL: ae me Mi ik

ey ‘ 4,
Uy Uj jprcr
way,
Sa, 200
rf My Pe
LP pa Sue / / pe ;

Lf”) 75 SS
00 TPLLANGER ndings 77 ; = SS Sas 100
; BALL

—— OBSERVED
—-—EYTRAPOLATED

t)
0
=i B® -60 ~50 -40 -30 -20 -10 Oo 10 20 30 sone Te 40
GHEARIN AIR TEMPERATURE IN CENTIGRADE
METABOLISM
B IROS BASAL® 100

4a0 400

TT RAO POV eAL

é ‘s, Ni
wo a7) Min (rq My’

ays
<Cih

oo ; :

70% = E FS) 10
TB OM EARTH AIR TEMPERATURE IN CENTIGRADE

Ficure 18.—Heat regulation and temperature sensitivity in arctic and tropic mammals
(top) and birds (bottom). (From Scholander, Hock, Walters, Johnson, and L. Irving,
Biol. Bull., vol. 99, figs. 10 and 11.)

other hand, started to increase their metabolism at temperatures only
a little below those normal in their natural environment.

By using this method of formulation it was found that the metabolic
measurements by a number of investigators working in temperate
climates produced curves showing the same pattern of relation between
metabolism, insulation, and temperature. The formulation was suc-
cessfully used by Hart (1952) for defining the character of the metab-
olism in cold of several kinds of wild and domesticated mice. The
formulation also served to compare the metabolism in summer and
winter of three species of mammals and of two birds in Alaska (L.
Irving, H. Krog, and M. Monson, 1955), to describe the metabolism
in cold of an Alaskan mountain goat (H. Krog, and M. Monson, 1954),
and to compare the endurance of cold by wild brown rats and by rats
of the white laboratory form (H. Krog, M. Monson, and L. Irving,
1955).

338 U. S. NATIONAL MUSEUM BULLETIN 217

Critical Temperature

The temperature at which an animal must increase its metabolic
heat production above the basal level is spoken of as the “critical tem-
perature,” and it differs among species, as shown in tables 18 and 19.
No small mammals or birds have been found with a low critical tem-
perature, since the mechanics of movement prevent them from wearing
enough fur or feathers for substantial insulation. Among the north-
ern mammals the arctic white fox (Alopex lagopus), weighing about
4 kg., is the smallest that we have found able to stand arctic cold with
only its basal metabolic rate (Scholander, Hock, Walters, Johnson,
and Irving, 1950). The distribution and habits of arctic hares (Lepus

-timidus) of about the same weight show that they withstand exposure
to any arctic cold, but snowshoe hares (Lepus americanus) obtain
some shelter in their brushy habitat (L. Irving, J. Krog, H. Krog, and
M. Monson, 1957). At Leningrad, Olnianskaya and Slonim (1947)
showed that arctic hares and several other arctic mammals conserved
their body temperature better in cold than some northern, but not
arctic, species. Smaller arctic and subarctic mammals such as weasels,
mice, lemmings, and ground squirrels seldom emerge for long from
their burrows and nests under the snow, where the protection from

TaBLe 18.—Critical temperatures at which the metabolic rate increases in some birds

Critical

Species Region Season Weight | tempera-

E ture

(°C.

Larus hyperboreus barrovianus (1) | Barrow Winter 1500 | below
—30
Perisoreus canadensis pacificus (1) | Barrow Winter 60 —5
Plectrophenax nivalis nivalis (1) Barrow Winter 40 12
Branta nigricans (2) Anchorage Winter 1118 6
Branta nigricans (2) Anchorage Summer 6
Corvus caurinus (2) Anchorage Winter 280 —10
Corvus caurinus (2) Anchorage Summer —10
Anser (domestic) (8) (4) Europe 4700 3
Gallus (domestic) (4) Europe 2000 26
Gallus (domestic) (9) America 2000 26
Columba (domestic) (5) Europe 400 28
Columba (domestic) (4) (6) Europe 20
Passer domesticus (7) Ohio 24 22
Troglodytes aedon (8) Ohio Summer 10 25
Nyctidromus (1) Panama 45 25
Pipra (1) Panama 12 20
Vidua paradisaea (3) Native to 13 32

Africa

SOURCES:
(1) Scholander, R. Hock, Walters, Johnson, and L. Irving, 1950, p. 237.
(2) Irving, H. Krog, and M. Monson, 1955.
(8) Terroine and Trautmann, 1927.
(4) Giaja, 1931. (7) Kendeigh, 1944,
(5) Kayser, 1930. (8) Kendeigh, 1939.
(6) Falloise, 1900. (9) Mitchell and Haines, 1927.

ARCTIC METABOLIC ECONOMY 339

TABLE 19.—Critical temperatures at which the metabolic rate increases in some
mammals in Alaska

{Explanation—(A) Arctic, (SA) (Subarctic.]

Critical tem-
perature (° C.)
Species Region Weight (kg.)
Winter | Sum-

Thalarctos maritimus (1) A (cubs) 9 0 0
Mustela rixosa (1) A 0. 04-0. 07 18
Vulpes vulpes (2) SA 5 —13 8
Alopex lagopus (1) A 3. 8-5. 5 —40
Canis familiaris (1) A (pups) 15 —25
Citellus undulatus (1) A 0. 870-1250 8
Tamiasciurus hudsonicus (2) SA 0. 165-0. 230 20 20
Dicrostonyx torquatus (1) A 0. 045-0, 056 15
Rattus norwegicus (3) SA 0. 200 23
Erethizon dorsatum (2) SA 4-7 —12 7
Lepus americanus SA 1 —12
Oreamnus americanus (4) SA 32 —20

Sources:
(1) Scholander, Hock, Walters, Johnson, and Irving, 1950.
(2) L. Irving, H. Krog, and M. Monson, 1955.
(3) H. Krog, M. Monson, and L, Irving, 1955.
(4) H. Krog and M. Monson, 1954.
(5) L. Irving, J. Krog, H. Krog, and M. Monson, 1957.

weather afforded by their behavior in seeking shelter probably keeps
their immediate environment at about their critical temperatures. It
appears that in general large arctic mammals have sufficient insulation
of their own to suit the climate in which they live. Smaller mammals
with inadequate insulation for arctic weather obtain shelter in the
burrows and nests habitual to their kind and by curling up to rest
in especially well protected sleeping places in effect insulate them-
selves by their behavior.

Under natural conditions smal] arctic mammals come out of their
shelters occasionally to run about on the snow. The tracks of ground
squirrels show that in spring and fall they are often out during
weather much colder than the temperatures which have been found ex-
perimentally to be critical. Tree squirrels (7 amiasciurus hudsonicus)
are seen occasionally in arctic forests during midwinter, lemmings
(Dicrostonyx torquatus) are seen in late winter running over the snow,
particularly in the years of their cyclic abundance. Clay Kaigelak
and I found a lemming scratching outside our camp one morning in
April on one of the Plover Islands, a scarcely perceptible sand bar
rising above the sea ice more than a mile from the shore east of Point
Barrow. The nights had been as cold as —30° C. and the days were
no warmer than —10° C. while the lemming ran out there over the
sea ice. Lively and pugnacious, it lived captive in the cold camp for

340 U. S. NATIONAL MUSEUM BULLETIN 217

another day. We crossed the tracks of an occasional lemming while
traveling some 20 miles at some distance from shore over the sea ice
and I found one about four miles from land dead but undamaged
by predators. These forays of lemmings turned out badly but they
show that while active these small and poorly insulated mammals
can for some hours withstand cold far below their critical tempera-
ture.

It is common to see the tracks of the arctic least weasel (Mustela
vizosa) in the snow of the arctic forest and over the tundra. I have
never followed them successfully very far before losing them where
they apparently returned beneath the snow. ‘The tracks of the larger
weasel (Mustela erminea) I have followed for a mile without reach-
ing the end of its continuous journey in cold winter air. They are
larger and have somewhat thicker fur than the least weasel, but I
doubt if their insulation could protect them with resting metabolism
in air colder than 0° C. Both species of weasels regularly expose
themselves while active, and they can probably pursue their intensive
activity for several hours at a rate sufficient to preserve their body
temperature by the heat of activity, although their insulation is not
sufficient for them long to endure cold while at rest.

Although some species have been observed to raise their metabolic
rates about six times in cold, others managed only about a 3-fold in-
crease during several hours of experimental exposure to cold. As yet
no climatic pattern in the possession of this faculty is shown among
tropical and arctic, small and large forms of mammals and birds,
but that it is, in part, a matter of metabolic disposition, is suggested
by the fact that aggressive wild brown rats (Zrattus norwegicus) of
Fairbanks raised their metabolism in experimental cold nearly twice
as much as white laboratory rats (H. Krog, M. Monson, and L.
Irving, 1955). And we have as yet no criteria for deciding whether
the metabolic increase originates in other tissues than the visibly active
muscles.

When the increase of metabolism below the critical temperature
accurately supplies the heat lost and maintains normal body tempera-
ture the line relating metabolism to air temperature is straight.
This is the situation postulated in the description of homoiotherms
as heat machines (Scholander, Hock, Walters, Johnson, and Irving,
1950). Hart (1952) found the body temperature of mice to be well
preserved by their muscular activity for a certain amount of cold
below their critical temperature. White rats regulated their body
temperature during short experimental periods in air temperatures
10° or 20° below their critical temperature, but in colder air their
body temperature dropped and the line showing elevation of
metabolism progressively fell below the theoretical line for

ARCTIC METABOLIC ECONOMY 341

homoiothermism (H. Krog, M. Monson, and L. Irving, 1955).
Porcupines in winter maintained body temperature and related metab-
olism to cold below their critical temperature better than they did
in summer (L. Irving, H. Krog, and M. Monson, 1955). In general,
well adapted arctic mammals preserve their body temperature very
accurately over ranges of cold which are far beyond the tolerance of
tropical forms (lL. Irving and J. Krog, 1954). As already men-
tioned, the small mammals find in their habits and behavior the sup-
plemental protection from the arctic climate which physiological
adaptation or adjustment cannot provide for them.

Stability of Basal Metabolic Rate

Although we have now no measurements of the temperature in the
burrows occupied by arctic tree squirrels, weasels, lemmings, mice,
voles, and shrews, these animals while resting probably are not exposed
to temperatures much lower than their critical temperature. If this
expectation is confirmed, they can rest without their metabolic ex-
penditure exceeding the basal level appropriate to animals of their
size. We have not found the basal metabolic rates of small or large
wild arctic mammals to differ from those of tropical or temperate
forms in any manner related to climate (Scholander, Hock, Walters,
and Irving, 1950; H. Krog and M. Monson, 1954; H. Krog, M. Mon-
son, and L. Irving, 1955; L. Irving, H. Krog, and M. Monson, 1955).
Mammalian basal metabolism, like body temperature, is a character-
istic not modifiable for climatic adaptation; and individuals of three
species (Vulpes vulpes, L'rethizon dorsatum, Tamiasciurus hudsoni-
cus) did not change basal metabolism significantly in passing from the
cold subarctic winter to the mild weather of summer at Anchorage
(L. Irving, H. Krog, and M. Monson, 1955).

After white laboratory rats have been exposed for a month to air
about 0° C. their resting metabolism is elevated some 60 percent above
the level characteristic of them when they are kept in a warm labora-
tory. This enabled them to survive cold better than the rats from a
warm room, but their reactions to low temperatures were not as well
balanced nor as effective as those of wild brown rats, living out of
doors in the Alaskan winter, whose basal metabolism did not differ
significantly from that of normal white rats. Although the white rats
acquired some extra tolerance of cold with their elevated resting meta-
bolic rate, it appeared that this adjustment resembled the physio-
logical alterations that compensate individuals for pathological con-
ditions or unnatural stress but which could not be regarded as a natural
adaptation proper for a race of animals (H. Krog, M. Monson, and
L. Irving, 1955).

342 U. §. NATIONAL MUSEUM BULLETIN 217

I view with suspicion of harmful consequences the prolonged expo-
sure of mammals to such cold as they can only meet by metabolism
which persistently stands high above the basal level. During periods
of activity mammals can regulate body temperature by utilizing the
heat which they produce above the basal level. Under experimental
conditions they regulate body temperature well for some hours in air
much colder than their critical temperature. Both the degree of cold
and apparently the duration of its tolerance are, however, limited.
During experimentation we have found that the time of endurance
of arctic winter cold by small arctic mammals is limited to a few
hours. It appears likely that animals of smallest size, having also the
higher critical temperatures, must soonest expend the metabolic
reserves available for activity. Apart from the question as to whether
the mammalian metabolic processes can run normally under such
stress of cold is the question of the stress upon the economy of nutri-
tion, for neither food nor time to obtain it are commonly disposable
except with limited possibility of expansion.

The Barrow glaucous gull did not increase its metabolism at
—30° C. and by extrapolation it is safe to locate its critical tempera-
ture as low as —40° C. It is probable that arctic gyrfalcons and
ravens have low critical temperatures, and it is reasonable to suppose
that arctic rock ptarmigan weighing 400 grams are about the smallest
birds with a critical temperature as low as arctic winter cold, for an
arctic Alaskan jay weighing 60 grams needed to elevate its metabolism
in air colder than —3° C. Asan arctic resident the jay had every in-
centive from its habits and environment to enlarge its insulation if it
were physically possible.

It is estimated that the smallest species of birds which can carry
insulation completely adapting them to arctic cold are about %49 the
weight of the smallest species of mammals fully adapted by their in-
sulation for arctic cold. Many northern and arctic birds weighing less
than 400 grams are constantly abroad in the coldest arctic weather (see
p-. 94), nor have I discovered habits which could give them shelter
as a substitute for their deficiency of bodily insulation. During much
of the arctic winter it would seem that they must exceed the basal
metabolic rate normal to homoiothermous animals of their size. In
this respect they appear not to be fully adapted to arctic cold. John
Steen (1958) has demonstrated that small Norwegian wild birds have
critical temperatures so high that through most of the winter they
cannot relax below twice normal metabolic rate.

Returning to view the relation between critical temperature and
normal geographical range, it appeared that the tropical mammals
and birds had critical temperatures about conforming to the climate
in their natural range (Scholander, Hock, Walters, Johnson, and

ARCTIC METABOLIC ECONOMY 343

Irving, 1950). A red fox and porcupines captive at Anchorage had
critical temperatures closely conforming with the mean environ-
mental temperature in winter and in summer, for by changing the
length of their fur they suited their metabolism to the temperature
of the season in the locality where they lived. Their adaptive varia-
tion in insulation shows that seasonal and geographical variation in
critical temperature in individuals of some species of mammals
correlate reasonably well with the climate to which they are exposed
(L. Irving, H. Krog, and M. Monson, 1955).

We were surprised, however, to find that the critical temperature
for a brant (Branta nigricans) and a fish crow (Corvus caurinus)
did not change seasonally. The crow’s critical temperature in winter
and summer at Anchorage corresponded to the January mean in
the warmer climate of Seward, where it had been captured from
among birds resident in the coldest part of their usual range. When
forced to remain in the colder winter of Anchorage, the critical tem-
perature of the crow and migratory brant remained the same as the
mean temperature of the warmer winter on the natural wintering
ground of the species. Neither bird showed any modification of its
critical temperature with season or with its exposure to a winter in
Anchorage much colder than it would naturally encounter.

There are other indications that under domestication birds do not
adjust their critical temperature to the climate in which they are
placed by man. Domestic geese at Strasbourg and Belgrade had
critical temperature near 0° C., according to the figures of Terroine
and Trautmann (1927) and Giaja (1931). The critical temperature
of domestic geese was above the midwinter temperatures of central
Europe, but birds of the genus Anser would seldom encounter below
freezing temperatures in their natural range, which usually extends
only as far north as water remains unfrozen.

Domestic fowl (Gallus) (Terroine and Trautmann, 1927; Mitchell
and Haines, 1927) and pigeons (Columba) (Kayser, 1930; Terroine
and Trautmann, 1927; Falloise, 1900) have high critical temperatures
in the temperate climate of central Europe and North America.
Domesticated fowl are considered to be derived from wild ancestors
living in warm climates and they appear to have retained the critical
temperatures appropriate to tropical birds. Nevertheless, fow] live
with some shelter in Alaska and we have kept pigeons outdoors dur-
ing winter in Barrow. In captivity the birds have an unlimited
supply of food available without effort on their part and they are
protected by cages from being disturbed. While relieving an animal
of the need for exertion to obtain food does not mitigate the cold,
it may reduce the effort necessary for existence so that the full natural
influence toward evoking climatic adaptation is lacking in captivity.

4694966023

344 U. S. NATIONAL MUSEUM BULLETIN 217

Although we have not yet found any birds having the adaptability for
seasonal cold which shows so clearly in the seasonal changes in the
fur of mammals, it does appear that birds of some arctic species are
adapted to cold, as is shown by their low critical temperatures.

Physiological Analysis of Insulation

Arctic land mammals, if they are large enough, are well protected
from cold by the thick covering of fur characteristic of land mam-
mals adapted to arctic life, and in some eases the ability to produce
this thick fur is an inherited character not determined by the environ-
ment. Some breeds of dogs, like the German shepherd can thicken
their fur in northern cold weather. Others, like short-haired pointers,
do not noticeably thicken their coat in cold seasons or when trans-
ported to the Arctic. Collie dogs in warm climates still have thick
fur and arctic sled dogs retain thick coats when taken to warm
climates. So the climatic adaptability of fur length varies among
the breeds of dogs selected under domestication.

We found seasonal adjustment of insulation to environmental tem-
perature in the arctic red fox and porcupine (L. Irving, H. Krog,
and M. Monson, 1955). In summer fur the red fox, with a critical
temperature of 8° C., was adapted to an environment only 30° colder
than its body. In winter at Anchorage its critical temperature was
—15° C., or 53° colder than its body. Arctic white foxes in winter,
with a critical temperature of —40° C., have insulation adaptive to
a temperature about 80° colder than their bodies. The winter in-
sulation of the arctic white fox and the summer insulation of the
red fox at Anchorage compare as 80 to 30. Some good-sized tropical
mammals had critical temperatures only 10° colder than their bodies,
and the effectiveness of their insulation compared with the winter
arctic white fox was as 10 to 80. The difference in these examples
adapted by natural distribution can be related to the thickness of
fur, a condition which varies with the climates in which they occcur
(Scholander, Walters, Hock, and Irving, 1950). |

It is quite possible for large arctic mammals to have fur ten times
thicker than that of tropical animals without hindering their move-
ments. By measurements of heat conductance through samples of
fur as much as 10-fold differences in physical insulation distinguished
the fur of well adapted arctic mammals from those in tropical Panama
(Scholander, Walters, Hock, and Irving, 1950). The critical tempera-
tures of the arctic glaucous gull and domestic fowl] differ by nearly
70°. The combination of all the physiological processes which make
up animal insulation must differ some 8-fold in these two species.
But arctic gulls do not have a feather cover which is eight times
thicker than that of hens, as would be necessary if thickness of

ARCTIC METABOLIC ECONOMY 345

feathers alone were the reason for the gull’s superior insulation. In
these two examples we must evidently look for other factors which
can modify the escape of heat.

It seems to be a common opinion that the feathers of arctic birds
are much thicker than those of tropical birds, for the effectiveness of
a physical insulator varies with its thickness. By inspection of sys-
tematically related arctic and tropical birds in the museum, I could
not find much difference in the thickness of contour feathers in the
prepared skins from related land birds of the warmest and coldest
regions or seasons.

In winter small arctic birds at rest erect their feathers until they
appear much enlarged and nearly spherical. It would not be possible
for a bird to keep this shape in flight, but I find that the individual
contour feathers of birds from the arctic are distinguishable from
those of a warm climate in having less rigid terminal barbs with softer
barbules containing extended fine processes. William Rowan remarked
to me upon this distinction of arctic feathers that a given mass of
them would retain more air than those from warm climates. When
erected on the bird, the arctic feathers retain air among the soft in-
terlocking barbules, whereas the feathers from warm climates readily
separate.

Taking this characteristic of feathers as a criterion for arctic birds,
I numbered several contour feathers from birds of twelve species of
migratory and resident Fringillidae taken at Anaktuvuk, and arranged
them in order of apparent usefulness for insulation as follows:

1. Pinicola enucleator alascen- Resident in winter
sis
2. Acanthis hornemanni exili- Resident in winter
es
3. ee flammea flammea_ Resident of interior Alaska in winter
4. Plectrophenax nivalis ni- Resident of interior Alaska in winter
valis
5. Calearius lapponicus alas- First arrival date April 29
censis
6. Spizella arborea ochracea First arrival date May 8
. Junco hyemalis hyemalis First arrival date May 9
. Zonotrichia leucophrys gam- First arrival date May 13
belii
9. Passerculus sandwichensis First arrival date May 13
anthinus
10. Passerella illiaca zaboria First arrival date May 19
11. Leucosticte tephrocotis teph- First arrival date May 22
rocotis
12. Calcarius pictus First arrival date May 27
I found that the winter resident birds ranked first and that the mi-
grants followed about in the sequence of their arrival dates in spring.

469496—60——24

346 U. S. NATIONAL MUSEUM BULLETIN 217

Several other judges to whom I explained the idea placed the feathers
in about the same order, so that it appears to have some significance.
It should be noticed that these comparisons were made with the feathers
of birds which had reached their summer condition and when the
arctic weather was warm enough to have the migratory examples pres-
ent for comparison, so that if this is an insulating characteristic of
feathers, it is retained in summer. The dissipation of heat, which
must be important for birds in summer is not facilitated by as much
thinning of the feathers as would anywhere near compensate for the
heat of the arctic summer. The heat is dissipated by some other pos-
tural disposition of the feathers or through other physiological chan-
nels.

Neither geographical nor seasonal variation in feather thickness
clearly adapts the insulation of birds to their climates, although dif-
ferences in structure exist which would permit the feathers to func-
tion by erection when at rest, with various effects useful in different
temperatures.

Fur can be regarded as freely modifiable for insulation, and that
seems to be its common function, but feathers serve primarily for
fight. Birds fly alike in all climates, and the requirements of aero-
dynamics which they encounter are unaffected by temperature. It is
to the specific mechanical and power characteristics of internal
muscular and bony mechanisms, and probably to the metabolic pro-
vision of power, characteristics not related to climate, that the feathers
must conform. Probably little modification of the thickness of a
given bird’s feathers is possible during flight; only when at rest is the
bird free to modify the insulation value, utilizing the finer structure
and regulating the position of the feathers to vary the thickness. As I
have said, however, this adaptive use for insulation is obscured and
perhaps repressed by the primary integration of the feathers into
mechanisms for flight that operate unaffected by climate or weather.

I suspect, too, that the necessity for the contour of diving and
swimming birds to conform to the requirements of the aquatic medium
as well as to flight may limit the climatic modifiability of the thickness
of their feathers for insulation. The combination of these various
functions in the feathers of birds precludes any simplified view that
their main use is for insulation.

Variability of Insulation in Animals

The fur clothing made by Eskimos preserves comfortable warmth
in arctic cold when the wearer is inactive or even at rest, but in a warm
house, or during the sunny part of a warm winter day, adequate
arctic clothing is unbearably hot, and even Eskimos, with all their
skill in the preparation and use of clothing, have not succeeded in

ARCTIC METABOLIC ECONOMY 347

making garments that serve well both purposes. If clothing is warm
enough for rest it seriously hampers heat dissipation during activity.
Arctic animals have solved this problem, for while wearing feathers
or fur sufficient for rest in cold they can endure heat and violent pro-
longed activity without apparent overheating. Just after they have
come out in spring with thick fur arctic grizzly bears (Ursus arctos)
when alarmed can run far up the steep rough slope of a mountain
with undiminishing speed. Wolves and caribou run at high speed
for long distances. Late in winter the sun shining on the mountain
cliffs where mountain sheep (Ovis daili) rest in their thick fur is un-
comfortably warm for a well clothed man. Arctic ptarmigan are the
swiftest flying grouse and make long flights during migration at
high speed.

This flexibility of the arctic animal’s insulation, designated one of
the important characteristics of physiological insulation (Scholander,
Hock, Walters, Johnson, and Irving, 1950), arouses the envy of arctic
man uncomfortably and even dangerously sweating from small
exertion in his clothing. How animals can in effect reduce their in-
sulation without removing their covering, and through what avenues
heat can escape in such variable and well regulated fashion, has been
one of the most interesting problems of physiology in the Arctic.

Sweating effectively regulates the disposal of surplus heat for some
animals in warm climates. In the Arctic, man, physiologically a
tropical animal, sweats violently during exertion, but under heavy
clothing the process is inefficient and not very useful. The ability to
sweat varies among mammalian species and its use is difficult to meas-
ure, but my impression is that it is not much employed by arctic
animals in cold weather, although in warm weather they probably
make some use of it, because at no time have I found their fur or
feathers moist, nor does frost collect on them as it does with such
embarrassment on the clothing of sweating man.

The Canidae utilize evaporative cooling by well regulated panting
in the Arctic, and in winter the cloud of moisture above a dog team,
like the “water smoke” over an open lead in the sea ice, is sometimes
visible at great distance. Caribou, reindeer, and men exhale moisture
which cools them in amounts varying with the amount of air breathed.
But without the well regulated cooling system involved in dogs’
panting, evaporation from the lungs does not serve as a flexible, well
regulated means for disposing of excess heat. With their air sacs birds
have large surfaces, but it is not clear that these surfaces are employed
for regulated cooling. In fact, the whole important subject of regu-
lated cooling for arctic animals is obscure. However, one system for
varying the escape of heat has not until recently received much con-
sideration. In it I can see some interesting uses and important con-
sequences for arctic life.

348 U. S. NATIONAL MUSEUM BULLETIN 217

Heat Loss through the Extremities

Some parts of birds and mammals protrude bare or but thinly
insulated, for in the Arctic, as in the Tropics, free movement of the
extremities is necessary, and certain sensory surfaces upon them must
be fully exposed to the environment which they are to perceive. The
bare foot pads of dogs and porcupines have been found to be near 0°
in cold weather, as have the hooves of caribou and the feet of gulls
and several other arctic birds (L. Irving and J. Krog, 1955). Noses
of mammals were also found to be cold.

The extremities derive most of their heat through the circulation
from the body, so that their low temperature is not by itself evidence
that the extremities are thereby saving heat. In the legs of gulls
(Larus glaucescens) a rapid decline in tissue temperature was found to
occur along the tibia under a thick covering of feathers, and the loca-
tion of the temperature change suggested that heat was not escaping
through the thick feathers, but that it was transferred from warm
arterial blood to the venous blood returning cool from the foot (L. Irv-
ing, 1951; L. Irving and J. Krog, 1955).

In the legs and noses of arctic mammals and in the legs of other
arctic birds were found temperature gradients similar to those ob-
served in the gulls. The extremities of mammals possess vascular
arrangements which appear suitable to act as heat exchangers for
the conservation of heat (Scholander, 1955, 1958). The bare tail of the
muskrat is cold, and aquatic mammals, in order to reduce their loss of
heat to the surrounding cold water, may have such vascular schemes
for heat conservation extending over as much of their surface as is
bare. These vascular heat exchangers probably vary in effectiveness
and in morphology among the species of birds and mammals.

That a vascular heat exchanger can change from a conserver to a
dissipator of heat is shown by the occurrence of temperatures almost as
warm as the body in the extremities of animals active in cold air
(L. Irving and J. Krog, 1955). In fact, given an efficient vascular
heat exchanger, the variation in surface temperature is a direct measure
of its usefulness in the regulation of heat loss. The range of variation .
in surface temperatures known to be possible for arctic animals is:

At the surface of the central part of the body 2°
At the surface of the skin over the body proper 6°
At the surface of the bare extremities 40°

By effecting these changes in temperature the “vascular insulation”
can vary the rate of heat conductance from a unit area of surface of
each of these regions in the relation 1:3:20. Where the bare surface
is large, as in the gull’s feet, there is provided an important component
in the total variable insulation of animals. The only physiological

ARCTIC METABOLIC ECONOMY 349

insulation available to the hairless surfaces of swine (L. Irving, 1956),
for example, or of aquatic animals lies in the vascular system of in-
sulation (L. Irving and Hart, 1956), for the insulation of fat and
tissue, while of some significance for heat conservation, is invariable
and is of no value in regulating dissipation of heat.

In the physiological insulation of the body, the vascular system can
have its greatest effectiveness at the freezing temperature, for the
tissue cannot become much cooler without freezing, and it provides
one of the means for regulated variation of the superficial insulation
on the extremities, unprotected by thick fur and feathers.

Adaptation of Tissue Function to Low Temperatures

In order that the extremities may be utilized in the economy of
animal heat their tissues must keep in normal operation such functions
as sensitivity at temperatures too cold for the tissues of the homoio-
thermous body to survive. An example of the adjustment of warm-
blooded tissues to functioning while cold was provided in the demon-
stration that the metatarsal portion of the peroneal nerve of herring
gulls (Larus argentatus) continued to show excitation and conduction
at 8° C. The central part of the nerve from within the warm body
failed at a higher temperature. These gulls had been kept out of doors
in winter at Boston. When they were kept in a warm room and with
warm water to swim in, the distal parts of the peroneal nerves failed
at about the same temperature as the central parts. Evidently the
lowest temperature at which these peripheral nerves will function
is modifiable by environmental temperature. It is, therefore, a clear-
cut adaptation which can be anatomically located in the terminal fiber
from centrally located nerve cells and can be defined in terms of
electrical measurements (Chatfield, Lyman, and L. Irving, 1953).

The homologous nerves of domestic hens kept with the gulls showed
no modifiability of their peripheral nerves by cold, nor were the legs
and feet of the hens cold, as were those of the gulls. It is interesting to
consider that these domesticated fowl, derived from tropical ancestry
appear to retain, along with their high critical temperature (table
18), legs and feet scarely adapted to cold climates.

The properties of some substances of warm-blooded tissues, such as
the high-melting-point fats common in the mammalian body, are
critically affected by cold. Fats from within the body of caribou melt
near 50° C. At body temperature, which is about 10° cooler, the fats
must be sufficiently fluid for normal functions. But at 0° C. these
fats are brittle hard. Since the marrow fat from the distal parts
of the caribou’s leg bones is found to melt at 10° C. it seems that
this distinction of peripheral from body fats would allow for the

350 U. S. NATIONAL MUSEUM BULLETIN 217

needed plasticity at the cold temperature often observed in the pe-
ripheral tissues (L. Irving, Schmidt-Nielsen, and Abrahamsen, 1957).

A similar gradient of melting points was observed in the fats from
the legs of herbivorous reindeer and porcupines, as well as in the
carnivorous fox, dog, and wolf from arctic Alaska. Among the large
arctic mammals it appears common that fats of low melting point
are selectively deposited in the parts of extremities where the tissues
are known to be cool. But a similar distribution of low melting fats
has also been found in the legs of cattle from temperate regions, and
the distribution of fats in the leg of a Panamanian brocket deer
(Mazama americana) was found to be identical with that observed
in homologous bones of arctic herbivores and carnivores. Selective
distribution of fats appears useful or even essential for arctic life,
but since it also occurs in temperate and tropical regions, it cannot
be an adaptation recently evoked for arctic life nor can it be a bad
condition for life in a warmer climate.

In the deposition of fats, selection, according to melting point,
takes place differentially in different anatomical parts. Since the
difference between a high and low melting fat is commonly based
upon the proportion of saturated to unsaturated fatty acid in the
natural mixture of fats, and since change in hardness is often ef-
fected by the relative amounts of oleic and stearic acids, which differ
only in hydrogenation, the difference in properties of fats is ascrib-
able to a biochemical step in fat synthesis.

Biological Significance of Tissue Modification

The composite function of insulation, described in terms of metab-
olism and temperature, shows, in proper physiological terms, that
the common metabolic economy of warm-blooded life in all climates
can be maintained in arctic cold. By comparison with tropical forms
adaptation to cold shows very clearly among arctic animals. But
this kind of adaptation does not provide explicit characters, by the
selective inheritance of which arctic races might become naturally
differentiated. The composite physiological functions are useful to
the animal, and the view of them is useful for the physiologist, but
they are not expressed in terms useful for the natural philosopher
speculating upon the methods which have established the adaptive
differentiation of animals. Hair, fur, and feather structures, on the
other hand, are visible characters produced by tissues, and inheritance
can be observed in them.

ARCTIC METABOLIC ECONOMY aor

The adaptability of the herring gull’s nerve to function in cold,
for example, is a necessity for the normal operation of the cold feet
of the gull, and it was not found in the nerves of hens. This condi-
tion is an important character of the composite insulative function,
and since it can be localized within a portion of a nerve cell and is
measurable by several physical dimensions, the systematic and geo-
graphical relations of animals might be nicely definable on this basis.
A character of this sort could be viewed as a unit in a system of in-
heritance upon which natural selection could operate.

Similarly, the selective deposition of low melting fats in distal
parts of appendages follows a good morphological pattern. It is at-
tributable to the biochemical reaction of dehydrogenation, and is ac-
cordingly distinguishable in the tissue chemistry. It is not demon-
strated as an adaptation to arctic life, but it represents the type of
modification which tissue substances of warm-blooded animals must
undergo to suit them for operation when cold. It is also the sort of
character which might turn out to suit the terms of genetics.

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10.—Retrospect

Observations during 12 years in northern Alaska have confirmed
my original opinion that Anaktuvuk Pass would be an ideal location
for studying the adaptation of animal populations to arctic life.

The Pass acts as a funnel through which move the bulk of the ter-
restrial vertebrate fauna of the region each spring and autumn, and
it also provides breeding sites and environment for many species.
It was the easily visible movement of birds and the obvious nature
of their biological purposes that lead me to deal primarily with
them, and to study them in other localities which, together with
Anaktuvuk, would illustrate their adaptation to all of interior arctic
Alaska and Yukon. In the prosecution of these studies, the native
peoples of the Arctic have played an important role. Their hos-
pitality has greatly eased the burden of working in remote regions,
while their extensive knowledge of the movements of the various
birds has helped me to distinguish the regularity common to popula-
tions from the deviations of occasional individuals.

I have found by observations in the field that the physiological
condition of birds can be correlated with season and behavior. Many
arctic nesting populations of migratory birds arrive fat after rapid
flights from distant wintering places. Each species arrives on its
own schedule. The individuals are in similar physiological prepara-
tion for breeding; they promptly separate into pairs and produce
young which grow to adult size; and then as individuals the parents
and young rejoin the populations ready to migrate southward.
Within each population the physiological states of individuals are
synchronized and the dependent social processes keep each population
distinct and in order, with little apparent conflict.

It is evident that the cycle of migration and nesting is physiologi-
cally anticipated and that the anticipatory processes are correlated to
meet the normal progress of the seasons. Normal seasonal cycles
vary geographically and have changed even in historic times. As
demonstrated in the early chapters, the geographical and temporal
extent of my observations have made possible an analysis of the kind
and degree of correlation that must exist between the lives of birds
and the environment in which they live. For many parts of the
world, the factors involved in such an analysis are so numerous and
their variations often so subtle that it would be difficult to develop

353

354 U. S. NATIONAL MUSEUM BULLETIN 217

a properly analytic approach. In the Arctic the situation is simpli-
fied. The number of species is small, none are passing through on
their way to higher latitudes, and the entire avifauna faces measur-
able and apparently irresistible external influences—a short summer,
extreme seasonal changes in temperature, and a resulting variation in
food supply. These factors impel arctic-nesting birds to regulate
their lives with strict regard for time and place.

The known geological history of arctic Alaska, moreover, indicates
that the populations of migratory land birds now breeding there
‘could not have existed in their present ranges until after the last
extensive glaciation had subsided about 10,000 years ago. This im-
plies that their present migratory routes are recent and that the
characters by which many arctic nesting populations are distinguished
taxonomically must have developed within that time span.

I have found no evidence, however, that the taxonomic characters
distinguishing these arctic population of birds are adaptive responses
to their arctic environment. Apparently these characters are the
result of isolation of the breeding populations. Such isolation could
lead to the production and establishment. of characters which are
ecologically indifferent and not necessarily adaptive—which survive
because they impose no important hardship on their bearers. These
characters may result from variability in a limited gene pool, such
as would be available to a fragment of a species isolated regularly
during the reproductive season, and they would not have become
established except under the special condition of isolation. In each
of several interglacial periods since the Pleistocene, new arctic popu-
lations of the several species of birds must have been reestablished
without essential modification, and it would appear that adaptability
to arctic conditions may have remained latent in many species during
glacial periods when it could not be fully expressed. Such cases do
not argue against the general probability of adaptive evolution.
They do illustrate the fact that visible characters distinguishing geo-
graphical races do not necessarily demonstrate an ever-present
progress toward adaptive evolutionary modification. The develop-
ment by warm-blooded creatures such as birds of an adaptability to
the rigorous demands of arctic conditions must have taken place
long ago, or it would not have been possible for many of these species
dwelling in more temperate latitudes to have extended their ranges
northward in the last post-glacial period.

Our studies have demonstrated in physiological terms the nature
of some of this adaptation to cold. Although the Arctic may seem
unfavorable to the human observer, arctic animals are adapted to
cold by insulation and behavior, so that they can, in most cases main-
tain their normal body temperature without expending special

RETROSPECT aap

metabolic effort. This physiological adaptability obviates environ-
mental stress. No animals, least of all small birds, can act in conflict
with arctic conditions. Individual and social behavior must direct
animals to localities, and at those seasons, when conditions are well
within the physiological tolerance of the population.

Our studies are still in progress. From them we can expect
continuing contributions to our knowledge of animal adaptation to
the arctic environment. Some of this knowledge, we can hope, will
have special meaning for the citizens of our newest State, Alaska,
and for all others who dwell in regions of intense cold.

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Appendix
(See chapters 2 and 8, pp. 322-324)

The weights, size of testes or ovaries, and fatness of common migra-
tory birds at Anaktuvuk are shown in the following figures, of which
21 to 32 (top) are for males and 32 (bottom) to 36 are for females.

On the left-hand ordinate weights have been referred to base 100 as
the average of all weights of aspecies. On the right-hand ordinate is
shown weight in grams.

Testes and eggs are graded as big when near the maximum size for
the species.

Fatness ranges from fat (F), medium fat (MF), little fat (LF), to
very little fat (VLF).

Explanation of symbols used within the plot:

(m.) average weight

(s.d.) standard deviation

(c.v.) coefficient of variation=s.d. x 100%
(F) mean of dates of first migrants’ arrival
(L) date ofstart of egg laying

369

370

EMPTY FOLLICLES, | EGG L Oo!

EMPTY FOLLICLE,

EGGS

EeGsS

EGGS

NESTLINGS

NESTLINGS

NESTLINGS

15 MAY 20

1 EGG LO

L

25 30 JUNE 4 9 14

MAXIMUM
TESTIS

MEDIUM
FAT

AVERAGB
WEIGHT
CV 2 65%

U. S. NATIONAL MUSEUM BULLETIN 217

Ficure 19.—Schedule and condition of Zonotrichia leucophrys gambelii during the breeding
cycle at Old Crow. Schedule is estimated from the date of observation (marked by
large open circle), allowing 1 day interval between each egg laid from date of first (L) to
date of beginning of incubation (I), 12 days to hatching (H), and 9 days to fledging (F).

APPENDIX 371

EGGS

EGGS

EGGS

EGGS

NESTLINGS

NESTLINGS

FLYING

FLYING

IS5MAY 20 25 30 JUNE 4 9 14 19

MAXIMUM
[e)
TESTIS.

_MEDIUM
FAT

e
AVERAGE
WEIGHT
CW 2 6-8 %

Ficure 20.—Schedule and condition of Passerella iliaca zaboria during the breeding cycle
at Old Crow. Schedule is estimated from date of observation, marked by large open
circle, allowing 3 days from laying of first egg (L) to date of beginning of incubation (I),
12 days to hatching (H), and 10 days to fledging (F).

372 U. S. NATIONAL MUSEUM BULLETIN 217

WEIGHT 10 MAY 20 31 JUNE 10 20 30 JULY 10 20 GRAMS

{20

TESTES 76g. cv.8.9%

BIG

MEDIUM

VLF

Ficure 21.—Pintail, @ (Anas acuta).

APPENDIX

WE IGHT 10 MAY 20 31 JUNE 10 20 30 JULY.

°

80 AYTHYA MLN.
m.9329 s4.63 9. ov 6.7%

VLF

10 MAY 20 31 JUNE 10 20 30 JULY
120
110
100
30

80 PLUVIALIS D. D. 4

. $:4.10.8g. c.v. 7.5%

TESTES m. 143.79. $:4.10.89. c.v. 7.5%

BIG °
co

VLF

10

10

20

20

373

900

800

Ficure 22.—Top: Greater scaup, o' (Aythya marila nearctica). Bottom: American golden

plover, & (Pluvialis dominica dominica).

374 U. S. NATIONAL MUSEUM BULLETIN 217

IGHT 10 MAY 20 31 JUNE 10 20 30 JULY 10 20

HETEROSCETUS |.
m. 100.99 Cv. 126%

10 MAY 20 31 JUNE 10 20 30 JULY 19 20

100

2S ° 70

TOTANUS F

oe m805¢ 5.46.79, cv.83 %

TESTES
BIG

MEDIUM

VLF

Ficure 23.—Top: Wandering tattler, o (Heteroscelus incanum). Bottom: Lesser yellow-
legs, o' (Totanus flavipes).

APPENDIX 375

WEIGHT 10 MAY 20 31 JUNE 10 20 30 JULY 10 20 GRAMS
Se
120
*- 100
110
$0
100
80
$0
EROLIA M,
80 m.86.19. sd7.9g9 cv91%

VLF

10 MAY 20 Si JUNE 10 20 60 JULY 10 20

°
eu EROLIA B. =e 3
m 88.89 6.43959 cvl02%

VLF

Ficure 24.—Top: Pectoral sandpiper, o (Erolia melanotos). Bottom: Baird’s sandpiper,
o& (Erolia bairdit).

376 U. 8. NATIONAL MUSEUM BULLETIN 217

t 10 MAY 20 31 JUNE 10 20 30 JULY 10 20 GRAMS

EROLIA MINUTILLA (M)-
m. 20.39 sd.l.7g. ov.8.1%

10 MAY 20 31 JUNE 10 20 30 JULY I0 20

GRAMS

WEIGHT
lo
120

10 Jo
100 100
904. a ; ° 190°
LIMNODROMUS Ss (M)
80 Ree eee 6
Sas m 99.89. 3475 9, os7.5 %
BIG

VLF = : a0

FIGURE 25.—Top: Least sandpiper, o (Zrolia minutilla). Bottom: Long-billed dowitcher,
o (Limnodromus scolopaceus).

APPENDIX 377

WEIGHT 10 MAY 20 31 JUNE 10 ‘ 20 30 JULY I0 20 6RAMS

VLE

WEIGHT 10 MAY 2O 31 JUNE 10 20 30 JULY I!10 20 GRAMS

COBIPES Ie
m31.9g9 sd1.8g cv.5.7 %

Figure 26.—Top: Semipaimated sandpiper, o& (Ereunetes pusillus). Bottom: Northern
phalarope, o& (Lobipes lobarus).

378 U. S. NATIONAL MUSEUM BULLETIN 217

T 10 MAY 20 3! JUNE 10 20 30 JULY IQ 20

90
EREMOPHILA A.A.

80
TESTES m. 36.69. $s.427q cv.7.4%

BIG .
MEDIUM

SMALL

VLF

MAY 20 31 JUNE 10 20 30 JULY 10 SEPT.

WEIGHT 10
%o
120

110

1090

80 TURDUS MM..
m.786g.s.46.1g9. cv.7.8 %

LF 000 0

VLF pcs co
a apa T See so mt tag) ni

Ficure 27.—Top: Arctic horned lark, o (Eremophila alpestris arctica). Bottom: Robin,
S (Turdus migratorius migratorius).

APPENDIX , 379

WEIGHT 10 MAY 20 3t JUNE 10 20 30 JULY 10 20 GRAMS
“lo

120

OENANTHE O. O.
m.23.99.8.d.2.5 9. cv lO %

WEIGHT 10 MAY 20 St gUNE 10 20 30 JULY 10 31 3 GRAMS

100
90
80 MOTACILLA FT.
TESTES mM™1639 %&'39 8.2%

81G 00 °

VLF

Ficure 28.—Top: Wheatear, o (Oenanthe oenanthe oenanthe). Bottom: Yellow wagtail
o' (Motacilla flava tschutschensis).

469496—60——26

380 U. S. NATIONAL MUSEUM BULLETIN 217

WEIGHT 10 MAY 20 31 JUNE 10 20 30 JULY 10 20 GRAMS

ANTHUS S.R. °
m2!1.6g¢s.d1.8 9 cv83%

(ome)
90 3

60
TESTES
BIG : °

VLF

ACANTHIS H.E.
a ml279. 6.41029 cv80%

BIG 00 000 0900 5

VLE

Ficure 29.—Top: Water pipit, o (Anthus spinoletta rubescens). Bottom: Hoary redpoll,
oO (Acanthis hornemanni exiltpes).

APPENDIX 381

WEIGHT 10 MAY 20 31 JUNE 10 20 30 JULY 10 20 GRAMS

100

30
AGANTHIS F. F.

wedee o m!I2.9g sdl.lg cv84%

BIG

co 00

VLF

WEIGHT 10 MAY 20 St JUNE 10 20 30 JULY 10 20 GRAMS

°
SPIZELLA A. O.
m.1849. s.d.1.1 g. v5.9 %

oo °

Ficure 30.—Top: Common redpoll, f (Acanthis flammea flammea). Bottom: Tree spar-
row, do! (Spizella arborea ochracea).

382 U. S. NATIONAL MUSEUM BULLETIN 217

WEIGHT 10 MAY 20 31 JUNE 10 20 30 JULY 10 20 GRAMS

100 25
90
80 ZONOTRIGHIA L.G,

TESTES m.25.6g. sd 1.79 cv.6.7% 20

008 ° o fore)

VLF

10 MAY 20 Si. JUNE 10 20 3O JULY !0 20

°
90 oO
AG GALGARIUS L. A.

m 28-69 sd 24g. cv.8:-4%

VLF

Ficure 31.—Top: Gambel’s wibioeaeened sparrow, co! (Zonotrichia leucophrys gambelit).
Bottom: Alaskan longspur, o! (Calcartus lapponicus alascensis).

APPENDIX 383

WEIGHT 10 MAY 20 31 JUNE 10 20 30 JULY 10 20 GRAMS
Ye -
120 35
110
30
100
90
25
80 GALCARIUS P
TESTES m.28.ig.s.¢15 9 cv5.3%

Sue ge ccccco aD

VLF

WEIGHT 10 MAY 20 31 JUNE 10 20 30 JULY 10 20

120

110

100

80 PLUVIALIS O.D.
m. 145.8g.°s'd.10.5g. ¢.v.7,2 %

VLF

Ficure 32.—Top: Smith’s longspur, @ (Calcarius pictus). Bottom: American golden
plover, @ (Pluvialis dominica dominica).

384 U. S. NATIONAL MUSEUM BULLETIN 217

WEIGHT 10 MAY 20 Si JUNE 10 20 30 JULY 10 20 GRAMS

° fe}

HETEROSCELUS |. (F)
mil6.1 9. $49.7 9 ¢v8.3 %

EROLIA BAIRDI! (F)
m 39.19 s4.4.1 g cvl0.4%

(2)

Ficure 33—Top: Wandering tattler, 9 (Heteroscelus incanum). Bottom: Baird’s sand-
piper, 9 (Erolia bairdit).

APPENDIX 385

WEIGHT 10 MAY 20 31 JUNE 10 20 30 JULY 10 20 GRAMS

Se
120
110 24
100 22
90 20
, EROLIA MINUTILLA
8 m.2L99 s.d.2.7q. cv.12.4 %

EGGS

VLF

10 Y 20 3! JUNE 10 20 30 JULY I

LOBIPES L.
m.35.29.s$¢d33g9 cv95%

Ficure 34.—Top: Least sandpiper, 2 (Erolia minutilla). Bottom: Northern phalarope
Q (Lobipes lobatus).

386 U. S. NATIONAL MUSEUM BULLETIN 217

WEIGHT 10 MAY 20 31 JUNE 10 20 30 JULY 10 20

° oe)
100- eo 8 130
feo) 5
907 . Po 12.0

F
AGANTHIS H.E.
m 128g s.d.0939. cv.7.2 %

VLF

WEIGHT 10 MAY 20 31 JUNE 10 20 3$O JULY 10 20 GRAMS
25

ZONOTRICHIA L G. 20
m 2339

VLF

Figure 35.—Top: Hoary redpoll, 9 (acanthis hornemanni exilipes). Bottom: Gambel’s
white-crowned sparrow, 9 (Zonotrichia leucophrys gambelit).

APPENDIX 387

10 MAY 20 31 JUNE 10 20 30 JULY 10 20°) )- eaages

CALCARIUS L.A.
m 26.09. s.d.2.19. cv.8.1 %

WEIGHT 10 MAY 20 31 JUNE 10 20 3O JULY t0 20 GRAMS

CALCARIUS PB
m.24.99

Ficure 36.—Top: Alaskan longspur, @ (Calcarius lapponicus alascensis). Bottom:
Smith’s longspur, 9 (Calcarius pictus).

iS ie Rome
all oinab aay panes tee

Sibi hike Auisuicel nea WiksStionscLal

se he snp apm mht hl 9 it a tay Sameer tibet Se ep el a att ea ae a te fbn

a

ARO nobis er ee ew yi Raebeanl ae, eae Tad Pe VEE elie paar Dcenc Kia hci

A i a oye
: ican! mine antee ttbag a aieSo “

ince bet fh SREY eon seaiees Sanita Sint a ig acm i = neato eens “6

teal Sates hearers anh enton a re

bah nnn ob,

Index

(Page numbers of principal entries in italics.)

A

Abel, Albert, 155, 207
Acanthis flammea, 107, 113, 114, 280, 307
flammea flammea, 115, 116 (table),
231 (table), 233 (table), 248, 264,
270, 288, 322, 345, 381 (graph)
flammea holboellii, 248
hornemanni, 280, 313
hornemanni exilipes, 112, 113, 115,
116 (table), 230, 231 (table), 232
(table), 248, 249, 264, 270, 288,
322, 323, 345, 380 (graph), 386
(graph)
Accipiter gentilis, 136, 180
gentilis atricapillus, 46, 247, 251,
253, 264
striatus, 180, 247
striatus velox, 136
Accipitridae (hawks, harriers), 46, 136,
180, 247, 259
Actitis macularia, 66, 193 (table), 194,
247, 307, 317
acuta, Anas, 37, 170, 245 (fig.), 246, 288,
303, 313, 322, 372 (graph)
adamsii, Gavia, 30, 246, 267, 288, 316
Adaptation to cold, 280, 302, 349
Adlakuk, Kobuk name for wandering
tattler, 138
aedon, Troglodytes, 309, 338
Aegolius funereus, 142, 207
funereus richardsoni, 85, 247, 253,
264
affinis, Aythya, 40 (table), 41, 135, 172,
173, 246, 260, 287 (fig.), 288, 307
Agelaius phoenicus arctolegus, 257
Ahalik, Nunamiut name for oldsquaw,
42
Ahaliknak, Nunamiut name for Western
harlequin duck, 43
Ahaluk, Indian name for oldsquaw, 176
Ahgook, Jesse, Eskimo man, 22, 33, 48,
131;,112
Aklaktak, Nunamiut name for buff-
breasted sandpiper, 75
Akpaksruayook, Nunamiut name for
red-breasted merganser, 46

Akpukshookti, Kobukmiut name for
sanderling, 140
alascanus, Haliaeetus leucocephalus, 47,
136, 181, 247, 253, 267
alascensis, Calcarius lapponicus, 123, 231
(table), 242 (table), 248, 259, 266,
269, 288, 303, 316, 322, 323, 345,
382 (graph), 387 (graph)
Lagopus lagopus, 54, 185, 247, 248,
249, 264, 324
Pinicola enucleator, 111, 231 (table),
248,'249, 250, 264, 345
Troglodytes troglodytes, 257
Alaudidae (larks), 88, 144, 209, 247, 259
alba, Crocethia, 76, 140, 247, 252, 267,
288, 316
albeola, Bucephala, 135, 246, 267
albifrons, Anser, 134, 269
albus, Lagopus lagopus, 57, 188
Alcedinidae (kingfishers), 142, 247, 259
Alcidae, 259
Aldrich, John W., and Duvall, Allen J.,
185
algistus, Bubo, 205
Alopex lagopus, 338, 339
alpestris, Eremophila,
(graph)
Amauligak, Nunamiut name for snow
bunting, 128

americana, Bucephala clangula, 174, 246,
267
Mareca, 38, 171, 246, 288
Mazama, 350
Oidemia nigra, 247, 267
americanus, Lepus, 338, 339
Oreamnus, 339

amnicola, Dendroica petechia, 110, 147,
148, 223, 224 (table), 226 (table),
248, 253, 269
Anaktuvuk Pass, 15-28, 17 (map)
birds of, 28-130, 248-255
status of birds at, 246-248 (table)
U.S. Post Office at, 15 (fig.)
weather and climate, 18 (fig.), 19
(fig.)

144, 313, 378

389

390

Anaruk kiviruk, Nunamiut name for
dippers, 95
Anas acuta, 37, 170, 245 (fig.), 246, 288,
303, 313, 322, 372 (graph)
carolinensis, 88, 170, 246, 288, 303
platyrhynchos platyrhynchos, 36, 169,
246, 288, 307
Anatidae (swans, geese, ducks), 33, 134,
159, 167, 247, 259
anatum, Falco peregrinus, 51, 183, 247
288
Anderson, Hannah, 127
Anderson, J. L., pilot, 26, 299
Anderson, Rudolph Martin, 30, 107,
128, 253, 256, 260
Anser albifrons, 134, 269
albifrons frontalis, 36, 166, 246, 288,
316
(domestic), 338, 343
Anthinus, 255
anthinus, Passerculus sandwichensis, 117,
231 (table), 234, 248, 269, 288,
345
Anthus spinoletta, 313
spinoletta pacificus, 221, 222
spinoletia rubescens, 107, 221, 248,
269, 303, 322, 380 (graph)
Aphriza virgata, 138, 247, 266
Aquila chrysaétos canadensis, 47, 136,
181, 247, 267, 288, 303
arborea, Spizella, 307
Arctic Alaska, interior of, 4-6
physiography of, 6-7
Arctic coast, birds of, 256-259
arctica, Gavia, 307
arcticola, Eremophila alpestris, 88, 144,
209, 247, 266, 269, 288, 303, 322
arcticus, Picoides, 86, 143, 247, 251, 264
arctolegus, Agelaius phoenicus, 257
Arctomys sp., 335
arctos, Ursus, 347
Arenaria interpres, 63, 138
interpres interpres, 63, 247, 265, 266,
270, 288
interpres morinella, 63, 265
argentatus, Larus, 141, 349
Armstrong, Edward Allworthy, 310
Arny, Samuel A., 90
Asio flammeus flammeus, 84, 247, 288
Atkaruak, Nunamiut name for young
gyrfalcon, 49
Atkinson, Ben, 4
atricapilla, Zonotrichia, 148, 248, 269

U. S. NATIONAL MUSEUM BULLETIN 217

atricapillus, Accipiter gentilis, 46, 247,
251, 253, 264
Parus, 145
Auksruak, Nunamiut name for red
phalarope, 76
auratus, Colaptes, 307
Avikiak, Nunamiut name for Wilson’s
snipe, 65
Avilyuktok, Nunamiut name for surf
scoter, 44
Aythinae (ducks), 167
Aythya affinis, 40 (table), 41, 135, 172,
173, 246, 260, 287 (fig.), 288, 307
marila, 185, 172, 313
marila nearctica, 39, 40 (table),
186, 172, 246, 260, 267, 288, 303,
322, 373 (graph)

B

Bailey, Alfred Marshall, 4, 5, 30, 63, 70,
79, 90, 101, 105, 107, 118, 141,
147, 165, 178, 191, 202, 221, 229,
256, 257

Baird, Spencer Fullerton, 102

bairdii, Erolia, 64, 70, 189, 193 (table),
198, 247, 269, 288, 303, 316, 317,
322, 323, 375 (graph), 384 (graph)

Baldwin, Samuel Prentiss, and Ken-
deigh, Samuel Charles, 325

barrovianus, Larus hyperboreus, 79, 200,
247, 267, 288, 316, 338

Barrow, birds from, 257 (table)

research at, 3-4
Point, 258

Bartholomew, J. G., Dawson, W. R.,
and O’Neill, E. J., 311

Bartholomew, J. G., and Herbertson,
Andrew John, 332

Bartramia longicauda, 65, 138, 247, 253,
269, 317

Bartsch, Paul, 320

baueri, Limosa lapponica, 266, 270

Bear, grizzly, 347

Beaver, 27

bendiret, Falco columbarius, 52, 247, 269,

288

Bent, A. C., 74, 104, 183, 212, 213, 317

Bergmann, Carl, 280

Bergmann’s rule, 280

Bernard, Claude, 310

bicolor, Iridoprocne, 80, 144, 211, 247,
253, 257, 269

INDEX

Birds, land, migrating in Alaska, Yukon
and western Mackenzie, 266 (ta-
ble), 269 (table)

present in winter, 264 (table)
residents of northern forests, 251
(table)

summer, in valley of Anaktuvuk
Pass, 253 (table)

winter, on American Pacific coasts,
267 (table)

winter, in valley of Anaktuvuk
Pass, 249 (table)

Bishop, Louis Bennett, 29, 185, 242, 256,
261

Blackbird, 111, 229

rusty, 111, 229, 254

Blanchard, Barbara D., 328

Bluebird, 97, 146, 216

Bluethroat, red-spotted, 101, 260, 274

Bombycilla garrula pallidiceps, 108, 222,
248, 251, 264

Bombycillidae (waxwings), 108, 222,
248, 259

Bonasa umbellus, 137, 185, 247

umbellus yukonensis, 264

borealis, Colaptes auratus, 85, 142, 207,

247, 253
Nuttallornis, 209, 247, 269
Phylloscopus, 147

boreophila, Spizella passerina, 257

brachyrhynchus, Larus canus, 80, 202,
247, 267, 288

Brandt, Herbert, 127

Brant, Pacific black, 35, 165, 202, 343

Branta canadensis, 34

canadensis canadensis, 263 (fig.)

canadensis parvipes, 164 (table), 165

canadensis taverneri, 34, 134, 162,
164 (table), 165, 246, 252, 265,
266, 269, 288, 316

hutchinsiz, 134

leucopaeia, 34, 134, 165

minima, 134, 165

nigricans, 35, 134, 165, 246,
288, 316, 338, 343

Breeding cycle, adaptation of,

and migration, 305

environmental influence on, 308

of species, 40 (greater scaup),
58 and 189 (Rock ptarmigan),
67-68 (wandering tattler), 61-62
(American golden plover), 75
(semipalmated sandpiper), 97 and

267,

308

391

Breeding cycle—Continued
216 (robin), 120-123 and 236
(white crowned sparrow), 123 and
126 (longspur), 177 and 179
(Scoter), 212 (cliff swallow), 217
(varied trush), 225 (yellow warb-
ler), 239 (foxsparrow)
synchrony of, 236, 239
Bridges, Esteban Lucas, 50
Brody, Samuel, 236
Brooding, 310
Brooks, Winthrop Sprague, 177, 256,
260
Brower, Charles, iv, 5, 257
Brower, Thomas, iv, v, 5, 24, 25, 37,
38, 45, 47, 60, 69, 72, 73, 79,
81, 88, 90, 91, 95, 99, 105, 107,
112, 114, 118, 121, 126, 138,
253, 257,275, 289
Brown, Harry, trader, 132, 257
Bruce, Robert, guide, 156, 158, 163,
174, 181, 204, 205, 207, 209, 220
Bryant, Joe, 181, 207
Bubo algistus, 205
saturatus, 205
virginianus lagophonus,
206, 247, 251, 253, 264
wapacuthu, 205
buccinator, Cygnus, 162
Bucephala, 168
albeola, 135, 246, 267
clangula, 260, 272
clangula americana, 174, 246, 267
islandica, 175, 246, 260, 267
Buckley, John L., 59, 158
Bufflehead, 135
Bumblebees, 21
Bunting, 111, 148, 230
snow, 127, 148, 230, 242, 250, 259
265, 270, 280, 293, 303, 307, 313
Burch, George, 157

Burleigh, Thomas
H. S., 227

Bushtits, 94, 145, 215
Buss, Irven O., 66
Buteo lagopus, 46, 180, 247, 288, 303, 307
lagopus s.johannis, 46
C
Cade, Tom, 46, 59, 90, 166, 252

Calcarius, 230, 255
lapponicus, 313

82, 141,

D., and Peters,

392

Calearitus—Continued
lapponicus alascensis, 128, 231
(table), 242 (table), 248, 259,
266, 269, 288, 303, 316, 322, 323,
345, 378, (graph), 383 (graph)
pictus, 125, 248, 269, 288, 303, 313,
322, 345, 383 (graph), 387
(graph)
calendula, Regulus calendula, 104, 147,
221, 248, 250, 253, 269
Campbell, Jack, 129
Canachites canadensis osgoodi, 53, 186,
247, 251, 264
canadensis, Aquila chrysaétos, 47, 186,
181, 247, 267, 288, 303
Branta, 34
Branta canadensis, 263 (fig.)
Grus canadensis, 60, 190, 247, 269,
288, 316
Perisoreus canadensis, 214
canagica, Philacte, 134
Canidae, 347
Canis familiaris, 339
Cantwell, John C., 145
caparoch, Surnia ulula, 83, 206, 247,
251, 264
Capella gallinago, 260, 307
gallinago delicata, 64,
(table), 247, 288
Caprimulgidae, 259
Caribou, 347
carolinensis, Anas, 38, 170, 246, 288, 303
Pandion haliaetus, 48, 136, 182,
247, 253
carolinus, Euphagus carolinus, 111, 229,
248, 254, 288, 307
Catoptrophorus semipalmatus semipal-
matus, 317
caurina, Megaceryle alcyon, 142, 247,
269
caurinus, Corvus, 338, 343
celata, Vermivora celata, 223, 224 (table),
248, 269, 307
Cepphus grylle mandtiz, 315 (fig.)
Charadriidae (plovers, turnstones, surf-
birds), 60, 137, 191, 247, 259
Charadrius semipalmatus, 60, 191, 247,
288, 303, 304, 307
vociferus, 247
vociferus vociferus, 61, 137, 191, 257
Charlie, Ben, 162, 181, 187
Charlie, Lazarus, 163, 217, 229, 274

192, 193

U. S. NATIONAL MUSEUM BULLETIN 217

Chen hyperborea hyperborea, 36, 167, 246,
267, 288, 316
Chickadee, Alaska, 95, 335
boreal, 215
brown, 145
gray-headed, 216
Yukon black-capped, 86, 94, 96,
112, 145, 250
Chilly cho, Indian name for rusty
blackbird, 230
Chin tettree, Indian name for pigeon
hawk, 184
Chinchityo, Indian name for pintail,
170
Chinechun, Indian name for peregrine,
183
Chit tryo, Indian name for Bonaparte’s
gull, 204
Chittese, Indian name for golden eagle,
181
Chizin, Indian name for bald eagle, 181
Chootzi, Indian name for waterthrush,
228
Chordeiles minor minor, 257
Chul rut tsit, Indian name for sharp-
shinned hawk, 180
Chut lut, Indian name for flicker, 207
Chut rhui chun tsik, Indian name for
rough-legged hawk, 181
Chut-tul, Indian name for
grouse, 185
Chya, Indian name for rock ptarmigan,
190
Cinclidae (dippers), 95, 146, 216, 247,
259
Cinclus mexicanus unicolor,
216, 247, 249, 264

ruffed

95, 146,

cinnamomea, Tringa solitaria, 66, 138,
193 (table), 195, 247, 260, 266,
269, 282, 288

Circus cyaneus hudsonius, 48, 181, 247,
253, 288

Ciiellus undulatus, 334, 339

Citellus sp., 335

clangula, Bucephala, 260, 272

Clangula hyemalis, 41, 175, 246, 267,

288, 303, 313, 316

Climate, and population, 12-14
arctic, 7-12
at Anaktuvuk Pass, 18, 19, 25
Northwestern Alaska, 278
seasonal progress of, 21

INDEX

clypeata, Spatula, 38, 135, 172, 246, 252,
288
coelebs, Fringilla coelebs, 328
Colaptes auratus, 307
auratus borealis, 85, 142, 207, 247,
253
cold, adaptation to, 280, 302, 349
Collins, Henry, 128
Columba (domestic), 338, 343
palumbus palumbus, 328
columbarius, Falco, 53, 184
columbianus, Olor, 33, 169, 246, 252, 267,
288, 316
Contopus sordidulus, 257
Cooke, Wells, W., 204
Cormorant, pelagic, 331 (fig.)
corniz, Corvus corniz, 328
cornutus, Podiceps auritus, 33, 133, 159,
246, 253, 267
coronata, Dendroica, 257
Corvidae, (jays, magpies, crows), 91,
214, 247, 259
Corvus caurinus, 338, 343
coraxz principalis, 92, 214, 247, 249,
250, 264
cornizx cornix, 328
Coues, Elliott, 215
Court, A., Sissenwine, N., and Mitchell,
G., 8., 332
Crane, 60, 190, 191, 252, 313
lesser sandhill, 60
Crocethia alba, 76, 140, 247, 252, 267,
288, 316
Crossbill, white-winged, 117, 230,
251
Crow, 91, 214
Crowberry, 189
Curlew, bristle-thighed, 64
currucotdes, Sialia, 257
cyanocephalus, Euphagus, 257
Cygnus buccinator, 162

D

Dale, Robert F., 297

Dall, William H., and Bannister, H. M.,
66, 102, 137, 162, 166, 167, 174,
175, 179, 184, 199, 233, 271, 272

dalli, Ovis, 347

Dawson, W. R., 334

Deer, Panamanian brocket, 350

deglandi, Melanitta, 43, 135, 176, 247,
267, 288, 303

Melanitia, deglandi, 287 (fig.), 313

234,

393

Deignan, Herbert, vi
Delacour, Jean, 35
Delacour, Jean, and Zimmer, John T.,
134, 164
delicata, Capella gallinago, 64, 192, 1938
(table), 247, 288
Dendrocopos pubescens, 148
pubescens nelsoni, 86, 143, 247, 249,
250, 264
villosos, 143
Dendroica coronata, 257
coronata hooveri, 110, 148, 223, 224
(table) 226 (table), 248, 253,
266, 269
petechia, 147, 268, 307
petechia amnicola, 110, 147, 148,
223, 224 (table), 226 (table), 248,
253, 269
striata, 223, 224 (table), 226 (table),
248, 268, 269, 272, 307
striata lurida, 223, 227
Dicquemare, Philippe, collector, 159,
175, 176
Dicrostonyx torquatus, 339
Dillon, L. S., map from, 279
Dippers, 95, 146, 216, 249, 250
dispars, Lymantria, 281
Distribution, 245-261
changed by glaciation 263 ff.
of species, 167 (duck), 192 (sand-
piper), 221-222 (pipit), 223
(wood warbler), 244 (snow bunt-
ing)
Dixon, Joseph Scattergood, 66, 256, 260,
271, 275
dixont, Melanitta, 177, 178 (table)
Dog, 339, 344, 348
domesticus, Passer, 338
dominica, Pluvialis, 192, 274
Pluvialis dominica, 61, 247, 269,
288, 303, 316, 317, 322, 323, 373
(graph), 383 (graph)
dorsatum, Erethizon, 339, 341
Dowitcher, long-billed, 72, 139, 376
(graph)
Dryas, 21
Duck, 33, 49, 133, 159, 167, 255, 256,
260, 313
eider, 135, 168
greater scaup, 39, 135, 172, 260,
373 (graph)
harlequin, 43, 176
king eider, 128

394

Duck—Continued
lesser scaup, 41, 135, 172, 173, 260,
287 (fig.)
pintail, 37, 41, 170, 171, 179, 245
(fig.), 303, 312, 372 (graph)
scaup, 172, 312, 325
Steller’s eider, 43
spectacled eider, 245 (fig.)
Dunlin, red-backed, 72, 139

E

Eagle, American golden, 47, 136, 181
Northern bald, 47, 136, 181
Eknikauktok, Nunamiut name for Stel-
ler’s eider, 43 ©
Eksrahgotilik, Nunamiut name _ for
Canada goose, 35
elasson, Gavia immer, 29
Empetrum (crowberry), 189
Empidonaz traillii, 208
traillit trailliz, 247, 269
Epilobium (fireweed), 21
Eremophila alpestris, 144,
(graph)
alpestris arcticola, 88, 144, 209, 247,
266, 269, 288, 303, 322
alpestris hoyti, 210.
Erethizon dorsatum, 339, 341
Ereunetes pusillus, 64, 74, 193 (table),
199, 247, 288, 303, 304, 309, 313,
316, 317, 322, 377 (graph)
erminea, Mustela, 340
Erolia alpina pacifica, 72, 189, 247, 253,
267, 288, 316
bairdii, 64, 70, 189, 193 (table),
198, 247, 269, 288, 303, 316, 317,
322, 323, 375 (graph), 384
(graph)
fuscicollis, 70, 247, 253, 316
melanotos, 64, 69, 193 (table), 197,
247, 269, 288, 316, 323, 375
(graph)
minutilla, 64, 71, 193 (table), 198,
247, 288, 303, 304, 322, 323, 376
_ (graph), 385 (graph)
erythrogaster, Hirundo, 257
Hirundo rustica, 90, 247, 253, 269
Esox lucius, 271
Euphagus carolinus carolinus, 111, 229,
248, 254, 288, 307

Euphagus cyanocephalus, 257
evura, Parus hudsonicus, 95, 215

313, 378

U. S. NATIONAL MUSEUM BULLETIN 217

excurbitor, Lanius, 223

exilipes, Acanthis hornemannt, 112, 113,
115, 116 (table), 117, 230, 231
(table), 232 (table), 248, 249, 264,
270, 288, 322, 323, 345, 380
(graph), 386 (graph)

F

Falco columbarius, 53, 184
columbarius bendiret, 62, 247, 269,
288
peregrinus, 136, 307
peregrinus anatum, 61, 183, 247, 288
rusticolus, 182
rusticolus obsoletus, 49, 247, 249, 264
sparverius, 137, 257
sparverius sparverius, 68, 247, 253
Falcon, 49, 136, 182
Falconidae (falcons), 49, 136, 182, 247,
259
familiaris, Canis, 339
farleyi, Parus hudsonicus, 215
fasciatus, Picoides tridactylus, 86, 143,
207, 247, 251, 264
Fat, metabolism of, 326
and migration 116 (redpoll), 123
(longspurs), 232 (grosbeaks),
321, 328
color, 87 (phoebe), 101 and 220
(wheatear), 206 (hawk-owl), 243
(snow bunting)
tissue, adaptation of function to
low temperature, 349
Feinstein, Bernard, vi, 113, 220, 223,
227, 244
Finch, 111, 148, 230
gray-crowned rosy, 112, 131 (fig.),
148
Firth, H. A., 310
fischeri, Lampronetta, 245 (fig.)
flammea, Acanthis, 107, 113, 114, 280, 307
Acanthis flammea, 116, 116 (table),
231 (table), 233 (table), 248, 264,
270, 288, 322, 345, 381 (graph)
flammeus, Asio flammeus, 84, 247, 288
flava, Motacilla, 147, 313
flavipes, Totanus, 64, 68, 193 (table),
196, 247, 260, 269, 288, 303, 304,
307, 374 (graph)
Flicker, yellow-shafted, 85, 142, 207
Flint, Richard Foster, 11, 12, 14

Flocking, 42 (oldsquaw), 98 (robin), 114
243 snow

and 312 (redpoll),
bunting), 319
Flocks, formation of, 312, 313
Fly, black, 21
Flycatcher, olive-sided, 209
Traill’s, 208
tyrant, 86, 104, 208
Fox, Arctic white, 338
red, 343, 344

INDEX

Friedmann, Herbert, v, vi, 34, 63, 118,

121, 134, 148, 228
Fringilla coelebs coelebs, 328

Fringillidae (grosbeaks, finches, spar-

rows, buntings), 79, 88, 111, 112,
148, 230, 231 (migration table),

232, 248, 259, 306, 311

frontalis, Anser, albifrons, 36, 166, 246,

288, 316
Frost, Freddie, 162
Frost, Stephen, 169, 216

fulicarius, Phalaropus, 76, 247, 267, 288,

316, 317
funereus, Aegolius, 142, 207

fuscicollis, Erolia, 70, 247, 253, 316

G

Gabrielson, Ira N., vi, 113, 122
gallinago, Capello, 260, 307
Gallus (domestic), 338, 343

gambelii, Zonotrichia leucophrys, 119,

231 (table), 235 (table), 238
(table), 248, 269, 288, 303, 322,
323, 345, 370 (graph), 382

(graph), 386 (graph)

Gavia adamsii, 30, 246, 267, 288, 316

arctica, 307

arctica pacifica, 31, 149 (fig.), 158,

246, 267, 288

immer, 29, 157, 246, 267, 288

immer elasson, 29

stellata, 32, 159, 246, 267, 288
Gaviidae (loons), 29, 157, 246, 259

gentilis, Accipiter, 136, 180
Giaja, A., 343

Giddings, James Louis, Jr., 5, 258

Glaciation, 7-13, 16, 152, 153, 263, 264,

266, 278, 279
glaucescens, Larus, 348
Gnatcatcher, 102, 147, 221

Godfrey, W. Earl, 94, 95, 215
Godwit, 76, 140, 252, 270

469496—60——27

395

Goldeneye, American, 168, 174, 260

Barrow’s, 175
Goldschmidt, Richard, 281
Goose, 38, 133, 159, 255, 313

Canada, 134, 162, 263 (fig.)

emperor, 134

snow, 36, 163, 167

Taverner’s, 252

white-fronted, 36, 134, 166
Goshawk, 46, 136, 250
Grebe, 32, 133, 159

horned, 32, 33, 133, 159

red-necked, 32, 133, 159

Grinnell, Joseph, 5, 66, 86, 99, 106, 109,
114, 132, 133, 134, 135, 136, 137,
138, 141, 143, 145, 146, 147, 148,

174, 185, 233, 241, 293
Grosbeak, 111, 148, 230, 335

pine, 94, 111, 118, 230, 232, 250

Gross, Alfred, 183, 212
Grouse, 53, 137, 184

Hudsonian spruce, 53, 59, 185

ruffed, 137, 185
sharp-tailed, 59, 137, 184
timber, 59

Gruidae (cranes), 60, 190, 247, 259

Grus canadensis canadensis, 60, 190, 247,

269, 288, 316
Guillemot, 315 (fig.)
Gull, 79, 140, 200, 255, 348, 349
Bonaparte’s, 204
glaucous, 141, 200, 342
herring, 741, 201, 351
mew, 80, 202
Ross’s, 81
Sabine’s, 81, 141
short-billed, 141
Thayer’s, 202, 284
Western glaucus, 79, 80
white-plumaged, 79
guttata, Hylocichla guttata, 257
Gyrfalcon, 49, 182, 249, 250, 335

H
Haldeman, Jack C., 4

Haliaeetus leucocephalus alascanus, 47,

136, 181, 247, 253, 267
Hare, Arctic, 338
snowshoe, 338

Harper, Francis, 57
Harriers, 46, 136, 180
Hart, J. S., 337, 340

396

Hawk, 46, 136, 180
duck, 49, 136
marsh, 48, 181, 253
pigeon, 49, 52, 53, 184
rough-legged, 46, 49, 180, 303
sharp-shinned, 136, 180
sparrow, 53, 137
hemastica, Limosa, 140
‘Henkelman, Carl, v, 24
Heteroscelus incanum, 64, 67, 138, 247,
266, 270, 288, 316, 322, 323, 374
(graph), 384 (graph)
Hibernation, arctic, 335
himantopus, Micropalama, 73, 139, 247,
252, 269, 316
Hirundinidae (swallows), 89, 144, 210,
247, 259, 261, 268
Hirundo erythrogaster, 257
rustica, 144, 145
rustica erythrogaster, 90, 247, 253,
269
histrionicus, Histrionicus, 42, 176, 246,
267, 288
Histrionicus histrionicus, 42, 176, 246,
267, 288
Hock, Raymond, iv, 20, 73, 95, 107, 128,
129, 326, 327, 334, 336
holboellii, Acanthis flammea, 248
holbéllii, Podiceps grisegena, 32, 1338, 169,
246, 267
Homoiothermism, 312, 332
hooveri, Dendroica coronata, 110, 148,
223, 224 (table), 226 (table), 248,
253, 266, 269
hornemanni, Acanthis, 280, 313
hoytt, Eremophila alpestris, 210
hudsonicus, Numenius phaeopus, 64, 64,
193 (table), 194, 247, 267, 288,
316
Parus hudsonicus, 96, 145, 216, 247,
251, 264
Tamiasciurus, 339, 341
hudsonius, Circus cyaneus, 48, 181, 247,
253, 288
Huggins, R. A., 309, 310
Hummingbird, 321, 327
hutchinsts, Branta, 134
hyemalis, Clangula, 41, 176, 246, 267,
288, 303, 313, 316
Junco hyemalis, 118, 231 (table),
234 (table), 248, 253, 269, 303,
345

U. S. NATIONAL MUSEUM BULLETIN 217

Hylocichla guitata guttata, 257
minima, 146, 272, 307
minima minima, 99, 219 (table),
247, 269
ustulata, 146
ustulata incana, 146, 218, 247, 261,
266, 269
hyperborea, Chen hyperborea, 36, 167,
246, 267, 288, 316
hyperboreus, Larus, 141, 276
hypopolia, Petrochelidon pyrrhonota, 212,
247, 269
I

Icteridae (blackbirds), 111, 229, 248,
259
Iklikvik, Numamiut name for fox spar-
row, 122
tliaca, Passerella, 307
llyirgik, Nunamiut name for rough-
legged hawk, 46
immer, Gavia, 29, 167, 246, 267, 288
incana, Hylocichla ustulata, 146, 218,
247, 261, 266, 269
incanum, Heteroscelus, 64, 67, 138, 247,
266, 270, 288, 316, 322, 323,
374 (graph), 384 (graph)
Incubation, 308-309
Ingstad, Helge, 25
interpres, Arenaria, 63, 188
Arenaria interpres, 63, 247, 265,
266, 270, 288
invictus, Lanius excubitor, 108, 248, 269,
288, 303
Iridoprocne bicolor, 80, 144, 211, 247,
253, 257, 269
Trirgik, Nunamiut name for Northern
shrike, 109
Irving, Laurence, 20, 21, 22, 129, 132,
142, 311, 327, 336, 348, 349
Irving, Laurence H., 23, 104, 144, 325
Irving, Laurence, and Hart, J. 8., 349
Irving, Laurence, and Krog, John, 237,
238, 239, 241, 309, 311, 325, 327,
332, 334 (table), 337, 338, 348
Laurence, Krog, John, and
Monson, M. W., 341, 343, 344
Irving, Laurence, and Paneak, Simon,
21, 28, 103, 106, 138, 139, 144,
146, 147, 312, 318
Irving, William, v, 21, 25, 26, 29, 30,
45, 49, 54, 68, 80, 87, 111

Irving,

INDEX

irvingi, Leucosticte tephrocotis, 112, 131
(fig.), 248, 265, 266, 269, 288
islandica, Bucephala, 175, 246, 260, 267
Isongnatcheak, Nunamiut name for
long-tailed jaeger, 79
Isongngakluk, Nunamiut name
Pomerine jaeger, 77
Ixoreus naevius, 307
naevius meruloides, 217 (table), 247,
266, 269

for

J

Jaeger, 77, 140, 200
long-tailed, 1 (fig.), 23, 69, 78, 200
parasitic, 77, 200
pomerine, 77, 140
Jay, 79, 91, 94, 184, 214, 335
Alaska, 91, 112, 249, 342
Johnson, Ernie, Eskimo man, 26, 52,
129, 327
Johnson, Richard F., 20, 336
Joseph, Paul, 187
Junco hyemalis hyemalis, 118, 231
(table), 234 (table), 248, 253,
269, 303, 345
Junco northern slate-colored, 118

K

Kadgagiak, Nunamiut
Sabine’s gull, 81

Kadgivik, Nunamiut name for willow
ptarmigan, 54

Kaigelak, Clay, 22, 339

Kaiyiorgon, Nunamiut name for north-
ern phalarope, 77

Kakena, Elijah, Eskimo man, 22, 25, 29,
33, 39, 65, 77

Kakena, May, 48

Kaklodook, Kobukmiut name for lesser
scaup, 135

Kaklutuk, Nunamiut name for scaup,
40, 41

Kakmakloak, Nunamiut name
Ross’s gull, 81

Kaksrauk, Nunamiut name for red-
throated loon, 32

Kalloksioyuk, Nunamiut name for the
osprey, 48

Kallorgosiksook, Nunamiut name for
Smith’s longspur, 126

Kaloree, Indian name for widgeon, 172

Kangok, Nunamiut name for snow
goose, 36

name for

for

397

Karlstrom, Thor, N. V., 280

Karplus, Martin, 100, 310

Katu, Indian name for horned lark, 210

Kaviksakruak, Nunamiut name for
gray-crowned rosy finch, 112

Kay, Joe, 162, 172, 174, 175, 176, 180,
182, 185, 193, 204, 206, 216, 220,
221, 223, 242, 243

Kayatavaurak, Nunamiut name for
junco, 118

Kayatavok, Nunamiut name for pine
grosbeak, 111, 118

Kellogg, Remington, v

Kendeigh, Samuel Charles, and
Baldwin, Samuel Prentiss, 309,
310, 325

Kennicotti, Phylloscopus borealis, 102,
147, 248, 260, 266, 270, 316

Kessel, Brina, 113

Kessel, Brina, and Cade, Tom J., 66,
111, 252

Kessel, Brina; Cade, Tom; and Schaller,

George B., 102, 146, 165, 175

Brina; Murie, Olaus;

Schaller, George B., 182, 184

Khut traluk, Indian name for kinglet,
221

Khut tsa luk, Indian name for Bohemian
waxwing, 222

Kidgaviatchaurak, Nunamiut name for
western pigeon hawk, 53

Kidgavitch kiriat, Nunamiut name for
American peregrine, 51

Kidigavitch kiringet, Nunamiut name
for goshawk, 46

Killdeer, 61, 137, 191

Killik Pass, 27 (map)

Kilyaktalik, Nunamiut name for long-
billed dowitcher, 73
Kimitkoilyak, Nunamiut name

sanderling, 76
Kingfisher, 142
Kinglet, 102, 147, 221, 268
ruby-crowned, 104, 147, 250
Kipilugoksiyook, Nunamiut name for
western solitary sandpiper, 66

Kirik, Nunamiut name for Alaska jay,
91

Kitgaviroak, Nunamiut name for gyr-
falcon, 49

Kkya notetuthga, Indian name for
arctic tern, 204

Kessel, and

for

398

Kobuk, birds of, 133-148, 255-256
comments on, 131-133
status of birds at, 246-248 (table)
Kodrakoruk, Nunamiut name for semi-
palmated plover, 60
Kogruk, Nunamiut name for whistling
swan, 33
Kookeh, Indian name for snow geese,
167
Koruaknak, Nunamiut name for Ameri-
can widgeon, 38
Korualorgosik, Nunamiut name for
_ green-winged teal, 38
Korugak, Nunamiut name for pintail,
37
Koyapigaktoruk, Nunamiut name for
eastern robin, 99
Krog, John, iv, 25, 26, 33, 34, 41, 53,
55, 63, 71, 77, 83, 104, 107, 108,
112, 124, 289, 325, 337
Krog, John, and Monson, M. W., 338,

340, 341

Kukuzu, Indian name for snow bunting,
242

Kwit kkyo zyo, Indian name for wag-
tail, 221

Kwi tsi chi, Indian name for gyrfalcon,
182

Kyekyzez, Indian name for myrtle

warbler, 226
Kyha, Indian name for Canada goose,
165
L

Lack, Davis Lambert, 309, 311, 318
lagophonus, Bubo virginianus, 82, 141,
2065, 247, 251, 253, 264
Lagopus lagopus, 57
lagopus alascensis, 54, 185, 247,
248, 249, 264, 324
lagopus albus, 57, 188
mutus nelsoni, 57, 188, 189 (table),
247, 249, 264
mutus rupestris, 190
rupestris nelsoni, 249
lagopus, Alopex, 338, 339
Buteo, 46, 247, 288, 303, 307
Lagopus, 57
Lampe, Chester, Eskimo man, 129
Lampronetta fischert, 245 (fig.)
Laniidae (shrikes), 108, 223, 248, 259

U. S. NATIONAL MUSEUM BULLETIN 217

Lanius excubitor, 223
excubitor invictus, 108, 248, 269,
288, 303
lapponica, Limosa, 76, 189, 247, 252,
288, 316
lapponicus, Calcarius, 313
Laridae (gulls, terns), 79, 140, 200,
247, 259
Lark, 88, 144, 209, 255
horned, 88, 101, 144, 209, 303,
378 (graph)
Larus argentatus, 141, 349
argentatus smithsonianus, 80, 200,
202, 247, 253
argentatus thayert, 202, 284
canus brachyrhynchus, 80, 202, 247,
267, 288
glaucescens, 348
hyperboreus, 141, 276
hyperboreus barrovianus, 79, 200,
247, 267, 288, 316, 338
philadelphia, 204, 247, 267
lathami, Parus cinctus, 145, 215, 247, 264
Lavoisier, Antoine Laurent, 336
Lawrence, George N., 166
Laying, 305-308, 316-18
Lea, John, vi
Leffingwell, Ernest de K., 30, 39
Letpoa ocellata, 310
Lemming, 339, 340, 341
lepida, Tachycineta thalassina, 89, 210,
247, 269
Lepus americanus, 338, 339
timidus, 338
leucopaeta, Branta, 34, 134, 165
leucophrys, Zonotrichia leucophrys, 122,
307, 328
Leucoptera, 230
Loxia leucoptera, 117, 231 (table),
234, 248, 251, 264
Leucosticte tephrocotis, 148
tephrocotis irvingt, 112, 131 (fig.),
248, 265, 266, 269, 288
tephrocotis littoralis, 113
tephrocotis tephrocotis, 112, 345
Lichen, 21
limnaeus, Seiurus noveboracensis, 223,
224 (table), 228
Limnodromus scolopaceus, 64, 72, 139,
247, 266, 269, 288, 316, 322, 376
(graph)

INDEX

Limosa hemastica, 140
lapponica, 76, 140, 247, 252, 288
lapponica baueri, 266, 270
Lincoln, Frederick Charles, 213
lincolnii, Melospiza lincolnii, 231 (table),
241, 248
Linklater, Irvin, 158, 159, 173
Linsdale, Jean Myron, and Sumner, E.
L., Sr., 325
littoralis, Leucosticte tephrocotis, 113
Liva liva, Nunamiut name for all small
sandpipers, 75
Liva livakpaurak, Nunamiut name for
semipalmated sandpiper, 75
Liva livaurak, Nunamiut name for least
sandpiper, 72, 75
Llano, George A., v
lobatus, Lobipes, 77, 199, 247, 267, 288,
303, 317, 377 (graph), 385 (graph)
Lobban, Mary, 128, 157, 165, 204
Lobipes lobatus, 77, 199, 247, 267, 288,
303, 317, 377 (graph), 385 (graph)
longicauda, Bartramia 64, 138, 247, 253,
269, 317
longicaudus, Stercorarius, 1 (fig.), 78,
200, 247, 267, 288, 316
Longspur, 255
Alaskan, 101, 122, 123, 230, 242,
259, 303, 382 (graph), 387 (graph)
Smith’s, 126, 383 (graph), 387
(graph)
Loon, 29, 157
common, 29, 30, 31, 157, 159
Pacific arctic, 31, 149 (fig.), 158
red-throated, 32, 159
yellow-billed, 30, 158
Lord, Peter, 159, 163, 167, 193, 201, 203
Lozia leucoptera leucoptera, 117, 231
(table), 234, 248, 251, 264

lucius, Esox, 271

ludoviciana, Piranga, 257

Lupine, 21

lurida, Dendroica striata, 223, 227

Luscinia svecica, 146
svecica svecica, 101, 260, 266, 270

Lymantria dispars, 281
M

MacGinitie, George, 4
Mackworth-Praed, Cyril Winthrop, and
Grant, Claud H. B., 274

399

macularia, Actitis, 66, 193 (table), 194,
247, 307, 317
Magpie, 91, 214
Malikak, Nunamiut name for horned
grebe, 33
Malirgik, Nunamiut name for Pacific
arctic loon, 31
Mallard, common, 36, 169, 171, 173, 305
mandtit, Cepphus grylle, 315 (fig.)
Maptigak, Mory, 46
Mareca americana, 38, 171, 246, 288
martla, Aythya, 135, 172, 313
maritimus, Thalarctos, 339
maxililiant, Riparia riparia, 90
Mazama americana, 350
McCabe, Thomas Tonkin, and Miller,
Alden H., 228, 328, 329
McConnell, R. G., 10
McDonald, Neil, 193
McElvey, Webb, collector, 156, 157
McLenegan, S. B., 132, 134, 137, 139,
140, 142, 143, 145
Megaceryle alcyon caurina, 142, 247, 269
Mekeana, Homer, 108
Melanitta deglandi, 43, 135, 176, 247,
267, 288, 303
deglandi deglandi, 287 (fig.), 313
dizont, 177, 178 (table)
perspicillata, 44, 178, 247, 267, 288
melanotos, Erolia, 64, 69, 193 (table), 197,
247, 269, 288, 316, 323, 375
(graph)
melodia, Melospiza, 318
Melospiza lincolnit lincolnii, 231 (table),
241, 248
melodia, 318
Merganser, red-breasted, 45, 179, 305
Mergus serrator serrator, 45, 179, 247,
267, 288, 303
meruloides, Ixoreus naevius, 217 (table),
247, 266, 269
Metabolism, 331, 336, 338, 340, 341
Arctic metabolic economy of warm
blooded animals, 331-351
Mice, 341
Micropalama himantopus, 73, 139, 247,
252, 269, 316
Microtus, sp., 182
Migration, arrival from, 289-302
and nesting, biological aspects of,

315-329
and origins, 263-285, 288 (table)

400

Migration—Continued
climate, and taxonomic differentia-
tion, 276-285, 277 (map), 279
(map)
departure on, 312
distance of, 291
duration of migratory behavior,
302-305
energy resources for, 320-329, 322
(table), 323 (table), 326 (table)
influence of seasonal phenomena on,
294-301
laying and, 305
maturity and, 205
mean date of arrival from, 294
memory, homing, and, 283
molting and, 299
of species, 54 (willow ptarmigan), 86
(downy woodpecker), 91 (jay),
94 (chickadee), 112 (pine gros-
beak), 115 (redpoll), 157-158
(loon), 162-163 (Canada goose),
165-166 (brant), 167 (snow
goose), 168 (duck), 169 (Mal-
lard), 170 (pintail), 175 (old-
squaw), 176 (scoter), 176 (white-
winged scoter), 193 (sandpiper),
194 (whimbrel), 200-203 (gull),
210 (swallow), 220 (wheatear),
222 (pipit), 244 (snow bunting),
276 (eider duck)
regularity of, 294-297, 299, 304
reproductive cycle and, 303 (table)
reproductive rate and, 319-320
routes of, 1538, 266, 271-276, 291
(table)
sex and, 305
synchronism of, 283, 304
temperature and, 294, 300
weight during, 323, 328
migratorius, Turdus migratorius, 97, 216,
247, 269, 303, 307, 313, 323
(table), 378 (graph)
minima, Branta, 134, 165
Hylocichla, 146, 272, 307
Hylocichla minima, 99, 219 (table),
247, 269
minor, Chordetles minor, 257
minutilla, Erolia, 64, 71,193 (table), 198,
247, 288, 303, 304, 322, 323, 376
(graph), 385 (graph) .
Mirgiaksyook, Nunamiut name for para-
sitic jaeger, 78

U. S. NATIONAL MUSEUM BULLETIN 217

Misapsak, Nunamiut name for western
tree sparrow, 119

Misikakak, Nunamiut name for black-
capped chickadee, 94

Mitchell, H. H., and Haines, W. T., 343

Mitkotailyak, Nunamiut name for Arc-
tic tern, 82

Monson, M. A., 162, 325, 327, 337

Moore, J. A., 281

Moore, Terris, President University of
Alaska, 26, 28, 119, 131

Moreau, Reginald Earnest, 183, 309

morinella, Arenaria interpres, 63, 265

Morry, John, 46, 65

Morry, Riley, son of John Morry, 103

Moses, John, 205, 207

Mosquito, 21

Motacilla flava, 147, 313

flava tschuischensis, 105, 248, 260,

266, 270, 303, 316, 323, 379
(graph)

Motacillidae (wagtails, pipits), 105, 147,
221, 248, 259

Mouchet, Father, 173, 175, 192, 194

Mountain, Joseph C., 4

Munro, Jonathan Alexander, 106, 166

Munro, J. A., and Cowan, Ian McTag-
gart, 194, 195, 197, 202, 211, 212,
221, 222

Murdock, John, 4, 316

Murie, Olaus, 5, 165, 166, 180, 215

Murphy, R. C., 284

Murray, Alexander Hunter, 163, 165,
166, 167

Muskrat, 27

Mustela erminea, 340
rizosa, 339, 340

N

naevius, Ixoreus, 307

Nakostikyi, Indian name for pintail, 170

Nakrulik, Nunamiut name for horned
lark, 89, 144

Nanum kanockdoroagna, Kobukmiut
name for upland plover, 66, 138

Napaktom Kadgia, Nunamiut name for
Hudsonian spruce grouse, 53

Nastok, Indian name for great gray owl,
142, 207

Nastotesul, Indian name for boreal owl,
207

INDEX

401

Natakcho, Indian name for common | nivalis, Plectrophenax nivalis, 127, 148,

mallard, 169
Natsik, Indian name for white-crowned
sparrow, 235
Nattak, Nunamiut name for American
great gray owl, 84
Nauyatcheak, Nunamiut name for mew
gulls, 81
Nauyavak, Nunamiut name for white-
plumaged gull, 79
Neakoktoakruk, Nunamiut name for
American hawk-owl, 83
nearctica, Aythya marila, 39, 40 (table),
135, 172, 246, 260, 267, 288, 303,
322, 373 (graph)
nebulosa, Strix, 84
Strix nebulosa, 142, 206, 247, 251,
264
Nelson, Edward William, 141, 172, 174,
202
nelsoni, Dendrocopos pubescens, 85, 148,
247, 249, 250, 264
Lagopus mutus, 57, 188, 189 (table),
247, 249, 264
Lagopus rupestris, 249
Nesting, 302-312
Asiatic birds in Alaska, 260
late, 318
maritime birds, 260
northern limits of, 167, 168, 210,
251
repeated, 318
seasonal conditions and, 289
territory, 50 (gyrfalcon), 90 and 91
(swallows), 95 (dipper), 58 and
59 (rock ptarmigan), 225 (yellow
warbler), 240 (fox sparrow), 236
(white crowned sparrow)
Nestlings, growth of, 310-312
Nice, Margaret Morse, 308
nigra, Oidemia, 135
nigricans, Branta, 35, 134, 165, 246, 267,
288, 316, 338, 343
Niklivik, Nunamiut name for white-
fronted goose, 36
Niksaktongik, Nunamiut name for rock
ptarmigan, 57
Ni kut itsi, Indian name for phoebe, 208
Nipailyutak, Nunamiut name for short-
eared owl, 84

Nityitin, Indian name for lesser scaup,
173

231 (table), 242, 248, 250, 259,
266, 270, 288, 303, 316, 338, 345

Niviokruksioyuk, Nunamiut name for
northern gray-cheeked thrush, 99

Nordenskiéld, Nils Adolf Erik, 335

norwegicus, Rattus, 339, 340

notabilis, Seiurus noveboracensis, 223,
224 (table), 227 (table), 248, 269

Notsik, Indian name for horned grebe,
159

noveboracensis, Seiurus, 307

Nukisirgak, Nunamiut name for north-
western horned owl, 82

Nuklinagak, Nunamiut name for Pacific
black brant, 35

Numenius phaeopus hudsonicus, 64, 66,
193 (table), 194, 247, 267, 288,
316

Nungaktuakruk, Nunamiut name for
Gambel’s white-crowned sparrow,
121

nuttalli, Zonotrichia leucophrys, 238

Nuttallornis borealis, 209, 247, 269

Nuvuksruk, Nunamiut name for Baird’s
sandpiper, 71

Nya, Indian name for white-winged
scoter, 177

Nyctea scandiaca, 83, 205, 247, 249, 250,
253, 264

Nyctidromus sp., 338

O

Oakeson, Barbara, 120, 122, 193, 236,
237, 239, 241, 305, 306, 311, 323,

328, 329

obsoletus, Falco rusticolus, 49, 247, 249,
264

ocellata, Leipoa, 310

ochracea, Spizella, arborea, 118, 231

(table), 235, 248, 269, 288, 303,
345, 381 (graph)
Odgillimakadga, Kobukmiut name for
sharp-tailed grouse, 137
Odgillyim kadgia, Nunamiut name for
timber grouse, 59
oenanthe, Oenanthe oenanthe, 100, 146,
220, 247, 260, 266, 270, 303, 316,
323, 379 (graph)
Oenanthe, oenanthe oenanthe, 100, 146,
220, 247, 260, 266, 270, 303, 316,
323, 379 (graph)

402

Ogiuguk, Nunamiut name for common
mallard, 36
Oidemia nigra, 135
nigra americana, 247, 267
Okiotak, Nunamiut name for gyrfalcon,
49
Oklaktak, Numamiut name for spotted
sandpiper, 66
Okpik, Nunamiut name for snowy owl,
83
Okpisoyuk, Nunamiut name for Savan-
nah sparrow, 117
ols Crow, 161 (fig.)
i, & birds of, 157-244, 256
: g comments on, 149-157
E&? status of birds at, 246-248 (table)
Old Hugo, Eskimo man, 65
Oldsquaw, 41, 175, 176, 256
olivacea, Piranga, 257
Olnianskaya, R. P., and Slonim, A. D.,
338
Olor columbianus, 33, 159, 246, 252, 267,
288, 316
Oporornis tolmiet, 257
Oreamnus americanus, 339
Osgood, Cornelius, 150
osgoodit, Canachites canadensis, 53, 186,
247, 251, 264
Osprey, 48, 136, 182
Ovingoayook, Nunamiut name for lesser
yellowlegs, 69
Ovis dalli, 347
Owl, 82, 141, 205
boreal, 85, 142, 207
great gray, 84, 142, 176, 206
great horned, 82, 141, 206, 251
hawk, 83, 206, 207
short-eared, 84, 109, 205
snowy, 83, 205, 250, 253, 318

P

pacifica, Hrolia alpina, 72, 189, 247, 253,
267, 288, 316
Gavia arctica, 31 (fig.), 158, 246,
267, 288
pacificus, Anthus spinoletta, 221, 222
Perisoreus canadensis, 91, 214, 247,
249, 264, 338
Pakagik, Nunamiut name for white-
winged crossbill, 117
pallidiceps, Bombycilla garrula, 108, 222,
248, 251, 264
pallidior, Plectrophenax nivalis, 244

U. 8. NATIONAL MUSEUM BULLETIN 217

pallidus, Plectrophenax, 128
palumbus, Columba palumbus, 328
Pandion, haliaetus carolinensis, 48, 1386,
182, 247, 253
Pandionidae (ospreys), 48, 136, 182, 247,
259
Paneak, Raymond, 96
Paneak, Robert, 71, 96
Paneak, Roosevelt, son of Simon, 68
Paneak, Simon, 22, 23, 24, 25, 26, 28,
29, 30, 33, 34, 37, 38, 43, 45, 46,
48, 49, 51, 54, 55, 59, 61, 65, 66,
68, 70, 72, 73, 76, 77, 80, 85, 87,
90, 92, 93, 95, 96, 101, 102, 103,
104, 105, 110, 111, 112, 121, 128,
129, 132, 145, 148, 187, 237, 243,
254, 290, 297, 335
Paneak, Susie, wife of Simon, 48, 66, 67,
68, 105, 117, 121
Papiktook, Nunamiut name for Ameri-
can marsh hawk, 48
paradisaea, Sterna, 82, 204, 247, 267,
288, 303, 313, 316
Vidua, 338
parasiticus, Stercorarius, 77, 200, 247,
267, 288, 303, 316
Paridae (titmice, verdins, bushtits), 94,
145, 215, 247, 259
Parkes, Kenneth Carroll, and Amadon,
Dean, 275
Parulidae (wood warblers), 110, 147,
223, 248, 259, 261
Parus atricapillus, 145
atricapillus turnert, 94, 247, 249,
250, 264
cinctus lathami, 145, 215, 247, 264
hudsonicus evura, 95, 215
hudsonicus farleyt, 215
hudsonicus hudsonicus, 96, 145, 216,
247, 251, 264
parvipes, Branta canadensis, 164 (table)
Passer domesticus, 338
Passerculus sandwichensis anthinus, 1 1 a
231 (table), 234, 248, 269, 288,
345
Passerella, 230
tliaca, 307
iliaca unalaschcensis, 257
iliaca zaboria, 122, 231 (table), 238
(table) 248, 269, 288, 345, 371
(graph)
Paviaktook, Nunamiut name for pec-
toral sandpiper, 70

INDEX

Pearson, Oliver P., 321, 327, 335
Pedicularis (lousewort), 21
Pedioecetes phasianellus, 59, 137, 251
pelagicus, Phalacrocorax, 331 (fig.)
Peregrine, American, 51, 182, 183, 213
peregrinus, Falco, 136, 307
Perisoreus canadensis canadensis, 214
canadensis pacificus, 91, 214, 247,
249, 264, 338
perspicillata, Melanitta, 44, 178, 247,
267, 288
petechia, Dendroica, 147, 268, 306
Peterson, Roger Tory, 70
Petrochelidon pyrrhonota, 90, 145, 254,
257
pyrrhonota hypopolia, 212, 247, 269,
307
Péwé, Troy L., 10
Peyton, Leonard, 155, 156, 157, 183,
191, 207, 208, 209, 225, 227
Peyton, Sidney B., 32, 67, 68, 133, 138,
156, 157, 183, 191, 218, 225, 227,
229, 234
Phalacrocoracidae, 259
Phalacrocorax pelagicus, 331 (fig.)
Phalarope, 76, 140, 199
northern, 77, 199, 377 (graph),
385 (graph)
red, 76, 140
Phalaropodidae (phalaropes), 76, 140,
199, 247, 259
Phalaropus fulicarius, 76, 247, 267, 288,
316, 317
phasianellus Pedioecetes, 59, 137, 251
Philacte canagica, 134
philadelphia, Larus, 204, 247, 267
Phoebe, Say’s, 144
Yukon Say’s, 87, 208
Phylloscopus borealis, 147
borealis kennicotti, 102, 147, 248,
260, 266, 270, 316
Picidae (woodpeckers, wrynecks), 85,
142, 207, 247, 259
Picoides arcticus, 86, 143, 247, 251, 264
tridactylus fasciatus, 86, 143, 207,
247, 251, 264
pictus, Calcarius, 125, 248, 269, 288,
303, 313, 322, 345, 383 (graph),
387 (graph)
Pike, North American, 271
pileolata, Wilsonia pusilla, 110, 148,
223, 229, 248, 269, 288
Pinguicola, 21

403

Pinicola, 230
enucleator alascensis, 111, 231
(table), 248, 249, 250, 264, 345
Piorgak, Nunamiut name for Alaska
yellow wagtail, 106
Piorgavik, Nunamiut name for Ameri-
can water pipit, 108
Pipit, 105, 147, 221, 255, 268, 303
American water, 101, 107, 380
(graph)
Pipra sp., 338
Piranga ludoviciana, 257
olivacea, 257
Pitelka, Frank Alois, 72, 73, 306, 319
platyrhynchos, Anas platyrhynchos, 36,
169, 246, 288, 307
Plectrophenax nivalis nivalis, 127, 148,
231 (table) 242, 248, 250, 259,
266, 270, 288, 303, 316, 338, 345
nivalis pallidior, 244
pallidus, 128
Plover, 60, 71, 137, 191, 256, 260, 313,
319
black-bellied, 63
American golden, 61, 98, 192, 273,
274, 305, 373 (graph), 383 (graph)
semipalmated, 60, 191
upland, 65, 138
Pluvialis dominica, 192, 274
dominica dominica, 61, 247, 269,
288, 303, 316, 317, 322, 323, 373
(graph), 383 (graph)
Podiceps auritus cornutus, 33, 133, 169,
246, 253, 267
grisegena holbéllii, 32, 133, 159, 246,
267
Podicipedidae (grebes), 32, 133, 159,
246, 259
Polysticta stelleri, 43
pomarinus, Stercorarius, 77, 247, 253,
267, 288, 316
Porcupine, 341, 343, 344, 348
Porcupine Valley, near Old Crow, 160
(fig.)
Porsild, A. E. and Robert, 4, 166, 170,
172, 178, 194, 201, 209, 211, 229,
234, 260, 278

Potentilla, 21

Potokioluk, Nunamiut name for Alas-
kan longspur, 123

Preble, Edward Alexander, 166
Predators, 50, 58, 98, 183, 184, 213

404

Principalis, Corvus corax, 92, 214, 247,
249, 250, 264
Procellariidae, 259
Ptarmigan, 53, 137, 184, 347
Nelson’s rock, 47, 185, 188, 189
(table), 249, 342
white-tailed, 184, 188
willow, 46, 49, 54, 56 (table), 58,
81, 184, 185, 188, 248, 335
pubescens, Dendrocopos, 143
pugetensis, Zonotrichia leucophrys, 237,
238, 305
pusilla, Wilsonia, 257, 307
Wilsonia pusilla, 223, 224 (table),
228 (table), 248, 269
Ereunetes, 64, 74, 193 (table), 199,
247, 288, 303, 304, 309, 313, 316,
317, 322, 377 (graph)
pyrrhonota, Petrochelidon, 90, 145, 254,

257, 307
R
Races, non-adaptive characters of, 281,
284

isolation and, 282
Rana sylvatica, 281
Rand, Austin Loomer, 29, 66, 106, 166,
167, 172, 174, 175, 178, 180, 182,
191, 192, 193, 197, 199, 202, 220,
221, 222, 229, 233, 242, 256, 260
Ranunculus (anemone), 21
Rat, white, 340, 341
wild brown, 340, 341
Rattus norwegicus, 339, 340
Rausch, Robert, v, 24, 25, 47, 80, 89,
95, 96, 101, 108, 112, 129
Raven, 79, 128, 335
northern, 92, 250
Ray, Patrick Henry, 4
Redpoll, 107, 109, 112, 113, 119, 148,
232, 270, 312, 320, 325
common, 115, 232, 2335, 280, 291,
381 (graph)
hoary, 113, 230, 232, 248, 280,
291, 293, 335, 380 (graph), 386
(graph)
Regulus calendula calendula, 104, 147,
221, 248, 250, 253, 269
Rhododendron, 21
Rhodostethia rosea, 81

Richardson, Sir John 91,
166, 167, 254, 271

162, 163,

U. S. NATIONAL MUSEUM BULLETIN 217

richardsoni, Aegolius funereus, 85, 247,
253, 264
Ridgway, Robert, 127
Riparia riparia, 145, 257
riparia maximiliant, 90
riparia riparia, 89, 144, 211, 247,

253, 269
riparia, Riparia, 145, 257
Riparia riparia, 89, 144, 211,

247, 253, 269

Riseitivay, Indian name for snowy owl,
205

Rissa tridactyla, 276

rizosa, Mustela, 339, 340

Robin, Constable P. A., 155, 187

Robin, eastern, 97 (table), 99, 101, 216
(table), 230, 319, 378 (graph)

rosea, Rhodestethia, 81

Ross, James, 293

Ross, Sir John 127, 184, 185, 229

rubescens, Anthus sptinoletta, 107, 221,
248, 269, 303, 322, 380 (graph)

Rulland, Frank, 22, 25, 33, 65

rupestris, Lagopus mutus, 190

rustica, Hirundo, 144, 145

rusticola, Scolopax rusticola, 328

rusticolus, Falco, 182
S

sabini, Xema, 81, 141, 247, 253, 267, 276
Salomonsen, Finn, 128, 148, 244, 293
s.johannis, Buteo lagopus, 46
Sanderling, 64, 76, 140, 252
Sandpiper, 65, 66, 79, 137, 138, 148,
192, 255, 256, 260, 305, 309,
313, 315, 317, 319
Baird’s, 71, 75, 77, 139, 198, 375
(graph), 384 (graph)
buff-breasted, 75, 139
least, 71, 198, 376 (graph), 385
(graph)
pectoral, 70, 197, 375 (graph)
semipalmated, 71, 72, 74,
199, 377 (graph)
solitary, 138, 192, 195, 260
spotted, 66, 194
stilt, 73, 139, 252, 253
western solitary, 66, 281
white rumped, 70

Td;

saturatus, Bubo, 205
saya, Sayornis, 144

INDEX

Sayornis saya, 144
saya yukonensis, 86 (table), 208,
247, 266, 269, 288, 316

scandiaca, Nyctea, 83, 205, 247, 249, 250,
253, 264

Scholander, Per F., 3, 20, 22, 129, 280,
325, 326, 336, 348

Scholander, P. F., Hock, R., Walters,
V., Johnson, F., and Irving, L.,
342, 344, 347

Scholander, Susan Irving, 320

scolopaceus, Limnodromus, 64, 72, 139,
247, 266, 269, 288, 316, 322, 376
(graph)

Scolopacidae (woodcock, snipe, sander-
lings), 64 (table), 138, 192, 193
(table), 247, 259, 317, 318

Scolopax rusticola rusticola, 328

Scoter, 256

common, 135
surf, 44, 173, 178, 251, 252
white-winged, 43, 46, 135, 168, 172,
176, 287 (fig.)
Sea-level isotherms, 10 (map), 11 (map)
Seiurus noveboracensis, 307
noveboracensis limnaeus, 223, 224
(table), 228
noveboracensis notabilis, 223, 224
(table), 227 (table), 248, 269
semipalmatus, Catoptrophorus semipal-
matus, 317
Charadrius, 60, 191, 288, 303, 304,
307

serrator, Mergus serrator, 45, 179, 247,
267, 288, 303

Sha sove, Indian name for tree swallow,
211

Shachtayolik, Eskimo name for fox
sparrow, 122

Shai tso ve, Indian name for bank
swallow, 212

Shattso, Indian name for cliff swallow,
213

Sheffler, W. J., 157

Sheldon, Charles, 32, 34, 43, 48, 67, 86,
90, 98, 109, 128, 132, 133, 134,
135, 136, 138, 139, 140, 141, 142,
143, 144, 145, 146, 147, 148

Shelesnyak, M. C., 3
Shinjee, Indian name for longspur, 242

Shishenetyi, Indian name for semi-
palmated plover, 191

405

Shoolishookruk, Kobukmiut name for
red-necked grebe, 32
Shooloktachailak, Kobukmiut name for
Bohemian waxwing, 108
Shoveler, 38, 135, 172, 252
Shrew, 341
Shrike, 108, 223, 313
Northern, 109
Sialia currucoides, 257
Sigoktovak, Nunamiut name for whim-
brel, 65
Silyirisoktok, Nunamiut name for wan-
dering tattler, 67
Sit tri gichi zzeh, Indian name for
Traill’s flycatcher, 209
Skua, 77, 140, 200
Skutch, Alexander Frank, 311
smithsonianus, Larus argentatus, 80, 200,
202, 247, 253
Snipe, 64, 138, 192, 260
North American, 192
Wilson’s, 64, 69, 193
Snow, and birds, 20
cover for mammals, 335, 338
fall at Anaktuvuk, 19
ptarmigan burrows in, 55, 186
Snyder, Lester Lynne, 166, 201
Solitaires, 97, 146, 216
solitaria, Tringa solitaria, 196, 282, 307
Songakpalutunygik, Nunamiut name
for willow’s warbler, 102, 147
sordidulus, Contopus, 257
Sousik, Big Joe, Eskimo man, 137
Sparrow, 111, 148, 256
eastern white-crowned, 122
fox, 117, 122, 230, 235, 239, 272, 307
Gambel’s white-crowned, 117, 119,
236, 239, 305, 307, 382 (graph),
386 (graph)
golden-crowned, 148, 325
Lincoln’s, 241
Savannah, 105, 117, 234, 268
song, 318
western tree, 118, 235, 239, 307,
312, 314, 381 (graph)
white-crowned, 311, 328, 329
sparverius, Falco, 137, 257
Falco sparverius, 53, 247, 253

Spatula clypeata, 38, 135, 172, 246, 252,
288

Spetzman, Lloyd, v
spinoletia, Anthus, 313

406

Spizella arborea, 307
arborea ochracea, 118, 231 (table),
235, 248, 269, 288, 303, 345, 381
(graph)
passerina boreophila, 257
Sprunt, Alexander, Jr., 320
Squatarola squatarola, 62, 247, 267, 288,
316
squatarola, Squatarola, 62, 247, 267, 288,
316
Squirrel, ground, 334, 338, 339
tree, 339, 341
Stapleton, Robert H., 24
Steen, John, 342
Stejneger, Leonhard H., 220
stellata, Gavia, 32, 159, 246, 267, 288
stelleri, Polysticta, 43
Stercorariidae (jaegers, skuas), 77, 140,
200, 247, 259
Stercorarius longicaudus, 1 (fig.), 78, 200,
247, 267, 288, 316
parasiticus, 77, 200, 247, 267, 303,
316
pomarinus, 77, 247, 253, 267, 288,
316
Sterna paradisaea, 82, 204, 247, 267, 288,
303, 313, 316
Stoney, George M., 21, 132
striata, Dendroica, 223, 224 (table), 226
(table), 248, 268, 269, 272, 307
striatus, Accipiter, 180, 247
Strigidae (typical owls), 82, 141, 205,
247, 259
Striz nebulosa, 84
nebulosa nebulosa, 142, 206, 247,
251, 264
Sturnus vulgaris vulgaris, 328
subruficollis, Tryngites, 64, 75, 139, 247,
269, 288, 316, 317
Suksangik, Nunamiut name for redpoll,
115
Surfbirds, 60, 137, 138, 191
Surnia ulula caparoch, 83, 206, 247, 251,
264
Sutton, George, 82, 86, 95, 205
svecica, Luscinia, 146
Luscinia svecica, 101, 260, 266, 270
Swallow, 89, 128, 144, 210, 255, 261, 268
bank, 89, 90, 144, 145, 211, 253, 254
barn, 90, 144, 145
cliff, 90, 91, 145, 183, 184, 211,
212, 254
rough-winged, 183

U. S. NATIONAL MUSEUM BULLETIN 217

Swallow—Continued
tree, 89, 90, 144, 145, 211
violet-green, 89, 90, 144, 210
Swan, 33, 134, 159, 252, 313
whistling, 33
Swarth, Harry Schelwald, 128, 167, 172,
174, 177, 179, 196, 197, 199,
202, 204, 213
Sya, Indian name for varied thrush, 218
sylvatica, Rana, 281
Sylviidae (Old World warblers, gnat-
catchers, kinglets) 102, 147, 221,
248, 259
Syo, Indian name for robin, 217

T

Tachycineta thalassina, 144, 145, 257
thalassina lepida, 89, 210, 247, 269

Tagatsil, Indian name for least sand-
piper, 198

Taka, Indian name for ptarmigan, 188

Tako, Indian name for rock ptarmigan,
190

Takpilyakruk, Nunamiut name for
boreal owl, 85

Talikvak, Nunamiut name for the kill-
deer, 61

Talivikeak, Nunamiut name for turn-
stone, 63

Taloo, Indian name for hoary redpoll,
233

Tamiasciurus hudsonicus, 339, 341

Tani cho, Indian name for scaup, 172

Tarui, Indian name for swan, 162

Taruikaka, Indian for green-winged
teal, 171

Tasingik, Nunamiut name for black-
billed loon, 29

Tatoo, Indian name for raven, 214

Tattidgak, Nunamiut name for lesser
sandhill crane, 60

Tattler, wandering, 67, 69, 138, 139,
374 (graph), 384 (graph)

taverneri, Branta canadensis, 34, 134,
162, 164 (table), 165, 246, 252,
265, 266, 269, 288, 316

Tazyah, Indian name for Wilson’s
snipe, 193

Tcheekeekak, Indian name for fox
sparrow, 239

Tchichika, Indian name for boreal
chickadee, 215

INDEX

Tchichitoo, Indian name for hawk-
owl, 208
Tchikikeekeejay, Indian name for junco,
235
Tchinkee, Indian name for tree sparrow,
235
Teal, blue-winged, 168
green-winged, 170
Techuh, Indian name for lesser yellow-
legs, 197
Techyo, Indian name for white-fronted
goose, 166
Teevay, Indian name for pine grosbeak,
232
Teggetesel, Indian name for pectoral
sandpiper, 198
Teggetselve, Indian name for semi-
palmated sandpiper, 199
Tekkin, Indian name for red-necked
grebe, 159
Temperature, body, of Arctic animals,
332, 333 (table), 334, 337 (table),
338, 339 (table), 341, 342
tephrocotis, Leucosticte, 148
Leucosticte tephrocotis, 112, 345
Tern, 79, 140, 200
arctic, 82, 204
Terroine, Emile F., and Trautmann,
Simone, 343
Tesitet kyi, Indian name for Barrow’s
goldeneye, 175
Tetnjyo, Indian name for whimbrel, 194
Tetrao urogallus, 57, 324
Tetraonidae (grouse, ptarmigan), 53,
137, 184, 247, 259
Tetre la, Indian name for surf scoter, 179
Tetrik, Indian name for shoveler, 172
Tetyet kkya, Indian name for herring
gull, 201
Thalarctos maritimus, 339
thalassina, Tachycineta, 144, 145, 257
thayeri, Larus argentatus, 202, 284
Thraupidae, 259
Thrush, 97, 146, 216, 256, 261, 268
northern gray-cheeked, 99,
219, 272
olive-backed, 146
Swainson’s, 218, 261
varied, 217
Thuk, Indian name for osprey, 182
Thulvit, Indian name for arctic loon, 158
Tikmiakpaurak, Nunamiut name for
European wheatear, 100

146,

407

Tikmiakpuk, Nunamiut name for
American golden eagle, 47

timidus, Lepus, 338

Tinbergen, Niko, 308, 319

Tissue function adapted to low temper-
ature, 349-350

modification, biological

cance of, 350-351

Titimkotom, Indian name for Canada
jay, 214

Titmouse, 94, 145, 215

Tizinkee, Indian name for crossbill, 234

Tizya, Moses, trapper, 187

Tobuk, David, Eskimo man, 85, 137

Todlik, Nunamiut name for eastern
American golden plover, 61

Todlivak, Nunamiut name for black-
bellied plover, 63

tolmiei, Oporornis, 257

Tolugaknek, Nunamiut name for tree
swallow, 89

Tolungiksyaurak, Nunamiut name for
rusty blackbird, 111

Tongargagruk, Nunamiut name for
white-winged scoter, 44

Tootlik, Nunamiut name for yellow-
billed loon, 30

Torhatoruk, Nunamiut name for godwit,
76

torquatus, Dicrostonyx, 339

Totanus flavipes, 64, 68, 193 (table), 196,
247, 260, 269, 288, 303, 304, 307,
374 (graph)

Tovi, Indian name for goldeneye, 174

Townsend, Charles Haskins, 66, 132,
133, 134, 135, 137, 141, 147

Toyuk, Nunamiut name for downy
woodpecker, 85, 143

Toyukpuk, Nunamiut name for three-
toed woodpecker, 85, 86, 143

traillit, Empidonaz, 208

Empidonaz traillii, 247, 269
Traill’s flycatcher, 208

Traruk, Indian name for spotted sand-
piper, 195

Trevug, Indian name for northern
phalarope, 199

tridactyla, Rissa, 276

Tringa solitaria cinnamomea, 66, 138,
193 (table), 195, 247, 260, 266,
269, 281, 288

solitaria solitaria, 196, 282, 307

signifi-

408

Troglodytes aedon, 309, 338
troglodytes alascensis, 257

Troglodytidae, 259 ;

Tryngites subruficollis, 64, 75, 189, 247,
269, 288, 316, 317

tschutschensis, Motacilla flava, 105, 248,
260, 266, 270, 303, 316, 323, 379
(graph)

Tsetso, Indian name for yellow warbler,
226

Tsetso khekui, Indian name for Wilson’s
warbler, 228

Tsi kwut go katshilyi, Indian name for
shrike, 223

Tsi tut kwilik, Indian name for harle-
quin, 176

Tta shait sove, Indian name for violet-
green swallow, 210

Ttha Tze, Indian name for wheatear,
220

Ttrah, Indian name for merganser, 180

Ttretetere, Indian name for common
loon, 158

Ttsun tratesit, Indian name for the
brant, 166

Ttze kug, Indian name for parasitic
jaeger, 200

Tue, Indian name for solitary sand-
piper, 195

Tuluak, Nunamiut name for northern
raven, 92

Turdidae (thrushes, solitaires, blue-
birds), 97, 146, 216, 247, 259, 261

Turdus migratorius migratorius, 97, 216,
247, 269, 303, 307, 313, 323
(table), 378 (graph)

Turner, Lucien McShan, 86, 105

turnert, Parus atricapillus, 94, 247, 249,
250, 264

Turnstones, 60, 63, 137, 138, 191

Tutchun Tsya, Indian name for three-
toed woodpecker, 208

Tyrannidae (tyrant flycatchers), 86,
144, 208, 247, 259, 261

Tyrannus tyrannus, 257

tyrannus, Tyrannus, 257

Tzecho, Indian name for marsh hawk,
182

Tzi chi tlio, Indian name for Swainson’s
thrush, 219

Tzi choh, Indian name for goshawk, 180
Tzi rzui, Indian name for dipper, 216

U. S. NATIONAL MUSEUM BULLETIN 217

Tzi vit tich kwatle, Indian name for
orange-crowned warbler, 224

Tzintzie, Indian name for gray-cheeked
thrush, 220

Tzivi, Indian name for olive-sided fly-
catcher, 209

Tzi vit sitik kwarzui, Indian name for
blackpoll warbler, 227

U

umbellus, Bonasa, 137, 185, 247

unalaschcensis, Passella iliaca, 257

undulatus, Citellus, 334, 339

unicolor, Cinclus mexicanus, 96, 146, 216,
247, 249, 264

Urner, Charles A., 304

urogallus, Tetrao, 57, 324

Ursus arctos, 347

ustulata, Hylocichla, 146

V

Veesay, Indian name for great horned
owl, 205
velox, Accipiter striatus, 136
Verdins, 94, 145, 215
Vermivora celata celata, 223, 224 (table),
248, 269, 307
Vetch, 21
Vidua paradisaea, 338
villosos, Dendrocopos, 143
virgata, Aphriza, 138, 247, 266
vociferus, Charadrius, 247
Charadrius vociferus, 61, 137, 191,
257
Vole, 341
von Zedlitz, Otto, 56, 324, 328
Voous, Karel H., 320, 328
vulgaris, Sturnus vulgaris, 328
Vulpes vulpes, 339, 341
vulpes, Vulpes, 339, 341
Vyou, Indian name for mew gull, 204

W

Wagtail, 105, 108, 147, 221
Alaska yellow, 99, 100, 105, 108,
147, 260, 314, 315, 379 (graph)
Walters, Vladimir, v, 12, 20, 129, 325,
327, 336
wapacuthu, Bubo, 205

INDEX

Warbler, 255, 256
blackpoll, 227, 272
Kennicott’s willow, 110, 147, 260,
275, 283, 315
myrtle, 110, 147, 226
Old World, 102, 147, 221
orange-crowned, 224
pileolated Wilson’s, 110, 147, 228
willow, 102, 110
wood, 110, 147, 223, 261, 268
yellow, 110, 147, 225
Waterthrush, northern, 227, 229
Watson, C. E. 297
Waxwing, 108, 222
Bohemian, 108, 222
Weasel, 340, 341
least, 340
Weber, Neal, v, 24
Weight, coefficients of variance of, 324
diurnal changes in, 325
dowitcher, 73
laying females 191, 196, 219, 226,
236, 240
local differences in, 56, 57
migration and fat, 322, 328
sex differences in, 64
white-crowned sparrow, 121, 236
Weigold, Hugo, 321, 328
Wetmore, Alexander, v, 55, 89, 284,
311, 320, 332
Whaley, Robert, 157
Wheatear, 146, 220, 260, 265, 270, 274,
314, 379 (graph)
European, 100, 107, 122, 128, 315
Whimbrel, 65, 192, 194
Widgeon, American, 38, 171
Wien, Sig, President of Wien Alaska
Airways, iv, v, 5, 22
Willett, George, 29
Williams, William, 43, 52, 184

409

Williamson, Francis, collector, 156, 157,
180, 181, 191, 195, 205, 208, 209,
210, 217, 218, 219, 224, 226, 227,
241
Willow, 21
Wilsonia pusilla, 257, 307
pusilla pileolata, 110, 148, 223, 224
(table), 229, 248, 269, 288
pusilla pusilla, 223, 228 (table),
248, 269
Wolf, 347
Woodcock, 64, 192
Woodpecker, 85, 142, 145, 207
black-backed three-toed, 143
downy, 143, 208, 250
hairy, 143
ladder-backed three-toed, 86, 143,
207, 251
Nelson’s downy, 85, 94, 112
Wren, house, 309
Wrynecks, 85, 142, 207
Wynne-Edwards, Vero, Copner, 220

DOs Gay
Xema, sabini, 81, 141, 247, 253, 267, 276

Yellowlegs, lesser, 67, 68, 69, 196, 260,
374 (graph)
yukonensis, Bonasa umbellus, 264
Sayornis saya, 86 (table), 208, 247,
266, 269, 288

zaboria, Passerella iliaca, 122, 231
(table), 238 (table), 248, 269,
288, 345, 371 (graph)
Zonotrichia atricapilla, 148, 248, 269
leucophrys gambelit, 119, 231 (table),
236 (table), 238 (table), 248, 269,
288, 303, 322, 323, 345, 370
(graph), 382 (graph), 386 (graph)
leucophrys leucophrys, 122, 307, 328
leucophrys nutialli, 238
leucophrys pugetensis, 237, 238, 305

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