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Full text of "North American fauna"

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ABSTRACT 



INFORMATION is compiled for use in conserving and man- 
aging the band-tailed pigeon, largest native member of the 
family Columbidae in the United States. The report deals 
with the discovery of the bird and its appearance, distribution, 
migration, life history, nesting, habits, and game status, and 
contains a discussion of factors governing its abundance. 

Original information is presented on food habits, based on 
the laboratory analyses of 691 stomachs and/or crops of these 
birds. 

Mast (acorns and pine nuts) vsras found in 268 of the 
specimens and constituted 50.1 percent of the annual food. 
The availability of mast for food largely determines the migra- 
tion routes and wintering habitat, and indirectly determines 
the nature and extent of damage inflicted by these birds upon 
crops. 

The fondness of the bandtail for cultivated cherries and 
small green prunes forms the basis for many agricultural com- 
plaints. These ftuits were found in 98 birds and made up 11 
percent of the diet. Other fruits tff the rose family taken by 
the birds included those of blackberry, salmonberry, service- 
berry, toyon, and hawthorn, together making up 13.6 percent 
of the annual food. 

Cultivated grains formed 12.8 percent of the diet, showing 
the adaptability of the bird to modern conditions. Cultivated 
peas made up nearly 5 percent of the food. Although both grain 
and peas are frequently waste gleaned from harvested fields, 
many complaints of depredations arise. 

Other vegetable items found very acceptable by the bandtail 
include fruits of huckleberry, salal, elderberry, dogwood, cas- 
cara, and hackberry. The foods identified included 76 in- 
dividual items, representing 26 plant families. 

Animal food of the adult bandtail comprises less than one- 
fourth of one percent and appears to be taken only accidentally. 

Th'e bird is a voracious feeder and when large flocks attack 
a farm crop, severe damage may result. A summary of the 
bandtail's economic status is compiled from records of the 
Fish and Wildlife Service, and is followed by suggestions for 
crop protection. 

In winter, California harbors most of the West Coast popu- 
lation of bandtails which constitutes the greater part of the 
total, as birds from the Northwest commonly concentrate there. 
As a species, the band-tailed pigeon is little more than holding 
its own, with local increases in some States, local decreases in 
others. As it lays only one egg, its reproductive potential is low, 
and careful attention must be directed toward every practice 
that might cause further decreases. Shooting seasons and bag 
limits should not be liberalized. 

As to management, breeding stock must be carefully con- 
served over the entire range; the accepted principles and 
practices of forestry that will preserve or even increase the 
supply of oaks and pines, and the wild fTuits and berries that 
supplement mast as food, must be kept in operation; and suit- 
able nesting habitat must be maintained. 



UNITED STATES DEPARTMENT OF THE INTERIOR 

J. A. Kiiifc. Secretary 

FISH AND WILDLIFE SERVICE 
Albert M. Day, Director 



North American Fauna 58 



HABITS, FOOD, AND ECONOMIC STATUS OF 
THE BAND-TAILED PIGEON 



BY 



JOHNSON A. NEFF 




UNITED STATES 

GOVERNMENT PRINTING OFFICE 

WASHINGTON : 1947 



For sale by the Superintendent of Documents, U. S. Government Printing OfBce 
Washington 25, D. C. — Price 20 cents 



it. S. SUPERINTENDENT OF DOCUMENTS 

JUN 19 1947 



CONTENTS 



Page 

Introduction 1 

History of knowledge of ttie species.... 2 

Natural history 4 

Description 4 

Size 4 

Plumages 5 

Courtship 5 

Calls 5 

Behavior 6 

Nesting 6 

Nesting season 6 

Nest location and structure .... 9 

Communal nature in nesting 11 

Eggs 12 

Incubation 12 

Growth of the young 13 

Natural enemies and disease 17 

Distribution 17 

General range 17 

Summer range 18 

Winter range 24 

Migration 2fi 

Spring 2fi 

Autumn 27 

Routes of migration 29 



Page 

Value as a game bird 29 

Present status 33 

Management possibilities 36 

Agricultural relationships 37 

Depredations 37 

Cherries, grapes, and prunes.... 39 

Grain crops 43 

Walnuts 45 

Methods of crop protection 4.5 

Food habits 51 

General 51 

Use of mineral salts 54 

Results of laboratory examinations.. 54 

Seasonal food preferences 63 

Winter 63 

Spring 64 

Summer 6 1 

Autumn 6.') 

Discussion 6.t 

Use of gravel 68 

Influence of age or sex 68 

Feeding mannerisms 68 

Quantity of food taken 68 

Summary 69 

Bibliography 71 



HABITS, FOOD, AND ECONOMIC STATUS OF THE 
BAND-TAILED PIGEON 

By Johnson A. Xefj-, BiologLst. Economic liivcstiffations. Division of Wildlife 
Research, Fish and Wildlife Service 



INTRODUCTION 

For many years the band-tailed pigeon {Columha fasciatci 
fasciata) has pleasurably interested sportsmen and ornithologists 
of the western United States, but at times it has aroused the ire 
of agriculturalists. Pursued as a game bird in some parts of its 
range for more than 70 years (except for a period of complete 
protection from 1913 to 1932) , the band-tailed pigeon has, through 
its wariness and powers of flight, survived in sufficient numbers to 
be treated as a game species in a few far-scattered localities. 

Bandtails, blue pigeons, blue rocks, or wild pigeons, as they are 
called in different parts of their range, are discussed in many pages 
of the ornithological literature of the past 75 years, yet informa- 
tion is still inadequate for determination of the exact status of the 
species and for definition of practical methods for its management. 

Food and cover are the primary essentials for the welfare of any 
species. In few instances, however, does the availability of an ample 
food supply influence the habits, range, and distribution of a species 
as extensively as it does those of the band-tailed pigeon. This 
report summarizes, for practical use, the available information on 
the range, life history, habits, food, and game status of this bird, 
and records the findings from the examination of 691 stomachs 
and crops. 

Prior to 1937 the Food Habits Laboratory, of the Bureau of Bio- 
logical Survey (now part of the Fish and Wildlife Service) , had ac- 
cumulated 142 stomachs and gullets of the band-tailed pigeon, and 
^0 of these had been examined by Sterling Bunnell, E. R. Kalmbach, 
and Alexander Wetmore. Early in 1937 continuation of this study 
was assigned to the writer. The remaining 62 stomachs and crops 
were examined, 549 additional crops and /or stomachs were ac- 
quired and examined, and field observations were carried on in 
5 States. 



studv of th^^n, f- tni.t^^i '^^'^?'"^"*, °i services rendered and of interest shown in this 
fnflhifhfnUv T ",;; f '^^*'°" ',* '"■'-'''' .*° ^ ^'"°"P o*^ cooperators too numerous to ,nention 
assisteH bvVniW^ ''"^^ present employees of the Fish and Wildlife Se.-vice have 

ftPn, ,.^L ^""^'St'ns: specimens, analyzing crop and stomach contents, identifying food 
ilvi-^n^ Tnd'"fdiHn°'' of Vr '" official reports or by letter, and assisting in thl writing 
re\ising, and editing of this report. Valued ass stance has been given also bv virinns 
employees of the Forest Service and the Soil Conservation SeVvce^of the Un ted Sates 

s!rvtTTti,fv^ffZ%'Z^' n"'' °/ "'? °.^.l" ?^ ^"d'^" Aff'-^'^-s '-^"d the National Park 
service ot tlie United States Department of the Interior. Most of the western State Game 

Sens and" in ?n,^ni"]fin^ of their wardens individually, have cooperated In collecting ?peci 
r^dn \inl,^ ^ V ", u?'7"''*'°'?- Certain scientific institutions, among them the Colo- 
Northern ri^onn'nV/n^S^n^^^^^^^ *^' California Academy of Sciences, and the Museum of 
h=,\^-f !^ Arizona, and biologists of many of tJie universities within the ran-e of the 
banatail, as well as many bird students, have also contributed freely. " 



2 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

HISTORY OF KNOWLEDGE OF THE SPECIES 

The type specimen of the band-tailed pigeon was obtained on 
Plum Creek, near Castle Rock, Douglas County, Colo., by members 
of Long's expedition to the PfOcky Mountains, and described for 
the first time by Say in 1823. Eggs were first collected by Lieuten- 
ant Benson (Bendire 1892)1 near Fort Huachuca, Ariz., on 
September 25, 1885. 

It is impracticable, if not impossible, to compare the present 
numbers of the pigeons with those of any year in the early history 
of the West. The variation in degree of concentration in response 
to changing food supplies and the widespread wilderness range of 
the species make it difficult now to draw satisfactory conclusions 
from the scattered reports on abundance. 

Prior to 1913 small notice had been taken of the species as game 
in most States and little protection was afforded it. In the North- 
west most of the shooting was done during the spring months 
when the pigeons were concentrated in grainfields. In California, 
however, pigeon hunting, including market shooting, was de- 
veloped to a high degree. During the period before the automo- 
bile became a great factor in hunter transportation, gunning had 
occurred locally wherever pigeons concentrated. Although the 
number of hunters increased annually as the West became more 
densely settled, lack of transportation limited the hunters to rela- 
tively small areas. Ornithologists were few in number and widely 
scattered, hence there is no estimate of early-day pigeon abun- 
dance that would make possible comparison with present numbers. 
Early writers use such terms as "quite numerous," "common but 
never abundant," "very abundant," "common," "occasionally quite 
plentiful," and "fairly abundant." 

Only one instance has been noted that specified truly great num- 
bers. Bendire (1892) quotes Carpenter as follows : ". . . this species 
is most numerous near the mouth of the Columbia River, where 
immense flocks were to be seen from May to October 1865, which 
fairly rivaled those of the Passenger Pigeon." 

During the winter of 1911-12 there was an enormous ffight of 
band-tailed pigeons along the California coast from Paso Robles to 
Nordhoff", and sport and market hunting flourished. Chambers 
(1912) described conditions in that area, stating that one market 
hunter shipped some 2,000 pigeons to city hotels, that the morning 
train from San Luis Obispo to Los Olivos carried about 100 hunt- 
ers each Sunday morning, and that frequently the hunters took an 
average of 30 birds each. The number of pigeons killed in the 
large area covered by that winter concentration must have been 
enormous, for hunters came from long distances. Apparently the 
birds remained in the area until shot out. This unusual congrega- 
tion of birds and of hunters brought the dangers of the bandtail's 
situation to public notice, and in 1913 Grinnell in furthering the 
drive for protection published an excellent summary of the status 
of the species. 

As practically the entire band-tailed pigeon population of the 

1 Publications referred to parentlietically by date are listed in the Bibliography, p. 71. 



THE BAND-TAILED PIGEON 3 

West Coast wintered in a relatively small area in central and 
southern California, it was possible for concentrated shooting to 
decimate the numbers of the species. The instance described by 
Chambers aroused ornithologists to the great need of the birds for 
protection. 

The first action came in 1913 when in the appropriation act for 
the United States Department of Agriculture there was embodied 
what was commonly known as the Migratory Bird Law of 1913. 
This act asserted the authority of the United States Government 
over those birds, both game and nongame, that in the course of 
their northern and southern migrations passed through any of the 
States or that did not remain permanently within the borders of 
any State or Territory. Under this legislation the Department of 
Agriculture was directed to adopt suitable regulations to give ef- 
fect to the act, and when the regulations were approved and pro- 
claimed by the President on October 1, 1913, they embodied a 
closed season on certain species, including the band-tailed pigeon, 
until September 1, 1918. 

This act served its purpose though there was doubt that it could 
stand the test of constitutionality. Therefore the treaty with Great 
Britain for the protection of birds migrating between the United 
States and Canada was negotiated and was made effective by the 
adoption of the Migratory Bird Treaty Act, of July 3, 1918. Un- 
der this unquestionably constitutional act the closed seasons on 
certain species prescribed in 1913 were continued, and the band- 
tailed pigeon was afforded complete protection until 1932, during 
which period the birds gained slowly but consistently in numbers. 
Owing to the growing numbers of complaints by farmers in Cal- 
ifornia that pigeons were seriously damaging their cherries, the 
Secretary of Agriculture, pursuant to authority granted in the Mi- 
gratory Bird Treaty Act, issued an order on April 30, 1924, allow- 
ing the granting of permits for the destruction of band-tailed pi- 
geons that were attacking cherry crops in that State. In June 
1930 an amended order extended the same privilege to Arizona 
and Washington. Permits were issuable covering the period from 
May 15 to July 31, and the grounds for giving them included dam- 
age to cherries and other small fruits. It was provided also that 
the birds killed in protecting crops could be used as food by the 
landowner or lessee. Complaints of alleged depredations on agri- 
cultural products continued to increase, and the demand for per- 
mits grew until it was apparent that soon in some sections they 
would in effect create almost an open season and that during the 
breeding season of the birds. 

The pigeons continued to increase slowly but steadily in num- 
bers, and in 1932 an open shooting season was permitted in Ari- 
zona, California, New Mexico, Oregon, and Washington. In Colo- 
rado no open season was permitted until 1944, and that only in 12 
southwesterly counties. In Texas and Utah the birds are so sparse- 
ly distributed that no open shooting has been permitted. The open 
season in each State in which band-tailed pigeons are present in 
shootable numbers was limited to 15 days annually until 1942, 



4 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

when the period was extended to 30 days ; the possession bag limit 
in all localities has remained at 10 birds. 

NATURAL HISTOR\ 

DESCRIPTION 

The band-tailed pigeon is described by Grinnell, Bryant, and 
Storer (1918) as follows: 

Adults, both sexes. — Head pinkish brown or vinaceoiis (exact tint varying 
greatly among different individuals), darlcest and more purplisii on top 
and back of liead, more asliy on chiin and cheeics; Ijase of bill straw yellow, 
and black; naked eyelids, coral red; narrow collar around hind neck, white, 
averaging more conspicuous in males; broad area on sides and back of 
neck (below white collar), iridescent bronzy green; back, dark olive 
brown; rump and bases of tail feathers, dark bluish gray; ill-defmed band 
across middle of tail, dull black; terminal portions of tail feathers, drab, 
lightest on outer ones; outer surface of closed wing, chiefly light gray, the 
coverts narrowly margined with white; ilight feathers, brownish black; 
lining of wing and axillars, gray; under surface of flight feathers, dull 
brown; under surface of body, pinkish brown or vinaccous, deepest on 
breast and sides, paling to almost white on belly; under tail coverts, white; 
under surface of terminal portion of tail, whitish, distinctly lighter than 
upper surface of same; feet straw yellow. In some females the tone of 
coloration verges towards grayish rather than pinkish brown. Juvenile 
plumage. — Similar to that of adult, but vinaceous tinge wholly lacking; 
neck without white collar or iridescent bronzing; under surface dark 
brownish, with feather tippings of lighter color, giving a faintly scaled 
effect. 

Marks for field identification. — Largest of our wild pigeons (about the 
bulk of a domestic pigeon); general bluish coloration; distinct dark band 
across middle of square-ended tail; wings without white patches. 

SIZE 

Of 320 band-tailed pigeons examined by John C. Knox near 
Mountain Park, N. Mex., in June 1941, the heaviest adult weighed 
151/2 ounces, the smallest 8% ounces. D. M. Gorsuch, of the Unit- 
ed States Forest Service, weighed 6 birds collected at Williams, 
Ariz., between September 26 and November 8, 1941. Two adults 
weighed, respectively, 345 and 359 grams (or 12.16 and 12.31 ounc- 
es), and four young, respectively, 270, 270, 290, and 290 grams 
(or 9.52 and io.23 ounces). United States Game Management 
Agent Frank F. Foley and the writer examined 22 bandtails shot 
in Colorado between September 17 and 19, 1945; the average 
weight of 17 adults was 338 grams (or 11.2 ounces), the largest 
bird weighing 372 grams (or 13.1 ounces) , the smallest 292 grams 
(or 10.3 ounces), and the average weight of 5 young was 270 
grams (or 9.52 ounces) . A 17-day-old nestling in Colorado weighed 
140 grams (or 4.9 ounces). 

Ridgway (1916) gives the following measurements for the 
band-tailed pigeon as taken from skins, the figures representing 
in millimeters respectively the smallest and the largest measure- 
ments of the birds he examined, and, in parentheses, the average 
(25.4 mm.=l inch) : 

Adult males, 38 specimens.— Length, 342-400 (363) ; wing, 195-221 (208.6) ; 
tail, 122-151 (139.4); culmen, 16-20 (18); tarsus, 24.5-29.5 (27); middle toe. 
27.5-34 (30.4). 



THE BAND-TAILED PIGEON 5 

Adult foiinles, 28 specimens.— hengtlh ;:!34-3!t5 (338) ; wing, 193-218 (204) ; 
tail, 121-149 (134.1); culmen, 15.5-19.5 (17.G) ; tarsus, 24-28.5 (2(1); middle 
toe, 26.5-33 (29.6). 

PLUMAGES 

In addition to features mentioned in the technical description 
quoted, it is stated in the literature that the nestling has a yellow 
skin covered with sparse, cottony, white down. The nestling stud- 
ied in Colorado in 1945, however, was covered with cottony down 
of a peculiar shade of yellow and the skin was not noticeably yel- 
low. 

According to Bendire, juvenile birds apparently molt during 
their first fall. The molt of the adults is not well known. Among 
specimens collected for the present study some birds were identi- 
fied by the field collectors as juvenile, but on laboratory examina- 
tion their crops were found to contain active milk glands. Study 
of September-collected birds in Colorado in 1945 showed consid- 
erable molt under way. Several adult birds had lost most of their 
crown and face feathers and presented a quilled hedgehog appear- 
ance. It was also apparent that soon after the close of nesting 
activity there is a rather speedy reduction in the prominence of 
the white neck-ring, especially in the male birds. 

COURTSHIP 

Apparently little is recorded concerning the courtship of the 
bandtail. Bent (1932), quoting Swarth, indicates that cooing and 
other calls and spectacular circling glides by the male from a perch 
are among the details of courtship. Pearse (1935) describes an- 
other activity, apparently part of the courting behavior. Short 
flights in a hesitant, quivering manner, during which the bird 
seems almost to float, form the basis of this display. The tail is 
fully spread, and Pearse says that the tips of the wings appear to 
be held down. The display is accompanied by two separate very 
low calls, one of which he describes as being very like the modified 
chirping of a cricket. 

CALLS 

The deep, cooing notes of the band-tailed pigeon are distinctive. 
Once heard as they ring through the depths of some mountain 
canyon they will never be forgotten. Mrs. Florence Merriam Bail- 
ey (1902) terms the calls "owl-like." They are most often heard 
during the courting period, and Wales (1926) describes the tilt- 
ing of the head downward to horizontal, the inflation of the neck, 
and other actions incident to the display of that particular period. 
After nesting has begun, the cooing is more subdued and less fre- 
quent. 

A variety of low guttural call notes may be heard when the par- 
ents are together at or near the nest ; these are similar to the notes 
of the domestic pigeon at such times. The squab, when hungry, 
utters a thin, piping note. Neither of these latter calls can be 
heard for more than a few yards. 



6 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

BEHAVIOR 

Band-tailed pigeons are very gregarious, consorting in flocks at 
all seasons except during nesting ; even then communal tendencies 
are exhibited. They are fond of perching for long periods in the 
tops of tall dead or partially dead trees, but when approached drop 
quickly away to better concealment. The flight is strong and swift, 
and in descending mountain sides the birds may dive with wings 
partially or completely shut. During such swoops the noise of the 
wind through the rigidly held wing feathers is often audible for 
a long distance. When the bird takes flight, the wings may be 
clapped together over the back producing a sound that may be 
heard for some distance. This seems to be a form of signal, and 
not the result merely of intense exertion. 

NESTING 

NESTING SEASON 

The nesting period of the band-tailed pigeon is lengthy. During 
practically all the months pigeons may be noted in flocks. Even 
in the nesting season bands of 5 to 50 birds have been observed. 
These flocks were once considered to be nonbreeders, but it is now 
thought that they may be nesting birds. This habit, combined 
with the usual remote nesting habitat, makes it difficult to obtain 
accurate information on the number of broods a season and other 
nesting phenomena. Observers have recorded successive broods 
from the same nest, without proof, however, that the same adults 
were concerned. The long nesting period alone has led some observ- 
ers to believe that at least in the South some bandtails may rear 
two or possibly three broods. In the northern part of the bird's 
range it seems that normally only one brood is produced. 

In California, Grinnell, Bryant, and Storer (1918) list nests con- 
taining eggs or young from March 5 to August 23, and A. C. Ober- 
le (letter of April 15, 1935) says that nesting has been observed 
on the Laurette Ranch north of San Dimas, Calif., and in Brown's 
Flat, the first nesting starting about April 1. Eggs were observed 
into May. Apparently each nest contained but one egg at a time. 
Abbott (1927) quoted Bushnell's observation of a nest that con- 
tained one egg on March 8, 1925, and later a second egg that 
hatched in mid-May. Moran (1919) relates finding a nest with 
one well incubated egg on March 30, 1895, at the head of Lopez 
Canyon, San Luis Obispo County. Grinnell (1928) watched a 
squab being fed in a nest on September 29, 1927. Stillman (1928) 
describes a young bird that left its nest in San Diego County on 
October 2. Kloppenburg (1922) noted a nest in the Plumas Na- 
tional Forest that contained one egg in mid-September. Derby 
(1920) recorded one containing a naked squab on September 1, 
1920, at 6,500 feet elevation on the headwaters of Deer Creek, Se- 
quoia National Forest. Davis (1938) wrote of the nesting season 
near Eureka, and Michael (1928) of that in Yosemite National 
Park. These records seem to bound the outer limits for California. 



THE BAND-TAILED PIGEON 7 

Kitchin (Bent 1932) said that the nesting season in Washington 
lasted from April through June. Einarsen (letter of December 
15, 1939) reported a nest near Triton Cove, Hood's Canal High- 
way, that on August 24, 1939, contained a squab about a week old. 
Hagenstein (193(5) observed a nest containing a squab near Seattle 
on October 17, 1935. 

Gabrielson and Jewett (1940) report that in Oregon egg-laying 
is at its height late in May and in June, and give May 3 and July 
12 as the known extremes of the season. Later, Jewett (1941) re- 
corded a nest with two eggs in Yamhill County on September 30, 
1940. 

In Arizona, according to Bendire's correspondents, nesting oc- 
curred in nearly every month of the year. Poling, writing to him 
from Fort Huachuca of various nests, said, "I have taken young 
two or three months old in February, and since that time young 
and eggs enough to show that they lay and nest from December 
to August." Benson reported nesting of the bandtails in the Hua- 
chuca Mountains from early July to late October; Willard (1913) 
noted an October nest; and Vorhies (1928) one in September. On 
Pinal Mountain south of Globe, Ariz., Carlos Stannard reported 
nests as follows : August 18, 1940, one egg ; August 19, 1940, squab 
ready to fly ; August 23, 1940, a squab about 4 days old which was 
still in the nest on August 30 ; and on August 14, 1941, four nests, 
one with eggs, three with squabs between 1 and 3 w^eeks old. Con- 
siderable field work by the writer, accompanied by State game 
wardens, leads to the conclusion that in southern Arizona nesting 
most commonly begins early in May, though occasionally earlier. 

For New Mexico, Mrs. Florence Merriam Bailey (1928) lists 
a number of interesting records, chief among them being that of 
a nest containing one wellincubated egg on April 23, 1922, 16 miles 
northeast of Santa Fe, at 10,500 feet elevation, well above the snow 
line. There are also records for the Animas Mountains, Guad- 
alupe Mountains, Monument Pass, Black Mountains, and the Pe- 
cos, Sacramento, Sandia, Taos, Sangre de Cristo, Jemez, and Mo- 
gollon Ranges, the dates extending from June 3 to mid-August, 
and the elevations from 5,800 to 8,000 feet. Of 280 adult band- 
tails examined by John C. Knox at Mountain Park between June 
25 and July 10, 1941, 235 were males, 151 of them with testes in- 
dicating active breeding condition, and 45 wxre females, 22 of 
which contained well developed eggs. 

In Utah, Lee Griner records a bandtail nesting in June, and 
Clarence Cottam (1941) saw a juvenile recently out of the nest 
on July 24. In Texas, Ray Williams, State Game Warden, ob- 
served a pair of pigeons building a nest on April 22, 1933, in the 
Davis Mountains, and saw a nest containing an egg in the Chisos 
Mountains in July of that year. 

For Colorado there was until 1945 no definite breeding record 
substantiated by collection of eggs or young or by photographs of 
nests. The presence of the birds over a wide range, however, made 
it certain that they did nest. R. J. Niedrach (Niedrach and Rock- 
well 1929) of the Colorado Museum of Natural History collected 
a bird near Kittredge on June 20, 1928, that contained a fully 



8 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

formed egg, and in Jarre Canyon near Sedalia on September 11, 
1938, he took several birds that contained active milk cells in their 
crops. Several State game wardens and woodsmen relate having 
seen nests in earlier seasons, but none were reported at the 
time and hence are not verified. Warden Hall, of Montrose, re- 
ported that in 1943 on Tabeguache Creek in the Uncompahgre Na- 
tional Forest he saw a number of juvenile birds that were so imma- 
ture they could not have been long out of the nest. 




B-62847 

Figure 1.— Nest and day-old yomig of the band-tailed pigeon in a 
lodgepole pine on the Rampart Range, Pike National Forest, Colo., 
August li2, 11J45, the first recorded and authenticated nest of this 
species for the State. 



THE BAND-TAILED PIGEON 9 

On August 22, 1945, the writer found the first recorded and au- 
thenticated nest for the State. It was in a lodgepole pine on the 
summit of the Rampart Range, Pike National Forest, some 55 
miles south of Denver, and contained a day-old squab (fig. 1). 
On August 31, R. J. Niedrach found a second nest nearby contain- 
ing an older squab. On the basis of information available at the 
end of 1945 it seems that the major nesting season for Colorado 
covers the period July 1 to August 30, with a few pairs of band- 
tails nesting earlier or later. 

NEST LOCATION AND STRUCTURE 

The location of band-tailed pigeon nests varies greatly. 0. B. 
Johnson (1880), Irene G. Wheelock (1904), and Cooper (1880) 
report ground nests, but no other similar nestings have been re- 
corded. Johnson observed in the Willamette Valley, Oreg., nests 
ranging from the top of a stump 8 feet high to the dense branches 
of a fir at about 180 feet. C. E. Bendire (1892) quotes Ankeny as 
describing nesting in the Rogue River Valley, Oreg., on limbs of 
small firs, generally in dense thickets. William Lloyd wrote Ben- 
dire that in Texas and in Mexico nests were largely in oaks, and 
Stephens told him that in Arizona nests were usually in pines at 
heights of 8 to 20 feet from the ground. Benson and Poling report- 
ed nests in Arizona in pines, oaks, and mulberries, at heights of 
15 to 70 feet, and from 5,000 to 8,000 feet elevation in the moun- 
tains. 

E. A. Kitchin, in correspondence with Bent, said that nesting 
sites in Washington were mainly in dark fir trees, in the lower 
branches near the trunk, and averaged about 20 feet from the 
ground. Occasionally nests were found in alder or birch on hill- 
sides. Leo K. Couch informed the writer that around Willapa 
Harbor, Wash., the bandtails nest in Sitka spruce (Piccci sitchcn- 
sis) . Ira N, Gabrielson and Stanley G. Jewett (1940) report that 
in Oregon the nest is usually high in a coniferous tree. 

Grinnell, Bryant, and Storer wrote that most of the California 
nests were in oaks, at heights of 8 to 30 feet. One was reported in 
a Douglas fir and one in a California lilac (Ceanothus thyrsiflor- 
us). 

Fowler (1903) reported live oaks as favored locations for nests 
in the Huachuca Mountains of Arizona, the nests being built on 
low, horizontal limbs from 9 to 12 feet from the ground. In July 
1939 the writer, with George Peterson, Deputy State Game War- 
den, of Nogales, Ariz., visited several of the high oak canyons in 
the Tumacacori and Washington Camp Mountains near Nogales in 
search of bandtail nests. About 10 nests, deserted but of the cur- 
rent season, were inspected . All were in oak trees at heights of 
12 to 25 feet, in dense woods near the canyon floors. David 0. 
Scott, of the United States Forest Service, describes a nest found 
near Williams, Ariz., as being 15 feet up in a yellow pine and very 
loosely constructed of coarse oak sticks. 

The two occupied nests found in 1945 in Colorado, and several 
unused nests found nearby, were all in lodgenole pine forest at ele- 



10 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 



vations of approximately 8,400 feet. One nest was in a fork against 
the trunk, the others one to two feet out on flat Hmbs ; all were 
14 to 16 feet above the ground. All were in trees on rims or slopes, 
with lower vegetation downhill, so that the adults had room for a 
downward dive upon leaving the nest as they picked up momentum. 
The nests (figs. 2 and 3), typical of those of the pigeon family, 




B87I4M 

Figure 2. — Band-tailed pigeon on a typical nest in a pine tree on Maverick 
Mountain, Prescott National Forest, Ariz. (Photographed hy Floyil Schroeder, 
Fnited States Forest Service.) 

are usually frail platforms of dead twigs of oak, pine, or other 
plant stems. The ground nest reported by Johnson (1880) in the 
Willamette Valley was made of leaves and moss. A nest measured 
by Grinnell (Grinnell, Bryant, and Storer 1918) was 8% inches 
by 4 inches, exclusive of several protruding twigs. Frequently 
there are numerous gaps in their structure, the frailty of the nests 
being generally commented on in the literature. Nests observed 
by the w^riter near Nogales, Ariz., in 1939 and 1940 were thin 
platforms of oak twigs, so fragile as to cause one to wonder how 
eggs and young could remain on them. 

Willard (1916) , describing nest building in the Huachuca Moun- 
tains of Arizona, says that it was carried on only in the morning 
hours from sunrise to about 8 a. m., almost entirely by the female, 
and that she proceeded in a most lackadaisical manner. The peri- 
od from the placing of the first stick to the laying of the first egg 
was 6 days. 



THE BAND-TAILED PIGEON 



11 




FiGiTRE 3. — Nest aud young of the band-tailed pigeon in the humid coastal area 
of Tillamook County, Oreg., June 25, 1931. (Photographed by Alex Walker.) 

Bent quotes Kitchin as stating that bandtails are very fond of 
their old nesting sites and that they come back year after year to 
the same limb even though the previous nest has been disturbed. 
Kitchin records second nests built on the same spot where an earli- 
er nest of the current season had been robbed or destroyed, and 
cites one instance of a nest from which he collected the egg being 
followed by a second nest on the same site from which he again 
took the egg ; later in the season he found in the same place a third 
nest that he left intact. 

Noack (1916) describes the breeding of the band-tailed pigeon 
in captivity. The first egg laid was removed from the nest and was 
hatched by a domestic pigeon. Within a few days the bandtail laid 
another egg, which was hatched and the young bird raised. Then 
a third egg was laid and a second squab reared to maturity. 



COMMUNAL NATURE IN NESTING 

Usually the band-tailed pigeon nests in widely separated pairs. 
Fowler (1903), however, describes a community breeding ground 
near Fort Huachuca, Ariz., where some 35 pairs nested in a loose 
rookery ; in the area most thickly populated there was one nest to 
each 3 or 4 acres. Observations by George Peterson, Deputy State 
Game Warden, of Nogales, in 1936 in the Tumacacori Mountain 
range northwest of Nogales, Ariz., were of the same purport. The 
community nesting ground was in dense vegetation along the bot- 



12 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

torn of a steep canyon at an elevation of about 6,000 feet. Inspec- 
tion of recently deserted nests in that canyon and others by Peter- 
son and the writer in 1939 and 1940 showed that nests were irregu- 
larly spaced along the bottom of the canyon, sometimes within a 
few yards of each other, but usually fairly widely separated. 

Surprising examples of community nesting are contained in 
two notes transmitted by H. Garvin Smith, of the United States 
Forest Service, Tucson, Ariz. He says that in 1933, while camping 
in the Magdalena Mountains of New Mexico he found 14 bandtail 
nests in one large Douglas fir near a spring, and that in 1934 in 
the same tree he found 17 nests, one of which contained two eggs, 
the others one each. 

Fred Herman, of Sanborn Park, Montrose County, Colo., relates 
that a former Forest Service employee described a number of 
"colony trees" that he had seen among the thick, bushy conifers 
along the rimrock above Horsefly Creek Canyon north of Sanborn 
Park, and that he had also found the pigeons nesting in blowholes 
and on ledges in the sandstone rimrock along Big Red Creek Can- 
yon nearby. Several other Colorado game wardens have reported 
bandtail activity about similar rimrock areas and they feel sure 
that pigeons nest there. 

EGGS 

The usual, or normal, deposition is one egg to a nest, but in the 
literature are a number of records of two eggs. In a tabulation 
covering 18 nests in California, Grinnell, Bryant, and Storer 
(1918) report one nest containing two eggs and one containing 
two squabs. 0. B. Johnson (1880) observed two nests in the Willa- 
mette Valley, Oreg., that contained two eggs each. Bendire quotes 
Ankeny as saying that two eggs were laid in nests in the Rogue 
River Valley, and Lloyd as describing two-egg sets in Texas and 
in the Sierra Madre of Mexico. Cooper (1880) also said that the 
set was two eggs. On the other hand, Bendire quotes Benson, 
Stephens, and Poling as stating that only one egg to a nest was 
laid in the Huachuca Mountains of Arizona. Jewett (1941) re- 
ports a two-egg nest in Yamhill County, Oreg., on September 30, 
1940, and H. Garvin Smith writes that he has seen several two- 
egg nests in New Mexico. 

Ralph Morrow, Deputy State Game Warden, of Hilltop, Ariz., 
observed "several" nests in the Chiricahua Mountains, in each of 
which w^ere a half-grown squab and one egg. G. W. Peterson, 
Deputy State Game Warden, of Nogales, Ariz., said that during 
June 1936 in canyons of the Tumacacori Range northwest of No- 
gales he inspected 56 bandtail nests, 11 of which contained two 
eggs. 

The eggs of the band-tailed pigeon are elliptical-ovate, somewhat 
pointed, smooth, pure white, and with a slight gloss. A series re- 
ported by Bent averaged 39.7 by 27.9 millimeters in dimensions. 
(25.4 millimeters^l inch.) 

INCUBATION 

According to Bendire (1892), incubation lasts 15 to 18 days; 



THE BAxND-TAlLED PIGEON 13 

even longer periods should not be exceptional in some of the early- 
season nests at high altitudes where the weather is still very cold. 

GROWTH OF THE YOUNG 

Doves and pigeons differ from all other birds in feeding their 
nestlings a substance, usually called "pigeon milk," that is devel- 
oped in the crops of both the male and the female adult. It is 
creamy-colored and somewhat similar in appearance and consist- 
ency to curd. Feeding is by regurgitation, and for the first few 
days after the nestlings are hatched this "milk" is their only food. 

Little is known of the development of this secretion in the band- 
tail. Even in regard to the domestic pigeon the literature lacks in- 
formation on some phases that are of interest ; but since the domes- 
tic pigeon incubates for 18 days and the young remain in the nest 
for at least 30 days, it seems probable that the development of the 
pigeon milk and the duration of the supply may be similar in the 
two species. The most detailed discussion of the phenomenon of 
pigeon milk that the writer has found is one that is contained in 
the literature on the domestic pigeon, and is here briefly summa- 
rized. About the 8th day of incubation the lobes of the pigeon's 
crop begin to increase in size through multiplication of the cells 
of the outer, or proliferating, layer ; by the 12th day the walls of 
the lobes are obviously thickened and enlarged and on the 18th 
day they are at their greatest development. The inner layer of each 
lobe has been pushed farther away from the source of blood in the 
outer layer, and it begins to degenerate ; globules of fat develop in 
the cells, and whitish masses of dead cells begin to peel off. By 
the time the eggs hatch on the 18th day these greasy masses are 
sufficient to furnish the entire food of the squabs during their first 
few days of life. After the first few days the regurgitated ma- 
terial begins to contain some quantity of seeds and other solids 
that have been softened in the crop of the adults. If is said that 
the duration of the milk supply varies considerably in the different 
families of the domestic pigeon, and that in some it has ceased to 
form by the time the young are 7 days old, w^hereas in others ap- 
proximately half of the food is still "milk" when the young have 
reached 10 days of age. 

Bendire (1892) states that both sexes assist in the care of the 
young. "The young grow rapidly and are able to leave the nest 
when about a month old." Mrs. Wheelock (1904) says that the 
squab is "fed on a thin milky fluid, by regurgitation, for 20 days." 

Considerable food must be administered at a feeding, as the 
squabs maintain continued rapid growth with a minimum of pa- 
rental visits. Abbott (1927) published notes from A. E. Stillman, 
of San Diego County, Calif., dated September 17, 1922, as follows : 
"That day the female left the oak tree in the early morning and 
returned at twilight ; after quickly feeding the young she left 
again. Next day she left at daybreak and returned at sundown." 
This squab, according to Abbott, left the nest and tree on October 2. 

Under date of November 9, 1939, Carlos Stannard, of Phoenix, 
Ariz., wrote : 



14 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

We camped near the top of Pinal Peak, about eight miles south of 
Globe, Ariz., at about 7,500 feet. In a small yellow pine just over our tent 
was a band-tailed pigeon's nest with one young. It was about a week old 
on August 13. I watched the nest pretty steadily for a number of days; 
we were in that camp until August 31. No adult stayed on the nest during 
the day after we went there. And after the 16th no adult stayed on the nest 
during the night. The young left the nest on the 25th but did not leave 
the tree; it was still in the tree on the morning of the 31st. During the 
time we were there the young was fed once each day by the female, 
usually shortly after sunrise, and once by the male about 8 or 9 a.m. I 
banded all three. The adults seemed to pay no more attention to us than 
if we were cattle. 

After the discovery of the two nests in Colorado in August 1945, 
the writer made every effort to recheck existing information on 
the growth of the young. As R. J. Niedrach was making a photo- 
graphic record of these first Colorado nests, the adults were not 
disturbed as frequently as would have been desirable to obtain 
full data on growth, but some observations were made on the two 
nests. At one day of age the squab was a tiny helpless creature 
about 2 inches long, unable to hold up its head for more than a 
few seconds. The female parent was on the nest when it was 
found at 2 p. m. on August 22, and she remained on the nest dur- 
ing the daylight hours of August 24. The male was on the nest 
during most of the day of August 23. During the remainder of 
the study the adults followed this routine : The male returned to 
the nest between 8:45 and 9:30 a.m. each day and brooded the 
young bird during the day (fig. 4). The female returned to the 




Figure 4.— Male band-tailed pigeon on nest hovering a ten-day-old scpiab. 
Kanipart Range, Pike Xationul Forest, Colo. August 31, 1945. (Photo 
graphed by R. J. Niedrach.) 



THE BAND-TAILED PIGEON 15 

nest at any time between 3 :45 and 5 :15 p. m. She was not observed 
feeding the youngster, though she undoubtedly must have fed it. 
Watched from davv^n until dark, the tiny squab appeared to sleep 
until after the return of the male. During the first week about 
three feedings, all between noon and 3 p.m., seemed to be the 
schedule. As the youngster's food capacity grew, the number of 
feedings was reduced to two, and the period of regurgitation indi- 
cated that a large quantity of food was taken. Both feedings were 
about midday. 

Contrary to Stannard's observation, the adults continued to 
hover the squab, both day and night, until it was 20 days old. On 
that date they stopped abruptly and did not return to the nest 
during either the day or the night except to feed the squab. Each 
parent came to the nest once daily, between 10 and 11 a. m., fed the 
squab, and departed. Occasionally during the day one of the par- 
ents would visit the vicinity, scan the nest and its surroundings 
carefully, and then depart. 

During the first 10 days the young squab did not grow very 
rapidly. It slept most of the day and increased in size, but the 
feather development seemed to be slow. After about 12 days feather 
development began visibly to progress. At 17 days of age one of the 
squabs was well covered with feathers, the body feathers being 
about 15 mm. out of the sheath and the first primary 30 mm. out 
of the sheath; the tail feathers measured 28 mm. from the tip 
of the tail flesh to the tip of the feathers; the head was heavily 
pinfeathered but had no open feathers, and the sides were quite 
bare. The yellowish brown down adhered to the tips of the 
feathers, giving the squab a peculiar fuzzy appearance. At this 
age the youngster crawled clumsily about over the nest and 
snapped its beak furiously at the intruder. It weighed 140 grams, 
or 4.9 ounces. External examination indicated that the crop was 
well filled with pigeon milk. 

When the squab was 20 days old the tail feathers measured 42 
mm. in length and the first primary was 40 mm. out of the sheath ; 
the pinfeathers on the head were opening but the sides were still 
quite bare. When it was 23 days of age its outward appearance 
had changed little (fig. 5), but when the writer's hand approached, 
the young bird snapped its beak vigorously, struck with bent wing, 
and danced awkwardly about over the nest. When the squab was 
26 days old it weighed 243 grams and its tail measured 75 mm. 
For the first time it spent much time preening, apparently picking 
off the down that still adhered to the tips of the feathers. On that 
day also, one of the squabs began to exercise and spent much of 
the day walking about, for the first time venturing off the nest 
onto the nest branch, waving and flapping its wings, and craning 
and peering about with interest. This was repeated on the 27th 
day, and on the 30th day the bird was gone from the nest, tree, 
and immediate area. The other squab was not observed between 
its 20th and 26th day, but on the latter day it was sitting quietly 
on the nest without apparent interest in exercising. The nest was 
not again visited. 

Nowhere in the literature examined has the writer found any 

695766°— 47— 2 



16 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 




FiGUKE 5. — Young hand-tailed pigeon about 24 days old on nest. Note the 
higli degree of eaniontiage alTorded hy the nest, trunk and branches of 
the tree, and the low-sitting, quiet habits of the young bird. Rampart 
Range, Pike Naticnial Forest, Colo. September 3. 194:1. (Photograplied 
by R. J. Niedrach.) 

information on how long the lobes of the crops of the adult band- 
tails remain enlarged and thickened, with easily distinguishable 
milk cells, after the period of active feeding of 'milk' to the young. 
Knowledge of the length of time that the young bandtail is fed 
'milk' is incomplete, but one squab examined in 1945 appeared 
to have its crop well filled with the soft curdlike 'milk' on the 
17th day. Just how long this material, alone or in combination 
with softened seeds and grains, makes up a part of the food of 
the squab and how long thereafter the thickened, whitish milk-cell 
pads remain easily distinguishable in the lobes of the crop of the 
adult after the 'milk' has ceased to be an item of food for the young 
are not yet known. 

On numerous occasions adult pigeons collected for study or 
shot by hunters on dates well outside the normal nesting season 
have carried weW defined milk pads in their crops ; others have 
contained variable remnants of the thickened crop-lobe walls. In 
some instances observations indicated that the birds still had 
young in nearby nests ; in others that they apparently were mi- 
grants. Among such late-season records of bandtails containing 
well defined milk pads in their crops are those of birds taken in 
Washington, September 22 and 26 and October 2; in Oregon, 
September 8 and October 3 ; in Arizona, September 29 ; in New 
Mexico, September 23 ; and in Colorado, September 15 to 20. In 
southwestern Colorado in 1944 a majority of the birds examined 
by United States Game Management Agent Frank F. Foley from 



THE BAND-TAILED PIGEON 17 

September 16 to 20 contained this evidence of recent feeding of 
young, and in New Mexico as late as September 23, 9 out of 34 
birds examined showed a similar condition. Until the correlation 
between disappearance of the milk-cell pads and the 'weaning' 
of the squab by its parents is definitely established, accurate analy- 
sis of these observations is impossible. 

NATURAL ENEMIES AND DISEASE 

The few instances of molestation of bandtails recorded in the 
available literature are those by Willard (1916), who said that in 
Arizona "the Prairie Falcon and Cooper Hawk take considerable 
toll from the flocks;" by Kitchin (Bent 1932), who remarked that 
in Washington a gray squirrel occasionally took possession of a 
pigeon nest, using it as a foundation and adding to it to suit itself ; 
and S. D. Durrant, whose Hanna, Utah, specimen was struck 
from a flock by a sharp-shinned hawk (Cottam, letter of May 6, 
1940). McLean (1925) reports that a western goshawk pursued 
bandtails in Yosemite National Park. No doubt there is some pre- 
dation upon eggs and squabs, but no data are available. 

One adult pigeon collected near Durango, Colo., in July 1945 was 
found to contain at least 12 flatworms in its abdominal cavity. 

Sick pigeons were reported from Whidby Island, Wash., be- 
tween September 8 and 21, 1939. All were found close about wa- 
terholes. Reports from United States Game Management Agent 
Bach mention six birds found dead by Washington State game 
protectors and three still able to fly weakly, which were shot. Six 
specimens were sent to Dr. Karl F. Meyer, Hooper Foundation, 
University of California, who found no evidence of poisoning or 
of bacterial disease. Dr. Meyer reported hemorrhages in the giz- 
zard and evidence of acute irritation in the intestinal lining, but 
the causes were unknown. 

The band-tailed pigeon seems to be relatively free from natural 
enemies ; the birds feed on the ground with little apparent fear, 
and perch conspicuously in the tops of dead trees. One beneficial 
result of their flocking habit is that individual safety is enhanced 
by the combined watchfulness of all the birds of the flock. Man 
through shooting the birds and clearing and destroying their nest- 
ing range, appears to be the only enemy of importance. 

DISTRIBUTION 

GENERAL RANGE 

As recorded in the American Ornithologists' Union Check-List 
of North American Birds, fourth edition, 1931, the range of Co- 
lumha fasciata fasciata is as follows: 

Breeds in the Transition Zone from southwestern British Columbia, Mon- 
tana, and north-central Colorado south through the southwestern United 
States and Mexico to Guatemala, and east to western Texas. Winters from 
the southwestern United States southward. Accidental in North Dakota. 

Library research and correspondence have been undertaken to 



18 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

obtain a concept of the bird's range at the present time. The re- 
sults are here presented, beginning in the northwest and moving 
southward and eastward. 



SUMMER RANGE 

The summer range of the band-tailed pigeon is shown in figure 6. 




Seal 
200 



Figure 6. — The snmiiier raii.ce of the band-tailed pigeon. (In Canada, north to 
Bella Coola and Graham Island.) In Arizona, Colorado, New Mexico, Texas, 
and Utah, large expanses of range grassland and semi-arid dessert may 
separate the monutain ranges in which pigeons are found. The range as 
shown in tJie map covers the extreme distribution as known from existing 
records. 



THE BAND-TAILED PIGEON 



19 



In British Columbia, Taverner (1934) says that the species is 
found west of the Coast Range, north at least to Bella Coola on 
the mainland and Tow Hill, Graham Island, and Queen Charlotte 
group along the coast. According to Bent (1932), Courtenay and 
Chilliwack are the normal limits of range, hence the southern half 
of Vancouver Island may be considered the area normally in- 
habited by the birds. 

In Washington, as in Oregon, bandtails breed locally in varying 
density west of the Cascades, and probably erratically in the Tran- 
sition and Upper Sonoran Zones of the eastern slopes. The coast- 
al belt and the area adjacent to Puget Sound seems to be the cen- 
ter of their abundance in the State. John Finley, of the Fish and 
Wildlife Service, at Olympia, says that the most easterly record 
he has obtained w^as near Bingen, Klickitat County, on the Colum- 
bia River. 

In Oregon, Gabrielson and Jewett (1940) report that the spe- 
cies is common in the western part of the State, its greatest abun- 
dance being reached on the coast (fig. 7). Breeding in Oregon, 




B87I0IV1 

FiGUKE 7. — Typical band-tailod pisi'Oii habitat in a ■'burn'" in the Ort'j^dii coastal 
mountains, in Tillamook County, where the luxuriant vegetation offers both 
food and nesting cover. (Photographed by A. S. Einarsen, September 10, 
1940. ) 

as elsewhere, is erratic. Pigeons may rarely be seen on the east- 
ern slopes of the Cascades, and the only record for the State at 
any distance east of this range is of a lone, immature bird col- 
lected by Jew^ett in the Steens Mountains in Harney County on 
October 19, 1928. 

In California band-tailed pigeons nest locally and erratically in 
Transition and Upper Sonoran Zone areas west of the Sierra Ne- 
vada divide, chiefly in mountainous areas from Del Norte and Siski- 
you Counties in the north to the Laguna Mountains of San Diego 



20 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

County, Their choice of nesting habitat apparently depends to a 
considerable degree on temperature. In the southern mountains, 
in the Sierras, and in the more northerly interior mountains 
nesting areas are usually at high elevations, but near the coast 
and in the coastal mountains they are scattered widely at much 
lower elevations, including the coastal redwood belt. According 
to Captain W. J. Harp, California Division of Fish and Game, the 
pigeons nest locally in fair numbers near the small ranches of 
Humboldt and Del Norte Counties in northwestern California. 

Bandtails have been listed in so many publications on California 
birds that it is unnecessary to mention them all; Grinnell and 
Wythe (1927) describe the species as a summer resident locally 
in the coastal belt just north of San Francisco Bay; George Wil- 
lett (1933) says that in southern California it breeds in moderate 
numbers in the oaks of the Transition Zone, mostly above 3,000 
feet, south to southern San Diego County; Grinnell and Storer 
(1924) record it as being in Yosemite Valley during the nesting 
season; Derby (1920) describes a nest in the Sequoia National 
Forest; and Kloppenburg (1922) reports one in the Plumas Na- 
tional Forest. These records outline roughly the outer limits of 
the range within which band-tailed pigeons may be observed dur- 
ing the summer where habitat is favorable. 

On October 4, 1941, Luther C. Goldman, of the Fish and Wild- 
life Service, collected a migrating band-tailed pigeon at a point 
7 miles east of Calexico, 3 miles north of the Mexican line. This 
constitutes, so far as the writer knows, the first record of the spe- 
cies from the floor of the Imperial Valley. 

In Nevada, Leo K. Couch reported observing a mature band- 
tail near Success Divide, Duck Creek Range, Nevada National 
Forest, on November 4, 1943. 

In Utah, band-tailed pigeons have been observed at several points 
during the breeding season, and nests have been found in at least 
one district. Benson (1935) collected a pigeon in the Navajo 
Mountains on June 20, 1935, and Presnall (1935) described the 
bird as uncommon on the western rim of Zion National Park. Ac- 
cording to Cottam (letter of May 6, 1940), S. D. Durrant obtained 
a specimen killed by a hawk near Hanna, Utah, in 1930 ; the speci- 
men is now in the collection of the University of Utah. Cottam 
(1941) summarized the status of the species in Utah, stating that 
apparently the birds were annual, summer residents in the moun- 
tains of southern Utah although probably never abundant any- 
where in the State. He reported 20 bandtails seen on July 24, 
1940, at Oak Grove and Bitter Creek Canyon, on the south slope 
of Pine Valley Mountain, one of them believed to be a juvenile 
only recently out of the nest. He said that W. G. MacFarland saw 
a flock at Buckboard Flat, Blue Mountains, in June 1939 at about 
8,000 feet elevation. Lee Griner is reported to have observed two 
flocks totaling 32 birds in the same mountain range 8 miles west 
of La Sal, Utah, on August 18, 1937, at 8,200 feet elevation, and 
a flock of 57 birds the same day a mile east of North Creek, at 
7,800 feet. Griner also saw the birds and found their nests in June 
1937 at Oak Grove, Pine Valley Mountain. Evidence obtained by 



THE BAND-TAILED PIGEON 21 

Griner from local observers indicated that the birds had occurred 
there each summer for at least the last 6 years. In June 1931 and 
1932, Cottam reported that flocks of bandtails caused damage to 
fruit in a small cherry orchard at New Harmony on the east slope 
of Pine Mountain Valley, and Oscar Deming stated that in 1937 
the birds frequented this same mountain from mid-May to Octo- 
ber. 

In Colorado the bandtail has been found in suitable foothill 
and mountain environment in the Transition and Upper Sonoran 
life zones on both sides of the Continental Divide. Numerous rec- 
ords obtained from the Colorado Museum of Natural History, the 
Colorado State Game and Fish Commission, and the field person- 
nel of the Forest Service and the Fish and Wildlife Service, and 
reports from farmers and sportmen show that pigeons have been 
seen at some time in 42 counties. 

Those counties in which the pigeon population is considered as 
moderate to large include Archuleta, Conejos, Dolores, Douglas, 
Eagle, Gunnison, Huerfano, La Plata, Las Animas, Montrose, 
Ouray, Pitkin, Pueblo, Rio Grande, and San Miguel. Counties 
with small to moderate populations include Alamosa, Bent, Boul- 
der, Chaffee, Clear Creek, Costilla, Custer, Delta, El Paso, Fre- 
mont, Garfield, Gilpin, Grand, Hinsdale, Jefferson, Mesa, Mineral, 
Montezuma, Park, Pt-io Blanco, Saguache, San Juan, Summit, and 
Teller. In several other counties pigeons have been seen occasion- 
ally and intermittently ; these include Elbert, Jackson, Lake, Lari- 
mer, and Weld Counties. 

Reports indicate extreme erraticism in much of the Colorado 
range ; in some counties pigeons are more numerous now than for 
some years, while in others none has been seen for several sea- 
sons, and in others there is a noticeable fluctuation in numbers from 
year to year. Occasional reports from old residents indicate that 
fifty years ago large numbers of pigeons existed in some parts of 
Colorado. Food conditions undoubtedly explain many of these var- 
iations. The bulk of the birds at the present time seems to occur 
south of a line drawn from Montrose to Pueblo along United 
States Highway 50. 

A Forest Service report for the year 1942 shows that pigeons 
were observed during that year on the Cochetopa, Holy Cross. 
Montezuma, Rio Grande, Roosevelt, San Isabel, San Juan, and 
Uncompahgre National Forests. The total reported as actually 
counted was 3,032. The San Juan Forest led with 1,200 pigeons. 

Colorado is the northeastward margin of range for the species, 
and for that reason variations would be expected to occur, espe- 
cially in counties along the east base of the mountains and at the 
north. Extensive field work is necessary before the status and 
range of the bandtails can be definitely outlined for Colorado. 

In Arizona, Swarth (1914) gave the range of the species as 
follows : 

A common summer resident in suitable localities throushout the S'tate; 
that is, in the higher mountain ranges, breeding usually above 6,000 feet. 
Has been found in summer on San Francisco Mountain, Mount Graham, 
and the White, Mogollon, Santa Catalina, Huachuca, and Carmelita Moun- 
tains. 



22 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

Specimens were collected north of Williams, and a nest was 
found south of that town, in 1937, and specimens were also col- 
lected in 1941. South and southeast of Williams bandtails are 
seen in fair numbers annually at many places in the Prescott Na- 
tional Forest. They also occur in some numbers in Oak Creek Can- 
yon south of Flagstaff. They have been observed on numerous 
occasions in the Coconino, Tonto, and Sitgreaves National For- 
ests. In June 1941 Charles C. Sperry found them abundant near 
Jacob's Lake in the northern end of the Kaibab National Forest. 

Clifford C. Fresnall noted several pairs near Hilltop, San Carlos 
Indian Reservation, on May 15, 1941, and in June of the same 
year saw paired pigeons at several places in the Fort Apache In- 
dian Reservation. The birds are also frequently observed near 
McNary and Springerville, and in many places in Greenlee Coun- 
ty. They nest in the higher elevations of the Crook National For- 
est near Globe, on Pinal Peak, in the Pinaleno Mountains of Gra- 
ham County (fig. 8), and in the Santa Catalina, Chiricahua, and 




B87091VI 

FiGrRE S. — Band-tniled pijieon range in the oak-and-pine-coverecl Pinaleno Moun- 
tains of Arizona. 

Huachuca ranges to the southward. Large numbers of them were 
seen on numerous occasions in the mountains between Patagonia 
and Nogales and in the ranges to the west of Nogales. 

Mrs. Florence Merriam Bailey (1928) says that in New Mexico 
"the occurrence of the Band-tailed Pigeon at any given place seems 
to vary in different years according to the food supply. This is 
especially true of the oak-covered mountain slopes which may 
swarm with the birds when there is a heavy crop of acorns and be 
entirely deserted during a whole year when the acorn crop is a 
failure." She adds that the birds usually occurred at 6,000 to 
8,000 feet, but frequently ranged much higher in the autumn, and 



THE BAND-TAILED PIGEON 23 

gives examples of observations in the Mimbres Range at 9,000 feet 
and on Pecos Baldy and in the Jemez Mountains at 10,000 feet. 

L. W. Simmons, Deputy State Game Warden, informed L. J. 
Merovka, United States Game Management Agent, that he saw 
bandtails near Tererro on June 25 and July 7, 1940. He estimated 
that there were 250 of the birds, and said that they remained in 
the area until September 12. He also reported 115 on Cow Creek 
near Upper Colomas and 100 near Lower Colomas on July 15, 
1940. W. M. Wilson wrote Merovka on October 2, 1940, that 
pigeons came into the Burro Mountains early in May 1939, nested, 
and left early in October. The birds concentrated well up the 
mountains. In 1940 pigeons were again present but widely scat- 
tered. Fred Sherman, of Deming, also reported to Merovka many 
past observations of bandtails in the Burro Mountains and the 
Black Range, and in 1940 some observations near White Signal in 
the Burro Mountains, and in the Black Range near Mimbres, Finos 
Altos, and Kingston. 

H. Garvin Smith, of the United States Forest Service, has sup- 
plied from his notes records ranging back to 1919, when he stat- 
ed that bandtails in large flocks almost ruined the cherry crop 
at High Rolls and Orchard Park. In 1925 he said that in the Black 
Range pigeons were everywhere, gorging on pifion nuts; that 
thousands passed over his camp flying to roost ; and that they were 
also reported from the San Mateo Mountains. By contrast, in 
1926 his notes record that pigeons were very scarce in the Mogo- 
llons and on the west side of the Black Range. He also observed 
nesting birds in the Magdalena Mountains in 1933 and 1934. 

David 0. Scott, ranger in the Carson National Forest, Penasco, 
wrote that he saw his first pigeon for the year on May 19, 1941 ; 
he concluded that bandtails were relatively scarce in that area, 
though there w^ere at times small groups of them in Santa Bar- 
bara Canyon and near Rio Pueblo. Dean M. Earl, of Carlsbad, 
reported that the only time he saw pigeons on the Lincoln Nation- 
al Forest was in the fall. 

Cliff'ord C. Presnall and other employees of the Fish and Wild- 
life Service saw about 400 bandtails on May 30, 1941, near White- 
tail School, on the Mescalero Indian Reservation, and single birds 
at other places on the reservation. D. Wood, Jr., one of the ob- 
servers, said that during previous employment on that reserva- 
tion, between 1931 and 1933, he saw many pigeons in small flocks, 
and that he believed they nested near Carizzo Springs. They have 
been reported from Sarca Canyon, Capulin Springs, and other 
places in the Sandia Mountains near Albuquerque. 

L. J. Merovka said that band-tailed pigeons are usually fairly 
numerous around Bland, Senorita, and La Jara Canyons in the 
Jemez Mountains, and that he saw 500 in the latter place in Sep- 
tember 1943. He also reported a flock from Stone Canyon in the 
Black Range, and on June 13, 1944, he noted a flock near Amalia, 
on the Costilla River, a few miles south of the Colorado State line. 

Few early references to the range of the species in western Tex- 
as were found. One specimen from Uvalde was examined : Van 
Tyne and Sutton (1937) listed the birds from Presidio and Brew- 



24 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

ster Counties; and Oberholser (1902) reported them from the 
Chisos, Guadahipe, and Davis Mountains. Ray Williams, State 
Warden, of Alpine, Texas, wrote Merovka that in the Trans- 
Pecos area bandtails ranged from 5,700 to 8,700 feet elevation in 
the Sierra Madre and the Glass Mountains of Pecos and Brewster 
Counties, in the Eagle Mountains of Hudspeth County, in the Di- 
ablo and Guadalupe Mountains, and in the Davis and Chisos rang- 
es. 

Casual records are listed by Carter and Trentoon (Nice 1924), 
including that of one bandtail killed near Crescent, Logan County, 
Okla., in 1905. Schufeldt (1912) reported the taking of a single 
specimen near Englevale, N. Dak., on June 2, 1912. Alcorn (1941) 
reported the collecting of the first specimen from Nevada near 
Fallon on October 17, 1940. 

In Mexico, according to information furnished by Frederick C. 
Lincoln, bandtails nest as far south as Durango (Otmapa Ranch), 
Puebla (Las Vegas), probably Vera Cruz (Jalapa), and the San 
Jose Mountains in Sonora. They are also found in Baja Califor- 
nia. Bandtails were seen by the writer in considerable numbers 
in June and July 1942 along the Rio de Los Alamos south of No- 
gales, Sonora, and near Cananea, and along the Rio Babasac near 
Cocospera. American sportsmen report them widely distributed 
in the mountains of northern Sonora. 

WINTER RANGE 

In the northern part of their range band-tailed pigeons are 
largely migratory, only a few scattered birds remaining during 
the winter in sheltered locations. An Audubon Society Christmas 
Bird Census in the Puget Sound area in 1935 tallied 30 pigeons, 
and W, H. Ransom, in a letter of April 15, 1939, said that "small 
bunches quite regularly winter around Medina and Bellevue, 
Wash., along the east side of Lake Washington." On January 18, 
1928, near Agnes, Oreg., a flock of about 150 pigeons feeding on 
madrona berries was reported to S. G. Jewett, and a few days 
later he found pigeon feathers along a trail in that vicinity. Jewett 
also observed a large flock feeding in stubble near Wolf Creek, 
Oreg., March 20, 1925, and A. W. Moore saw two birds on the Ne- 
halem River in Oregon on February 22, 1940. 

In California relatively few pigeons winter in the Sierras north 
of Yosemite National Park or in the coastal mountains north of 
San Francisco Bay. Hence the great majority of the bandtails 
breed from San Francisco Bay northward and concentrate in win- 
ter in west-central and southern California. The chief wintering 
area lies in the Sierras from Yosemite southward to Mexico, and 
in the coastal mountains from Santa Cruz and Santa Clara Coun- 
ties southward to the Tehachapi range and Los Angeles, usually 
centering in Monterey, San Luis Obispo, and Santa Barbara Coun- 
ties. In some years, the major wintering grounds may be from 
Los Angeles and San Bernardino Counties southward. 

Moderate but irregular numbers of pigeons occasionally winter in 
the counties just north of San Francisco Bay, and casual winter 



THE BAND-TAILED PIGEON 25 

resident bands may at times frequent the slopes on either side of 
the Sacramento Valley to Redding. 

The wintering range in California is closely correlated with 
food supply, but concentrated shooting during any one season un- 
doubtedly causes the population of the area affected to diminish 
during succeeding seasons even though food is abundant. Mrs. 
Mary Bartol (1940) described the (to her) immense population 
of the Mount Pinos area in Ventura County in 1933, and remarked 
that only one year since then had been characterized by a crop of 
pihon nuts and a consequent large population of pigeons ; she also 
said that in one of the intervening seasons the shooting area for 
southern California was in the oak country of the Santa Ynez 
Valley of Santa Barbara County. 

Fred H. Post, State Game Warden, of Salinas, Calif., told the 
writer that in the winter of 1932-33 the birds had been very abun- 
dant in the mountains of that county. In 1933 food was scarce 
in the high areas and the birds scattered everywhere. In 1934 
food was again plentiful and the pigeons were abundant in De- 
cember. Other State game wardens in California have described 
to the writer large midwinter concentrations in areas of abun- 
dant food and have told of the annual fluctuation in abundance 
and distribution occasioned by variation in the food supply. 

Grinnell and Wythe (1927) list the bandtail as a winter resi- 
dent in the San Francisco Bay region, irregular but sometimes 
occurring in large numbers and in widespread areas. Willett 
(1933) records it as sporadically plentiful in southern California 
in foothills and mountains up to snow line, and also as having 
been observed on Catalina Island in 1932. 

In Arizona, New Mexico, and Texas band-tailed pigeons from 
the northward and from the mountain ranges of those States move 
toward their southern boundaries or on into Mexico to winter. 
Variable numbers, however, remain in mountain ranges near the 
border as food supply and weather conditions permit. Mrs. Bailey 
(1928) reports them as resident some winters below the 5,000 
foot elevation near Clifi", N. Mex., and quotes Stokley Ligon as 
having observed them on Haut Creek, Socorro County, at 7,500 
feet during the winter of 1912-13. Ligon (1927) says that a heavy 
crop of pihon nuts has in a few instances held them throughout 
the winter in the Black Range. 

Fred Pickens, Deputy State Game Warden, and William Wood, 
of the United States Forest Service, saw 35 bandtails in the San- 
dia Mountains near Albuquerque, N. Mex., on December 30, 1940, 
and on the same day Fred Johnson of the same Service saw fully 
100 in a neighboring canyon. 

In Texas, Ray Williams, of Alpine, reports, "I have seen them 
stay in the Chisos Mountains all winter when there is plenty of 
food," and on March 22, 1941, G. W. Peterson, Deputy State Ward- 
en, of Nogales, Ariz., wrote that "band-tailed pigeons stayed 
with us all winter . . . Acorn crop was very good last year." 

South of the Mexican border, according to Frederick C. Lincoln, 
the band-tailed pigeon winters south to Guatemala and Chiapas. 
A subspecies, Viosca's pigeon (Columba fasciatu vioscae), appar- 



26 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

ently nonmigratory, is restricted to southern Baja California, and 
other subspecies occur in Central America. 

MIGRATION 

Periods and routes of migration of the bandtails are strongly 
influenced by availability of food and by weather phenomena, es- 
pecially temperature and rainfall. Owing to the vast areas of 
wilderness through which the birds pass, it is difficult to trace 
their movements, and over much of their range they appear to 
move in small flocks which often may pass unnoticed. 

SPRING 

Search of the literature gives the following "earliest" dates for 
the appearance of the bandtail in the northerly parts of its range : 
Colorado, Beulah, May 7 ; Oregon, Mercer, March 5 ; Washington, 
Clallam Bay, April 9; British Columbia, Courtenay, May 31. 

Leaving the wintering grounds in west-central and southern 
California late in winter, the northward-bound pigeons move 
slowly along the foothills of the Sierras and along the coast moun- 
tains. Frequently enormous flocks concentrate in areas of abun- 
dant food supply. The date of appearance at various California 
points varies with the season, but often large flights appear in 
the live-oak-covered foothills east of Sacramento late in February 
or early in March. Food supplies to a marked degree seem to con- 
trol the dates and routes of movement. 

Grinnell (1898) observed a large flock that remained until mid- 
June 1895 feeding on acorns near Pasadena, and Van Denburgh 
(1899) reported the birds as frequently remaining in large bands 
until mid-May near Palo Alto. Evidently migration is erratic and 
influenced by a number of factors which in few instances are well 
understood. 

R. J. Little, State Game Warden, reported in 1935 that the pi- 
geons usually arrived in Butte and Sutter Counties, Calif., in Feb- 
ruary. On February 15 he saw about 4,000 birds near Bangor, 
and on February 20 about 9,000 arrived at the small mountain 
uplift known as the Sutter Buttes, where they fed on acorns or on 
grain stubble in the adjacent valley fields. Nelson Poole, State 
Warden, described a large flight of pigeons, which late in Feb- 
ruary and early in March 1935 invaded the oak-covered hills be- 
tween Sacramento, Folsom, and Auburn. These were feeding as 
they made their way northward, and soon left the Sacramento 
area. Near Auburn, however, although the numbers diminished, 
pigeons were numerous until late in April. Just as the cherry 
growers began to worry about their presence they suddenly moved 
on, and no more than normal numbers remained. 

In the border States of Arizona, New Mexico, and Texas, it is 
more difficult to record the spring migration, as in favorable areas 
some of the birds frequently spend the winter. Ralph Morrow, 
Deputy State Game Warden, reported his first 1940 bandtail late 
in February in the Chiricahua Mountains. The usual spring rec- 
ords, however, are of birds observed late in April or early in May. 



THE BAND-TAir.ED PIGEON 27 



The autumn migration also is strongly influenced by weather 
conditions and availability of food. After the major breeding 
season, pigeons flock together and move about locally following 
successions of food crops. There is also a definite vertical move- 
ment from the mountains to foothill or valley lands. This verti- 
cal migration varies greatly in season and in the Sierras of Cali- 
fornia seems to be stimulated by snow, storm, and cold weather 
at the higher elevations as well as by local food diminution, some- 
times not occurring until December or January. 

The latest fall records in the North found in the literature 
(Bent 1932) are for British Columbia, October 29, and for Wash- 
ington, October 29. For Oregon, Bent (1932) lists the "latest" 
record as Newport, October 28. Jewett, in a letter of June 10, 
1937, gives the following dates : Harney County, October 19 ; Mult- 
nomah County, October 11 ; Tillamook County, September 17; and 
Jackson County, October 6. In 1937 the writer spent the period 
of September 10 to September 25 in western Washington. During 
that time moderate concentrations of pigeons were observed feed- 
ing in the area immediately adjacent to Puget Sound from Blaine 
and Sumas in Whatcom County southward to Olympia, and north- 
ward to Dungeness in Clallam County. Local game wardens in 
those areas, however, said that there had been a considerable dim- 
inution in the population of the bandtails late in August and in 
the early days of September. South of Hoquiam and west of Olym- 
pia in the area between the Cascades and the ocean almost no pi- 
geons were remaining on September 15, and local authorities stat- 
ed that the mass of the population had departed by September 1 
to 10. 

The southward migration in Oregon normally begins during the 
last days of August and is M^ell completed by September 20. Along 
the coastal mountains definite flyways exist, and from stations on 
them during the period of major migration it is possible for an 
observer to note band after band of pigeons passing south from 
daylight until dark. According to a number of observers, these 
birds "feed" their way along, the roosting place each evening 
lying farther south than that of the night before. 

A. S. Einarsen, in a letter of October 3, 1944, described a Wash- 
ington pigeon migration as follows : 

On a trip to the State of Washington on September 25, I ran into a 
migration, and at one point on the Cowlitz River north of Castle Rock, 
counted 960 birds which came to a point, settling in a few roosting trees 
and, incidentally, dropping down for a drink of water in a backwater pool 
before continuing their migration southward almost immediately. In my 
experience I had never before seen such a steady drift of bandtails at any 
one point as large or continuous. 

And in a brief special report dated March 30, 1943, Einarsen 
describes another migratory concentration point in Oregon thus : 

They may pause for seA'eral weeks at some gravel bar or watering hole, 
and often other flocks join them. These facts have been confirmed by field 
observations at Pigeon Butte in Oregon, 12 miles south of Corvallis and 3 
miles west of Bruce Station, one of many such points. Here an annual con- 



28 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

centration exists from about August 25 to October 10, enlarged by migra- 
tions from areas far to the north. Pigeons come into this area to drinli at a 
source of water supply and fly through a pass in great numbers. 

From reports of local observers and from personal field obser- 
vations the writer traced the 1937 migration from Benton and 
Tillamook Counties southward. From late August to mid-Septem- 
ber the bulk of the bandtail population ranged from Corvallis to 
the vicinity of Coquille and Myrtle Point. The writer followed the 
flight from September 29 to October 4, and in the northern part 
of the area found only scattered small groups. In the Myrtle 
Point-Powers district a light population was found, where 20 
days earlier several cooperators had reported enormous numbers. 
From the Roseburg-Coquille highway in Oregon south to north- 
central California stretches a wilderness that is so large and so 
sparsely traversed with modern highways that proper analysis of 
the bandtail migration through it is difficult. At numerous places 
within this great district the south-bound pigeons are reported 
almost annually as congregating in great numbers. In the 1937 
field trip the writer continued southward along the coastal route. 
At Brookings, Oreg., ranchers along the Chetco River reported 
on October 4 that pigeons had been very abundant in their stubble- 
fields until the onset of a v^ind and rain storm on September 29 
and 30. Later Einarsen wrote that from October 19 to 23, 1937, 
there had been a heavy concentration of pigeons feeding on ma- 
drona berries in the Applegate section of the PvOgue River Valley. 

In California it appears that the band-tailed pigeons on the 
coastal side of the northwest coast mountains congregate in flocks 
and commence to move south almost concurrently with the begin- 
ning of the movement in northwestern Washington and Oregon. 
State game wardens in Humboldt County, Calif., reported that 
late in August 1937 there had been a large population of pigeons 
in the area south of Fortuna and west of Garberville that had 
moved on southward down the coast prior to the arrival of the 
northern birds. Captain Harp, State Warden, said that in coastal 
Humboldt County the autumn withdrawal usually had begun by 
September 1 and that by mid-October of normal years none of 
the birds were to be found. 

In 1937 the flights of bandtails from Oregon and Washington 
were apparently reaching northeastern Mendocino, eastern Hum- 
boldt, and Trinity Counties by early October, as during that month 
predatory animal hunters at various localities in the area reported 
pigeons in great numbers, and by late October considerable flocks 
were observed in southern Mendocino, Lake, and Napa Counties. 

In the Sierras the autumnal flights appear to remain normally 
at high elevations, hence study of them is difficult. Deer hunters 
and State game wardens often report moderate concentrations in 
the area near Mount Shasta and west of Mount Lassen, and at 
times fairly large feeding concentrations are noted farther south 
along the range. In many seasons, however, the Sierra pigeon pop- 
ulation remains at from 3,000 to 7,000 feet elevations until Decem- 
ber. 

In 1939 the autumn on the West Coast was very mild, and the 



THE BANU-TAILED PIGEON 29 

band-tailed pigeon migration was delayed considerably in com- 
parison with average seasons. Pigeons were present in central 
Oregon in fairly large numbers all through September and into 
October, and a few yet remained in early November. Grinnell, 
Bryant, and Storer (1918) said that "in event of continued favor- 
able weather, the birds will often continue in their summer haunts 
until October or even November . . ." if food is available in these 
haunts. 

Mrs. Bailey (1928) compiled from the reports of several ob- 
servers a number of autumn records for New Mexico, among which 
were the following: September 15 and 16, 1914, Upper Blue River 
Canyon; September 17, 1915, Diamond Creek; and November 10, 
1914, 20 miles east of Silver City. W. M. Wilson, of Silver City, 
informed Lawrence J. Merovka that in 1940 pigeons remained 
in the area near Tyrone, N. Mex., until November 10, and that 
others were reported in the Black Range ; this, he said, was fully 
a month later than the noi'mal date of departure. 

ROUTES OF MIGRATION 

Except for evidence accumulated through observation of large 
flocks of band-tailed pigeons, there is little direct information con- 
cerning the routes of migration used by these birds. They appear 
to be very erratic, following those routes that offer satisfactory 
food for the migrating flocks. 

Extensive banding of these birds is needed to furnish more ex- 
plicit evidence relative to their migrations and to determine def- 
initely the winter range of the pigeons produced in the major 
breeding areas. Reed Ferris, of Beaver, Oreg., discovered that 
bandtails are readily trapped when feeding on the ground, and 
he has banded more pigeons than any one else. Banders in pigeon 
country should make every effort to mark these birds. 

Up to May 1, 1940, only 185 band-tailed pigeons had been band- 
ed, and only 5 return records were in the files of the Fish and 
Wildlife Service. Three of these were of birds that had been band- 
ed by Ferris at Beaver, Oreg., late in May 1932. One was shot 
at China Camp (Calaboose Canyon), Monterey County, Calif., in 
December 1932 ; another in the "Monterey Hills" on December 
10, 1933 ; and the third, 15 miles east of Gonzales, Monterey Coun- 
ty, on the same date. A pigeon banded at Carmel, Calif., in July 
1937 was shot on December 14, 1937, near Atascadero, Calif., and 
one banded at the State Game Farm, at Chino, Calif., in June 1937 
was found dead near Cucamonga, Calif., during the same month. 

VALUE AS A GAME BIRD 

Grinnell (1913) stated that "the value of the band-tailed pigeon 
as a true game bird is to be conceded without argument. Its pur- 
suit is of a different type from that offered by any other game spe- 
cies." "Stillhunter" (1907), an anonymous writer in southern Cal- 
ifornia, said that the best place for hunting pigeons was near a 



30 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

dead tree where the birds ahght, and that a .22 or .25-20 rifle 
should be used so that single birds could be procured without scar- 
ing the flock ; but that for sneaking up on the flocks a "duck gun" 
was used. Ten pigeons were considered a good day's bag. If the 
flesh was strong, owing to an acorn diet, soaking it in brine fla- 
vored with vinegar or lemon would remove the disagreeable taste. 
Mrs. Mary Bartol (1940) describes the hunting of bandtails in 
southern California, noting the long-range shooting and conse- 
quent wasting of ammunition and crippling of birds. She men- 
tions also the diff"erence in flavor of the flesh caused by the birds 
eating different foods. 

Since shooting seasons were resumed in 1932 after 20 years of 
total protection, several changes have occurred in the annual reg- 
ulations, each tending toward permitting shooting at a period 
when more pigeons might be killed. The history of these seasons 
from 1932 to 1945 is abstracted. 

Arizona.— 19S2-1^M, December 1-15; 1935-1938, October 16- 
30 ; 1939, October 1-15 ; 1940-1941, September 16-30 ; 1942-1945, 
September 16-October 15. 

California.— 19S2-ld41, December 1-15; 1942-1945, December 
1-30. 

Colorado. — 1944-1945, Counties of Archuleta, Dolores, Huerfano, 
La Plata, Las Animas, Montrose, Montezuma, Ouray, San Juan, 
San Miguel ; and the drainage of the North Fork of the Gunnison 
River in Delta and Gunnison Counties, September 16-October 15. 

New Mf'.T/co.— 1932-1934, November 1-15; 1935-1939, October 
1-15; 1940-1941, September 16-30; 1942-1945, September 16-Oc- 
tober 15. 

Oregon.— 19S2-19S8, October 16-30; 1939-1941, September 1- 
15 ; 1942-1945, September 1-30. 

Washington.— 19Z2-19M, October 16-30; 1935-1941, September 
16-30 ; 1942-1945, September 16-October 15. 

The very slow rate of increase of this pigeon was apparently 
sufficient to maintain its population under early conditions, but it 
is amazing that the species has stood up so well and so long against 
the combination of modern firearms, good roads into wilderness 
areas, speedy automobile transportation, and the continued trend 
toward shooting seasons during the period when the birds are 
most abundant. 

In winter, California is largely responsible for the fate of the 
bulk of the present band-tailed pigeon population of the Pacific 
Coast. The first open shooting season after 20 years of protection 
in that State occurred in 1932, continuing from December 1 for 
15 days. The writer was not afield during that period but has re- 
ceived numerous verbal reports from hunters. In the Santa Lucia 
range of Monterey County, Calif., in the Monterey division of the 
Los Padres National Forest, there was, on December 1, 1932, a 
heavy wintering population of pigeons. The road leading to Tassa- 
jara Hot Springs resort, at a place close to 5,000 feet elevation, 
crosses a narrow saddle locally called China Camp, which sep- 
arates two deep canyons, Miller and Calaboose. Oak and pine trees 



THE BAND-TAILED PIGEON 



31 



are abundant on the slopes near the summit of the mountains, 
and a short distance north of the saddle several wide and fairly 
open oak flats occur. Madrona and manzanita are abundant along 
the canyons, and water is available in at least one of the canyons. 

According to sportsmen, on the first day of the 1932 open sea- 
son countless pigeons flew across China Camp saddle from Miller 
to Calaboose Canyons and back again. Hunters quickly congre- 
gated and band-tailed pigeon shooting became once again a lead- 
ing sport. Owing to the roughness of the terrain it was extreme- 
ly difficult for the game officers properly to patrol the pigeon area 
and no criticism of them is implied. Certain hunters told the writ- 
er of watching gunners who shot all day long, assisting others 
to obtain their limits, and left piles of birds on the ground. 

In 1934 there was again an ample food supply in the area ad- 
jacent to China Camp saddle, and a large flight of pigeons gath- 
ered there and in the adjacent flats known as Chews Ridge and 
White Oak Flat. The shooting season opened on December 1, and 
the writer visited the area on December 7, viewing the flight 
and methods of hunting and conferring with wardens and 
hunters. Since the birds flew to and fro continuously, it was prac- 
tically impossible to estimate the numbers present, but the flight 
on December 7, after 6 days of shooting, was not unusually large. 
When there was little shooting at China Camp saddle, the birds 
flew up and down the two canyons ; when numbers of hunters oc- 
cupied the saddle, the birds scattered out and flew widely over the 
mountain-top flats. 

On December 9 (Sunday) the writer again went to the shooting 
area. The entire mountain abounded with hunters, automobiles, 
and guns. Gunners covered every opening in the forest and shots 
poured into the pigeon flight from all directions and elevations. 
At China Camp saddle the picture was not pleasant, as ten men 
shot where one would have been enough. Sportsmanship was vir- 
tually absent. In the continued fusillade of long-range shots, many 
wounded pigeons plunged to earth or fluttered down to alight in 
shrubbery or trees. Owing to the steepness of the slopes and their 
dense vegetation, the loss of birds was very high, possibly as many 
as five pigeons being lost or mortally wounded for every bird 
picked up by a hunter. 

State game wardens made partial surveys of the kill during this 
1934 season, usuafly counting only the full-limit bags taken off" the 
area. On December 9, however, the writer assisted three State 



Tat.le 1. — Data on pigeon kill, Tasmjara District, Monterey County, Calif.. 19S'i 



Date 


lypQ of count 


Number of 
hunters checked 


Number of ba? 
limits checked 


Recorded 
kill 


Deo 1 




185 
412 
•123 
60 
143 
830 


1.5(1 

28lt 

60 

30 

85 

445 


1.500 


Dec. 2 


do 


2.800 


Dee 5 


do 


600 


Dec. 7. .. 


. . .do 


300 


Dec 8 


do 


850 


Dec. 9 




6,404 









695766°— 47— 3 



32 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

game wardens in a complete roadside inspection at the foot of the 
mountain. The record of kill on the various days is given in table 
1, the figures for December 1 to 8 being generously furnished by 
the State game wardens covering the district. 

In conversation with wardens and several scores of hunters, 
December 7 to 10, 1937, the writer learned the reactions of the 
gunners of this district. The vast majority denounced the reckless 
shooting and the wastage of birds. Numerous hunters expressed 
the opinion that the band-tailed pigeon could not maintain its ex- 
istence in the face of such shooting as occurred in the China Camp 
area in 1932 and 1934. Fully 20 percent voluntarily declared for 
regulations to reduce the kill. 

Since 1934 there has been no such large concentration of birds 
at that place. In some seasons there has been a shortage of food, 
but to the writer it seems that some of the scarcity of birds can 
be attributed to the terrific shooting of 1932 and 1934. It is for 
the welfare of the species, therefore, that usually the winter pop- 
ulation in California has not concentrated in any such small areas. 

The writer observed shooting practices on Whidby Island, in 
the Snohomish River delta area, and at other places in Washing- 
ton in 1937. There was no parallel to the California situation, the 
shooters being widely scattered. The most frequently observed 
method of shooting, however, left much to be desired in the matter 
of "sport," as hunters lay in wait and shot at pigeons that perched 
in the tops of tall Douglas fir snags. 

The food of the pigeons during the general period of shooting 
greatly aff'ects the flavor of the flesh of the birds. In the Puget 
Sound area during the September open season the major food is 
peas from the stubblefields, and the birds are said to be of fine 
flavor. Mrs. Bartol (1940) pointed out the difference in flavor 
of California birds that fed on pihon nuts from those that ate 
acorns. By far the majority of pigeons killed in California have 
fed on acorns or on the fruits of the madrona. Their flesh is fla- 
vored by those foods, and old birds are very tough. Entirely too 

Table 2. — Baml-tailrd i)if/eo» kiU iv CaJipirnia, 193'/ and 1935: State ami thir- 
teen Jeadiiiff counties; I 



State and county 



CALIFORNIA (ENTIRE STATE) 
Counties: 

Monterey 

Santa Cruz 

San Luis Obispo 

Santa Clara 

Calaveras 

Tulare 

Santa Barbara ■ 

Los Angeles 

Sonoma 

Fresno 

San Diego 

Humboldt 

Riverside 



51.05fi 

21,961 

5,81!) 

1,216 

1,075 

567 

528 

520 

515 

512 

506 



19.35 3 



34.136 

10.855 

1,439 

608 

542 



454 

678 



1.048 
405 
405 



^ Compiled from hunting'-license application blanks by the California Division of Fish 
and Game. 

= Published in the California Conservationist 1 (8): 3. August 1936. 

8 Published in the California Conservationist 2 (8): 20. August 1937. 



THE BAND-TAILED PIGEON 



33 



many of the birds killed are destined ultimately for the garbage 
can ; no specific evidence can be given, but at numerous times the 
writer has heard statements that could have no other meaning. 
A relatively small percentage of the gunners hunt pigeons be- 
cause they like to eat the birds; the majority are in the field for 
the sport of shooting, the thrill of killing game, and the enjoy- 
ment of the high wild country. 

In table 2, based on data compiled by the California Division 
of Fish and Game, is given the band-tailed pigeon kill for the sea- 
sons of 1934 and 1935, as reported on applications for hunting li- 
censes for the ensuing year. The table gives the total for the en- 
tire State and for each of the 13 leading counties. The figures are 
conservative, for few hunters will report more than they actually 
kill and some make no report at all. Table 3, based on data from 
the same source, shows the 1934 and the 1935 kill of each of Cal- 
ifornia's game birds. 

Tables. — (lame bird kill in. Valifoniia, I'JS'i and 1935^ 



Game bird 



Band-tailed pj^eoii .... 
Western nuniinin^ dove. 

Quails, all species 

Pheasant 

Ducks, all species 

Geese, all species 



51,(156 


34,136 


580,110 


539,722 


500,481 


916,589 


25,220 


40,616 


389,247 


304,882 


40,278 


35,096 



• Compiled from hunting license application hlanks by tlie California Division of Fish and 
Game; published in the California Consei-vationist 2 (s) : 20, Aujrust li»37. 

Writing of Pigeon Butte, near Corvallis, Oreg., Einarsen says : 

For years this pass has been popular with hunters. Because of the ter- 
rain and the usual eagerness of the hunter, most of the shots are at long 
distances, which results in a crippling loss averaging more than 60 per- 
cent of the pigeons bagged. When the number of birds killed (at each pass) 
is more than 600, as in 1942 at this pass, the seriousness of the slaughter 
can be recognized. 

The band-tailed pigeon will never have wide abundance. It is hunted on 
uneven ground, where a high percentage of crippled birds are lost and little 
effort is made to recover them. Five months after the season it was still 
possible to pick up birds which had died of injuries in large numbers 
around Pigeon Butte. 

Far too few hunters know that band-tailed pigeons decoy well, 
and in many localities far better sport could thus be obtained with 
an accompanying great reduction in the number of crippled or 
dead birds lost. Decoying the bandtails into forest openings or 
fields is far preferable to shooting them at waterholes or gravel 
bars, pot-shooting perched birds out of dead snags, or to the aver- 
age mountain-pass shooting in rough, brushy terrain. 



PRESENT STATUS 



Under conditions that prevailed between 1932 and 1942 it ap- 
peared that the band-tailed pigeon held its own under the 15-day 



34 xXORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

shooting season and low (10-bird) possession bag limit; there 
are those who state emphatically that in certain localities it was 
not maintaining itself, but was slowly decreasing. It is too soon 
to analyze the effect of the 30-day shooting season established in 
1942, as wartime conditions of ammunition shortage and gasoline 
and tire rationing markedly affected pigeon shooting. Where num- 
bers of the birds are to be found not far from hunting centers 
there may have been some reduction, but in other districts pigeon 
concentrations occur only in wilderness areas where during the 
war period few hunters could reach them. 

In several States where there is an open season, it is set at a 
period when many of the birds have already moved southward, 
and there has been local clamor for a change of the season to co- 
incide with the presence of peak populations. Such requests for 
changes in shooting seasons should be closely scrutinized for, if 
granted, they may lead to great increases in the kill in States in 
which the pigeon population is too low. Seasons of good shoot- 
ing result in a great increase in the number of shooters afield ; re- 
ports of poor shooting soon result in decreased hunting. 

Earlier summaries of the status of the band-tailed pigeon are 
those of Chambers (1912), Grinnell (1913). and Taylor (1924). 
Owing to the extreme erratic nature of the species, accurate 
appraisal of its status over all the range is extremely difficult. The 
writer's own observations and reports of others are summarized 
for their interest in this connection. 

A. S. Einarsen in a report, dated January 6, 1936, stated that 
"regardless of what may be said to the contrary, the population 
of band-tailed pigeons in the Eraser Valley, the Puget Sound coun- 
try, the islands of Puget Sound, the river valleys west of the Cas- 
cade Mountains in Washington State, and in the favored nesting 
areas west of the Cascades in Oregon shows a tremendous de- 
crease from that of the 1934 season . . ." Pearse (1940) said that 
bandtails were reduced to a precarious state on Vancouver Island 
and blamed overshooting in the winter habitat. Miller, Lumley, 
and Hall (1935), writing of San Juan in Puget Sound, state that 
numbers are decreasing rapidly, 

Einarsen. in a letter of October 3, 1944, reported that "the mi- 
gration in Oregon, generally speaking, has not been as great as in 
previous years, but there have been a few birds passing through 
throughout the entire month." 

Eield investigations in the Puget Sound area between Septem- 
ber 13 and 25, 1937, indicated that the population of bandtails 
was from 50 percent to less than 10 percent of what it had been 
at the same date in 1936. This means little, however, in the ab- 
sence of correlated data on weather and food conditions for the 
period concerned. In Whatcom County the 1936 population was 
reported as hea\T, that of the spring of 1937 as normal, and that 
of the autumn of 1937 as very light. Snohomish County estimates 
placed the 1937 population at about 10 percent of that of 1936. In 
Skagit County, although scattered groups of birds were present, 
no pigeon concentrations were observed in September 1937. 

San Juan reports indicated a normal 1937 season, except that 



THE BAND-TAILED PIGEON 35 

the autumn migration had occurred earlier than usual. Similar 
reports were obtained from Thurston, Grays Harbor, and Mason 
Counties. Pacific County reported an increased population in the 
spring of 1937 and an early autumn departure. Pierce County 
had a very large spring population, which dispersed to nest and 
did not reassemble. Lewis County experienced a marked increase 
during the 10 years previous. In Clallam County there were large 
spring concentrations during 1935, 1936, and 1937, and in some 
arf;as an abundant summer population also. Officials agreed that 
there was an increase in the area known as the Black Hills dis- 
trict near Elma, Shelton, and Olympia. Game protectors in all 
the counties named (except two) reported annually increasing 
shooting. 

On September 27, 1937, United States Game Management Agent 
Gerow said that although numerous complaints of crop depreda- 
tions had been received, the consensus of opinion was that pigeons 
were decreasing in numbers in Oregon. Alex Walker, of Beaver, 
Oreg., reported on September 30, 1937, that at best the pigeon 
population in Tillamook County was standing still and on March 
30, 1940, he was of the opinion that the birds were definitely de- 
creasing. Coincident with the late 1939 autumn migration in Or- 
egon, a considerable increase in hunting was reported in some 
parts of the State. 

Allen C. Oberle, of La Verne, Calif., wrote on February 15, 1935, 
that in the San Dimas district of Los Angeles County there was 
a flock of about 350 pigeons where in 1926 there had been only 
three pairs. Lawrence W. Saylor (formerly with the Fish and 
Wildlife Service) wrote on January 15, 1940, that bandtails did 
not become at all common in the Ben Lomond section of the Santa 
Cruz Mountains of California until about 1932, when in a few 
seasons the flocks increased from a dozen birds to a hundred or 
more. Warden C. E. Holladay, of San Jose, Calif., said in 1934 
that the pigeon population of Santa Clara County had doubled in 
6 years. 

From these and numerous recent reports it appears that the 
breeding population of band-tailed pigeons in California is hold- 
ing its own in all sections ; each year there are reports of pigeons 
nesting in areas where they had not been known to nest for years, 
and these seem to indicate that the California breeding population 
is slowly increasing and extending its range. 

In Colorado, Bergtold (1928) called the bandtail an Infrequent 
summer resident, and in the Denver area Niedrach (Niedrach 
and Rockwell 1929) reported the birds as regular summer resi- 
dents in small numbers. Studies of the past two or three seasons 
substantiate a statement by E. R. Kalmbach to the effect that 
these birds seem to have decreased in numbers in northern Colo- 
rado, for few birds have been observed in the northern half of the 
State. On the other hand, in the southern part of the State and 
in a few more northerly localities, the birds seem slowly to have 
increased. Frank F. Foley, United States Game Management 
Agent, believes that there has been a constant but small annual 
increase for the past 10 years, especially in southern Colorado; 



36 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

and Barry C. Park, of the United States Forest Service, says that 
the pigeon in Colorado is increasing slowly but constantly and is 
extending its range each year. 

Inquiry among game officials and sportsmen in Arizona from 
1938 to 1941 leads the writer to believe that the pigeon population 
of that State is slowly increasing. Recent seasons of drought 
caused food failure in many parts of Arizona, and this resulted 
in very erratic movements of the birds. Very few birds are shot 
in this State as the season opens after the bulk of the pigeons of 
the southern part of Arizona have moved into Mexico, and the 
birds are widely scattered. 

Ligon (1927) said that the bandtails did not appear to increase 
in New Mexico but remained at about a constant level, and that 
the population was insufficient to justify an open shooting season. 
No recent general statement is available, but the reports from 
various localities in New Mexico indicate that the bandtail there, 
as in Arizona, may locally be slowly increasing. 

From Texas, Ray Williams, of Alpine, wrote in November 
1940 : "For 25 years I have observed them in this territory and I 
do not see any increase in their numbers. They will never be too 
plentiful . . . They . . . just about hold their own." 

Winter populations are most difficult to appraise owing to the 
wandering habits of the birds and to the wide range of country 
in which concentrations may occur. No recent information from 
wintering range indicates a decrease, nor is there any reliable 
evidence of any marked increase. 

Considering actual pigeon populations alone, without reference 
to any other phase, it is felt that only in the Pacific Coast States 
are there sufficient numbers of birds to justify hunting. In Ari- 
zona, New Mexico, Colorado, Texas, and Utah, the habitats and 
habits of the birds may be such that few birds are killed and little 
damage results from open shooting ; yet there is danger of again 
decimating the species. Shooting must be rigidly controlled. Local 
increases may easily be decimated by concentrated shooting in 
more southerly migratory or wintering areas. 

It is of the utmost importance that Federal and State wildlife 
officials proceed with caution in liberalizing shooting privileges on 
this species in any part of its range. It is even more important 
that they carefully note the year to year status of the species over 
its entire range so that the need for reduction in numbers made 
necessary by local conditions may be anticipated and hunting con- 
trol measures applied in sufficient time to prevent gross destruc- 
tion that would require a total closing of the season such as was 
necessary in 1913. 

MANAGEMENT POSSIBILITIES 

It is inevitable that during a study such as this many questions 
that relate directly to management of the species should be raised. 
From a few sources have come direct inquiries concerning meth- 
ods that might permit development of a sufficient population of 
band-tailed pigeons so that more shooting might be allowed. 



THE BAND-TAILED PIGEON 37 

Most important of all management practices must be the care- 
ful guarding of the species through continued stringent protec- 
tion, with short shooting seasons and small bag limits. Crop dep- 
redations must be controlled or prevented with the minimum 
destruction of the pigeons. The species cannot withstand liber- 
alized shooting in more than a few districts, and because of its 
restricted range, excessive liberalization at any point is undesir- 
able. 

Through cooperation of the local agencies concerned, most of- 
ten the United States Forest Service and the State Game Depart- 
ments, shooting should not be permitted at all in places where 
large concentrations of the birds are exposed to highly destruc- 
tive hunting, examples of which have been mentioned (p. 31). 

Normally, the bandtail is a native of the primitive forest and 
rough, wild country, usually at fairly high elevations. Manage- 
ment practices that in any manner assist in maintaining primitive 
areas in this country aid the bandtail ; among these are fire pro- 
tection, erosion control, restriction of human intrusion, and re- 
forestation. 

Few birds are controlled so completely by available food sup- 
plies as is the bandtail. Hence, management of food supplies might 
be expected to form part of any management program. Full 
study of the food habits of the bird, however, fails to elicit opti- 
mism as to the possibilities in that direction. 

If any area is to attract and hold a winter population of band- 
tails, it must afford a bountiful supply of acorns or piiion nuts. 
Although the pigeons feed upon wild fruits and berries, in part, 
throughout the winter, mast is necessary as winter food. Hence 
any program looking toward increasing the winter population of 
pigeons must include increases in the mast supply. 

During summer the food of the bandtail includes a wide range 
of wild and cultivated fruits and in some districts, acorns and 
grains. The use of cultivated fruits and grains may be largely 
eliminated from consideration as a practice intended to increase 
pigeon populations, for, except in local areas, an increase in plant- 
ing of these would entail reduction in the forest or would 
increase the depredation problem. 

In brief, it seems that only three practices are of real value in 
the management of the band-tailed pigeon : conservation of our 
present population ; preservation of primitive and marginal wil- 
dernesses and woodland to offer habitat ; and forestry, including 
fire protection and reforestation of already denuded lands, using 
wherever practicable species of plants that are of known value as 
bandtail foods. There are strong possibilities in the planting of 
wild fruits, such as mulberry, in high mountain park areas where 
these trees might thrive and where food is now scarce. 

AGRICULTURAL RELATIONSHIPS 

DEPREDATIONS 

Depredations on agricultural crops by band-tailed pigeons al- 
though sporadic may be serious. They are generally local and vary 



38 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

greatly from season to season. They sometimes involve numbers 
of birds so large as to be amazing, and again may concern only 
a few pairs. Their occurrence is so erratic as to be impossible to 
forecast, although in some areas slight to moderate damage may 
be done annually. The supply of natural wild foods is a determin- 
ing factor in the occurrence of crop damage because of its effects 
on both pigeon concentrations and routes of migration. With nat- 
ural foods plentiful in the mountains and wilderness there is less 
necessity for the pigeons to feed extensively on cultivated crops. 

Many of the complaints against band-tailed pigeons come from 
ranchers whose production is relatively small, or they may con- 
cern only the loss of garden or orchard crops destined for home 
consumption. Noticeable attack on the products of large-scale 
farming operations is infrequent. Depredations on small acre- 
ages are more quickly noticed, have more vital importance, and 
concern a far greater number of individuals. Very frequently 
even a moderate loss in a cash crop like cherries may be keenly 
felt by the farmer concerned, and if the loss is heavy, it may mean 
financial disaster even though only a few trees are damaged. Even 
so, far too great a percentage of the complaints against the pigeons, 
as against other game birds, upon close analysis, are found to be 
unjustified. In many instances the desire to shoot pigeons for 
food will be found to be the underlying motive for the complaint. 

In the literature are a number of records of band-tailed pigeons 
feeding on agricultural crops or their residues but only a few can 
be mentioned here. Fisher (1893) reported bandtails foraging 
in barley stubble in 1891, and Oilman (1903) noted large flocks 
of them in barley stubble in Riverside County, Calif., in March 
1901. Grinnell (1913) mentions good-sized flocks on newly sown 
barley fields near Palo Alto in January 1901 and near Santa Mon- 
ica in February and March of that year, and Barnes (1916) dis- 
cusses alleged damage to grain. Anonymous news notes in 1924 
and 1930 reported damage to cherries and grapes. Burtch (1930) 
discussed damage to grapes, and McAtee (1932) abstracted com- 
plaints against the species. 

Taverner (1926) writes that "they are especially partial to 
peas and are said to pull up the sprouting seeds ... As they are 
large birds, each one intent on filling a capacious crop, their pow- 
er for damage is not small. In the autumn they alight upon the 
stooked grain and may take a considerable toll of it." 

Grinnell and Storer (1924) describe bandtails feeding near the 
edge of a newly planted grainfield on the floor of Yosemite Val- 
ley in April 1916. Grinnell, Bryant, and Storer (1918) cite a 
single bird taken near Crescent City, Calif., on May 15, 1916, 
whose crop contained 509 kernels of barley, 23 of oats, 6 of corn, 
and fragments of acorn. Jewett reported depredations on newly 
planted fields of oats in Tillamook County, Oreg. Grinnell, Dixon, 
and Linsdale (1930) record a few bandtails coming in pairs or in 
small groups to cherry trees near Manton, Tehama County, Calif. 
(June 4, 1936). Kobbe (1900) said that they fed in wheatfields in 
Pacific County, Wash. 



THE BAND-TAILED PIGEON 39 

Munro (1924) reported on an investig-ation of band-tailed pi- 
geon damage in British Columbia as follows : 

On June 9, 1923, I had occasion to investigate a report that band-tailed 
pigeons were causing damage to sprouted wheat on a small bush farm in 
the Sooke district. The farm in question was found to include a portion of 
a large beaver meadow — one of the few open areas in this heavily wooded 
region — the balance comprising rough timbered hillside, and a wooded ra- 
vine through which flows a small stream. About eight acres of the meadow 
had been seeded to wheat and oats by hand and as always is the case with 
this method of sowing, a large percentage of the seed was on the surface. 
This exposed seed had germinated. 

Pigeons commenced feeding on the wheat field shortly after my arrival, 
so, in order to study them at close quarters — for they are invariably wild 
when in the open — I made a careful stalk through the wooded ravine and 
reached, unobserved, a suitable hiding place at the edge of the field. From 
this position it was seen that 53 pigeons were feeding. Usually in flocks 
of this size small detachments from the rear keep flying over the main flock 
to alight in front of the foremost birds, but on this particular day the birds 
kept their formation — ■ an undulating blue ribbon — and slowly moved 
across the field in my direction until a scant 60 yards distant, when they 
suddenly arose, circled several times, and then dropped on another part of 
the field — there to spread out immediately and commen-ce feeding as be- 
fore. From the several dead trees amongst the green timber behind me 
came other birds, singly as a rule, and joined the feeding band. Close ob- 
servation with binoculars showed that only surface seed was being taken, 
the young plants from buried seed were not pulled up. 

Mrs. Florence Merriam Bailey (1928) writes: 

On the west side of Moreno Valley, on July 4, 1919, Mr. S. E. Piper dis- 
covered that two or three thousand Pigeons had congregated along the bor- 
ders of a deep cove. He was attracted to the place by heavy shooting on the 
part of the ranchmen — mainly foreigners who said that the birds destroyed 
their young grain, especially barley. On examining several areas from which 
the birds rose, Mr. Piper could find no indication that they were either dig- 
ging or pulling the young grain: and barley found in the gizzard of one was 
old stained grain evidently gathered from the surface or about old stack 
or shock stands. It was evident tliat the hunters were seeking justification 
for sliooting them, though several averred that they found them unfit to 
eat. 

The following accounts of band-tailed pigeon depredations were 
reported by ranchers, State game wardens, United States game 
management agents and other representatives of the Fish and 
Wildlife Service (formerly the Bureau of Biological Survey), and 
other persons. These reports include in some instances not only 
the original complaint but also a statement of the results of in- 
vestigations by officials. A few of them have appeared in print, 
but most of them have never before been published. 

CHERRIES, GR.4PES, AND PRUNES 

On August 20, 1920, E. R. Kalmbach, of the Bureau of Biologi- 
cal Survey (now the Fish and Wildlife Service), recorded that 
ranchers near Lookingglass, Oreg., reported severe damage to cher- 
ries by band-tailed pigeons. 

Forest Supervisor Blair, writing from Glenwood Springs, Colo., 
May 29, 1920, transmitted reports of pigeon damage to cherries 
on several ranches in that area. One rancher w^ho complained of 
very severe losses said that scarecrows were a failure and that 



40 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

men without guns could not keep the birds out. Mr. Blair quoted 
several other small ranchers as saying that the tops of the cher- 
ry trees were completely stripped of fruit. 

On June 27, 1921, W. F. Kubichek, of the then Bureau of Bio- 
logical Survey, made a study of the situation as reported in 1920. 
He described the area concerned as a narrow belt between the Col- 
orado River and the mountains beginning 10 miles east of the 
town and widening to the west. After a thorough inspection Ku- 
bichek reported (unpublished manuscript) that bandtails were 
seriously destructive in only a few orchards and in general were 
doing less damage than were other species of birds. In an orchard 
where severe damage had been reported he found a few pigeons 
feeding and was greatly surprised at their fearlessness and the 
extreme difficulty he had in driving them from the trees. His con- 
clusions were that in most instances it was the size and conspicu- 
ousness of the pigeons that alarmed the owners and that actual 
damage by the birds was not great. 

In May 1921, Charles C. Sperry, of the Biological Survey, in- 
vestigated complaints of depredations on fruits by birds in Oregon. 
In his report (unpublished manuscript) several complaints against 
bandtails are included. In Lane County, Oreg., three fruit grow- 
ers claimed measurable loss of cherries through damage by pi- 
geons, and in Douglas County, Oreg., two out of four growers in- 
terviewed made the same report. From Multnomah, Benton, Linn, 
Marion, Polk, Yamhill, Clackamas, Columbia, Washington, Was- 
co, Umatilla, and Jackson Counties no reports of damages were 
received. 

On June 25, 1923, Ray C. Steele, United States Game Warden, 
wrote of a visit to a cherry orchard near Riddle, Oreg. Pigeons 
were abundant, some trees being literally filled with them. They 
were very shy and flew when approached, preventing the collect- 
ing of specimens. The ground was covered with cherries, about 
10 percent of them showing bill marks, the remainder having 
been knocked from the trees as the birds alighted or fed in the 
branches. 

On July 20, 1923, B. R. Britton, United States Game Warden, 
reported damage to cherries in the Teseque Valley on the west- 
ern slopes of the Sangre de Cristo Mountains of New Mexico. On 
July 12 on the Williams ranch, he saw trees that appeared bluish 
because of the many pigeons in them. On some trees the fruit 
was completely devoured, on others the tops were stripped. On 
this 84-acre ranch Britton estimated a loss of 20 to 30 percent of 
the cherry crop. Thirty trees near the margin were completely 
stripped. One bird was watched as it picked 20 cherries and 
dropped about one-third of them as it attempted to swallow them. 

On March 7, 1924, a fruit grower of Paradise, Calif., wrote to 
the Biological Survey regarding pigeon depredations : "My cher- 
ry season lasts six or seven weeks . . . Last season I spent forty 
dollars for ammunition and had to hire a guard for four weeks ; at 
that I lost over 1,500 pounds of cherries." Another orchardist of 
Paradise, also in 1924, reported : "For the last three years our 
cherry crop in this section has been destroyed by wild pigeons . . . 



THE BAiND-TAILED PIGEON 41 

I speak for about 20 growers who estimate their losses at from 
one-fifth to three-fourths of the crop." In a letter of June 24, 1925, 
an orchardist of Fortuna, Calif., said that pigeons had taken be- 
tween 1 and IVi- tons of cherries that year. 

In a letter of June 19, 1925, George Tonkin, United States Game 
Warden, reported several complaints against pigeons. The fol- 
lowing comments are abstracted from his letter: 

In the Paradise district, 80 miles almost due north of Sacramento, there 
are many pigeons. There is said to be a roost and breeding ground in the 
vicinity of this district and the pigeons are there nearly the entire year. I 
found only three farmers in this district who complained about the birds 
to any extent. . . . These ranchers have frightened the birds away even 
though they had, up to the time of my visit, been unable to get any old- 
fashioned black powder. One of them told me that a bullet from his 30-3 
rifle fired across the orchards had a very good effect. 

At another nearby ranch Tonkin found pigeons feeding in cher- 
ry trees close to the ranch house, and it was apparent no effort had 
been made to frighten them away. Five shots from his revolver 
drove fully 75 birds from the orchard. Upon investigating the 
complaint of a rancher near Inwood, Calif., in June 1925, Tonkin 
found the small cherry orchard in a wooded section completely 
surrounded by heavy forests. Near Mount Shasta a tenant re- 
ported pigeons attacking his strawberries. Here Tonkin saw many 
pigeons flying about, but as the ranch chickens had free access to 
the strawberry patch, he did not feel that the pigeons should be 
held responsible for the alleged damage. 

An orchardist of Dunsmuir, Calif., on May 7, 1926, wrote of 
the bandtails: "They are in this section by the thousands, and 
you can stand and shoot and they will fly away a few feet and 
circle right back. We have lost our entire crop of cherries but 
about 60 pounds out of 1,000 this year." 

On June 21, 1928, United States Game Warden Tonkin reported 
his investigations of several pigeon complaints. Near Garberville, 
Mendocino County, Calif., he inspected a small orchard of fine 
cherries completely surrounded by heavy redwood forests. The 
rancher was shooting at, and killing a few of the pigeons to pro- 
tect the crop. Near Fortuna, Scotia, and Carlotta, in Humboldt 
County, Tonkin visited other similar orchards. In one of them the 
rancher or his sons maintained constant patrol with a .410 gage 
shotgun. Pigeons were numerous ; wild berry patches in the tim- 
ber attracted large numbers, which at times invaded the cherry 
orchards. 

Recommending that an open season on band-tailed pigeons be 
permitted in California, Tonkin said : "It seems reasonable to be- 
lieve that the wild band-tailed pigeons have been (and are now) 
a serious menace to the cherry and grain crops in Humboldt Coun- 
ty. Under present conditions I do not believe that the killing of 
wild pigeons in the cherry orchards can be stopped so long as the 
farmers feel that it is impossible to drive them out by other 
means." 

A fruit grower of Gilroy, Calif., wrote on August 27, 1930 : 
"This spring they (the bandtails) came in here by the hundreds 



42 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

and fed on green primes, stripping some trees before we realized 
what they were doing. The State game warden estimated that in 
6 miles along the foothills they did damage amounting to from 
$2,000 to $2,500 to the prune orchards." 

The superintendent of a ranch near Arvin, Kern County, Calif., 
reported on March 18, 1930, that wild pigeons were damaging the 
extensive vineyards on the ranch. The local State game warden, 
county agricultural agents, and Biological Survey officials were 
notified. In 1929, a heavy tonnage of grapes had not been har- 
vested. These had dried to raisins on the vines, and in March were 
on the ground as the result both of natural fall and of the seasonal 
pruning of the vines. On March 21, 1930, deciduous fruit trees 
on the property were in full blossom and the grapevines had al- 
ready put out tender shoots that were from 2 to 4 inches long; 
these shoots carry both the leaf and the blossom for the current 
season. The pigeons frequenting the vineyards were so numerous 
as to arouse great interest. Capt. E. P. Brownlow and Lester Ar- 
nold, Warden, of the California Division of Fish and Game, made 
a detailed study of the flight and estimated that 200,000 pigeons 
were involved. Investigation disclosed that the birds roosted on 
Bear Mountain, some 6 miles to the eastward, and that the acorns 
and other wild foods in the foothill area had been almost entirely 
cleaned up. In the vineyard the enormous flocks of bandtails would 
alight on the trellised grapevines, then drop to the ground to 
feed on the waste raisins ; in doing this they broke ofi" the ten- 
der new growth carrying the current season's crop. In the near- 
by deciduous fruit trees they alighted at times in such numbers 
as to break branches and to knock off the blossoms. The defense 
measures undertaken are discussed under Methods of Crop Pro- 
tection (p. 46). The situation was described by Burtch (1930). 
The depredations of the pigeons continued from March 18 to and 
including April 2. Officials of the ranch estimated the loss of 
Malaga grapes alone at 500 pounds a day, and the loss of peaches 
and plums at 300 pounds a day. They estimated a total loss of 
$5,000 for the grapes and $1,000 for peaches and plums. In addi- 
tion, it was said the owners of the ranch and the State Division 
of Fish and Game together expended about $2,000 in attempting 
to drive the pigeons from the vineyards. 

County Game Warden Fredericksen, of Gilroy, Calif., reported 
that during May 1934 five prune orchards near Gilroy were dam- 
aged. In one of these the pigeons flew across nearly 2 miles of 
almost solid prune orchard and virtually destroyed the crop in a 
10-acre block of sugar prunes. 

For many years pigeons have attacked cherry crops in the vi- 
cinity of Mountain Park, Otero County, N. Mex., and during June 
and July 1939 J. S. Ligon, of the Fish and Wildlife Service, inves- 
tigated conditions there. He reported that the pigeons were not 
so abundant as during some other years, but that damage was even 
heavier than usual owing apparently to a food scarcity in the 
Sacramento Mountains. Because of drought, less grain had been 
produced in the lower altitudes, and the pigeons concentrated on 



THE BAND-TAILED PIGEON 43 

the cherries, which, in that locality, ripen before most of the nat- 
ural wild foods. 

At intervals late in June and early July of 1940 and 1941, J. C. 
Knox, United States Game Management Agent, of Albuquerque, 
inspected the Mountain Park cherry orchards. He reported dam- 
age to about 20 orchards, totaling about 2,500 trees, scattered 
about in narrow mountain canyons. During his 1940 investiga- 
tions he estimated that he did not see more than 250 pigeons, but 
"when 50 to 100 of them alight in one tree at a time, they either 
eat or bruise all the cherries in a very short time, and some of 
these cherries sell for 25 to 30 cents a pound." In 1941 there was 
a great increase in the number of pigeons attacking the cherry 
orchards. Knox estimated that there were 1,000 bandtails. 

GRAIN CROPS 

In August 1921, Webb Toms, Assistant Warden, of the Califor- 
nia Division of Fish and Game, reported depredations by band- 
tailed pigeons on wheat in the San Luis Rey Valley of San Diego 
County. One rancher claimed that wheat enough to fill 20 sacks 
had been taken. Damage was inflicted only while the wheat was 
in the shock, and pigeons covered the shocks so densely as to break 
ofi" many of the heads which dropped to the ground and were wast- 
ed. In contrast, 16 years later (1937), E. H. Glidden, State Game 
Warden, of San Diego, Calif., said that "depredations by wild 
pigeons are unknown here." 

On July 22, 1923, B. R. Britton, United States Game Warden, 
investigated a complaint of pigeon damage near the top of the 
White Mountains of New Mexico. Thirty acres of winter wheat 
were ready to harvest. The surrounding country was in a wild 
state, covered with scrub oak and scattered pines ; the elevation 
was about 8,000 feet. Britton estimated a 5-percent loss of the 
grain, mostly in small patches near the edges of the field close to 
large pine trees. In these places the stalks were beaten down, 
the heads broken away, and the grain stripped off. The owner 
later said that as soon as the grain was shocked the bandtails came 
in even greater numbers and stripped the shocks. 

On September 26, 1924, an agriculturist of Sumas, Wash., re- 
ported that he had 7 acres of fall wheat planted, that the pigeons 
had been eating there since planting time, and that they were even 
pulling up sprouted plants. 

During April 1932, the writer recorded two instances in which 
migrating flocks of bandtails crossing the Sacramento Valley, Cal- 
if., alighted in a newly seeded rice field and ate the broadcast seed 
until driven out. In January 1934, a large flight of pigeons invaded 
the Pacheco Pass district of California. Ranches there are more 
or less marginal in nature, and complaints of severe damage to 
grain were received. Permits for killing the pigeons committing 
the depredations were issued, and local wardens reported 3,300 
pigeons killed, a far greater number than was justified by the 
damage. 

Frank Poley, United States Game Management Agent, on Sep- 



44 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

tember 26, 1941, investigated complaints of depredations on wheat 
in the Sanborn Park area near Norwood, Colo. On one ranch he 
estimated a loss of about one-third of the shocked wheat. Similar 
losses were noted on several adjacent ranches. 

From April 28 to May 15, 1925, Ira N. Gabrielson investigated 
alleged depredations by band-tailed pigeons in two Washington 
Counties, San Juan Island and the mainland areas in Whatcom 
County. Both areas were extensive pea-growing districts, total- 
ing from three to four thousand acres. Of the Sumas district of 
Whatcom County, Gabrielson wrote : "Whatever the condition may 
have been at other seasons, there was no damage at all in this dis- 
trict in the spring of 1925. There were no pigeons in the district 
nor had there been any up to the date of my departure (May 11). 
Testimony agreed that when the pigeons came in, they ate all the 
peas left on top of the ground. The claim was made that these peas 
on top of the ground M'ould grow if left there. Most of the farmers 
refuted this, saying that the peas on top of the ground, while they 
might sprout, would never make good plants." During his study 
Gabrielson observed a field in which there were many peas on top 
of the ground ; because of favorable weather, these had sprouted, 
but on the day of his inspection the sun shone brightly and the 
ends of the sprouts were blackening and shriveling. 

Residents of the Sumas area told Gabrielson that pigeons had 
been exceedingly abundant in the spring of 1924, and that this 
had led to numerous complaints. 

On the islands Gabrielson found a more difficult situation. Most 
of the peas were grown on San Juan Island, and only a few on Or- 
cas and Lopez Islands. The industry there had developed rapidly 
after its start in 1922, and in 1925, 1,000 acres of peas were being 
grown for the cannery. The soil was largely moisture-retaining, 
stiff clay, hence hard to work and cloddy, and the drills left more 
peas exposed than on lighter soils. This condition led to the nu- 
merous complaints against pigeons. 

It was the practice of most of the farmers to run a clod-masher 
over the fields after seeding was completed. The only loss attribut- 
able to the pigeons was the eating of peas from among the clods 
in the interval between seeding and clod-mashing. Farmers al- 
most unanimously agreed that peas left on top after the masher 
passed were valueless. 

Farmers on those islands who grew grain or peas for seed told 
Gabrielson that the bandtails did considerable damage at harvest 
time through alighting on the shocks and eating all the seed they 
could reach. 

An orchardist of Sequim, Wash., on April 17, 1935, said that in 
1933 he had sufi'ered a total loss of five acres of peas and that he 
had a thin stand on the remainder of his planting ; also that many 
bandtails were present at the time of his reporting and severe 
damage would probably occur. When a Biological Survey investi- 
gator inspected the property the pigeons had already departed, 
but there was evidence that large patches of peas had been cleaned 
up. The orchardist told the investigator that he grew peas for 
seed and that the greatest damage was inflicted on the ripening 



THE BAND-TAILED PIGEON 45 

seed crop in August when the birds alighted on shocks, bent the 
curing plants, and shelled them onto the ground. 

S. J. Handron, State Game Protector, of Hoquiam, Wash., re- 
ported on May 24, 1937, that between two and three thousand 
pigeons were feeding on pea plantings near Elma, Wash., and on 
September 21, 1937, Fred Rice, State Game Protector, of Port 
Angeles, Wash., told the writer that for the past 3 years the spring 
concentration of bandtails had been exceptionally heavy and that 
severe damage to oats and peas had been sustained. 

WALNUTS 

In February and March 1932, a large concentration of pigeons 
near Exeter, Calif., caused unusual damage. The birds gathered 
at dusk in an English walnut grove in such numbers as to break 
branches from the trees. By counting pigeons in several of the 
trees Capt. 0. P. Brownlow, in charge of the State Game Patrol 
in that area, estimated that at one time there were in excess of 
25,000 bandtails in that grove. 

METHODS OF CROP PROTECTION 

Methods of effectively and economically protecting crops from 
damage by band-tailed pigeons have been more or less extensively 
studied by certain farmers and by conservation officials. In the re- 
ports of field men of the Fish and Wildlife Service are numerous 
notes on this phase of the pigeon problem. 

Gabrielson in his 1925 report wrote of a rancher near Belling- 
ham, Wash.: 

He stated that the issuing of permits would not help any as the average 
farmer did not have the time to stand around and shoot these birds, and 
that they could not kill enough of them to make any difference. His belief 
was that some sort of frightening device would be of much greater value. 
He found that firing a gun from his front porch frightened the pigeons 
for an hour or two. He started in with a 12-gage shotgun . . . [he] found 
the 22 rifle just as effective. 

In 1926, George Tonkin, United States Game Warden, reported 
on a method of driving pigeons from cherry orchards : 

On the morning of May 16 I arrived at an orchard at Paradise [Calif.] 
before 5:30 a. m. I had a 10-gage shotgun and some shells loaded with 8 
drams of black powder, known as the yacht cannon load. I demonstrated to 
several ranchers that day that a shot flred from this gun, loaded with black 
powder and without shot, would frighten the pigeons from their orchards 
and from the roosting places in the oak groves for a distance of one-third 
to one-half mile from the place where the gun was fired. But the birds either 
returned in a short time or another flock took their place, probably the 
latter case. At any rate, it is necessary for a rancher to keep a guard in his 
orchard from daylight until dark. On some days the pigeons appear to have 
left the country; perhaps on the following day a flock will visit the orchard 
every few minutes. It is a great hardship for farmers who have a large 
orchard some distance from their residence, as a few minutes' absence from 
the orchard may mean the stripping of cherries from several trees. 

B. R. Britton, United States Game Warden, thus described his 
attempts at protecting grainfields from pigeons in the Teseque 
Valley, near Santa Fe, N. Mex., in 1923 : 

I purchased some black powder shells and returned to the Williams 
farm, arriving about 2:30 p. m. I took the shotgun shells and cut the shot 



46 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

portion of them away, and began patrolling the orchard, firing on the pigeons 
whenever I was close to them, but found this had little effect. Continuous 
patrol by myself and hired men had the effect of keeping the birds moving. 
At daylight on Friday morning I found the birds roosting in the cotton- 
woods adjoining the orchard, and slipping under these trees I fired blank 
charges up through them. This was continued all day, and I found that when 
I could get close enough to the trees in which the birds were roosting or 
feeding to get the smoke and report of the gun practically among them 
that it had a terrifying effect. I found that the pigeons would avoid a tree 
in which they had been disturbed in this manner. 

Constant patrol was maintained from 5 a. m. to 7 p. m.. someone being 
in the orchard all the time. I mysel'i gave particular attention to the resting 
places of the birds along the edge of the orchard. Part of the time I used 
a revolver and part of the time a 30-4 rifle, simply discharging them into 
the air. I found that the report of the rifle or revolver had a more terrifying 
effect on the birds owing possibly to the sharper report. I also tried fluting 
or cutting the revolver bullets, which resulted in a screeching sound as they 
passed through the air. By Monday the birds had practically deserted the 
Cottonwood trees adjacent to the orchard as resting places (except in the 
very early morning) and rested in the scrub cedar and pifion trees on the 
hillsides. Prom there they were routed by firing the rifle into the dry 
hillside in the vicinity of the roosting trees; this caused a cloud of dust 
which kept the birds moving. The birds then moved into a canyon behind 
a hill, and I followed them there and finally succeeded in driving them out 
of there. By Wednesday morning there apparently remained only about 30 
or 40 birds and these were very wild, flying high, and if they dropped into 
the orchard, would rise on the approach of any one. On Wednesday after- 
noon there was a violent thunder storm accompanied by a high wind, and 
about 5 p.m. a flock of about 2 00 pigeons appeared and were persistent in 
settling in the orchard, acting in the same manner as those found in the 
orchard at first, having to be driven 'from the trees in which they settled 
to feed. I might state that the orchard as viewed from the hillsides ap- 
peared like the madrone woods in California, which were favorite feeding 
places for the wild pigeons; and in my opinion it was this bright coloring 
that attracted passing birds. Thursday morning there appeared only the 
remnants of the flocks that were at first jtresent, the flock of birds that had 
appeared on Wednesday evening apparently having passed on. 

In July 1937, the writer was collecting specimens of bandtails 
in the mountains near Santa Cruz, Calif. One mountain ranch 
visited contained about 20 fine trees of sweet cherries, all loaded 
with a heavy crop of good fruit. Among the birds, of six species, 
noted feeding on the cherries were approximately 30 to 50 pigeons. 
In one instance about a half dozen of these birds alighted in the top 
of a tree on which two fruit pickers on short ladders were work- 
ing. They fed until one of the pickers, after drumming on his pail 
and yelling, descended to the ground and threw clods at them. 
Shooting incident to collecting specimens continued intermittently 
for an hour, and the pigeons at no time retreated farther than 
the tops of some tall sequoias and other conifers adjacent to the 
cleared farm land. 

James A. Blair, Forest Supervisor, at Glenwood Springs, Colo., 
writing on May 29, 1920, quoted a rancher in that area as having 
tested the effectiveness of scarecrows in preventing damage by 
pigeons. The rancher said that his cherry pickers tried to frighten 
away the pigeons, but that no method except the use of a shotgun 
would cause them to do more than circle to another part of the 
orchard. 

In attempts to frighten away the enormous flocks of pigeons that 
were damaging the vineyard on the ranch in Kern County, Calif., 



THE BAND-TAILED PIGEON 47 

in 1930, previously mentioned (p. 42), various methods were 
tested. Among these was that of trying to feed the pigeons in the 
mountains near their roosting grounds. There 1,000 pounds of 
barley and 500 pounds of raisins were scattered. The pigeons are 
said to have taken this food, but they were so numerous as to make 
the method ineffective. An airplane was then used in an attempt 
to drive the birds from the vineyard, but without success. The 
California Division of Fish and Game and the owners of the ranch 
cooperated in hiring 16 men to patrol the vineyards and in furnish- 
ing the necessary ammunition. After a week's trial, the flocks 
appeared to be little diminished in numbers and the damage was 
said to be increasing, owing to the increasing length of the grape 
shoots and the budding out of later varieties of grapes. 

On March 30, fifty sportsmen were invited to assist in the drive. 
A number of pigeons were killed and although the rest were kept 
stirred up, they were not discouraged but kept alighting in the 
vineyard. As a result of publicity some 300 sportsmen assembled 
on March 31 and April 1 ; yet the birds continued to come in by the 
thousands. On April 2, about 500 men appeared at the vineyard 
with about 200 rounds of ammunition each, and it was said that "a 
bombardment such as has not been heard since the First World 
War took place from daylight to about noon." Not a pigeon was 
given a chance to alight. 

On April 3, hundreds of hunters appeared but no pigeons came. 
A survey of the vineyard by officials revealed fewer than a dozen 
birds during the entire forenoon, and inspection of the roost indi- 
cated that they had deserted the area. It was estimated that be- 
tween 5,000 and 7,000 pigeons were killed. 

These reports indicate the problems that develop in herding off 
pigeons by gunfire when they range in numbers from a few birds 
to almost a quarter million. As intimated by Tonkin and Britton, 
and fully substantiated by the experience of countless farmers 
and game officials, timing the protective effort is most important. 
Control work should be started at the first evidence of crop loss, 
not after the pigeons have fed long enough to become accustomed 
to the area. It is far easier to discourage the birds at the begin- 
ning of an attack than after the flight has developed to large 
proportions. 

Herding off pigeons with gunfire remains the standby, with 
advantages and disadvantages as herein pointed out. It is costly ; 
it requires a large quantity of black-powder ammunition and con- 
stant patrol by gunners whose numbers depend on the size of the 
area to be patrolled. Rifle fire, although effective, is too danger- 
ous in most localities. 

As already reported, scarecrows appear to have no effect on 
the pigeons. Airplane herding was of no avail. Feeding the pi- 
geons in their mountain haunts proved possible, but was so costly 
that it was impracticable. 

John C. Knox, United States Game Management Agent, in June 
and July of 1940 and 1941, made extensive and fairly successful 
tests of frightening devices in the cherry orchards of Mountain 
Park, N. Mex. He described the results obtained with the use of 

695766°^7— 4 



48 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

the automatic acetylene flash gim2 known to be of vaKie in pre- 
venting- depredations by various species of birds. 

One of these "guns" was operated near High Rolls in an or- 
chard of 30 cherry trees. Before its installation 40 pigeons had 
fed in these trees morning and evening for more than 10 days. 
When the exploder was timed so that explosions occurred at 10- 
minute intervals, the pigeons were kept away from the trees, and 
when it was regulated so that explosions occurred every 3 minutes 
the jays, robins, orioles, and grosbeaks were controlled to an esti- 
mated 80 percent. At the end of the 1941 test, Knox concluded 
that the use of 10 or 12 acetylene flash guns together with a 
limited amount of concurrent shotgun shooting would effectively 
solve the problem in that area. 

An adaptation of the "flagging" system commonly used against 
horned larks in truck crops in California^ was tested. In 1940, in 
an orchard of 150 trees a cord was stretched over the tops of 
the trees in each row. Between and near each tree streamers made 
of white wrapping paper were tied to the cord so that they waved 
in the rather constant breeze and made considerable noise. For 
7 days after the installation of this device, no pigeons fed in the 
orchard. Then a severe rain and wind storm destroyed the papers. 
In 1941 streamers made from a durable kraft crepe paper with 
an asphalt inner binding were tested. This paper proved very 
successful during dry weather in frightening the pigeons. It with- 
stood rain and wind, but when wet it made no noise. Apparently 
it was the crackling and popping of the streamers rather than 
their motion that frightened the pigeons, for when the crepe paper 
streamers ceased to make noise, they lost their effectiveness and 
the pigeons returned. 

A spotlight beacon useful in frightening away night-feeding 
ducks^was tested against the day-feeding pigeons, but proved in- 
effective, as did also strips of paper, cloth, and bright tin hanging 
in the trees. 

Knox demonstrated that tree covers made of tobacco cloth were 
both effective and economical. Cover for trees of various sizes were 
made at costs ranging from 50 cents to $3 each. Because pigeons 
prefer the higher branches, the covers were so constructed as to 
cover only the upper parts of the tree, but where other birds are 
plentiful extension of the covers to protect the entire tree might 



" The acetylene flash gun is a commercially manufactured device consisting of a carbide, 
water, combustion, and flash chambers. A controlled flow of water entering the carbide 
chamber forms acetylene gas. Wlien pressure forces release of the gas. it is ignited by 
a pilot light and explodes with a report similar to that of a shotgun, and with a blinding 
flash of light. By means of the valve controlling the water flow, the apparatus can be 
set to explode at almost any desired frequency. Information on where this and other bird- 
frightening devices can be obtained can be furnished by the Fish and Wildlife Service, 
Department of the Interior, Chicago 54, 111. 

3 Protecting crops from damage by horned larks in California. By Johnson A. Neff, 
Biologist. U. S. Dept. Agr., Bureau of Biological Survey Wildlife Research and Manage- 
ment Leaflet BS-61, lO pp., illus., September 1936. [Processed.] 

* Protecting field crops from waterfowl damage by means of reflectors and revolving 
beacons. By F. M. Uhler, Biologist, and Stephen Creech. Game Management Agent. U. S. 
Dept. Int., Bureau of Biological Survey Wildlife Leaflet BS-149, 6 pp., illus. November 
1939. [Processed.] 



THE BAND-TAILED PIGEON 49 

be well worth while. As the covers would be in use only about 30 
days of each year, if carefully handled they should last for several 
seasons. The writer on many occasions has observed similar covers 
used to protect vines and fruit trees, the most elaborate being a 
gas-pipe-chicken-wire structure covering large sweet cherry trees. 

Near Hermosa, Colo., in 1942, Game Management Agent Foley 
found the acetylene flash gun very ineffective unless it was moved 
about the orchard several times daily and was accompanied by 
liberal gunfire. Pigeons were seen feeding in the tree adjoining 
the flash gun after it had been operating for almost an hour. In 
the same area, in 1943, white Very signal flares were tested and 
proved most efi'ective in frightening pigeons from the orchard. 
Their use was restricted by the great fire hazard in all places 
where the ground cover was inflammable. Pyrotechnic 3-inch flash 
bombs such as are widely used for frightening ducks were also 
tested in cherry orchards and proved to be very satisfactory, hav- 
ing greater value than gunfire. 

Studying the depredations on cherries at Cloudcroft, N. Mex., 
Frank C. Knox, Game Agent, found that a small number of wild 
mulberry trees grew in the mountain canyons near the cherry 
orchards. The fruit on these trees normally ripened about 2 weeks 
after the earliest cherries were ripe, but before the late cherries 
ripened. He first noticed that the pigeons were more easily driven 
from the cherry orchards after the mulberries began to ripen. 
Then, in 1943, weather conditions caused the early cherries and 
the mulberries to ripen at the same time; a large part of the pi- 
geons remained in the canyons feeding on mulberries and did not 
come to the cherry orchards until the mulberry crop was exhausted, 
at a time when most of the cherry crop had been harvested. Simi- 
lar conditions existed near Paonia, Colo., in 1945. 

Knox's observations led him to believe that the planting of early 
ripening mulberries in the canyons of the Cloudcroft area would 
go far toward reducing the cherry damage there. Such a method 
of prevention might well be practicable in districts in which the 
pigeon population is moderate to small, but it is the writer's opin- 
ion that it would not prove eff'ective in areas in which the pigeon 
population is very large, for severe crop depredations have been 
noted in many instances in areas where pigeons were abundant 
and where the native wild food supply was far greater than was 
necessary for the birds. The effectiveness of the method prob- 
ably depends largely on local conditions. 

Studies of methods of crop protection by means of deterrent 
and frightening devices have not kept pace with field needs, and 
further extensive experimentation is desirable. Man's ingenuity in 
devising noise-makers and frightening devices has scarcely been 
tapped, and much can doubtless be accomplished along this line. 

Each complaint of pigeon depredations should be carefully in- 
vestigated by either State or Federal officials, and when damage 
is found to have occurred, the victims should be given every prac- 
ticable assistance. 

Because of the fondness of band-tailed pigeons for agricultural 
crops, good management decrees that the pigeon population should 



50 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

not be permitted to increase unreasonably; therefore carefully 
planned open seasons should be continued so that shooting will act 
as a population control, particularly in areas in which pigeons are 
abundant and depredations severe. Open shooting seasons, how- 
ever, are not the remedy for specific instances of depredation for 
because of the habits of these birds control of depredations by 
hunting could be realized only through general and widespread 
population reduction, which is undesirable. 

The issuance of permits to kill pigeons that are damaging crops 
has never been completely successful. Earlier authorizations al- 
lowed the owner or lessee to use the birds so killed for food, and 
as a result a great demand for permits developed, the desire to 
hunt being the primary motive. Hence permits no longer include 
the privilege of utilizing the birds killed as food, and significantly 
there has been a great decrease in the number of complaints. Dur- 
ing recent seasons it has been possible in some areas through agree- 
ment between Federal and State officials to require the permit- 
tee to preserve the pigeons killed for delivery to charitable institu- 
tions or hospitals. This has effected further decrease in the num- 
ber of requests for permits. 

Another defect of the permit system is that shooting usually is 
allowed only on the area on which damage is occurring; but, as 
has been pointed out by Britton in New Mexico, shooting the pi- 
geons at their roosting and perching sites may aid greatly in reduc- 
ing the damage. These roosts and da^'time perching trees may 
be near the crop attacked or may be miles distant. 

Where important crop losses occur, it would often facilitate pro- 
tection if the owner were permitted to shoot the birds on his prop- 
erty while at the same time game wardens or men under their im- 
mediate and close supervision traced the flight of the pigeons and 
continued the frightening process at perching and roosting places 
until a change in flight lines or the habits of the birds brought re- 
lief. This procedure, however, would require a large personnel 
and a great expenditure of funds. 

There can be no question that serious agricultural losses sus- 
tained by persons on account of depredations by band-tailed pi- 
geons must be controlled. On the other hand, conservation sentiment 
demands that there be no excessive or unwarranted destruction of 
the birds. Pigeon depredations affect a large number of persons, 
especially in mountain, valley, or wilderness areas close to abun- 
dant bandtail populations. In many cases there is a distinct loss, 
but in many others the damage is magnified. Regardless of the 
degree of damage incurred or of the justification for control of 
the birds on the individual farm, studies of methods of prevention 
of damage have lagged, and there are few economical, practicable, 
and highly successful methods that can be recommended. Where 
the farmer will get out with his gun at daylight and keep the 
pigeons from feeding in his field until dark usually only one to 
three days are required to break up even the worst of the normal 
pigeon attacks. 



THE BAND-TAILED PIGEON 51 

FOOD HABITS 

GENERAL 

The literature on the band-tailed pigeons contains many refer- 
ences to their feeding habits, mostly based on field observations. 
Huey (1913) describes their feeding on manzanita berries in Cali- 
fornia, Willard (1916) on acorns of Querctis eiyioryi in Arizona, 
Oilman (1903) in grain stubble in California, and Taverner 
(1934) on peas and grain in British Columbia. Bendire (1892) 
quotes Carpenter relative to the feeding of the pigeons on the salm- 
onberry (Rubus nidkanus) along the Columbia River, and Kobbe 
(1900) confirms the report. Belding comments on their feeding 
on acorns and oak buds in California, Ankeny on their eating 
acorns in the Rogue River Valley of Oregon, and Lloyd on their 
consumption of wild grapes in western Texas and of acorns there 
and in Mexico. From Arizona Benson reported to Bendire that 
acorns were taken after July 15, and Poling mentioned mulber- 
ries, although not definitely stating that the fruits were eaten. 

Dawson (1923) said the bandtails fed on elderberries (Sambu- 
cus), cascara {Rhamnus pursJiiana) , cofleeberries (Rhamnus cali- 
fornicus), huckleberries (Vuccinium), salal (Gaultheria) , salm- 
onberries (Ritbiis), madrona berries (Arbutus), and Christmas- 
berries (Photinia), but that acorns were their favorite food. 

Gabrielson and Jewett (1940) summarize the food of the birds 
in Oregon as consisting of acorns, mountain ash berries, berries of 
various species of Rubus, elderberries, and currants ; kinnikinnick, 
dogwood, and other fruits ; grains ; and peas and other legumes. 
They mention that after nesting time, when the birds flock to- 
gether, they feed on the fruits of salal, salmonberry, blackberry, 
and other wild fruits ; Jewett has stated that they are particularly 
fond of the cascara berry (Rhamnus purshiana) ; and Gabrielson 
frequently observed them apparently feeding on the seeds of Lu- 
pinus lyalli. 

Mrs. Bailey (1928) summarizes the food of the species as fol- 
lows : "Mainly nuts, especially acorns, and berries, including wild 
currant, chokecherry, wild cherry, juniper, madrone, manzanita, 
raspberry, blackberry, elderberry, huckleberry, salmonberry, cof- 
feeberry (Rhamnus calif ornica) , Christmasberry, and black goose- 
berry; wild grapes, flowers and leaf buds, sycamore balls, pine 
buds, seeds, and needles, and grain (mostly waste) ; together with 
grasshoppers and other insects." 

Grinnell, Bryant, and Storer (1918), in Game Birds of Califor- 
nia, say that "the amount of food available to the pigeon appears 
to be the main controlling factor in its distribution. This is more 
particularly true in winter, though probably to some extent in sum- 
mer also." They pointed out that the food consisted mainly of 
nuts and berries, and that the plants that bear these are frequent- 
ly alternate-year, or intermittent, crop-bearers. 

Of 22 food records summarized by these authors, 10 give acorns 
as the chief item. Specifically, these included the acorns of Quer- 
cus Kigrifolia, Q. ivislizenii, Q. kelloggii, and Q. cknjsolepis. Ber- 
ries of the madrona (Arbuttcs menzies'ii) were said to be an at- 



52 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

tractive autumn food. Fruits of certain species of manzanita(Ar'c- 
tostaphylos) were reported as taken from the time they were first 
formed until very late in the season when fully ripe. During the 
autumn, fruits of the coffeeberry, elderberry (Sambucus glauca) , 
and chokecherry {Prunus demissa) were mentioned as favored 
foods, and in the winter, berries of the toyon (Photinia cLrbutifo- 
Ha) were taken. 

Buds and blossoms also were taken. Dean (1904) reported the 
bandtails feeding on manzanita buds, and there are many records 
of their eating oak buds and flowers. Evermann (1886) found 
35 of the ball-like flowers of the sycamore in the crop of a single 
pigeon. Pine seeds were recorded as bandtail food by Belding 
(1890) and Grinnell (1905). Cultivated grains, including wheat, 
barley, oats, milo maize, and field corn, have been listed in nu- 
merous instances. Many miscellaneous items are recorded, includ- 
ing wild peas, dogwood berries, hazelnuts, alder seeds, and juni- 
per berries. In New Mexico, H. W. Henshaw (1886) observed the 
pigeons feeding on fruits of the elder {Sambucus racemosa) and 
acorns of the scrub oak {Quercus undulata) . 

J. A. Munro, Chief Federal Migratory Bird Officer, of British 
Columbia, reported in 1941 on the examination of the food of 13 
bandtails collected in British Columbia between 1923 and 1933. 
The crop of a bird collected in June contained approximately 346 
kernels of wheat. Of 12 birds collected in September, the crops 
of 6 contained field peas, and those of the others such items as 
acorns of Quercus garryana, seeds of Conius nuttalli, Sambucus 
glauca, Gaultho ia shallon, and oats. 

Several writers have commented on the great volume of food 
that can be taken by a single pigeon. Grinnell, Bryant, and Storer 
mention an acorn dropped by a flying pigeon that measured nearly 
1 by IV2 inches, and also record a pigeon killed in Del Norte Coun- 
ty, Calif., whose crop contained 534 kernels of barley, oats, and 
corn. 

Van Rossem (1914) reported finding pigeons in a dying condi- 
tion, their crops pierced by acorns that they had swallowed. John 
G. Traub, of the Fish and Wildlife Service, who ranched for a 
time in San Luis Obispo County, Calif., told the writer of killing 
apparently healthy pigeons that upon examination were found to 
have points of acorns protruding through the crop wall ; the feath- 
ers were matted with crop juices, an indication that this condition 
had existed for several days, yet the pigeons had apparently con- 
tinued to feed and fly vigorously and normally. 

E. H. Glidden, State Game Warden, of San Diego, Calif., wrote 
on December 10, 1937, "You will find the chief diet of wild pigeons 
at this time of the year in this district to be acorns ; however, one 
may at times find the young shoots of pines in their crop, especial- 
ly on Palomar and the Mesa Grande Mountain. During the spring 
they feed almost entirely on elderberries." 

W. H. Ransom, United States Game Management Agent, said in 
1937 that madrona and mountain ash berries were supposed to be 
choice winter foods for the small number of pigeons that wintered 
about Seattle, Wash., and that in June the bandtails fed on ripened 



THE BAND-TAILED PIGEON 53 

wild strawberries in great numbers, especially on Sand Island 
near Willapa Harbor. 

Lawrence W. Saylor wrote: "I once kept an injured bandtail 
for two weeks . . . and it w^ould eat as many as 30 madrona berries 
a day ; it also accepted and ate huckleberries ( Vaccinium ovatwni ) , 
coffeeberries, cultivated currants, and strawberries, and on several 
occasions readily took grasshoppers when offered. . . . Apparently 
the tips of Douglas fir branches are eaten (by the wild birds), as 
I have stood under the trees and watched the birds pick at and 
swallow the tender tips of the twigs." 

There is frequent mention of insects as food of these pigeons, 
but it would seem that most of these references trace to a field 
examination made by Vernon Bailey, on July 29, 1903, of a pigeon 
killed by him at Pecos Baldy, N, Mex., of which he said, "The giz- 
zard was full of insects, mainly grasshoppers and the larvae of a 
wasplike insect." Despite this observation, and those of Saylor 
just mentioned, pigeons as a group eat little animal food. 

Several writers have described feeding mannerisms of band- 
tails. Oilman (1903) says that "instead of spreading out they 
kept together alternately walking and flying. Those behind would 
fly a few feet ahead of the advance line, alight, and walk along 
picking up grain until other rear ones would fly ahead and it came 
their turn again." 

Huey (1913) stated that a flock observed feeding on manzanita 
in San Diego County arrived a little after sunrise and left be- 
tween eight and nine o'clock in the morning ; in the afternoon the 
birds returned about four and left again at dusk. Willard (1916) 
described the bandtails feeding on acorns in Arizona : "They would 
walk out on the slender branches till they tipped down, then, hang- 
ing by their feet, would secure an acorn, and drop off to alight on 
a branch lower down." 

In mid-June 1941, pigeons numbering possibly 12 to 20 came 
from the Pinaleno Mountains to a group of mulberry trees on a 
ranch 7 miles south of Pima, Ariz., to feed upon the ripening 
fruits. The distance from the mulberry trees to the oak-pine zone 
high up the side of Mount Graham must be at least 10 miles. Two 
specimens of bandtails were obtained here, giving the first rep- 
resentation of mulberries in stomachs in the entire study although 
this fruit was known to be well accepted. 

In May 1932, the writer saw two migrating bands of pigeons 
flying across the broad Sacramento Valley of California stop to 
feed upon seed rice in two newly planted fields. 

During the course of intermittent field work on band-tailed pi- 
geons extending from 1936 through 1941, the writer has personally 
observed, or has received reports from cooperators who have ob- 
served, the birds feeding on most of the food already mentioned 
in the abstract of the literature on pigeon foods. In some instances 
no specimens of bandtails were obtained, hence the particular item 
does not appear in the tabulations of the result of stomach exami- 
nation ; in other cases the item may be represented in unnaturally 
small ratio owing to lack of adequate collecting in a habitat pro- 
viding that particular food. 



54 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

USE OF MINERAL SALTS 

In 1937, many Puget Sound observers noted that the pigeons 
frequently flew to mud flats at the margin of the Sound to drink 
salt water even though fresh-water streams were available. In 
this connection W. H. Ransom (in a letter of April 15, 1939) 
writes : "Near Cathlamet, Wash,, along the Columbia River there 
is a cliff containing saline deposits of some sort, and the game 
protector stationed there a few years ago told me of seeing scores 
of pigeons gathered there in late summer, while groups of them 
fluttered in the air as close to the vertical clifl" as they could get, 
all the while picking at the salt-bearing earth." 

Frank B. Wire, of the Oregon Game Commission, told the writ- 
er that many years ago he had frequently hunted pigeons at a 
salt spring that attracted the birds in considerable numbers. Ben- 
dire (1892) quotes Anthony relative to a large salt spring south 
of Beaverton, Oreg., where pigeons were always to be found in 
large numbers, Kloppenburg (1922) writing of the bandtails in 
the Plumas National Forest in California says that "they can usu- 
ally be found near mineral springs, especially soda and sulphur 
springs." 

This habit may be analogous to that of the mourning doves that 
are attracted to salt in the Southeastern States to such extent 
that the use of salt is included as a prohibited method of baiting. 
In Arizona the writer has observed western white-winged doves 
picking at salt blocks in cattle feed lots. Salt was used as bait also 
in connection with the shooting of the now extinct passenger pi- 
geon. 

RESULTS OF LABORATORY EXAMINATIONS 

The following report covers the laboratory examination of 691 
band-tailed pigeon crops and/or stomachs. This material admittedly 
is inadequate, some States being represented by only a few speci- 
mens. In California, the pigeons occur at some time of year in 
most of the more than 50 counties, but only 18 of them are repre- 
sented in this study, and 194 out of 267 stomachs were salvaged 
from the bags of hunters during the 15-day December open season ; 
likewise, 113 of the Washington specimens were procured from 
hunters during the 15-day September open season, in 1937. 

For the purpose of this report the crop and stomach, or the crop 
or stomach-, of an individual bird is considered a unit specimen. 
Every efl'ort was made to obtain both crop and stomach, but since 
many specimens were obtained from hunters, it was found diflS- 
cult to obtain both, and in many instances only the crop or the 
stomach was obtained. Regularly cooperating observers, how- 
ever, sent in both crop and stomach. 

The food percentages in this report have been computed by the 
standard volumetric method described by Cottam,-"^ each item be- 



^ Economic ornithology and the correlation of laboratory and field methods. By Oarence 
Cottam, Biologist. U. S. Dept. Agr., Bureau of Biological Survey Wildlife Research and 
Management Leaflet BS-30, 13 pp., illus. January 1936. [Processed.] 



THE BAND-TAILED PIGEON 



55 



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California (18) 

Colorado (3) 

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Texas ( 1 ) 


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56 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 



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THE BAND-TAILED PIGEON 



57 



ing measured after drying. Although representation varies 
greatly, the month is used as the period for the computations. For 
plant names the most recent publication on taxonomic botany for 
each of the States has been used. 

Of the specimens available, 25 had such incomplete data that 
they were useless, although information obtained from them may 
be mentioned ; 27 others were so nearly empty that they were 
eliminated from the tabulations. Hence the percentages of food 
volume and frequency of occurrence are based on the remaining 
639 specimens. 

In the examination of these, 76 separate classes of plant food 
items were listed, including unidentified mosses, vegetable debris, 
unidentifiable vegetable fragments, and rodent scats composed en- 
tirely of vegetable debris. The identified plant-food items repre- 
sent 26 plant families. In addition, 10 occurrences of insect frag- 
ments, comprising only a trace of the whole food, were recorded. 
Hence the species may be considered almost wholly vegetarian. 
Gravel in stomachs is figured in its ratio to the total content, while 
food item percentages are calculated after the grit has been re- 
moved. 

In table 4 is summarized by States the material available for 
each month ; the number in parentheses following the name of the 
State refers to the number of counties represented by the stom- 
achs collected. 

In table 5 are listed the general classes or families of food items 
represented, by months, with the composite percentage by volume 
of all species within the family, based on a total of 639 stomachs 
and crops. An added column gives the percentage of frequency of 
occurrence of the combined species of each family. 

The percentages by volume and frequency of occurrence of the 

Table 6. — Percentage by volume and frequency of occurrence of the food items 
that totaled one percent or more of the annual food of 639 bandtails, based 
OH the examination of their stonutciis and crops 



Food item 



Quercus agri folia, live oak acorns 

Que)-cus sp.. uniflentifled acorns 

Avena sativd. cultivated oats 

Prmiiis, cultivated cherries 

Pisinn sativmn. g'arden peas 

Triliciim aestir)iiii. wlieat 

Pin us edulis, pinon nuts 

Quercus kelloggii, California black oak acorns. 

Quercus sp., blossoms of oak 

Rhamnvs purshiana, cascara fruits 

Arbutus menziesii, madrona berries 

Prunus domesficus, cultivated prunes 

Cornus nuttrdlii, dogwood fruits 

Arctostaphylos, sp., inanzanita seeds and tlowei 
Quercus wislizenii, interior live oak acorns... 

Pinus ponderosa, yellow pine seeds 

Srnnbucus glrnicrt, elderberry fruits 

Quercus garrynmi. Orejjon white oak acorns. . 

Prunus sp., wild cherry fruits 

Miscellaneous vejietable debris 

Qaultheria sfioUon, salal fruits 

Quercus emoryi, Emory oalc acorns 



I'ercentage by 


Percenta 


ge by frequency 


volume 


of 


occurrence 


17.5 




11.1 


13.4 




18.2 


7.4 




5.8 


7.1 




12.4 


4.8 




7.4 


5.(1 




10.3 


4.0 




1.9 


3.9 




1.6 


3.5 




1.3 


3.0 




2.8 


3.0 




6.1 


3.0 




3.0 


2.8 




11.7 


2.6 




3.1 


2.2 




2.0 


2.(1 




.5 


l.S 




6.4 


1.7 




.8 


1.5 




6.0 


1.5 




4.4 


1.0 




3.8 


1.0 




1.7 



58 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

food items that totaled 1 percent or more of the annual food of 
the bandtails, as based on the examination of the 639 stomachs 
and crops, are shown in table 6. 

Pinaceae (6.4 poxent). — Buds of the Sitka spruce (Picea sit- 
chensis) were found in a May-killed Oregon bird, and staminate 
aments of an unidentified conifer in another. Unidentified pine 
seeds were found in one California bird killed in December. Seeds 
of the yellow pine (Pinus ponderosa) were found in two Califor- 
nia birds collected in June and December, respectively. One bird 
obtained in July in the Kaibab National Forest, Ariz., contained 
625 seeds of this pine, measuring 47 cubic centimeters (4 cubic 
centimeters=l teaspoonful) . 

Nuts of the pinon (Pinus edulis) were found in 12 Arizona 
birds collected in September, October, and November. These nuts 
were the only food in 11 of the birds and composed 94 percent of 
the food of the other. Nuts of the single-leaf piiion (Pinus mono- 
phyUa) composed the entire food of 9 pigeons and 90 pertent of 
the food of a tenth collected in California in December. One bird 
had taken 60 of the nuts which displaced 45 cubic centimeters. 

Ciipressaceae (trace). — One pigeon killed in Arizona in Sep- 
tember contained 270 staminate buds of an unidentified juniper 
(Junipe7'us) . 

Gramincac (12.8 percent). — Cultivated grains make up a mod- 
erate portion of the food of the band-tailed pigeon in or near farm- 
ing areas. Wheat occurred in 66 birds collected during 7 months 
of the year in 4 States and averaged 5.0 percent of the annual 
food. Oats were found in 37 birds collected in 3 States during 5 
months and formed 7.4 percent of the annual food. Barley oc- 
curred in 12 birds collected in 2 States during 4 months and 
formed 0.4 percent of the annual food. Field corn was found in 
only 2 September birds from Colorado. 

Unusual quantities of grain were found in a few crops ; one 
contained 725 kernels of wheat, one 200 kernels of barley, and 
another 660 whole kernels of oats. 

Seeds of Poa sp., wild oats (Avenci fatua), darnell (Lolhim 
temidentmu), and needlegrass (Stipa sp.) were each found in 
single stomachs of birds taken in April, May, and June in Oregon 
and California. The family Gramineae contributed 12.8 percent of 
the annual food. 

Liliaceae (0.5 of 1 percent). — Seeds of an unidentified yucca 
were found in one California bird collected in June, and seeds of 
sotol (Dasjilivion wheeleri) (fig. 9) in three August and two Sep- 
tember birds from southeastern Arizona. Combined they made 
up one-half of 1 percent of the annual food. 

AmcuiiUidaceae (0.8 of 1 percent). — Ten birds collected in July 
and August in southern New Mexico contained the anthers of an 
agave (Agave parryi) (fig. 10), aggregating 0.8 of 1 percent of 
the annual food, and one Arizona bird killed in August contained 
125 seeds of a Smilacina, probably stellaria. 

Fagaceae (US. 7 percent). — The acorns and flowers of oaks con- 
stitute the major food of the band-tailed pigeon. Staminate flow- 
ers of oak were found in four January and four March birds from 



THE BAND-TAILED PIGEON 



59 





FiGiUiE 9.— Pi.iieous from the oak canyons of the Southwest 
often feed on the seeds of the sotol {Dasi/Iirion whecleri), 
which grows in the adjacent high desert. Pinal County, 
Ariz. (Photographed by H. L. Croclvett, July 1941.) 

California. Acorns or fragments of acorns were found in 233 
birds collected during 10 months and from every State represented 
except Texas. Oak products totaled 43.7 percent of the annual food. 
Acorns of the California live oaks were most frequently taken ; 
those of the coast live oak (Quorns agrifoUa) occurred in 71 
stomachs and averaged 17.5 percent of the year's food, and those 
of the interior live oak (Q. ivislizenii) were found in 13 birds col- 



60 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 




B87I2M 

FiGTTRE 10. — The flowers of this tree-like agave (Agave parri/i) 
furnish summer food for iiigeons in the Southwest. The 
agaves grow in the higher desert mountains, sometimes very 
close to pigeon-nesting habitat. Head of Mills Canyon, Pinal 
Mountains, Ariz. (Photographed by H. L. and Ruth Crock- 
ett, July 26, 1936.) 

lected during the period November to March. Other species repre- 
sented were the California black oak (Q. kclloggii), blue oak (Q. 
douglasii) , Oregon white oak {Q. garryana) , Emory oak (Q. em- 
oriji), white-leaf oak (Q. hypoleuca) , and the valley oak (Q. lo- 
bata ) . Fragments of acorns not further identified were found in 
116 birds and averaged 13.4 percent of the annual food. The abil- 



THE BAND-TAILED PIGEON 61 

ity of these birds to consume quantities of acorns has been de- 
scribed. The crop of one specimen examined contained 22 acorns 
of the coast Uve oak, displacing 38 cubic centimeters. 

Juglandaccae (trace). — One California bird killed in January 
contained flowers of the walnut (Juglans). 

Mijricaceae (trace). — Parts of the seeds of the wax myrtle (My- 
rica californica) occurred in one California specimen. 

Ulmaceae (trace). — Seeds of an unidentified hackberry (Cel- 
tis) were taken from the stomach of a bird collected at Uvalde, 
Texas, in December. Pigeons collected in the Capitan Mountains 
of New Mexico had eaten fruits of Celt is reticulata. 

Loranthaceue (0.2 of 1 percent). — The pine mistletoe (Arceuth- 
obium) was represented in nine December stomachs from Cali- 
fornia and three July stomachs from New Mexico by fragments 
of both fruiting and vegetative parts. 

Moraceae (0.1 of 1 percent). — Fruits of the mulberry (Morus 
alba) occurred in two June birds from Arizona, and seeds of an 
unidentified mulberry in one July bird from New Mexico. 

Rosaceae (13.6 percent). — This family, producing many famil- 
iar wild fruits and berries, is well represented in the bandtail's 
diet. Seeds of wild blackberries or raspberries were found in 14 
birds; eight were unidentified; and salmonberry (Rubus specto/- 
bilis) occurred in four, and a wild blackberry (Riibus macrope- 
talus) in two birds. Rose hips were found in one stomach. 

Wild cherries, including Primus emarglnata and P. melunocar- 
pa, were found in 44 specimens collected in Washington, Oregon, 
Colorado, and New Mexico. Fruits of Prunus ewarghiata oc- 
curred in 38 Washington specimens taken in July, August, and 
September, and averaged 1.5 percent of the annual food. 

Seeds of the serviceberry (Amelanchier aluifolia) occurred in 
five September birds, and those of the hawthorn (Crataegus doug- 
lasii) in four July birds from Oregon. Seeds of the wild straw- 
berry (Fragaria) were found in two May specimens, and fruits 
of the toyon (Photinia arbutifolia) in four December birds. 

Cultivated prunes were found in 19 birds collected in May in 
the Willamette Valley of Oregon, and averaged 3 percent of the 
annual food. Cultivated cherries occurred in 79 birds collected 
from May to August in Washington, Oregon, California, New 
Mexico, and Colorado, and formed 7.1 percent of the total food. 
Both the sweet cherry (Prunus avium) and sour cherry (P. cer- 
asus) were represented. The Pacific coast fruits were entirely 
sweet cherries, but in Colorado and New Mexico both varieties 
were included in the food of the pigeons. The combined products 
of the Rosaceae family averaged 13.6 percent of the annual food. 

Leguminosae (If.8 percent). — Seeds of clover (Trifolium),\\x- 
pine (Lupinus), and trefoil (Lotus), and leaves of TrifoUum were 
all found in the crop of a single California bird. 

Cultivated peas had been taken by 44 bandtails collected in 
Washington. Those found in seven April and May stomachs had 
been gleaned from freshly seeded fields and averaged slightly more 
than 2 percent of the annual food; the significance of such feed- 
ing has already been discussed (p. 44). During July, August, and 



62 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

September peas are taken mostly from the stubble of harvested 
fields. Occasionally the birds may attack a field that is to be har- 
vested for seed, causing severe losses. For the entire year, culti- 
vated peas, waste or valuable, averaged 4.8 percent of the food. 

Geraniaceae {trcice) . — A few tiny leaves of the alfilaria (Er odi- 
um) had been eaten by one California bird. 

Euphorbiaceae (trace). — Seeds of the turkey mullein (Eremo- 
carpus setigerus) were found in one specim.en. 

Anacardiaceae (0.5 of 1 percent). — Seeds of two species of su- 
mac were found. Those of Rhus emoryii occurred in the stomachs 
of five specimens from New Mexico, and those of R. trilobata in 
one bird from Arizona, making up 0.5 of one percent of the an- 
nual food. 

Rhanmaceae (3.1 percent). — The berries of the cascara (Rham- 
nus purshiana) are a favorite food in Oregon; these fruits were 
taken from 18 Oregon specimens collected from June to Septem- 
ber and averaged 3 percent of the annual food. Seeds of the coffee- 
berry (R. calif ornwa) occurred in one California stomach. Fruits 
and seeds of the lote bush (Condalia lycioides) were identified 
from a single Arizona specimen. Products of this family averaged 
3.1 percent of the annual food. 

Vitaceae (0.7 of 1 percent) . — Fruits and seeds of the wild grape 
(Vitis arizonica) composed the major food of three August speci- 
mens collected in the Huachuca Mountains of Arizona, and formed 
0.7 of one percent of the annual food. 

Malvaceae (trace). — A single seed, determined as Sidalacea sp., 
was found in one Oregon specimen. 

Araliaceae (trace). — A number of green berries of Aralia hu- 
milis were identified from the stomach of a single Arizona bird. 

Umhelliferae (trace). — Seeds of the gambleweed (Sanicula 
menziesH) occurred in one California specimen taken in Decem- 
ber. 

Cornaceae (2.8 percent). — The fruits of the dogwood are a 
highly favored food, being found in 75 stomachs from Washing- 
ton and Oregon, and averaged 2.8 percent of the annual food. 
Those of the mountain dogwood (Cornus nuttallii) are the most 
commonly taken. 

Ericaceae (6.7 percent). — The heath family includes four 
groups, the fruits of which are relished by band-tailed pigeons 
and averaged 6.7 percent of the annual food. 

In the Pacific Northwest the fruits of the salal (Gaultheria shal- 
lon) were taken by 24 birds from Washington and Oregon and 
averaged just 1 percent of the annual food. Fruits of the madrona 
(Arbutus menziesli) were eaten by 39 California birds and com- 
posed 3 percent of the annual food. 

Flowers and fruits of the manzanita (Arctostaq)hylos) occurred 
in 20 specimens taken during every month from March to August 
in Oregon, California, and Arizona, and furnished 2.6 percent of 
the annual food. Fruits of two species of huckleberry (Vaccinium 
ovatuni and V. delicosum) were identified, the first from a single 
Oregon bird, the second from two Washington birds. 

Sokmaceae (trace). — Seeds of Solarium sp., were found in one 



THE BAND-TAILED PIGEON 63 

Colorado specimen. 

Caprifoliaccue {3.2 percent). — Fruits and flowers of the elder- 
berry (Sanibucus) were found in 70 stomachs and averaged 3.2 
percent of the annual food. Of these, the fruits in 41 specimens 
were identified as those of the blue elderberry {Satnhucus glauca) 
and averaged 1.8 percent of the food; fruits in seven other speci- 
mens were identified as red elderberries (Scvmbucus callicarpa) . 

Compo^itae {trace). — One pigeon collected in California in De- 
cember contained seeds of the tarweed {Madia). 

Miscellaneous {1.5 percent). — Bits of moss were taken from 
one specimen. Fragments of oak galls were often found in pigeons 
that had been feeding on acorns. Unidentifiable vegetable frag- 
ments and debris were recorded from 17 specimens and averaged 
1.3 percent of the annual food. One Oregon bird contained 12 
whole rodent scats ; the scats were entirely vegetable in compo- 
sition and were tentatively identified as those of the wood rat 
{Neotoma) . 

Animal Foods. — All the insect material in the stomachs appeared 
to have been accidentally or incidentally taken ; it was present in 
only 10 stomachs or crops. 

One oak twig gall very similar to an acorn in size and appear- 
ance was found to contain larvae of gall flies {Cynipidue). A 
staphylinid larva (rove-beetle) and an adult Anthrenus (skin 
beetle), each in single stomachs, might have been taken inciden- 
tally in feeding on other items. 

One stomach contained 7 honey ants {Prenolepis iTnparis) and 
a quantity of tiny leaflets of clover, and it is reasonable to assume 
that the ants were upon the leaflets when they were taken. One 
fire ant {Solenopsis) occurred in a crop otherwise filled with wild 
berries, and one acrobat ant {Cremastogaster) was also identi- 
fied : both are frequently found about ripe fruits. 

The larvae of a Tineid moth were found in one crop, traces of 
cocoon silk in two, and fragments of the elytra of a beetle in one. 
With all confidence the conclusion may be drawn that the band- 
tailed pigeon feeds only rarely upon insects. 

SEASONAL FOOD PREFERENCES 

In order to set forth more clearly the relation of foods to band- 
tailed pigeon migration and distribution, food items identified 
in the present study have been segregated and tabulated in four 
seasonal classifications: winter, spring, summer, and autumn. The 
seasons, arbitrarily designated, are most nearly accurate for the 
California district. 

WINTER 

The period designated as winter includes the months of Novem- 
ber, December, and January. Normally, by November most of 
the pigeons of the Pacific coast have reached California and are 
settled in their winter habitat, except for descent from higher to 
lower elevations caused by snowfall. Soon after the end of Janu- 
ary a movement northward includes part of the wintering popu- 

695766°— 47— 5 



64 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

lation. 

The stomachs and crops of 214 birds collected in four States 
during this period were available for examination. Acorns were 
taken by 172 of the 214 birds and averaged 77.2 percent of the 
winter food, with oak blossoms adding another 2.2 percent. Pine 
nuts also played an important part, contributing 8.5 percent to 
the winter food. Fruits of the madrona (Arbutus menziesii) were 
found in 37 of the birds and formed 4.8 percent of the season's 
food. Christmasberries, or toyon berries (Photinia arbutifolia) , 
made up 3.4 percent, and wheat from stubblefields 2.3 percent, of 
the winter food. These were the only foods that averaged more 
than 1 percent. 

Consideration of all data, both field and laboratory, leads to the 
conclusion that mast, acorns, and pine nuts are the basic foods 
necessary to maintain a population of band-tailed pigeons through 
the winter, and that these birds will not remain in numbers in 
areas that do not provide these foods. 

SPRING 

The period designated as the spring season covers February, 
March, and April. Beginning usually during February, the con- 
centrations of bandtails that winter in central or southern Cali- 
fornia start to move slowly northward, and by the end of the pe- 
riod the majority of the pigeons of the west have reached their 
breeding range or are close to it. 

Only 21 specimens were available for this period, collected in 
three States. Oak products, chiefly acorns, led in the spring foods ; 
they were found in 13 specimens and averaged 62.2 percent of the 
season's food. Cultivated grain, including wheat, oats, and barley, 
and cultivated peas made up 25.4 percent of the food. This season 
covers at least part of the seeding period for certain of these crops, 
and in other areas winter wheat and barley is ripening by the end 
of the spring. 

Other foods that averaged more than 1 percent during this peri- 
od included fruits of the manzanita (Arctostaphi/los) , 4.8 percent, 
and of the madrona (Arbutus menziesii), 7.4 percent, the latter 
being found in only two stomachs. 

SUMMER 

During the summer period (May, June, and July) the majority 
of the pigeons are nesting, though some may not begin to nest 
until June, and may continue brooding until well after the end of 
July. Others may begin nesting before May and may be away 
from the nest and moving about before the end of July. 

Examination of 197 specimens collected in six States furnished 
the data for the calculations for the summer period. Cultivated 
grains—wheat, oats, and barley—ranked high, averaging 25.7 per- 
cent, and were found in 80 specimens. It is certain that much of 
the grain was gleaned from stubblefields after harvest, and was 
therefore of little economic value. Cultivated cherries and prunes 
averaging 40.2 percent, led the summer food items, and occurred 



THE BAND-TAILED PIGEON 65 

in 95 specimens; domestic prunes were taken during May while 
still very small; the cultivated cherries included both sweet and 
sour varieties and were taken at ripening season. Complaints 
agamst pigeons arise most frequently from their pilfering in cher- 
ry and prune trees. 

Wild fruits of many kinds ripen during this period, and they 
are taken in wide variety. These include blackberries, raspberries, 
wild cherries, strawberries, elderberries, and fruits of the dog- 
wood, hawthorn, lote-bush {Condalia) , and others. Individually 
they composed from 1 to as high as 3 percent, and collectively 16.9 
percent, of the summer food. The blossoms and berries of the man- 
zanita and the berries of the sumac form an additional 5.8 percent. 

Acorns comprise 5.6 percent of the summer food ; some of these 
are gleaned from the ground under the oaks, but in the southern 
part of the range the acorns of the Emory oak and other species 
are reaching maturity and are taken from the trees. Pine nuts 
also enter the diet again as the new crop becomes available. 

AUTUMN 

During August, September, and October, the arbitrarily desig- 
nated autumn period, migration on the Pacific coast is under way 
and m many cases is completed ; to considerable extent the same 
is true m the Rocky Mountain States and the Southwest. The 
crops and stomachs of 207 specimens collected in six States were 
available for this period. 

With the ripening of the new acorn crop, these nuts rose in im- 
portance once more to 26.5 percent of the food and were found in 
35 birds. Pine nuts made a further advance in the diet, averag- 
mg 9.2 percent and being found in 11 birds. Cultivated grains 
(wheat, oats, and barley) and cultivated peas were found in 71 
specimens and averaged 13.7 percent of the food; most of this 
gram is taken from stubblefields. Wild fruits of many varieties 
continued to play an important part in the pigeon's diet. Fruits 
of the dogwood (Cornus) rose to 11.3 percent, elderberries to 7 
percent, cascara (Rhamnus) to 8.2 percent, salal (Grmlthcna) to 
3.3 percent, wild grape (Vitis) to 2.8 percent, wild cherries to 6.0 
percent, and a number of others ranked above 1 percent. 

DISCUSSION 

The seasonal food preferences of band-tailed pigeons as deter- 
mined by the examination of 639 stomachs and crops are shown 
m table 7. The table lists the items that totaled 1 percent or more 
of the food for each of the four seasons and gives the percentage 
of their frequency of occurrence. The number in parentheses aft- 
er the designation of the season refers to the number of specimens 
examined. 

It will be noted that during three of the four seasons, oak prod- 
ucts, largely acorns, and pine nuts, combined, led all other food 
items by a considerable margin. The availability of these nuts 
to a very large degree determines the distribution of the band- 
tailed pigeon ; a number of the oaks bear acorns only at two-year in- 



66 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 



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68 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

tervals, and both the acorn and the pine nut crops are frequently 
affected by adverse weather conditions. The winter range of the 
pigeon and its migration routes both northward and southward 
are directly influenced by the presence or absence of these nuts. 
Band-tailed pigeons have developed a liking for cultivated 
grains and peas, as well as for cultivated prunes and cherries, and 
are sufficiently adaptable to have learned to congregate in areas 
where these crops are grown ; yet study of the data available does 
not indicate that these crops are at all necessary for the mainte- 
nance of the present population. Wild fruits during the summer 
season and a plentiful supply of mast suffice to maintain them. 

USE OF GRAVEL 

As mentioned earlier, the grit or gravel found in the stomachs 
examined was measured according to its ratio to the total stomach 
content. Reference to table 5 shows that the volume of grit ranged 
from 7.5 percent in July to 31.4 percent in December. In exam- 
ining stomachs of band-tailed pigeons, it becomes evident that the 
birds do not use gravel in the digestion of foods that have a hard 
pit, as wild and tame cherries and seeds of dogwood ; only rarely 
do stomachs containing these pits also contain gravel. Apparently 
the pits are softened by the digestive juices and broken by mus- 
cular action, the fragments then serving as grinding material. 

INFLUENCE OF AGE OR SEX 

No nestlings were included in the present food study. A small 
number of flying juveniles were examined, but a survey of the 
tabulations showed no difference in their feeding habits from those 
of adult birds, nor did the feeding habits of the adult male differ 
from those of the adult female. 

FEEDING MANNERISMS 

During this study the writer observed feeding mannerisms that 
have been described by others. The alternate walking and ffying 
manner of feeding in grain stubble as described by Oilman (1903) 
was noted in several localities, and the birds hanging by their feet 
to reach food at the tips of branches, as recorded by Willard (p. 
53) was observed in connection with their feeding on acorns and 
elderberries. 

With respect to feeding hours, in some instances at least, Huey's 
statement (p. 53) was found to be correct. At other seasons feed- 
ing appeared to be intermittent all day long, periods of feeding to 
satiation alternating with hours of perching quietly in some tall 
dead tree. 

QUANTITY OF FOOD TAKEN 

The quantity of food that can be taken at one feeding is almost 
beyond belief, and after heavy feeding a digestive period of 2 or 3 
hours is needed before the bird is able to fly about in normal fash- 



THE BAND-TAILED PIGEON 69 

ion. Owing to the usually contracted state of the stomach, it is 
difficult to estimate the normal capacity of the organ ; however, it 
appears that few band-tailed pigeon stomachs contain more than 
10 cubic centimeters of food and gravel, and that about 15 cubic 
centimeters is the maximum (4 cc.=l teaspoonful). 

The following itemizations are of actual crop contents measured 
by displacements when fully dried, and indicate the volume of food 
that a pigeon crop can hold : 227 whole garden peas, 67.6 cc. ; 622 
seeds of Pinus ponderosa, 47 cc. ; 60 seeds of Pimis monophylla, 
45 cc. ; 86 seeds of Finns cdulis, 34 cc. ; 22 whole California live oak 
acorns, 38 cc. ; and 56 whole Emory oak acorns, 40 cc. Unmeas- 
ured, but in every instance forming the entire content of a single 
crop were the following items, each the largest of its group: 725 
whole kernels of wheat, 660 whole kernels of unhulled oats, 104 
whole berries of toyon, or Christmasberry, 270 whole berries plus 
550 seeds of elderberry, 56 whole fruits of madrona, 26 whole cul- 
tivated cherries, 69 whole wild cherries, 104 fruits of cascara sa- 
grada, and 80 whole fruits of dogwood. 

SUMMARY 

This report on the band-tailed pigeon (Cohimba fasciata fnsci- 
ata ) , largest member of the pigeon family now found in the United 
States, is based on field studies in Arizona, California, Colorado, 
New Mexico, Oregon, and Washington, and on reports from ob- 
servers in Nevada, Texas, and Utah. Observations from British 
Columbia and Mexico are also included. 

Personal observations and reports received indicate that in Ore- 
gon, Washington, and California the bandtail either has shown a 
definite decrease or its numbers are static. In California the sum- 
mer population seems to be increasing, but the winter population 
is no more than holding its own. In Arizona, Utah, and New Mex- 
ico the species appears to be slightly on the increase. In Colo- 
rado it seems that the population may be dropping back slightly 
from a moderate high reached two or three seasons back. In Tex- 
as the evidence indicates that the birds have not increased in the 
past 25 years and are barely holding their own. 

On California largely depends the future of the band-tailed pi- 
geon, as the bulk of the population spends the winter in that State. 
Owing to the response of the birds to an abundance of food, con- 
ditions may arise permitting excessive slaughter. Recognition 
should be made of this fact in designating bag limits, seasons, and 
shooting areas. 

Adult bandtails weighed during this study ranged from 10.3 to 
15.5 ounces. In length, the bird averages about 15 inches. 

Within the United States the nesting season varies with the 
climate of the district inhabited, and in the 8 States where nest- 
ing is now known to occur specific records cover every month from 
March through October. There was no authentic nesting record 
for Colorado until 1945, when two nests were discovered. Nesting 
locations vary almost as widely as does the forest cover within the 
range. The nest is typical of the frail nests of all members of the 



70 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

dove and pigeon group. 

Normally only one egg is laid. Incubation is said to be 14 to 18 
days. For 20 days after hatching, the adults carefully brooded 
the squabs under observation in Colorado in 1945. The male came 
to the nest about 9 a. m. and cared for the squab until between 4 
and 5 p. m., when the female returned and cared for the youngster 
until the next morning. When the squab was 20 days old both par- 
ents ceased brooding, and thenceforth each came to the nest only 
once daily to feed the young bird. 

At 17 days of age the nestling weighed 4.9 ounces, and its middle 
tail feathers measured only 28 mm. ; at 26 days of age it weighed 
8.5 ounces and the tail measured 75 mm. as compared with the av- 
erage 140 mm. of the adult bird. The squab left the nest between 
its 28th and 30th days. 

Discussion of food habits and economic status covers testimony 
obtained from numerous field observers as well as the results of 
the laboratory examination of 691 stomachs and/or crops. Of the 
stomachs and /or crops examined 639 contained sufficient food to 
serve in computations of the diet. 

Mast (largely acorns and pine nuts) furnished the largest single 
element of the food of the bandtail ; it was present in 268 of the 
specimens studied and averaged 50.1 percent of the total food. 
So important is mast to the welfare of the bandtail that it deter- 
mines not only the bird's route of migration and wintering 
grounds, but indirectly the nature and extent of the damage these 
birds inflict on crops. 

The birds' fondness for cultivated cherries and prunes was in- 
dicated by the presence of these fruits in 98 of the birds exam- 
ined, comprising 11 percent of the diet. Other related fruits ob- 
tained from wild sources (blackberries, strawberries, serviceber- 
ries, and berries of toyon and hawthorn) increased the percentage 
of food referable to the family Rosaceae to 13.6. 

Cultivated grains (wheat, oats, and barley) comprised 12.8 per- 
cent of the food, and their consumption reflects the bird's ability 
to adapt itself to the changing conditions of agriculture. A liking 
for cultivated peas, w^hich composed nearly 5 percent of the food, 
is the basis for local concentration of the birds and at times the 
cause of damage. 

Other items of vegetable origin acceptable to the bandtails are 
huckleberries, fruits of salal and other members of the heath fam- 
ily (Ericaceae), elderberries (Caprifoliaceae) , and fruits of dog- 
wood (Cornaceae) and cascara and other kinds of Rhamnaceae. 

The animal food ingested by the adult bandtail appears to be 
taken accidentally; it comprises less than one-fourth of one per- 
cent of the total food. No nestlings were examined, but juveniles 
that had left the nest showed food preferences similar to those of 
the adults. 

Establishment of isolated farms in otherwise primitive areas 
invites the possibility of damage by pigeons as well as other wild 
creatures and should be discouraged. Farmers now located in areas 
where pigeons cause damage to crops can adequately protect their 
crops by diligently following procedures recommended. 



THE BAND-TAILED PIGEON 71 

As the reproductive potential of the bandtail is low (in some 
parts of its range only one clutch is laid and this normally is com- 
posed of one egg) , it cannot withstand severe drains on its num- 
bers. For that reason demands for more liberalized shooting 
should be carefully scrutinized. 

Management of the bandtail must be built on the premise of 
safeguarding the species from threatened decimation, yet there 
should be recognition of the fact that severe crop damage may be 
inflicted and effective remedial measures must be available. 

Looking toward a long-time program of management, the basic 
population must be carefully watched and undue decimation 
through hunting prevented. Those accepted principles of forestry 
that will prevent forest destruction, and that will preserve the 
oaks and pines, and the wild fruits and berries that supplement 
mast as a food, will be advantageous to the band-tailed pigeon. 

BIBLIOGRAPHY 

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1927. Notes on the nesting of the band-tailed pigeon. Condor 29: 

121-123. 

Alcorn, J. Ray. 

1941. New and additional Nevada bird records. Condor 4o : lls-119. 
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1902. Handbook of birds of the western United States. 511 pp., 
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1928. Birds recorded from the Santa Rita Mountains in soutliern 

Arizona. Pacific Coast Avifauna 15, 60 pp., illus. 
192S. Birds of New :M('xi('o. S()7 pp., illus. Santa Fe, N. Mcx. 
Barnes, K. P. 

1916. Band-tailed pigeons alleged destruction of grain. Calif. 
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Bartol, Mary. 

1940. Hish. wild, and hniidsoni«\ Outdoor Life 8.") (4) : 2()-27. 107-10;t. 
Belding, Lymax. 

1879. A partial list of the birds of central California. U. S. Nat. 

Mus. Proc. 1: 388-499. 
1890. Land birds of the Pacific district. Calif. Acad. S'ci. Occas. 
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Bendire, Charles Emu,. 

1892. Life histories of North American birds with special reference 
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Benson, Seth BmiTRAxr. 

1935. Biological reconnaissance, Navajo Mountains, Utah. Univ. Calif. 
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Bent, Arthur Cle'V'eland. 

1932. Life histories of North American gallinaceous birds. U. S. Nat. 

Mus. Bull. 162, 490 pp., illus. 

Beegtold, William Harry. 

1912. October birds of the Gila River, New Mexico. Auk 29: 331. 
1928. A guide to the birds of Colorado. 207 pp., illus. Denver. 

Brooks, Allan Cyrh.. and Swarth. Harry Schkiavaldt. 

19 25. A distributional list of the birds of British Columbia. 
Pacific Coast Avifauna 17, lijS pp., illus. 
Bltrtch, Lewis. 

1930. Wild pigeons — Kern County, California. Calif. Dept. Agr. 
Monthly Bull. 19 (.5) : 37r)-376. 



72 NORTH AMERICAN FAUNA 58, FISH AND WILDIJFE SERVICE 

Chambers. Wii.t.ik Lke. 

1912. Who will save the band-tailed pigeon? Condor 14: lOS. 
Cooper, James Graham. 

1880. On the migration and nesting habits of West Coast birds. 
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COTTAM, Cl.\!!ENCK. 

1941. Indigo hunting and band-tailed pigeon in Utah. Condor 48: V2: 
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1938. Nesting dates in Humboldt Bay region. Condor 40: 182-183 
Dawson, William Leon. 

19l'3. The hiids of California. Student's ed. 3 vols.. 2.121 pp. 
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Dean, W. F. 

19 04. A few notes on bird life at Three Rivers, Tulare County, 
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Derby, Wii.LiA>t F. 

1920. Band-tailed pigeon nests in Sequoia National Forest. 
Calif. Fish and Game 6: 182. 

Evermann, Barton Warpen. 

1886. A list of the birds observed in Ventura Countv, California 
Auk 3: 86-94. 

Fisher, Albert Kfndkuk. 

1893. Birds of the Death Valley expedition. North Anier. Fauna 7, 
158 pp. 

Fowler, Fredkhick Hall. 

1903. Stray notes from southern Arizona. Condor 5: 68, 71. 
Gabeie2,son, Ii!a Noel, and Je^vett. Stanley GoitnoN. 

1940. Birds of Oregon. 650 pp., illus. Portland, Oreg. 
Oilman, Marshall French. 

1903. More ahout the Imiid-tailed pigeon (Culiimba fanciatd). 
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Gbinnell, Joseph. 

1898. Birds of the Pacific slo])o of Los Angeles County. Pasadena 

Acad. Sci. Pub. 2, 52 pp. 
1905. Siummer biids of Mount Pinos, California. Auk 22: 378-391. 

1913. The outlook on conserving the band-tailed pigeon as a game 

bird of California. Condor 15: 25-40. 
1915. A distributional list of the birds of California. Pacific 

Coast Avifauna 11, 217 pp. 
1928. September nesting of the band-t.-iilod jjigeon. Condor 3(1: 126. 

Bryant, Harold Child: and Storei:. Thacy Iiiwin. 

1918. The game birds of California. 642 pp. Berkeley. 

DixoN. Joseph ScattT':rgood : and Linsdale, .Tean ^Iyi:on. 



1930. Vertebrate natural history of a section of northern California 

through the Lassen Peak region. Univ. Calif. Pub. Zool. 35. 
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and Stouek. Ti;a( y Irwin. 



1924. Animal life in the Yosemite . . . 741 pp., illus. Berkeley. 
and Wythe. IMargaket Wilhel:mina. 



19 2 7. Directory to the bird life of the San Francisco Bay region. 
Pacific Coast Avifauna 18, 160 pp.. illus. 

Hagen stein, Walter M. 

1936. Late nesting of the band-tailed pigeon. Murrelet 17: 21-22. 

Henshaw, Henry Wktherbee. 

1886. Birds of the Upper Pecos River, New Mexico. Auk 3: 80. 

HuEry, I>awrence Markham. 

1913. With the band-tailed pigeon in San Diego County. Condor 15: 
151-153. 



THE BAND-TAILED PIGEON 73 

Hunter, Joseph Sr.AVTdN. 

1936. Kill of game in State is compiled (1934-35 fiscal vear). 

Calif. Conserv. 1 (8): 3. 

1937. Kill of game in State is compiled (1935-36 fiscal year). 

Calif. Conserv. 2 (8): 20. 

JEWETT, STANT.ET GoIDON. 

1941. Late nesting of the band-tailed pigeon. Condor 43: 78. 

and Gahkielson, Ira Noel. 

1929. Birds of the rortlaiid, Orcjion, area. I'licific Cojisl Avifauna 19. 
55 pp., illus. 

Johnson, O. R. 

1880. List of the birds of the Willamette Valley, Oiegon. Amei-. Nat. 
July, pp. 638-639. 

KL0PPENBri;G, H. A. 

1922. Band-tailed pigeons abundant in Phimas National Forest. Calif. 
Fish and Game 8: 57. 

KOBBE, WiT.LlAM HOFF>rAN. 

1900. The birds of Cape Disappointment, Washington. Auk 17: 349-358 

LiGON, JAMKS S'KIKET.Y. 

1927. Wildlife of New Mexico: its conservation and management. 

212 pp., illus. Santa Fe. 

McAtee, Waedo I>ek. 

1932. The need for studies in bird control in California. Calif. 
Dept. Agr. Monthly Bull. (4-5-0) : 2G9-28t). 
McLean, Donald Dudley. 

192.J. A western goshawk scatters Yosemite's band-tailed pigeon colony. 
Yosemite Nature Notes 4: 103. 

MiCHAEX, Charles Wilso.x. 

1928. Nesting time of band-tailed pigeons in Yosemite Valley. 

Condor 30: 127. 

Mili.er, Robekt ('T'XNiN(jnA.\r ; Lumley, Ellswouth L. ; and Hall. F. S. 
1935. Birds of the 3an Juan Islands, Washington. Murrelet 16 (3): 
51-65. 

MoRAN, Nathan. 

1919. Nesting of the band-tailed pigeon. Calif. Fish and Game 5: 160. 

MuNRO, James Aibxander. 

1922. The band-tailed pigeon in British Columbia. Canadian Field-Nat. 

36: 1-4. 
1924. Miscellaneous bird notes from Vancouver Island, 1923. 
Canadian Field-Nat. Ss : 149-l.m 
Nice, Margaret ]\Iorse, and Nice. Leonard Blaine. 

1924. The birds of Oklahoma. Univ. Oklahoma Bull., new series 20, 
Univ. studies 286, 122 pp., illus. May 15. Norman, Okla. 

NiEDRACH. Robert J., and Rof kwell. TiOhfrt B. 

1929. Birds of Denver and Mountain Parks. Colo. Mus. Nat. Hist. 

Pop. Ser. 5, p. 89. 

NoACK, H. K. 

1916. Band-tailed pigeons bred in captivity. Calif. Fish and Game 2: 
212. 

Oblrholsee. Harry Chui-.ch. 

1902. Some notes from west Texas. Auk 19: 300. 

Pearse, Theed. 

1935. Display of the band-tailed pigeon. Murrelet 16 (3): 71-72. 
1940. Precarious status of the band-tailed pigeon on Vancouver 
Island. Murrelet 21: 10-11. 

Presnall, Cltbtord Charles. 

19 35. Birds of Zion National Park. Utah Acad. Sci. 12: 201. 
RiDGWAY, Robert. 

1916. The birds of North and Middle America. U. S. Nat. Mus. Bull. 
50, part 7, pp. 288-291. 



74 NORTH AMERICAN FAUNA 58, FISH AND WILDLIFE SERVICE 

Shufei.dt, Robert Wilson. 

1912. The band-tailed pigeon in North Dakota. Auk 29: 539-540. 

Stillhuntee. 

19 07. South coast shooting. IX: The band-tailed pigeon. Western 
Field 11: 200-202. 

Stilljian, a. E. 

1928. Nesting of the band-tailed pigeon. Amer. Forests. May, 
pp. :^(J7-208. 
SwAKTH, Harry Schelwaldt. 
1904. Birds of the Hnachuca Mountains. Arizona. Pacific Coast Avi- 
fauna 4, 70 pp. 
1914. A distributional list of the birds of Arizona. Pacltic Coast 
Avifauna 10, 133 pp. 
Taverner, Pexcy Algernon. 

1926. Birds of western Canada. Canada Dept. Mines, Victoria 
Memorial Mus. Bull. 41 (Biol. Ser. 10), 380 pp., illus. 
Ottawa. 
1934. Birds of Canada. Canada Dept. Mines. Nat. Mus. Bull. 72 (Biol. 
Ser. 19), 445 pp., iUus. Ottawa. 
Taylor, Walter Penn. 

19 2 4. The present status of the band-tailed pigeon on the Pacific 
coast. Calif. Fish and Game 10: 1-9. 

Van DenbL'EGH, .John. 

1899. Notes on some birds of S^anta Clara County, California. Amer. 
Philos. Soc. Proc. 38: 157-180. 

Van Rossem, Adrian Joseph. 

1914. Notes from the San Bernardino Mountains. Condor 16: 145-146. 

Van Tyne, Jocelyn, and Sutton, George Misch. 

193 7. The birds of Brewster County, Texas. Univ. Michigan Mus. Zool. 
Misc. Pub. 37, illus. 
VoRHiES, Charles Taylor. 

1928. Band-tailed pigeons nesting in Arizona in September. Condor 30: 
253. 

Wales, Joseph Howe. 

1926. The coo of the band-tailed pigeon. Condor 28: 42. 

Wheelock, Irene Grosvenor. 

1904. Birds of California. 578 pp. illus. Chicago. 

Willard, Frances Cottt.e. 

1913. Some late nesting notes from the Huachuca Mountains, Arizona. 

Condor 15: 41. 
1916. Nesting of the band-tailed pigeon in southern Arizona. Condor 
18: 110-112. 

Willet, George. 

1933. A revised list of the birds of southwestern California. Pacific 
Coast Avifauna 21, 204 pp. 



THE BAND-TAILED PIGEON 



75 



INDEX 



Abundance, early day, 2. 

present status, 33. 
Acetylene flash finn, 4S. 49. 
Agricultural relationships, 3, 37-45. 

Belmvior, 6. 

calls. 5. 

courtship, 5. 

flight, 6. 

flocking, 6. 

perching, 6. 
Berries as food, 38-43, 51-53, 55-63. 

Calls, 5. 

Care of young, 13-15. 
Colunibd fasciota fnxrinfa. 1. 
Communal nesting, 11, 12. 
Control of Damage, 45-50. 

by acetylene flash gun, 48, 49. 

by flash bombs, 49. 

by paper streamers, 48. 

by shooting, 45^7. 

by signal flares, 49. 

by tree covers, 48, 49. 

permits for. 3. 49, 50. 

proper timing, 47. 
Cooper's hawk, 17. 
Courtship, 5. I 

Decoys, use of. 33. 

Depredations, cherries, 3, 38-43. 

grains. 38. 43^5. 

grapes. 42. 

prunes, 42. 

walnuts, 45. 
Description, adult, 4. 

nestling. 5. 
Development, feather, 15. 
Disease, 17. 
Distribution, general, 17. 

map, 18. 

summer, 18. 

winter, 24. 

Eggs, 12. 

description, 12. 

discovery, 2. 

number, 6, 12. 

size, 12. 
Enemies, 17. 

Falcon, prairie, 17. 
fasciala fa.sriafa. Cohimha, 1. 
Feather development, 15. 
Feeding mannerisms, 68. 
Flash bombs, 49. 
Flight, 6. 
Flocking, 6. 
Food, autumn, 65. 

berries, 38-13, 51-53, 55-68. 

general, 51-71. 

grains, 43-45, 51-53, 55-58, G3-68. 

insect, 53, 63. 

mast, 37, 38-i3, 51-53. 55-59, 63-68. 

quantity taken, 68, 69. 

seasonal preferences, 63. 



spring, 64. 
summer. 64. 
winter, 63. 

Grains as food, 43-45. 51-53, 55-58, 

63-68. 
Gravel, use of, 68. 
Growth of young, 13-15. 

Hawk. Cooper's. 17. 

goshawk, 17. 

l)rairie falcon. 17. 

sharp-shinned, 17. 
Hunting, 2. 30, 31, 32. 33. 

bag limit, 3. 

effects of, 2, 31. 32. 33. 

practices, 2. 3. 30-33. 

seascm trends, 34, 36. 

seasons, 3, 30. 

wastage In, 31-33. 

Incubation period, 12. 
Kill, records of, 31-33. 
Location of nests, 9. 

Management, 36. 

tire prevention, 37. 

increasing food supply, 37, 49. 

reforestation, 37. 

stringent protection. 37. 

wilderness maintenance, 37. 
Mast as food. 37-43, 51-53, 5;5-59, 

63-68. 
Measurements. 4, 5. 
Migration, autumn, 27. 

routes of, 29. 

spring, 26. 
Molts. 5. 

Nesting, 6-12. 

seasons. 6-7. 
Nest location, 9. 

structure, 9-10. 

Paper streamers, 48. 
Parasites, internal, 17. 
Perching, 6. 

Permits for control, 3. 49, 50. 
Pigeon, band-tailed, 1. 

discovery of, 2. 

blue. 1. " 

blue rock, 1. 

history, 2. 

milk, 13, 16. 

wild, 1. 
Plumages, adult, 4. 

nestling. 5. 
Prairie falcon, 17. 
Predators, 17. 
Protection, 3, 30. 36. 

Quantity of food taken, 68. 69. 

Range, summer. 18. 

winter, 24. 
Records, banding, 29. 



76 NORTH AMP:RICAN FAUNA 58, FISH AND WILDLIFE SERVICE 



Routes of migration. 29. 
Salt, use of. r>4. 
Season of nesting. ."> 7. 
Sharp-.shinned hawk. 17. 
Shooting, control hy. 4"— 47. 
Signal flares, 49. 
Squirrel, gray. 17. 
Streamers, paper. 48. 
Structure of nests. 9, 10. 

Tree covers. 48. 49. 

Use of decoys. '.'3. 
gravel, 68. 



.salt, -j4. 
Value as game. 29. 

Weights. 4. In. 
Western goshawk. 



17. 



Young, 13-16. 

activity of. l-"*. 
brooding of, 15. 
care of. 13-16. 
feather grow^th, 15. 
food of, 13. 
growth of. 13-16. 
we:'ght of, 4. 15. 



* U. S. GOVERNMENT PRINTING OFFICE: 1947-695766 






UNITED STATES DEPARTMENT OF THE INTERIOR 

FISH AND WILDLIFE SERVICE 



THE POCKET GOPHERS 
(GENUS THOMOMYS) OF ARIZONA 



NORTH AMERICAN FAUNA 59 




UNITED STATES DEPARTMENT OF THE INTERIOR 
J. A. Krug, Secretary 

FISH AND WILDLIFE SERVICE 
Albert M. Day, Director 



North American Fauna 59 



THE POCKET GOPHERS 
(GENUS THOMOMYS) OF ARIZONA 



BY 

EDWARD A. GOLDMAN 




UNITED STATES 

GOVERNMENT PRINTING OFFICE 

WASHINGTON : 1947 



For sale by the Superintendent of Documents, U. S. Government Printing Office 
Washington 25, D. C. — Price 15 cents 



U. S. SUPtRlMTENBENT Of OOCUMEMl* 

JUL 17 1941 



CONTENTS 



Page 

Introduction 1 

Groups of Arizona pocket gophers 1 

Economic status 6 

Species and subspecies of Thomomys 
bottae group 7 



Page 
Subspecies of Thomomys baileyi group.. 31 
Subspecies of Thomomys talpoides group 32 
Subspecies of Thomomys umbrinus group 33 



THE POCKET GOPHERS (GENUS THOMOMYS) 
OF ARIZONA 

By Edward A. Goldman, formerly Senior Blohxji.st, Biological Surveys, Division 
of Wildlife Research, Fish and Wildlife Service 



INTRODUCTION 

Pocket gophers of the genus Thomomys inhabit practically ev- 
ery part of Arizona, from the desert lowlands near the Delta of 
the Colorado River to near the tops of the highest mountains, in- 
cluding San Francisco Mountains at timber line, where the alti- 
tude is approximately 11,500 feet. Between these extremes in 
altitude the sedentary pocket gophers occupy every sort of habitat 
in which it is possible to burrow. In general, soft or sandy soil is 
preferred, but in some places there is so little soil that the exca- 
vated material consists almost entirely of small stones.. In the 
desert regions some soils are so hard and compact that excavation 
is difficult, and with plants widely spaced the food supply is ev- 
idently precarious. 

The following review of the pocket gophers of Arizona is pri- 
marily a distributional study, based mainly on the collections in 
the United States National Museum, including the extensive Bio- 
logical Surveys collection. For the loan of specimens in their 
charge, however, the writer is indebted to Laurence M. Huey, Nat- 
ural History Museum, San Diego, Calif. ; the late Dr. Joseph 
Grinnell and Dr. E. Raymond Hall, then of the Museum of Ver- 
tebrate Zoology, University of California, Berkeley, Calif. ; and to 
Dr. W. H. Burt, Museum of Zoology, University of Michigan, Ann 
Arbor, Mich. The maps were drawn by Mrs. Katheryne C. Tabb, 
of the Fish and Wildlife Service. No effort has been made to in- 
clude the many published references to pocket gophers in the 
State, most of which would add little or nothing to present knowl- 
edge of distribution as determined by specimens examined. Col- 
ors mentioned are mainly from Robert Ridgway, "Color Standards 
and Color Nomenclature," 1912. 

GROUPS OF ARIZONA POCKET GOPHERS 

The pocket gophers of Arizona are assignable to four appar- 
ently distinct groups, as follows: (1) The Thomomys bottae group, 
(2) the Thomomys baileyi group, (3) the Thomomys talpoides 
group, and (4) the Thomomys umbrinus group. 



2 NORTH AMERICAN FAUNA 5<J, FISH AND WILDLIFE SERVICE 

THOMOMYS BOTTAE GROUP f 

The Thomomys bottae group (fig. 1) embraces most of thei 
pocket gophers of Arizona. It is a remarkable assemblage of 
closely related forms, ranging from the coast of California through 
several southwestern States and east to Texas. The Colorado River 
and its canyons form an effective barrier limiting the distribution 
of pocket gophers in western Arizona, but one subspecies, albatus, 
forms a connecting link across the Delta of the Colorado River, 
where shifting channels transfer local populations from one side 
to the other. The group is also represented north of the Colorado 
River, in northwestern Arizona. 

The Thomomys bottae group presents characters as follows: 
Size variable from rather large to small ; form robust ; colors of 
upper parts varying from nearly white to cinnamon buff and rich 
ochraceous tawny, more or less mixed with black; post-auricular 
black spots small; mammae normally 4 pairs (2 pectoral and 2 
inguinal). Skull of rather heavy proportions; rostrum broad; 
nasals only slightly tapered and gradually narrowed posteriorly, 
the ends usually truncate; zygomata heavy, the maxillary arm ex- 
panded to form a prominent external angle near line of contact 
with jugal; interparietal not extending posteriorly beyond plane 
of suture between parietals and supraoccipital ; ossified external 
auditory meatus moderately developed; upper incisors broad and 
heavy, usually decurved in vertical plane at anterior ends of nasals. 

Members of the bottae group are evidently extraordinarily re- 
sponsive to environmental and genetic influences combined with 
the isolation factor, as studies reveal that each principal moun- 
tain range or valley tends to support its more or less peculiar 
form, marked by variations in combination of details of a pattern 
of general characters which is maintained with surprising uni- 
formity throughout the group. The intergradation of forms is 
clearly exhibited in some cases, and in others, in which it is not 
shown by the material at hand, the quantitative characters pre- 
sented are those known to be of subspecific value elsewhere, and 
are assumed to warrant the use of a subspecific name. Despite the 
large number of subspecies that have been described, very few 
names can be relegated to synonjiny. It is evident, however, that 
there is now an approach to the limit of the number of forms that 
can be recognized in Arizona. 

Isolation is evidently an important factor in the evolution of 
the many forms. Distribution is irregular. Local colonies, which 
may consist of numerous individuals, develop in areas where the 
soil is soft, as along a wash in the bottom of a desert valley, or in 



THE POCKET GOPHERS 




Figure 1. — Distribution of species and subspecies of Tbomomys bottae 

Arizona : 



group in 



1. 


T. 


b. 


virgineus. 


13. 


r. 


b. 


chrysonofus. 


26. 


r. 


fa. pusillus. 


2. 


T. 


b. 


nicholi. 


14. 


T. 


fa. 


sufasimihs. 


27. 


T. 


fa. grahamensis 


3. 


T. 


b. 


planhostris. 


15. 


T. 


fa. 


pafuius. 


28. 


T. 


b. extenuatus. 


4. 


T. 


b. 


trumbuUensis. 


16. 


T. 


fa. 


mutafaiJis. 


29. 


T. 


fa. catalinae. 


5. 


T. 


b. 


absonus. 


17. 


T. 


fa. 


cervinus. 


30. 


T. 


fa. parvulus. 


6. 


T. 


b. 


aureus. 


18. 


T. 


fa. 


pinalensis. 


31. 


T. 


fa. hueyi. 


7. 


T. 


b. 


peramp7us. 


19. 


T. 


fa. 


aUenus. 


32. 


T. 


fa. coiiinus. 


8. 


T. 


b. 


deserforum. 


20. 


T. 


fa. 


albatus. 


33. 


T. 


fa. modicus. 


9. 


T. 


b. 


fulvus. 


21. 


T. 


fa. 


depaupeiatus. 


34. 


T. 


alexandiae. 


10. 


T. 


fa. 


hualpaiensis 


22. 


T. 


fa. 


phasma. 


35. 


T. 


suboles. 


11. 


T. 


fa. 


desitus. 


23. 


T. 


fa. 


aridicola. 


36. 


T. 


muralis. 


12. 


T. 


fa. 


opezosus. 


24. 
25. 


T. 
T. 


fa. 
fa. 


growlerensis. 
comobabiensis. 


37. 


T. 


haiquahalae. 



4 NORTH AMERICAN FAUNA 59, FISH AND WILDLIFE SERVICE ! 

a mountain meadow. In such places the burrows may be close to- 1 
gether, with many mounds of excavated earth dotting the surface! ; 
and tunnels interspersed. Numerous mounds may mark the oper- 
ations of one or several individuals, but studies indicate that each 
burrow occupant, male or female, is solitary, except during the 
mating season and during the time that the young must remain 
in the same tunnels with their mother. Colonies may be more or; 
less isolated by areas of nearly impervious soil or by rock forma- 
tions, with breeding connections between them uncertain. Below the 
upper walls of the Grand Canyon one form, muralis, was found I 
living in isolated strips of soil only a few feet wide, bounded above |( 
and below by vertical cliffs about 300 feet high. It presents rath-i 
er well marked characters, and in this and a few similar casesJ 
the use of full specific names seems warranted, although the close 
relationship to a neighboring form is also clearly evident. 

Response to environmental conditions is shown by the markedi 
tendency of pocket gophers to assume the colors of the soils ini 
which they burrow. This tendency is shared with many othert 
mammals, but is especially noteworthy in the forms of the Tho- 
moniys bottae group. Pocket gophers of this group inhabiting 
light-colored sand are usually light shades of buff, varying to al- 
most white ; those living in reddish soils assume ruf escent tones ; 
and those from blackish terrain, especially dark volcanic soils, 
are usually deeper reddish or dark brown, and may vary to black 
in some specimens. Occasional melanistic individuals may occur 
anywhere, but black or blackish specimens are more prevalent in 
lava areas. Pocket gophers inhabiting soft, sandy ground or al- 
luvial bottomlands, where food is more abundant, or more acces- 
sible, tend to be larger than those inhabiting more rocky or ster- 
ile areas. 

THOMOMYS BAILEYI GROUP 

The Thomomys bnileyi group (fig. 2), as recognized, embraces 
several races ranging in westei'n Texas, parts of southern New 
Mexico (except the Rio Grande Valley), northern Chihuahua, and 
west to southeastern Arizona, where the group is represented by 
T. b. mearnsi. In some of the more important characters these 
races agree with those of the bottae group, but the zygomata are 
slender with slight expansion of the maxillary arm near the line 
of contact with the jugal, the rostrum is of shallow depth, and 
the upper incisors are strongly procumbent. Representatives of 
the two groups taken in the same vicinity appear to be distinct. 



THE POCKET GOPHERS 




Figure 2. — Distribution of subspecies of Thomomys talpoides, T. umbiinus. and 
r. baiieyi groups in Arizona: 



1. r. /. kaibabensis. 

2. r. /. fossor. 



3. 7". u. proximus. 

4. T. u. burti. 

5. T. u. queicinus. 



6. T. u. inteTmedius. 

7. T. b. mearnsi. 



THOMOMYS TALPOIDES GROUP 

The Thomomys talpoides group (fig. 2) is now known to con- 
stitute an assemblage of numerous forms which ranges as a whole 
from Saskatchewan, Alberta, and southern British Columbia 
southward in the Rocky Mountain region, and meets the distribu- 
tion area of the bottae group in Nevada, southern Utah, northern 
Arizona, and northwestern New Mexico. In Arizona the group 
favors the higher mountains and is represented by T. t. kaihaheii- 



6 NORTH AMERICAN FAUNA 59, FISH AND WILDLIFE SERVICE 

sis of the Kaibab Plateau and by T. t. fossor which occurs in the 
Tunitcha and Lukachukai Mountains in the extreme northeastern 
corner of the State. 

The members of the Thomomys talpoides group are similar in 
size to those of the bottae group, but the upper parts in general 
are dull cinnamon buff, the back usually overlaid with rusty, the 
black post-auricular spots are larger and the general coloration is 
duller in tone. The mammae vary from four to six pairs (2 abdomi- 
nal pairs normally present in typical talpoides). The skull is elon- 
gated and slender ; rostrum long and narrow, the sides more deeply 
excavated than in other groups, leaving the roots of upper incisors 
clearly outlined; nasals rather broad anteriorly, gradually nar- 
rowing posteriorly; zygomata slender, depressed posteriorly, the 
jugals sloping upward to points of contact with maxillae ; ossified 
external auditory meatus large and prominent; interparietal ex- 
tends posteriorly beyond plane of posterior border of parietals; 
upper incisors narrow and thin, decurved about as in bottae group. 

THOMOMYS UMBRINUS GROUP 

The Thomomys umbrinus group (fig. 2), composed of small 
pocket gophers, is widely dispersed in Mexico. Like the bottae 
group, it is subdivided into numerous geographic races, several 
of which are restricted to the mountains of southeastern Arizona. 

In color of upper parts the members of the umbrinus group are 
between cinnamon brown and russet, varying to sayal brown in 
proximus, the back usually deep black along the median line; 
black post-auricular spots usually large and confluent with black 
of back in some specimens. The skull is slender, with brain case 
smoothly rounded, the temporal ridges inconspicuous or absent; 
rostrum short, and moderately broad; nasals distinctly wedge- 
shaped, narrowing posteriorly, the ends usually emarginate ; zygo- 
mata slender, the sides nearly parallel; ossified auditory meatus 
small ; upper incisors relatively broad and heavy, decurved in ver- 
tical plane at anterior ends of nasals in Arizona forms, strongly 
procumbent in numerous Mexican races. 

ECONOMIC STATUS 

The pocket gophers of all groups consume plant food, but their 
widespread burrowing activities, which tend to stir the soil, are, 
under natural conditions, beneficial to plant growth and thus in- 
directly to other animals dependent on plants. Erosion of the 
land surface may, however, be started by water entering the 
tunnels, especially where these extend up and down steep slopes. 
Beneficial habits are, therefore, offset by some destructive ef- 



THE POCKET GOPHERS 7 

fects, and pocket gophers seem rather neghgible in the biotic 
complex. The desert forms are limited in numbers by the food 
supply and by natural enemies. Where desert land is brought 
under irrigation and cultivation, however, the food supply is 
greatly increased, predation is lessened, and the pocket gopher 
population may be expected to rise inordinately unless effective 
control measures are adopted. On cultivated lands severe dam- 
ages result from crops directly consumed, or roots severed, and 
from the tunnels which often cause breaks in irrigation ditches. 

SPECIES AND SUBSPECIES OF THOMOMYS 
BOTTAE GROUP 

THOMOMYS BOTTAE VIUGINEUS Goldman 
ViBGiN Valley Pocket Gophek 

Thomomys bottae virgineus Goldman, Biol. Soc. Wash. Proc. 50: 133, Sep- 
tember 10, 1937. 

Type. — From Beaverdam Creek, near confluence with Virgin 
River, at Littlefield, northwestern Arizona (altitude 1,500 feet) ; 
collected by Luther C. Goldman, October 16, 1936. 

General characters. — A cinnamon-buff form, similar to cen- 
tralis of eastern Nevada, but skull relatively narrower, more 
elongated, the zygomata less widely spreading, more distinctly 
bowed inward near middle of jugals. Similar to nicholi of the 
Shivwits Plateau region, but somewhat darker, and skull differ- 
ing in about the same characters as from centralis. 

Measurements. — Average of three adult male topotypes: Total 
length, 232 (232-232) ; tail, 74 (72-75) ; hind foot, 31.5 (31-31.5) 
millimeters. Two adult female topotypes, respectively, 207, 210; 
62, 58 ; 28, 29.5 millimeters. 

Distribution and habitat. — The range of virgineus includes the 
Virgin River Valley below the canyon traversed by the river in 
breaking across the fault line marked by the Beaverdam Moun- 
tains on one side, and the Grand Wash Cliffs on the other, just 
above the type locality. East of the Virgin Valley this form is 
found in an extremely arid section near Pakoon Spring, along 
Grand Wash. The general area inhabited is well down in the 
Lower Sonoran Zone. On the bottomlands along Beaverdam 
Creek and the Virgin River, pocket gophers are numerous 
enough in some places to be destructive to alfalfa, which is grown 
on a limited scale. 

THOMOMYS BOTTAE NICHOLI Goldman 

Shivwits Plateau Pocket GoPHiai 

Thomomys bottae nicholi Goldman, Jour. Wash. Acad. Sci. 28 (7): 337, July 
15, 1938. 

696493°— 47— 2 



8 NORTH AMERICAN FAUNA 59, FISH AND WILDLIFE SERVICE 

Type. — From 20 miles south of Wolf Hole (road to Parashonts), 
Shivwits Plateau, Mohave County, Ariz, (altitude 5,000 feet) ; 
collected by Luther C. Goldman, August 6, 1937. 

General characters. — A light, cinnamon-buff subspecies of me-i 
dium size. Closely allied to trumbullensis of the neighboring Mount i 
Trumbull lava area, but paler buff, the back less mixed with ! 
black. Similar to virgineus of the Virgin River Valley below the 
break through the Beaverdam Mountains, but somewhat darker; 
skull differing in detail, the frontals broader, zygomata more 
widely spreading, and the jugals not distinctly bowed inward as 
in virgineus. 

Measurements. — Two adult male topotypes, respectively : Total 
length, 229, 208 ; tail, 65, 59 ; hind foot, 27.5, 28 millimeters. Two 
adult females from Saint George, Utah, respectively: 208, 204; 
71, 63 ; 27, 27.5 millimeters. 

Distribution and habitat. — The range of this pocket gopher is 
the Shivwits Plateau region, on the terrace between the Hurri- 
cane Ledge on the east, and the Grand Wash Cliffs on the west, 
north of the Grand Canyon, northwestern Arizona. The general 
area, at 4,500 to 5,000 feet, is in the lower part of the Upper So- 
noran Zone, but slopes down at the north end to 2,500 feet al- 
titude in the Lower Sonoran Zone at Saint George, in the Virgin 
River Valley, just across the Utah boundary. The pocket gophers 
occur irregularly in small, widely scattered colonies, on land com- 
monly overgrown with Atriplex and sagebrush {Artemisia tri- 
dentata) . 

THOMOMYS BOTTAE PLANIROSTRIS Burt 

ZioN Pakk Pocket Gopher 

Thoniomys pcrpallidns planirostris Burt, Biol. Soc. Wash. Proc. 44: 3.S, May 
8, 1931. 

Type. — From Zion National Park, Utah (altitude 4,400 feet) ; 
collected by A. Brazier Howell, May 4, 1920. 

General characters. — Size medium but form robust ; colors rich, 
the upper parts distinctly tawny, little modified by dark-tipped 
hairs. Allied to absonu^ of House Rock Valley, but colors bright- 
er, more tawny, and proportions heavier ; skull more massive, with 
heavier dentition. 

Measurements (from Burt). — Average of eight adult male topo- 
types: Total length, 238.3 (222-261); tail, 75.6 (66-83); hind 
foot, 32.4 (31-34) millimeters. Average of eight adult female topo- 
types: 215 (205-228); 71 (61-78) ; 30.9 (30-33) millimeters. 

Distribution and habitat. — The Zion Park pocket gopher is found 
at 4,500 feet on the broad Upper Sonoran, Atriplex overgrown, ter- 



THE POCKET GOPHERS 9 

-ace near Fredonia and the vicinity of Kanab Wash to westward. 

This brightly colored subspecies apparently ranges into Arizona 
through the Short Creek Valley along the western and southern 
base of the Vermilion Cliffs west of Fredonia. The burrows in 
! sandy soil were noted at intervals for miles west of Fredonia. 
jvVhile planirostris and absomis occur at points not far apart, and 
I might be expected to meet along Johnson Creek east of Fredonia, 
i these animals are very local in distribution and may not be in di- 
jrect contact. Both forms occur in Zion National Park, but their 
! ranges in the park appear to be completely separated. The floor 
')f the narrow valley, at 4,400 feet altitude, in the park, is the type 
j locality of pkmirostris, while absomis is found at 5,500 to 5,700 
jeeet in the heads of small canyons and on the plateau near the 
!9ast entrance above the barrier formed by the escarpment or east- 
:ern wall of the valley. At Fredonia pocket gophers enter alfalfa 
i fields, but are not sufficiently numerous to be very destructive. 

THOMOMYS BOTTAE TRUMBULLENSIS Hall and Davis 

Mount Trumbxjll Pockbt Gopher 

iThonwmvft hottar fnimhnllensis Hall and Davis, Biol. Soc. Wash. Proc. 47: 
51. February 9, 1934. 

Type. — From 3 miles south of Nixon Spring, Mount Trumbull, 
Mohave County, Ariz, (altitude 6,500 feet) ; collected by Seth B. 
Benson, May 26, 1933. 

General characters. — A dark-colored lava-area-inhabiting sub- 
species. Upper parts normally between cinnamon and cinnamon- 
buff, usually rather heavily mixed with black. Of 38 specimens 
examined, 7 are deep glossy black, except the feet and the tip of 
the tail, which are white. Similar to nicholi of the closely adjoin- 
ing, but lower, Shivwits Plateau, and to absonus of House Rock 
Valley, but color darker than either, the upper parts more heavily 
mixed with black. Compared with planirostris, typical in Zion 
National Park, Utah, but which also occurs at Fredonia, Ariz., 
trumbullcnsis is smaller and darker, the upper parts more exten- 
sively mixed with black, less tawny ; skull slenderer, with lighter 
dentition. 

Measurements. — Average of three adult male topotypes : Total 
length, 234 (228-238) ; tail, 76 (71-83); hind foot, 30.5 (30-31) 
millimeters. Two adult female topotypes, respectively: 210, 204; 
68, 56 ; 28, 26.5 millimeters. 

Distribution and habitat.— The range of this subspecies is the 
lava and cinder area centered about Mount Trumbull on the pla- 
teau near the north rim of the Grand Canyon. The area lies 
mainly along the boundary between the Upper Sonoran and Tran- 



10 NORTH AMERICAN FAUNA 59, FISH AND WILDLIFE SERVICE 

sition Zones at 6,000 to 6,500 feet, but extends up to 7,00,0 feel 
on the slopes of Mount Trumbull and Mount Logan, and burrows 
were observed at about 4,500 feet altitude on the prominent cindei 
cone at the lower end of Toroweap Valley. As pointed out by Hal 
and Davis (op. cit. : 52), the darker color of this subspecies, com- 
pared with the neighboring geographic races, may indicate a re 
sponse to environmental conditions. 

THOMOMYS BOTTAE ABSONUS Goldman 

House Rock Valley Pocket Gopher 

Thotnomijs perptillidus absonns Goldman, Join*. Wash. Acad. Sci. 21 (17) : 425 
October 19, 1931. 

Type. — From Jacobs Pools, House Rock Valley, Coconino Coun-i 
ty, northern Arizona (altitude 4,000 feet) ; collected by E. A,\ 
Goldman, June 7, 1931. 

General characters. — A dull grayish buffy subspecies of me- 
dium size with a narrow, slender skull. Closely allied to planiro- 
stris of Zion National Park, Utah, and the vicinity of Fredonia, 
Ariz., but somewhat slenderer, less tawny, and skull less massive, 

Measurements. — Average of three adult male topotypes : Total^ 
length, 231 (228-234) ; tail, 77 (74-82); hind foot, 31 (30-32.5)' 
millimeters. Two adult female topotypes, respectively: 210, 217; 
69, 70 ; 30, 29 millimeters. 

Distribution and habitat. — The range of absonus in Arizona is* 
probably restricted to House Rock Valley, but extends north ini 
Utah to the eastern entrance of Zion National Park, at 5,700 fee1< 
altitude. 

House Rock Valley occupies a broad depression with a gener- 
ally level bottom lying in the Upper Sonoran Zone along the north- 
ern side of the Marble Canyon of the Colorado River. This reach 
of the river bisects the interior basin of which House Rock Val- 
ley is the northern half, above the upper entrance to the Granc 
Canyon, and forms a barrier limiting the distribution of this sub- 
species and of most of the other mammals of the region. The 
bottom of House Rock Valley is gashed by side canyons of the 
Colorado, and the dispersal of this pocket gopher is much re- 
stricted even here. It has been found inhabiting soft sand extend- 
■ng for several miles out over the floor of the valley from Jacobs. 
Pools, a spring at the western base of the escarpment markingj 
the great fault line known as the Vermilion Cliffs. Burrows are 
common along the highway through low-growing shrubby vege- 
tation, largely Atriplex bushes and Coleogyne ramossissima, s 
dominant species on poor soils. But the pocket gophers feed tc 



THE POCKET GOPHERS 11 

a considerable extent on the roots and tender growing tops of the 
large white poppy {Avffemone). 

THOMOMYS BOTTAB AUREUS Allen 

Painted Desert Pockp^t Uopher 

Tlunnomya aureus Allen, Amer. AIiis. Nat. Hist. Bull. 5: 49, April 2S, 1893. 
Thomdmya lafirostri.-t Morriam. Proe. P>i(il. S(wc. Wnsh. 14: 107, July 19, 1901. 

Type from Tanner Crossing, about 'A miles above Cameron, Little Colorado 

River, Coconino County, Ariz. 

Type. — From Bluff, San Juan County, Utah; collected by 
Charles P. Rowley, May 12, 1892. 

General characters. — A large, cinnamon-buff or golden-colored 
subspecies. Closely allied to and probably intergrading with per- 
amplus of the higher mountains of the northeast corner of Ari- 
zona. 

Measurements. — Average of four adult male topotypes : Total 
length, 241 (232-252) ; tail, 73 (69-80) ; hind foot, 31 (30-32) milli- 
meters. Average of four adult female topotypes: 222 (215-229) ; 
66 (64-72; 30 (30-31) millimeters. 

Distribution and habitat. — The range of aureus extends from 
the San Juan Valley, Utah, into northeastern Arizona, along the 
valley of Chin Lee Creek, and embraces as a whole the major part 
of the Painted Desert region to the Little Colorado River. Over 
this area, however, the pocket gophers are very irregularly dis- 
tributed in more or less isolated colonies that favor the more fer- 
tile ground, usually in the valleys, in the Upper Sonoran Zone. 
Specimens have been obtained at localities varying in altitude 
from about 4,000 feet along the Little Colorado River to 6,500 
feet in the Pueblo Colorado Valley at Ganado. The dominant veg- 
etation of the region occupied consists largely of Atriplex bushes 
or sagebrush (Artemisia tiidentata) , with a scattered growth 
of nut pines and junipers along the valley borders. Thomomys 
lutirostris was based on a single specimen from Tanner Crossing, 
near Cameron, on the Little Colorado River. Efforts to obtain 
topotypes, made by various collectors including the writer, have 
been unsuccessful. The locality is a very barren one, made more 
so by overgrazing by domestic stock in recent years, and if pocket 
gophers still occur they must be rare as we found no trace of 
them. The skull of the type specimen of latirostris, an old male, 
has a very broad rostral portion and is believed to be abnormal, 
as no such character appears in specimens from Tuba City, Wins- 
low, and Oraibi, which are in the same general faunal area. Be- 
tween the range of aureus along the valley of the Little Colorado 
River and that of the widely different subspecies fulvus of the 



12 NORTH AMERICAN FAUNA 59, FISH AND WILDLIFE SERVICE 

Mogollon Plateau is an arid belt in which pocket gophers have noi j 
been detected. 

THOMOMYS BOTTAE PBRAMPLUS Goldman 

TuNiTCHA Mountain Pocket Gopher 

Thomomys fulvus peramplus Goldman, Jour. Wash. Acad. Sci. 21 (17): 423 
October 19, 1931. 

Type. — From Wheatfields Creek, west slope of Tunitcha Moun-il 
tains, Apache County, northeastern Arizona (altitude 7,000 feet) j 
collected by Paul Trapier, June 23, 1927. 1 

General characters. — A large, dark-colored subspecies, closeljl: 
resembling apache of northern New Mexico, but upper parts dull 
er, the sides vinaceous-buff instead of ochraceous-buff ; skull mon 
elongated; nasals longer and broader, less wedge-shaped poste- 
riorly. Contrasting strongly with the lighter cinnamon or cinna- 
mon-buff tones of aureus, which inhabits parts of the neighbor- 
ing desert region, in dark, dull coloration, but cranial characters 
indicate close relationship and the two probably intergrade in 
places along the basal slopes of the mountains. 

Measurements. — ^Average of four adult male topotypes: Tota' 
length, 246 (240-255) ; tail, 80 (60-90) ; hind foot, 35 (34-37) 
millimeters. Average of six adult female topotypes : 230 (225-240) 
75 (65-88) ; 32 (31-33) millimeters. 

Distribution unci habitat — The Tunitcha Mountain pocket go- 
pher is known from 7,000 feet altitude on Wheatfields Creek up 
to 8,000 feet on the upper slope of the range. It also occurs at 
7,000 feet altitude in the valley at St. Michaels on the easternni 
side of the Defiance Plateau. This pocket gopher probably occurs 
irregularly in suitable places throughout this high mountainous 
section of the State. It gives way, however, at the higher eleva-^'j 
tions in the Tunitcha Mountains to Thomomys talpoides fossor, 
which tends toward chestnut color, with large, conspicuous, black 
ear patches. The general habitat of peramplus is in the yellow 
pine belt of the Transition Zone. 

THOMOMYS BOTTAE DESERTORUM Mebriam 

Detrital Vallett Pocket Gopher 

Thomomys desertorum Merriam, Biol. Soc. Wash. Proc. 14: 114, .Tuly 19, 1901, 

Type. — From Mud Spring, Detrital Valley, Mohave County^ 
Ariz. ; collected by Vernon Bailey, February 21, 1889. 

General characters. — A small, tawny subspecies, similar in color 
and closely allied to desitus of the Big Sandy River Valley, but 
smaller. 

Measurements. — An adult male : Total length, 200 ; tail verte- 



THE POCKET GOPHERS 13 

3rae, 68 ; hind foot, 26 millimeters. Average of four females : 190 ; 
30 ; 25.5 millimeters. 

Distribution and habitat. — This little pocket gopher occupies 
the Lower Sonoran desert region of the broad Detrital Valley and 
neighboring areas lying mainly at about 3,500 feet altitude in the 
angle" formed by the bend of the Colorado River north and west 
of the Hualpai Mountains in the northwestern part of the State. 
It ranges to a somewhat higher elevation in the Upper Sonoran 
Zone, however, near its southern limit in the Chemehuevis or Mo- 
have Mountains. East of the northern end of the Hualpai Moun- 
tains it doubtless intergrades with desitus. The burrows are lo- 
cated in hard upland soil among tree yuccas and a varied assort- 
ment of other desert vegetation, including cactuses. 

THOMOMYS BOTTAE FULVUS (Woodhouse) 

Fulvous Pocket Gopher 

ileoDii/.s ftilvus Woodhouse, Acad. Nat. Sci. Phila. Proc. 6: 201, 1852. 
Thoniomys bottae nasutus Hall, Biol. Soc. Wash. Proc. 45: 96, June 21, 1932. 

Type from West Fork of Black River, Apache County, Ariz, (altitude 7,550 

feet) ; collected by Annie M. Alexander, June 14, 1931. 

Type. — From San Francisco Mountains, Coconino County, 
Ariz. ; collected by S. W. Woodhouse, October 1851. 

Genend cJia.racters. — Pocket gophers of medium size, distin- 
guished in the region south of the Grand Canyon by dark, rusty 
brown coloration associated with that of the dark lava soil they 
inhabit. Skull with wide-spreading zygomata and small auditory 
bullae. Closely allied to mutabilis of the Verde Valley, but dark- 
er, the upper parts more extensively mixed with black; skull less 
massive ; basicranial region narrower ; auditory bullae smaller. 

Measurements. — Average of five males : Total length, 219 ; tail 
vertebrae, 70 ; hind foot, 30 millimeters. Average of five females : 
209 ; 66 ; 29.2 millimeters. 

Distribution and habitat. — The fulvous pocket gopher is the 
most widely dispersed of the numerous representatives of the 
Thomomys bottae group found within the State. Its distribution 
area occupies the whole of the elevated Coconino and Mogollon 
Plateau regions, extending from the southern rim of the Grand 
Canyon southeastward to the White Mountains and on far into 
New Mexico. A spur from the main range reaches south through 
the high country to the Bradshaw Mountains west of the Verde 
River Valley. This pocket gopher ranges mainly in the yellow 
pine forests of the Transition Zone above 5,000 feet altitude, but 
ascends into the Canadian Zone on San Francisco Mountains and 
the White Mountains. The general region is marked by cinder 



14 NORTH AMERICAN FAUNA 59, FISH AND WILDLIFE SERVICE 

cones and old lava beds, indicating volcanic activity in the past 
The decomposing cinders and lava produce dark-colored soils thaii 
are reflected in the color tones not only of the pocket gophers bui 
of many of the other small mammals. Along the southern side of 
the Mogollon Plateau fulvvs intergrades with mutabilis, which 
ranges at lower levels. On the long gradual desert slope from th( 
top of the plateau toward the Little Colorado River pocket gophen 
are generally absent, and no direct connection with aureus, which 
inhabits parts of the Little Colorado Valley and Painted Desert ' 
is apparent. 

In the yellow pine forests the pocket gophers occur irregularlj 
in colonies wherever there is sufficient soil for their excavations 
but they favor the soft soil of open grassy meadows. 

THOMOMYS BOTTAE HUALPAIENSIS Goldman 

HuALPAi Mountains Pocket GoPHEa? 

Thnmomys hottae hualpaicnsis Goldman, Jour. Wash. Acad. Sci. 26 (3): 114 
March 15, 1936. 

Type. — From Hualpai Peak, Hualpai Mountains, Mohave Coun- 
ty, Ariz, (altitude 7,000 feet) ; collected by E. A. Goldman, Oc- 
tober 6, 1917. 

General characters. — A light ochraceous buffy subspecies oil 
medium size. Allied to desitus of the adjoining valley of the Bigi 
Sandy River; similar in size but paler; brain case lower, nasals 
more wedge-shaped. Compared with desertor'um of the Detrital 
Valley: considerably larger and paler. 

Measurements. — The type, an adult male: Total length, 245; 
tail, 78 ; hind foot, 31.5 millimeters. 

Distribution and habitat. — Known only from 6,500 to about 
7,500 feet altitude in the Transition Zone on the slopes of the 
Hualpai Mountains. The burrows occur in soft spots in yellow 
pine and oak timber. 

THOMOMYS BOTTAE DESITUS Goldman 

Big Sandy Rivek Poc'Kp:t Gophek 

Tho})i(»inis hottae dcsitiis Goldman, Jour. Wash. Acad. Sci. 26 (3) : 113, March 
15, 1936. j 

Type. — From Big Sandy River, near Owen, Mohave County, 
Ariz, (altitude 2,000 feet) ; collected by E. A. Goldman, Septem- 
ber 21, 1917. I 

General characters. — A medium-sized, tawny subspecies ; color' 
about as in desertorum of the neighboring Detrital Valley region, 
but size much larger. Size about as in fulvus of the Mogollon Pla- 



THE POCKET GOPHERS 15 

:eau, but color lighter, clearer tawny, the back less mixed with 
3lack. 

Measwements. — An adult male : Total length, 230 ; tail verte- 
brae, 70; hind foot, 30.5 millimeters. An adult female: 210; 62; 
29.5 millimeters. 

Distribution and habitat. — This pocket gopher occupies the Big 
Sandy River Valley. Thence it ranges east to Kirkland, Yavapai 
County. Its distribution area is in the Lower Sonoran Zone at 
2,000 to 4,000 feet altitude. In the Big Sandy River Valley desitus 
is confined mainly to the loose sand along the broad alluvial river 
bottom, the excavated mounds of earth often appearing close to 
the edge of the water where the stream is bordered by willows 
and Baccharis bushes. 

THOMOMYS BOTTAE OPEROSUS Hatfield 
Peeples Valley Pocket Gopher 

Thomornvs hottae opcroxus Hatfield, Chicago Acad. Sci. Bull. 6 (8) : 151, .Tan- 
iiary 12, 1942. 

Type. — From Peeples Valley, 6 miles north of Yarnell, Yavapai 
County, Ariz, (altitude 4,400 feet) ; collected by Roy Komarek, 
May 30, 1937. 

Gene7'al characters. — From original description : Size large ; tips 
of hairs on back cinnamon to middorsal area which is blackish; 
sides pinkish buff; top of head blackish; ears surrounded by 
black ; skull broad, with widely spreading zygomata. Differs from 
fulvus in larger size, more widely spreading zygomata, greater 
mastoid breadth, and heavier dentition. Differs from mutabilis 
in darker color, with more black on nose, occiput, and back. Dif- 
fers from patulus and desitus in darker color, longer tail, and 
more widely spreading zygomata. 

Measurements. — From original description: Average of three 
adult males: Total length, 232.3 (228-237); tail, 81.3 (80-82); 
hind foot, 29 millimeters. Average of eight adult females: 216.5 
(202-234) ; 71.4 (65-79) ; 27.4 (26-29) millimeters. 

Distribution and habitat. — No specimens of this subspecies have 
been examined by the writer. It is known only from the type lo- 
cality and appears to be a local race, perhaps restricted to Peeples 
Valley, which is somewhat isolated, midway between the desert 
country of southern Arizona and the high plateau to the north- 
ward. 

THOMOMYS BOTTAE CHRYSONOTUS Grinitoll 
Golden Pocket Gophex 

Thomomys chrysonotus Grinnell, Univ. Calif. Pub. Zool. 10: 174, June 7, 1912. 

Thomomys fulvus flavidus Goldman, .Jour. Wash. Acad. Sci. 21 (17) : 417, 

October 19, 1931. Tyi>e from Parker, Yuma County, Ariz, (altitude 350 feet). 

696493°— 47— 3 



16 NORTH AMERICAN FAUNA 59, FISH AND WILDLIFE SERVICE 

Type. — From Ehrenberg, Yuma County, Ariz.; collected by 
Frank Stephens, March 27, 1910. 

General characters. — A large ochraceous-buff or golden yellow- 
ish subspecies with an angular massive skull. Similar to albntus. 
but upper parts ochraceous-buff instead of pinkish buff; skulls 
much alike; auditory bullae larger, more inflated in chrysonotus. 

Measurements. — An adult male: Total length, 249; tail verte- 
brae, 83; hind foot, 33 millimeters. An adult female: 224; 67 ;i 
31.5 millimeters. 

Distribution and habitat. — The golden pocket gopher occupies 
the alluvial bottomlands along the east side of the Colorado Rive» 
from near Ehrenberg north to Parker, and invades the adjoiningi 
gravelly mesa overgrown with creosotebush (Covillea glutinosa){ 
to some extent. The opposite, or western, side of the Colorado 
River Valley is inhabited by Thomomys bottae riparius, the rivei 
serving as a barrier between the two forms. Within the main 
range of the golden pocket gopher on the bottomlands the bur- 
rows are numerous in the soft alluvial soil and extend down in 
places into the arrowweed {Pluchea sericea) belt near the edge 
of the water. 

THOMOMYS BOTTAE SUBSIMILIS Goldman 

Harquahala Mountain Pocket (Jopher 

Thomomys fulvus subsimilis Goldman, Biol. Soc. Wash. Proc. 46: 71, Apri" 
27, 1933. 

Type. — From Harquahala Mountains, Yuma County, Ariz, (al- 
titude 3,000 feet) ; collected by E. A. Goldman, October 14, 1917. 

General characters. — A very small cinnamon-buff animal with 
a weakly developed skull. Similar to desertorum of the Detrital. 
Valley region, but still smaller and paler; skull more delicate ini 
structure. 

Measurements.— The type, an adult female: Total length, 186 ;• 
tail vertebrae, 60 ; hind foot, 25 millimeters. 

Distribution and habitat. — This tiny pocket gopher is knowni 
only from a single specimen from 3,000 feet altitude in the Har 
quahala Mountains. The mountain slopes are rocky, with little( 
soil, but the animal is likely to be found in the softer spots up tci 
near the summit at about 5,000 feet altitude. The mountains arei 
Lower Sonoran in faunal character up to near the top, where ai 
few Upper Sonoran Zone elements appear. 

THOMOMYS BOTTAE PATULUS Goldman 

Hassayampa Valley Pocket Gopher 

Thomomys bottae patulus Goldman, Jour. Wash. Acad. Sci. 28 (7): 341 July 
15, 1938. 



THE POCKET GOPHERS 17 

Type. — From bottomland along Hassayampa River, 2 miles be- 
low Wickenburg, Maricopa County, Ariz, (altitude 2,000 feet) ; 
collected by Luther C. Goldman, September 16, 1937. 

General characters. — A large subspecies, similar to cervinus of 
the Salt River Valley, but upper parts more vivid in color, near 
cinnamon or cinnamon-bufF instead of vinaceous-buff or fawn; 
skull shorter, less angular. Somewhat resembling mutabilis of 
the Verde River Valley and desitus of the Big Sandy River Valley, 
but larger and paler than either. 

Measurements. — An adult male and an adult female topotype, 
respectively: Total length, 240, 215; tail, 80, 60; hind foot, 31, 
29 millimeters. 

Distribution and habitat. — Known only from the type locality, 
but probably has an extensive range in alluvial soil along the val- 
ley of the Hassayampa River. The subspecies patidus is abun- 
dant in alfalfa fields, where it becomes somewhat troublesome to 
farmers. The distribution of pocket gophers appears to be dis- 
continuous in the arid areas bordering the Hassayampa River 
Valley. 

THOMOMYS BOTTAE MUTABILIS Goldman m 

Vebde Valley Pocket Gopher 

Thomomys fuhms mufabirni Goldman, Biol. Soc. Wash. Proc. 46: 75, April 
27. 1933. 

Type.- — From Camp Verde, Yavapai County, Ariz, (altitude 
3,200 feet) ; collected by Walter P. Taylor, July 25, 1916. 

General characters. — A medium-sized, cinnamon-buff subspecies. 
Closely allied to fulvus, but paler, the upper parts less mixed with 
black; skull more massive; basicranial region broader; auditory 
bullae larger. 

Measurements. — An adult male: Total length, 236; tail verte- 
brae, 71 ; hind foot, 31 millimeters. Average of 7 adult females : 
216 ; 68 ; 29 millimeters. 

Distribution and habitat. — The Verde Valley pocket gopher in- 
habits the valleys and lower slopes of the mountains in the Verde 
and Salt River drainages along the southern side of the Mogollon 
Plateau from Camp»Verde east to the Gila Mountains, Graham 
County. Vertical range from about 2,500 to 4,500 feet. This sub- 
species inhabiting a region arid in general character favors the 
softer soils along streams. 

THOMOMYS BOTTAE CERVINUS Allen 

Fawn-colored Pocket Gopher; Phoenix Pocket Gophek 

Thomomys cervinus Allen, Amei-. Miis. Nat. Hist. Bull. 7: 208, June 29, 1895. 



18 NORTH AMERICAN FAUNA 59, FISH AND WILDLIFE SERVICE 

Type. — From Phoenix, Maricopa County, Ariz, (altitude 1,000 
feet) ; collected by J. Diefenbach, October 20, 1894. 

General chamcters. — A large vinaceous-buff or light fawn-col- 
ored subspecies. Similar to albatus of the lower Colorado River 
Valley, but larger; color darker (upper parts near pale pinkish 
buff or pinkish buff in albatus) ; skull more elongated, with rela- 
tively narrower brain case ; auditory bullae relatively larger. Al- • 
lied to modicus of the Altar and Santa Cruz Valleys, but larger; 
color paler (near wood brown, varying to tawny in modicus) ; 
skull of similar proportions, but much larger, more massive. 

Measurements. — Average of three adult male topotypes: Total 
length, 253 (251-255) ; tail, 84 (77-90) ; hind foot, 34.5 (34-36) 
millimeters. Average of three adult female topotypes: 246 (239- 
255); 78 (73-81); 34 (33-34.5) millimeters. 

Distribution and habitat. — The range of this large pocket go- 
pher is in the Salt River and Gila River Valleys, near Phoenix, and 
southwest along the latter watercourse to Gila Bend, where it in- 
tergrades with the desert subspecies, aridicola. The general area 
is in the Lower Sonoran Zone at from 700 to 1,000 feet altitude.. 
The fawn-colored pocket gopher is associated with such native 
vegetation as the mesquite {Proso^is juliflora), catsclaw {Aca- 
cia greggii), and paloverde {Cercidium torreyanum), but with 
the rapid development of agriculture it has invaded the fields and 
multiplied greatly in numbers. Owing to the extent of its depre- 
dations in an important agricultural area, this pocket gopher 
should probably be rated as the most destructive of the numerous 
geographic races that occur within the State. 

THOMOMYS BOTTAE PINALENSIS Goldman 
Pinal Mountains Pocket Gopher 

Thomomys bottae pinalensis Goldman, Jour. Wash. Acad. Sci. 28 (7): 342", 
July 15, 1938. 

Type. — From Oak Flat, 5 miles east of Superior, Pinal Moun- 
tains, Ariz. ; collected by Walter P. Taylor, May 22, 1924. 

General characters. — A very small cinnamon subspecies, with 
a narrow, slenderly formed skull. Most closely allied to mutabilis 
of the adjoining region to the north, but much smaller and dark- 
er ; skull smaller, narrower, less massive. 

Measurements. — The type, an adult female : Total length, 195 ; 
tail, 56 ; hind foot, 24 millimeters. 

Distribution and habitat. — The burrows of this small subspecies 
were noted in numbers in various places along the highway be- 
tween Superior and Globe across the upper slopes of the Pinal 



THE POCKET GOPHERS 19 

Mountains. The animal favors the softer soil in small openings in 
the oak woods of the Upper Sonoran Zone. 

THOMOMYS BOTTAB ALIENUS Goldman 
Upper Gila Valley Pockett Gopher 

Thnviomys Imttae alienus Goldman, Jour. Wash. Acad. Sci. 28 (7): 338, July 
15, 1938. 

Type. — From Mammoth, San Pedro River, Pinal County, Ariz. 
(altitude 2,400 feet) ; collected by E. A. Goldman, November 4, 
1936. 

Geneml charncters. — A large, rich rufescent subspecies, allied 
to cervinus of the Salt River Valley, but smaller ; upper parts near 
cinnamon instead of vinaceous-buff. Similar to toltecus of the 
Casas Grandes Valley, northwestern Chihuahua, but color more 
vivid ; skull with lower brain case ; upper incisors less projecting 
forward. Larger, less distinctly tawny than mutabilis of the Rio 
Verde and Salt River drainages. 

Measurements. — Average of four adult male topotypes: Total 
length, 240 (230-254) ; tail, 68 (58-80) ; hind foot, 32 (30-33.5) 
millimeters. Average of four adult female topotypes: 212 (207- 
218) ; 58 (57-59) ; 28 (27.5-28.5) millimeters. 

Distribution and h^ibitat. — The general range of this large sub- 
species is along the bottoms of the lower San Pedro River Valley 
near Mammoth, and the bottoms of the Gila River Valley above 
the confluence of the Gila and San Pedro as far as Redrock, New 
Mex. The area is in the Lower Sonoran Zone from about 2,000 to 
3,500 feet or a little more in altitude. These pocket gophers are re- 
stricted rather closely to the fertile, alluvial lands along the rivers, 
overgrown in the natural state with a heavy stand of mesquite 
(Prosopis juliflo7^a) timber. 

Economic status.^— MMoh of the alluvial land along the San Pe- 
dro and Gila Rivers is cultivated under irrigation, alfalfa being 
one of the principal crops. The pocket gophers feed upon both the 
roots and tops of alfalfa and tend to increase in number where 
a bountiful supply of food is thus provided. Where they become 
very numerous the alfalfa crop is materially reduced. At Safford 
the farmers complained of damages to alfalfa, and also of breaks 
in irrigation ditchbanks caused by gopher tunneling. 

THOMOMYS BOTTAE ALBATUS Grinnell 

Whitish Pocket Gopher 
Thomomys alhatus Grinnell, Univ. Calif. Pub. Zool. 10: 172. June 7, 1912. 

Tijpe. — From west side of the Colorado River at old Hanlon 
Ranch near Pilot Knob, Imperial County, Calif. ; collected by Jo^ 
seph Dixon, May 7, 1910. 



20 NORTH AMERICAN FAUNA 59, FISH AND WILDLIFE SERVICE 

General chm-acters. — Distinguished by extremely pallid color- 
ation and rather large size. Upper parts nearly uniform pale pink- 
ish buff, in some specimens appearing almost white. Closely al- 
lied to phasm<i of the Tule Desert region, but larger; skull more 
massive. 

Measurements. — An adult male: Total length, 272; tail verte- 
brae, 100; hind foot, 35 millimeters. An adult female: 264; 91; 
34 millimeters. 

Distribution and habitat. — This nearly white subspecies inhab- 
its both sides of the lower Colorado River Valley from the vicin- 
ity of Yuma, at least, to points in the Delta. It burrows in soft 
alluvial soil, and local occurrence points to the transfer of colonies 
from one side to the other with the frequently changing channels 
of the river. Across the lowlands of the Delta, therefore, the gen- 
eral range of cdbatus appears to be the connecting link in a chain 
of subspecies of the Thomomys bottae group extending from the 
Pacific coast to the Rio Grande Valley in Texas. 

THOMOMYS BOTTAE DEPAUPERATUS Grinnell and Hill 

TiNAJAS Altas Pocket Gopher 

Thomomys perpalUdus depawperatus Grinnell and Hill, Jour. Mammal. 17 (1): 
4, February 17, 1936. 

Tyve. — From the east base of the Tina j as Altas Mountains, 7 
miles south of Raven Butte, Yuma County, Ariz, (altitude 1,150 
feet) ; collected by Annie M. Alexander, January 17, 1934. 

General characters. — Distinguished by pale pinkish buff colora- 
tion combined with small size. Color about as in the geographic 
neighbors albatus and phasma, but smaller than either (much 
smaller than albatus) ; skull weaker in structure, less angular 
than in phasma; zygomata more slender ; nasals relatively shorter. 

Measurements. — A representative female (from original de- 
scription) : Total length, 188; tail, 60; hind foot, 28 millimeters. 

Distribution xind habitat. — Known only from a small colony in 
the desert along the eastern base of the Tina j as Altas Mountains, 
a southern extension of the Gila Mountains, and about 4 to 5 miles 
north of the Mexican Boundary. It is reported to inhabit gravel- 
ly soil along the margins of washes, where the mesquite (Prosopis 
juliflora) and catsclaw {Acacia greggii) are among the dominant 
woody plants. 

THOMOMYS BOTTAE PHASMA Goldman 

Tule Desert Pocket Gopher 

Thomomys fulvus phasma Goldman, Biol. Soc. Wash. Proc. 46: 72, April 27, 1933. 

Type.— From 2 miles south of Tule Tank, Tule Desert, Yuma 



THE POCKET GOPHERS 21 

County, Ariz, (altitude 1,200 feet) ; collected by E. A. Goldman, 
December 8, 1913. 

General characters. — One of the palest of the known forms of 
the genus. Upper parts near pale pinkish buff, scarcely modified 
by dark-tipped hairs. Closely allied to ulbatus of the Colorado 
Delta region, but smaller ; skull less massive. 

Measurements. — An adult female : Total length, 199 ; tail verte- 
brae, 66 ; hind foot, 29 millimeters. 

Distribution and habitat. — The Tule Desert pocket gopher oc- 
curs irregularly so far as known in small, local colonies on the 
low Lower Sonoran plains from the vicinity of the type locality 
near the Mexican Boundary northwest to Wellton, Yuma County. 
Its habitat on the desert is in one of the most arid regions in North 
America. The burrows are found along washes and in open stands 
of desert vegetation, including the creosotebush (Covillea triden- 
tata), ironwood (Olneya tesota), paloverde (Cercidium torreya- 
num), and giant cactus {Carnegiea gigantea) . Numerous mounds 
of earth pushed out near together at about the same time are evi- 
dences of periodical activity, apparently following rains, which 
are infrequent in the region. Periodical excavation of earth on 
an extensive scale is a characteristic of pocket gophers in general 
that seems emphasized in this and some of the other desert forms. 
During the long intervals between rains the animals remain in the 
deeper underground workings, and there is little evidence of activ- 
ity on or near the surface. At such times tunnels near the surface 
are generally plugged for several feet, or show signs of disuse. 
To obtain specimens of the gophers may involve considerable la- 
bor, as under such conditions it may be necessary to dig a trench 
2 to 3 feet deep with a shovel in order to reach the used tunnels 
where traps can be set with excellent results. 

THOMOMYS BOTTAE ARIDICOLA Huey 

Gila Bend Pocket Gopher 

Thomomys bottae aridicola Huey, San Diego Soc. Nat. Hist. Trans. 8 (25): 
354, June 15, 1937. 

Type. — From 10 miles south of Gila Bend (or, exactly, on Ajo 
Railroad right-of-way, about 2 miles north of Black Gap), Mari- 
copa County, Ariz, (altitude 900 feet) ; collected by Laurence M. 
Huey, February 1, 1936. 

General characters. — A medium-sized, light buffy, desert sub- 
species. Similar to cervinu^ of the Salt River Valley, but smaller 
and color of upper parts shading toward cinnamon-buff instead 
of vinaceous-buff . 

Measurements. — Type, an adult female : Total length, 212 ; tail, 
63 ; hind foot, 29 millimeters. 



22 NORTH AMERICAN FAUNA 59, FISH AND WILDLIFE SERVICE 

Distribution and habitat. — The type and a topotype of this pock- 
et gopher came from close along the Ajo Railroad, 2 miles north oi 
Black Gap, and 10 miles south of Gila Bend, Maricopa County, 
Ariz. The animal appears to be one of the isolated forms that 
exist as local colonies in the wide expanse of Lower Sonoran des- 
ert. 

THOMOMYS BOTTAE GROWLERENSIS Huet 

Growler Valley Pockbtt Gopher | 

Thomomys hottae growlerensis Huey, San Diego Soc. Nat. Hist. Trans. 8 (25)- 
353, June 15, 1037. 

Type. — From 7 miles east of Papago Well, Pima County, Ariz, 
(or, exactly, along a well wooded desert wash on the southwestern 
side of a range of hills in the southern end of Growler Valley ; the 
Agua Dulce Mountains form the southern boundary of this local- 
ity and are not far distant) ; collected by Laurence M. Huey, 
March 16, 1937. 

General characters. — A desert form, closely allied to phnsTna 
of the Tule Desert to the west, but darker, deeper pinkish buff or 
yellowish in color. Much lighter colored in comparison with modi- 
cus of the Altar Valley to the east. 

Measurements.— Type, an adult male: Total length, 208; tail, 
62 ; hind foot, 30 millimeters. 

Distribution and habitat. — The known range of this denizen 
of the desert includes several localities from the southern end of 
Growler Valley and Quitobaquito north to Bates Well in Growler 
Pass, between the Growler Mountains and Bates Mountains, all 
in the Lower Sonoran Zone. The burrows are usually found in 
soft soil along gravelly wooded washes where the iron wood (01- 
neya Desoto) , mesquite (Prosopis), catsclaw (Acacia greggii), and 
paloverde (Cercidium torreyanum) are among the dominant 
woody plants. 

THOMOMYS BOTTAE COMOBABIENSIS Huey 

CoMOBABi Pocket Gopher 

Thomomys lottae comobabiensis Huey, San Diego Soc. Nat. Hist. Trans. 8' 
(25): 354, June 15, 1937. 

Type.— From 5 miles northwest of Sells, Pima County, Ariz, 
(altitude 2,400 feet) ; collected by Laurence M. Huey, March 22,, 
1937. 

General characters. — A medium-sized, distinctly brownish sub- 
species. Similar in color to modicum of the Altar Valley, but some- 
what smaller; skull with larger, more fully inflated, auditory 
bullae. 

Measurements.— The type, an adult female ; Total length, 215 ; 



THE POCKET GOPHERS 23 

tail, 70 ; hind foot, 28 millimeters. 

Distribution and habitat.— Known only from the type locality 
on the basal slope of the Comobabi Mountains. Like some of the 
other desert representatives of the genus, this form appears to be 
very local in distribution. More abundant material for study may 
show close alliance to modicus to which specimens from Sells are 
referred. 

THOMOMYS BOTTAK PUSILLUS Goldman 

Coyote Moxjntain Pocket Gopher 

Thomomys fulvus pusillus Goldman, Jour. Wash. Acad. Sci. 21 (17): 422, 
October 19, 1931. 

Type. — From Coyote Mountains, Pima County, Ariz, (altitude 
3,000 feet) ; collected by E. A. Goldman, September 4, 1915. 

General characters. — A small, rich ochraceous-tawny form with 
a slender, delicate skull and large, fully distended, auditory bullae. 
Mammae, pectoral two pairs, inguinal two pairs. Allied to modi- 
cus of the neighboring valleys, but much smaller; color more 
tawny. 

Measurements. — The type, an adult female : Total length, 201 ; 
tail vertebrae, 65 ; hind foot, 27.5 millimeters. 

Distribution and habitat. — As the name implies, this is a very 
small pocket gopher. It is known only from a spot where a little 
soil had accumulated at 3,000 feet altitude in the exceedingly 
rocky Coyote Mountains. It is associated with the catsclaw {Aca- 
cia greggii), mesquite {Prosopis juliflora), and other vegetation 
of the Lower Sonoran Zone. 

THOMOMYS BOTTAE GRAHAMENSIS Goldman 

Graham Mountains Pocket Gopher 

Thomomys fulvus grahamensis Goldman, Jour. Wash. Acad. Sci. 21 (17) : 420, 
October 19, 1931. 

Type. — From Graham Mountains (=Pinaleno Mountains), Gra- 
ham County, Ariz, (altitude 9,200 feet) ; collected by Ernest G. 
Holt, June 7, 1914. 

General characters. — A dark, high-mountain subspecies, resem- 
bling fulvus of the Mogollon Plateau region, but upper parts near 
ochraceous-buff instead of ochraceous-tawny, but as in fulvus 
rather heavily mixed with black ; skull narrower, with zygomata 
less widely spreading. About like collirius of the Chiricahua Moun- 
tains in color ; skull narrower, but interorbital region broader. 

Measurements. — An adult male: Total length, 231; tail verte- 
brae, 71 ; hind foot, 29 millimeters. An adult female : 228 ; 76 ; 28 
millimeters. 

696493°— 47— 4 



24 NORTH AMERICAN FAUNA 59, FISH AND WILDLIFE SERVICE 

Distribution and habitat. — This subspecies is known only from 
the forested upper slopes (6,100 to 9,200 feet altitude) of the 
Graham Mountains, which like other ranges of the general region, 
rise steeply, island-like, from arid surrounding plains and valleys. 
This pocket gopher burrows in soft soil in the Transition Zone, 
and is numerous in the grassy high mountain meadows bordered 
by firs (Pseudotsuga) and spruces (Picea) in the Canadian Zone 
along the crest of the range. Pocket gophers from Fort Grant at 
the west base are closely allied to grafmmensis, but are much paler 
and are referred to extenuatus. Along the arid, eastern basal 
slopes of the Graham Mountains pocket gophers are scarce or ab- 
sent, and grahamensis contrasts strongly with alienus, which is 
abundant along the Gila Valley. 

THOMOMYS BOTTAE EXTENUATUS Goldman 

Sulphur Springs Valley I'ocket Gopher 

Thomomys hottae extenuatus Goldman, Biol. Soc. Wash. Proc. 48: 149, Oc- 
tober 31, 1935. 

Type. — From Willcox, Cochise County, Ariz, (altitude 4,000 
feet) ; collected by Vernon Bailey, November 27, 1889. 

General characters. — This small cinnamon-buff or light tawny 
pocket gopher, with strongly decurved upper incisors and large 
auditory bullae, is allied to several neighboring forms from all 
of which it differs in combination of size, color, and skull struc- 
ture. Similarity in color of exte^iuatus to alieniLS of the lower el- 
evations along the San Pedro and Gila River Valleys is evident, 
but the smaller general size and more swollen auditory bullae of 
extenuatus are distinctive. Extenuutus is about the same in size 
as, and evidently related to, the neighboring high-mountain forms 
grahamensis, collinus, catalinae, and hueyi but differs from 
all in lighter color and cranial details. In size and color extenua- 
ttis closely approaches Thomomys baileyi mearnsi, and the two oc- 
cur together in places ; extenuxitus may be recognized by the high- 
er brain case and heavier dentition, and the upper incisors are 
more strongly recurved instead of projecting forward beyond the 
nasals as in mearnsi. 

. Measurements. — An adult male and an adult female topotype, 
respectively: Total length, 203, 198; tail, 67, 67; hind foot, 27.5, 
29 millimeters. 

Distribution and habitat. — The range of extenuatus embraces 
the desert plains and the basal mountain slopes bordering the Sul- 
phur Springs Valley, and adjoining valleys near the top of the 
Continental Divide, extending east into the San Simon Valley, 
north to Fort Grant, and west across the upper part of the San 



THE POCKET GOPHERS 25 

Pedro Valley to Oracle. The general area lies near the boundary 
between the Upper and Lower Sonoran Zones. In this region, as 
near Willcox, the ranges of extenuatus and meamsi, which are re- 
garded as representatives of distinct species, interdigitate, and 
careful examination of skulls may be necessary to distinguish 
them. 

THOMOMYS BOTTAE CATALINAE Goldman 

Santa Catalina Pocket Gophek 

Thomomys fvlrvs catalinae Goldman, Jour. "Wash. Acad. Sci. 21 (17): 419, 
October 19, 1931. 

Type. — From Summerhaven, Santa Catalina Mountains, Pima 
County, Ariz, (altitude 7,500 feet) ; collected by E. A. Goldman, 
August 6, 1923. 

General characters. — A small, notably dark-colored subspecies, 
the upper parts tawny, heavily mixed with black. Closely allied 
to hueyi of the Rincon Mountains, but darker in color, the upper 
parts more profusely mixed with black, and the under parts hav- 
ing a darker buff tone. Closely resembles collinus of the Chirica- 
hua Mountains in dark color, but skull flatter and narrower ; zygo- 
mata less widely spreading. 

Meusurements. — Average of four adult male topotypes: Total 
length, 211 (204-220) ; tail vertebrae, 64 (58-72) ; hind foot, 28.5 
(27.5-30) millimeters. Average of four adult female topotypes: 
202 (196-212) ; 59 (55-62) ; 25 (24-26) millimeters. 

Distribution and habitat. — This richly colored pocket gopher 
is restricted to the upper slopes of the Santa Catalina Mountains, 
where it burrows in the soft, gravelly soil in rather open stands of 
timber, largely yellow pines and oaks, in the Transition Zone at 
7,000 to 8,000 feet altitude. It also occurs, however, among Cana- 
dian Zone elements nearer the summit of Mount Lemmon. The 
oak belt along the northern basal slope of these mountains, at Or- 
acle, is inhabited by pocket gophers obviously closely related to 
the present form, but that in color are more like extenuattts, to 
which they are referred. The Santa Catalina Mountains are con- 
nected across a saddle or pass with the Rincon Mountains, the 
upper slopes of which are occupied by the closely related, but pal- 
er, form hueyi. Mountainous masses of nearly solid rock tend to 
separate the two, and the well marked, small, local form 'parvulus 
is interposed in a series of shallow, rock-bound depressions filled 
with stony soil in the pass. Pocket gophers appear to be absent 
in a broad, arid belt along the southern side of the Santa Catalina 
Mountains, and no evidence of intergradation with modicus, 
which is common along the Santa Cruz River near Tucson, is found. 



26 NORTH AMERICAN FAUNA 59, FISH AND WILDLIFE SERVICE 

THOMOMYS BOTTAE PARVULUS Goldman 

Intbrmountain Pocket Gopheb 

Thomomys hottae parvtdus Goldman, Jour. Wash. Acad. Sci. 28 (7) : 339, 
July l'5. 1938. 

Type. — From the pass between the Santa Catalina and Rincon 
Mountains, Pima County, Ariz, (altitude 4,500 feet) ; collected by 
Luther C. Goldman, June 5, 1937. 

General characters. — A very small cinnamon or tawny subspe- 
cies; mammae, pectoral two pairs, inguinal two pairs. Allied to 
catalinue of the upper slopes of the closely adjoining Santa Cata- 
lina Mountains, and to hueyi of similar proximity in the Rincon 
Mountains, but much smaller than either, color lighter, more in- 
clining toward tawny. 

Measurements. — Two adult male topotypes, respectively: Total 
length, 203, 211; tail, 60, 57; hind foot, 27, 25 millimeters. Two 
adult female topotypes, respectively: 190, 188; 56, 55; 25, 25 
millimeters. 

Distribution and habitat. — These diminutive pocket gophers are 
apparently restricted in range to gravelly pockets in the granitic 
formation in the pass between the Santa Catalina and Rincon 
Mountains. Here they are numerous, although the soil is so thin 
and scanty that gopher excavations consist largely of pebbles. 
Massive rock exposures in the vicinity may be effective barriers 
limiting distribution. The area, at 4,000 to 4,500 feet altitude, is 
near the boundary between the Upper Sonoran and Lower Sonoc 
ran Zones, as shown by overlapping floral elements. The Upper 
Sonoran Zone is represented by the lower edge of the oaks (Quer- 
cus arizonica and Q. emoryi), but along these is a thin stand 
of mesquite (Prosopis juliflora), catsclaw {Acacia greggii), and 
desertwillow (Chilopsis linearis). 

THOMOMYS BOTTAE HUEYI Goldman 

KiNcoN Mountains Pocket Gopher 

Thomomyfi hottae hueyi Goldman, Jour. Wash. Acad. Sci. 28 (7) : 340, July 
15, 1939. 

Type. — From Spud Rock Ranger Station, Rincon Mountains, 
Pima County, Ariz, (altitude 7,400 feet) ; collected by Laurence 
M. Huey, June 17, 1932. 

General characters. — A small, cinnamon subspecies, closely al- 
lied to catalinae of the adjoining Santa Catalina Mountains, but 
upper parts less profusely mixed with black, and under parts a 
lighter buff tone. Larger than its near neighbor parvulus, which 
occupies the pass between the Rincon Mountains and Santa Cata- 
lina Mountains ; color darker, less tawny. 



THE POCKET GOPHERS 27 

Measurements. — Two adult male topotypes, respectively: Total 
length, 220, 220 ; tail, 62, 66 ; hind foot, 30, 29 millimeters. Two 
adult female topotypes, respectively : 198, 196 ; 60, 60 ; 26, 27 mil- 
limeters. 

Distribution and habitat. — In the Rincon Mountains hueyi is 
known from the Transition Zone near the top at 7,400 to 7,900 
feet altitude. Pocket gophers that appear to be referable to this 
subspecies are also found at about 7,000 feet altitude in Ramsay 
Canyon and in the head of Miller Canyon in the Huachuca Moun- 
tains. The same subspecies may inhabit the Whetstone Mountains 
and elevated plains between the two localities mentioned. In the 
Rincon Mountains massive rock exposures apparently separate the 
habitat of hueyi from that of parvulus in the pass connecting this 
range with the Santa Catalina Mountains. 

THOMOMYS BOTTAE COLLINUS (Joi.dman 
Chiricahua Mountain Pocket Gophek 

Thomomj/s fulvus collinus Goldman, Jour. Wash. Acad. Sci. 21 (17): 421, Oc- 
tober 19, 1931. 

Thotnomys uvibrinus chiricahuae Nelson and Goldman, Jour. Mammal. 15 (2): 
117, May 15, 1934. Type from Pinery Canyon, Chiricahua Mountains, 
Ariz, (altitude 7,500 feet). 

Type. — From Fly Park, Chiricahua Mountains, Cochise County, 
Ariz, (altitude 9,000 feet) ; collected by A. K. Fisher, June 10, 
1894. 

General characters. — A dark, high-mountain subspecies, resem- 
bling catalinae of the Santa Catalina Mountains and grahamen- 
sis of the Graham Mountains, but skull differing in more widely 
spreading zygomata and other details. Pectoral mammae, normal- 
ly two pairs, but they may vary to one pair. 

Measurements. — Average of five adult males : Total length, 213 
(207-222) ; tail vertebrae, 58 (50-60) ; hind foot, 28.3 (27-30) mil- 
limeters. 

Distribution und habitat. — Like some other high-mountain forms 
of the general region, this pocket gopher is restricted to a single 
mountain range, and in this case the Chiricahua Mountains. It 
ranges from the mouths of Cave and Turkey Creeks at about 5,000 
feet altitude on the eastern and western sides, respectively, to the 
extreme summit of Fly Peak, at 9,700 feet, but is most abundant 
in the soft, dark soil of mountain meadows, such as Rustler Park, 
at 8,500 feet altitude in the Canadian Zone. Specimens from the 
lower elevations are somewhat paler and approach extenuatus of 
the Sulphur Springs Valley region. A few specimens from 7,500 
feet in Pinery Canyon are small, and as only one pair of pectoral 
mammae was found, they were described as Thomomys umbrinus 



28 NORTH AMERICAN FAUNA 59, FISH AND WILDLIFE SERVICE 

chiricahuae, but the number of these mammae proves to vary from 
normal in some individuals. Additional specimens of collinus in- 
dicate that the characters ascribed to chiricahuae are within the 
range of individual variation in that form. 

THOMOMYS BOTTAE MODICUS Goldman 

Altar Valley Pockbtt Gopheb 

Thomomys fulvus modicus Goldman, Jour. Wash. Acad. Scl. 21 (17): 418, 
October 19, 1931. 

Type. — From La Osa (near Mexican Boundary), southern end 
of Altar Valley, Pima County, Ariz. ; collected by E. A. Mearns 
and F. X. Holzner, December 14, 1893. 

General characters. — A dark-colored subspecies of medium size. 
Closely allied to cervinus of the Salt River Valley, but smaller ; , 
upper parts near wood brown or cinnamon, varying to rich tawny ■ 
instead of vinaceous-buff or fawn color; skull more slender. 

Measurements. — Average of four adult male topotypes: Total 
length, 214 (204-222) ; tail vertebrae, 67 (55-75) ; hind foot, 28 
(25.5-30) millimeters. Average of six female topotypes : 208 (198- 
224) ; 69 (60-74) ; 27.5 (26.5-28) millimeters. ; 

Distribution and habitat. — The general range of this subspecies 
includes the Altar Valley, the upper part of the Santa Cruz River 
Valley, and neighboring valleys and desert plains as far west as 
Sells, and east to Fort Huachuca. The altitudinal limits are from 
about 2,500 to about 4,500 feet, mainly in the upper part of the 
Lower Sonoran Zone. The vegetation consists prominently of 
mesquite (Prosopis juliflora), catsclaw {Acacia greggii), palo- 
verde (Cercidium torreyanum) , creosotebush {Covillea triden- 
tata), and many cactuses. The gophers range up in places, how- 
ever, along the basal slopes of the mountains into the lower edge of 
the oaks (Quercus emoryi and Quercus arizonica), marking the 
Upper Sonoran Zone. Distribution, as in many other forms of 
the group, is not continuous. These pocket gophers exhibit a pref- 
erence for the softer soils along streams and dry washes, which 
may be separated by many miles of unoccupied desert. Along the 
lower slopes of the Santa Rita and Huachuca Mountains, modicus 
meets the range of Thomomys umbrinus proximus, regarded as a 
representative of a distinct species. 

THOMOMYS ALEXANDRAE Goldman 

Navajo Pocket Gopher 

Thomomys alexandrae Goldman, Jour. Wash. Acad. Sci. 23 (10): 464, Oc- 
tober 15, 1933. , 

Type. — From 5 miles southeast of Rainbow Lodge, near Navajo 



THE POCKET GOPHERS . 29 

Mountain, Coconino County, Ariz, (altitude 6,200 feet) ; collected 
by E. A. Goldman, June 16, 1933. 

General characters. — An apparently distinct species of the 
Tho7nomys hottue group, allied to aureus of the adjoining desert 
region, but decidedly smaller; color much duller, near cinnamon- 
buff instead of rich ochraceous-tawny ; skull flatter and slen- 
derer, with more widely separated temporal ridges. 

MeasureTYients. — An adult male : Total length, 210 ; tail, 60 ; 
hind foot, 28 millimeters. An adult female: 215; 70; 27 milli- 
meters. 

Distribution and habitat. — The Navajo pocket gopher is com- 
mon in places on the nearly flat sagebrush-covered mesa at 6,200 
feet altitude south of Navajo Mountain. It has also been recorded 
by Benson (Univ. Calif. Pub. Zool. 40: 449, Dec. 31, 1935) from 
near Soldier Spring at 8,600 feet on Navajo Mountain, just across 
the Utah line. The species appears to be isolated in a somewhat 
triangular area between the precipitous walls of Navajo and Pin- 
to Creek canyons which diverge to the Colorado River. Along 
the narrow divide between the upper courses of these creeks 
the solid bedrock formation is nearly bare of soil for miles. None 
of the characteristic gopher mounds was seen, and this barren 
ridge, as well as the canyons, may have served as an effective bar- 
rier isolating the habitat of alexandrae for thousands of years. 

THOMOMYS SUBOLES Goij)MAn 

Searchlight Ferry Pocket Gopheb 

Thomonivft fidvus suboles Goldman, Biol. Soc. Wash. Proc. 41: 203, December 
18, 1928. 

Type. — From Old Searchlight Ferry, Colorado River, north- 
west of Kingman, Mohave County, Ariz, (altitude 600 feet) ; col- 
lected by Luther C. Goldman, September 26, 1923. 

General characters. — A small, light-colored species, allied to 
desertorum of the Detrital Valley, but more ochraceous-tawny; 
skull more angular, narrower, but heavier in detail; maxillary 
arms of zygomata much broader, with acutely projecting lateral 

angles; auditory bullae more compressed laterally, less rounded. 

Measurements. — An adult male: Total length, 227; tail verte- 
brae, 75; hind foot, 30 millimeters. An adult female: 194; 62; 
26.5 millimeters. 

Distribution and habitat. — In an embayment of the escarpment 
flanking the Colorado River, near Old Searchlight Ferry, above 
Pyramid Canyon, and northwest of Kingman, Ariz., are alluvial 
bottoms extending for several miles until interrupted by cliffs 



30 NORTH.AMERICAN FAUNA 59, FISH AND WILDLIFE SERVICE 

rising abruptly from the water. The bottoms, consisting of soft, 
sandy soil, are overgrown with mesquite {Prosopis), catsclaw 
{Acacia greggii), and other Lower Sonoran Zone vegetation. The 
pocket gophers have become isolated here in a narrow belt be- 
tween the river and the escarpment, which arises steeply to the 
crest of a rocky ridge at about 3,500 feet altitude. These pocket 
gophers have evidently found their restricted habitat congenial 
as attested by their numbers. The distribution and habitat of sub- 
oles in relation to other species have been discussed in detail by 
Grinnell and Hill (Jour. Mammal. 17 (1) : 7-10, Feb. 17, 1936), 
who refer to the occurrence of a quite different subspecies, Tho- 
momys bottue centralis, on the opposite side of the Colorado River, 
although the two forms live under similar conditions of soil, cli- 
mate, and food. 

THOMOMYS MURALIS Goldman 

Grand Canyon Pocket Gopheb 

Thommnys muralis Goldman, Jour. Wash. Acad. Sci. 26 (3) : 112, March 15, 
1936. 

Type. — From lower end of Prospect Valley, Grand Canyon, 
Hualpai Indian Reservation, Ariz, (altitude 4,500 feet) ; collected 
by E. A. Goldman, October 3, 1913. 

General characters. — A diminutive ochraceous buffy or some- 
what tawny species, similar in size and in color to, and closely 
resembling, desertorum, but cranial characters distinctive; brain 
case more rounded and inflated, the basicranial region tending 
to bulge more prominently posteriorly; frontal region broader; 
upper incisors more strongly recurved. Differs from fulvus in 
lighter color, and smaller size, the skull more delicate in structure 
and exhibiting a departure in about the same details as from des- 
ertorum. 

Measurements. — Type, an adult male: Total length, 194; tail, 
64; hind foot, 26 millimeters. Two adult female topotypes, re- 
spectively : 182, 190 ; 57, 56 ; 24.5, 25.5 millimeters. 

Distribution and habitat. — Isolated on terraces along the inner 
gorge below the outer rim in Prospect Valley, a lateral pocket 
within the Grand Canyon, near the eastern end of the Hualpai 
Indian Reservation. The geographic isolation of muralis in the 
Grand Canyon appears to be complete, and characters presented 
suggest full specific rank. In places it was found inhabiting strips 
of soil on ledges only a few feet wide, bounded above and below 
by vertical cliffs hundreds of feet high. ZoTial range : Upper So- 
noran. 



THE POCKET GOPHERS 31 

THOMOMYS HARQUAHALAE Grinnexl and Bill 

Ranbgras Plain Pocket Gophejb 

Thomomys harquahalae Grinnell aud Hill, Jour. Mammal. 17 (1) : 7, Feb- 
ruary 17, 1936. 

Type. — From Ranegras Plain, 10 miles west of Hope, Yuma 
County, Ariz, (altitude about 1,250 feet) ; collected by Louise 
Kellogg, February 27, 1934. 

General characters. — A large pinkish buff species; skull with 
widely spreading zygomata and anteriorly projecting upper inci- 
sors. Not very closely allied to any other known form, and there- 
fore accorded full specific status. Similar to chrysonotus of the 
Colorado River Valley, but paler (cinnamon-buff in chrysmiotus) ; 
zygomata wider ; upper incisors strongly procumbent instead of 
strongly decurved; auditory bullae shorter, more rounded, less 
projecting below plane of basioccipital. 

Measurements. — An adult male: Total length, 236; tail verte- 
brae, 69; hind foot, 31 millimeters. An adult female: 210; 62; 
29.5 millimeters. 

Distribution and habitat. — Known only from a colony along the 
highway where it crosses the lowest part of the broad open grassy 
Ranegras Plain west of Hope. The soil in which these pocket 
gophers burrow is compact in texture and difficult to perforate, 
suggesting a possible special use for the forward-curving incisors. 
Many mounds, marking excavations in the highway embankment, 
where food appears to be scanty, indicate that the animals wel- 
come a change from the hard soil of their natural habitat. 

SUBSPECIES OF THOMOMYS BAILEYI GROUP 

THOMOMYS BAILEYI MEARNSI Bailey 
Me.\rns Pocket Gopher 
Thomomys mearnsi Bailey, Biol. Soc. Wash. Proc. 27: 117, July 10. 1914. 

Ti/pe. — From Gray's Ranch, Animas Valley, southwest corner 
of Grant County, N. Mex. ; collected by E. A. Goldman, August 
10, 1908. 

General characters. — This subspecies, a near relative of typical 
haileyi of western Texas, requires close comparison with extenu- 
atus, a form of the bottae group in Arizona, as the ranges of the 
two meet or interdigitate. In cinnamon-buflf or light tawny color- 
ation meanisi resembles extenuatus, and for differential charac- 
ters recourse must be had to the skull, which is very similar in 
general, but relatively broader with a somewhat lower, flatter 
brain case, a combination of characters apparently indicating 
group relationship. In nnearnsi the dentition is lighter, the upper 



32 NORTH AMERICAN FAUNA 59, FISH AND WILDLIFE SERVICE 

incisors narrower, more projecting forward beyond the nasals. 

Measurements. — Type, adult male, and an adult female topo- 
type, respectively: Total length, 220, 201; tail, 67, 65; hind foot, 
31, 29 millimeters. 

Distribution and habitat. — The range of mearnsi extends from 
the Animas Valley, southwestern New Mexico, into the elevated 
plains region mainly in the lower part of the Upper Sonoran Zone 
of southeastern Arizona. Specimens were obtained in soft moist 
ground along a small stream at San Bernardino, near the Mexican 
Boundary and on the open plain near Willcox in the Sulphur 
Springs Valley. While mearnsi and extenvxitus appear to be typ- 
ically quite distinct, their general ranges meet or interdigitate 
and specimens from San Simon Valley suggest the possibility of 
hybridism. The exact habitat relations of the two forms remain, 
therefore, to be determined. 

SUBSPECIES OF THOMOMYS TALPOIDES GROUP 

THOMOMYS TALPOIDES KAIBABENSIS Goldman 

Kaibab Plateau Pocket Gopher 

Thomomys fossor kaibabensis Goldman, Jour. Wash. Acad. Sci. 28 (7): 333, 
July 15, 1938. 

Type. — From DeMotte Park, Kaibab Plateau, Ariz, (altitude 
9,000 feet) ; collected by Luther C. Goldman, September 10, 1937. 

General characters. — Resembles fossor of southwestern Colo- 
rado, but larger, less ruf escent ; skull with more widely spreading 
zygomata; interparietal smaller; auditory bullae larger; upper 
incisors broader, less recurved. 

Measurements. — An adult male and an adult female topotype, 
respectively: Total length, 238, 230; tail, 58, 77; hind foot, 31, 30 
millimeters. 

Distribution and habitat. — The Kaibab Plateau pocket gopher 
appears to be restricted to the higher parts of the well forested 
Kaibab Plateau, which rises island-like from arid plains or broad 
valleys toward the north and presents a sheer face alonge the 
north side of the deepest part of the Grand Canyon. This pocket 
gopher occurs irregularly in local colonies mainly in the Canadian 
Zone at 8,500 to 9,000 feet altitude. These animals are numerous 
in the soft soil in DeMotte Park near the top of the plateau, an 
open grassy meadow several miles in length, bordered by fir, 
spruce, and aspen forest. Specimens have also been taken near 
the northern rim of the Grand Canyon. 

General habits. — In DeMotte Park, on the Kaibab Plateau, when 
the deep snow of winter melts away, many lines of earth 5 to 10 



THE POCKET GOPHERS 33 

feet in length, with branches, are revealed radiating from the en- 
trances to the burrows of the gophers. These cylindrical dumps, 
in addition to the usual mounds, are evidence of burrowing opera- 
tions late in winter that seem to be followed in spring by a period 
of comparative inactivity. 

THOMOMYS TALPOIDES FOSSOR Allen 
Rocky Mountain Pocket Gopher 
Thomomys fossor Allen, Amer. Mus. Nat. Hist. BaiU. 5: 51, April 28, 1893. 

Type.— From Florida, La Plata County, Colo, (altitude 7,200 
feet) ; collected by Charles P. Rowley, June 25, 1892. 

General characters. — Small pocket gophers, characterized by the 
chestnut brown overtone of the top of head and back and con- 
spicuous black ear patches; skull (compared with bottae group) 
narrow, with widely separated temporal ridges ; interparietal tri- 
angular; zygomata depressed posteriorly, the jugal sloping up- 
ward to join maxilla anteriorly ; auditory meatus large and promi- 
nent. 

Measurements. — An adult male and an adult female, respec- 
tively, from the Lukachukai Mountains: Total length, 212, 210; 
tail, 70, 60 ; hind foot, 30, 28 millimeters. 

Distribution and habitat.— The Rocky Mountain pocket gopher 
is a denizen of the upper slopes of high mountains. In Arizona 
fossor is known only from about 8,000 to 9,000 feet altitude, main- 
ly in the Canadian Zone, near the tops of the Tunitcha and Luka- 
chukai Mountains in the northeastern corner of the State. Here 
its range seems to meet that of Thomomys bottue peramplus, 
which extends from the lower slopes upward to about 8,000 feet, 
but the two forms are not known to occupy the same local terrain. 

SUBSPECIES OF THOMOMYS UMBRINUS GROUP 

THOMOMYS UMBRINUS PROXIMUS Burt and CAMPBPa.L 

Arivaca Pocket Gopher 

ThomotmiH hurti pro.rhuiifi Burt ;iiid Campbell. ,Jour. Mainninl. 15 (2) : 151, 
May 15, 1934. 

Type. — From Old Parker Ranch (Pickett's Ranch on U. S. 
Geological Survey topographic map, Patagonia Quadrangle, edi- 
tion of August 1905) , altitude 4,800 feet, west slope of Santa Rita 
Mountains, Pima County, Ariz.; collected by W. H. Burt, June 
9, 1931. 

General characters. — A small, pale russet-colored form, similar 
to Thomomys umbrinus burti of the upper slopes of the Santa Rita 
Mountains. Some specimens are not very unlike Thomomys bottae 
modicus of the neighboring plains, representing a distinct species ; 



34 NORTH AMERICAN FAUNA 59, FISH AND WILDLIFE SERVICE 

pectoral mammae, sometimes two pairs, as in modicus; skull mort 
slender, with narrower nasals and smaller auditory bullae. Simi- 
lar in size to burti, but upper parts paler, less deep russet, with a 
less well defined, less uniformly darkened median dorsal area: 
skull very similar. 

Measurements. — Type, an adult female: Total length, 193; 
tail, 61 ; hind foot, 25 millimeters. Two adult males, respectively, 
from Fort Huachuca : 200, 195 ; 59, 54 ; 27, 26 millimeters. 

Distribution and habitat. — ^A few specimens have been taken 
at the type locality on the west slope of the Santa Rita Mountains, 
at the Empire Ranch, east of these mountains, at Fort Huachuca 
and at Arivaca. At Arivaca, W. P. Taylor collected two specimens 
on the same day, one of which is referred to proximus and the 
other to typical modicus, a representative of a species regarded 
as distinct. Of 20 specimens from the Empire Ranch, 19 are nearly 
typical modicum, but one is referred to proximus. This single indi- 
vidual was also obtained by Taylor on the same day and at the 
same recorded altitude (4,632 feet) as an example of modicus. 
The occurrence of two species of pocket gophers regarded as dis- 
tinct in such close proximity is unusual. Zonal range : Upper Sono- 
ran, as indicated by open stands of oaks {Quercus emoryi and( 
Quercu^ arizonica) ; altitude, 4,500 to 4,800 feet. 

THOMOMYS UMBRINUS BURTI Huey 

Santa Rita Mountain Pocket GoPHEa; 

Tliomoviys hurti Huey, San Diego Soc. Nat. Hist. Trans. 7 (15): 158, July 
28, 1932. 

Type. — From Madera Canyon, Santa Rita Mountains, Santa i 
Cruz County, Ariz, (altitude 6,000 feet) ; collected by W. H. Burt, 
May 29, 1931. 

Gene7'al characters. — A small, dark subspecies, with a coloration 
unusual except in closely allied forms ; upper parts between cinna- 
mon and cinnamon-brown or russet, becoming uniformly blackish 
along the moderately broad, well defined median area from top of 
head to rump; skull small, brain case smoothly rounded; nasals 
^wedge-shaped, emarginate posteriorly; auditory bullae small; 
mammae, pectoral one pair, inguinal two pairs. Closely allied to 
inter-medius of the upper slopes of the Huachuca Mountains ; color 
slightly paler, dentition heavier. 

Measurements. — An adult male and an adult female topotype, 
respectively : Total length, 217, 200 ; tail, 60, 61 ; hind foot, 27.5, 
26 millimeters. 

Distribution and habitat. — Restricted to the Santa Rita Moun- 
tains, southern Arizona. Zonal range : from 4,500 feet altitude in 



THE POCKET GOPHERS 85 

:;he Upper Sonoran Zone near the mouth of Madera Canyon up to 
8,000 feet in the Transition Zone near the summit. The burrows 
are located in the softer soil, usually in small open meadows. 

THOMOMYS UMBRINUS QUERCINUS Hurt and Campbell 

Pa.iarito Mot^ntain PocKErr (Jophkr 

Thomonnis Intrtl qiirrchms Burt and Cjinii»l)(>Il, Jour. Mainmal. 15 (2) : 150, 
May 15, 1934. 

Type. — From Pena Blanca Spring, Pajarito Mountains, Ariz. 
(altitude 4,500 feet, near Mexican Boundary, north of Monu- 
ment 128) ; collected by Berry Campbell, July 15, 1933. 

General characters. — A small, cinnamon subspecies ; mammae, 
pectoral one pair, inguinal two pairs. Closely allied to proximus, 
but still smaller ; coloration of the same pattern and general tones ; 
skull smaller, more delicate in structure; nasals slightly shorter, 
reaching anterior plane of zygomata. 

Measurements. — An adult male topotype : Total length, 198 ; 
tail, 60 ; hind foot, 27 millimeters. Type, female, and an adult fe- 
male topotype, respectively: 193, 182; 61, 56; 25, 25 millimeters. 

Distribution and habitat. — The known range of this close rela- 
tive of proximus is from 4,500 feet altitude at the type locality 
in the Pajarito Mountains to about 6,000 feet in the pass over the 
summit of the Patagonia Mountains. Both localities are in the 
oak belt in the Upper Sonoran Zone. At Pena Blanca Spring 
burrows indicating a small colony were found in gravelly soil 
along a broad wash. At the time of the writer's visit in June the 
ground was very dry, no fresh gopher excavations were in evi- 
dence, and specimens were difficult to obtain. In the pass over the 
Patagonia Mountains, where the slopes are steep and brush-cover- 
ed, considerable search failed to reveal the excavations of any 
gophers except those of the single individual taken. 

THOMOMYS UMBRINUS INTERMEDIUS Mearns 

HuACHUcA Mountain I'ocket Gopher 

Thomomys fulvus intermedius Mearns, U. S. Nat. Mus. Free. 10: 719, July 
30, 1897. 

Type. — From summit of the Huachuca Mountains, southern 
Arizona (altitude 9,000 feet) ; collected by F. X. Holzner, Sep- 
tember 6, 1893. 

General characters. — A small, dark subspecies, closely allied 
to burti of the Santa Rita Mountains ; color slightly darker ; skull 
very similar, but nasals longer; dentition lighter. 

Measurements. — Type: Total length, 200; tail, 66; hind foot, 
26 millimeters. 



36 NORTH AMERICAN FAUNA 59, FISH AND WILDLIFE SERVICE 

Distribution and hubitat.—The type, from 9,000 feet altitudJ 
near the summit of the Huachuca Mountains, is the only speci-^ 
men examined and definitely assigned to this form. Specimensi 
from Fort Huachuca, near the north base of the mountains, arei 
referred to proximiis. 

4 



THE POCKET GOPHERS 



INDEX 



37 



[Principal pag-e references to a spec 

absonns, Thomomys l)ottae, '.i. 8, 10. 

Thomomys perpdJlidii.s, 10. 
nlhfttiis, Thoi)iomi/s, 19. 

Thomomys bottae, 2, 3, 16, 18, 19, 
20, 21. 
alexandrae, Thomomys, 3, 28. 
alienus, Thomomys hottae, 3 19, 24. 
apache, Thomomys bottae, 12. 
aridicohi, Thomomys bottae, 3, 21. 
Arivaca pocket gopher, 33. 
aureus, Thomomys, 11. 

Thomomys bottae, 3, 11, 12, 14, 20. 

baileyi, Thomomys, 1, 4, 5. 

Big Saudy River pocket gopher, 14. 

bottae, Thomomys, 1, 2, 6. 

bnrti, Thomomys, 34. 

Thomomys umbrinus, 5, 33, 34. 

catalinae, Thomomys bottae, 3, 24, 25, 
26, 27. 

Thomomys fulvus, 25. 
centralis, Thomomys bottae, 7, 30. 
cervhius, Thomomys, 17. 

Thomomys bottae, 3, 17, 21, 28. 
Characters, 2. 
Chiricahiia Mountain pocket gopher. 

27. 
chiricahuae, Thomomys iinihrimis. 27. 
chrysonotns, Thomomys, 15. 

Thomomys bottae", 3, 15, 31. 
coUinus, Thomomys bottae, 3, 23, 24, 27. 

Thomomys fulvus, 27. 
Comobabl pocket gopher, 22. 
comobableusis, Thomomys bottae, 3, 22. 
Coyote Mountain pocket gopher, 23. 

depauperatus, Thomomys liottae, 3, 20. 

Thomomys perpaUidus. 20. 
desertorum, Thomomys, 12. 

Thomomys bottae, 3, 12, 14, 16, 29, 
30. 
desitus, Thomomys bottae, 3, 12, 14, l;!, 

17. 
Detrital Valley pocket gopher, 12. 

Economic status, 6. 

extenuatus, Thomomys bottae, 3, 24, 
25, 27, 31, 32. 

Fawn-colored pocket gopher, 17. 
flavidns, Thomomys fulvus. 15. 
fossor. Thomomys. 33. 

Thomomys "talpoides, 5, 12, 33. 
Fulvous pocket gopher, 13. 
fulvus. Gcomys. 13. 

Thomomys bottae, 3, 11, 13, 14, 15, 
17, 23, 30. 

Qeomys fulvus, 13. 

Gila Bend pocket gopher, 21. 

Oolden pocket gopher, 15. 

Uraham Mountains pocket gopher, 23. 



ies in Itoldfaoe; synonyms in ifalir] 

grahamensis, Thomomys bottae, 3, 23, 
24, 27. 

Thomomys fulvus. 23. 
Grand Canyon pocket gopher, 30. 
Group, Thomomys baileyi, 1, 4, 5 (dis- 
trib. map), 31. 

Thomomys bottae, 1, 2, 3 (distrib. 

map), 7. 
Thomomys talpoides, 1, 5 (disliib 

map), 32. 
Thomomvs umbrinus, 1, 5 (distrib, 
map), 6, 33. 
Growler Valley pocket gopher, 22. 
growlerensis, Thomomys l)ottae, 3, 22. 

Harquahala Mountain pocket gopher, 

16. 
harquahalae, Thomomys, 3, 31. 
Hassayampa Valley pocket gopher, 16. 
House Rock Valley pocket gopher, 10. 
Huachuca Mountain pocket gopher, 35 
Hualpal Mountains pocket gopher, 14. 
hualpaiensis, Thomomys bottae, 3, 14. 
hueyl, Thomomys bottae, 3, 24, 26. 

intermrdius, Thomomys fulvus. 35. 

Thomomys umbrinus. 5, 34, 35. 
Intermountain pocket gopher, 26. 

Kaibab Plateau pocket gopher, 32. 
kaihabrnsis. Thomomys fossor. 32. 
Thomomys talpoides, 5, 32. 

Infirostris, Thom,om,ys, 11. 

Mearns pocket gopher, 31. 
mrnrusi, Thomom,ys, 31. 

Thomomys baileyi, 5, 24, 25, 31. 
modicus, Thomomvs bottae, 3, 18, 22, 
23, 25. 28, 33, 34. 

Tho-momys fulvus, 28. 
Mount Trumbull pocket gopher, 9. 
muralis, Thomomys, 3, 4, 30. 
mutabilis, Thomomys bottae, 3, 14, 15, 
17, 18, 19. 

nasutus, Thomomys "bottae. 13. 
Xava.io pocket gopher, 28. 
nicholi, Thomomys bottae, 3, 7. 9. 

operosus, Thomomys bottae, 3, 15. 

Painted Desert pocket gopher, 11. 
Pa.1arito Mountain pocket gopher, 34. 
parvulus, Thomomys bottae, 3. 26. 27, 
patulus, Thomomys bottae, 3, 15, 16. 
Peeples Valley pocket gopher, 15. 
peramplus, Thomomys bottae, 3, 11, 12, 
33. 

Thomomys fuhrus, 12. 
phasma, Thomomys bottae, 3, 20, 22. 

Thomomys fulvus, 20. 
Phoenix pocket gopher, 17. 
I'inal Mountains pocket gopher, 18. 



38 NORTH AMERICAN FAUNA 59, FISH AND WILDLIFE SERVICE 



pinalensis, Thomomys bottae. 3, 18. 
planirostris, Tliomomys bottae, 3, 8, 9 
10. 

Thomomys perpallidus, 8. 
Pocket gopher, Arivaca, 33. 

Big Sandy River, 14. 

Chirieahua Mountain, 27. 

Comobabi, 22. 

Coyote Mountain, 23. 

Detrital Valley, 12. 

fawn -colored, 17. 

fulvous, 13. 

Gila Bend, 21. 

golden, 15. 

Graham Mountains, 23. 

Grand Canyon, 30. 

Growler Valley, 22. 

Harquahala IMountain, 16. 

Hassayampa Valley, 16. 

House Rock Valley, 10. 

Huachuca Mountain, 35. 

Hualpai Mountains, 14. 

intermountain, 26. 

Kaibab Plateau, 32. 

Mearns, 31. 

Mount Trumbull, 9, 

Navajo, 28. 

Painted Desert, 11. 

Pajarito Mountain, 35. 

Peeples Valley, 15. 

Phoenix, 17. 

Pinal Mountains, 18. 

Ranegras Plain, 31. 

Rincon Mountains, 26. 

Rocky Mountain, 33. 

Santa Catalina, 25. 

Santa Rita Mountain. 34. 

Searchlight Ferry, 29. 

Shivwits Plateau, 7. 

Sulphur Springs Valley, 24. 

Tinajas Altas, 20. 

Tule Desert, 20. 

Tunitcha Mountain, 12. 

Upper Gila Valley, 19. 

Verde Valley, 17. 

Virgin Valley. 7. 

whitish, 19. 

Zion Park, 8. 
proximns, Thomomi/s hurti, 33. 

Thomomys umitrinus, 5, 33. 34, 35. 
pusillus, Thomomys bottae, 3, 23. 

Thomomys fillvus, 23. 

qverichms, Thomomys iurti, 3n. 
Thomomys umbrinus, 5, 35. 

Ranegras Plain pocket gopher, 31. 
Rincon Mountains pocket gopher, 26 
nparius, Thomomys bottae, 16. 
Rocky Mountain pocket gopher, 32. 

Santa Catalina pocket gopher, 25 
^34^ ^ita Mountain pocket gopher, 

Searchlight Ferry pocket gopher, 29 
fehivwits Plateau pocket gopher 7 
suboles, Thomomys, 3, 29. 



suboles, Thomomy.s — Continued. 

Thomomys fiilvus, 29. 
subsimilis, Thomomys bottae, 3, 16. 

Thomomys fiilvus, 16. 
Sulphur Springs Valley pocket gophei 

talpoides, Thomomys, 1, 5, 6. 
Thomomys alhatus, 19. 4 

alexandrae, 3, 28. i 

aureus, 11. 

baileyi, 1, 4, 5. 

baileyi mearnsi, 5, 24, 25, 31. 

bottae, 1, 3 (distrib. map), 6. 

bottae absonus, 3, 8 10. 

bottae albatus, 2, 16, 18. 19. 20, 21 

bottae alienus, 3, 19, 24. 

bottae apache, 12. 

bottae aridicola, 3, 21. 

bottae aureus, 3, 11, 12, 14, 29. 

bottae catalinae, 3, 24, 25, 26, 27. 

bottae centralis, 7, 30. 

bottae cervinus, 3, 17, 21, 28. 

bottae chrysonotus, 3, 15, 31. 

bottae collinus, 3, 23, 24, 27. 

bottae comobabiensis, 3, 22. 

bottae depauperatus, 3, 20. 

bottae desertorum, 3, 12, 14, 16 29 
30. 

bottae desitus, 3, 12, 14, 15, 17. 

bottae extenuatus, 3, 24, 25, 27 31 
32. 

bottae fulvus, 3, 11, 13, 14, 15, 17 
23, 30. 

bottae grahamensis, 3, 23, 24, 27. 

bottae growlerensis, 3, 22. 

bottae hualpaiensis, 3, 14. 

bottae hueyi, 3, 24, 26. 

bottae modicus, 3, 18, 22 23 ''5 
28, 33, 34. ' " ' 

bottae mutabilis, 3, 14, 15, 17, 18 
19. 

bottae nasutus, 13. 

bottae nicholi, 3. 7, 9. 

bottae operosus, 3, 15. 

bottae parvulus, 3, 26, 27. 

bottae patulus, 3, 15, 16. 

bottae peramplus, 3, 11, 12, 33. 

bottae phasma, 3, 20, 22. 

bottae pinalensis, 3, 18. 

bottae planirostris, 3, 8, 9, 10. 

bottae pusillus, 3, 23. 

bottae riparius, 16. 

bottae subsimilis, 3, 16. 

bottae toltecus, 19. 

bottae trumbullensis, 3, S, 9. 

bottae virgineus, 3, 7, 8. 

burti. 34. 

burti proximus, 33. 

burti queschius, 35, 

cervinus, 17. 

chrysonotus, 15. 

desertorum, 12. 

fossor, 32. 

fossor kaibabensis, 32. 

fulvus catalinae, 25. 

fulvus collinus, 27. 

fulvus flavidus, 15. 



THE POCKET GOPHERS 



59 



rhomomys — Oontinned. 

fulvns grahamensi^<. 23. 
fuli'iis intermedins, 35. 
fulvns modicns, 2.S. 
fnJvHS nnitabiUs, 17. 
fiilriis peraiiipJus. 12. 
fulvns phasDia, 20. 
fulvns pnsillns, 23. 
fulvns snboles. 29. 
fulvus snhsimilis, 16. 
liarquahalae, 3, 31. 
latirostris, 11. 
mearnsi, 31. 
muralis, 3, 4, 30. 
perpaUidus ahsoiius, 10. 
perpallidiis depanpcratus, 20. 
perpallidns planirostris, 8. 
snboles, 3, 29. 
talpoides, 1, 5, 6. 
talpoides fossor, 5, 6, 12, 33. 
talix)icles kaibabensis, 5, 32. 
umbrinus, 1, 5, 6. 



umbrinus bnrti, 5, 33, 34. 

umhriiius chiricnhunr, 27. 

umbrinn.s intermedins, 5, 34, 35. 

nnihiinns proximns. 5, 33, 34, 35. 

iniibrinns qnercinns, 5, 35, 
Tinajas Altas pocket gopher, 20. 
toltecns, Thomomys bottae, 10. 
Tnle Desert pocket gopher, 20. 
Tunitcha Monntain pocket goiilier. 12. 
trumbnllensis, Thomomys bottae, 3, 8, 
9. 

nmbrinns, Thomomys, 1, 5, 6. 

Upper Gila Valley pocket gopher, 19. 

Verde Valley pocket gopher, 17. 
Virgin Valley pocket gopher, 7. 
vii'ginens, Thomomys bottae, 3, 7, 8. 

Whitish pocket gopher, 19. 

Ziou Park pocket gopher, 8. 



U. S. GOVERMENT PRINTING OFFICE: 1947-696493 



-!( 



Sfo :{, Gj 



UNITED STATES DEPARTMENT OF THE INTERIOR 

FISH AND WILDLIFE SERVICE 



RACCOONS OF NORTH AND MIDDLE AMERICA 



NORTH AMERICAN FAUNA 60 




UNITED STATES DEPARTMENT OF THE INTEKIOR 
Oscar L. Chapman, Secretary 

FISH AND VilLDLIFE SERVICE 
Albert M. Day, Director 



North American Fauna 60 



RACCOONS OF NORTH AND MIDDLE AMERICA 



BY 
EDWARD A. GOLDMAN 



With 

Foreword, Appendix, and Revision 

of Bibliography 

By 

Hartley H. T. Jackson 




UNITED STATES 

GOVERNMENT PRINTING OFFICE 

WASHINGTON : 1950 



For sale by the Superintendent of Documents, U. S. Government Printing Office 
Washington 25, D. C. - Price 45 cents 



U. S. SUPERINTENDENT OF DOCUMENT^ 

MOV 8 1950 



FOREWORD 

The present monograph, The Raccoons of North and Middle Amer- 
ica, was completed by its author, P^dward Alphonso Cioldman, in 
December 1940, and earh' in 1941 was submitted for pul)lication in the 
North American Fauna series. Advent of World War II delayed its 
printing, and at the time of the death of Major Goldman, September 2, 
1946, he had been so engrossed in the preparation of his manuscript 
on Biological Investigations in Mexico tliat he had not revised the 
raccoon manuscript. In the meantime one new subsj)ecies Procyon 
lotor niegalodous Lowery had lieen descril)ed and several papers relating 
to raccoons had been published. Since Major Goldman's death 
another subspecific name Procyon lotor man'tiniits Dozier has aj^peared 
and a few other papers have l)een pul)lished. 

In tliis hnal review and analysis of the manuscript it is believed de- 
sirable to leave Goldman's views and expressions as little changed as 
possible. Accordingly all editing has been done with this in view and 
only such changes made as would clarify and collate the text, or make 
consistent abbreviations and citations. Some important items to be 
noted have been indicated and explainefl in footnotes. No deletions 
of pertinent matter have tieeii made. The bibliography (p. 87) has 
been extended to include literature published to date. The two 
subspecies, Procyon lotor niegolodous Lowery (1943, p. 225) and Procyon 
lotor ninritiinus Dozier (1948, p. 286), descriptions of which may be 
-found in the appendix (p. 84), have been included in the distributional 
map of species and subspecies of the subgenus Procyon (fig. 1) but have 
not been included in the list of North American species and subspecies, 
with type localities (p. 27') , in the key to species and subspecies (p. 29), 
or in the general discussion throughout the text. 



ILLUSTRATIONS 

Plate Page 

1. Pacific Northwest Raccoon (Frocyon lotor pacificus) Frontispiece 

2. Skins (dorsal view) Procyon lotor lotor, Procyon cancrivorus pcniainensis^ 109 
W. Skulls (lateral view) Procyon lotor lotor, P. I. hirtus 11] 

4. Skulls (lateral view) Procyon lotor litoreus, P. I. elucus 113 

5. Skulls (lateral view) Procyon lotor incautus, P. maynardi 115 

6. Skulls (lateral view) Procyon lotor excelsus, P. I. psora 117 

7. Skulls (lateral view) Procyon lotor hernandezii, P. I. pintiilns 119 

8. Skulls (lateral view) Procyon pygmaeus, P. insulari.s insularis 121 

9. Skulls (dorsal view) Procyon lotor lotor, P. I. hirtus 123 

10. Skulls (dorsal view) Procyon lotor litoreus, P. I. elucus 125 

11. Skulls (dorsal view) Procyon lotor incautus, P. maynardi 127 

12. Skulls (dorsal view) Procyon lotor excelsus, P. I. psora 129 

13. Skulls (dorsal view) Procyon lotor hernandezii, P. I. puniilus 131 

14. Skulls (dorsal view) Procyon pygmaeus, P. insularis insularis 133 

15. Skulls (ventral view) Procyon lotor lotor, P. 1. hirtus 135 

16. Skulls (ventral view) Procyon lotor litoreus, P. I. elucus 137 

17. Skulls (ventral view) Procyon lotor incautus, P. maynardi 139 

18. Skulls (ventral view) Procyon lotor excelsus, P. I. psora 141 

19. Skulls (ventral view) Procyon lotor hernandezii, P. I. pumilus 143 

20. Skulls (ventral view) Procyon pygmaeus, P. insularis insularis 145 

21. Skulls (dorsal view) Procyon lotor elucus (male), P. I. elucus (female) 

P. I. lotor (lateral view of mandibular ramus) ^ _ 147 

22. Skulls (dorsal and ventral views) Procyon cancrivorus panamensis 149 

Figure 

1. Map, showing distribution of species and subspecies of subgeiuis Procyon. 24 

2. Map, showing distribution of subgenus Euprocyon (Procyon cancrivorus 

panamensis) in Panama 81 

VI 



fN .^/^ 




N 



.^. 



THE RACCOONS OF NORTH AND MIDDLE 
AIMERICA 

By EDWARD A. GOLDMAN, Senior Biologist, Biological Surveys, Branch of 

Wildlife Research 



INTRODICTION 

The raccoons, genus Procyon, colloquially known as ''coons," belonji: 
to the carnivorous family Procyonidae, which also includes the Ameri- 
can genera Na^^ua, Nasuella, Bassaricyon, and Potos, and the Old 
World genera Ailurus and Ailuro'poda of the sul)faniily Ailiuinae. 

The members of the Procyon lotor group (subgenus Procyon), with a 
transcontinental range from southern Canada to Panama, except in 
parts of the Rocky Mountain region, and including those inhabiting 
several distant islands, are among the most familiar and characteristic 
of North American mammals. This group is not known to occur 
south of Panama. It is overlapped in the Isthmian region by the 
so-called crab-eating raccoons of the subgenus Euprocyoii, which 
range from that northern limit as far south as Paraguay in South 
America. The raccoons have been greatly reduced in numbers or 
have disappeared in many formerly wooded sections, owing to clearing 
and intensive human occupation. Despite adverse conditions, how- 
ever, they have maintained themselves in many places with remarkable 
tenacity. Trapping for other fur bearers may have reduced the 
northern fringe to some extent, btit the general range of the group has 
been little diminished. At the present time raccoons reach their 
northern limit in regular occurrence on Vancouver Island, B. C. 

The continental forms of the subgenus Procyon constitute a com- 
pact assemblage of closely allied geographic races all assignable to 
Procyon lotor. Complete intergradation is evident in numerous cases 
and the relative value and combination of characters presented indi- 
cate such close relationships that it can safely be assumed where lack 
of material leaves gaps in the known ranges. 

In the present revision of the raccoons are treated the North Amer- 
ican continental species as far as the eastern border of Panama and the 

PLATE 1 
Pacific Northwest Raccoon {Procyon lotor pacijicus). 



2 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

species on outlying islands along- both the Atlantic and the Pacific 
coasts. Thirty species and sul)species are recognized. Twenty-nine 
of these are assigned to the sul^genus Procy(yn and one to the subgenus 
Euprocyon. 

The revision is based mainly on a study of raccoon material in the 
collection of Biological Surveys, Fish and Wildlife Service, and in 
other collections in the United States National Museum. These and 
358 specimens borrowed from other museums make a total of 1,337 
examined. The assemblage included the types or topotypes of most 
of the known species and subspecies. 

For the loan of specimens the writer is especially indebted to Dr. 
Thomas Barbour, Museum of Comparative Zoology, Cambridge, 
Mass.; the late Dr. Joseph Grinnell, Museum of Vertebrate Zoology, 
Berkeley, Calif.; Dr. W. H. Osgood, Chicago Natural History Mu- 
seum, Chicago, 111.; Dr. H. E. Anthony, American Museum of Natural 
History, New York City; Di-. R. M. Anderson, National Museum of 
Canada, Ottawa, Canada; the late Oldfield Thomas of the British 
Museum (Natural History) ; Francis Kermode, Provincial Museum, 
Vancouver, British Columbia; Dr. L. R. Dice, Museum of Zoology, 
University of Michigan, Ann Arbor, Mich.; and the late D. R. Dickey, 
Pasadena, Calif. Grateful acknowledgment is also due to Percy Shu- 
feldt, La Cueva, N. Mex., for the generous donation of specimens 
collected by him in Campeche, Mexico. Notes on his examination of 
specimens in the British Museum have been kindly furnished by Dr. 
Remington Kellogg, United States National Museum, Washington, 
D. C. Stanley P. Young, Fish and Wildlife Service, Washington, 
D. C, generously supplied the photograph for the frontispiece. 

Dr. E. W. Nelson became keenly interested in the raccoons, as shown 
by his work on those inhabiting the Florida Keys (1930a). ^ During 
the same time and in the following year new sul)species were described 
jointly by Nelson and the wi-iter in preparation for a revision of the 
group; but other projects claimed attention and our collaboration 
could not be carried beyond this preliminary stage. 

HISTORY 

The raccoons represent a highly successful i)ranch of a well-developed 
phylogenetic tree. Their ancestry has been traced far back to the 
genera Phlaocyon and Cynodictis of the Lower Miocene or Oligocene 
periods. Early progenitors of these animals probably also gave rise 
to such divergent modern families as the Canidae, the LTrsidae, and the 
Ylustelidae. For detailed discussion of the phylogenetic relation- 
ships of the raccoons see the authors listed in the Bibliography (p. 87), 
especially Wortman and Matthew (1899, p. 109), Matthew (1930, p. 

' Publications referred to parenthetically by date are listed in the Bibliography, pp. 87-106. 



RACCOONS OF NORTH AND MIDDLE AMERICA 3 

129), and Gregory (1933, p. 83). The genus Procyon was well repre- 
sented in the early Pleistocene of North America, when it already 
ranged across the present United States from the Atlantic to the 
Pacific. Among Pleistocene species described were Procyon priscus 
Le Conte (1848, p. lOG) from Illinois, Procyon simus Gidley (1906, p. 
553) from California, and Pi'ocyon nanus Simpson (1929, p. 575) from 
Florida. 

The name "raccoon" is derived from Indian appellations of the 
animal, which have been variously rendered as "aroughcun," 
"arathkone," and "arakun." The familial* abbreviation "coon" is 
in general colloquial use in the United States. An animal as common 
and conspicuous and possessing such peculiar and interesting traits as 
the raccoon could not remain long unobserved by explorers and set- 
tlers in its country, and as it became better known it was accorded a 
prominent place in the folkloi-e of the United States. 

The earliest reference to a raccoon fouml in the literature is by 
Captain John Smith (1612, p. 13), who in describing the animals of 
Virginia says: '^Tlwre is a beast they call AnnKjIicun, much like a 
haehjer, but vseth to live on trees as Squirrels doe." This reference 
was closely followed by that of Purchas (1614, p. 761) in describing 
the same region. 

Under the name "Mapach," and apparently as "Tepe Maxtlaton," 
and perhaps under others, tlie raccoon was recorded by Hernandez 
(1651, tract 1, pp. 1, 9) in southern Mexico. The voyager around the 
world, Dampier (1729, p. 276), mentions the abun(hince of these 
animals on the Tres Marias Islands, off western Mexico, which he 
visited in 1686. The early systematic term Vulpi affiiiis Americana 
was applied by Ray (1693, p. 179), in connection with a generalized 
description of animals probably representing both the subgenera 
Procyon and Euprocyon then undifferentiated and very imperfectly 
known from both North and South America. Quaint descriptions of 
the raccoon in the Carolinas were pul)lished by Lawson (1718, p. 121), 
and by Catesby (1743, p. XXIX). Hans Sloane (1725, p. 329) credits 
the animal to Jamaica as follows: "The Racoons are commonly here 
in the mountains, and live in hollow fiddlewood Trees, from whence 
they make Paths to go to seek Sugar Canes, which is their chief, if not 
only Sustenance." No specimens are available from Jamaica, and if 
this record was well founded it seems strange that it has not been 
supplemented by others. 

Evidently noting the general resemblances, Linnaeus closely asso- 
ciated the raccoon with the bear in the 1740 edition of his Systema 
Naturae (p. 35) as Ursus cauda eloiKjata in contradistinction to the 
true bear, Ursus cauda abrupta. Under the same name in 1747 (pp. 
277-289, table 9, figs. 1 and 2) he pubUshed a lengthy description of 



4 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

the raccoon accompanied by the earhest ilhistrations seen by the 
author. Of these, figure 1 is a sketch of the entire animal. In figure 
2 attention is directed to the strongly developed and peculiarly 
formed os penis, or baculum. 

The accounts of the raccoon in Pennsylvania and New Jersey by 
Peter Kalm in 1753 (Benson 1937, pp. 52-53, 111, 242-243) formed a 
part of the basis for Linnaeus' short description of Ursus lotor in the 
tenth edition of his Systema Naturae (1758, p. 48). Recognizing 
distinctive characters, Storr (1780, p. 35) used Procyon as the generic 
name for the group typifted by Ursus lotor Linnaeus. G. Cuvier (1798, 
p. 113) described Ursus cancrivorus, the crab-eating raccoon from 
Cayenne which later became the type of the subgenus Euprocyon Gray 
(1864, p. 705). Only a few new North American species or subspecies 
were added during the nineteenth century by Wagler (1831, p. 514), 
Gray (1842, p. 261), Baird (1857, p. 215), Bangs (1898a, p. 219; 1898b, 
p. 92), and Merriam (1898, p. 17; 1899, p. 107). Short papers descrip- 
tive of new forms by Merriam (1900, p. 151; 1901, p. 101), ALller 
(1911, p. 3), Meanis (1914, pp. 63-66), Hollister (1914, p. 142), 
Goldman (1913, p. 15), Nelson (1930a, pp. 7-10), and Nelson and 
Goldman (1930a, p. 82; 1930b, pp. 453-459; 1931a, pp. 17-20; 1931b, 
p. 308) have since appeared. 

RACCOON NAMES NOT CLEARLY ASSIGNABLE 

The following names that have been proposed for species of the 
raccoon are unrecognized or unassigned owing to the author's inal)ility 
to associate them with any particular region, or because of some 
obvious defect in status. If the type specimens of any of these are 
extant, it is possible that any such accompanied by skulls, may afford 
clues to their identity; because of the range of individual variation in 
subspecies, however, there is likely to be considerable uncertainty. 
Skins subject to fading over a period of many years are of very limited 
value for comparative purposes, beyond the determination of the 
subgenus. 

Procijon nivea Gray, Charlesworth's Mag. Nat. Hist., vol. 1, p. 580, 1837. "In- 
habits North America, Texas." "Fur soft, silky, white. Tail one-colored." 
No type specimen designated. Doubtless based upon an albino, as suggested 
in the original description. At least two subspecies occur in Texas. Name 
unidentifiable. 

Procyon hrachipirus Wiegmann, Archiv fiir Naturgesch., dritter jahrgang, erster 
band, p. 369, 1837. "Patria: Antillae?" Based on two specimens said to have 
come from the West Indies (see pp. 354-355), but their place of origin was 
regarded as uncertain by the describer, as shown by the notation. Figured by 
Wagner in Schreber's Saugthiere (p. 143 C). The plate illustration is of two 
brownish animals, the tail shown in one as quite short. 



RACCOONS OF NORTH AND MIDDLE AMERICA 5 

Procyon obscurus Wiegmann, Archiv fiir Naturgesch., dritter jahrgang, erster band, 
p. 370, 1837. "Patria ignota." Figured by Wagner in Schreber's Saugthiere 
(p. 143 D). The plate illustration is of a very dark-colored animal. This seems 
to be unidentifiable. 

[Procyon brachyurus] var. fusca Burmeister, Verzeichniss Zool. Mus. Univ. Halle- 
Wittenberg Saugeth., Vogel Amphib., 1850, p. 13. Based on Procyon obscurus 
Wiegmann and Procyon obscurus Wagner, in Schreber's Saugthiere, SuppL, 
vol. 2, p. 159, 1841, without description. 

[Procyon lotor] var. melanus Gra.y, Proc. Zool. Soc. London, 1864, p. 704. No type 
locality indicated. No type specimen designated. "Nearly black." 
Unidentifiable. 

[Procyon lotor] var. albina Gray, Proc. Zool. Soc. London, 1864, p. 704. (Nomen 
nudum.) 

Pr[ocyon] hernandezi castaneus de Beaux, Zool. Anzeiger, vol. 35, p. 624, April 26, 
1910. From Mexico. Based on a specimen (No. 357) which had been in 
the Royal Zoological Museum, Florence, Italy, since 1857. According to 
the description in part (p. 621), "Die Korperfarbe ist ein echtes und rechtes 
Kastanienbraun mit prachtvoU silbrigem Glanze." The color of raccoons is 
so variable that a single specimen does not afford reliable differential char- 
acters. The color described is unusual for a raccoon and might be due to 
fading or to erythrism. If the skin is accompanied by a skull, comparison of 
the latter with those of the several geographic races known to occur in 
Mexico might afford a clue to identity. 

Pr[ocyon] lotor rufescens de Beaux, Zool. Anzeiger, vol. 35, p. 625, April 26, 1910. 
Type locality unknown ("?Heimat"). Type specimen not designated. 
About 10 specimens said to have been examined. Body color inore or less 
suffused with rich red brown. Apparently not identifiable. 

Pr[ocyon] l[otor] flavidus de Beaux, Zool. Anzeiger, vol. 35, p. 626, April 26, 1910. 
Type locality "Southern United States?" Type specimen not designated. 
Only one skin examined. Color dirty yellow. Hairs of back neither ringed 
nor tipped with black. Apparently not identifiable. 

Pr[ocyon] hudsonicus Brass, Aus dem Reiche der Pelze, p. 564, April 1911. No 
type designated. Described as "sehr gross und granbraun.". Apparently 
based upon commercial skins assumed by the describer to be from Hudson 
Bay where no raccoons occur. The name is therefore unidentifiable. 

HABITS 

Few North American animals are endo^ved with more interesting or 
attractive ways than the raccoons. The general habits, as recorded 
by many observers, seem everywhere to be very similar for the mem- 
bers of each of the two subgenera. In Panama, Procyon and 
Euprocyon share to some extent the same local habitat, both favoring 
the vicinity of swamps and streams and both being addicted to the 
crab-eating habit as shown by stomachs examined. But Procyon 
seems to be more arboreal than Euprocyon, and the two probably 
depart materially in general behavior. Dr. Thomas Barboin* informed 
the author that the local representatives of both sidjgenera have been 
kept in captivity at the biological station on Barro Colorado Island 



6 NORTH AMERICAN FAUNA 6 0, FISH AND WILDLIFE SERVICE 

in Gatiin Lake, Canal Zone, and that he has noted that Procyon 
"washes" its food in the characteristic manner while Euprocyon does 
not. The writer's own general observations indicate that under 
natural conditions Procyon does not regularly wash its food and 
suggest that washing may be limited mainly to food supplied to 
animals in captivity. As Euyrocyon has a very restricted range in 
North America, the present discussion is limited chielly to the mem- 
bers of the typical subgenus Procyon. 

Much has been written on the life history of the raccoon of the 
eastern United States which may be assumed to apply, with some 
reservations, to all members of the Procyon lotor group. The peculiar 
habits of the raccoon began to attract the attention of the settlers 
during the early colonial period, as is shown by the following cpiaint 
and somewhat fantastic account of this animal in the Carolinas by 
Lawson (1718, p. 121): 

The Raccoon is of a dark-gray Colour; if taken young, is easily made tame, but 
is the drunkenest Creature living, if he can get any Liquor that is sweet and 
strong. They are rather more unlucky than a Monkey. When wild, they are 
very subtle in catching their Prey. Those that live in the Salt-Water, feed much 
on Oysters which they love. They watch the Oyster when it opens, and nimbly 
put in their Paw, and pluck out the Fish. Sometimes the Oyster shuts, and 
holds fast their Paw till the Tide comes in, that they are drown'd, tho' they swim 
very well. The way that this Animal catches Crabs, which he greatly admires, 
and which are plenty in Carolina, is worthy of Remark. When he intends to 
make a Prey of these Fish, he goes to a Marsh, where standing on the Land, he 
lets his Tail hang in the Water. This the Crab takes for a Bait, and fastens his 
Claws therein, which as soon as the Raccoon perceives, he, of a sudden, springs 
forward, a coiisideral)le way, on the Land, and brings the Crab along with him. 
As soon as the Fish finds himself out of his Element, he presently lets go his 
hold; and then 1h(» Raccoon encounters him, by getting him cross-wise in his 
Mouth, and devours him. There is a sort of small Land-Crab, which we call a 
Fiddler, that runs into a Hole when any thing pursues him. This Crab the 
Raccoon takes by putting his Fore-Foot in the Hole, and pulling him out. With 
a tame Raccoon, this Sport is very diverting. The Chief of his other Food is all 
sorts of wild Fruits, green Corn, and such as the Bear delights in. This and the 
Possum are much of a Bigness. The Fur makes good Hats and Linings. The 
Skin dress'd makes fine Womens Shooes. 

More accurate early descriptions of the animal in Pennsylvania 
and New Jersey are those of Kalm (Benson 1937, pp. 52-53): 

The quadruped, which ])r. Linnc in the memoirs of the Royal Academy of 
Sciences has descrilied by the name of Ursus cauda elongata, and which he calls 
Ursiis Lotor, in his Systema Naturae, is here called a raccoon. It is found very 
frequently and destroys many chickens. It is hunted by dogs, and when it runs 
up a tree to save itself a man climbs up after it and shakes it down to the ground, 
where the dogs kill it. The flesh is eaten and is reputed to taste well. The bone 
of its male parts is used for a pipe cleaner. The hatters purchase their skins and 
make hats of them, which are next in quality to those of beavers. The tail is 
worn round the neck in winter and therefore is likewise valuable. 



RACCOONS OF XORTH AND MIDDLE AMERICA 7 

And quoting Kalm further (Benson 1937, pp. 242-243): 
I have already mentioned something of the raccoon; I shall here add more of 
the nature of this animal and its mode of living in its habitat, in a place which is 
properly its native country [vicinity of the then village of Raccoon at or near the 
present town of Swedesboro, N. J.]. The English call it everywhere by the name 
of raccoon, which name they have undoubtedly taken from one of the Indian 
nations: the Dutch call it hespan, the Swedes, espan, and the Iroquois, attigbro. 
It commonly lodges in hollow trees, lies close in the daytime, never going out 
except on a dark, cloudy day; but at night it rambles and seeks its food. I 
have been told by several people that in bad weather, especially when it snows 
and blows a storm, the raccoon lies in its hole for a week without coming out 
once; during that time it lives by sucking and licking its paws. Its food consists 
of the several sorts of fruit, and corn, while the ears are soft. In gardens it often 
does a great deal of damage to the apples, chestnuts, plums, and wild grapes, 
which are its favorite food; to the poultry it is very cruel. When it finds the 
hens on their eggs, it first kills them, and then eats the eggs. It is caught by 
dogs, which trace it back to its nest in hollow trees, or by snares and traps, in 
which a chicken, some other bird, or a fish is put for bait. It generally brings 
forth its two or three young in May when it prepares its nest. Some people eat 
its flesh. It leaps with all its feet at once; on account of this and of several other 
qualities many people here reckoned that it belonged to the genus of bears. The 
skin is sold for eighteen pence at Philadelphia. I was told that the raccoons 
were not nearly so numerous as they were formerly; yet in the more inland parts 
they were abundant. I have mentioned before the use which the hatters make 
of their furs, that they are easily tamed, and that they like sweetmeats, etc. 
Of all the North American wild quadrupeds none can be tamed so easily as this 
one. 

In regard to th(> duration of life in the raccoons under natm-al 
conditions, no information is no\v at hand. Such data shoidd become 
avaihibh' in the futtn-e through the tagging or otherwise marking of 
animals cajjtiu'ed and liberated. According to Flower (1931, p. 177), 
a male raccoon lived in the Rotterdam Zoological Garden from Sep- 
tember 30, 1890 to May 6, 1900, 9 years, 7 months, and 6 days, and 
an albino was in the London Zoological Garden from May 6, 1884, 
to February 27, 1898, 13 years, 9 months, and 21 days. He also 
mentions a crab-eating raccoon that lived in the London Zoo 1.5 
years, 10 months, and 5 days. Lowery (1936, p. 19) quotes Clau<k^ 
Odum of Bernice, La., who said that he kept a raccoon in captivity 14 
years. 

FOOD AND GENERAL ACTIVITIES 

Throughout the vast range of the group, raccoons favor the vicinity 
of water in forested regions; but they also occm- along streams travers- 
ing open desert areas. Although raccoons are truly omnivorous, 
feeding to a considerable extent on a great variety of plant substances 
such as acorns, beechnuts, berries, persimmons and other fresh fruits 
of many kinds, and corn in the. "milk" stage, most of their food is 
obtained in or near shallow water in swamps and marshes, and along 



8 NORTH AMERICAN FAUNA 6 0, FISH AND WILDLIFE SERVICE 

the shores of streams, lakes, and brackish Lag-oons, and even along the 
sea coasts, as in the Florida Keys and other islands. In such places, 
frogs, ^mall fishes, crayfish, crabs, clams, oysters, insects, small 
mammals, reptiles, and other animal foods are sought, as shown by 
the characteristic telltale footprints revealing the course of nocturnal 
wanderings and by stomach examinations. 

As water recedes to lower, levels and pools become detached, fish, 
of which raccoons are very fond, are more readily captured by them. 
Referring to some water holes near Lake Drummond, Dismal Swamp, 
Va., in October 1895, A. K. Fisher reported: "Judging from the 
tracks about these pools, as many as a dozen must have come every 
night to feed on the fish imprisoned therein. The heads of catfish, 
pike, eels, and perch were found in abundance under the bushes and 
along the edges where the raccoons had dropped them." According 
to Mary J. Rathbun (1918, p. 401), the fiddler crab {Uca pugilator) 
is the main food of the raccoon on the bay shores next to the Gulf 
in Texas. Young birds and eggs in the nest are often taken, and 
departing from the usual acpiatic hal>itat, the raccoons occasionally 
make raids on the farmer's poultry. 

A complete list of the miscellaneous items composing the diet of 
raccoons would be exceedingly long and would vary in accordance 
with the season and with local conditions. On Key I^argo, Fla., 
E. W. Nelson found the raccoons feeding extensively on the ripening 
fruit of the marlberry (Icacorea paniculata) in March. The taking 
of dry l)erries may be resorted to when more acceptable food supplies 
are insufficient. Examination of stomach contents has revealed the 
hard seeds of the hackberry and juniper berries in Texas raccoons and 
Vernon Bail(\y found these animals feeding upon manzanita {Arcto- 
stajjhylos) berries in California. The eating of grasshoppers has been 
reported in Texas. 

Although raccoons enter the water freely, much time is spent in 
patrolling the muddy shores. Closely crowded tracks, suggesting the 
imprints of human baby hands and feet, often mark the lines of least 
resistance up and down the banks of streams or through swamps, and 
well worn trails are gradually formed, disappearing in places at the 
edge of the water where it was necessary for the animals to wade or 
swim, and reappearing again on the farther side. In addition to 
water, trees, especially hollow ones affording shelter, are almost indis- 
pensable for the well-being of most raccoons. There seems to be evi- 
dence that the clearing of timber, especially the cutting of the large 
shelter trees needed for refuge and hibernation, has been an important 
factor in reducing the numbers of these animals in the northern part 
of their range. In the warmer southern territory, where hibenuition 
does not occur, shelter trees are evidently not so essential. Mangiove 



RACCOONS OF NORTH AND :^11DDLE AAIERICA 9 

swamps, with no large trees within many miles, are regularly inhabited 
by large mmibers of raccoons that seem able to forego supplies of fresh 
water. Although hollow trees are favored for the shelter afforded, 
holes in banks and rocky ledges are also occupied, especially in locali- 
ties where such trees are few or absent. Raccoons are mainly noc- 
turnal in their search for food, but they sometimes come out during 
the day, and are especially fond of sunning tliemselves, usually sprawled 
in a variety of postures on the larger upper liml)s of trees. 

SENSES AND INSTINCTS 

The sensory organs in raccoons are evidently highly developed. 
Many observers accord these animals a reputation for great curiosity 
and cunning, and a cleverness or adroitness, involving a high order of 
general intelligence. As a result of experiments Cole (1907, p. 261) 
concluded that "in the rapidity with which it forms associations the 
raccoon seems to stand midway between the monkey and the cat. In 
the complexity of the associations it is able to form it stands nearer 
the monkey." It is remarkable, as pointed out by Stock (1929, p. 288), 
that although Procyon occurred in California during the Pleistocene, no 
member of the family has been found in the Rancho La Brea deposits. 
This is probably due to the caution of raccoons in approaching and 
investigating water holes or such natural traps as the miry, sticky tar 
pits presented. 

The senses and instincts of raccoons, as exhibited by animals in 
captivity, have been carefully studied and well described by Cole 
(1912), who concluded that although most of the senses are strongly 
developed, that of smell is less utilized than the others. His results 
seem worth quoting at length: 

The most conspicuous behavior of the raccoon seetns to be associated with the 
sense of touch, which is liighly developed in the pahn of the forepaw and the tip 
of the nose. During their hours of activity the animals were most often busy in 
exploring with their paws the floor and objects on the floor of the room in which 
they were kept. . . . Dark places, as your pocket or a knothole, are explored by 
touch hundreds of times. . . . Notwithstanding the strength of the raccoon in 
clinging and climbing, no touch is softer or more gentle than that of his forepaw'S 
when engaged in this investigating activity. 

An evidence that the nose is sometimes used for pure touch is the fact that these 
animals frequently investigated the experimenter's hands, and even his face, with the 
nose. This also seemed to be an affair of pure curiosity and quite breathless. . . . 
Occasionally thej' would both touch a strange object with the nose and sniff 
at it also. . . . 

The raccoon's taste for sweets is especially marked. All other foods were 
promptly deserted for cane sugar by my animals. . . . My raccoons avoided all 
food which had a purely sour taste, yet ripe apples and peaches were eaten which 
have for human taste a slightly acid tang along with the sweet flavor. Unlike 
herbivorous animals the raccoon refuses to taste salt. . . . 

876119° — 50 2 



10 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

Next to sugar the raccoons preferred boiled beef and they were ahnost equally 
fond of uncooked apples, peaches, plums, and cherries. My animals never ate 
the raw lieef which we offered them a few times. Some raccoons have been forced 
to eat it but they do not appear to thrive on such food. My raccoons were often 
seen to catch and eat flies. They would eat grains of corn, even when dry and 
hard, if they were hungry. Bread made of either corn-meal or flour was readily 
accepted. It seems evident, therefore, that the raccoon in his native haunts lives 
upon forest fruits and buds, and upon flies, beetles, minnows, etc. . . . 

So far as I could observe the raccoons did not often employ the sense of smell, 
though this may have been due to their captive condition. In no case did they 
seem to find pieces of meat on the floor by means of smell. If one of them saw 
a small piece of meat dropped in the hay on the floor he would search for it care- 
fully but beyond a distance of a few inches he did not seem to smell it. They 
found small pieces of loaf sugar on the floor ciuite as promptly as they did meat, 
yet from the standpoint of the human sense of smell sugar has no odor. 

In one case smell was evident. When the animals were to be fed the basin of 
food was usually placed on the step while the door was being unlocked. During 
this time all of the raccoons sniffed noisily at the crack beneath the door. When 
it was opened, however, they looked for the food basin. So in this case smell was 
evident only when sight could not be used. 

The studies of Cole indicated that the raccoon has a keen sense of 
sight. In regard to hearing he says: 

This appears to be the special protective sense of the raccoon. The slightest 
sound produced (1st) perfect immobility, and (2d) fear and scurrying to the 
highest part of their place of confinement. . . . 

Every sound at a distance was listened to intently for several seconds after the 
experimenter had ceased to hear it. On one occasion all the raccoons became 
still and yet the observers could hear no sound. Investigation showed that a 
man was trundling a wheelbarrow over the grass plot at least 100 yards distant 
from the house in which the raccoons were kept. 

The sound caused by dropping on the floor a piece of meat, one-half the size of 
a grain of corn, was often heard by each of the animals. They turned directly 
toward the source of the sound. Hence they not only hear faint sounds but 
localize them well. Localization was further tested by putting raccoon No. 3 in 
a large box with a solid back. The experimenter then scratched on the outside of 
the back of the box with a small stick. The raccoon turned directly to the spot. 
The place was changed some two feet. He tiu-ned instantly to the new place and 
grasped with both forepaws at the exact spot. He did this repeatedly. His 
behavior suggests that localization of sound is much more definite than that of the 
human ear. His grasping at the spot might indicate that the raccoon catches 
some small prey partly by the aid of hearing. . . . 

On the principle that animals which make souTids hear sounds we may, in con- 
nection with hearing, mention the sounds which the raccoon is capable of making. 
A warning growl always accompanied eating when they were fed. When hungry 
they sometimes emit a sound about midway between a whine and a purr, "a 
whimpering cry." This sound is well-known to woodsmen and is far more char- 
acteristic of the young than of the adult animal. . . . When forcibly held their 
whining and growling is somewhat similar to that of a dog. In fighting the animal 
gives short, sharp barks as he snaps. 

Turning once more to the whining-purr, there is less and less of it (in captivity) 
as the animals grow older and only long waiting for food produces it. In the 



RACCOONS OF NORTH AND MIDDLE AMERICA 11 

forest it may be used as a call to others. A very young raccoon, making this cry 
from loneliness or in search of its mother, will cease to make it if gently stroked or 

scratched. 

Of the climbing- habit Cole says: 

This instinct involves the sense of support, which is present before the raccoon 
):)ossesses either the strength or the muscular co-ordination necessary for climbing, 
and the impulse to cling to any support. The sense of support is best described by 
an example. When raccoon No. 5 was probably not more than two weeks old I 
placed him one day upon the top of a small closed bo.x six inches high. He groped 
o^•er the top of this box . . . with his forepaws extended, feeling the way. But 
1 he moment his paws felt the edge of the box the animal shrank back and began to 
grope in another direction. Again he would find the edge and again shrink back 
and start anew. Apparently at this age vision did not serve to show him that he 
might safely droj) to the floor. It seems likely that this impulse enables the young 
raccoon to remain safely in a high nest, even though it were not enclosed. . . . 

When the raccoon is a month old it is able to sustain its weight by clinging to a 
support by any one of its paws and this it does instinctively. ... As soon as the 
young raccoon can walk well any bush or free arouses his impulse to climb. At 
first there is some awkwardness and two of our animals were seen to fall from a 
small tree, when al)out eight weeks old. A little later they could hardly be 
dislodged at all. . . . As Brehm states, the raccoon often climbs along a branch 
with his back down "like a sloth or an ape"". . . . My raccoons always laid hold of 
one l)ough before releasing the other. They go from one bough to another very 
ciuickly yet they rely much more on their strength than on their agilit}^ 

Cole agrees with other observers in the conclusion that raccoons are 
very playful: 

One would sit for a long time and play with his hind feet or the tip of his tail. 
Three were ol)served to play in this fashion for one and a ciuarter hours, with 
almost no pause. While my animals had to work twice a day for their food I 
observed only momentary play, or perhaps curiosity, as the tendency to pick up a 
straw or bit of cornhusk and roll it for a moment between their forepaws. In 
some degree, therefore, their play seems to depend on the possession of surplus 
energy. When well rested they played roughly with each other in mock fights, 
running and seizing each other gently with the teeth, rolling over and over in their 
tussles. In this play they would often climb to the shoulder of the observer, 
whereby he may learn both the strength of their grip and the sharpness of their 
claws. They also make a pretense of biting your hand in play, a characteristic 
reaction of the pet raccoon. 

In this connection Cole quotes Beckmann, as follows: 

"In the munberless leisure hours which every captive raccoon has he does 
thousands of things in order to dispel the tedium. Now, he sits upright in a 
secluded corner, and with a most earnest expression he is busied in trying to tie 
a straw around his nose. Now, he plays thoughtfully with the toes of his hind 
foot, or snatches after the wagging end of his long tail. At another time he lies 
on his l)ack and has a whole heap of hay or dry leaves hugged against his belly and 
he tries to tie down this loose mass by drawing his tail tightly over it with his 
forepaws." 



12 NORTH AMERICAN FAUNA 6 0, FISH AND WILDLIFE SERVICE 

The impulse to follow which seems to be inherent in young animals 
of many kinds was noted by Cole, who says: 

After learning to walk, the raccoons would all follow me, or anyone else, with 
the utmost eagerness. If I ran they struggled through the grass at their best 
rate, giving the instinctive cry more and more shrilly as I got further away from 
them, and ceasing to give it when they overtook me. In the middle of the 
seventh month this instinct to follow began to wane. When released from their 
place of confinement each one tended to go on an exploring tour of his own and 
to make for a nearby tree. At this time they would still follow if called. A 
month later no one of the four would follow at all, and their period of infancy 
was past. 

Althoug-h Cole regards the raccoon as especially good-natured, 
". . . yet anger or ferocity was observed in these animals at about 
the twelfth week of their age. Though scrupulous care was taken to 
keep the animals tame they became fierce if they were left without 
being handled for a few days. In the fighting attitude the ears are 
laid back, the head lowered and the posterior portion of the body 
sharply humped up. Growling and luifleshing the teeth accompany 
this fighting attitude and, when provoked the raccoon is an ugly 
fighter." He found that his raccoons showed fear by starting at 
sounds, and the sudden darkening of the room caused by the door 
blowing shut produced in young animals a panic for a moment. 
Indifference to each other's behavior was marked. No certain 
evidence of the sexual instinct was noted by Cole until the twelfth 
month. 

In regard to the practice of washing food that caused Linnaeu§ to 
apply the name l(>f<»\ and the Germans Waschbdr, Cole says: "My 
raccoons did not always dip their food in water. No doubt this was 
partly due to their being fed together so that they formed the habit 
of eating rapidly. . . . Nevertheless, I do not believe that the 
raccoon in his native state will carry food very far for the purpose of 
'washing it'." Whitney (1931, p. 35) comments on this point as 
follows: "Unquestionably the most common error into which writers 
have fallen in regard to the habits of raccoons is that the raccoon 
washes most of the food that he eats ... in the wild state the 
raccoon washes almost nothing that he eats." He regards the error 
as due to observations made on animals in confinement. It is obvious 
that the washing of many kinds of food taken by raccoons, especially 
at a distance from water, would be impracticable. It is probable that 
under natin-al conditions raccoons wash only shellfish and other food 
gathered in or aljout water, the washing being often necessary to 
remove sand or other gritty matter. 

Concerning the sleep of raccoons, Cole writes: 

There are two rather characteristic positions in sleeping. In one the animal 
lies on his back with his forepaws placed over his eyes. A young raccoon, when 



RACCOONS OF NORTH AND MIDDLE AMERICA 13 

held and somewhat frightened, also puts both forepaws over his eyes, thus giving 
a somewhat comical appearance, suggestive of "hiding its face in its hands." 
Another position in sleep consists in rolling the body almost into a ball with the 
lop of the head placed flat on the floor between the forelegs. In this position even 
I he ears are hardly visible. Though the animal does sleep in other positions 
ihese two are most common. It would seem that the raccoon sleeps best, there- 
fore, with his eyes not only closed but covered, and for protection he depends 
most upon his lofty nest and its concealment from enemies. 

Among mental attributes of the raccoon, both Davis (1907, p. 486) 

and Whitney (1933, p. 112) regard curiosity as an outstanding 

characteristic. 

BREEDING 

The period of gestation in Procyon loior, at k^ast in the northern 
part of its range, has l)een (k^termined by various authorities to be 
al)out 9 or 10 weeks. One of the more definite records is that of 
Gan(k>r (1928), rehiting in a singk instance to Procyon lotor psora in 
southern Cahfornia. An animal kept in captivity was mated January 
27 to 29 and the yoimg w^ere born April 3. Another record is that of 
Brow^n (1936) of 69 days from first coition. 

A litter of small yoimg was collected l)y E. A. Preble at Tuckerton, 
N. J., Jinie 23; one taken by B. V. Lilly at iVbbeville, La., suggests 
that in sotithern localities the season may be more irregidar. Rac- 
coons breed btit once a year and four yoimg are usually produced at 
a birth, but the nimiber may vary from two to six. In regard to 
breeding in the Adirondacks of northern New York, Merriam (1884, 
p. 94) says: 

The Raccoon makes its home high up in a hollow of some large tree, preferring 
a dead limb to the trunk itself. It does little in the way of constructing a nest, 
and from four to six young are commonly born at a time, generally early in April 
in this region. The young remain with the mother about a year. 

The act of copidation, rarely recorded in raccoons under natm'al 
conditions, was witnessed by the author on Blackbeard Island, Ga., 
April 19, 1939. From a point of vantage on high groimd in the woods 
a mated pair, imconscious of his presence, was observed in short 
grass in the open marsh about 75 yards away. With a pair of field 
glasses a very clear view of the animals in bright sunshine was obtained 
at short range. When first seen at 2 :05 p. m., the female, surmoimted 
hy the male, was in a standing position and sexual conjunction 
appeared to be already complete. Rhythmical movements of the 
hind qiutrters of the male were interrupted by periods of qinet. Several 
times he shifted position slightly from one side to the other, but 
remained most of the time w^ith his head resting near the median line 
of the back of the female. The latter remained passive imtil at the 
end of about half an hour by the watch she laid her ears back and tin-ned 



14 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

her head, showing her teeth and apparently snarhng at the male, 
although even at the short distance no sound was heard. The male 
quietly slipped from her, and both animals moving only a few feet 
immediately resumed their search for small crabs that were numerous 
in the marshy ground. The fur on the lower part of the back of the 
female had become considerably rumpled, but this was ignored by her 
in the search for food. 

hibp:knation 

The winter activities of raccoon vary in southern and northern 
latitudes. In the southern United States and southward the raccoons 
are active thi-oughout the year. In the North these animals become 
torpid, and there is a kind of hibernation or partially suspended 
animation, similar to that of the bears and only approximating the 
deep lethargic winter sleep of some other northern animals. In 
describing behavior in winter in Canada, Wesley Mills (1892, sec. 4, 
p. 50) refers to W. Yates, of Hatchly, Ontario, and says: "This 
observer has made some very interesting observations on a tame rac- 
coon (Procyon lofor). This creature lived in a hollow log lined with 
straw and drowsed away the greater part of December and January, 
leaving any food placed before him unnoticed. The raccoon is 
known to spend the greater part of the winter in hollow elm trees 
in this part of the country, and Mr. Yates points out that the cutting 
down of most of these trees resulted in the raccoons betaking them- 
selves to underground burrows including those once occupied by 
foxes." According to Seton (1929, p. 252): "In the Red River 
Valley [Canada], the sleep lasts from mid-November to early March." 
Concerning hibernation in the Adirondacks of northern New York, 
Merriam (1884, p. 93) writes: "The Raccoon hibernates during the 
severest part of the winter, retiring to his nest rather early, and 
appearing again in February or March, according to the earliness or 
lateness of the season. Disliking to wade through deep snow he 
does not come out much till the alternate thawing and freezing of the 
surface, suggestive of coming spring, makes a crust upon which he 
can run with ease." 

ECONOMIC STATUS 

Raccoons are naturally prolific, and owing to very extensive geo.- 
graphic range and adaptability the forms of Procyon lotor constitute 
a wildlife asset of major recreational and economic importance. In 
the extreme scarcity of money in pioneer days raccoon skins supplied 
an important element in helping the people to maintain their existence. 
In 1788 (Chase 1911) the residents of a mountain section in Tennessee 
organized the local "State of Franklin." Money was scarce, and the 



RACCOONS OF NORTH AND MIDDLE AMERICA 15 

salaries of public officials were paid in animal skins, including the 
following: "... secretary to his excellency, the governor, 500 rac- 
coon [skins] ; . . . clerk of the house of commons, 200 raccoon [skins] ; 
members of assembly, per diem, 3 raccoon [skins]; . . ." Through- 
out the pioneer days raccoon slvins were regular articles of barter. 
'I'he slvins were especially popular for making caps and coats, the 
latter use extending to the present time as garments for both men and 
women. Although their original numbers have greatly decreased, 
owing to the reduction or elimination of suitable habitat incident to 
human encroachment, raccoons have persisted where many other 
native animals have become extinct. Aside from the fur produced, 
iheir value in providing excellent nocturnal sport for an army of 
hunters and exercise for the "coon" dogs nearly throughout the 
forested sections of the country is well known. This hunting asset 
is becoming better appreciated by vState game commissions and 
sportsmen's associations, and the lil)eration of raccoons in suitable 
places is a regular part of the annual program of wildlife management. 

The meat, especially of young raccoons, is an accepted article of 
food in some parts of the country and is very palatable. During the 
early days in California, according to Newberry (1855, p. 47), raccoons 
in considerable numl>ers were sold in the San Francisco market, 
commanding a price of one to three dollars each. 

Raccoons are destructive to human interests in some places to a 
limited extent. Of the economic status of the animal in its typical 
region, Pennsylvania and New Jersey, Rhoads (1903, p. 182) says: 

Dr. Warren reports answers from correspondents which condenin this animal 
as a stealer of fish, especially trout. Others say it does not catch many of these 
but is after crayfish chiefly. His raids on nesting turkeys I can vouch for, the 
eggs being sucked. His destruction of poultry is occasionally severe and he likes 
green maize ears dearly. No doubt he is a destroyer of birds' nests, eggs and 
young, both terrestrial and arboreal. He catches some mice, but being a slow 
sort of fellow, prefers more leisurely employment. On this account, he is quite a 
vegetarian, grapes, nuts, fruits and certain vegetables falling to his share. His 
furs for warmth and his carcass for food about compensate for the direct losses 
sustained by humanity in his depredations. 

In the Gulf Coast Region, where raccoons still al)oini(l, conditions 
are described by Kopman (1921, p. 28) thus: 

One of the principal foods of the raccoon in Louisiana is crayfish. Among 
vegetable foods, corn in the milk, persimmons, wild grapes, and palmetto berries 
are very acceptable to the "coon." As a destroyer of poultry the raccoon is 
often a great nuisance, and it takes many wild birds. These animals are estab- 
lished on many of the bushy islands of the coast, and they eat the eggs and young 
of the seabirds and other aquatic species breeding there. On Marsh Island and 
other bird and game preserves on the coast owned by the State, the Department 
of Conservation has had to provide for systematic trapping of the raccoon. . . . 



16 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

In considering the natural enemies of birds, Forbiish (1916, pp. 
24-25) discusses the raccoon as follows: 

There is some evidence to the effect that the raccoon robs birds' nests, but it 
is not numerous enough now in settled regions to be very destructive. Its fond- 
ness for green corn has not endeared it to the farmer, and the sportsman and 
angler believe that it destroys game and fish. Add to these alleged reasons for 
its destruction the increasing price for its skin in the market and we can see why 
the "coon" is not destined long to be a great factor as an enemy of birds, except 
possibly on lands where all animals are protected. 

An early mention of the raccoon in New England is by Josselyn (1672), 
who says: "The Raccoon liveth in hollow trees, and is about the size 
of a Gib Cat; they feed upon Mass, and do infest our Indian Corn 
very much; they will be exceeding fat in Autumn; their flesh is some- 
what dark, but good food roasted." 

Corn is grown extensively throughout much of the range of the 
raccoon, and perhaps more complaints are lodged against the animal 
for damages to this staple crop than to any other human interest. 
Cornfields adjoining woodland inhal)ited by raccoons may be in- 
vaded, usually for only a short distance, about the time that the 
ears reach the "milk" stage. The stalks are pulled down, or the 
ears stripped oft" and partly eaten and left scattered over the ground. 
In some of the most serious instances noted by the writer more than 
one-half of the corn was destroyed on areas several acres in extent. 
Other grain crops (as, for example, kafir corn) and fruits and vege- 
tables of many kinds are also subject to some injury. Personal 
observations have shown that raccoons sometimes become nocturnal 
despoilers of the nests of waterfowl. 

General observations over a wide range indicate that the depre- 
dations of raccoons are sporadic in relation to himian interests, 
involve few individuals, and are usually so limited and local in extent 
that they are quite negligible. The removal of one or two oft'enders 
by trapping or shooting will put a stop to the raids in most cases. 
In a very few instances systematic trapping may be necessary to reduce 
a local raccoon population that has l)ecome too numerous and destruc- 
tive. The isolated cases of damages sustained are, in general, far 
outweighed by the asset value of the species. 

The northern subspecies of Procyon lotor are among the most 
important fur bearers, but pelts of the forms of the crab-eating 
raccoon, Procyon (Euprocyon) cancrivorus, are of little value, owing 
to the short, thin, bristly character of the pelage. 

Some idea of the numbers of raccoon {Procyon lotor) pelts that have 
been handled as furs may be gained from estimates based upon sta- 
tistical studies made in 1925 by Frank G. Ashbrook of the then Bureau 
of Biological Survey, United States Department of Agriculture 
(now part of the Fish and Wildlife Service, Department of the In- 



RACCOONS OF NORTH AND MIDDLE AMERICA 17 

terior), and Horace J. McMiillen, of the then National Association of 
the Fur Industry (Ashbrook and McMullen 1925). The data 
gatliered from the principal fur auctions in the United States and in 
London, and from raw fur receiving houses indicated that the average 
yearly production of raccoon pelts for the 10 years preceding 1925 
was 600,000 to 1,000,000. These figures were also taken to represent 
the average yearly consumption for the period stated. [The most 
recent (1948) information compiled by the Fish and Wildlife Service 
indicates an annual take of from 1 to IK million pelts in the United 
States.! 

The raccoon has an assured place as one of the more important 
American fur-bearing animals and will continue to hold this position 
so long as it can be maintained in suitable numbers. [It is outnum- 
bered only by the muskrat, opossum, and skunk in pelts taken.] 

The natural supply of raccoon furs is being gradually reduced 
through the general encroachment of civilization upon the range of 
the animal. Aside from trapping for the fur, hunting for sport with- 
out adequate regulations, and harassment by dogs, the drainage of 
water areas and the cutting of timber, especially the older trees afford- 
ing convenient sheltering hollows, have resulted in conditions unfa- 
vorable for raccoons. Displaying wonderful adaptability, raccoons 
still maintain themselves even in many well-settled areas, often in the 
viciuity of human habitations or even large cities, with a persistence 
truly remarkable. Experiments have been conducted on raising 
raccoons in captivity. Much should be done, however, to further 
better management of the raccoon in the wild, not only as an important 
fur bearer and for the sport afforded in its chase, but as a characteristic 
American animal of outstanding general interest owing to its peculiar 
and attractive habits. 

GENERAL CHARACTERS 

The raccoons as a whole present a narrow range of variation in 
external appearance. The general color pattern, including the black 
facial mask and the barred tail, is everywhere very similar, even for 
the two subgenera. The subgenus Procyon, embracing the nmnerous 
forms of the typical North American group, is, however, easily dis- 
tinguished by the normal, or backward direction of the hair on the 
nape, by the presence of underfur, and by the grayish forearms and 
thighs. In Euprocyon, on the other hand, the pelage of the nape is 
reversed, undei'fur is absent, and the forearms and thighs are usuall}^ 
blackish instead of grayish. 

The normal number of mammae seems to be six in both Procyon and 
Euprocyon, but has been found to vary to eight in the latter subgenus. 
Cranial and dental subgeneric distinctions are pointed out in the 
treatment of subgeneric characters. 



18 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

In both siibgenera the plantigrade structure of the feet is an out- 
standing feature. The fore feet somewhat resemble tiny hands, with 
long fingers opposable to a high degree, possessed of great strength, 
and yet capable, in Procyon at least, of being used with a remarkable 
deftness and delicacy of touch. The digits of the hind feet are much 
less opposable, and the imprints of the broad flattened soles along 
muddy shores may be likened to those of a small child's feet. Al- 
though the favorite haunts of the members of both subgenera are in 
the vicinity of water and much time is spent upon the ground, Procyon, 
as compared with Ewprocyon, is provided with claws that are narrower, 
sharper, more compressed laterally, and strengthened by greater ver- 
tical depth at the base, better adapting this subgenus for climbing and 
a more arboreal life. In Panama, where the two subgenera occur 
together, the crab-eating habit is shared in common, but may be in- 
dulged in to a greater extent by Euprocyon than by Procyon. The 
broader, less trenchant cusps in the molariform teeth of Euprocyon, 
as compared with those of Procyon, are better fitted for crushing hard 
substances. Along the coast of Salvador, mangrove swamps are in- 
habited by a local form, Procyon lotor dickeyi, which feeds extensively, 
perhaps principally, upon crabs. The abrasive effect of such a diet 
on the teeth of a member of the typical subgenus is there strikingly 
shown by the early wear and ra])id shearing off of the crowns of the 
molars, leaving the premolars comparatively little affected. In some 
of the older specimens of dickeyi the molar crowns are reduced until a 
mere shell remains near the roots. This may, however, be due to 
some unusual local condition as such rapid or extensive wear has not 
been observed anywhere else. 

The l)lack mask varies somewhat in extent, and some forms are 
])aler than others, but owing to general uniformity in pattern of color- 
ation in each subgenus, recourse must usually be had to size and to 
cranial and dental modifications in tracing the relationships of species 
and subspecies. 

In the subgenus Procyon most of the sutures of the skull ar(^ easily 
traced at birth. Among the earliest sutures to close are those of the 
basicranial segment surrounding the foramen magnum. The supra- 
occipital, exoccipitals, and basioccipital are all firmly united, and the 
sutures have disappeared before the permanent dentition is fully in 
place. The union between these bones and the remainder of the sivull, 
however, remains distinctly visible until finally closed later with ad- 
vancing age. The jugals unite with the maxillae earlier than with 
the squamosals. Progressive obliteration extends to the maxillo- 
premaxillary sutures and to the median line between the frontals, 
while the parietal sutures remain <Hstinct. The closure of the pai-ietal 
sutures may be taken as an indication of maturity. In old age all the 



RACCOONS OF NORTH AND MIDDLE AMERICA 19 

bones of the skull become coalesced, among the last to unite firmly 
being the nasals and the mandibles. A well-developed, sometimes 
high and trenchant, sagittal crest commonly present in the older males 
is less frequent and less prominent in the females; but in many old 
adults of both sexes the temporal impressions do not unite to form a 
crest. The deciduous dentition is retained only a short time. The 
permanent middle incisors appear before the molars. 

In the continental forms of the subgenus Procyon subspecific dis- 
tinctions rest upon combinations of relatively slight characters, 
indicating close relationships. Although the characters do not stand 
out very conspicuously as a rule, and due allowance must be made for 
individual variation, they are maintained with a fair degree of con- 
stancy over areas often of considerable extent. Some of the more 
extreme forms of the intergrading series are very similar in external 
appearance, but are difl'erentiated by well-marked details of cranial 
structure. Skull characters, rather than color, must therefore be 
relied upon in determining systematic relationships. In dental 
sculpture all the forms are very similar, but they vary greatly in the 
size of the teeth and, to some extent, in the form of the molar crowns. 

In tracing the relationships of the numerous forms of the subgenus 
Procyon the principal characters of taxonomic value are the following: 
General color, whether light or dark, plain grayish, or suffused with 
ochraceous buff, or varying shades of rusty rufous; relative develop- 
ment of the black mask, whether continuous across middle of face, 
extent of black postauricular spots, and of white facial markings; 
general form of the skull (especially of the brain case and the frontal 
profile), massiveness, development of ])ostorbital processes and of 
zygomata, width of i)alate, size of auditory bullae; size and relative 
length and breadth of large molariform teeth. The males are usually 
deci(k'dly larger than the females in all dimensions, but the sexes agree 
closely in details of cranial structure. 

PELA(;K AM) MOLT 

The pelage differs widely in the subgenera Procyon and Euprocyon, 
as pointed out in the treatment of subgeneric characters. In the 
subgenus Procyon it is longer, softer, and much denser than in Eupro- 
cyon, the denseness being largely due to the fine underfur, which 
dift'ers in texture from the longer overfur or guard hairs, and which is 
absent in Eu))rocyon. Owing to the dift'erences in density and texture 
of the hairs, Euprocyon is of little value for the fur. 

The annual molt in the subgenus Procyon extends over a lengthy 
period during the sunnncr, at least in the more northern and more 
heavily furred subspecies. The new pelage, rather short in the fall; 



20 NORTH AMERICAN FAUNA 6 0, FISH AND WILDLIFE SERVICE 

becomes long:er in the winter. In the subgenus Etiprocyon — inhabiting 
tropical countries — no definite seasonal molt seems apparent. 

VARIATION 

Variation in the raccoons is assignable to several categories, of 
which perhaps the most obvious are geographic and individual. 

GEOGRAPHIC VARIATION 

The raccoons are believed to intergrade throughout the vast range 
of the species Procyon lotor on the North American mainland, and the 
component subspecies are the expression of geographic variation in 
size, weight, color, and minor details of structure in response to 
environmental and genetic influences. Some of the insular forms 
present a greater degree of differentiation, evidently due to isolation, 
and are regarded as distinct species. The largest form of the genus, 
Procyon lotor excelsus, inhabits interior valleys, principally the Snake 
River Valley in southeastern Washington, eastern Oregon, and 
southern Idaho. Large, but less extreme, geographic races occupy 
the other Western States and the mainland of Middle America. 
These give way to smaller subspecies in the eastern United States, 
the minimum size being reached by those living on the Florida Keys. 
Maria Madre and Maria Magdalena of the Tres Marias Islands 
Group off western Mexico are occupied by Procyon insularis, a large 
form regarded as specifically distinct from the mainland animal. 
Small species of raccoons inhabit New Providence Island in the 
Bahamas, and Guadeloupe and Barbados Islands of the Lesser 
Antilles. The smallest species of raccoon known was well named 
Procyon pygmaeus from Cozumel Island, Yucatan. 

Geographic variation in color in the raccoons is limited mainly to 
the general tone and to the relative development of the black mask 
and other facial markings. The paler subspecies, such as Procyon 
lotor -paUklus, inhabit the thinly timbered desert areas in the Colorado 
River Valley and adjoining territory, while darker races have de- 
veloped in the eastern United States and in densely forested regions 
of heavy precipitation in Central America. In considering the pallid 
coloration of raccoons from desert areas, as along the Colorado River, 
it should be understood that these animals are restricted to the 
vicinity of water, yet they share the general pallor that is a marked 
characteristic of the mammals of the region as a whole. 

IINDIVIDUAL VARIATION 

By individual variation reference is made to all the degrees of 
divergence from a typical mean exhibited by large series of conspecific 
skins and skulls from any given locality. In the raccoons the range 
of this variation in size, color, and cranial details is about the same as 



RACCOONS OF NORTH AND MIDDLE AMERICA 21 

that for which due allowance must be made in other groups of carni- 
vores. Since the subspecies of Procyon lofor are geographic races 
with confluent geographic ranges, an unusually large individual of a 
small form may be similar in size to an unusually small individual of 
a large form, and color and cranial details may vary in comparabk' 
ratio. Owing to individual variation, some specimens, especially 
from unknown localities, may be difficult to identify subspecifically. 
They may usually be distinguished, however, by the combination of 
characters presented. 

Apparently, abnormal individual variations in general color are 
common in the raccoons. A half-grown example (No. 253823, U. S. 
Natl. Mus.) from Nelson County, Va., exhibits an apparent case of 
erythrism. The usual black facial mask, postauricular spots, dark 
bands on the tail, and the normally dark tips of hairs are light yel- 
lowish Ijrown; the usual white areas tend toward creamy white. In 
a specimen from Santee, S. C. (No. 178391, U. S. Natl. Mus.), the 
usual black tips of the hairs over the back and the dark tail rings are 
light brownish. The dark facial markings are also inclined toward 
brown instead of black, and the basal color of the fur in general is 
lighter than normal. 

No definiteh^ melanistic raccoons have been examined, but in a 
specimen from Red Bluff, Cahf. (No. 14466, U. S. Natl. Mus.), there 
is an intensification of the overlying black on the upper parts, due to 
the unusual extent of the black on the tips of the hairs and the cor- 
responding reduction of the light sul)apical zone on these hairs. As 
a result the back appears to be almost solid black. The usual light 
bars are present on the tail, and the white facial markings are normal. 
The occurrence of albino raccoons is reported from time to time. 
An adult male from Paducah, Ky. (No. 151657, U. S. Natl. Mus.), is 
pure white except on the nape where the white is suffused with 
yellowish. 

The weight of the northern raccoons undoubtedly varies consider- 
ably according to the season; the animals become very fat in the fall, 
especially in regions where they must hibernate. The more southern 
raccoons that are active throughout the year do not accumulate so 
great a store of fat, and even their shorter pelage would weigh less. 
The weight diff'ers, of course, in accordance with size in animals of 
comparable age, sex, and condition in the various species and sub- 
species. Comparatively few weights, however, appear to have been 
reliably recorded and are available for comparison. Whitney (1931, 
p. 31) reports the taking of more than 300 raccoons (Procyon lotor 
lotor) in Massachusetts and Connecticut during a 7-year period. 
Of the 300, every one that appeared to be uncommonly large was 
weighed on accurate scales. The largest weighed 22 pounds and 10 



22 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

ounces. Wliitney believed, however, that the average would be 
about 1.3 pounds as the weis-hts included those of a good many animals 
taken in the fall that had been born in the spring of the same year 
and had not been able to attain a weight of much more than 10 
pounds. Eighteen raccoons regarded by Wliitney as of uniformly 
greater weight were taken by him in the fall near Brunswick, Maine. 
Eight of these weighed more than 23 pounds each, the largest, 27 
poimds, including, a sack estimated to weight three-fourths of a 
pound. 

A large, fat, adult male raccoon {Procyon lotor hirtus) collected by 
Vernon Bailey (1923, p. 124) at Elk River, Minn., November 5, 
1886, was recorded by him as weighing 30)^ pounds. Another male 
of the same subspecies taken at Fargo, N. Dak., by O. J. Murie, 
November 9, 1919, weighed 24 pounds. Average animals from the 
same localities would undoubtedly weigh much less. 

Weights of specimens of Procyon lotor elucus, which is active through- 
out the year and does not l)ecome so fat as the more northern sub- 
species, were obtained in winter by E. A. Mearns on Saw Grass 
Island, Catfish Creek, Polk County, Fla. Five adult males from the 
island ranged from 10 to 12 pounds in weight, the average being 11 
pounds. The weight of three adult females from the same locality 
ranged from 7.7 to 10 pounds, the average being 9 pounds. Weights 
of the diminutive raccoon Procyon lotor auspicatus, of Key Vaca, 
one of the Florida keys, were obtained late in winter by E. W. Nelson. 
Five adult males were found by him to range in weight from 4 to 6 
pounds, with an average of 5.3 pounds. Two adult females from the 
same locality weighed 4 and 5 pounds, respectively. 

Individual variation in cranial and dental development is extensive 
in scope and may render difficult the determination of some speci- 
mens, especially if from unknown localities. The variations are 
noticeable especially in the form of the brain case and frontal profile, 
relative prominence of post orbital processes, size of auditory bullae, 
and size of large molariform teeth. Dental abnormalities are pre- 
sented in a few cases. In two individuals, one of Procyon lotor 
litoreus from Saint Simon Island, Ga., and the other of Procyon 
gloveralleni of Barbados, Lesser Antilles, the first premolars in both 
jaws are absent. Supernumerary teeth sometimes suggest early 
division of the dental matrix. In a skull of Procyon lotor hernandezii 
from Colima two canines are present on one side in the upper jaw, one 
somewhat smaller being posterior to the normal canine in the space 
usually occupied bv the first premolar which is absent. On the side 
opposite the double canines the first premolar is also absent, there 
being a hiatus l)otween the canine and second premolar. The 
mandible is normal. 



RACCOONS OF NORTH AND MIDDLE AMERICA 23 

EXPLANATIONS 

MEASUKEIVIENTS 

All measurements of specimens are in millimeters. The weights 
given are in ])()vm(ls. Adult males usnally exceed adult females in 
dimensions, and the measurements are, therefore, ])resented according- 
to sex. In some cases so few nearly typical examples are available 
that the measurements given may not represent the normal range 
of individual variation, and too broad generalizations, therefore, 
should not be based on them. 

The external measurements, unless otherwise stated, were taken 
in the flesh by the collector, as follows: 

Total length. — Nose to end of terminal tail vertebra. 

Tail vertebrae. — Upper base of tail to end of terminal vertebra. 

Hind foot. — Heel to end of longest claw. 

The following cranial measurements were taken with a vernier 
caliper l)y tlie author: 

Greatest length. — Length from anterior tip of premaxillae to supra- 
occipital in median line over foramen magnum. 

fondylobasal length. — Length from anterior tip of premaxillae to 
posterior plane of occipital condyles. 

Zygomatic breadth. — Greatest distance across zygomata. 

Interorbital breadth. — Least distance between orbits. 

Least -width of pedatal shelf. — Width l)etween outer sides of palate 
at constriction behind posterior molars. 

Maxillary tooth row. — Distance from fi-ont of canine to back of 
posterior molar at alveolar border. 

Crown length of wpper carnassial. — Greatest length of crown of upper 
carnassial along outer side. 

Crown wielth of upper carnassial. — Greatest transverse diameter of 
crown of upper carnassial. 

COLORS 

Owing to the banding of the individual hairs, raccoons present 
coarsely blended colors difficult to segregate. For this reason very 
limited use has been made, in quotation marks, of names of colors 
from Ridgway's "Color Standards and Color Nomenclature, 1912." 
In the description of colors generally understood, modifying or com- 
parative terms have been employed in naming tones, many of which 
are not well defined. 

SPECIMENS EXAIMINED 

Specimens examined, unless otherwise indicated, are in the collec- 
tions of the United States National Museum, including the Biological 
Surveys collection. 



24 NORTH AAIERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

USE OF KEY TO SPECIES AND SUBSPECIES 

The key to the species and subspecies of the subgenus Procyon is 
based largely on the geographic ranges, as trenchant characters 
cannot be assigned to intergrading geographic races, and most of 
the insular forms treated as species are imperfectly known. The key 
supplements the map (fig. 1) in affording a clue to the identification 
of particular specimens from known localities. 




Fk; 

1. P. 

2. P. 

3. P. 

4. P. 

5. P. 

6. P. 

7. P. 

8. P. 

9. P. 

10. P. 

11. P. 



IRE 1. — Distribution of species and subspecies 



lutor lotor. 
I. hirtii.s. 
L varius. 
I. litoreu--<. 
I. solutu.-i. 
I. eluciis. 
I. mar inns. 
I. inesperatns. 
!. auspirntus. 
I. incaiitiis. 
I. fuscipes. 



12. P. I. )nexicanus. 

13. P. I. pallidum. 

14. P. I. psora. 

15. P. I. pacificus. 

16. P. I. excelsjis. 

17. P. I. vancouverensis. 

18. P. I. grinuelli. 

19. P. I. hernandezii. 

20. P. I. shufcldti. 

21. /''. /. dickct/i. 

22. P. I. crassideas. 



of the subgenus Procyon: 

23. P. I. pumilus. 

24. P. insnlaris insularis. 

25. P. i. vicinus. 

26. P. niaynardi. 

27. P. pygniaeus. 

28. P. minor. 

29. P. gloveralleni. 

30. P. I. )naritimus. 

31. P. I. megalodous. 



RACCOONS OF NORTH AND MIDDLE AMERICA 25 

Genus PROCYON Storr 

Procijon Storr, Prodr. Meth. Mammal., p. 35. 1780. Type Ursus lotor Linnaeus. 

Campsiurus Link, Beytr. Naturg. 1 (2): 87, 1795. Type Ursus Intor Linnaeus 
(see Hollister, p. 146, 1915). 

Lotor Geoffrey and Cuvier, Mag. Enc. 2: 187, 1795. 

Lotor Oken, Lehrb. Naturg., S'^-- Theil., 2t<= Abth., p. 1080, 1816. 

Euprocyon Gray, Zool. Soc. London Proc. 1864: 705 (subgenus). Type Ursus 
cancrivorus Cuvier. 

Moinprocyonus Herrera, Sin. Vulg. Cient. Vert. Mexicanos 1899: 18. 

Euprocyon Goldman, Smithsn. Misc. Coll. 60 (22): 16, Feb. 28, 1913 (genus). 

Euprocyon Hollister, U. S. Natl. Mas. Proc. 49 (2100): 146, Aug. 13, 1915 (sub- 
genus) . 

Distribution. — Southern Canada to southern Brazil and northern 
Argentina, and some of the outlying islands. 

Generic characters. — Form robust; head broad, with short, pointed 
muzzle ; ears medium-sized, pointed ; upper lip hairy across median line ; 
soles of feet naked, smooth, without well-developed digital pads; digits 
free, very long, tlip first more than half the length of the second; claws 
nonretractile; tail shorter than body, cylindrical, distinct!}^ annulated; 
baculum long, curved and bilobed distally; mammae 6, arranged in 
three pairs, as follows: pectoral, 2; abdominal, 2; inguinal, 2. 

Skull broad and massive; rostrum broad; brain case broad posteri- 
orlv, tapering gradually anteriorly; interorbital and postorbital con- 
strictions mock'rate; postorbital processes of maxillae usually more 
developed than postorbital processes of frontals; sagittal crest high 
and trenchant in some old adults, absent in others, the temporal ridges 
not uniting along median line. Alastoid processes long, stout, strongly 
everted, rounded distally; hamular processes rounded, with knob-like 
ends; auditory bullae large, inflated on inner side, the outer side 
sloping gradually to external auditory meatus. Mandible heavy, 
inferior border evenly rounded; symphysis short; coronoid process 
rising high and curving backward over condyle. 

Dental formula: i 3/3 c 1/1 j^m 4/4 m 2/2 = 40. 

Dentition heavy; molar crowns moderately high, with prominent 
cusps; first and second upper premolars simple unicuspids; third upper 
premolar with a high conical principal cusp and a postero-internal 
shelf-like cingulum sometimes bearing a small cusplet ; crown of fourth 
upper premolar sul)quadrate, about as long as broad, with five prin- 
cipal cusps; crown of first upper molar usually slightly broader than 
long, with four princi])al cusps; second upper molar subtriangular, with 
three principal cusps; crown of first lower molar elongated, sub- 
rectangular, with five distinct cusps. First upper premolar with a 
single root; second and third upper premolars 2-rooted; fourth upper 
premolar 3-rooted. Incisors heavy, the crowns more or less distinctly 
grooved when unworn. Canines oval in cross section at alveoli, 

870119°— 50 3 



26 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

conical, without distinct grooves, the upper canines not strongly 
everted. 

Eejnarks. — The genus Procyon is readily distinguished from the other 
Hving genera of Procyonidae. It is most closely allied to Nasua, but 
differs strikingly in external appearance as well as internal structure. 
It shares with Nasua the general pattern of white and black facial 
markings, hairy mid-section of upper lip, and annulated tail, but 
departs in more robust form, shorter snout, pointed ears, free digits, 
short front claws, and short, cylindrical instead of tapering tail. 
Important similarities in cranial structure and dental details, espe- 
cially the molar cusps, are apparent, l^ut the skull diverges notably in 
its short and broad, instead of narrow and elongated outlines. Among 
other cranial characters that distinguish Procyon from Naxua are the 
greater breadth of the palate between the molars in relation to breadth 
of bony palate behind molars (palate nearly parallel-sided throughout 
its length in Nasua) ; upper molariform tooth rows curved posteriorly, 
instead of being nearly straight; mastoid processes much more promi- 
nent; and canines more rounded and conical, instead of flattened and 
saber-like, with trenchant anterior and posterior edges. The genus 
Procyon differs from the genus Nasuella in about the same characters 
as from Nasua. 

The genus Procyon is more distantly related to the genus Bas- 
saricyon which it approaches in general type of dentition, but with 
which it contrasts strongly in color and in many important structural 
details. Procyon is a much larger, more heavily built animal than 
Bassaricyon, which also exhil)its a departure in color, pelage, and other 
external features. In Bassaricyon the color is more uniform, the face 
somewhat grayish but lacking the black mask and white markings of 
Procyon and the general body color ochraceous tawny. The general 
pelage is much denser, softer, and has a silky quality very unlike that 
of Procyon. The tail is longer in Bassaricyon, flattened instead of 
cylindrical, and is indistinctly annulated. The ears are more rounded 
in Bassaricyon than in Procyon, and a median projection of the 
rhinarium extends across the lip to the mouth. The skulls of Procyon 
and Bassaricyon are somewhat similar in general form, but dift'er in 
many important features. Contrasted with that of Bassaricyon, the 
skull of Procyon presents points of difl'erence including the following: 
Much larger, more massive (thin-walled and delicate in Bassaricyon) ; 
brain case less inflated; orbits relatively smaller; postorbital processes 
much less, and mastoid processes much more, developed; canines 
without distinct grooves (canines with two distinct longitudinal 
grooves on inner, and two on outer, surfaces in Bassaricyon). 

Compared with the genus Pofos, the most aberrant American mem- 
ber of the family as currently recognized, the genus Procyon dift'ers so 



RACCOONS OF NORTH AND MIDDLE AMERICA 27 

widely that the commonly accopterl family alignment seems open to 
some question. Poios contrasts strongly in nearly uniform coloration, 
rounded ears, and long, tapering, short-haired, prehensile tail. The 
rhinnrium in Potos traverses the upper lip more as in BasHaricyon. In 
the skull of Potos similarly striking contrasts with Procyon are evident. 
The teeth may be regarded as somewhat similar in general form hut 
there the resemblance ends. The molar crowns in Potos are lower and 
much simpler than in Procyon, Ix'ing nearly flat and without well- 
developed cusps at any age, the posterior molars almost completely 
opposed, above and below. Anterior premolars, present in Procyon, 
are absent in Potos, a condition correlated with the shortening of the 
rostrum in tlie latter genus. The canines, normally rounded and 
without distinct grooves in the adult stage in Procyon, are flattened and 
saber-like, with deeply grooved sides in Potos. The mandible in Potos 
is remarkable for its depth and long, early -fused symphysis, the space 
between the rami anteriorly U-shaped instead of V-shaped, as is usual 
in the group. The lower border of the ramus, convex in Procyon, is 
concave in Potos, owing to lateral compression and downward expan- 
sion of the angle. Among other differential cranial features of Potos 
are the parallel-sided palate, peculiar flat bullae, and complete absence 
of the mastoid processes so well developed in Procyon. 

The genus Procyon requires no close comparison with the Old World 
procyonid genera Ailurus and Alluropoda of the subfamily Ailurinae. 
The characters of the Old World genera and their relationship to the 
other procyonids have recently been discussed by Gregory (1936) and 
by McGrew (1938). Among other important references bearing on 
the classification of the Procyonidae are Hollister (1915), and Pocock 

(1921). 

KEY TO SUBGENERA 

a'. Pelage of two kinds, long guard hairs and short, soft underfur; hair on nape 
normal, not directed forward; palate extending behind posterior molars a 
distance of more than one-fourth length of palate Procyon (p. 28) 

a^. Pelage coarse and wiry, without underfur; hair on nape directed forward; 
palate extending behind posterior molars a distance of less than one-fourth 
length of palate Eii procyon (p. 80) 

LIST OF NORTH AMERICAN SPECIES AND SUBSPECIES, WITH TYPE 

LOCALITIES 

Subgenus PROCYON Storr 

Procyon lotor lotor (Linnaeus) Pennsylvania (p. 33). 

lotor hirtus Nelson and Goldman Elk River, Minn. (p. 37). 

lotor varius Nelson and Goldman Castleberry, Ala. (p. 38). 

lotor litoreus Nelson and Goldman Saint Simon Island, Ga. (p. 40). 

lotor solutiis Nelson and Goldman Hilton Head Island, S. C. (p. 41). 

lotor el veil s Bang?. Oak Lodge, Brevard County, Fla. 

(p. 42). 
lotor niarinus Nelson Chokoloskee, Fla. (p. 44). 



28 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

LIST OF NORTH AMERICAN SPECIES AND SUBSPECIES, WITH TYPE 
LOCALITIES— Continued 

Subgenus PROCYON Storr— Continued 

Procxjon lotor inesperatus Nelson Upper Matecumbe Key. Fla. (p. 46) . 

lotor auspicatus Nelson Marathon, Key Vaca. Fla. (p. 47). 

lotor incautus Nelson Torch Key, Fla. (p. 48). 

lotor Juscipes Mearns Fort Clark, Tex. (p. 49). 

lotor mexicanus Baird Espia, Chihuahua, Me.xico (p. 52). 

lotor pallidus Merriam New River, Colorado Desert, Calif. 

(p. 54). 

lotor psora Gray Sacramento, Calif, (p. 56) . 

lotor pacificus Merriam Lake Keechelus, Wash. (p. 58). 

lotor excehus Nelson and Goldman „ ^ . Owyhee River, Greg. (p. 60). 
lotor vancouverensis Nelson and Gold- Vancouver Island, British Colum- 
man. bia (p. 61). 

lotor qrinnelli Nelson and (ioldinan La Paz, Baja Calif, (p. 62). 

lotcr hernandezii Wagler_ Valley of Mexico, Mexico (p. 64). 

/o/o/s/(M/e/rfh' Nelson and Goldman La Tuxpeiia, Campeche, Mexico 

(p. 65). 

lotor dickeyi Nelson and Goldman Barra de Santiago, Salvador (p. 67). 

lotor crassidens Hollister Talamanca, Costa Rica (p. 69). 

lotor pumilus Miller Ancon, Panama (p. 70). 

insiilaris insularis Mevvisim Maria Madre Island, Tres Marias 

Islands, Mexico (p. 72). 
insularis viciniis "Selson SLud Gokhnan _ Maria Magdalena Island, Tres 

Marias Islands, Mexico (p. 73). 
»iay/(orr// Bangs New Providence Island, Bahamas 

(p. 75). 

pygmaeus Merriam Cozumel Island, Yucatan (p. 76). 

minor Miller Pointe-a-Pitre, Guadelou])e Island, 

Lesser Antilles (p. 77). 
(7/ot'e;-a//c/;i Nelson and Goldman Island of Barbados, Lesser An- 
tilles (p. 79). 

Subgenus EIJPROCYON Gray 
Procyon caiicrivoniN panainensis (Goldman).^ Gatun, C. Z. (p. 82). 

Subgenus PROCYON Storr 
[Rpfprences under Gonus Procyon .Storr, \). 2r\] 

Distribution. — Nearly transcontinental from southern Canada to 
Panama; occurring also on some of the outlying islands. 

Sub<ieneric characters. — Contrasted with the subgenus Euprocyon: 
Pelage longer and of two kinds — long coarse guard hairs and short, 
soft underfiu-; hair on nape normal, not directed forward; claws nar- 
rower, more compressed laterally, of greater vertical depth at base, 
and more sharply pointed. Bony palate extending behind posterior 
molars a distance of more than one-fourth the total length of palate. 
Molariform teeth, except first premolars, smaller, with narrower, 
more sharply pointed cusps ; connecting ridges between principal cusps 
higher, more trenchant. 



RACCOONS OF NORTH AND MIDDLE AMERICA 29 

Remark'^. — The subgenus Procyon overlaps the range of the sub- 
genus Ewprocyon in Panama, but the characters pointed out are quite 
distinctive. 

Key to Species and Subspecies of the Subgenus PKOCYON 

ITypical adults] 
rt'. Geographic range continental. 
h^. Geographic range eastern Uniteci States and sonthern Canada, west to near 
longitude 90°. 
c^. Size smaller; hind foot usually less than 120 nun.; geographic rang(» excluding 
greater part of Florida. 
d^. Color darker; geographic range southeastern Canada and the North- 
eastern States, mainly north of latitude 35° .P. /. lotor (p. 33). 

d'-. Color paler; geographic range Southeastern States, mainly south of 

latitude 35° P. I. varius (p. 38). 

C-. Size larger; hind foot usually more tlian 120 mm.; geographic range greater 

part of Florida P. I. elucus (p. 42). 

b-. Geographic range not including eastern United States far beyond longitude 

90°. 

c^. Geographic range mainly east of longitude 105°, and north of latitude 22°. 

(/'. Color darker, more suffu.sed with buff; pelage longer; geographic range 

mainly upper Mississippi and Missouri River drainage__P. I. hirtus (p. 37). 

d'. Color paler, more suffused with gray; pelage shorter; geographic range 

mainly Texas and northeastern Mexico P. I. fuscipes (p. 49). 

C-. Geographic range not mainly east of longitude 105° and north of latitude 22°. 
d^. Geographic range mainly west of the Sierra Nevada and Cascade Range. 
gi. Color darker; geographic range mainly southwestern British Columbia, 

western Washington, and western Oregon P. I. pacific us (p. 58). 

e-. Color paler; geographic range mainly California P. I. psorn (p. 50). 

d^. Geographic range not mainly west of the Sierra Nevada and Cascade 
Range, 
gi. Size larger; geographic range mainly Snake and Humboldt River drainage 

in Idaho, Oregon, and Nevada P. I. excelsus (p. 60). 

e^. Size sinaller; geographic range not including Snake and Humboldt River 

drainage in Idaho, Oregon, and Nevada. 
p. Color paler; geographic range Colorado River drainage 

P. I. paUidus (p. 54). 
/-. Color darker; geographic range not including Colorado River drainage. 

gK Cieographic range southern Baja California P. I. grinnelli (p. 62). 

g-. Geographic range not including southern Baja California. 
h^. Geographic range mainly north or west of Isthmus of Tehuantepec. 
i^. Color paler; geographic range northwestern Mexico south to about 

latitude 22° P. I. mexicanus (p 52) . 

i^. Color darker; geographic range high tableland and coastal regions of 
Mexico from about latitude 22° to Isthmus of Tehuantepec 

P. I. hernandezii (p. 64). 
h^. Geographic range mainly south or east of Isthmus of Tehuantepec. 
iK Color paler; geographic range mainly north of latitude 14°, including 

Yucatan Peninsula P. I. shiifeldti (p. 65). 

i^. Color darker; geographic range mainly south of latitude 14°. 
ji. Skull less massive; known geographic range southwestern coast of 
Salvador P. I. dickei/i (p. 67). 



30 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

j2. Skull more massive; known geographic range excluding southwestern 
coast of Salvador. 
k^. Skull longer and narrower; dentition heavier; known geographic 
range Costa Rica, Nicaragua, and Honduras 

P. I. crassidens (p. 69). 
k^. Skull shorter and broader; dentition lighter; known geographic 

range Panama P. /. pumilus (p. 70). 

a^. Geographic range insular (at least in part). 
b^. Geographic range off Pacific coast. 

cK Georgaphic range Vancouver Island P. I. vancouverensis (p. 61). 

C-. Geograjjhic range off west coast of Mexico [P. insularis and subspecies]. 
dA. Color paler; geographic range Maria Madre Island, Nayarit 

P. i. insularis (p. 72). 
d'. Color darker; geographic range Maria Magdalena Island, Nayarit 

P. i. vicinus (p. 73). 
62. Geographic range off Atlantic coast (at least in part), 
c^ Geographic range far offshore islands (Bahamas and Lesser Antilles), 
f/i. Cleograjihic range Bahama Islands (New Providence Island) 

P. maynardi (p. 75). 
d'^. Geographic range Lesser Antilles, 
gi. Color darker; upper carnassial longer than broad; geographic range 

Barbados Island P- gloveralleni (p. 79). 

e-. Color paler; upper carnassial shorter than broad; geographic range 

Guadeloupe Island P. minor (p. 77). 

c^. Geographic range coastal islands (at least in part). 
d^. Size larger; hind foot more than 90 mm.; geographic range southern 
P'lorida Keys and islands and coasts of Georgia and South Carolina, 
e^ Geographic range southern Florida Keys (at least in part). 
/I. Color darker; geographic range very close to or extending to Florida 
coast. 
g^. Size larger; total length (adult male) more than 700 mm.; geographic 
range Upper Matecumbe and other keys near base of main Florida 

chain P- /■ inesperatus (p. 46). 

gS. Size smaller; total length (adult male) less than 700 mm.; geographic 
range keys of Ten Thousand Islands group and adjacent coast 

P. /. uiarinns (p. 44). 

p. Color paler; geographic range outer half of Florida Keys. 

g^. Size larger; hind foot (adult male) more than 110 mm.; palatal bridge 

extending on median line beyond plane of last molars more than 12 

mm.; geographic range Big Pine Key group, near extreme end of 

Florida chain P. /. incautus (p. 48). 

g-. Size smaller; hind foot (adult male) less than 110 mm.; palatal bridge 
extending on median line beyond plane of last molars less than 12 

mm.; geographic range Key Vaca P. /. auspicatiis (p. 47). 

e'^. Geographic range islands and coasts of Georgia and South Carolina. 
p. Dentition heavier; crown length of upper carnassial usually more than 
9 mm.; geographic range Saint Simon Island, neighboring islands, and 

coast of Georgia P. /• Utoreus (p. 40). 

p. Dentition lighter; crown length of upper carnassial usually less than 
9 mm.; geographic range Hilton Head Island, neighboring islands, and 

coast of South Carolina P. /• solutus (p. 41). 

d2. Size smaller; hind foot about 90 mm. or less; geographic range Cozumel 
Island, east coast of Yucatan P. pygniaeus (p. 76). 



RACCOOXS OF NORTH AND :MIDDLE AMERICA 31 

PROCYON LOTOR GROUP 

Distribution. — Transcontiiiental (except in parts of the Rocky 
Mountain region) from southern Canada to Panama, and ishxnds as 
far distant as the Tres Marias off the west coast of Mexico and the 
Bahamas and Lesser Antihes, West Indies. Altitudinal range is from 
sea level up along streams to about 5,000 feet in parts of the Rocky 
Mountain region (a few animals reaching as high as 8,500 feet eleva- 
tion), and to more than 9,000 feet in the mountains near Ajusco south 
of the Valley of Mexico. It occupies the Tropical, Austral, Transition, 
and lower part of Canadian Zones. 

(liaracters. — Contrasted with Procyon cancrivorus and related forms: 
Pelage of nape inclined backward; pelage consisting of two distinct 
kinds of hairs — soft, dense, velvety underfur, and longer, stift'er, 
projecting overlying hairs; throat and postauricular areas blackish; 
cusps of larger molariform teeth relatively high and trenchant, with 
distinct connecting ridges. 

Eemarks. — The Procyon lotor grou]) includes P. lotor and subspecies 
of the mainland from Canada to Panama and closely adjacent islands. 
To the group may also conveniently be referred several more distant 
insular forms regarded as specifically distinct, but closely allied, as 
shown by similarity in important characters. These inhabit the Tres 
Marias Islands off the west coast of Mexico, Cozumel Island off Yuca- 
tan, and several rather widely separated islands of the West Indies. 
How the particular West Indian islands now inhabited were reached 
by raccoons and why these animals do not occur on many other 
islands of the archi])elago where conditions seem similarly suitable are 
interesting subjects for speculation. Sloane (1725, p. 329) referred to 
the occurrence of the animal in Jamaica, as follows: "The Raccoons 
are commonly here in the Mountains, and live in hollow fiddlewood 
Trees, from whence they make paths to go to seek Sugar Canes, which 
is their chief, if only Sustenance." As there appear to be no later 
records and as Sloane referred vaguely to various authors who described 
raccoons elsewhere, he probably confused Jamaica with some other 
island. 

The members of the group as a whole differ among themselves in 
tone of coloration, but the pattern of color markings is essentially the 
same, and all forms are much alike in general external appearance. 
They require close comparison as a group only with the crab-eating 
raccoon, Procyon {Euprocyon) cancrivorus, the range of a northern 
representative of the latter being overlapped in Panama. The char- 
acters that have been mentioned, however, readily separate the two 
groups. 



32 NORTH AAIERICAX FAUNA 60. FISH AND WILDLIFE SERVICE 

PROCYON LOTOR (Linnaeus) 

[Synoiiyiiiy unilcr subspecies] 

Disfribution. — Transcontinental (except in the Rocky Mountain 
region) from southern Canachi to Panama, and islands near mainland 
coast. The altitudinal and zonal ranges have been given under the 
distribution of the various subspecies. 

General characters. — Size variable, general color of upper parts 
ranging from iron grayish to blackish, more or less suffused with 
ochraceous buff, especially on nape and lighter caudal rings; pelage 
long, full, and soft in the northern subspecies, much shorter, thinner, 
and stiff er in the more southern forms. Similar in external appearance 
to, but differing in cranial characters from, distant insular animals 
recognized as belonging to distinct species. 

Color. — Upper parts in general varying from iron grayish to blackish, 
more or less suffused with bufT, rusty, or "orange rufous," especially on 
nape, the general tone depending much upon the relative width and 
distribution of light subapical })ands and black tips of long hairs; 
dorsum more or less heavily overlaid with black, tending to thin out 
and become grayer along sides; top of head varying mixtures of l:)lack 
and white or gray, producing a grizzled effect; face with a sharply 
delimitefl black mask usually reaching from cheeks across eyes and 
muzzle, with median extensions downward to rhinarium and upward 
on forehead, more or less interrupted between the eyes, however, in 
some forms; facial mask bordered above by conspicuous white lines 
extending from near middle of forehead backward under ears or to 
sides of neck; sides of muzzle, lips, and chin white; tufts of stiff, 
whitish vibrissae 50 to 100 millimeters in length, arising from sides of 
muzzle, and smaller, less cons])icuous tufts arising, one on each side 
over eyes and sides of cheeks; under parts, in general, thinly overlaid 
with long grayish or buffy over hairs, only partially concealing the 
dense brownish underfur; throat crossed by a distinct blackish or 
brownish area, separated from facial mask by narrow white lines 
extending posteriorly from muzzle; ears clothed with short grayish or 
buffy hairs, with black areas varying in size and distinctness at 
posterior base; forearms and thighs similar to under parts, but hind 
legs more or less distinctly blackish near heels; fore feet whitish; hind 
feet usually whitish, but dusky of ankles sometimes extending down 
on metatarsus; toes of hind feet with a few grayish or dusky bristles 
usually extending beyond ends of longest claws ; tail above with five to 
seven conspicuous black rings and a black tip, alternating with broader 
grayish or buffy rings, the black rings less sharply defined and some- 
times interrupted below. 



RACCOONS OF NORTH AND MIDDLE AMERICA 33 

Remarks. — Procyon lot or is divisible into 25 geographic races whicli 
on the mainland form a closely intergrading series. The species 
attains its largest size in P. I. excelsus of the Snake River Valley in 
southeastern Washington, eastern Oregon, and southern Idaho, and 
the smallest forms are from the Florida Keys. The palest subspecies 
inhabit the hot arid delta of the Colorado River and adjoining regions, 
and the darkest have developed in the regions of heavy precipitation 
in Central America. 

PROCYON LOTOR LOTOR (Linnaeus) 

Eastern Raccoon 

[f//-s?/s] lotor Linnaeus, Syst. Nat. (ed. 10) 1: 48, 1758. 

[Meles] lotor Boddaert, Elenchus Animal 1 : 80, 1784. "Habitat in America." 

L[oior] vulgaris Tiedemann, Zoologie. Zu seinen Vorlesungen entworfen, erster 

band, Mensch und Saugthiere, p. 380, 1808, (part). From North America, 

Mexico, and the Antilles. 
Procyon annulatus G. Fischer, Zoognosia 3: 177, 1814 (part). "Habitat in Ameri- 

cae maritimis." 
Procyon lotor Illiger, Abhand. Konig Akad. Wissensch. Berlin, 1804-1811, pp. 

70, 74, 1815. 
Procyon gularis Hamilton Smith, Jardine's Nat. Lib. 15: 222, 1848. From State 

of New York. 

Type locality. — Pennsylvania. ^ 

Ty2)e. — Not known to exist. 

Distribution. — Nova Scotia, southern New Brunswick, southern 
Quebec, and southern Ontario south through the eastern United 
States to North Carolina, and from the Atlantic coast west to Lake 
Michigan, Indiana, southern Illinois, western Kentucky, and probably 
eastern Tennessee. Lower Austral to Canadian Zones. 

General characters. — A rather small, dark form with long, full, soft 
pelage; skull with moderately h"gh, narrow frontal region, and weak 
or obsolescent postorbital processes. Similar to P. I. hirtus of Minne- 
sota, but much smaller; pelage less extremely long, and less suffused 
with ochraceous buff; skull smaller. Differs from P. I. soliitus of 
Hilton Head Island and the coastal region of South Carolina in darker, 
less grayish coloration, more elongated skull, and other cranial 
details. Resembles P. I. litoreus of Saint Simon Island and the coastal 
region of Georgia, but pelage longer and softer, and cranial characters, 
especially the much smaller molariform teeth, distinctive. Mtich 
like P. I. varius of Alabama, but larger, usually darker, and pelage 
much longer; skull larger and of heavier proportions. 

Color. — Upper parts, in general, varying shades of buffy grayish 
(becoming ochraceous buff or rusty rufous on nape and across shotd- 

-Type locality fixed by Thomas, Proc. Zool. Soe. London, 1911, p. 140. 



34 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

ders in some individuals) overlaid with black, the general tone due 
mainly to black-tipped hairs with a lighter subterminal zone, the dark 
brownish underfur showing through to some extent; sides of body 
somewhat lighter, the black tips of hairs shorter or black-tipped hairs 
less numerous than on median dorsal area; top of head mixed black 
and white or grayish, giving a coarsely grizzled effect; black mask 
enclosing eyes, but more or less discontinuous on middle of face where 
a blackish median line is more or less distinctly isolated by lighter 
lateral lines; upper surface of muzzle usually brownish; facial mask 
bordered above by rather broad and conspicuous whitish lines extend- 
ing posteriorly across cheeks to sides of neck; sides of muzzle, lips, 
and chin white; under parts, in general, thinly overlaid with long 
grayish or bufiy overhairs only partially concealing the dense under- 
fur, wdiich varies from near wood brown to chestnut bown; throat 
crossed by a brownish or blackish area, separated from facial mask 
laterally by narrow whitish or buffy lines extending posteriorly from 
muzzle; ears densely clothed inside and out with short, whitish or 
buffy hairs, mei-ging with the general pelage on external basal portion; 
black postauricular patches usually large and conspicuous; forearms 
and thighs similar to under parts, but hind legs more or less distinctly 
blackish near heels; fore and hind feet, including toes, wdiitish, the 
soles black and naked; tail above with five to seven narrow black 
rings and a black tip, alternating with broader grayish or ochraceous 
buffy rings, the black rings less sharply defined and sometimes inter- 
rupted below. Young (in first pelage) : Color markings as in adults, 
but top of head, nape, and postauricular spots nearly pure brownish 
black, in contrast to the lighter, generally buffy, tone of dorsum, over 
which the black-tipped hairs beginning to appear are still incon- 
spicuous. 

Cranial characters. — Skull rather small, with moderately high, nar- 
row frontal region; brain case depressed near fronto-parietal suture; 
postorbital processes of frontals small or obsolescent; postorbital 
processes of jugal well developed. Very similar to that of P. I. hirtus, 
but much smaller, less massive; sides of frontals behind orbits usually 
more deeply indented or constricted, the result being that sides of 
brain case are more rounded or bulging, less tapering anteriorly; 
dentition relatively the same. Not very unlike that of P. I. solutus, 
but longer and relatively narrower; frontal region usually narrower; 
palatal shelf longer, extending farther posteriorly beyond posterior 
molars; dentition usually somewhat lighter, the large molariform teeth 
rather narrow, but maxillary tooth row longer as a rule, owing to wider 
spacing of premolars. Similar in general form to that of P. I. litoreus, 
but smaller, and dentition relatively much lighter, the difference most 
marked in the molariform teeth. Compared with that of P. I. Darius, 



RACCOONS OF NORTH AND MIDDLE AMERICA 35 

the skull of p. 1. lofnr is larger and heavier; jiigal broader; sides of 
front als behind orbits usually more deeply indented or constricted; 
maxillary tooth row longer; posterior upper premolar and upper 
carnassial usually distinctly larger. 

Measurements. — Adult female from Liberty Hill, Conn.: Total length, 8.32 milii- 
nicters; tail vertebrae, 247; hind foot, 118. Adult female from Minerva, N. Y. : 805; 
225; 105. Adult male from Granville, Nova Scotia: 837; 240; 116. Two adult 
males from Dismal Swamp, Va., respectively: 800, 860; 245, 285; 115, 110. Adult 
female from Dismal Swamp, Va. : 800; 250; 115. Skull: Adult female from Lib- 
erty Hill, Conn.: Greatest length, 114.4; condylobasal length, 109.8; zygomatic 
breadth, 74.1; interorbital breadth, 23; least width of palatal shelf, 14.8; maxillary 
tooth row (alveoli), 41.7; upper carnassial, crown length, 8.8, crown width, 9.2. 
Male and female from Adirondack Mountains, N. Y., respectively: Greatest 
length, 117, 110.9; condylobasal length, 112.9, 107.4; zygomatic breadth, 71.8, 
67.9; interorbital breadth, 22.7, 23.3; least width of palatal shelf, 16.2, 16.2; max- 
illary tooth row, 42.4, 41.9; upper carnassial, crown length, 8.3, 8.3, crown width, 
8.9, 9.3. Male and female from Dismal SVamp, Va., respectively: Greatest 
length, 116, 111.6; condylobasal length, 109.2, 105.7; zygomatic breadth, 76.4, 
68.5; interorbital l)readth, 25, 23.3; least width of palatal shelf, 16.2, 16.5; maxil- 
lary tooth row, 41.9, 39.7; upper carnassial, crown length, 8.8, 8.5, crown width, 
9.2, 9.1. 

Remarks. — Although individual variation is considerable, and due 
allowance should be made for it, the general characters of P. I. lotor 
are maintained with a fair degree of constancy throughout its range. 
Specimens from the northern part of the area have somewhat longer 
pelage and average somewhat darker than those from the southern 
part, but individuals contrasting strongly in color, some very dark 
and others light in tone, may be found at the same locality. Inter- 
gradation with P. I. hirtus, P. I. solutus, P. I. litoreus, and P. I. 
varius is evident, but the transition from one geographic race to another 
seems to be rather abrupt. [See also appendix, p. 84.] Specimens of P. 
I. lotor from Belleville, 111., and New Richmond, Mich., approach 
hirtus and might be referred to that form. Those from Dismal Swamp, 
Va., and eastern North Carolina are variable; some being near-typical 
lotor, while others grade toward solutus. 

[Meles] lotor Boddaert, L[otor] vulgaris Tiedemann, and Procyoii 
annulatus G. Fischer are substitute names for [Ursus] lotor Linnaeus. 
Procyon gularis Hamilton Smith was based on a live individual "in the 
State of New York," in which the "whole throat was black." There 
is no reason to assume that the animal differed from the typical form 
of the region, in which the amount of black on the throat is quite 
variable. 

Specimens examined. — Total number, 181, as follows: 

Connecticut: Liberty Hill, 3;^ West Greenwich, 2 (skulls only); exact locality un- 
known, 1 (skull only). 
District of Columbia: Washington, 1 (skull only). 

3 Mus. Comp. Zool. 



36 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

Illinois: Belleville, 2 (skulls only); Olive Branch, 2 (skulls only);'' Rosiclare, I.'* 
Indiana: Bicknell, 2 (skulls only); Culver, 1 (skull only); Pitcher Lake, Posey 

County, 1 (skull only) f Porter County, 1 (skull only) ; Russiaville, 1 (skull 

only); Salamonia, 1 (skull only).< 
Kentucky: Mammoth Cave, 1; Paducah, 1. 
Maine: Bethel, 4 (3 skulls without skins) ;•' Bucksport, 2 (skulls only) •,^ Ellsworth, 

1 (skin only);' Greenville, 1;' Penobscot River, 2; Umbagog Lake, 1 (skull 

only) :' LTpton, L' 

Maryland: Blackwater National Wildlife Refuge, Dorchester County, 5 (1 skull 
without skin) [referable to P. L maritimus, see appendix, p. 85]; Bowie, 1 
(skull only) ; Branchville, 1 ; Cabin John, 1 ; Jefferson, 2 (1 skull without skin) ; 
Laurel, 7 (skulls only); Marshall Hall, 1 (skull only); Patuxent River, 1 
(skull only). 

Massachusetts: Ayer, 1 (skull only);^ Brookhaven, 1 (skull only); Pepperell, 1 
(skull only);' Stockbridge, 1 (skull only).' 

Michigan: Constantine, 1 (skin only) ;^ Detroit, 1 (skull only); New Richmond, 1 
(skull only). 

New Hampshire: (Jssipee, 1 (skull only). 

New Jersey: Tuckerton, 3. 

New York: Adirondack Mountains, 9 (skulls only); Essex County, 4 (skulls 
only) ; Fort Montgomery, 2 (1 skin without skull) ;^ Hastings, 1 ;'' Hastings 
on Hudson, 4 (2 skins without skulls) ;^ Lake George, 1 (skull only) ; Lewis 
County, 1 (skull only); Leyden, 1 (skull only); Locust Grove, 1; Long 
Island, 1;" Monroe County, 1 (skull only) ;^ Minerva, 3;' Netherwood, 1;^ 
Piseco, 1; Saint Lawrence County, 2 (1 without skin and 1 without skull); 
Schoharie, 1 (skin oiily) :^ Schroon Lake, 1 (skull only); Severance, 8 (skulls 
only); Sing Sing, !() (skulls only); West Point, 2 (skins only). 

North Carolina: Asheville, 6 (5 skulls without skins); Coinjock, 1 (skin only); 
Highlands, 1 (skin only); Moore County, 1 (skull only); Pisgah National 
Forest (Bent Creek), 3 (skulls only); Raleigh, 1 (skull only). 

Nova Scotia: Bridgetown, 1;' Digby, 1; Granville, 3.^ 

Ontario: Credit River, 1 (skull only) ;^ Preston, Waterloo County, 1 (skull only).^ 

Pennsylvania: Allegheny County, 2 (1 without skin and 1 without skull) ; Carlisle, 
7 (skulls only): Chester County, 1 (skull only). 

Vermont: Newfane, 1 (skull only). 

Virginia: Amelia County, 1 (skull only); Buckingham County, 2 (skins only); 
Chesterfield, 1 (skull only); Chesterfield County, 1 (skull only); Clarke 
County, 1 (skull only); Dismal Swamp, 5 (1 skull without skin); Fredericks- 
burg, 4 (skulls only); Gunstou, 2 (skulls only); Morrison, 1 (skull only); 
Nelson County, 1 (skin only); Smith Island, 2; Warwick County, 1 (skull 
only); Washington, 4 (1 skull without skin). 



3 Mus. Comp. Zool. 
Chicago Mus. Nat. Hist. 
•Univ. Michigan Mus. Zool. 

* Mus. Vert. Zool. 

" Amer. Mus. Nat. Hist. 

* Natl. Mus. Canada. 



RACCOONS OF NORTH AND MIDDLE AMERICA 37 

PROCYON LOTOR HIRTUS Nelson and Goldman 
Upper Misslssippi Valley Raccoon 

Procyon lotor hirtus Nelson and Goldman, Jour. Mammal. 11 (4): 455, Nov. 11, 
1930. 

Tyjye locality. — Elk River, Sherburne County, Minn. 

Type. — No. 187926, male adult, skin and skull. United States 
National Museum (Merriam collection) ; collected by Vernon Bailey, 
March 4, 1886. 

Distribution. — Upper Mississippi and Missouri River drainage areas 
from the eastern slopes of the Rocky Mountains east to Lake Michi- 
gan, and from southern Manitoba and probably southwestern 
Ontario and southeastern Alberta south to southern Oklahoma and 
Arkansas. Overlapping- divisions of Upper Austral and Transition 
Zones; entering Canadian Zone to a limited extent near Lake Superior. 

General characters. — A large, dark subspecies with long, full, soft 
pelage, usually suffused with ochraceous buff"; skull with high, narrow 
frontal region, and weak or obsolescent postorbital processes. Similar 
to /'. /. lotor of the eastern United States, but much larger; pelage 
longer and usually more suffused with ochraceous buff. Size about 
as in P. I. Juscipes of Texas, but color darker, the pelage much longer 
and denser, more suffused with bull' instead of grayish, the light 
subapical zone of hairs over upper parts less extensive and permitting 
the under color to show through ; skull differing in slight details. 

Color. — Similar to P. I. lotor but usually more suft'used with 
ochraceous buff". 

Cranial characters. — Skull very similar to that of P. I. lotor in general 
form, but much larger, more massive; })rain case usually more tapering 
anteriorly, the sides of frontals diagoiuilly below and behind post- 
orbital processes less deeply indented or constricted; postorbital 
processes of frontals weakly developed, or obsolescent, as in lotor. 
About the same in size and in most important details as P. I. fuscipes, 
but interorl)ital and postorl)ital regions usually narrower; frontal area 
similarly high, but usually less flattened, with a narrower, more 
distinct, V-shaped median depression. 

Measurements. — An adult male from Fargo, N. Dak.: Total length, 880 mm.; 
tail vertebrae, 265; hind foot, 125. St<ull: Type: Greatest length, 127.1; 
condylobasal length, 122.2; zygomatic breadth, 80.5; interorbital breadth, 25.8; 
least width of palatal shelf, 15.8; maxillary tooth row (alveoli), 45.8; upper 
carnassial, crown length, 8.8, crown width, 9. 

Remarks. — The raccoon of the upper part of the Mississippi Valley 
is readily distinguished from its eastern relative, P. I. lotor, by much 
larger size, especially of the skulls. It is less easily separated from 
P. I. fuscipes of Texas, which is of about the same size, but typical 
specimens differ in color and in cranial details as pointed out. Inter- 



38 NORTH AMERICAN FAUNA 6 0, FISH AND WILDLIFE SERVICE 

gradation is evident, but in the sum of characters presented these 
widely ranging forms are quite distinct. 

Specimens examined. — Total number, 61, as follows: 

Colorado: Arkins, 1 (skull only); Cherry Creek, Arapahoe County, 2 (1 skull 

without skin); Estelene, 1; Las Animas, 1; Loveland, 1; Tuttle, 1; Wray, 4 

(1 skin without skull; 3 skulls without skins). 
Illinois: Chicago (Jackson Park), 1; Henderson County, 1; Joliet, I (skin only). 
Iowa: Keosanqua State Park, Van Buren County, 1.' 
Kansas: Manhattan, 1 (skull only). 
Minnesota: lieltranii County, 1; Elk River (type locality), 13 (6 skulls without 

skins). 
Missouri: Independence, 1 (skull only); Marble Cave, 1 (skull only). 
Nebraska: Beemer, 2 (skulls only); Haigler, I (skull only); Johnstown, 1 (skull 

only); Republican Fork, Platte River, 1 (skull only); Valentine, 1 (skull 

only): without exact locality, 1 (skull only). 
New Mexico: Bear Canyon, Raton Range, I (skull only); Raton Range (mouth 

of Trinchera Pass), 1. 
North Dakota: Fargo, 1; Grafton, 1 (skull only); Towner, 1. 
Oklahoma: Fort Cobb, 1 (skull only): Frederick ("20 miles from"), 1 (skin 

only); Mount Scott, 4 (1 skull without skin); Redfork, 1 (skin only). 
Texas: Canadian, 1 (skull only). 
Wisconsin: Delavan, G (5 skulls without skins): Okee, 1 (skull only); without 

exact locality, 1 (skull only). 
Wyoming: New Haven, 1. 

PROCYOX LOTOR VARIl'S Nelson .\nd Goldman 
Alabama Raccoox 
Prori/on lolor !<(tn'iis Nelson and (ioldnian, .lour. Mauimal. II (4): 456, Nov. 11, 
1930. 

Type localif!/. — Castlel^erry, Conecidi Comity, Ala. 

Type. — No. 158246, female adult, skin and skidl, United States 
National Museum (Biological Surveys collection) ; collected by A. H. 
Howell, October 10, 1908. 

Distribution. — Extreme southw^estern Kentucky, Tennessee, Missis- 
sippi, northern Louisiana, Alabama, northwestern Florida, and western 
Georgia. Mainly Ijower Austral Zone. 

General cha/acters. — A small subspecies most closely resembling 
Procyon lotor lotor, but smaller, usually paler, pelage much shorter, 
and skull differing in detail. Differing from P. I. elucus of Florida in 
paler color, rather decidedly smaller size, and in cranial features. 
Similar to P. I. fuscipes of Texas in color, but much smaller, with a 
different skull. 

Color. — Upper parts in general light buffy grayish, with a light 
ochraceous buffy suffusion along median dorsal area, l^ecoming more 
intense on nape and shoulders, thinly overlaid with black; sides 
clearer gray, the black-tipped hairs of dorsum thinning out ; top of 
head mixed brownish black and gray; facial mask brownish black, 

'Iowa State College. 



RACCOONS OF NORTH AND MIDDLE AMERICA 39 

becominp: rusty brownish on median line between eyes, and oehraceous 
buft'y on upper surface of muzzle; sides of muzzle, lips, and chin 
white; under parts in gcnieral thinly ovei'laid with buft'y grayish; 
throat patch brownish black; ears grayish with small black patches 
at posterior base; legs grayish, becoming whitish on feet; tail above 
with about five or six black rings and a black tip, alternating with 
light oehraceous buffy rings, the dark rings becoming bufty and less 
distinct below. 

Cranial characters. — Skidl small and slender, with weak or obsoles- 
cent postorbital processes of frontals; very similar to that of P. I. 
lotor, but smaller and more delicate in structure; jugal narrower; 
sides of frontals diagonally behind and below postorbital processes 
usually less deeply indented or laterally constricted; maxillary tooth 
row shorter; posterior upper premolar and upper carnassial usually 
distinctly smaller; very similar to that of P. I. elucus in general form, 
but rather decidedly smaller; brain case narrower, frontal region 
flatter, less "humped." Compared with that of P. I. fuscipes the skull 
is much smaller, more slender, with narrower frontal region, and 
postorbital processes of frontals (not very prominent \i\ fuscijpes) less 
developed. 

Measurements. — Type: Total length, 720 mm.: tail vertebrae, 218: hind foot, 
103. An adult male from Hurricane, Ala.: 772: 258; 109. Skull: Type: 
Greatest length, 104.8; condylobasal length, 98.6; zygomatic breadth, 64.4; 
interorbital breadth, 22.5; least width of palatal shelf, 14.6; maxillary tooth row 
(alveoli), 39.8; upper carnassial, crown length, 7.8, crown width, 7.9. 

Remarks. — The Alabama raccoon agrees more closely in combina- 
tion of characters with typical lotor than with any of the other known 
subspecies, although its distril)ution area constitutes a wedge, sepa- 
rating the ranges of elucus, fuscipes, and litoreus. [See appendix, 
p. 84.] Intergradation with these forms is evident, but the lines of 
demarcation between them appear to be fairly sharply drawn. 

Specimens examined. — Total luimber 57, as follows: 

Alabama: Ashford, 2 (1 skull without skin); Barachias, 5 (skulls only); Castle- 
berry, 2; Huntsville, 1; ?Iurricane (4 miles north), 2; Orange Beach, 
11 (9 skulls without skins); Perdido Bay, 1 (skull only); Sylacauga, 2 
(skulls only). 

Florida: Apalachicola, 5 (skulls only). 

Georgia: Geneva, Talbot County, 1 (skull only); Juniper, Talbot County, G:'" 
Nashville, 2 (1 skin without skull); Talbot County, 5 (skulls only). 

Kentucky: Hickman, 2. 

Louisiana: Baton Rouge, 1;^' Morrow, Saint Landry Parish, 1 (skull only); 
Mississippi River (mouth), 1 (skull only) " [referable to P. I. megalodous, 
see appendix, p. 84]. 

Mis.sissippi: Bay Saint Louis, 1 (skin only); Saucier, 1; '- Washington, 2. 

Tennessee: Arlington, 1; Big Sandy, 1; Clarksville, 1. 

1" Mus. Comp. Zool. 

" Louisiana State Univ. Mus. 

'2 Southern Forest Expt. Sta. Collection. 



40 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

PROCYON LOTOR LITOREUS Nelson and Goldman 

Saint Simon Island Raccoon 

Procyon lotor litoreus Nelson and Goldman, Jour. Mammal. 11 (4): 457, Nov. 
11, 1930. 

Ty2)e locality. — Saint Simon Island, Glynn Comity, Ga. 

Type. — No. 2450, adult (prol)ably male), skull only. United States 
National Museum; collected by Samuel W. Wilson; entered in museum 
catalog, Aug-ust 7, 1856. 

Distribution. — Coastal strip and islands of Georgia. Austrori- 
parian division of Lower Austral Zone. 

General characters. — Size medium and color dark, much as in 
Procyon lotor elucus oi Florida; length and texture of pelage about the 
same; skull differing in detail, especially in the much heavier (Umi- 
tition. Similar to P. I. lotor of Pennsylvania in color, but pelage 
shorter, more bristly, and cranial characters distinctive. Differing 
from P. I. sol nt lis of South Carolina in more buffy or brownish colora- 
tion and heavy dentition. 

Color. — Al)Out as in P. I. elucus. 

Cranial characters. — Skull similar in general form to that of P. I. 
elucus, but frontal region narrower and flatter, the sides usually more 
compressed or abruptly indented behind postorbital processes, leaving 
the brain case bulging laterally instead of tapering gradually into 
orbit as in elucus; dentition much heavier throughout, the difference 
being most noticeable in the large molariform teeth. Differing from 
that of P. I. sohitus in larger size and much heavier dentition. Com- 
pared with that of P. I. lotor tlie skull is usually larger, with much heav- 
ier dentition; postorbital processes of frontal weak or obsolescent as 
in lotor and elucus. 

Measurements. — Adult topotype: Hind foot (dry skin) 107 mm. SlcuH: Type: 
Greate^st length, 116.6; condylobasal length, 109.4; zygomatic breadth, 72.9: 
interorbital breadth, 22.2; least width of palatal shelf, 15.8; maxillary tooth 
row (alveoli), 43.6; upper carnassial, crown length, 9.6, crown width, 9.9. 

Remarks. — The remarkably heavy dentition readily distinguishes 
P. I. litoreus from all others of the group inhabiting the eastern 
United vStates. Its large teeth are equalled elsewhere in the group 
only in some of the larger subspecies of the Western States and south- 
ern Mexico and Central America. Specimens from vSaint Simon 
Island appear to reach the maximum in dental development, but are 
closely approached by those from the adjacent mainland. 

Specimens examinecl. — Total number, 25, as follows: 

Georgia: Altamaha River (mouth), 5 (skulls only); Mcintosh County, 2 (skulls 
only); Ossabaw IslaTid, 1 (skin only): Saint Simon Island, 15 (2 skins, 13 
skulls); Thunderbolt Creek, Chatham County, 2. 



RACCOONS OF NORTH AND MIDDLE AMERICA 41 

PROCYON LOTOR SOLUTUS Nelson and Goldman 

Hilton Head Island Raccoon 

Procyon lotor solulus Nelson and Goldman, Jour. Mammal. 12 (8) : 308, Aug. 24, 
1931. 

Ty2>e locality. — Hilton Head Island, Beaufort County, S. C. 

Ty2)e. — No. 256027, male adult, skin and skull. United States 
National Museum; collected by W. L. Brown, December 10, 1930. 

Distributtoii. — Coast region and islands of South Carolina. Lower 
Austral Zone. 

General characters. — Size rather small; color grayish; black mask 
uninterrupted across face in two of every three individuals; winter 
pelage rather long and dense. Similar, in general to P. I. litoreus of 
Saint Simon Island and the coast region of Georgia, but color clearer 
gray, less inclining toward ])uflFy or brownish; skull characters, espe- 
cially the much lighter dentition, distinctive. Dilfering from P. I. 
lotor in more gra^dsh general coloration, less elongated skull, and other 
cranial details. 

Color. — Upper parts in general grayish, rather heavily overlaid 
with black, especially on median dorsal area ; rather small nape patch 
suffused with ochraceous buff; top of head mixed black and gray, the 
gray pi-edominant; black mask usually continuous across middle of 
face, prolonged upward along median line to middle of forehead and 
downward over middle of muzzle to nose; sides of muzzle, lips, and 
chin white; under parts in general thinly overlaid with silvery gray, 
the dark l)rownish under color showing through; throat patch black- 
ish; ears gray, with l)lack ])atches at posterior base; limbs similar to 
under parts, becoming brownish gray on feet, the hind legs with pure 
black areas on outer side above heels; tail with about six black rings, 
narrowest near base, and a black tip, alternating with light ochra- 
ceous buffy rings. 

Cranial characters. — Skull of medium size, rather broad, short, and 
light in structure. Similar in general to that of P. I. litoreus but 
smaller; dentition very much lighter. Compared with that of P. I. 
lotor the skull is shorter and relatively broader; frontal region usually 
broader and flatter; ])alatal shelf shorter; dentition usually somewhat 
heavier, but maxillary tooth row shorter as a rule, the premolars more 
closely crowded. 

Measurements. — An adult male from Bulls Island, S. C. : Total length, 803 
mm.; tail vertebrae, 244; hind foot, 117. Two adult females from Bulls Island, 
S. C, respectively: 635, 749; 193, 260; 105, 107. Skull: Type: Greatest length, 
111.7; condylobasal length, 108.2; zygomatic breadth, 75.1; interorbital breadth, 
24.1; least width of palatal shelf, 16.4; maxillary tooth row (alveoli), 42.1; upper 
carnassial, crown length, 8.8, crown width, 9.2. 

87r>110°— 50 4 



42 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

Remarks. — The raccoon of Hilton Head Island and neighboring 
islands, and the adjacent mainland is readily distinguished from P. /. 
litoreus by much lighter dentition. Differentiation of this insular form 
is apparently due to isolation, its typical habitat being separated from 
the mainland by a broad and rather deep channel. Some skulls of 
this form closely resemble some of those of P. I. varius, the general size 
and dentition being very similar. Closely compared with those of 
varius, however, the skulls are usually shorter and broader, the frontal 
region broader and flatter, the brain case more rounded and inflated, 
and the cheek teeth somewhat larger. In addition, the longer pelage, 
grayer coloration, and more complete black facial mask appear to be 
distinctive. 

Sj^ecimens examined. — Total number, 31, as follows: 

South Carolina: Bulls Island, Charleston County, 11 (8 skins without skulls); 
EddinKs Island, 1; Edisto Island, Charleston County, 5; ^^ Hilton Head Island 
(type locality), 9 (6 skulls without skins); Hunting Island, 1; Saint Helena 
Island, 1; Santee, 2; Yeniassee, Hampton County, 1 (skull only).!^ 

PROCYON LOTOR ELUCUS Bangs 
Florida Raccoon 

Procyon lotor elucvs Bangs, Boston Soc. Nat. Hist. Proc. 28 (7) : 219, March, 1898. 

Tyi:)e locality. — Oak Lodge, on a peninsula opposite Micco, Brevard 
County, Fla. 

Type. — No. 3502, old male adult, skin and skidl. Museum of Com- 
parative Zoology (Bangs collection) ; collected by Outram Bangs, 
February 15, 1895. 

Distrihiffion. — Peninsular Florida, except southwestern part in- 
habited by P. I. 7narinus, north to extreme southern Georgia; grading 
into P. I. varius in northwest Florida. Tropical and Austroriparian 
division of Lower Austral Zone. 

General characters. — A medium-sized, generally dark-colored sub- 
species, with a deep, rusty rufous nuchal patch prominent in many 
typical examples; skull characterized especially by greatly inflated 
frontal vactiities usually giving the upper outline a decidedly "humped" 
appearance. Much like P. I. litoreus of Saint Simon Island, Ga.; 
general size, color, length and texture of pelage about the same; skull 
differing in detail, especially the much lighter dentition. Similar to 
P. I. varius of Alabama, but color usually darker, size decidedly larger, 
and cranial characters distinctive. Differing from P. I. marinus of the 



Mus. Conip. Zool. 



RACCOONS OF NORTH AND MIDDLE AMERICA 43 

Ten Thousand Islands and P. I. inesperatus of Upper Matecumhe Key 
in larger size, and the more elevated frontal re(2,'ion of skull. 

Color. — About as in P. I. litor-eus. Very similar to P. I. lofor but 
averag'ino- somewhat paler, the hairs over median dorsal area with 
shorter black tips and the nape more regularly and deeply suffused 
with rusty or orange rufous. 

Cranial characters. — Skull similar to that of P. I. litoreus, but frontal 
region broader, higher arched, or more "humped"; sides of frontals 
usually less compressed or abruptly indented behind orbits, leaving the 
brain case tapering more gradually anteriorly, instead of bulging 
laterally as in liforeus; dentition much lighter, the difference most 
noticeable in the molariform teeth. Decidedly larger than those of 
P. I. lofor, P. I. varius, P. I. marinus, or P. I. inesjjeratuSjWith brain case 
broader, and frontal region more ''humped"; postorbital processes of 
frontals obsolescent or small as in other eastern forms. 

Measurenieti.ts. — Type (from original description): Total length, 892 mm.; tail 
vertebrae, 286; hind foot, 125. An adult male topotype: 800; 244; 120. Average 
of five adult males from Saw Grass Island, Catfish Creek, Polk County, Fla. : 812 
(790-850); 259 (240-280); 126 (125-129); weight (pounds), 11 (10-12). Average 
of three adult females from same locality: 758 (745-770); 245 (235-255); 121 
(117-123); weight (pounds), 9 (7.7-10). Skull: Average of five adult males from 
Saw Grass Island, Catfish Creek, Polk County, Fla.: Greatest length, 119.4 
(113.7-123); condylobasal length, 114.5 (110.3-117.1); zygomatic breadth, 74.1 
(72-76.6); interorbital breadth, 23.7 (22.3-24.8); least width of palatal shelf, 16 
(14.9-17.3); maxillary tooth row (alveoli), 43.2 (41.7-44.6): upper carnassial, 
crown length, 8.7 (8.4-9.1), crown width, 8.9 (8.8-9.2). Average of three adult 
females from same locality: Greatest length, 113.6 (112.1-115.2); condylobasal 
length, 108.8 (106.8-110.4); zygomatic breadth, 68.4 (66.6-70); interorbital 
Ijreadth, 24 (23.3-24.5); least width of palatal shelf, 16.4 (15.4-17.6); maxillary 
tooth row, 41.9 (41.3-42.7); upper carnassial, crown length, 8.8 (8.7-9), crown 
width, 8.8 (8.6-9). 

Remarks. — In P. I. elucus the inflation of the frontal sinuses reaches 
its extreme development giving the skidl a "humjicd" appearance in 
outline, a character shared to some extent with other forms including 
P. I. lofor. Typical examples of elucus are quite dark in color, and the 
rusty rufous suffusion of the nape, appearing irregularly in many 
sul)species, is more prevalent and of a deeper and richer tone than is 
usual in the group. Intergradation with litoreus, imrius, and marinus 
is evident, and while not clearly indicated probably occurs with in- 
€sj)erafus which is known only from the mangrove-fringed islands along 
the southeast coast of the peninsula of Florida. Specimens from as 
far south as Naples on the west coast and Cutler, Dade County, are 
clearlv referable to elucus. 



44 XORTH AMERICAN FAUNA 6 0, FISH AND WILDLIFE SERVICE 

Specimens examined. — Total number, 127, as follows: 

Florida: Allenhurst, 1 (skull only); Aiicilla River, G (skulls only); Black Point, 
Dade County, 1; Blue Cypress Lake, Osceola County, 1 (skull only); Blue 
Springs, 1 (skin only) ; !•* Buena Vista, 1 (skull only); Citronelle, 4; ^^ Cutler, 
Dade County, 9 (1 skull without skin) ; '< Englewood, Sarasota County, 1; 
Enterprise, 1 (skin only) ; i« Fort Kissimmee, 18 (14 skulls without skins) ; 
Gainesville, 3;is Homosassa, 1 (skull only); Kissimmee, 1; Kissimmee River, 
1; Lake Cypress, 1; Lake Harney, 5 (1 skin without skull); Lake Hatch-ne- 
haw, 15 (skulls only): Lake Kissimmee, 8 (7 skulls without skins); Lake 
Monroe, 1; Matanzas Lilet, 1 (skull only); Miami, 1; Micco, 1 (skull only); 
Naples, 3 (skulls only); New Berlin, 5; '» Oak Lodge (type locality on penin- 
sula opposite Micco), 2 (1 skull without skin) ; '^ Orlando, 1 (skin only); is Royal 
Palm Hammock, 1 (skull only); San Mateo (5 miles northeast), 1 (skin only); 
Saw Grass Island, Catfish Creek, Polk County, 12; Snapper Creek, Dade 
County, 2; Tarpon Springs, 3 (2 skins without skulls; 1 skull without skin);'^ 
Taylor Creek, 2 (skins only) ; ^^ Wilson, 1; Welaka, 1 (skin only).i< 

Georgia: Fargo, 8 (5 skulls without skins) ; Okefenokee Swamp, 2 (1 skull without 
skin) . 

PROCYON LOTOR MARINUS Xelson 

Tex Thousaxd Islands Raccoon 

Procyon lotor mariniis Nelson, Smithsn. Misc. Collect. 82 (8): 7, July 10, 1930. 

TyjJe locality. — Near Chokoloskee, Collier County, Fla. 

Type.—^o. 254989, male adult, skin and skull. United States Na- 
tional Museum; collected by E. W. Nelson, February 28, 1930. 

Distribution. — Keys of the Ten Thousand Islands Group, and ad- 
joining mainland of southwestern Florida from Cape Sable north 
through the Everglades to Lake Okeechobee (Ritta). Tropical Zone. 

General characters. — A very small subspecies with heavy dentition. 
Not very unlike P. I. elucus and P. /. inesperatus in color, but smaller 
than either, and cranial characters, especially the relatively larger 
posterior upper premolar and carnassial, distinctive. Decidedly 
darker than P. I. auspicatus or P. I. incautus and cranial characters 
quite different. 

Color. — Similar to that of P. I. elucus, but somewhat grayer, especi- 
ally on the head, the back usually less heavily overlaid with black; 
rusty nape patch averaging less strongly marked, often obsolescent; 
under parts and light rings on tail paler, less ochraceous bulTy; black 
mask more restricted. 

Cranial characters. — Skull much smaller and more delicately propor- 
tioned than in P. I. elucus, frontal area much more depressed; brain 
case more rounded; posterior upper premolar and carnassial relatively, 
and sometimes actually, larger; palatal shelf about the same. Very 
similar in general form to that of P. I. inesperaius, but smaller, witli 

'■I Mus. Comp. Zool. 

'■' Amer. Mns. Nat. Hist. 

16 Four in Chicago Mus. Nat. Hist.; one in Mus. Comp Zool. 



RACCOONS OF NORTH AND MIDDLE AMERICA 45 

relatively and often actually larger (especially broader) posterior upper 
premolar and carnassial. Differing from those of P. I. auspicatus and 
P. I. incautus mainly in usually broader frontal region and much larger 
posterior upper premolar and carnassial. 

Measurements. — Type: Total length, 665 mm.; tail vertebrae, 222; 
hind foot, 105; weight (pounds), 7. Two adult male topotypes, 
respectively: 642, 655; 214, 200; 100, 98; weight (pounds), 7, 8. Two 
adult female topotypes: 610, 613; 200, 192; 93, 93; weight (pounds), 
5, 5.5. Skull: Type: Greatest length, 105.9; condylobasal length, 
101.8; zygomatic breadth, 64.8; interorbital breadth, 22.3; least width 
of palatal shelf, 13.9; maxillary tooth row (alveoli), 40; upper carnas- 
sial, crown length, 9, crown width, 9.1. Two adidt male topotypes: 
Greatest length, 101.3, 106.7; condylobasal length, 98, 101.8; zygo- 
matic breadth, 65.5, 69.5; interorbital breadth, 20.4 23.3; least width 
of palatal shelf, 14.9, 15; maxillary tooth row, 39.8, 40.5; upper carnas- 
sial, crown length, 8.9, 8.7, crown width, 9.7, 9.5. Two adult female 
topotypes: Greatest length, 93.7, 94.7; condylobasal length, 90.1, 
91.8; zygomatic breadth, 58.3, 60.6; interorbital breadth, 20.5, 20.3; 
least width of palatal shelf, 14.2, 14.1; maxiUary tooth row, 36.4, 38; 
upper carnassial, crown length, 8, 8.8, crown width, 9.2, 9.1. 

Remarks. — P. I. marinus is one of the smaller sul)species of raccoons 
that have developed neai- the southern end of the peninsula of 
Florida, not dift'ering much in size from P. I. auspicatus and P. I. 
incautus. It appears to be limited to the great maze of mangrove- 
covered or -bordered islands, or keys, known as the "Ten Thousand 
Islands" where raccoons are present in great numl:)ers, and to parts 
of the adjoining Everglades region. Specimens from Ritta at the 
southern end of Lake Okeechobee appear to be referable to marinus. 
Most of the islets mentioned are covered by the sea to a depth of 
from 3 to 4 feet at each high tide, and are totally devoid of fresh 
water. As most of these keys have no large trees to afford hollows 
and no dry land the raccoons must make their homes on top of the 
mangrove roots where they are forced to retreat by the incoming tide. 
Specimens from Cape Sable show gradation toward P. I. elucus, which 
ranges south to the eastern part of Dade County along the eastern 
side of the peninsula. Although evidently closely related to elucus, 
which occupies a different, but adjoining habitat, marinus maintains 
its distinctive characters with remarkable constancy. 

Specimens examined. — Total number, 49, as follows: 

Florida: Cape Sable, 3; Chokoloskee (type locality), 138; Coon Key, Ten 
Thousand Islands, l;i" Flamingo, Monroe County (skulls only), 3; ''* Ritta, 4 
(skulls oiilv). 



•" Amer. Mus. Nat. Hist. 
1' Mus. Comp. Zool. 



46 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

PROCYON LOTOR INESPERATUS Nelson 

Matecumbe Key Raccoon 

Frori/on Intor inesperatiis Xelsoii, Smithsn. Misc. Collect. 82 (8): 8, July 10, 1930. 

Ty2)e locality. — Upper Matecumbe Key, Monroe County, Fla. 

Type. — No. 255037, male adult, skin and skull. United States 
National Museum; collected by E. W. Nelson, March 19, 1930. 

Distribution. — Key Largo Group, embracing fringing keys along 
the southeast coast of Florida, from Virginia Key south to Lower 
Matecumbe Key. Tro]3ical Zone. 

General characters. — Closely allied to P. I. elucus of adjacent main- 
land, but averaging smaller and grayer; skull flatter. Differs from 
P. I. marinus, P. /. auspicaius, and P. I. incautus, representatives of 
neighboring groups of Florida keys, in its larger, more robust form, 
and in the combination of color and cranial characters. 

Color. — Much as in P. I. elucus but usually somewhat grayer, 
especially on head and face; black mask more restricted, the upper 
surface of muzzle paler; dorsum rather heavily washed with black, 
and rusty rufous nuchal patch well marked as in elucus; dark rings on 
tail distinct, and light rings often strongly buffy. 

Cranial characters. — Skull similar to that of P. I. elucus, but frontal 
area markedly depressed, instead of highly arched, or "humped." 
Differing from those of P. I. marinus, P. I. auspicaius, and P. I. incautus 
in larger size and more massive proportions; posterior upper premolar 
and carnassial actually, and therefore relatively, decidedly smaller 
than in P. I. marinus. Compared further with those of auspicaius 
and incautus, the palatal shelf extends farther behind the posterior 
molars than in the former and the frontal region is usually broader 
than in either. 

Measuretiients. — Type: Total length, 730 mm.: tail vertebrae, 250; hind foot, 
115; weight (pounds), 8.5. Adult male from Key Largo: 795; 222; 124; weight 
(pounds), 12. Adult female from Lower Matecumbe Key: 648; 228; 102; weight 
(pounds), 5. Skull: Type: Greatest length, 114; condylobasal length, 108.1; 
zygomatic breadth, 68.2; interorbital breadth, 23.1; least width of palatal shelf, 
15.2; maxillary tooth row (alveoli), 41; upper carnassial, crown length, 9.1, crown 
width, 9.6. 

Remarks. — Only a short distance separates the insular habitat of 
the present subspecies from the adjacent Florida mainland which is 
occupied by P. I. elucus. Nevertheless specimens from the various 
keys of the Key Largo Group differ somewhat in color as pointed out, 
and the skulls may at once be recognized by appreciably smaller size 
and more flattened frontals. The skulls of those from Key Largo 
and Virginia Key are larger than those from the more distant Upper 
and Lower Matecumbe Keys, and in this respect grade toward the 



RACCOONS OF NORTH AND MIDDLE AMERICA 47 

mainland animal. The motor highway from Miami to Key West, 
connecting keys by fills or viadncts enabling raccoons to pass from one 
key to another will doubtless result, through interbreeding, in the 
blending and obliteration of the interesting cbai'acters that now 
distinguish the various races of the island chain. 
Specimens examined. — Total number, 25, as follows: 

Florida (Key Largo Group): Elliotts Key, 7;" Key Largo, 5;^° Lignum Vitae 
Key, 1; Lower Matecumbe Key, 7; Plantation Key, 2; Upper Matecunibe 
Key, 1 (type); Virginia Key, 2. 

PROCYON LOTOR AUSPTCATUS Nelson 

Key Vaca Raccoon 

Procijon lotor (luspiratus Nelson, Sniithsn. Misc. Collect. 82 (8): 9, July 10, 1930. 

Tyj)e locality. — Marathon, Key Vaca, Monroe Cotnity, Fla. 

Tyj^e.—'No. 255080, male adult, skin and skull. United States 
National Musetmt; collected by E. W. Nelson, March 28, 1930. 

Distribution. — Key Vaca and doubtless closely adjoining keys of 
the Key Vaca Group, a central section of the main chain off the 
sotithern coast of Florida. Tro])ical Zone. 

General characters. — A very small, pale sid)species; skull with a 
narrow, but rounded brain case. Similar in size to P. /. marinus of 
the Ten Thousand Islands and P. I. incautus of the Big Pine Key 
Groui), btit decid(>dly paler than the former and differing in cranial 
details from both. Distinguished from P. I. inesperatus of Upper 
Matectmibe Key, by smaller size, much paler color, and by cranial 
characters. 

Color. — Very pale, similar to that of P. I. incautus, but not qtiite 
so extreme, much paler throughotit than P. I. inesperatus or P. I. 
marinus, the u])])er parts tistially thinly overlaid with rusty brownish, 
and the underfur of a lighter brownish tone than in inesperatus or 
marinus; black facial mask more restricted ; dark rings on tail narrower, 
more brownish, but usually distinct all aroimd. 

Cranial characters. — Skull very small, with a short palatal shelf 
and moderately heavy dentition. Similar to that of P. I. marinus, 
but somewhat smaller, with shorter palatal shelf, and lighter dentition. 
Smaller than that of P. I. inesperatus, with brain case relatively nar- 
rower, palatal shelf shorter, pterygoids less divergent posteriorly. 
Compared with that of P. I. incautus the skull is smaller, with shorter 
palatal shelf and narrower zygomata. 

Measurements. — Type: Total length, 644 mm.; tail vertebrae, 214; hind 
foot, 99; weight (pounds), 5.5. Average of five adult male topotypes: 657 
(634-700); 236 (214-275); 100 (96-107); weight (pounds), 5.3 (4-6). Two adult 

1^ Mus. Comi). Zool. 

-" Two in Mus. Comp. Zool. 



48 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

female topotypes, respectively: 603, 620; 212, 232; 83, 97; weight (pounds), 4, 5. 
Skull: Type: Greatest length, 100; condylobasal length, 94.7; zygomatic breadth, 
64.5; interorbital breadth, 19.4; least width of palatal shelf, 13.9; maxillary tooth 
row (alveoli), 37.4; upper carnassial, crown length, 7.8, crown width, 8.6. Aver- 
age of five adult male topotypes: Greatest length, 102.1 (99.9-105.9); condylo- 
basal length, 97 (94.3-101); zygomatic breadth, 63.5 (60.2-66.2); interorbital 
breadth, 20 (18.8-22.7); least width of palatal shelf, 14 (13.4-15); maxillary 
tooth row, 38 (36.4-38.9); upper carnassial, crown length, 7.8 (7.6-8.1), crown 
width, 9 (8.7-9.3). Two adult female topotypes: Greatest length, 93.6, 97.5; 
condylobasal length, 89.4, 94; zygomatic breadth, 59.8, 58.7; interorbital breadth, 
18.8, 19; least width of palatal shelf, 14, 13.8; maxillary tooth row, 35, 36.6; 
upper carnassial, crown length, 7.4, 7.7, crown width, 8.3, 8.6. 

I\e7ym)'ks. — The Key Vaca raccoon is one of the most salient in 
characters of the subspecies inhabiting the Florichi Keys. It re- 
seni])les P. I. hiccmtux in pale coloration, but departs from all in 
combination of cranial features. Its range is the most restricted of 
any of the Florida races. 

SpecimenK examined. — Thirteen, from ty])e locality. 

rKOC'YOX LOTOR INCAUTUS Xelson 

ToR( 11 Kky Raccoon 

Prorynn Ininr iitcaiitiis Xelsoii, Sinithsn. Misc. Collect. S2 (8): 10, .July 10, 19.30. 

Type locality. — Toi-ch Key, Big Pine Key Oi'oup, Monroe County, 
Fla. 

Type.— No. 2550()0, male a(hdt. skin and skull, United States 
National Museum; collected by El. W. Nelson, March 24, 1930. 

Disfribufion. — Big Pine Key Group, near southwestern end of 
chain of Flori(hi keys. Tropical Zone. 

General characfers. — A small, very pale std)species, palest of the 
Florida forms, with skull highly arched, and narrow l)etween orbits. 
Closely resembling P. I. auspicafus of Key Vaca in color, but cranial 
chai-acters, especially the nari'ower, high frontal region, distinctive. 
Decidedly paler than P. J. mnrirnis or P. 1. ineKperatus of Upper 
Matecumbe Key, and skull (hirering in important (U'tails. 

Color. — Very pale, similar to that of P. I. auspieatw'<, but averaging- 
even paler, especially on head and face, the black mask more restricted, 
more distinctly interrupted between eyes, the whitish areas cor- 
i'es])ondingly extenchHl and more compl(>tely isolating the dusky 
median streak; upj^er surface of muzzle light buffy; rusty nuchal 
patch conspicuous, inclining toward yellowish in worn pelages; dark 
rings on tail rusty brown, as in auspicatus, but usually broader. 

Cranial character.^. — Cranium small, with narrow, highly arched 
frontal region and light dentition. Averaging larger than that of 
P. I. auspicafu.s, with frontal region narrower, usually more highly 
arched; palatal shelf extending farther behind plane of last molars; 



RACCOONS OF NORTH AND MIDDLE AMERICA 49 

pterygoids more divergent posteriorly; molariform teeth smaller. 
Similar in general to those of P. I. marinus and P. I. inesperafus, 
but distinguished by narrower frontal region and smaller molariform 
teeth. 

Measurements. — Type: Total length, 694 mm.; tail vertebrae, 263; hind foot, 
118; weight (pofinds), 8.5. Average of five adult males from Big Pine Key and 
No Name Key: 710 (656-738): 247 (216-273); 111 (108-113); weight (pounds), 
8 (7.5-9.5). Average of four adult females from Torch Key (type locality), 
Boca Chica Key, and No Name Key: 688 (660-720); 240 (226-253); 105 (103- 
110); weight (pounds), 6.1 (5.5-6.5). Skull: Type: Greatest length, 111; 
condylobasal length, 104.7; zygomatic breadth, 66.7; interorbital breadth, 19.8: 
least width of palatal shelf, 14.9; maxillary tooth row (alveoli), 38.8; upper 
carnassial, crown length, 7.8, crown width, 8.5. Average of seven adult males 
from Big Pine Key and No Name Key: Greatest length, 109.8 (105-113.8) 
condylobasal length, 102.9 (97.8-106.2); zygomatic breadth, 69.4 (62.5-78.1) 
interorbital breadth, 21.8 (19.6-23.9); least width of palatal shelf, 14.8 (13.9-16) 
maxillary tooth row, 39.2 (38.1-40.4); upper carnassial, crown length, 7.8 (7.6-8), 
crown width, 8.6 (8-9). Average of four adult females from Torch Key, Boca 
Chica Key, and No Name Key: Greatest length, 104.9 (101-107.7); condylobasal 
length, 94 (96.5-100.5); zygomatic breadth, 61.9 (60.7-64.2); interorbital breadth, 
21.1 (20-22.2); least width of palatal shelf, 14.4 (14-15); maxillary tooth row, 
38 (37.4-38.5); upper carnassial, crown length, 7.5 (7.3-7.8), crown width, 8.3 
(7.8-8.8). 

Remarks. — The home of this race of raccoons is on the group of 
Florida Keys farthest from the mainland. As is the case with the 
other Florida Key raccoons they live mainly, and sometimes entirely, 
in mangrove swamps without access to fresh water except during rains. 
The brilliant light of their environment may have affected their 
general color more than the others, as suggested by their pale, faded 
tints. In general form and proportions the skull resembles that of 
P. I. elucus rather more closely than those of its geographically nearer 
insular relatives. It is much smaller, however, and suggests a minia- 
ture of that of the mainland animal. 

Specimens examined. — Total number, 33, as follows: 

Florida: Big Pine Key, 16; ^i Boca Chica Key, 2; Geiger's Key, 2; 22 Key West, 3 
(1 skull without skin); ^3 No Name Key, 5; Stock Island, 3; ^3 Torch Key 

(type locality), 2. 

PROCYON LOTOR FUSCIPES Mearns 

Texas Raccoon 

Procijon lotorfiiscipes Mearns, Biol. Soc. Washington Proc. 27: 63, March 20, 1914. 

Type locality. — Las Moras Creek, Fort Clark, Kinney County, Tex. 
(altitude 1,011 feet). 



-' Ten in Mus. Coiiip. Zool. 

" Mus. Comp. Zool. 

-' One iu Mus. Comp. Zool. 



50 NORTH AAIERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

Type. — No. 63055. male achilt. skin and skull, United States Na- 
tional Museum; collected by Edgar A. Mearns, February 6, 1893. 
Original number 2273. 

Distribution. — Texas, except extreme northern and western parts, 
southern Arkansas, Louisiana, except delta region of Mississippi, and 
south into northeastern Mexico, including Coahuila and Nuevo Leon, 
to southern Tamaulipas, Austroriparian and Lower Sonoran divisions 
of Lower Austral Zone. 

General characters. — A large, dark grayish subspecies, with pelage of 
medium length and texture; skidl with high, moderately broad frontal 
region and weakly developed postorbital processes. Size about as in 
P. /. hirtus; of Minnesota, but color grayer, less suffused with buff; 
mask more uniformly black and continuous across face and on upper 
surface of muzzle; pelage much shorter and less dense. Similar to 
P. I. mexicanus of Chihuahua, but decidedly darker, and cranial 
characters distinctive. Resembling P. I. varius of Alabama in general 
color, but somewhat grayer and much larger, with a different skull. 
Similar in size to P. I. hernandezii of the Valley of Mexico, but less 
grayish, the postauricular spots larger, more conspicuous; skull less 
flattened and differing in detail. 

Color. — Similar to that of P. I. varius, but averaging somewhat 
grayer, less suft'used with buff; mask usually more uniformly black and 
continuous across middle of face and on upper surface of muzzle to 
nasal pad. 

Cranial characters. — Size and general proportions of skull nearly as 
in P. I. hirtu.s, but interorbital and ])ostorbital regions usually l)roader; 
frontal region similarly high, but usually flatter, with a less distinct, 
V-shaped, median depression. vSimilar in size to that of P. I. mexi- 
canus; frontal region similarly high behind plane of postorbital 
processes, but more elevated anteriorly, the upper outline more convex; 
brain case more depressed near fronto-parietal suture; interorbital and 
postorbital regions usually narrower; postorbital processes of frontals 
shorter, the upper margin of orbit less deeply concave. Compared with 
that of P. I. varius the skuU is much larger and heavier, with broader 
frontal region. Not very unlike that of P. I. hernandezii, but less 
flattened, the frontal region more elevated; brain case usually more 
depressed near fronto-parietal suture; postorbital processes of frontals 
less prominent, the upper margin of orbit less deeply concave; posterior 
upper premolar and upper carnassial usually smaller. 

Measurements. — Type: Total length, 900 inin.; tail verteV>rae, 290; hind foot, 
132. Two adult males from Laredo, Tex., respectively: 860, 850: 298, 275; 
136, 131. An adult male and female from Sabinas, Coahuila: 922, 760; 330, 260; 
136, 116. Skull: Type: Total length, 130.2; condylobasal length, 125; zygomatic 



RACCOONS OF NORTH AND MIDDLE AMKHICA 51 

breadth, 84.4; interorbital breadth, 26.9; least width of palatal shelf, 16.2; maxil- 
lary tooth row (alveoli), 47.4; upper carnassial, crown length, 8.5, crown width, 
9.6. An adult male and female from Sabinas, Coahuila: Greatest length, 130.6, 
117.7; condylobasal length, 123.9, 112.1; zygomatic breadth, 82.8, 75.5; inter- 
orbital breadth, 24.4, 24; least width of palatal shelf, 18.1, 15.9; maxillary tooth 
row, 46.9, 43.3; upper carnassial, crown length, 8.9, 8.7, crown width, 10, 9.4. 

Remarks. — P. I. fuscipes requires rather close comparison with P. I. 
hliius to the north and P. I. hernandezii to the south, but typical 
specimens difrer in combination of characters as pointed out. It is 
readily distinguished from P. I. varius by much larger size and from 
P. I. mexicanus by darker color. Intergradation with all of these may 
safely be assumed. In typical fuscipes, however, the facial mask 
usually extends as a broad, uniformly black area across the face as in 
P. I. mexicanus and western subspecies in general, instead of being 
more or less distinctly interrupted by whitish longitudinal lines, one on 
each side near inner angle of eye, tending to isolate a narrow, elongated 
black median patch as in hirtus, varius and other eastern continental 
forms. In fuscipes the brain case, on the other hand, is somewhat 
depressed near the fronto-parietal suture and the postorbital processes 
of the frontals are veiy" short or obsolescent, characters shared with 
hirtus, varius and the eastern subspecies in contrast with mexicanus and 
the more western and southern continental forms in which the depres- 
sion of the brain case is less evident, and the postorbital processes of 
the frontals are well developed. 

Specimens examined. — Total numljer, 100, as follows: 

Coahuila: Muzquiz, 1; opposite Langtry, Texas, 3; Sabinas, 3. 

Louisiana: Abbeville, 8 (3 skins without skulls); Abbeville (24 miles southwest), 
3; Iowa, 1 (skull only); Lake Ridge, 1; Morgan City, 1; Tallulah, 12 (10 skulls 
without skins). [All specimens except those from Lake Ridge and Tallulah 
probably are referable to P. I. megalodous.] 

NuevoLeon: Monterrey, 1. 

Tamaulipas: Alta Mira, 1 (skull only); Bagdad, 1; Camargo, 3; Marmolejo, 1;-* 
Matamoros, 3 (2 skulls only). 

Texas: Angleton, 1; Aransas National Wildlife Refuge, Refugio County, 2; 
Aransas County, 2 (skulls only) ; Broome, 1 (skin only) ; Brownsville, 1 ; 
Canyon, 2; Carlsbad (10 miles east), 3; Columbia, 1 (skull only); Corpus 
Christi, 2; Dickinson Bayou (opposite Galveston), 1; Eagle Pass, 1; Fort 
Clark, 2 (including type); Cirady, 1 (skull only); Kerrville, 2 (skulls only); 
Kountze, 1; Langtry, 1; Laredo, 5 (3 skins without skulls); Liberty, 1 (skull 
only) ; Lomita Ranch, 2 (skulls only) ; Long Point, 1 (skull only) ; Los Ratones, 
Zapata County, 1; Mason, 4; Matagorda, 6 (5 skulls without skins); Padre 
Island, 1 (skull only); Port Lavaca, 1 (skull only); Rankin, 2; Sour Lake, 4 
(3 skulls without skins); Texarkana (10 miles northwest), 2; Washington 
County, 1 (skull only); Water Valley, 2 (skuUs only). 



2^ Univ. Michigan Mus. Zool. 



52 jS'ORTH AMERICAN FAUNA 60. FISH AND WILDLIFE SERVICE 

PROCYOX LOTOR MEXICAXUS Baird 
Mexican Raccoon 

Procyon lotor, variete mexicaine I. Geoffroy-Saint Hilaire, Voy. sur la Venus, 
Zoologie, p. 125, pL VI, 1855. From Mazatlan, Sinaloa, Mexico. 

Procyon hernandezii var. mexicana Baird, Mammal. Xorth Amer., p. 215, 1857. 

Procyon lotor mexicanus Mearns, Biol. Soc. Washington Proc. 27: 65, Mar. 20, 
1914. 

Tyi^e locality. — Espia, northwestern Chihuahua, Mexico. 

Tyjie. — No. 2018, probably female, adult, skull only (originally ac- 
companied by skin which cannot now be found), United States 
National Museum; collected by C. B. R. Kennerly, April 1855. 

Disfribution. — New Mexico, except northeastern and northwestern 
parts, southeastern Arizona, western Texas, and south through 
Chihuahua, eastern Sonora, Sinaloa and Durango to northern Nayarit, 
Mexico. Lower Sonoran to Transition Zone. 

General characiers. — One of the palest subspecies of the group; skull 
with broad frontal area highly arched behind plane of well-developed 
postorbital processes. Color and general size about as in P. I. ■pallidus 
of the Colorado River Valley, but skull usually broader, especially 
betw^een orbits, and differing in other slight details. Decidedly paler 
than P. I. fiiscipes of Texas, or P. I. hirtus of Minnesota, and combina- 
tion of cranial characters quite different. Similar in general to 
P. I. hernandizn of the Valley of Mexico but paler, the upper parts 
less extensively overlaid with black; skull more highly arched and 
presenting other distinctive features. 

Color. — Upper parts in general coarsely grizzled iron grayish and 
under parts light buffy about as in P. I. jmUuIus; black mask broad 
and uninterrupted across face; rusty nuchal patch usually absent, 
but faintly incHcated in occasional specimens. 

Cranial characters. — Skull most closely resembling that of P. I. 
imllidus, Ijut brain case, frontal area and palatal shelf usually broader; 
front als rather high behind plane of postorbital processes as in pallidus; 
tooth rows usually shorter. Compared with those of P. I. fuscipes 
and P. I. hirtus the frontal region is similarly high behind plane of 
postorbital processes, but less elevated anteriorly, the frontal outline 
descending in a more nearly straight line with nasals; brain case less 
depressed near frontoparietal suture; interorbital and postorbital 
areas usually broader; postorbital processes of front als longer, the 
upper margin of orbit more deeply concave. Contrasted with that 
of P. I. hernandezii the skull is of similar size, but less flattened, the 
frontal region more elevated behind plane of postorbital processes; 



RACCOONS OF NORTH AND MIDDLE AMERICA 53 

interorbital and postorbital areas usually broader; maxillary tooth 
row shorter; posterior upper premolar and carnassial smaller. 

Measurements. — Adult male from Loehiel, Santa Cruz River, Ariz.: Total 
length, 895 mm.; tail vertebrae, 365; hind foot, 121. Adult female from Deming, 
N. Mex. : 840; 305; 125. Adult male and female from Fort Lowell (near Tucson), 
Ariz., respectively: 890, 820; 325, 305; 131, 125. Skull: Adult male from 
Loehiel, Ariz., and adult female from Deming, N. Mex.: Greatest length, 121.1, 
120.6; condylobasal length, 114.6, 115.4; zygomatic breadth, 77, 78; interorbital 
breadth, 24.2, 24.2: least width of palatal shelf, 55.7, 55.9; maxillar,y tooth row 
(alveoli), 42.5, 42.7; upper carnassial, crown length, 8.7, 8.6, crown width, 9.7, 9.7. 
Adult male and female from Fort Lowell, xAriz.: Greatest length, 123.3, 116.2; 
condylobasal length, 115.6, 110.9; zygomatic breadth, 83.5, 78.8; interorbital 
breadth, 28.8, 25.3; least width of palatal shelf, 17.6, 16.4; maxillary tooth row, 
42.7, 43.6; upper carnassial, crown length, 8.5, 8.8, crown width, 9.5, 9.6. 

Remarks. — P. I. mexicanus shares extremely pale coloration with 
P. I. palluhis of the Colorado River Valley, typical examples of the 
two being externally indistinguishable. They are evidently very 
closely allied, some specimens being practically identical, but the 
combination of cranial characters pointed out usually serves to 
separate them as geographic races. The specimen described by 
Baird and regarded as the type of mexicanus is a fully adult, but 
undersized in(livi(Uial, probably a female (greatest length of skull, 
113) which does not properly reflect the true characters of the sub- 
species as shown by other specimens from the type region. The 
type locality was originally given as Espia, Sonora. Examination 
of Boundary Survey reports and statements by C. B. R. Kennerly, 
the collector, show that it was taken at Espia, shown on modern 
maps in extreme northwestern Chihuahua. The error was doubtless 
due to lack of infoi-mation in regard to the exact location of the 
Sonora-Chihuahua boundary. To this subspecies are referred speci- 
mens from Mazatlan, Sinaloa. A specimen from Mazatlan formed 
the basis of the detailed description by Geoffroy-Saint Hilaire (1855, 
p. 125) of a Mexican raccoon, "variete mexicaine," which, however, 
he did not name. A specimen from northern Nayarit is referred to 
mexicanus^ but those from localities farther south seem more properly 
assignable to P. I. hernandezii. Specimens from the upper part of 
the Gila River Valley grade toward and might be referred to paUidus. 
The series of 47 specimens from Escuinapa, Sinaloa, affords an unusual 
opportunity for the study of individual variation. 

Specimens examined. — Total number, 87, as follows: 

Arizona: Fort Huachuca, 1; Fort Lowell, 2; Loehiel, 1; San Bernardino Ranch, 
Cochise County, 1; Santa Catalina Mountains, 1 (skull only); Santa Rita 
Mountains (McCleary's Ranch), 1. 

Chihuahua: Casas Grandes, 1; Colonia Diaz, 1; E.spia, 1 (type, skull only); San 
Luis Mountains, 1. 



54 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

Durango: Rancho Saiituario (northwestern Durango), 1 (skull only).^^ 

Nayarit: Acaponeta, 1. 

New Mexico: Alcalde, 7 (skulls only): Central, Grant County, 1 (skull only) ; 

Chloride, 1; Deming, 1; Gila National Forest, 2 (skulls only); Magdalena 

Mountains, 1; Redrock, 1; Rinconada, 1; Santa Rosa, 2; Velarde, 3 (skulls 

only). 
Sinaloa: Escuinapa, 47 (12 skins without skulls; 10 skulls without skins) ; ^sMazat- 

lan, 2; Rosario, 1. 
Sonora: "N. Sonora, Lumholtz Expedition", 1 (skin only); ^s Oputo, 2. 
Texas: El Paso, 1 (skull only). 

PROCYON LOTOR PALLIDUS Merriam 
Colorado Desert Raccoon 

Procyon paUidus Merriam, Biol. Soc. Washington Proc. 13: 151, June 13, 1900. 

Procyon lotor ochraceus Mearns, Biol. Soc. Washington Proc. 27: 64, Alar. 20, 
1914. Type from Sonoyta River, Sonora, Mexico, near Quitobaquito, 
Pima Co., Ariz., No. 59900, male subadult, U. S. Natl. Mus. ; collected by 
Edgar A. Mearns, February 7, 1894. 

Ty2)e locality. — New River, Colorado Desert, Imperial County, 
Calif. 

Tyjje. — No. 99272, female adult, skin and skull. United States 
National Museum (Biological Surveys collection) ; collected by Frank 
Stephens, October 16, 1899. 

Dlsfribufion. — Colorado and Gila River Valleys and adjoining 
territory from tlu> delta north to northeastern Utah, and east to 
western Colorado and northwestern New Mexico. Mainly Lower 
Sonoran division of Lower Austral Zone, but ranging up along streams 
into Transition Zone. 

General characters. — One of the palest subspecies of the group; 
skull with narrow frontal area highly arched behind plane of postorbi- 
tal processes. Color and general size about as in P. I. mexlcanus of 
Chihuahua, but skull usually narrower, especially between orbits, 
and differing in other slight details. Decidedly paler, more ashy 
gray than P. /. psora of the Sacramento Valley; skidl with upper 
outline rising more prominently behind plane of postoi'])ital processes. 
Similar to P. I. (jrinnelli, l)ut slightly paler, and cranial characters, 
especially the more abruptly sloping frontal profile from apex behind 
plane of postorbital processes, distinctive. 

Color.- — About as in P. I. mexicanus. 

Cranial characters. — Skull closely resembles those of P. I. mexicanus 
and P. I. j^sora but brain case, frontal area and palatal shelf usually 
narrower; frontals high behind posterior plane of postorbital processes 
as in mexicanus (flatter and rising less prominently in psora) ; tooth 



-' AnuT. Mus. Nat. Hist. 



RACCOONS OF NORTH AND AIIDDLE AMERICA 55 

rows usually loniixn- than in mexicanus. Compared with P. I. grinnelli 
the brain case and interorbital region are narrower, and the anterior 
frontal outline descends in a more nearly straight line from apex 
innncdiately behind the postorbital processes — upper outline of 
frontals a more evenly convex curve in grinnelli. 

Measurements. — Type: Total length, 855 iiiiii.; tail vertebrae, 295: hind foot, 
128. Two adult males from Colorado liiver, Mexican Boundary, Ariz., respec- 
tively: 950, 875: 405, 340; 135, 126. An adult female from same locality: 845; 
305; 128. Skull: Type: Greatest length, 117.6; condylobasal length, 111.1: 
zygomatic breadth, 77.8; interorbital breadth, 25.5; least width of palatal shelf, 
13.5; maxillary tooth row (alveoli), 43.4: upper carnassial, crown length, 8.8, 
crown width, 9.6. Two adult males from Colorado River, Mexican Boundary, 
Ariz.: Greatest length, 133, 126.6; condylobasal length, 122.4, 120.6; zygomatic 
breadth, 79.1, 78.6; interorbital breadth, 25.3, 24.8; least width of palatal shelf, 
15.4, 16.8: maxillary tooth row, 46.6, 43.7; upper carnassial, crown length, 9.7, 
8.6, crown width, 10.4, 9.5 An adult female from same locality: Greatest length, 
119.6; condylobasal length, 114.1; zygomatic breadth, 77.1; interorbital breadth, 
23.4; least width of palatal shelf, 14.6; maxillary tooth row, 41.7; upper carnassial, 
crown length, 8.7, crown width, 9.3 

Remarks;. — As the name indicates the present subspecies is charac- 
terized by light coloration, a feature shared with P. I. mexicanus. 
The close alliance of P. I. pallidus and P. I. mexicanus is obvious but 
differing combinations of cranial characters seem to warrant their 
recognition as distinct, but not strongly marked forms. The type of 
paUidus is an unusually pale specimen as shown by comparison with 
others from localities so near that they must be regarded as typical. 
The type of P. I. ochraceus is a subadult male in rather faded pelage 
to which the name is doubtless due. It is not satisfactorily separable 
from P. I. pallidus. 

Specimens examined. — Total nmnber, 29, as follows: 

Arizona: Colorado River, Mexican Boundary, 3; Fort Apache (25 miles south- 
east), 1; Lakeside, 1 (skull only); Mellen, 1 ; ^e Phoenix, 1; Springerville (3 
miles northwest), 2; Tempe, 1; Topock, 1; Wupatki National Monument, 
Coconino County, 1. 

Baja California: Calexico (11 miles southeast), 1; Cocopah Mountains, 3 (skulls 
only); New River (5 miles south of Mexicali), 1 (skin only); Pascualitos 
Laguiia, 1; exact locality unknown, 1 (skull onl}-). 

California: Colorado River (5 miles below Needles), 1;-'' Colorado River (near 
Pilot Knob) , 1 ; -^ Colorado River (20 miles north of Picacho) , 1 ; -'' New 
River, Colorado Desert, 1 (type); Pilot Knob, 1; Potholes, 1. -^ 

Colorado: Navajo River, Archuleta County, 1 (skin only). 

Sonora: Sonoyta River, near Quitobaquito (type of ochraceus), 1. 

Utah: Pine Valley, 1 (skull only); Saint George, 1. 



26 Mus. Vert. Zool. 



56 NORTH AMERICAN FAUNA 6 0, FISH AND WILDLIFE SERVICE 

PROCYON LOTOR PSORA Gray 
California Raccoon 

Procyon psora Gray, Ann. Mag. Nat. Hist. 10 : 261, Dec. 1842. 

Procj/on lotor cah'fornicus Mearns, Biol. Soc. Washington Proc. 27: 66, Mar. 20, 
1914. Type from ocean beach near last Mexican Boundary Monument 
(No. 258), San Diego County, Calif.. Xo. 60675, female subadult, U. S. Natl. 
Mus.; collected by Frank Xavier Holzner, July 16, 1894. 

Type locality. — Sacramento, Sacramento County, Calif. 

Type. — Perhaps in British ■Museum; collected by Captain Belcher. 

Distnbution. — California, except extreme northwest coastal strip, 
the northeastern corner and southeastern desert region, ranging south 
through northwestern Baja California to San Quintin; extreme west- 
central Nevada (Wilson Canyon, east slope of Sierra Nevada). 
Lower Austral, Upper Austral, and Transition Zones. 

General characters. — A large, moderately dark form with a broad, 
rather flat skull. Very similar to P. I. pacificus of Washington, but 
averaging paler; skull usually more elongated and differing in detail. 
Decidedly darker, less ashy gray than P. /. paUidus of the Colorado 
River Valley, and cranial characters distinctive. Similar in general 
to P. I. excel sus of the Snake River Valley, but much smaller and 
usually darker; skull relatively narrower. 

Color. — Similar in general to that of P. I. lotor, but upper parts 
grayer, less suffused with buff under the overlying black-tipped hairs; 
light rings on tail less buffy; rusty nuchal patch usually absent or less 
prominent; black mask continuous across face, as in western forms in 
general, instead of more or less distinctly interi'upted on either side of 
median line as in lotor. 

Cranial characters. — Skull closely resembles that of P. I. pacificus but 
usually more elongated, the brain case less fully expanded, especially 
anteroexternally ; interorbital region narrower; upper profile rather 
flat and postorl)ital processes of frontals well developed as in 
pacificus. Similar to that of P. I. excelsus, but smaller, relatively 
narrower. Compared with that of P. I. pallirlus, the skull is of similar 
size, but relatively broader, with flatter frontal region. 

Measurement. — Adult male from Tehama, Calif.: Total length, 880 mm.; tail 
vertebrae, 277; hind foot, 138. Adult female from Wheatland, Cahf.: 870; 300; 
120. Adult male and female from Nicasio, Calif., respectively: 901, 820; 348, 
312; 132, 121. Skull: Adult male from Tehama and adult female from Wheat- 
land, Calif.: Greatest length, 124.8, 120.4; condylobasal length. 120, 113; zygo- 
matic breadth, 82.6, 79.2; interorbital breadth, 26.3, 26.8; least width of palatal 
shelf, 17.6, 16.2; maxillary tooth row (alveoli), 44.2, 43.9; upper carnassial, 
crown length, 8, 8.6, crown width, 9.3, 9.6. Adult male and female from Nicasio, 
Calif.: Greatest length, 124.5, 1 18.7; condylobasal length, 116.6, 1 14.2; zygomatic 
breadth, 82, 78.4; interorbital breadth, 24.2, 25.8; least width of palatal shelf, 
16.4, 16.1; maxillary tooth row, 43.2, 41.8; upper carnassial, crown length, 8.8, 
8.6, crown width, 9.7, 9.3. 



RACCOONS OF NORTH AND MIDDLE AMERICA 57 

Remarks. — P. I. psora has commonly been treated by authors as 
specifically distinct from P. I. lotor of the eastern United States. 
This is not so surprising as direct comparison of the skulls of these 
widely separated subspecies reveals rather striking differences, especi- 
ally the much larger general size, and broader, flatter frontal region 
with much more prominent postorbital processes of psora. These 
differences, however, are completely bridged by the intervening forms. 
P. I. psora passes gradually into P. I. pacijicus in northern California 
and southwestern Oregon, and some specimens from those regions 
might with similar propriety be referred to either form. Some 
specimens from near the type locality of P. I. callfornicus suggest 
gradation toward P. I. pallidus, but general comparisons indicate that 
calif ornicus cannot satisfactorily be separated from psora. 

Specirnens examined. — Total number, 198, as follows: 

Baja California: Laguna Hansen, 1; -^ San Quintin, 1; San Ramon (mouth of 

Santo Domingo River), 2; '-" San Telmo, 1 (skin only) ; '-" Valle de las Palmas, 

1 (skin only).-' 
California: Areata, 2; ^7 Baird, 1; Bakersfield, 1; -" Banta, 1 (skull only); Berkeley, 

4; 2" Bodfish, 1; Bradley, 3 (1 skull without skin, 2 skins without skulls); 

Camp Meeker, 1; Carbondale, 1;-" Carlotta, 1;-" Cassel, 4 (skulls only); 

Cazadero, 1 (skull only); ^' Chico, 4; Colusa, 2 (1 skull without skin); Covelo, 

1 (skull only) ; Cuddeback, Humboldt County, 2 (skulls only) ; -" Cuyamaca 
Mountains, 1;-' Cypress Point, Monterey County, 1;-" Dyerville (5 miles 
south), 1; Eel River (southwest of South Yollo Bolly Mountain), 2; El 
Portal, 2; ^^ Eureka, 1; 2' Fort Tejon, 2 (1 skull only); Gazelle (5 miles east), 

2 (skulls only) ; Glen Ellen, 1; Grass Valley, 3; ^^ Grizzly Island, 1; -' Gualala, 
1; 27 Hay Fork, Trinity County, 1; 27 Helena, 3; '-' Hoopa Valley, 1; Hum- 
boldt Bay (Carson's Camp, Mad River), 1; Inverness, 1 (skull only); Isa- 
bella, 1; -" Jolon, 3; Julian, 2; 27 Kern River (25 miles above Kernville), I; 
Klamath River, Siskiyou County, 1 (skull only); Knight's Landing (near 
type locality), 1; La Jolla, 1; -' Lake Merced, 1; 27 Lassen Creek, 1; Layton- 
ville, 2; Lierly's Ranch, Mendocino County, 1; Little Browns Creek, Trinity 
County, 2 (skulls only); Little Shasta, 3; Lockwood, 2; Marysville Buttes, 
2; 27 McCloud River (near Baird), 3 (2 skulls without skins); 27 Mendota, 1; 
Menlo Park, 1 ; ~~ Mission, Santa Inez, 1 ; Mohave River, 1 (skull only) ; 
Monterey, 2; Mount Diablo, 1; 27 Mount Saint Helena, 2; Mount Sanhedrin, 
2 (1 skull without skin); 27 National City, 2; Nelson, 1 (skull only); Nicasio, 
10 (5 skulls without skins) ; Orland, 1 (skin only) ; Ornbaun Spring, Hum- 
boldt County, 1 (skull only) ; -' Pacific Ocean beach, near Monument 258, 
Mexican Boundary, 1 (type of californicus) ; Paine Creek (Dale's Ranch), 
Tehama County, 4 (2 skins without skulls); 27 Pescadero, 5; 27 Pine City, 2 
(skulls only) ; Pitt River, Shasta County, 1 (skull only) : Placerville, 2 (skulls 
only); 27 Pleyto, 2 (skulls only); Point Pinois, 1; Point Reyes, 4 (2 skulls 
without skins); Portola, 2; 27 Portola Lake, San Mateo County, 1: 27 Posts, 
1; Red BluflF, 1 (skin only); Rio Dell, 1 (skull only); Rockport, 1 (skull only): 
Round Mountain, 3 (skull only); Rumsey, 1; 27 Saint John, Glenn County, 2; 
San Emigdio, 3; San Francisco, 1 (skin only); San Luis Obispo, 3; Sausalito, 1 
(skull only) ; 27 Shasta County, 1 ; Shasta Valley (6 miles east of Edgewood) , 



27 Mus. Vert. Zool. 

876119°— .50- 



58 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

1; Siielling, 3; ^^ Soqiiel Mill (40 miles east of Raymond), 1; South YoUo 
Bolly Mountain, 1; Spalding, Eagle Lake, 3; ^^ Spenceville, 1;-^ Stockton, 
14; -'" Suisun Marsh, 1; ^^ Tehama, 1; Three Rivers, 1; Tower House, Shasta 
County, 1; -^ Trinidad, 1; " Union Island, 1; Vacaville, 1; ^7 Victorville, 1; 2? 
Wawona, 1 (skull only); Weaverville, 2 (skulls only); " Wheatland, 1; Willow 
Lake, Plumas County, I; "-'' Winthrop, 1 (skull only); Wolf, 1.27 
Nevada: Wilson Canyon, east slope of Sierra Nevada, 1. 

PROCYON LOTOR PACIFICUS Merriam 

Pacib'ic Northwest Raccoon 

Procyon psora pacifica Merriam, North Amer. Fauna 16: 107, Oct. 28, 1899. 
Proei/oti proteus Brass, Aus dem Reiche der Pelze, p. 564, 1911. West coast from 
Puget Sound to the Cascade Mountains. (Not Procyon proteus Allen, 1904.) 

Type locality. — Lake Keechelus, Kittitas County, Wash, (altitude 
8,000 feet). 

Type. — No. 93137, adult [female], skin and skull, United States 
National Museum (Biological Surveys collection) ; collected by C. 
Hansen, January 15, 1898. 

Distribution. — Southwestern British Columbia, except Vancouver 
Island, northern, central, and western Washington, western Oregon, 
and extreme northwestern California. Upper Austral and Transition 
Zones. 

General characters. — A dark subspecies with a relatively broad, flat 
skull. Most closely resend)ling P. I. psora of the Sacramento Valley, 
Calif., but darker; skull relatively shorter and broader. Similar to 
P. I. excelsus of the Snake River Valley, southeastern Oregon, but much 
smaller, darker, and cranial characters distinctive. Decidedly larger 
than P. I. vancouverensis of Vancouver Island, and skidl tliffering in 
detail. 

Color. — As in P. I. vancouverensis. Much as in P. I. psora, but darker, 
the top of head and long guard hairs over upper parts in general more 
extensively black; subapical light bands of hairs somewhat narrower, 
tending to permit the basal color, which is of a darker tone (near dark 
cinnamon brown), to show through; mask unmixed black continuous 
across face and over upper surface of muzzle; rusty nuchal patch 
usually absent or inconspicuous. 

Cranial characters. — Skull comparatively short, broad and flat; 
interorbital space very broad; postorl)ital processes of frontals well 
developed. Resembling that of P. I. psora, but usually less elongated, 
the brain case more fully expanded, especially anteroexternally; inter- 
orbital region broader. Similar to that of P. I. excelsus, but smaller, 
with brain case more rounded. Very similar in general form to that 
of P. I. vancouverensis, but much smaller throughout; brain case 



Mus. Vert. Zool. 



RACCOONS OF NORTH AND MIDDLE AMERICA 59 

I'clatively larger, more inflated; nasals broader, narrowing more 
al)ni])tiy to a point posteriorly; pterygoids longer, more diverging 
l)osteriorly; maxillary tooth rows longer, the individual teeth much 
larger. 

Measurements. — Adult female from Steilacoom, Wash.: Total length, 830 mm.; 
tail vertebrae, 270; hind foot, 129. Skull: Type (9) and an adult female from 
Steilacoom, Wash., respectively: Greatest length, 11.3.8, 114.2; condylobasal 
length, 106.8, 109.2; zygomatic breadth, 79.9, 81.2; interorbital breadth, 2(3.8, 
27.3; least width of palatal shelf, 14.9, 1().9; maxillary tooth row (alveoli), 41.4, 
41.3: upper carnassial, crown length, 8.G, 7.9, crown width, 9.4, 10. Average of 
five adult males from Lake Cushman, Wash.: Greatest length, 119.5 (116.1-123.1) ; 
condylobasal length, 112.6 (109.5-116.5); zygomatic breadth, 81.7 (78.8-84.7); 
interorbital breadth, 26 (25.1-27.1); least width of palatal shelf, 16.8 (16.2-17.6); 
maxillary tooth row, 43.6 (42.8-44); upper carnassial, crown length, 9.1 (8.8-9.4), 
crown width, 9.8 (9.5-10). 

Remarks. — The j^resent subspecies is the raccoon of the Pacific 
Northwest coastal and Cascade Range regions, extending in small 
nimibers into the interior along the Columbia River Valley to north- 
eastern Washington. Like the regional representatives of other 
groups of mammals it is characterized by dark coloration. Compared 
with P. I. lotor of the eastern United States, which is also dark in 
color, the upper parts in pacijicus are more heavily overlaid with black, 
the light subterminal bands of the longer hairs and the light rings of 
the tail are grayer, less buffy or yellowish, the top of the head is 
blacker, and the mask is more uniformly black and continuous across 
the face. P. I. jxtcijicus intergrades with P. I. psora in southwestern 
Oregon and northwestern California, and with P. I. excelsus east of the 
Cascade Mountains in Washington and Oregon. Procyon proteus 
Brass was assigned to the raccoon of the west coast from Puget Sound 
to the Cascade Moimtains, which is within the range of typical P. I. 
pacificus. The name is also preoccupied by Procyon proteus Allen, 
applied to a crab-eating raccoon in South America. 

Specimens examined. — Total number, 82, as follows: 

British Columbia: Hastings, 1; Port Moody, 3 (skulls only). 

California: Crescent City, 5 (4 skulls without skins). 

Oregon: Big Summit Prairie, Ochoco National Forest, 1 (skull only); Blue River, 
1 (skull only); Bridge, 1; Colly wash Burn, 1; Estacada, 3 (2 skulls without 
skins); Glendale, 3 (skulls only); Glide (24 miles east), 1; Glide (14 miles 
northeast), 1; Grant's Pass, 7 (32 miles south, 5 [1 skull without skin]; 43 miles 
northeast, 2 [1 skin without skull]); Hardman, 2; Pistol River (North Fork), 
Curry County, 1; Port Orford, 5 (skulls only); Remote, 1 (skull only); 
Riverside, 2. 

Washington: Easton, 1; Hoodsport, 4 (skulls only); Lake Cushman, 10 (skulls 
only) ; Lake Keechelus, 1 (type) ; Mount Vernon, 2 (skulls only) ; Orcas Island, 
1 (skull only); Skokomish River, Olympic Mountains, 2 (skulls only); 
Steilacoom, 5 (1 skin without skull): Tieton, 1; Toppenish, 3; Trout Lake, 
south base of Mount Adams, 5 (skulls only); Whidby Island, 1 (skull only). 



60 NORTH AMERICAN FAUNA 6 0. FISH AND WILDLIFE SERVICE 

PROCYON LOTOR EXCELSUS Nelsox and Goldman 

Snake River Valley Raccoon 

Procyon lotor excehvfi Nelson and Goldman, Jour. Mammal. 11 (4): 458, Nov. 11, 
1930. 

Type locality. — Owyhee River, Oreg., 10 miles west of Fairylawn, 
Owyhee County, Idaho. 

Type. — No. 236214, old male adult, skin and skull, United States 
National Museum (Biological Surveys collection) ; collected by J. W. 
Fisk, April 15, 1920. 

Distribution. — Snake River drainage in southeastern Washington, 
eastern Oregon, and southern Idaho, the Humboldt River Valley, 
Nev., and river valleys of northeastern California. Mainly Upper 
Sonoran Zone. 

General characters. — Size largest of the group; color rather pale, 
similar in color to P. I. psora of California, but usually paler, and much 
larger, with skull differing in detail. Closely allied to P. I. pacijicus 
of Washington, but much larger, decidedly paler, top of head much 
grayer, and cranial characters distinctive. 

Color. — Upper parts of body in general very light buffy grayish, 
with a light ochraceous buffy suffusion along median dorsal area, 
becoming pronounced on nape, moderately overlaid with black; sides 
clearer gray, the overlying black-tipped hairs less numerous than on 
dorsum; top of head a grizzled mixture of black and gray; face with the 
usual black mask and white markings; under parts in general thinly 
overlaid with buft'y grayish, the light brownish undertone showing 
through; throat patch dark brownish; ears grayish with black patches 
at posterior base; limbs grayish, the hind legs with small, unmixed 
brownish areas on outer sides near heels; tail above with about six 
black annulations and a black tip, alternating with somewhat broader, 
light buffy rings, the dark rings usually becoming indistinct below. 

Cranial characters. — Skull similar to that of P. I. psora, but larger 
and more angular; frontal region broader, generally flattened, and 
postorbital processes well developed, as in psora. Compared with 
that of P. I. pacijicus the skull is larger, with brain case relatively 
more elongated; frontal region broad as in pacijicus. 

Measurements. — No skin inea.surements available. Skull: Type: Greatest 
length, 136.5 mm.; condylobasal length, 125.8: zygomatic breadth, 89.1; inter- 
orbital breadth, 30.1; least width of palatal shelf, 17.2; maxillary tooth row 
(alveoli), 47; upper carnassial, crown length, 9, crown width, 10.6. 

Bemarks. — The present form is easily distinguished from all others 
of the group by the large size and massive development of the skull. 
No close cranial comparisons with forms east of the Rocky Mountains 
are necessary as this race differs notably in the much larger size, and 



RACCOONS OF NORTH AND MIDDLE AMERICA 61 

broad, flat frontal region, with prominent postorbital processes — 
frontal region generally high and narrow, and postorbital processes 
weak, or obsolescent, in forms east of Rocky Mountains. 
Specimens examined. — Total number, 32, as follows: 

California: Parker Creek, Modoc County, 1.^** 

Idaho: Bruneaii. 1 (skull only); Emmett, Gem County, 1; Forest (Deer Creek), 
1 (skin only); Hagerman, 4 (1 skull without skin); Lost Valley Reservoir, 
head of Wieser River (altitude 5,000 feet), Adams County, 1 (skin only); 
Preuss Mountains, 1 (skull only) ; Stanley Lake, Custer County (altitude 
8,500 feet), 1 (skin only); Three Creek, 2 (1 skin without skull; 1 skull without 
skin) . 

Nevada: Golconda, 1 (skull only); Montello, L 

Oregon: Adel, 2; Dry Creek, Malheur County, 1 (skull only); Enterprise, 1 
(skin only); Harper (8 miles east), 1 (skull only); Huntington, 1 (skull only); 
Imnaha, 1 (skull onhO ; Owyhee River (type locality, 10 miles west of Fairy- 
lawn, Idaho), 2; Rome, 1; Tupper, 1 (skull only); VaTisycle, 1 (skull only). 

Washington: Alpowa, 1 (skull only); Garfield County, 1; Touchet, 1; Wallula, 1; 
Washtucna, 1. 

PROCYON LOTOR VANCOUVERENSIS NelsoxX and Goldman 

Vancouver Island Raccoon 

Frocyon lotor rancouverensis Nelson and Goldman, Jour. Mammal. 11 (4): 458, 
Nov. 11, 1930. 

Tyi^e localiiy. — Quatsino Sound, Vancouver Island, British Colum- 
bia, Canada. Transition and Canadian Zones. 

Tyije. — No. 135457, male adult, skull only, United States National 
Museum (Biological Surveys collection); collected by Charles Sheldon, 
November 1904. 

Distribution. — Known only from Vancouver Island. 

General characters. — A dark subspecies most closely allied to P. I. 
pacijicus of Washington, but decidedly smaller, and cranial details 
distinctive. 

Color. — An adult (winter pelage) from Beecher Bay, Vancouver 
Island: Upper parts in general grayish, heavily overlaid with black; 
small nape patch sufi'used with ochraceous buff; top of head mixed 
black and gray, the black predominating; face with brownish black 
mask, the dark color extending down along middle of muzzle to nose; 
sides of muzzle, lips, and chin white; under parts, in general, thinly 
overlaid with btiff gray, the dense brown underfur showing through; 
throat patch brownish, mixed with gray along median line; ears gray- 
ish, with black patches at posterior base; limbs similar to under parts, 
becoming soiled whitish on feet, but hind legs with unmixed, dark 
brownish areas on outer sides above heels; tail with six narrow black 
rings and a black tip, alternating with broader grayish rings, the black 
rings interrupted on under side near base. 

2s Mus. Vert. Zool. 



62 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

Cranial characters. — Skull rather small, short, low, broad, and flat, 
with well-developed postorbital processes. Very similar in general 
form to that of P. /. pacjjicux, Init much smaller throughout; brain case 
relatively smaller and less inflated; nasals narrower and more attenuate 
posteriorly; pterygoids shorter, less diverging posteriorly; maxillary 
tooth rows shorter, the individual teeth much smaller. 

Measurements. — An adult from Beecher Bay, Vancouver Island: Hind foot 
(dry skin), 112 mm. Skull: Type: Cireatest length, 116; condylobasal length, 
108.9; zygomatic breadth, 77.5; interorbital breadth, 25.4; least width of palatal 
shelf, 16.5; maxillary tooth row (alveoli), 40.2; upper carnassial, crown length, 8.3, 
crown width, 8.9. 

Remarks. — The Vancouver Island raccoon is a well-marked sub- 
species. It requires close comparison only with P. I. pacijicus of the 
adjacent mainland. 

S])ecimens examined. — Total numl)er, 40, as follows: 

Vancouver Island, B. C: Alberni Valley (Hall's Eanch), 1; 29 Beecher Bay, 3 (2 

skulls without skins) ; ^o Cadboro Bay, 1 (skull only) ; ^ Errington, 1 ; ^^ Fort 
Rupert, 1 (skull only):" French Creek, 1 ; -s Little QuaHcum River, 1 ; ^9 
Mount Tolmie, 1 (skull only) ; ^o Parksville, 2; -'» Quatsino Sound (type 
locality), 21 (skulls only): San Josef River Valley, 1 (skull only); Sooke, 2 
(skulls only) ; 3" Victoria, 1 (skull only) ; ^o exact locality unknown, 3 (skulls 
only).3« 

PROCYON LOTOR GRINNKLLI Nelson and Goldman 

Baja California Raccoon 

Prncyon lotor (/rinnelli Nelson and Goldman, Jour. \\'ashington Acad. Sci. 20 (5): 
82, Mar. 4, 1930. 

Type locality. — La Paz, Baja California, Mexico. 

Type. — No. 147181, male adult, skin and skidl. United States 
National Museum (Biological Surveys collection); collected by E. W. 
Nelson and E. A. Goldman, February 15, 1906. 

Distribution. — Southern Baja California from the Cape region north 
at least to San Ignacio. Tropical and Lower Sonoran Zones. 

General characters. — A large, pale sid^species with a rather broad, 
high, evenly arched skull. Similar to P. I. pallidus of the Colorado 
Desert, but slightly darker and cranial characters, especially the more 
evenly arched profile of s*lvull, distinctive. Compared with P. I. psora 
of the Sacramento Valley, general color paler, more grayish, less 
deeply suft'used with huft", the long black guard hairs over dorsum less 
in evidence; top of head grayer, less heavily mixed with black; black 
areas at posterior base of ears smaller; skull with frontal region more 
highlv arched. 



-* Mus. Vert. Zool. 

3fi Provincial Mus., Britisli Columbia. 

31 Anier. Mus. Xat. Hist. 



RACCOONS OF NORTH AND MIDDLE AMERICA 63 

Color. — Upper parts in general coarsely grizzled iron grayish, the 
median dorsal area faintly suffused with pale buff, becoming pro- 
nounced on back of neck, rather thinly overlaid with black; top of head 
gray, mixed with black, prochicing a grizzled effect; face with solid 
black mask; white facial markings as usual in the group; under parts in 
general overlaid with very pale buffy grayish, the brown undertone 
showing through; ibi'oat patch blackish; ears gi-ayish, with rather 
small black patches at posterior base; limbs similar to under parts, but 
becoming whitish on feet; hind legs with small, pure brownish areas on 
outer side near heels; tail with the usual annulations and black tip, 
the light rings pale cream buff and the narrower dark rings (6 to 7) 
consisting of black-tipped hairs with an underlying buffy suffusion; 
dark rings less evident on under side of tail and scarcely complete, 
tending to fade out on median line, except near tip. 

Cranial characters. — Skull similar to that of P. I. palluius, but brain 
case and interorbital region broader; frontals rising higher anteriorly, 
the upper outline a more evenly convex curve — anterior frontal outline 
descending in a more nearly straight line from apex immediately 
behind postorl)ital processes in pallidus] dentition about the same. 
Compared with that of P. I. psora the skull is less flattened, the frontal 
region more highly arched; brain case rather broad and other cranial 
details much as in psora. 

Measurements. — Type: Total length, 913 mm.; tail vertebrae, 335; hind foot, 
132. Skull: Type: Greatest length, 122.1 ; condylobasal length, 115.5; zygomatic 
lireadth, 77.9; interorbital breadth, 24.3; least width of palatal shelf, 16.7; maxil- 
lary tooth row (alveoli), 44.1 ; upper carnassial, crown length, 8.8, crown width, 9.3. 

Remarks. — Raccoons are dependent upon water for existence, and 
owing to exceedingly arid concHtions in the central section of Baja 
California their general range is interrupted for consideral)le distances. 
The form here described, which occupies the southern half of the 
peninsula, differs rather markedly in coml)ination of characters from 
both of the more northern stil)species, P. I. psora and P. I. pallidus. 
It recpiires no very close comparison with P. I. mexicanus of the 
adjacent mainland of Mexico, which in general, is paler, with the black 
postain-icular spots obsolescent, and sktdl notably (k'pressed in frontal 
region. 

Specimens examined. — Total nimiber, 11, as follows: 

Baja California, Mexico: La Paz (type locality), 3;^- Mount Miraflores, 3; ^^ 
San Ignacio, 5. 



32 One (skull only) in Mus. Vert. Zool. 

33 Two in Amer. Mus. Nat. Hist. 



64 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

PROCYON LOTOR HERNANDEZII Wagler 

Mexican Plateau Raccoon 

Pr[ocyon] hernandezii Wagler, Isis 24: 514, 1831. 

Procyon lotor hernandezii Allen, Amer. Mus. Nat. Hist. Bull. 3: 176, Dec. 10, 1890. 

Type locality. — Valley of Mexico, Mexico (specimens from Tlalpam 
regarded as typical). ^^ 

Type. — Not designated. 

Distribution. — Southern part of tableland or plateau region of 
Mexico and adjoining coasts, from Nayarit, Jalisco, and San Luis 
Potosi, south to near the Isthmus of Tehuantepec. Altitudinal range 
from sea level to about 8,000 feet. Tropical to Transition Zone. 

General characters. — A large, dark grayish subspecies; skull some- 
what flattened, with narrow frontal region and slender, wide spreading 
zygomata; dentition heavy. vSimilar in general to P. I. fuscipes of 
Texas, but upper parts grayer; skull flatter and differing in detail. 
Decidedly darker than P. I. mexicanus of Chihuahua, the upper parts 
more extensively overlaid with l)lack, and cranial characters distinc- 
tive. Differing from P. I. shuj'eldti of Campeche in longer pelage; top 
of head darker and back more heavily overlaid with black ; skull more 
slender. 

Color.- — Very similar to that of P. I. fuscipes but still grayer, less 
buffy beneath overlying black; black postauricular spots smaller. 
Young (in first pelage) : Similar to lotor of corresponding age, but top 
of head and postauricular areas less extensively brownish black, and 
black mask continuous across face (mask more or less interrupted 
between eyes in lotor) ; feet dark brownish instead of buffy. 

Cranial characters. — Skull size about as in P. I. fuscipes but more 
flattened above, the frontal region less elevated, and brain case less 
depressed near fronto-parietal suture; postorbital processes of frontals 
usually longer, narrower, more acutely pointed; upper margin of orbit 
usually more deeply concave; posterior upper premolar and upper 
carnassial usually larger. Similar to that of P. I. mexicanus, but 
flatter, the frontal region less elevated; interorbital and postorbital 
regions usually narrower; maxillary tooth rows longer; posterior 
upper premolar and upper carnassial larger. Compared with that 
of P. I. shufeldti the skull is more slender, less massive; interorbital 
and postorbital regions narrower; dentition about the same. 

Measurements. — Adult male from Tlalpam, Valley of Mexico, Mexico: Total 
length, 905 mm.; tail vertebrae, 283; hind foot, 122. Two adult males from 
Jalpan, Queretaro, and Patzcuaro, Michoacan, respectively: 894, 872; 340, 308; 
129, 127. Two adult females, Tetela del Volcan, Morelos, and El Chico, Hidalgo: 
860,825; 300, 264; 120, 122. Skull: Adult male from Tlalpam, Mexico: Greatest 



3* Type locality fixed by Nelson and Goldman, Biol. Soc. Washington Proc. 44: 17, Feb. 21, 19.31. 



RACCOONS OF NORTH AND MIDDLE AMERICA 65 

length, 122.9; condylobasal length, 116.9; zygomatic breadth, 86; intororbital 
breadth, 25.2; least width of palatal shelf, 16.8; maxillary tooth row (alveoli), 
45.7; upper carnassial, crown length, 9.2, crown width, 10.3. Two adult males 
from Acambaro, Michoacan, and Jalpan, Queretaro: Greatest length, 128.3, 
123.7; condylobasal length, 124.8, 116; zygomatic breadth, 86.2, 83.1; interorbital 
breadth, 24.2, 23.9; least width of palatal shelf, 16.4, 16.4; maxillary tooth row, 
45.2, 44.2; upper carnassial, crown length, 9.3, 9.1, crown width, 10, 9.3. Two 
adult females from Tetela del Volcan, Morelos, and El Chico, Hidalgo: Greatest 
length, 118.1, 114; condylobasal length, 114.8, 109.3; zygomatic breadth, 79.3, 
76.8; interorbital breadth, 24.7, 22.1; least width of palatal shelf, 16.4, 15.8; max- 
illary tooth row, 44.2, 42.3; upper carnassial, crown length, 9.4, 8.2, crown width, 
10, 9.4. 

Jiemarks. — The range of P. I. hernandezii in southern Mexico is 
tiaiiscontineiital, and while mainly at 4,000 to 6,000 feet on the table- 
land of the interior it extends from sea level along the tropical coasts 
to 8,000 feet altitude on the slopes of the mountains bordering the 
Valley of Mexico. It intergrades on the north in eastern Mexico 
with P. I. fusci/pes and in western Mexico with P. I. mexicanus. 
Toward the southeast its range meets that of P. I. ■•<hufeidti. 

Specimens examined. — Total number, 50, as follows: 

Colima: Armeria, 1; Colima, 5 (3 skulls without skins); IManzanillo, 8 (3 skulls 

without skins). 
Guerrero: Papayo, 2; Tlapa, 1, 
Hidalgo: El Chico, 1. 
Jalisco: Arroyo de Plantinar, 1;^^ Atemajac, 3 (1 skull without skin); Barranca 

Ibarra (Canyon de Oblatos), Rio Grande de Santiago, 1; Garabatos, 1;^^ Las 

Canoas, 2;^' Zacoalco, 1; Zapotlan, 2 (1 skull without skin). 
Mexico: Ajusco, Distrito Federal, 1 (skull only); Tlalpam, Distrito Federal, 1. 
Michoacan: Acambaro, 2 (skulls only); Patzcuaro, 2 (1 skull without skin); 

Querendaro, 3 (2 skulls without skins). 
Morelos: Tetela del Volcan, 1. 
Nayarit: San Bias, 1. 
Oaxaca: Cuicatlan, 1. 

Queretaro: Jalpan, 2 (1 skull without skin). 
San Lui.s Potosi: Hacienda la Parada, 1; San Luis Potosi, 1. 
Veracruz: Jico, 3; ]\Iirador, 2 (1 skin without skull). 

PROCYON LOTOR SHUFELDTI Nelson and Goldman 
Campeche Raccoon 

Procyon lotor shufeldti Nelson and Goldman, Biol. Soc. Washington Proc. 44: 17, 
Feb. 21, 1931. 

Type locality. — La Tuxpena, Champoton, Campeche, Mexico. 

Type. — No. 177546, male adult, skin and skull, United States 
National Museum (Biological Surveys collection) ; collected by Percy 
W. Shufeldt, April 20, 1911. 

Disfributioti. — From the Isthmus of Tehiiantepec east through 
Chiapas, Tabasco, Campeche, Yucatan, Quintano Roo, British 

35 Amer. Mus. Nat. Hist. 



66 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

Honduras, and Guatemala to western Honduras; limits of range 
unknown. Tropical Zone. 

General characters. — A large, rather pale, short-haired subspecies, 
with massive skull. Similar in general to P. I. hernandezii of the 
Valley of Mexico, but pelage shorter, color duller, top of head grayer 
and back less modified by black-tipped hairs; black postauricular 
spots (small in hernandezii) still less distinct; skull more massive and 
differing in detail. Size about as in P. I. crassidens of Costa Rica, but 
color decidedly paler and grayer, the upper parts less heavily overlaid 
with black, and the subterminal light zone of longer hairs more 
extended and thus affecting the general tone; skull less flattened. 
Differing from P. I. dickeyi in larger size, much grayer color, and in 
cranial characters. 

Color. — Upper parts in general usually light buffy gray, with rather 
thinly distri])uted overlying black-tipped hairs resulting in a coarsely 
grizzled blend; nape patch rusty rufous; sides lighter, the black tips of 
hairs inconspicuous; top of head clearer gray, mixed with black, lacking 
the light buffy tone suffusing the back; black mask across face extend- 
ing downward along median line of muzzle to nose and upward to 
middle of forehead; lines bordering mask above, sides of muzzle, lips, 
and chin white as usual in the group; under parts in general thinly 
overlaid with very light bufl'y hairs, the light ])rownish underfur show- 
ing through, but short and scarcely concealing the skin; throat patch 
brownish; cars grayish; black postauricular spots small and incon- 
spicuous; limbs similar in color to under parts, but over hairs denser, 
becoming dull whitish on feet; hind legs with outer sides of ankles 
brownish; tail above with seven to eight narrow blackish rings and a 
black tip, alternating with light ochraceous buffy rings, less distinct 
and tending to become confluent below, especially near base. Varying 
in some specimens from paler and grayer to darker, with dorsum more 
profusely overspread with black, and rusty rufous nape patch indis- 
tinct or absent. Young (in fii'st pelage): Similar to P. I. hernandezii, 
but paler above, especially the top of head, which is scarcely differ- 
entiated from back. 

Cranial characters.- — Skull similar in size to that of P. I. hernandezii, 
but more massive; interorl)ital and postorbital regions broader; denti- 
tion about the same. Similar in size and angularity to that of P. I. 
crassidens, but less flattened, the frontal region higher arched behind 
postorbital processes; dentition and other cranial details about as in 
crassidens. Compared with that of P. I. dickeyi the skull is decidedly 
larger, more massive; sagittal and lambdoid crests heavier, thicker and 
less trenchant; palate broader; auditory bullae usually larger. 

Measurements. — Type: Total length, 874 mm.; tail vertebrae, 292; hind foot, 
116. An adult female topotype; 909; 296; 128. Skull: Type: Greatest length, 



RACCOONS OF NORTH AND MIDDLE AMERICA 67 

126.1; condylobasal length, 118.7; interorbital breadth, 26.8; least width of palatal 
shelf, 16.8; maxillary tooth row (alveoli), 45.1; upper carnassial, crown length, 
9.6, crown width, 9.8. 

Remarks. — The g-eiioral range of the present su})species embraces the 
peninsula of Yucatan and adjoining territory as far south and west as 
the Isthmus of Tehuantepec. Like the representatives of other widely 
ranging subspecies inhabiting the general region it is characterized by 
pale colors. Occasional specimens, however, as one from Huilotepec 
(near Tehuantepec) , Oaxaca, have the upper parts more heavily over- 
laid with black, indicating gradation toward the darker Central 
American forms. It is closely allied to P. I. hcrnamlezii, but the 
characters pointed out are distinctive. 

Specimens examined. — Total number, 23, as follows: 
British Honduras: El Cayo (near San Lorenzo), l.^"* 
Campeche: La Tuxpena (type locality), 3. 
Chiapas: San Vicente, 1 (skull only). 

Guatemala: El E.'^pino, 1; northern Guatemala (exact locality unknown), 1. 
Honduras: Santa Barbara, 1.^" 

Oaxaca: Huilotepec, 7; San Mateo del Mar, 1 (skull only); Tehuantepec, 4.^8 
Tabasco: Montecristo, 1. 
Veracruz: ]\Iinatitlan, 1. 
Yucatan: Chichen Itza, 1. 

PROCYON LOTOR DICKEYI Nelson and Goldman 
Salvador Raccoon 

Proci/on lotor dickei/i Nelson and Cioldman, Biol. Soc. Washington Proc. 44: 18, 
Feb. 21, 1931. 

Type locality. — Barra de Santiago, Department of Ahuachai)am, 
Salvador. 

Type. — No. 12796, male adult, skin and skull, collection of Donald 
R. Dickey; collected by G. D. Stirton, April 14, 1927. 

Distribution. — Coast region of southwestern Salvador and probably 
of southeastern Guatemala; limits of range unknown. Tropical Zone. 

General characters. — A dark-colored subspecies (one of the darkest 
of the group) of medium size; skull short and light in structure. Color 
about as in P. I. crassidens of Costa Rica; size similar, but skull of 
lighter proportions, and differing in important details. Similar in 
general to P. I. shufeldti of Campeche, but smaller, and much darker, 
the upper parts more heavily overlaid with black; cranial characters 
distinctive. 

Color. — Upper parts in general grayish, heavily and rather uniformly 
overlaid with black extending well down along sides; light subterminal 
zone of longer hairs narrow^ and dark undercolor showing through in- 



36 Univ. Michigan Mus. Zool. 

3' Amer. Mus. Nat. Hist. 

3* One sliin witliout skull, one skull without skin, .\mor. Mus. Nat. Hist. 



68 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

tensifying general dark tone; top of head clearer gray, heavily mixed 
with black, producing a somewhat grizzled effect, the black predom- 
inating; black facial mask extending downward on median line to nose 
and upward to middle of forehead; white supraorbital lines short and 
narrow, ending under ears instead of continuing posteriorly to sides of 
neck as in shufeldti and more northern forms; sides of muzzle, lips, and 
chin white; under parts in general thinly overlaid with buffy white, 
the underfur light brownish, sparse and only partially concealing the 
skin beneath; throat patch brownish black; ears grayish; black post- 
auricular spots small, tending to blend with dark tone of back; fore- 
arms dull grayish, becoming soiled whitish on feet; outer surfaces of 
hind legs similar to sides of body, becoming brownish black near heels 
and soiled whitish on feet; tail above with about seven blackish rings, 
rather indistinct near base, and a black tip, alternating with rich 
ochraceous buffy rings, tending to blend along median line below. 

Cranial characters. — Skull characterized by thin-walled, delicate 
structure, with weakly developed sagittal and lambdoid crests. Most 
closely resembling that of P. I. cra'Ssidens, but of lighter proportions; 
frontal region less flattened; palate much narrower, a character very 
noticealile in the lesser distance between cheek tooth series; jugal more 
slender; dentition heavy, much as in cras.sirlens. Compared with that 
of P. I. shvfelfJti the skull is decidedly smaller and less massive ; frontal 
region of similar elevation; sagittal and lambdoid crests weaker, 
thinner and more trenchant; palate narrower; auditory bullae usually 
smaller; dentition about the same. 

Measurements. — Type: Total length, 840 mm.; tail vertebrae, 310; hind foot, 
115. Average of four adult male topotypes: 840 (800-870) mm.; 297 (.300-340); 
114 (110-120). Average of eight adult female topotypes: 782 (730-790); 300 
(280-340) ; 1 10 (105-1 20) . Skull: Type and an adult male topotype, respectively : 
Greatest length, 114.7, 108.3; coiidylobasal length, 108.4, 102.7; zygomatic 
breadth, 79.3, 76.2; interorbital breadth, 23.9, 22.5; width of palate between last 
molars, 19.5, 20.9; least width" of palatal shelf, 16, 15.3; maxillary tooth row 
(alveoli), 41.7, 40.7; upper carnassial, crown length, 7.7, 8.5, crown width, 9, 8.9. 
Average of seven adult female topotypes: Greatest length, 116.1 (113-122.5); 
co-;dylobasal length, 109.7 (107.5-115.9); zygomatic breadth, 74 (70.2-80); inter- 
orbital breadth, 23.9 (22.9-25.3); width of palate between last molars, 19.7 (18.4- 
21); least width of palatal shelf, 15.2 (14.7-15.7); maxillary tooth row^ 43.7 
(42.7-45.7); upper carnassial, crown length, 9.2 (8.7-10), crown width, 10 
(9.5-11). 

Remarks. — P. I. dickeyi is the most northern of the known Central 
American subspecies, all of which are characterized by darker color 
than their more northern relatives. It appears to be a highly special- 
ized mangrove-inhabiting race as specimens from the interior only a 
short distance away are markedly dift'erent and nearer to crassidens. 
In external appearance this subspecies is similar to P. I. crassidens, but 
the cranial features are quite distinctive. The rusty rufous nape 



RACCOONS OF NORTH AND MIDDLE AMERICA 69 

patch often present in more northern forms is absent or only faintly 
indicated in some individuals. In the type locality it was found by 
the collector living among mangroves where specimens were obtained 
by shooting. Examination of stomach contents revealed crabs, which 
appear to be the principal food. In all of the skulls, including that of 
a young individual about two-thirds grown, the large cheek teeth are 
much more worn than is usual in raccoons of corresponding ages. This 
excessive wear, greatest on the molars, is due evidently to the abrasive 
character of the food. The delicate cranial structure and rapid reduc- 
tion of the molars also suggest that malnutrition resulting from an 
imperfect diet, or incomplete mastication of food, may have been 
responsible for the development of the peculiar characters of this 
localized race. 

Specimens examined. — Total luunber, 22, as follows: 
Guatemala: Exact locality unknown, 5. 

Salvador: Barra de Santiago, Department of Ahuachapam (type locality), 17 (4 
skins without skulls). ^^ 

PROCYOX LOTOR CRASSIDENS Hollister 
Costa Rican Raccoon 

Proci/on lotor crassidens Hollister, Biol. Soc. Washington Proc. 27: 142, July 10, 
1914. 

Type locality. — Talamanca, northeastern Costa Rica. 

12191 
Type. — No. , axhilt [male?], skin and skidl, United States 

National Museum; collected by William M. Gabl). Original ntimber 
14. 

Distribution. — Costa Rica, Nicaragua, Salvador, except south- 
western coast region and probably Honduras, except western part; 
probably extending into western Panama. Tropical Zone. 

General characters. — One of the darkest known forms of the group; 
closely resembling P. I. jnimilus of Panama and P. Z. dickeyi of Salvador 
externally, but cranial characters distinctive. 

Color. — About as in P. I. dickeyi, the dorsum heavily overlaid with 
black extending well down the sides; white supraorbital lines distinct, 
but short and disappearing under the ears as in dickeyi. 

Cranial characters. — Skull similar to that of P. I. dickeyi, but more 
massive; frontal region more flattened; palate much broader, the 
tooth rows more widely separated; dentition heavy much as in 
dickeyi. Compared with that of P. I. pumilus the skull is larger, 
relatively longer, narrower, and less extremely flattened; inter- 
orbital and postorbital regions iiarrowers; postorbital processes of 



" Donald R. Dickey collection. 



70 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SEBVICE 

front als shorter, broader and more obtusely pointed; dentition similar 
but usually heavier. 

Measureiuents. — An adult male from Jalapa, Nicaragua: Total length, 950 mm.; 
tail vertebrae, 310; hind foot, 120. An adult male from San Rafael del Norte, 
Nicaragua: 880; 250; 110. Skull: Type: Greatest length, 125.5; condylobasal 
length, 122.9; zygomatic breadth, 75.8; interorbital breadth, 25.8; width of 
palate between last molars, 24.1; least width of palatal shelf, 17.3; maxillary 
tooth row (alveoli), 47.3; upper carnassial, crown length, 10, crown width, 10.7. 

Remarks. — P. I. crassidens is similar to P. I. purnilus and P. I. 
dickey I in external appearance, all sharing an extremely dark colora- 
tion. While closely allied to the forms mentioned, the cranial char- 
acters presented are quite distinctive. Some specimens from the 
interior and southeastern coast region of Salvador, quite near the 
restricted range of dickeyi, are distinctly grayer than typical crassidens, 
and in this character, as in cranial details, grade toward shurfeldti. 

Sj)ecrm.ens examined. — Total number, 18, as follows: 
Costa Rica: L]l Sauce Peralta, 1; Talamanca, 1 (type); exact locality unknown, 1. 
Nicaragua: Jalapa, 2;^° San Rafael del Norte, 2;^" Vijagua, 1.^" 
Salvador: Barrios Mine, Morazan, 1;-" Colima, Cuscatlan, 1;^' Lake Guija, l;''^ 
Puerto del Triunfo, Usulutan, 1;*' Rio Goascoran, La Union, 1;^^ Rio San 
Miguel, 3;*' San Pedro Mine, Morazan, 1;" Volcan San Miguel, 1." 

PROCYON LOTOR PUMILUS Miller 

ISTHMI.W R.\CCOON 

Procyon pumihis Miller, Biol. Soc. Washington Proc. 24: 3, Jan. 28, 1911. 

Type locality. — Ancon, Panama. 

Type. — No. 171983, young adult [female?], skin and skull. United 
States National Museum; collected by Allan H. Jennings, 1910. 

Distribution. — Panama and the Canal Zone from Porto Bello west 
to Boqueron, Chiriqui, limits of range unknown. Tropical Zone. 

General characters. — Closely allied to P. I. crassidens of Costa Rica; 
color very similar; skull shorter, relatively broader and flatter. 

Color. — Very dark, the upper parts heavily overlaid with black 
about as in P. I. crassidens, but white supraorbital lines usually less 
distinct, somewhat obscured by dusky hairs. 

Cranial characters. — Skull smaller, shorter, relatively broader, and 
still flatter than that of P. I. crassidens; interorbital and postorbital 
regions broader; postorbital processes longer, narrower, more acutely 
pointed; dentition lighter, especially the cheek teeth distinctly smaller. 

Measurements. — Adult male from Porto Bello, Panama: Total length, 920 mm.; 
tail vertebrae, 350; hind foot, 125. Adult female froni Gatun, Canal Zone; 
831; 292; 123. Skull: Adult male from Porto Bello, Panama and adult female 
from Gatun, Canal Zone, respectively: Greatest length, 113.5, 113.2; condylo- 



w Amer. Mus. Nat. Hist. 

<• Donald R. Dickey collection. 



RACCOONS OF NORTH AND MIDDLE AMERICA 71 

basal length, 110.8, 110.5; zygomatic breadth, 81.1, 80.6; interorbital breadth, 
20.8, 24.8; least width of palatal shelf, 15.1, 14.8; maxillary tooth row, 44, 41.9; 
upjier carnassial, crown length, 8.9, 7.6, crown width, 9.6, 9. 

Ucmarkx. — The range of P. I. putnilus marks the known extreme 
southern hmit of the Proci/on lotor group. Its distribution area 
overhips that of Procyon ccnicrirortis panamensis, the so-ealled crab- 
eating raccoon, tlie two occui'ring in tlie same locahties in the Canal 
Zone and vicinity. From the hitter it is easily distinguished extern- 
ally by its smaller size, more slender proportions, grayish instead of 
blackish forearms and thighs, presence of underfur, and the normal 
iuclination backward of the pelage of the nape which in the cral)- 
eating raccoon is reversed. The skull is recognizable especially 
l)y the smaller molars, with more pointed instead of rounded cusps. 
Although the dentition of pumilus is not so well fitted as that of the 
crab-eating raccoon for crushing hard substances such as crabs, it 
shares with it the crab-eating habit, at least to some extent, as shown 
by the examination of stomach contents. 

P. I. i)um.ilus is most closely allied to P. I. crassidens. In external 
appearance some specimens of the two are practically indistinguisha- 
ble, although the white supraorbital lines are usually less distinct in 
})umUu-'<: but the skull is notable for its shortness; and in the general 
flatness, and length of the postorbital processes it reaches the extreme 
development presented in the group. Material now available, 
including a series of six topotypes (Balboa and Ancon, the type locality, 
ai-e contiguous), shows that this raccoon is not so very small as the 
type, an unusiutlly tnider-sized and not fully adidt individual, seemed 
to indicate. 

Specimens examineel. — Total ninnber, 15, as follows: 

Canal Zone: .\ncon, 1 (type); Ball^oa, 6;^- Gatvui, 4. 

Panama: Boqueron, 1;^^ Pedregal, 1;" Porto Bello, 2. 

PROCYON INSULARIS Merriam 

[References under sub-species] 

Distribution. — Tres Marias Islands, off west coast of Nayarit, 
Mexico. 

General characters. — A large, pale species, with short, coarse pelage 
and massive skull. Similar to adjacent mainland forms of P. lotor 
(P. I. mexicanus and P. I. hernandezii), but pelage shorter, more 
bristly, color inclining toward huffy instead of iron grayish, the back 
less overlaid with black; black postauricular spots much smaller, less 
conspicuous; skull more angular and dift'ering in important details. 



>■ Chicago Mus. Nat. Hist. 
" Amer. Mus. Nat. Hist. 
^* Mus. Comp. Zool. 



72 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

Color. — Upper parts in general light cream buff, the dorsal area 
thinly overlaid with black; nuchal patch undifferentiated or faintly 
indicated by a very pale buffy line ; sides lighter, the black-tipped hairs 
inconspicuous; top of head grizzled gray and black; black mask 
extending across face and along median line from nasal pad to middle 
of forehead; white supraorbital lines continuous to sides of neck; 
sides of muzzle, lips, and chin white; under parts, in general, thinly 
overlaid with very pale creamy buff", the light brown underfur show- 
ing through; throat patch brownish flecked with gray; ears grayish, 
the black patches at posterior base, usual in the group, obsolescent; 
limbs about like sides, becoming dull whitish on feet; hind legs 
brownish on outer sides near ankles; tail above with about seven 
black rings and a black tip, alternating with broader cream buffy 
or light ochraceous buffy rings, the dark rings interrupted below. 

Cranial characters. — Skull large, angular, and massive, with remark- 
ably heavy zygomata, the squamosal arm, especially, very broad 
anteriorly and extended vertically (as apparent when viewed from 
<^he side). Similar in general to that of P. lotor, especially , P. I. 
mexicanus and P. I. hernandezii, but more angular; zygomata broader 
and heavier, the squamosal arm broader anteriorly, more extended 
vertically; transverse squamosal portion of zygoma bearing a more 
conspicuous process on anterior border near posterior end of jugal; 
palatal shelf relatively narrower, the lateral borders more deeply 
concave; postorbital processes of frontals well developed as in mexi- 
canus and hernandezii; large molariform teeth narrower; crown of 
second upper molar subquadrate, instead of subtriangular, the inner 
border more evenly rounded. 

Remarks. — P. insularis is clearly allied to P. I. mexicanus and 
P. I. hernandezii of the adjacent mainland and was regarded by its 
describer as a subspecies of the widely ranging continental animal. 
The characters pointed out are so trenchant, however, that its position 
in the group is better expressed by according it specific rank. It is 
subdivisible into two closely related insular forms. 

PROCYON INSULARIS INSULARIS Merriam 
MARfA Madre Island Raccoon 
Procyon lotor insularis Merriam, Biol. Soc. Washington Proc. 12: 17, Jan. 27, 1898. 
Type locality. — Maria Madre Island, Tres Marias Islands, off west 
coast of Nayarit, Mexico. 

Type. — No. 88978, old male, skin and skull, United States National 
Museum (Biological Surveys collection) ; collected by E. W. Nelson 
and E. A. Goldman, May 10, 1897. 



RACCOONS OF NORTH AND MIDDLE AMERICA 73 

Distribution. — Known only from Maria Madre Island. Tropical 
Zone. 

General characters. — Closely resembling P. i. vicinus of Maria Mag- 
dalena Island, but dorsum less conspicuously overlaid with black, and 
top of head grayer; cranial characters distinctive. 

Color. — About as set forth for the species as a whole, differing only 
slightly from P. i. ricinus in the somewhat lesser amount of overlying 
black. 

Cranial characters. — Skull very similar to that of P. i. vicinus, but 
brain case less highly arched; lambdoid crest more broadly spreading, 
not rising so high over foramen magnum; basioccipital, basisphenoid, 
and palatal shelf broader; palatal ridges (extending posteriorly to 
]iterygoids) more widely separated; pterygoids thicker, the posterior 
ends more strongly everted and knob-like; maxillary arm of zygoma 
with lower external border projecting as a distinct process separated 
from outer alveolar border of molars by a deep notch (process absent 
in vicinus) ; zygomata very broad and heavy as in vicinus; foramen 
magnum more decidedly wider than high (more nearly circular in 
vicinus); dentition about the same. 

Measurcment.s. — Type: Total length, 854 mm.; tail vertebrae, 286; hind foot, 
132. An adult male topotype: 840; 264; 128. Sktdl: Type and an adult male 
topotype, respectively: Greatest length, 121.8, 119; condylobasal length, 114.6, 
114; zygomatic breadth, 86.4, 82.5; interorbital breadth, 27.8, 27.2; least width of 
palatal shelf, 15.4, 14.3; maxillary tooth row, 43.6, 42.2 (alveoli): upper carnas- 
sial, crown length, 8.6, 8.6, crown width, 9.3, 9.1. 

Reiriarks. — P. i. insularis requires close comparison only with P. i. 
vicinus of Maria Magdalena Island. Whil(> cranial details appear to 
be cpiite distinctive these insular forms are much alike in external 
appearance. In the few specimens available, however, the black over- 
lying the dorsum — rather thin in nicinu'^ — is further reduced in 
insulari'^, leaving a coarsely grizzled effect. 

Specimens examined. — Six, all from the type locality. 

PROCYON INSULARIS VICINUS Nelso.\ and Goldman 
Maria Magdalena Island Raccoon 

Procyon insulans iiciniis Nelson and (Joldinan, Biol. Soc. Washington Proc. 44: 
20, Feb. 21, 1931. 

Tyjje locality. — Maria Magdalena Island, Tres Marias Islands, 
Nayarit, Mexico (altitude 250 feet). 

Tyjje.—Ko. 88982, male adtdt, skin and skull. United States National 
Museum (Biological Surveys collection) ; collected by E. W. Nelson 
and E. A. Goldman, May 27, 1897. 

Distribution. — Known only from Maria Magdalena Island. Trop- 
ical Zone. 

876119°— 50 6 



74 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

General character.^. — A pale subspecies with short, coarse pelage. 
Closely reseinl)ling Procyon i. inxvlaris of Maria Madre Island, but 
dorsum more conspicuously overlaid with black, and top of head some- 
what darker; cranial characters distinctive. 

Color. — Upper parts in general light cream buff, the dorsal area 
rather thinly overlaid with black; sides lighter, the black-tipped hairs 
inconspicuous; top of head gray mi.xed with l)lack, giving a grizzled 
effect ; black mask across face extending downward to nose and up- 
ward on median line to middle of forehead; white supraorbital mark- 
ings normal; sides of muzzle, lips, and chin white; under parts in gen- 
eral thinly overlaid with very pale creamy buff', the light brown under- 
fur showing through; throat patch brownish flecked with gray; ears 
grayish, the black patches at posterior base, usual in the group, obso- 
lescent; legs about like sides, becoming dull whitish on feet; hind legs 
brownish on outer sides near ankles; tail above with seven black rings 
and a black tip, alternating with broader cream buff' rings, the dark 
rings interrupted below. 

Cranial characters. — Skull very similar to that of P. i. insularis, but 
brain case more highly arched; lam])doid crest rising higher over fora- 
men magnum; basioccipital, basisphenoid, and palatal shelf narrower; 
palatal ridges (extending posteriorly to pterygoids) less widely sep- 
arated; pterygoids thinner, the posterior ends less everted; maxillary 
arm of zygoma normal, the lower external border not projecting and 
forming a distinct process separated from outer alveolar border of 
molars by a deep notch; zygomata very bioad and heavy, as in insu- 
laris; foramen magnum more nearly circular (more decidedly wider 
than high in insularis); dentition about the same. 

Mrannrenientfi. — Type: Total length, 904 mm.; tail vertebrae, 313; hind foot, 
135. Skull: Type: Greate.st length, 120; coiidyloba.sal length, 115.2; zygomatic 
breadth, 84.6; interorbital breadth, 27.7; least width of palatal shelf, 14.1; ma.xil- 
lary tooth row (alveoli), 42.6; upper carnassial, crown length, 8.7, crown width, 9.2. 

Remarks. — As might be expected P. i. ricinus is closely allied to its 
near geographic neighbor, P. i. insularis of Maria Madre Island, and 
requires no very close comparison with any other form. It is distin- 
guished externally from adjacent mainland forms, P. I. mexicanus and 
P. I. hernandesii, by shorter, coarser pelage, the general color inclining 
toward buffy instead of grayish, and the black postauricular spots 
obsolescent; the skidl dift'ers in numerous important details, especially 
the higher arched brain case, much broader, heavier, zygomata, nar- 
rower palatal shelf, and narrower carnassials. 

Si^ecimens examined. — Two, from the type locality. 



RACCOONS OF NORTH AND MIDDLE AMP]RICA 75 

PROCYON MAYNARDI Bangs 
Bahama Raccoon 
Procijon inaynardi Bangs, Biol. Soc. Washington Proc. 12: 92, Apr. 30, 1898. 

Type locality. — New Providence Island, Bahamas. 

Type. — No. 7750, male young, skin and skull, Museum of Com- 
parative Zoology (collection of E. A. and O. Bangs); collected by 
Herbert L. Claridge, August 1897. 

Distribution. — Known only from New Providence Island, Bahamas. 
Tropical Zone. 

General characters. — A small, medium (hirk-colored species with a 
slender, delicate skull, narrow palatal shelf, and light dentition. 
J^imilar in general to P. I. incautus of the extreme southern Florida 
l<eys, Init color darker, and cranial characters, especially the small 
teeth, distinctive. Somewhat similar in size to P. minor of Guade- 
loupe Island, Lesser Antilles, but apparently somewhat paler in color 
and skull differing notably in the narrowness of the palatal shelf. 

Color. — Upper parts in general grayish, becoming ochraceous buffy 
on nape and over shoulders, moderately overlaid with black, thinning 
out along sides; top of head a grizzled mixture of gray and black; black 
mask interrupted between eyes, a dusky median patch extending to 
forehead somewhat isolated by lighter lateral lines, as in P. I. lotor; 
upper surface of muzzle ochraceous buffy; supraorbital lines, sides of 
muzzle, lips, and chin white; ears grayish, with black patches at 
posterior base; under parts thinly overlaid with grayish; limbs similar 
to under parts, the hind legs blackish near ankles; tail with five or six 
black rings and a black tip, alternating with ochraceous buffy rings. 

Cranial characters. — Skull very similar in outline and general propor- 
tions to that of P. I. incautus, but palatal shelf narrower, the sides 
distinctly concave (sides more nearly parallel in incautus) ; nasals 
narrower posteriorly; auditory bullae slightly larger, more inflated; 
posterior upper premolar and carnassial slightly smaller. Similar in 
size to that of P. minor, but brain case narrower, less flattened above; 
palatal shelf decidedly narrower, the sides more concave; nasals nar- 
rower between anterior processes of frontals; auditory bullae slightly 
larger; dentition similar, but molariform teeth broader. 

Measurements. — Adult Male topotype: Total length, 713 mm.; tail vertebrae, 
240; hind foot (dry skin), 100. Stcidl: An adult male and an adult female (topo- 
types), respectively: Greatest length, 105.9, 103.5; condylobasal length, 101.7, 
101; zygomatic breadth, 79.3, 64.6; interorbital breadth, 21.9, 22.1; least width of 
palatal shelf, 12.2, 13.3; maxillary tooth row (alveoli), 38.7, 38.9; upper carnassial, 
crown length, 7.6, 7.6, crown width, 8.4, 8.5. 

Remarks. — The Bahama form is closely related to the raccoons of the 
Florida Keys as shown in the skull by agreement in form and general 
proportions, especially the height of the frontal region, with slight 



76 NORTH AMERICAN FAUNA 6 0, FISH AND WILDLIFE SERVICE 

development of postorbital processes, and the depression of the bram 
case near the fronto-parietal suture. The differential characters, 
however, warrant its recognition as a distinct species. In describing 
P. maynardi Bangs (1898b, p. 92) says: "There is no tradition among 
the inhabitants of Nassau that the raccoon was ever introduced upon 
the island. . . . The raccoon is abundant upon Nassau [New Provi- 
dence] but Mr. Maynard believes that it does not exist upon any of the 
other islands of the Bahama group." According to the Acting 
Colonial Secretary Charles P. Bethel, the raccoons on New Providence 
Island have decreased during recent years owing to the destruction by 
hurricanes of fruit trees that afforded a food supply. 

Specimens examined. — Three, ^^ all from New Providence Island. 

PROCYON PYGMAEUS Merriam 

CozuMEL Island Raccoon 

Procyon pygmaeus Merriam, Biol. Soc. Washington Proc. 14: 101, July 19, 1901. 

Tyj^e locality. — Cozumel Island, Yucatan, Mexico. Tropical Zone. 

Type. — No. 108511, male subadult, skin and skull. United States 
National Museum (Biological Surveys collection) ; collected by E. W. 
Nelson and E. A. Goldman, April 14, 1901. 

Distribution. — Known only from Cozumel Island. 

General characters. — Smallest known species of the genus, with short, 
bristly, grayish pelage; skull with short, posteriorly rounded nasals and 
very small teeth. Somewhat similar in color and texture of pelage to 
P. I. sltufeldti, of the adjacent mainland, but so much smaller and 
cranial characters so distinctive that close comparison is not required. 

Color. — Upper parts in general light buffy gray, the median dorsal 
area suffused with pale buff, becoming more pronounced and approach- 
ing ochraceous buff on a narrow nuchal patch in some specimens, 
rather thinly overlaid with black; top of head clearer, grizzled gray and 
black, lac^'ing light buffy tone suff'using back; black mask becoming 
brownish and usually more or less mixed with gray on middle of face, 
the gray admixture invading also the dark median streak extending to 
the forehead; upper surface of muzzle brownish; lines bordering mask 
above, sides of muzzle, lips, and chin white; under parts, in general, 
thinly overlaid with light buft'y hairs, the light brownish underfur 
showing through; throat patch dark brownish, clearly defined; ears 
grayish or light buffy; postauricular spots brownish, small and incon- 
spicuous; legs similar to untler parts, becoming dull whitish on feet, 
the hinil legs with outer sides of ankles clearer brownish; tail with six 
or seven narrow, brownish or blackish rings and a black tip, alternating 
with broader ochraceous buffy rings, the dark rings ill-defined on 
under side. 

*5 One in Amer. Mus. Nat. Hist.; one in Mus. Comp. Zool. 



RACCOONS OF NORTH AND MIDDLE AMERICA 77 

Cranial characters. — Skull small, short, and flattened, with relatively 
short, narrow rostrum, short nasals, broad frontal region and brain 
case, and light dentition. Somewhat similar in general form to that 
of P. I. shvfeldti, but departing widely in the smaller size; rostrum 
relatively shorter and narrower; frontal region relatively broader; 
nasals relatively shorter, more rounded, less acutely pointed pos- 
teriorly; postorbital processes of frontals well-developed and upper 
border of orbit distinctly concave as in shufeldti; teeth similar in 
sculpture, but relatively much smaller, the first and second upper 
premolars more widely spaced, and the last molar with a narrower 
internal lobe. Compared with those of P. maynardi and P. minor, 
the skull is smaller, with rostrum shorter, frontal region flatter and 
relatively broader than in either; nasals shorter, broader and more 
rounded posteriorly; palatal shelf narrow much as in maynardi (much 
narrower than in minor); auditory bullae smaller than in either; 
postorbital processes of frontals more developed; dentition similar 
but lighter. 

Measurements. — Type: Total length, 667 mm.; tail vertebrae, 230; hind foot, 
90. Adult femalet opotype: 665; 250; 97. Skull: Type and adult female 
topotype, respectively: Greatest length, 100, 96.7; condylobasal length, 93.7, 
91.9; zygomatic breadth, 58.8, 60.8; interorbital breadth, 19.5, 19.8; least width 
of palatal shelf, 12.5, 12.3; maxillary tooth row (alveoli), 35.3, 35.5; upper car- 
nassial, crown length, 6.8, 7, crown width, 7.8, 8. 

Remarks. — P. pygmaeus, as the name suggests, is distinguished by 
its small size. The general flattening of the cranium, especially the 
flatness and breadth of the frontal region, the development of the 
postorbital processes of the frontals, and the slight depression of the 
brain case near the fronto-parietal suture, indicate relationship to the 
raccoon of the adjacent mainland as might be expected, rather than 
to any of the West Indian species. Striking differences from the 
mainland animal in size and in other more important characters, 
however, point to long isolation in its insular habitat. The teeth arc 
remarkably small, the second upper premolar especially, being reduced 
in size and separated from the third upper premolar by a distinct gap. 

Specimens examined. — Five, all from the type locality. 

PROCYON MINOR Miller 
Guadeloupe Island Raccoon 
Procyon minor Miller, Biol. Soc. Washington Proc. 24: 4, Jan. 28, 1911. 

Type locality. — Pointe-a-Pitre, Guadeloupe Island, Lesser Antilles. 

^ ^^ 38417 

lype. — No. ^ -, male young (permanent canmes not quite fully 

in place), skin and skufl. United States National Museum; coflected 
by L. Guesde. Received from the THerminier Museum. 



78 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

Distribution. — Known only from Guadeloupe Island. Tropical 
Zone. 

General characters. — A small, rather dark species with a slender, 
delicate skull, remarkably broad palatal shelf, and very light dentition. 
Similar to P. maynardi, of the Bahamas, but apparently somewhat 
darker in color, and cranial characters, especially the much broader 
palatal shelf, distinctive. 

Color. — Upper parts in general grayish, becoming "ochraceous buff" 
on nape and shoulders, the dorsum heavily overlaid with black; 
sides paler and almost silvery gray, the dark-tipped hairs thinning out; 
top of head whitish mixed with black, with the usual grizzled effect; 
black mask continuous across face in one specimen, somewhat inter- 
rupted between eyes in another; sides of muzzle, lips, chin, and 
supraorbital lines white; ears grayish, with large, conspicuous black 
patches at posterior base; under parts thinly overlaid with grayish, 
the light })rown underfur showing through; throat patch blackish; 
forearms, hind legs and feet grayish, similar to sides, the ankles 
blackish; tail with about seven black rings and a black tip, alternating 
with ochraceous bufl'y rings. 

Cranial characters. — Skull similar in size and general form to that 
of P. maynardi, but brain case broader and flatter; palatal shelf 
much broader, the sides forming nearly straight parallel lines (sides 
more concave in maynardi) ; nasals broader between anterior processes 
of f rentals; auditory bullae slightly smaller; dentition similar, but 
molariform teeth narrower. 

Measurements. — No reliable skin measurements available. SkuU: Type and a 
subadult topotype, respectively: Greatest length, 101.6 mm., 104.5; condylobasal 
length, 94.5, 98.5; zygomatic breath, 55.3, 62; interorbital breadth, 18.1, 19.8; 
least width of palatal shelf, 16.5, 15.5; maxillary tooth row (alveoli), 38.5, 37.3; 
upper carnassial, crown length, 7.6, 7.6, crown width, 8.1, 8.1. 

Remarks. — Although widely separated geographically, P. minor 
appears to be more nearly related to P. maynardi than to any other 
known form. The elevation of the frontal region, absence or slight 
prominence of the postorbital processes of the frontals, and the 
depression of the brain case near the fronto-parietal suture are 
characters denoting alliance with inaynardi and the raccoons of the 
Florida region. It requires no close cranial comparison with P. 
gloveralleni of Barbados, the skull of which is distinguished at a glance 
by the larger molariform teeth. 

Specimens examined. — Two, the type, and a topotype. '''^ 



<8 Mus. Comp. Zool. 



RACCOONS OF NORTH AND MIDDLE AMERICA 79 

PROCYOX OLOVERALLENI x\elson and Goldman 
Bakbados Raccoon 

Procyou (iJorcraJleni Nelson and Goldman, Jour. Mammal. 11 (4): 453, Nov. 11, 
1930. 

Type locality. — Island of Barbados, Lesser Antilles, West Indies. 

Type. — No. 18591, young male, skin and skull, Museum of Com- 
parative Zoology; collected by Sir Francis Watts, 1920. 

Disfribufion. — Known only from the Island of Barbados. Tropical 
Zone. 

General characters. — A small, dark species, with a short, delicately 
formed skull. Similar in color to Procyon minor, of Guadeloupe 
Island, Lesser Antilles, but sides of body and limbs in type specimen 
darker, owing to more numerous black-tipped hairs (black-tipped 
hairs thinning out and sides of body and limbs more grayish in minor) ; 
cranial characters, especially the much heavier dentition, distinctive. 
Contrasting with P. maynardi, of New Providence Island, Bahamas, in 
darker general color and widely different skull. 

Color. — Type: Upper parts in general near "light ochraceous buff" 
(most intense on nape and shoulders) rather heavily overlaid with 
black, becoming lighter buff, less obscured by black on sides of body 
and limbs; top of head buffy gray, mixed with black; face with solid 
black mask and usual white markings; the upper surface of muzzle 
black to nose; sides of muzzle, lips, and chin white; under parts thinly 
overlaid with buffy grayish; throat patch brownish black, thinly 
overlapped by ochraceous buffy hairs, here reversed as usual in the 
group; ears buffy grayish with black patches at posterior base; 
ankles dusky all around; feet soiled buffy whitish; tail with four 
narrow black rings and a black tip alternating with light ochraceous 
])uffy rings, the subterminal black ring and tip nearly coalescent. 

Cranial characters.- — Skull similar in general to that of P. minor, 
but frontal region broader and flatter; postorbital processes more 
prominent; palatal shelf narrower; teeth very similar in sculpture, 
but crown of upper carnassial tending to be longer than broad, a 
condition unusual in the group. Compared with that of P. maynardi 
the skull differs in about the same characters as from minor, except 
that the palatal shelf is decidedly broader. 

Measurements. — Type: Hind foot (dry skin), 89 mm. SlcuU: Type: Greatest 
length, 94; condylobasal length, 89.2; zygomatic breadth, 53.4; interorbital 
breadth, 18.7; least width of palatal shelf, 13.6; maxillary tooth row (alveoli), 
37; upper carnassial, crown length, 9.8, crown width, 9. Two adult topotypes, 
No. 267380, female, and No. 267381, sex undetermined, United States National 
Museum, respectively: Greatest length, 113.2, 109.8; condylobasal length, — , 105.3; 
zygomatic breadth, — , 69; interorbital breadth, 24.3, 24.3; least width of palatal 
shelf, 15, 14.4; maxillary tooth row, 40, 39.8; upper carnassial, crown length, 
8.6, 8.7, crown width, 8.9, 8.6. 



80 NORTH AMERICAN FAUNA 6 0, FISH AND WILDLIFE SERVICE 

Remarks. — Dr. Glover M. Allen (1911, p. 221) recorded the occur- 
rence of raccoons in Barbados and referred to Griffith Hughes (1750, 
p. 66) who, writing in the middle of the 18th century, mentioned a 
law of the Island providing a bounty for their destruction. In the 
absence of specimens for study the animal was tentatively referred 
by Allen to "Procyori (?) cancrivorus G. Cuvier." Subsequent efforts 
by him to obtain representatives resulted in the collection, in 1920, 
of the young individual later made the type of a new species bearing 
his name. 

The type specimen was so young when collected that the permanent 
premolars and canines, although well advanced, are not in fidl func- 
tional position. In the type the first premolars, both deciduous and 
permanent, are absent in both jaws, an abnormality observed else- 
where only in the large-toothed form P. I. litoreus, inhabiting Saint 
Simon Island, Ga. Since the original description was published 
two specimens, in the exhibit collection of the United States National 
Museum, taken by the Reverend Barnett about 1867 have attracted 
attention and have been dismounted. The skulls show full maturity. 
One specimen, No. 267380, had been marked female, and the other, 
No. 267381, slightly smaller, is probably of the same sex. The 
molariform teeth are rather large, but not so large as in the type. 
In the upper carnassial a tendency toward equal to or greater length 
than width of the crown is e.xhi])ited, a condition sometimes presented 
in P. I. ])umihis of Panama. In the broad frontal region and well- 
developed postorbital processes the relationship of (jloreralleni to the 
raccoons of Central America is also suggested, but it dift'ers widely in 
other respects. 

Raccoons, formerly abundant, and said to favor a rugged region 
on the south sitk^ of the island, have apparently been reduced to or 
near extermination. In response to a formal inquiry the American 
Consul, Frederick W. Baldwin, wrote July 13, 1932: "Very few rac- 
coons now exist in Barbados and specimens would be extremely 
difficult to obtain." 

Specimens examined. — Three, all from the type locality. 

Subgenus EUPROCYON Gray 

[References under Genus Procyon Storr, p. 25] 

Distribution. — Southern Costa Rica, western Panama, and northern 
Colombia to southern Brazil. 

Subgeneric characters. — Contrasted with subgenus Procyon: Pelage 
shorter, underfur absent; hair on nape directed forward; claws broader, 
less compressed laterally, of lesser vertical diameter at base, and more 
bluntly pointed. Bony palate extending behind posterior molars a 
distance less than one-fourth total length of palate. Molariform 



RACCOONS OF NORTH AND MIDDLE AMERICA 



81 



teeth, except first premolars, larger and more massive, with broader, 
more rounded and bluntly pointed cusps; connecting ridges between 
principal cusps lower, less trenchant. 

Remarks. — The subgenus Euprocyon overlaps the range of the 
subgenus Procyon in Panama, but the two differ so conspicuously 
in appearance that no very close comparison is necessary. 

PROCYON CANCRIVORUS (G. Cuvier) 

Ursus cancrivorus G. Cuvier, Tabl. Elem. de I'Hist. Nat. des Aniinaux, 1798, 

p. 113. Type from Cayenne, French Guiana. 
Procyon cancrivorus Desmarest, Diet. Hist. Nat. 29: 93, 1819. 

Distribution. — (See under subgenus Eujirocyon.) 

General characters. — (See subgeneric characters under subgenus 
Euprocyon.) 

Color. — General dorsal area varying from ashy gray to ochraceous, 
more or less heavily overlaid with l)lack; ears, supraorbital streaks, 
and sides of muzzle whitish; black mask, usual in the group, extending 
across face to cheeks, including orbits, and the median line from fore- 
head to nose; under parts varying from pale gray to yellowish or 
ochraceous; outer sin-faces of forearms and thighs usually blackish; 
feet varying from gray to brown; tail with about seven or eight alter- 
nating black and gray or yellowish rings and a black tip. 

Cranial characters. — (See subgeneric characters under subgenus 
Euprocyon.) 

Rfmarks. — Few specimens of Procyon cancrivorus have been avail- 
able for study, l)ut general comparisons indicate that the sjiecies has 





w 


^ 


t 


Seal 


\^^yZ^/ 


"^ 




80 100 



Figure 2. — Distribution of Procyon cancrivorus pananiensis (subgenus 

Euprocyon) . 



82 NORTH AMERICAN FAUNA 6 0, FISH AND WILDLIFE SERVICE 

a wide range in South America, somewhat parallehng the great range 
of P. lotor in North America. A single subspecies extends into the 
region under review. 

PROCYON CANCRIVORUS PANAMENSIS (Goldman) 
Panama Crab-Eating Raccoon; Mapachin 

Euprocyon cancrivorus panamensis Goldman, Smithsn. Misc. Collection 60 (22) : 
15^ Feb. 28, 1913. 

Ti/2)e locality. — Gatun, Canal Zone, Panama. 

Type. — No. 171669, female adult, skin and skull. United States 
National Museum (Biological Surveys collection); collected by E. A. 
Goldman, June 21, 1911. 

Distribution. — Southern Costa Rica, western Panama to near the 
Colombian boundary; doubtless reaching Colombia, but range in that 
country undetermined. Tropical Zone. 

General characters. — A dark Panama representative of the species 
ranging widely in South America. Similar in size to P. c. proteus of 
northern Colombia, but general color less tawny; cranial details dis- 
tinctive. Differing from P. c. cancrivorus, of Cayenne, in darker color 
and in cranial characters. 

Color. — Ground color over dorsum varying from ashy gray to ochra- 
ceous buffy or yellowish ochraceous, heavily overlaid with black; top 
of head grizzled black and gray, the black predominating; sides of 
muzzle, and rather restricted supraorbital streaks, white or grayish 
white; facial area, including orbits, interorbital space, lower part of 
cheeks, and median line from forehead to nose, nearly clear black; 
under parts, including base of tail, varying from pale ochraceous buff 
to yellowish ochraceous, becoming more or less grayish white on throat, 
chin, and lips; ears well-clothed with whitish or yellowish hairs, dark- 
ening gradually on upper base by encroachment of body color; outer 
sides of hind legs and ankles all around deep glossy black; fore legs 
black or dark brownish all around ; feet thinly clothed with short hairs 
varying from brownish to grayish ; tail with seven or eight alternating 
black and grayish or yellowish rings and a black tip, the proximal rings 
more or less interrupted along median line below. 

Cranial characters. — In general form the skull closely resembles that 
of P. c. cancrivorus, but palate more elongated, lower surface of basioc- 
cipital more convex, the lateral margins turning downward and partly 
covering auditory bullae; nasals broader; auditory bullae broader, 
more inflated posteriorly; dentition about the same. Contrasted with 
that of P. c. proteus the skull differs in longer palate, and anteriorly 
broader, posteriorly narrower nasals. 

Measxirements. — Type: Total length, 950 mm.; tail vertebrae, 350; hind foot, 
142. Skull: Type: Greatest length, 130; condylobasal length, 125.8; zygomatic 



RACCOONS OF NORTH AND MIDDLE AMERICA 83 

breadth, 83.3; interorbital breadth, 25.7; least width of palatal shelf, 17.7; maxil- 
lary tooth row (alveoli), 48.3; upper carnassial, crown length, 10.2, crown width, 11. 

Remarks. — Comparison of the Panama series and South American 
material from various locahties, including a specimen from northern 
Brazil assumed to be near typical P. c. cancr'morus of Cayenne, and 
the type and two topotypes of P. c. proteus, of northern Colombia, indi- 
cates that the Panama animal is a well-marked geographic race. The 
close agreement in dentition and the other essential characters, how- 
ever, point to probable intergradation of all forms of cancrivorus. P. 
c. panamensis overlaps the range of Procyon lotor pumilus in Panama 
where both occur at the same localities, but may readily be distin- 
guished by the reversed pelage of the nape, absence of underfur, and 
the blackish instead of grayish forearms and thighs. 

Specimens examined. — Total number, 7, as follows: 

Canal Zone: Gatun (type locality), 3. 

Costa Rica: Canas Gordo, 1.^^ 

Panama: Boquete, 1 (skull only);** Cana, 1; Porto Bello, 1. 



<' Amer, Mus. Nat. Hist. 
<8 British Mus. (Nat. Hist.). 



APPENDIX 

Descriptions of two subspecies that were not included in the Gold- 
man manuscript are here abstracted or copied almost verbatim from 
the original accounts of the respective authors, but as nearly as 
possible in conformity of treatment with the Goldman manuscript. 

PROCYON LOTOR MEGALODOUS Lowery 

Mississippi Delta Raccoon 

Procyon lotor megalodous Lowery, Occas. Papers Miis. Zool. Louisiana State Univ. 
13: 225, November 22, 1943. 

Type locality. — Marsh Island, Iberia Parish, Louisiana. 

Type. — No. 2321, male adult, skin and skull, Louisiana State Univ. 
Mus. Zool.; collected by Ted O'Neil and prepared by George H. 
Lowery, Jr., October 24, 1943. 

Distribution. — Coast region of southern Louisiana from St. Bernard 
Parish west to Cameron Parish. 

General characters. — A medium-sized raccoon in which the pelage is 
strongly suffused above with black and pale yellow. Skull massive 
and with extremely large molariform teeth, by which characters it is 
separable from the two geographically adjacent subspecies P. I. varius 
and P. I. fuscipes. 

Color. — Nearest to P. I. varius, but distinguished in its much more 
yellowish (less grayish) suft'usion on upper parts and greater concen- 
tration of black along mid-dorsal line; ears, pale areas of face, legs, 
flanks and under parts decidedly yellowish, not grayish as in P. I. varius. 
Also much more yellowish (less grayish) than P. I. fuscipes, with black 
of dorsal midline more pronounced. 

Cranial characters. — Skull dift'ering from both P. I. varius and 
P. I. fuscipes in the larger size of the molariform teeth ; also dift'ering 
from that of P. I. varius in its more inflated frontal region, and lesser 
interorbital breadth. 

Measurements. — Type: Total length, 804 mm.; tail vertebrae, 262; hind foot, 
128. Skull: None available except a long table of measurements of molariform 
teeth (Lowery 1943, pp. 228-229). 

In his discussion of P. I. megalodous Lowery (1943) has made the 
following comments : 

Remarks. — This new race of raccoon, which is an abundant inhabitant of the 
Louisiana coastal marshes, is so clearly separable from all other races of Procyon 

84 



RACCOONS OF NORTH AND MIDDLE AMERICA 85 

lotor that it is surprising it has not been described until now. Superficially, it 
resembles varius of northern and eastern Louisiana, Mississippi, and Alabama, but 
its much more ^yellowish pelage which is strongly suffused with black, and its 
massive skull and large molariform teeth clearly set it apart from that form. In 
coat color this new race bears no close similarity to P. I. fuscipes of Texas, being 
distinctive as outlined above. However, the two agree with respect to certain 
cranial characters. In both the skull is massive, the frontal "hump" distinct, and 
the postocular constriction evident, but the dentition of megalodous is so decidedly 
heavier that skulls of the latter are separable from fuscipes without much difficulty. 
Three skins from Grand Terre Island are decidedly yellower than anything else 
examined in the present coiuiection. The dark middorsal area is restricted to a 
narrow but heavily concentrated band of dark brown (no black) ; hence the yellow- 
ish pelage of the sides and flanks is less suffused with dark hairs than in other 
raccoon specimens. The pelage of these specimens lacks any vestige of gray or 
black, the hairs being either yellowish or brown. Although there is a definite 
tendency among marsh dwelling raccoons to assume a decided xanthochronistic 
appearance in late spring and summer, this condition is clearly associated with 
wear, stain, and fading. These latter factors are not at all evident in the Grand 
Terre Island specimens, which are in fresh fall pelage. Whether this island popu- 
lation merits taxonomic recognition is dependent upon how constant the above 
noted characters appear in additional material, which is not obtainable at present. 

Specimens examined. — Total number, 20 skins and 41 skulls, as 
follo\vs : 

Louisiana: Cameron Parish: Rockefeller Refuge, 1 skin without skull; near 
Sabine Wildlife Refuge, 19 skulls. Calcasieu Parish: Lake Charles, 1 skin 
with skull. Terrebonne Parish: Timbalier Island, 1 skin with skull. Si. Ber- 
nard Parish: Toca Village, 1 skin with skull; Belair, 9 skins and 10 skulls. 
Plaquemines Parish: Delta [National \\'ilfllifel Refuge below Pilottown, 3 
skins and 4 skulls. Jefferson Parish: Grand Terre Island, 3 skins with skulls 
and one miscellaneous skull. Iberia Parish: Marsh Island, skin with skull 
(type). 

PROCYON LOTOR MARITIMUS Dozier 

Coastal Marsh Raccoon 

Proci/on lotor maritimus Dozier, Jour. Mammal. 29 (3): 286, August 31, 1948. 

Type locality. — Blackwater National Wildlife Refuge, Dorchester 
County, Marjdand. 

Type.— No. 275,290, female adult, skin and skull, U. S. Natl. Mus. 
(Biological Surveys collection) ; collected by Herbert L. Dozier, 
December 3, 1946. 

In describing P. I. maritirnus Dozier (1948) wrote: 

Distribution. — Known only from marsh areas on the Delniarva Peninsula 
(Delaware, Maryland, and Virginia). 

General characters. — A small to medium, pale subspecies. Readily distinguished 
from typical Procyon I. lotor by its paler coloration; longer but more sparse guard 
hairs; much smaller size; more slender legs and general build; narrower and more 
pointed head; decidedly shorter, more pointed, and less prominently banded tail; 
and relatively much shorter caudal vertebrae. In general color and length of the 
subapical band of the guard hair it is perhaps nearest to Procyon I. mexicanus 
of Arizona, New Mexico, and Mexico, but is only about half the size of the latter 
and its tail is decidedlv shorter and less distinctlv marked. 



86 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

Color. — Upper portion in general pale buffy grayish, becoming light ochraceous 
buffy on shoulders, with rufous tinge on nape; the longer, black-tipped guard 
hairs over the median dorsal area are grouped on the back as unevenly colored 
streaks, somewhat undulating or serpentine in arrangement (in marked contrast 
to the darker general salt and pepper effect of typical Procyon I. lotor) ; light- 
colored subapical band of each guard hair extends a greater distance beyond the 
underfur tips than in lotor and results in a much paler or more yellowish cast to 
the outer pelage; guard hairs almost entirely white along the sides and lower 
parts, adding to the pale, shaggy general appearance of the animal; top of head a 
light grizzled mixture of gray and brownish-black; facial mask brownish-black 
and decidedly less prominent than in lotor; feet black, with gray hair dorsally, the 
claws dull black (description from live and freshly killed animals) ; tail with five 
rather narrow, brownish-black rings and a black tip, alternating with wider light 
ochraceous buffy rings, less clearly defined below. 

Pelage. — An uneven spread of hair with respect to average guard hair length is 
apparent, that of the middorsal area being in general slightly shorter than in the 
rest of the pelt, due probably both to sparser distribution of hair and to variation 
in fiber length. The very long, coarse guard hair is typical of this race and unique 
among eastern races of raccoons. Due to the length and sparseness of the guard 
hairs, coupled with possible decreased density of the underfur, there is a slight 
backward slant to the direction of the hair flow. In lotor the hair is more dense, 
shorter, erect, and more fluffy in appearance than in maritimus. 

Skull. — Of medium size, rather narrow and elongate. Compared with that of 
lotor, the skull is slightly smaller, less heavily built, distinctly narrower, and more 
elongate; interorbital portion more elongate, postorbital processes of frontals 
rather weakly developed or obsolescent, frontal area relatively narrow, flatter, 
markedly more sloped or depressed, and somewhat concave; palatal shelf decidedly 
narrower; posterior part of zygomata less arched dorsally. 

Measurements. — Type: Total length, 718 mm; tail vertebrae, 210; 
hind foot, 111. Adult male topotype (No. 275,296, U. S. Natl. Mus.) : 
Total length, 762; tail vertebrae, 254; hind foot, 102. Skull: Type: 
Greatest length, 112.2; condylobasal length, 107.9; zygomatic breadth, 
68.8; interorbital breadth, 23.6; least width of palatal shelf, 15.1; 
maxillary tooth row, 41.7. 

To quote Dozier further: 

Remarks. — This form appears well adapted for survival under the rather 
exacting requirements of our eastern tidewater, coastal marsh conditions. The 
medium size and slender build fit it for fast traveling in the marsh; the pale 
coloration blends well with the vegetation of its habitat during most of the 
season; and the long, coarse-haired pelage can withstand a lot of abrasive action 
from sharp-edged sedges and coarse grasses. In Procyon I. maritimus the length 
of the pale subapical band is the greatest yet recorded, nearest that in some 
western races of Procyon lotor, namely, excelsus, pacificus, psora, and me.ricanus. 
The guard hairs are the longest of any known race except hirtus of the Upper 
Mississippi Valley, in which the hairs average slightly longer. 

Specimens examined. — Total number, 34, as follows: 

Delaware: Rehoboth Bay, 1. 

Maryland: Blackwater National Wildlife Refuge, 29 (6 skins only); Crocheron, 1 

(skin only); Vienna, 1 (skin only). 
Virginia: Saxis Island, 2 (skins only). 



RACCOONS OF NORTH AND MIDDLE AMERICA 87 

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88 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

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General account of P. 1. meTicanii.^ in New Mexico, pp. .348-349. 
1933. Cave life of Kentucky. Amer. Midland Nat. 14 (5): 433-434. 
1936. The mammals and life zones of Oregon. North Amer. Fauna 55, 416 
pp., illus. June. 

Accounts of P. I. paciftcns and P. I. eicelstis, pp. 315-317. 

Baird, Spencer Fullerton. 

1857. Mammals of North America. Pacific R. R. Rept. 8, 757 pp., illus. 

General account of the raccoons, pp. 209-215; original description of Proajon hernandezii 

var. mezicniui ( = P. L meikanus), |). 215. 

Bangs, Outram. 

1898a. The land mammals of peninsular Florida and the coast region of 
Georgia. Boston Soc. Nat. Hist. Proc. 28 (7) : 157-235. March. 
Original description of /'. /. elucii.i, p. 219. 

1898b. A new raccoon from Nassau Island, Bahamas. Biol Soc. Washington 
Proc. 12: 91-92. April 30. 
Original description of P: mtii/ixirdi, p. 92. 

1901. Chiriqui Mammalia. Mus. Comp. Zool. Bull. 39 (2): 17-51. 
Barnes, Claude Teancum. 

1922. Mammals of Utah. Univ. Utah Bull. 12 (15): 1-66. April. 
Distribution of the raccoon in Utah, pp. 133-134. 



RACCOONS OF NORTH AND MIDDLE AMERICA 89 

Bennet, James Milton. 

1925. Dog, gun, and raccoon. Forest and Stream 95: 554. 
liENNiTT, Rudolf, and Werner O. Nagel. 

1937. A survey of the resident game and fur bearers of Missouri. Univ. 
Missouri Studies 12 (2): 1-215. April 1. 
Benson, Adolph B. 

1937. The America of 1750, Peter Kahn's travels in North America. The 

English version of 1770. Revised from the original Swedish and 
edited by Adolph B. Benson, with a translation of new material from 
Kalm's diary notes. Vol. 1, 380 pp., illus.; vol. 2, pp. 381-797, 
illus., folding map 1. New York. 

Accounts of the raccoon in Pennsylvania and New Jersey. Vol. 1, pp. 52-53, 111, faced 
witli illustration, 242-243. 

Bergtold, W. H. 

1925. Unusual nesting of a raccoon. Jour. Mammal. 6 (4): 280-281. 

lilLLINGS, E. 

1856. Natural history of the raccoon (Procyon lotor). Canadian Nat. and 
Geol. 1 (4): 253-260. September. 

An early general account. 

Black, John David. 

1938. Mammals of Kansas. Kansas State Board Agr., Thirteenth Bien. 

Rept., pp. 116-217. 
Distribution and habits, pp. 154-155. 

Blair, W. Frank. 

1938. Ecological relationships of the mammals of the Bird Creek Region, 

northeastern Oklahoma. Amer. ^Midland Nat 20 (3): 496. No- 
vember. 

1939. Faunal relationships and geographic distribution of mammals in Okla- 

homa. Amer. Midland Nat 22 (1): 104. July. 

1940. A contribution to the ecology and faunal relationship of the mammals 

of the Davis Mountain region, southwest Texas. Univ. Michigan 
Mus. Zool. Misc. Pub. No. 46, p. 23. June 28. 
Boddaert, Peter. 

1784. Elenchus Animalium. Vol. 1, 174 pp. Roterdami. 

Description of Procyon lotor under the substitute name [A/e/w] lotor, p. 80. 

Bole, B. P., Jr., and Philip N. Moulthrop. 

1942. The Ohio recent mammal collection in the Cleveland Museum of Natural 
History. Cleveland Mus. Nat. Hist. Sci. Pub. 5 (6) : 83-181. Sep- 
tember 11. 
Bradt, Glenn W. 

1946. The raccoon, masked clown of the woodlots. Michigan Conserv. 15 
(8) : 6-7. September. 
Brass, Emil. 

1911. Aus dem Reiche der Pelze. 709 pp. April. 

Descriptions of Pr[oajoii] hndsoniciis [unidentifiable] and Procyon profeus (=P. I. 
pacificus), p. 564. 

Brown, C. Emerson. 

1936. Rearing wild animals in captivity, and gestation periods. Jour. 
Mammal. 17 (1): 10-13. February 14. 
Gestation period in a raccoon in Philadelphia Zoo 69 days from fii-st coition, p. 13. 

876119°— 50 7 



90 NORTH AMERICAN FAUNA 6 0, FISH AND WILDLIFE SERVICE 

Brown, L. G., and Lee E. Yeager. 

1943. Survey of the Illinois fur resource. Illinois Nat. Hist. Survey Bull. 22 

(6): 462. September. 
Browne, F. C. 

1892. The coon in eastern Massachusetts. Forest and Stream 39: 405. No- 
vember 10. 

BURMEISTER, HERMANN. 

1850. Verzeichniss der im Zoologischen Museum der Universitat Halle- 
Wittenberg aufgestellten Saugethiere, Vogel und Amphibian. 84 pp. 

Lists three species of raccoon. Proposes new combination [Ptocuoii hrachyurus] var. 
fusca, p. 13, based upon Procyon ohscuruK. without description or comment. 

BuKT, William H. 

1946. The mammals of Michigan. 288 pp., 13 color pis., 107 te.xt figs., 67 

text maps. 
Butterfield, Robert T. 

1944. Tagged raccoons— 1943-44. Ohio Con.serv. Bull. 8 (7): 11. July. 
Cagle, Fred R. 

1949. Notes on the raccoon, Procyon lotor megalodous Lowery. Jour. Mam- 
mal. 30 (1): 45-50. February 14. 
Cahalane, Victor H. 

1928. A preliminary wild life and forest survey of southwestern Cattaraugus 
County, New York. Roosevelt Wild Life Bull. 5 (1): 81-98. 
March. 

1947. Mammals of North America. 682 pp., illus. Macmillan Co. 

Raccoon, pp. 156-161. 

Cahn, Alvin R. 

1921. The mammals of Itasca County, Minnesota. Jour. Mammal. 2 (2): 
68-74. 
Campbell, J. E. 

1945. The raccoon in North Dakota. North Dakota Outdoors 7 (8): 4. 

February. 
Cartwright, George. 

1792. A journal of transactions and events, during a residence of nearly 
sixteen years on the coast of Labrador. 3 vols. 

A nortliern record: On the Alexander River, a brancli of the St. Lewis River, shot a 
raccoon, August 22, 1774, vol. 2, p. 23. 

Cart, Merritt. 

1911. A biological survey of Colorado. North Amer. Fauna 33, 256 pp., 
illus. August 17. 

Distribution and habits in Colorado, pp. 193-194. 

Catesby, Mark. 

1743. The natural history of Carolina, Florida and the Bahamas. 2 vols. 

Describes the raccoon, p. XXIX, "An Account of Carolina and the Bahama Islands"; in 
appendix to vol. 2. 

Chamberlain, Montague. 

1884. List of the mammals of New Brunswick. New Brunswick Nat. Hist. 
Soc. Bull. 3: 39. 
Chase, Harry. 

1911. The SuUivan Law. Forest and Stream 77: 551-552. October 7. 

Refers, p. 552, to the use of raccoon skins in payment for services of the Secretary to the 
Governor of the "State of Franklm" in eastern Termessee. 



RACCOONS OF NORTH AND MIDDLE AMERICA 91 

Clarke, Frank I. 

1910. Game of British Columbia. Bur. Provincial Inforination British 
Cohimbia Off. Bull. No. 17: 29. 
Cole, H. E. 

1922. Wildlife in Baraboo Hills. Wisconsin Conserv. .'-5 (6): 9. January. 
Cole, Lawrence Wooster. 

1907. Concerning the intelligence of raccoons. Jour. Coinp. Neurol, and 
Psychol. 17 (3): 211-261. May. 
Records the results of rather elaborate experiments. 

1912. Observations of the senses and instincts of the raccoon. Jour. Animal 
Behavior 2 (5) : 299-309. September-October. 
Cory, Charles B. 

1912. The mammals of Illinois and Wisconsin. Field Mus. Nat. Hist., Zool. 
Ser. 11, Pub. 153, p. 396. 
Criddle, Stuart. 

1929. An annotated list of the mammals of Aweme, Manitoba. Canadian 
Field-Nat. 43: 156. October. 
Cross, E. C, and J. R. Dymond. 

1929. The mammals of Ontario. Royal Ontario Mus. Zool. Handbook No. 1, 
pp. 12-13. 
CuRRiE, John W. 

1949. This raccoon racket. Kentucky Happy Hunting Ground 5 (2) : 5, 19. 
March. 

Brief account of raising raccoons at Game P'arm of the Kentucky Game and Fish 
Commission. 

CuviER, Georges. 

1798. Tableau elementaire de I'histoire naturelle des animaux. 710 pp., 
illus. Paris. 

Original description of Ursvs cancrirnms {=Proc!/on cancrirorus). p. ]1.3, from Cayenne. 

Dalquest, Walter W. 

1948. Mammals of Washington. Univ. Kansas Pubs. Mus. Nat. Hist., vol. 2, 
444 pp. 
Discussion of the raccoon, pp. 179-182. 

Dampier, William. 

1729. A new voyage round the World. Vol. 1, 550 pp. 

Refers to the occurrence of raccoons on the Tres Marias Islands, off the west coast of 
Mexico, p. 276. 

Banner, E. Y. 

1931. Raccoons like candy. Mt. Rainier Nature Notes 9 (8): 8. 
Davis, Herbert Burnham. 

1907. The raccoon: A study in animal intelligence. Amer. Jour. Psychol. 
18 (4) : 447-487. October. 
An important detailed account of the results of experiments. 

Davis, W^illiam B. 

1939. The recent mammals of Idaho. 400 pp., illus. Caxton Printers, 
Caldwell, Idaho. 
An account of P. I. excelsus within the State, pp. 128-129. 



92 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

De Beaux, Oscar. 

1910. tjber eine neue Farbenspielart des Waschbaren. (Ein Beitrag zur 

Systeinatik des Proci/on lotor.) Zool. Anzeiger 35: 621-626. 

April 26. 

A confusing; general discussion of raccoons. Three new names, "Pr. I. flaiHdus" (p. 026), 
"Pr. lotor rufescens" (p. 625). and "Pr. hernandezi castaneus" (p. 624) are proposed, but 
appear to be unidentifiable. 

Dellinger, S. C, and J. D. Black. 

1940. Notes on Arkansas mammals. Jotir. Mammal. 21 (2) : 188. 
Desmarest, a. G. 

1819. Dictionaire histoire iiattirelle. Vol. 29, p. 93. 
Dice, L. R. 

1925. A stirvey of the mammals of Charlevoix County, Michigan, and 
vicinity. Univ. Michigan Miis. Zool. Occas. Papers No. 159, p. 20. 
April 11. 
Dionne, Charles Eusebe. 

1902. Les mammiferes de la Province de Quebec. 285 pp. 

Distribution and habits of raccoon, pp. 164-168. 

DoziER, Herbert L. 

1948. A new eastern marsh-inhabiting race of raccoon. Jour. Mammal. 29 
(3): 286-290. August 31. 

Description of Procyon lotor maritimns, from Blackwater National Wildlife Refuge, 
Dorchester County, Maryland. 

Dozier, Herbert L., Thora M. Plitt Haruy, and Merle H. Markley. 

1948. Fur characteristics of two eastern raccoons. Jour. Mammal. 29 (4) : 
383-393. December 31. 
Dymond, John Richardson. 

1928. The mammals of Lake Nijiigon region. Trans. Royal Canadian Inst. 
16 (pt. 2) : 239-250. 

Northern records of the raccoon, p. 210. 

Elliot, D. G. 

1901. A list of mammals obtained by Thaddeus Surber in North and South 
Carolina, Georgia, and Florida. Field Columbian Mus. Pub. No. 
58, Zool. Ser. 3 (4): 51-53. June. 

1903. A list of mammals collected by Ednnind Heller, in the San Pedro 

Martir and Hanson Laguna Mountains and the accompanying 
coast regions of Lower California. Field Cohimbian Mus. Pub. 
No. 79, Zool. Ser. 3 (12): 231. June. 

1904. Catalogue of mammals collected by E. Heller in southern California. 

Field Columbian Mus. Pub. No. bl, Zool. Ser. 3 (16): 316. March. 
1907. A catalogue of the collection of mammals in the Field Columbian 
Museum. Field Columbian Mus. Pub., No. 115, Zool Ser. 8: 
426-427. 
Enders, Robert Iv. 

1930. Some factors influencing the distribtition of mammals in Ohio. Univ. 
Michigan Mus. Zool. Occas. Papers No. 212, p. 9. April 23. 
Evermann, Barton \V.\^rren, and Howard Walton Clark. 

1911. Notes on the mammals of the Lake Maxinkuckee region. Washington 

Acad. Sci. Proc. 13 (1): 32. 



RACCOONS OF NORTH AND MIDDLE AMEJUCA 93 

Fischer, Gotthelf. 

1814. Zoognosia Tabulis Synopticis. Vol. 3, 732 pp. 

Description of Proci/nn anyuihilus ( = Ptocijoii Jotor), p. 177. 

Flower, Stanley Smyth. 

1931. Contributions to our knowledoe of the duration of life in vertebrate 
animals. V. A'laminals. Zool. Soc. London. I'roc. 1<)31 (pt. 1): 
145-234. 

The longevity of the raccoon in cai)tivity, yi. 177. 

FoRBUSH, Edw.\rd Howe. 

1916. The natural enemies of birds. Massachusetts Board Agr. Bull. 

No. 3, pp. 24-25. 

Consideration of the raccoon in relation to birds. 

Frantzius, a. von. 

1881. Los mamiferos de Costa Rica. Colecion de documentos para la 
historia de Costa Rica publicados por el Don Leon Fernandez. 442 pp. 
Raccoon, p. 412. 

Frye, 0. Earl, and Daniel W. Lay. 

1943. Fur resources and fur animals of Texas. Texas Came, Fish, and 
Oyster Comm. Bull. No. 25: 18-19. 
Gander, Frank Forrest. 

1928. Period of gestation in some American mammals. .Jour. Mammal. 
9(1): 75. February 9. 

Records the period of gestation in a raccoon under observation. 

Garman, H. 

1894. A preliminary list of the vertebrate animals of Kentucky. Essex 
List. Bull. 26 (1-3): 4. 

The raccoon as an enemy of bird life. 

Gaumer, George Franklin. 

1917. Monografia de los mamiferos de Yucatan. Pub. by Secretaria de 

Fomento, D. F. Mexico. 331 pp. 

Notes on tlie raccoon of Yucatan, pp. 210-215. 

Geoffroy-Saint Hilaire, Isidore. 

1855. Voyage sur la Venus. Zoologie. 355 pp., illus. 

Description of "Procynn lofor variete mexicaine," from Mazatlan, Sinaioa, pp. 125-132, 
pi. VI. 

Geoffroy-Saint Hilaire, Isidore, and G. Cuvier. 

1795. Memoire sur une nouvelle division des mammiferes, et sur les principles 
c}ui doivent servir de base dans cette sorte de travail. Mag. Encyc. 
2: 164-190. 
GiDLEY, James Williams. 

,1906. A fossil raccoon from a California Pleistocene cave deposit. U. S. 
Natl. Mus. Proc. 19: 553-554, illus. 

Original description of Procyon simus, p. 553. Similar in size to P. I. excelsus, but molari- 
form tooth rows more strongly curved outward near middle; crown of first upper premolar 
with outer side sloping inward posteriorly, instead of nearly parallel to antero-posterior 
axis of skull; posterior lower molar with a distinctly broader cusp, or "heel." 

Gill, Theodore N. 

1872. Letter on the alleged hybrid between a raccoon and cat. Amer. 
Nat. 6: 53-55. January. 



94 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

Gilpin, J. Bernard. 

1870. On the Mammalia of Nova Scotia. Trans. Nova Scotia Inst. Nat. 
Sci. 2 (pt. 1): 15. 
Goldman, Edward Alphoxso. 

1913. Descriptions of new mammals from Panama and Mexico. Smithsn. 
Misc. Collect. 60 (22): 1-20. February 28. 

Original description of E. c. paviarnensis { = Proc!/on (Eiiprocyon) cancrivorus panaviensis), 
p. 15. 

1920. Mammals of Panama. Smithsn. Misc. Collect. 69 (5): 1-309, illus. 

Accounts of P. I. pitmilus and P. c. paiuimensis. pp. 151-153. 
GOODPASTER, W. 

1941. A list of the birds and mammals of southwestern Ohio. Jour. Cin- 

cimiati Soc. Nat. Hist. 22 (3): 43. June. 
Goodwin, George Gilbert. 

1924. Mammals of the Gaspe Peninsula, Quebec. Jour. Mammal. 5 (4): 
253. 

1934. Mammals collected by A. W. Anthony in Guatemala, 1924-1928. 

Amer. Mus. Nat. Hist. Bull. 68, art. 1, 60 pp., illus. December 12. 
Records P. I. shnfeldli from El Espino and Finea Cipres, Guatemala. 

1935. The mammals of Connecticut. Connecticut State Geol. and Nat. 

Hist. Survey Bull. 53, 221 pp., illus. 
Distribution records in Connecticut, p. 59. 

1942. Mammals of Honduras. Amer. Mus. Nat. Hist. Bull. 79: 176. May 

29. 
1947. The carnivores. Part II. Raccoons and their allies. Audubon Nat. 
Bull., Ser. No. 18, Bull. No. 3, 4 pp. November. 
Gray, John Edward. 

1837. Charlesworth's Mag. Nat. Hist. 1: 580. 

Description of Procyon nivea which is unidentifiable. 

1842. Descriptions of some new genera and fifty unrecorded species of Mam- 
malia. Ann. Mag. Nat. Hist. 10: 255-267. December. 

Description of Procyon psora, p. 261. 

1864. A revision of the genera and species of ursine animals (Ursidae), 
founded on the collection in the British Museum. Zool. Soc. London 
Proc. 1864: 677-709. 

Includes accounts of the raccoons, pp. 703-706. Euprocyon proposed as a subgenus, p. 
705. 

Green, H. U. 

1932. Mammals of the Riding Mountain National Park, Manitoba. Can- 

adian Field-Nat. 46 (7): 149-152. October. 
Records extinction of raccoon in Riding Mountain section during past two decades, p. 

152. 

Gregory, Tappan. 

1936. Mammals of the Chicago region. (Program of Activities.) Chicago 

Acad. Sci. 7 (2-3) : 34. July. 
Gregory, William King. 

1933. Nature's wild dog show. New York Zool, Soc. Bull. 36 (4): 83-96, 

illus. July-August. 
Brief reference to origin of raccoon family during lower Oligocene period, p. 83. 



RACCOONS OF NORTH AND MIDDLE AMERICA 95 

Gregory, William King — Continued 

1936. On the phylogenetic relationships of the giant panda (Aihiropoda) to 

other arctoid Carnivora. Ainer. Mus. Novitates No. 878, pp. 1-20. 
August 8. 
Grinnell, Joseph. 

1914. An account of the nianimals and birds of the lower Colorado V'alley. 
Univ. California Pub. Zool. 12 (4): 51-294, illus. March 20. 
Distribution and habits of P. I. patlidus, pp. 260-262. 

1933. Review of the recent mammal fauna of California. Univ. California 
Pub. Zool. 40 (2) : 99. September 26. 
Grinnell, Joseph, Joseph Scattergood Dixon, and Jean Myrox Linsdale. 
1930. Vertebrate natural history of a section of northern California through 
the Lassen Peak Region. Univ. California Pub. Zool., vol. 35, 594 
pp., 181 text figs. October. 

Accoimt of P. 1. psora, pp. 460-462. 

1937. Fur-bearing mammals of California. 2 vols. Vol. 1, pp. 137-165. 

Univ. California Press, Berkeley. 
Grinnell, Joseph, and Jean Myron Linsdale. 

1936. Vertebrate animals of Point Lobos Reserve, 1934-35. Carnegie Inst. 
Washington, Pub. No. 481, p. 134. December 10. 
Hall, Archibald. 

1861. On the mammals and birds of the district of Montreal. Canadian 
Nat. & Geol. 6 (4) : 294. 
Description of specimen of raccoon. 

Hall, E. R. 

1946. Mammals of Nevada. 710 pp. Univ. California Press, Berkeley. 

Notes on raccoons, pp. 17.5-179. 

Hamilton, William John, Jr. 

1941. Notes on some mammals of Lee County, Florida. Amer. Midland 
Nat. 25 (3) : 688. May. 

1943. The mammals of eastern United States. 432 pp., illus. Ithaca, N. Y. 

Raccoons, pp. 118-12.5. 

Hardy, Manly. 

1907. The raccoon. Forest and Stream 69: 852. November 30. 

On raccoon characteristics. 

Harlan, Richard. 

1825. Fauna Americana: being a description of the mammiferous animals in- 
habiting North America. 318 pp. 
General account of the raccoon, pp. ,5.3-56. 

Harper, Francis. 

1920. Okefinokee Swamp as a reservation. Jour. Amer. Mus. Nat. Hist. 20: 

30. January-February. 
1927. The mammals of the Okefinokee Swamp region of Georgia. Boston 

Soc. Nat. Hist. Proc. 38: 297. 
1929. Notes on mammals of the Adirondacks. New York Mus. Handbook, 
No. 8, p. 69. May. 
Harrington, Lyn. 

1944. The masked bandit of the woods. Canadian Nature 6 (5) : 154, illus. 

November-December. 



96 NORTH AMERICAN FAUNA 6 0, FISH AND WILDLIFE SERVICE 

Hatt, Robert T. 

1924. Tlie land vertebrate communities of western Leelanau Comity, Mich- 
igan, with an annotated list of the mammals of the county. Papers 
Michigan Acad. Sci., Arts and Letters 3: 392. 
Hernandez, Francisci. 

165L Rerum medicarum Novae Hispaniae thesaurus sen plantarum ani- 
malium, mineralium Mexicanorum Histaria [Francisci Hernandez, 
et al.]. 

Accounts of the "Mapach," tract 1, cap. 1. p. 1 and of the "Tepe Maxtlaton," tract 1, 
cap. 28, p. 9 of an appendix of the general work; both apparently referable to the raccoon. 

Herrera, Alfred L. 

1899. Sinominia vulgar y cientifica de los principales vertebrados Mexi- 
canos. 31 pp. Mexico. 
Herrick, Clarence Luther. 

1892. Mammals of Minnesota. Minnesota Geol. Nat. Hist. Survey Bull. 7, 
299 pp., illus. 

Account of Procyon lotor ( = P. I. hirtiis), \)\). i:{9-145. 

Hibbard, Claude W. 

1933. A revised check list of Kansas mammals. Trans. Kansas Acad. Sci. 

36: 235. 
1944. A checklist of Kansas mammals, 1943. Trans. Kansas Acad. Sci. 
47: 61-88. 
Notes on raccoons, p. 67. 

HoLLisTER, Ned. 

1914. Descriptions of four new mammals from tropical America. Biol. 

Soc. Washington Proc. 27: 141-144. July 10. 

Original description of P. I. cra-fsuletis, p. 142. 

1915. The genera and subgenera of raccoons and their allies. U. S. Natl. 

Mus. Proc. 49 (2100): 143-150. August 13. 
An important contribution to knowledge of the classification of the group. 

Howell, Arthur H. 

1921. A biological survey of Alabama. North Amer. Fauna 45, pp. 34-35. 
Hughes, Criffith. 

1750. The natural history of Barbados. 314 p])., ilhis. London. - 
Refers to a bounty on raccoons, p. (ifi. 

Huntington, George S. 

1892. On the ileo-colic junction in Prori/on lotor and allied forms. Trans. 
New York Acad. Sci. 11: 50-53. 
Illiger. Johann Karl Wilhelm. 

1815. Ueberblick der saugthiere nach ihrer vertheilung uber die welttheile. 
Abhand. Konig Akad. Wissensch. 1804-1811: 39-159. Berlin. 
IvEY, DeA^'itt R. 

1948. The raccoon in the salt marshes of northeastern Florida. Jour. 
Mammal. 29 (3): 290-291. August 31. 
Jackson, H. H. T. 

1908. A preliminary list of Wisconsin mammals. Wisconsin Nat. Hist. 
Soc. Bull. 6: 13-34. 
Jaeger, E. C. 

1947. Use of the os phallus of the raccoon as ripping tool. Jour. Mammal. 
28 (3) : 297. August. 



RACCOONS OF NORTH AND MIDDLE AiMERICA 97 

Jameson, E. W., Jr. 

1943. Notes on the habits and siphonapterous parasites of the mammals 
of Welland County, Ontario. Jour. Mammal. 24 (2): 194-197. 
May 20. 
Johnson, Charles E. 

1916. A brief descriptive list of Minnesota mammals. Fins, Feathers and 
Fur, No. 8, p. 7. December. 
Johnson, M. S. 

1930. Common injurious mammals of Minnesota. Univ. Minnesota Agr. 
Expt. Sta. Bull. 259, p. 57. January. 
Jones, CIlenn. 

1946. The raccoon in Oklahoma. Oklahoma Game and Fish News 2 (12): 
4-5, 7. December. 
JossELYN, John. 

1672. New England's rarities discovered in birds, beasts, fishes, serpents, 
and plants of that country. (Reprint, 1865, 169 pp.) 
Refers to tW habits of the raccoon, p. .52, 1865 edition. 

Keeler, Charles A. 

1891. Notes on the colors of west coast mammals. Zoe 2: 212. 
Kellogg, Remington. 

1937. Annotated list of West Virginia mammals. U. S. Natl. Mus. Proc. 
84 (3022) : 443-479. October 7. 

Distribution and liabits of raccoon, ji. 4.51. 

1939. Annotated list of Tennessee mammals. U. S. Natl. Mus. Proc. 86 
(3051): 245-303. 
Distribution of raccoon within the State. 

Kerr, Robert. 

1792. The animal kingdom or zoological system of the celebrated Sir Charles 
Linnaeus, Class 1, Mammalia. 400 pp. 

An account of the raccoons under the names Ursus lotor and " L^ lotor meliiin.s," the latter 
a color phase, pp. 188-189. 

KOPMAN, H. H. 

1921. Wild life resources of Louisiana. Louisiana State Dept. Con.serv. 
Bull. No. 10, pp. 23-39. December. 
Contains an account of the raccoon in Louisiana, p. 28. 

Lanman, Charles. 

1856. Adventures in the wilds of the United States and British American 
Provinces. 2 vols. Vol. 1, p. 83. Philadelphia. 
Lawson, John. 

1718. The history of Carolina. 258 pp. 
An account of the raccoon, p. 121. 

Le Conte, John L. 

1848. Notice of five new species of fossil Mammalia from Illinois. Amer. 
Journ. Sci. and Arts (Silliman's Journal), ser. 2, vol. 5, art. 16, pp. 
102-107. May. 

New name, Procyon priscus, proposed, p. 106. 

Leraas, H. J. 

1942. Notes on mammals from west-central Minnesota. Joiu". Mammal. 
23 (3) : 344. August 14. 



98 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

Linnaeus (Linne'), Carl. 

1740. Systema Naturae, inquo naturae regna tria, secundum classes, ordines, 
genera, species, systematice proponuntur. Ed. 2, 80 pp., Stock- 
holniiae. 

Listed the raccoon with the bear as Ursus cauda elongata, p. 35. 

1747. Beskrifning Pa et Americanskt Diur, som Hans Konglige Hoeghet 
gifvit til underjoekning. Kongliga Svenska Vetenskaps Academiens 
Handlingar, vol. 8, pp. 277-289, tab. 9, figs. 1-2. 

A general description of the raccoon under the name Ursus cauda elongnta, including 
habits, and perhaps the first illustration of the animal and its os penis. 

1758. Systema Naturae, secundum classes, ordines, genera species, cum 
characteristibus, differentiis, synonymis, locis. Ed. 10, tonms 1, 
regnum animale, 824 pp. 
Original description of [ Ursus] lotor i^Procyon lotor lotor), p. 48. 

LowERY, Georoe H., Jr. 

1936. A preliminary report on the distribution of the mammals of Louisiana. 
Louisiana Acad. Sci. Proc. 3 (1): 1-39. March. 
Distribution and habits of raccoons, p. 19. 

1943. Check-list of the mammals of Louisiana and adjacent waters. Lou- 
isiana State Univ. Mus. Zool. Occas. Papers No. 13, pp. 213-257, 
illus. November 22. 
Raccoons, pp. 225-231. Description of Procyon lotor megalodous, p. 225. 

LlTTTRINGER, L. L., Jr. 

1931. An introduction to the mammals of Pennsylvania. Pennsylvania Bd. 
Game Comm'rs. Bull. 15: 29. 
Lyon, Marcus Ward, Jr. 

1936. Mammals of Indiana. Amer. Midland Nat. 17 (1) : 1-384. January. 
Maar, R. D. 

1947. Raccoons snap their own pictures. Field and Stream 52 (6) : 112-113, 
illus. October. 
MacFarlane, Roderick Ross. 

1905. Notes on mammals collected and observed in the northern Mackenzie 
River District Northwest Territories of Canada, with remarks on 
explorers and explorations of the far North. LT. S. Natl. Mus. Proc. 
28: 673-764, illus. June. 
Raccoon numbers and distribution, pp. 715-716. 

MacLulich, D. a. 

1936. Mammals of the Wanapitei Provincial Forest, Sudbury District, 
Ontario. Canadian Field-Nat. 50 (4): 57. April. 
Manville, R. H. 

1942. Notes on the mammals of Mount Desert Island, Maine. Jour. Mam- 
mal. 23 (4) : 393. November 14. 
Matthew, William Diller. 

1930. The phylogeny of dogs. Jour. Mammal. 11 (2): 117-138. 

Phylogenetic relationships of raccoons, p. 129. May 9. 

McGrew, p. O. 

1938. Dental morphology of the Procyonidae with a description of Cynarc- 
toides, gen. nov. Field Mus. Nat. Hist. Geol. Ser. 6 (22): 323-338, 
illus. October 31. 



RACCOONS OF NORTH AND MIDDLE AMERICA 99 

McKean, William T. 

1948. Winter foods of North Dakota predatory animals. North Dakota 
Outdoors 10 (8) : 6. February. 
McLean, Donald D. 

1934. Predatory animal studies. California Fish and Game 20 (1): 32. 
January. 
McMahan, C. a. 

1946. Trapping the raccoon. Texas Game and Fish 4 (12): 30. November. 
McQueen, A. S., and Hamp Mizell. 

1926. History of Okefenokee Swamp. 191 pp. Jacobs & Co., Clinton, S. C. 
Mearns, Edgar Alexander. 

1914. Descriptions of three new raccoons from the Me.xican boundary region. 
Biol. Soc. Washington Proc. 27: 63-67. March 20. 

Original descriptions of P. /. fuscipes. p. 63, P. I. ochracens (=P. 1. palUdus), p. 64, and 
P. I. californicus {^P. I. psora), p. 66. 

Merriam, Clinton Hart. 

1884. The mammals of the Adirondack Region. 316 pp. 

Includes an account of the raccoon, pp. 91-95. 

1898. Mammals of Tres Marias Islands off western Mexico. Biol. Soc. 

Washington Proc. 12: 13-19. January 27. 

Original description of Prncymi lolor itisnlaris (=:P. insiiliTis insnlaris), p. 17. 

1899. Results of a biological survey of Mount Shasta, California. North 

Amer. Fauna 16, 179 pp., illus. October 28. 
Original description of Proci/oii psora pacifica (=P. hjfor pacificiis), p. 107. 

1900. Descriptions of two new mammals from California. Biol. Soc. ^^'ashing- 

ton Proc. 13: 151-152. June 13. 

Oiiginal description of Procyon pallidus (=P. lofor pallidiis), p. 151. 

1901. Six new mammals from Cozumel Island. Biol. Soc. Washington Proc. 

14: 99-104. July 19. 

Original description of Procyon pygniaevs, p. 101. 

Miller, Gerrit S., Jr. 

1900. Key to the land mammals of northeastern North America. New York 

State Mus. Bull. 8 (38) : 137. 
1905. Mammals of the Bahama Islands. (The Bahama Islands.) Geog. Soc. 
Baltimore, pp. 371-384, illus. 

Extended account of P. maynardi, pp. .376-:379. 

1911. Descriptions of two new raccoons. Biol. Soc. ^^'ashington Proc. 
24: 3-6. January 28. 

Original descriptions of Procyon pnniihis {=Procyon lotor piiniiliis), p. 3, and Procyon 
minor, p. 4. 

1924. List of North American recent mammals, 1923. U. S. Natl. Mus, 
Bull. 128, 673 pp. 
Species of Procyon, pp. 107-110. 

Mills, Wesley. 

1892. Hibernation and allied states in animals. Trans. Royal Soc. Canada 
10 (sec. 4): 49-66 (1893). 
Describes the hibernation of a tame raccoon, p. 50. 



100 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

MoHR, Carl O. 

1947. Table of equivalent populations of North American small mammals. 

Amer. Midland Nat. 37 (1): 224. January. 
Morris, R. F. 

1948. The land mammals of New Brunswick. Jour. Mammal. 29 (2): 168. 

May. 
MosBY, Henry 8. 

1947. Virginia animals everyone should know: The raccoon. Virginia 
Wildlife 8 (2) : 8-9, 20. February. 
MuRiE, Adolpu. 

1935. Mammals from Guatemala and Honduras. Univ. Michigan Mus. 
Zool. Misc. Pub. 26, 30 pp., illus. July 15. 
Records P. I. shufeldli from El Cayo, British Honduras. 

Nagel, W. O. 

1943. How big is a 'coon'? Missoiu'i Conserv. 4 (7) : 6-7. October. 
Necker, Walter L. and Donald F. Hatfield. 

1941. Mammals of Illinois. An annotated check list with keys and bibli- 
ography. Chicago Acad. Sci. Bull. 6 (3): 46. May 15. 
Nelson, Arnold B., and Harvey I. Scudder. 

1947. The raccoon as a predator on turtles. Jour. Mammal. 28 (4): 406. 
Nelson, Edward William. 

1899. Natural history of the Tres Marias Islands, Me.xico. North Amer. 
Fauna 14, 97 pp., illus. April 29. 
Distribution and habits of the raccoons, p. 17. 

1930a. Four new raccoons from the keys of southern Florida. Smithsn. 
Misc. Collect. 82 (8): 1-12, illus. July 10. 

Original descriptions of P. I. marhmx, p. 7; P. I. iiiesperntus, p. 8; P. I. rnispicdfiis. p. 9; 
and P. I. incautus, p. 10. 

1930b. Wild animals of North America: intimate studies of big and little 
creatures of the mammal kingdom. 1930 ed., 254 pp., illus. 
General account of the raccoon, pp. 59-60. 

Nelson, Edward William, and Edward Alphonso Goldman. 

1930a. A new raccoon from Lower California. Jour. Washington Acad. Sci. 
20 (5) : 82-83. March 4. 
Original description of P. I. grinnelli, ji. S2. 

1930b. Six new raccoons of the Procyon lotor group. Jour. Mammal. 11 (4): 
453-459. November 11. 

Original descriptions of Procyon gloreralleni. p. 45:^: P. I. hirtu.f, p. 455; P. I. rariiis, p. 456; 
P. I. litoreiis, p. 457; P. I. eiccl.sns. p. 458; and P. I. iiincoiirereiisis, p. 458. 

1931a. Three new raccoons from Mexico and Salvador. Biol. Soc. ^^'ashing- 
ton Proc. 44: 17-22. February 21. 
Original descriptions of P. I. shufeldti, p. 17, P. 1. dkkeyi, p. 18, and P. i. ricinus. p. 20. 

1931b. A new raccoon from South Carolina. Jour. Mammal. 12 (3): 308- 
309. August 24. 
Original description of J\ I. soliitiis, p. 308. 

1932. The type locality of Ursus lotor Linne. Jour. Mammal. 13 (4) : 367. 
November. 



RACCOONS OF NORTH AND MIDDLE AMERICA 101 

Newberry, J. S. 

1855. Report upon the mammals, in routes in California and Oregon explored 
by Lieutenants R. S. Williamson and Henry L. Abbott, Corps 
Topographic Engineers in 1855, Rept. Expl. and Surv. R,. R. route 
Mississippi River to Pacific Ocean, 1854-1855. Vol. 6, pt. 4, 
No. 2, chap. 1, pp. 35-72. 
Raccoons sold for food in San Francisco market, p. 47. 

Norton, Arthur H. 

1930. The mammals of Portland, Maine, and vicinity. Portland Soe. 
Xat. Hist. Proc. 4(1): 24-25. 
Okex, Lorexz. 

1816. Lehrbuch der Naturgeschichte. Dritter Theil, Zoologie, Zweite 
Abth. Fleischthiere, pp. 386-387. 
Orr, Robert T. 

1943. Mammals of the Clearwater Mountains, Idaho. California Acad. 

Sci. Proc, 4th ser. 23 (35) : 521. August 18. 
Osgood, F. L., Jr. 

1938. The mammals of Vermont. Jour. Mammal. 19 (4): 437. November 
14. 
Packard, A. S., Jr. 

1866. List of vertebrates observed at Okak, Labrador, by the Reverend 

Samuel Weiz. Boston Soc. Nat. Hist. Proc. 10: 269. 
1891. The Labrador coast. 513 pp. New York. 
Pearson, Leonard, and B. H. Warren. 

1897. Diseases and enemies of poultry. Special Report. 749 pp. (Pub. 
by authority Pennsylvania State Legislature.) 

Chickens killed by raccoons and other habits injurious to human interests in Pennsyl- 
vania described, pp. 481-484. 

Pike, Zebulon M. 

1895. Expeditions of Z. M. Pike 1805-1807. 2 vols. Edited by Elliott 
Coues. 
PococK, Reginald Innes. 

1921. The external characters and classification of the Procyonidae. Zool. 
Soc. London Proc. 1921: 389-422. 
A discussion of the genera and subfamilies. 

PooLE, Arthur J., and Viola S. Schaxtz. 

1942. Catalog of the type specimens of mammals in the United States 

National Museum, including the Biological Surveys collection. 

U. S. Natl. Mus. Bull. 178, pp. 1-705. 
Pope, Clifford H. 

1944. Attainment of sexual maturity in raccoons. Jour. Mammal. 25 (4): 

91. November. 
Prince, E. E. 

1906. Zoological report. Trans. Ottawa Field Nat. Club. 22: 58. June. 
PuRCHAS, Samuel. 

1614. Purchas his pilgrimage. Or relations of the world and the religions 
observed in all ages and places discovered from the creation unto the 
present. 918 pp., 8th "booke", chap. 5. 
An early reference to the raccoon in Virginia, p. 761. 



102 NORTH AMERICAN FAUNA 6 0, FISH AND WILDLIFE SERVICE 

Rand, A. L. 

1933. Notes on the mammals of the interior of western Nova Scotia. Cana- 
dian Field-Nat. 47 (3) : 45. March. 

1943. History of the raccoon (Procyon lotor L.) in Nova Scotia. Canadian 

Field-Nat. 57 (4-5) : 95. April-May. 

1944. The recent status of Nova Scotia fur bearers. Canadian Field-Nat. 

58 (3) : 90. May-June. 
1948. Mammals of the eastern Rockies and western plains of Canada. Nat. 

Mus. Canada Bull. No. 108, 237 pp., illus. 
Rathbun, Mary Jane. 

1918. The grapsoid crabs of America. U. S. Natl. Mus. Bull. 97. 445 pp., 

illus. (1917). 

Reference to fiddler crab as food of raccoon, p. 401. 

Rausch, Robert. 

1947. Suggestions for the handling of certain mammals. Jour. Wildlife 
Mangt. 11 (2): 189. April. 
Ray, John. 

1693. Synopsis methodica animalium. Quadrupedum. 336 pp. London. 
Includes an early reference to the raccoon, p. 179 

Rhoads, S. N. 

1885. Tenacity and ferocity in the raccoon. Amer. Nat. 19 (8): 823-824. 
1894. Contributions to the mammalogy of Florida. Acad. Nat. Sci. Phila- 
delphia Proc, p. 155. 
1903. The mammals of Pennsylvania and New Jersey. 266 pp., illus. 
Account of Procyon lotor, pp. 182-183. 

Richardson, John. 

1829. Fauna Boreali-Americana. Part 1, p. 36. London. 
Ridgway, Robert. 

1912. Color standards and color nomenclature. 44 pp., 53 color pis. 

Washington, D. C. 
Rinker, George C. 

1944. Os clitoridis from the raccoon. Jour. Mammal. 25 (4): 91-92. 
Rowley, John. 

1902. The mammals of Westchester County, New York. Abst. Proc. Linn. 
Soc. New York, Nos. 13-14, p. 53. 
Rust, H. J. 

1946. Mammals of northern Idaho. Jour. Mammal. 27 (4): 313. Novem- 
ber. 
Sansom, Joseph. 

1820. Travels in lower Canada. Voyages and Travels 3 (1) : 49. 
Saunders, William Edwin. 

1932. Notes on the mammals of Ontario. Trans. Royal Canadian Inst. 18 
(pt. 2, JMo. 40) : 271-309. July. 

Notes on distribution and habits of raccoon. A few reported on north shore of Lake 
Superior: rare north of latitude 47°, p. 281. 

SCHOONMAKER, W. J. 

1929. Notes on some mammals of Allegany State Park. Jour. Mammal. 10 
(3): 247. August. 
Scott, William Berrymax. 

1913. A history of land mammals of the Western Hemisphere. 693 pp., illus. 

Origin and relationships of Procyonidae, pp. 546-548. 



RACCOONS OF NORTH AND MIDDLE AMERICA 103 

Seton, Ernest Thompson. 

1929. Lives of game animals. Bears, coons, badgers, skunks, and weasels. 
Vol. 2 (pt. 1), 367 pp., illus. 
Includes an illustrated accoiuit of the raccoon, pp. 230-256. 

Sherman, H. B. 

1936. A list of the recent land mammals of Florida. Florida Acad. Sci. 
Proc. 1: 110. 
SHIR.A.S, George, 3d. 

1920. A raccoon explores new country. Forest and Stream 90 (1) : 10, 11, 44. 
.January. 
Shufeldt, Robert Wilson. 

1915. On the taxonomy of the Procyonidae. Science (n. s.) 41 (1062): 691- 
692. May 7. 

A superfamily Procyonidea is divided into two families— the Procyonidae and the Poto- 
sidae, with the former divided into tliree subfamilies. 

Simpson, George Gaylord. 

1929. Pleistocene mammalian fauna of the Seminole Field, Pinellas County, 

Florida. Amer. Mus. Nat. Hist. Bull. 56 (art. 8): 561-599. Feb- 
ruary 12. 

Original description of Procyon nanus, p. 575, which differed conspicuously from living 
species of the rep;ion in greater length in proportion to width of first upper molar. 

Sloane, Hans. 

1725. A voyage to the Islands Madera, Barbados, Nieves, St. Christophers, 
and Jamaica with the natural history of the herbs and trees, four- 
footed beasts, insects, reptiles, etc., of the last of these Islands. Vol. 
2, 499 pp., illus. 
Records the raccoons (apparently erroneously) in Jamaica, p. 329. 

Smith, Charles Hamilton. 

1848. Introduction to Mammalia. Jardine's Naturalist's Lilirary 15: 17- 
313, illus. 

Description of Proci/oii qiilarh ( = P.l. Mot), p. 222, based on a live specimen "in the State 
of New York," and of /'. cancrirnriis, p. 223. 

Smith, Captain [John]. 

1612. A map of Virginia. With a description of the country, the commodi- 
ties, people, government, and religion. 109 pp. 

Contains the earliest reference to the raccoon found in literature. Tlie animal is recorded 
under the name aro7ighc)in, p. 13. 

Smith, Ronald W.\rd. 

1940. Tlie land mammals of Nova Scotia. Amer. Midland Nat. 24 (1): 

225. July. 
Snyder, L. L. 

1930. A faunal investigation of King Township, York County, Ontario. 

Trans. Royal Canadian Inst. 17 (pt. 2): 175. 

1941. The mammals of Prince Edward County, Ontario. Univ. Toronto 

Studies, Biol. Ser. No. 48, p. 21. 
Snyder, L. L., and E. B. S. Logier. 

1931. A faunal investigation of Long Point and .icinity, Norfolk County, 

Ontario. Trans. Royal Canadian Inst. 18 (pt. 1) 39: 131. 



104 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

SoPER, J. Dewey. 

1923. The mammals of Wellington and Waterloo counties, Ontario. Jour. 

Mammal. 4 (4) : 251. November. 
1942. Mammals of Wood Buffalo Park, northern Alberta and District of 
Mackenzie. Jour. Mammal. 23 (2) : 126 May 14. 
Squires, W. Austin. 

1946. Changes in mammal population in New Brunswick. Acadian Nat. 
2 (7): 31. May. 
Stephens, Frank. 

1906. California mammals. 351 pp., illus. 

Accounts of P. I. psora, P. I. pacifica, and P. I. pallidus, pp. 225-228. 

Stock, Chester. 

1929. A census of the Pleistocene mammals of Rancho La Brea, based on 
the collections of the Los Angeles Museum. Jour. Mammal. 10 (4) : 
281-289, illus. November. 

Points out, p. 288, the remarkable absence of Procyonidae. 

Stone, Witmer, and William Everett Cram. 

1902. American animals, a popular guide to the mammals of North America 
north of Mexico, with intimate biographies of the more familiar 
species. 318 pp., illus. 
A good account of the common raccoon, pp. 247-254. 

Storr, Gottlieb Conrad Christian. ' 

1780. Prodromus methodi mammalium. 43 pp., Tubingen. 

Procyon employed as a generic name for the first time, p. 35. 

Struwing, Nels J. 

1948. North Dakota fur harvest report, 1947-48. North Dakota Outdoors 
11 (2): 11. August. 
Stuewer, Frederick VV. 

1943a. Reproduction of raccoons in Michigan. Jour. Wildlife Mangt. 7 (1): 

60-73. January. 
1943b. Raccoons: Their habits and management in Michigan. Ecol. 

Monog. 13: 203. April. 
1948. Artificial dens for raccoons. Jour. Wildlife Mangt. 12 (3): 296-301. 
July. 
Sturgis, Robert S. 

1939. The Wichita Mountains Wildlife Refuge. Chicago Nat. 2 (1): 12. 

March. 
Surber, Thaddeus. 

1932. The mammals of Minnesota. Minnesota Dept. Conserv. Bull., Div. 

of Game and Fish, p. 48. 
Svihla, Arthur, and Ruth Dowell Svihla. 

1940. Annotated list of the mammals of Whitman County, Washington. 

Murrelet 21 (3) : 54. September-December. 
Swanson, G. a., T. Surber, and Thomas S. Roberts. 

1945. The mammals of Minnesota. Minnesota Dept. Conserv. Tech. Bull. 
No. 2, p. 63. 
Swarth, Harry Schelwald. 

1912. Report on a collection of birds and mammals from Vancouver Island. 
Univ. California Pubs. Zool. 10 (1): 1-124, illus. 

Distribution and habits of raccoon, pp. 108-109. 



RACCOONS OF NORTH AND MIDDLE AMERICA 105 

SWENSON, SiDEL B. 

1931. The mammals of Minnesota. Fins, Feathers and Fur, No. 94, p. 7. 
February. 
Taylor, Walter Pe.vn, and W. T. Shaw. 

1927. Mammals and birds of Mount Rainier National Park. 249 pp., illus. 
OccuiTenco and Iiabits of /'. /. pucijica, pp. 14-4.'). 

Tev'is, Lloyd, Jr. 

1947. Summer activities of California raccoons. Jour. Mammal. 28 (4): 
323-332. December 1. 

TiEUEMANN, D. FrIEDRICH. 

1808. Zoologie. Zu seinen Vorlesungen entworfen, erster Band, Mensch und 
Saugthiere. 610 pp. 

Description of PTOcyon lotor under tlie substitute name L[otoT\ nihitiris, p. 380. 

Tucker, William J. 

1930. Yearbook on Texas conservation of wildlife 1929-30. Texas Game, 

Fish, and (Jyster Comn., p. 37. 
Tyrrell, J. B. 

1889. Catalogue of the Mammalia of Canada exclusive of the Cetacea. 
Canadian Inst. Proc, 3rd ser. 6: 76. 
Wac;ler, Johann Georg. 

1831. Einige Mittheilungen liber Thiere Mexicos. Isis 24: 510-533. 

Description of P. hernandezii, p. 514. 

Wagner, Johann Andreas. 

1841. In Schreber's Die Saugthiere. Supplement, Abth. 2, 558 pp. 

Treatment of the genus Procyon, pp. 153-162, and refers to illustrations: Proctjon lotor, pi. 
143, P. hernandezii, pi. 143 A, P. brachyurus, pi. 143 C, and P. ohscurus, pi. 143 D, published 
in 1778. The latter two species are unidentifiable. 

Warren, Edward Royal. 

1910. The mammals of Colorado. 300 ])]). G. P. Putnam's Sons, New York. 

Account of Procyon lotor {=P. I. fiirliix] in the State, pp. 219-220. 

1942. Mammals of Colorado. Their habits and distribution. 330 pp. 
Univ. of Oklahoma Press. 
Whitlow, Wayne B., and E. R. Hall. 

1933. Mammals of the Pocatello region of southeastern Idaho. Univ. 
California Pub. Zool. 40 (3) : 246. 
\\ iiiTNEY, Leon F. 

1931. The raccoon and its hunting. Jour. Mammal. 12 (1): 29-38. Feb- 

ruary 12. 
a good general account of the raccoon in New England. 

1933. The Raccoon — some mental attriljutes. Jour. Mammal. 14 (2): 108- 
114. May 15. 
WiEGMANN, Ar. Fr. Aug. 

1837. Ueber die Gattung Procyon. Archiv fiir Naturgesch., drifter Jahrgang, 
erster Band, pp. 353-372. 

An early review of the genus in whicli five species are recognized, two of these, Procyon 
brachyurus (p. 369) and Procyon oliscurus (p. 370) described as new. Neither of these seems 
to be clearly identifiable. 



87(ill'J°— 50- 



106 NORTH AMERICAN FAUNA 60, FISH AND WILDLIFE SERVICE 

Williams, M. Y. 

1942. Notes on the fauna of Bruce Peninsula, Manitoulin and adjacent 
islands. Canadian Field-Nat. 56 (6) : 92. September. 

WlLLI.\MS, S. R. 

1909. On hibernation in the raccoon. Ohio Nat. 9 (6): 495-496. April. 
Wood, Fraxk Elmer. 

1910. A study of the mammals of Champaign County, Illinois. Illinois State 

Lab. Nat. Hist. Bull. 8: 573. 
Wood, N. A. 

1911. Results of the Mershon E.xpedition to the Charity Islands, Lake 

Huron. Michigan Acad. Sci. 13th Ann. Rept., p. 133. 

1912. Notes on Michigan mammals. Michigan Acad. Sci. 14th [Annual] 

Rept., p. 164. 
1914. An annotated check-list of Michigan mammals. Univ. Michigan Mus. 

Zool. Occas. Papers No. 4, p. 9. April 1. 
1922. The mammals of Washtenaw County, Michigan. Univ. Michigan 
Mus. Zool., Occas. Papers, No. 123, p. 10. 
Wood, Normax A., and L. R. Dice. 

1924. Records and distribution of Michigan mammals. Papers Michigan 
Acad. Sci., Arts and Letters 3: 437-438. 
WoRTMAN, J. L., and William Diller Matthew. 

1899. Ancestry of certain members of the Canidae, the Viverridae, and 
Procyonidae. Amer. Mus. Nat. Hist. Bull. 12: 109, 131-138. 
Wright, A. H., and S. E. R. Si.mpson. 

1920. The vertebrates of the Otter Lake Region, Dorset, Ontario. Canadian 
Field-Nat. 34: 167. December. 
Yeager, Lee E. 

1942. Coal-stripped land as a mammal habitat, with reference to fur animals. 

Amer. Midland Nat. 27 (3): 621-622. May. 
Yeager, Lee E., and R. G. Rennels. 

1943. Fur yield and autumn foods of the raccoon in Illinois River bottom 

lands. Jour. Wildlife Mangt. 7 (1): 4.5-60. January. 



PLATES 



J 



PLATE 2 

A. Procyon [Proct/ou] lotor lotor (Liiuiacnis) ; male adult; Lake Drummond, Dismal 

Swamp, Va. (No. 75255, U. S. Natl. Mus., Biological Surveys collection.) 
Note long pelage, uniformly directed l)ackward. 

B. Procyon [Eiiprocyon] cancrivorxs panamensis Goldman; female adult; Gatun, 

Canal Zone, Panama. (No. 171229, U. S. Natl. Mus., Biological Surveys 
collection.) Note short pelage, reversed on nape. 

108 



North American Fauna 60, Fish and Wildlife Service PLATE 2 







Skins of the subgenera Procj/on and Eiiprocijon. 



109 



PLATE 3 

[Two-thirds natural size] 

A. Procyon [Procyon] lotor lotor (Linnaeus): [male] adult; Sing Sing, N. Y. (No. 

129146, U. S. Natl. Mus., Biological Surveys collection.) 

B. Procyon [Procyon] lotor hirtus Nelson and Goldman; male adult; Elk River, 

Minn. (No. 18792G, U. S. Natl. Mus., Merriam collection.) 

110 



NORTH AMERICAN FAUNA 60. FiSH AND WILDLIFE SERVICE PLATE 3 




..i^fT^., 






m 




Skulls of Piocyon, subgenus Procyon. 



Ill 



PLATE 4 

[About three-fourths natural size] 



Proci/on [Prori/on] lotor lUoreus Nelson and Goldman; type; [male] adult; 

Saint Simon Island, Ga. (No. 2450, U. S. Natl. Mus.) 
Procyon [Procj/on] lotor ehicus BaiiKs; male adult; Fort Kissimmee, Fla. 

(No. 64002, U. S. Natl. Mus., Biological Surveys collection.) 



112 



North American Fauna 60. Fish and Wildlife Service PLATE 4 




f 




Skulls of P roc yon, subgenus Procyon. 



113 



IM.ATE 5 

[AhotU Ihicr-fourths natural sizo] 

Proci/oii [Proryon] lotor incaiitiix Nelson; type; male adult: Torch Key, Fla. 

(No. 255060, U. S. Natl. Mus., Biological Surveys collection.) 
Procijon [Proci/on] mminardi Bangs; male adult; New Providence Island, 

Bahamas. (No. 121905, U. S. Natl. Mus.) 

114 



NORTH AMERICAN FAUNA 60, FlSH AND WILDLIFE SERVICE PLATE 5 



■^K^ 



^■ 





JCa, -• 

o 



^ 



Skulls of Procyon, subgenus Procyon. 



115 



PLATE 6 

[About four-flfths natural size] 

A. Procyon [Procyon] lotor excelsus Nelson and Goldman; type; male adult; 

Owyhee River, Oreg., 10 miles west of Fairylawn, Idaho. (No. 236214, 
U. S. Natl. Mus., Biological Svirveys collection.) Distinguished by very 
large size. 

B. Procyon [Procyon] lotor psora Gray; male adult; Nicasio, Calif. (No. 187936, 

U. S. Natl. Mus., Merriam collection.) 

116 



North American Fauna 60, Fish and Wildlife Service PLATE 6 




,,-^^ 




Skulls of P roc you, subgenus Procyon. 



117 



PLATE 7 

[Throe-fourths natural size] 

A. Procijon [Froci/on] lotor henianfJezii Wagler; male adult; Tlalpam, Vallev of 

Mexico. Mexico. (No. 51151, U. S. Natl. Mus., Biolooical Surveys 
collection.) 

B. Procyon [Proci/on] lotor pinnilua Miller; male achilt; Porto Bello, Panama. 

(No. 171484, U. S. Natl. Mu.s., Biological Surveys collection.) 

118 



NORTH American Fauna 60, Fish and Wildlife Service PLATE 7 




1>': 



f 




I) % 



Skulls of Procijoii, subgenus Procijon. 



119 



PLATE 8 

[Five-sixtlis natural size] 

A. Procyon [Procyon] pygmaeus Merriani; type; male voung adult; Cozumel 

Island, Yucatan, INIexico. (No. 108511, "U. S. Natl. Mus., Biological Surveys 
collection.) 

B. Procyon [Procyon] insiilaria insidaris Merriam; male adult; Man'a Aladre 

Island, Nayarit, :\Iexico. (No. 88978, U. S. Natl. Mus., Biological Surveys 
collection.) Note heavy zygomatic arch. 

120 



North American Fauna 60, Fish and wildlife Service plate 



f^'^ 





Skulls of Procijon, subgenus Procyon. 



121 



876119°— .")() 9 



[Five-si'VfUths natural size] 

A. Proci/on [Proei/on] lotor lolor (Liimaevis) ; [male] adult; SiiiK Sing, X. Y. (No. 

121) 146. r. S. XaM. Mus., Biological Surveys oollectioii.) 

B. Procyon [Prociion] lotor hirtiis Nelson and Goldman; male adult: Klk River, 

Minn. (No. 187".»2(), V . S. Natl. Mus., Merriam collection.) 

122 



North American Fauna 60, Fish and Wildlife Service PLATE 9 




V 






'^•p^/^/^i 





Skulls of Procijon, subgenus Procyou. 



123 



PLATE 10 

(Throe-fourths natural sizoj 

A. Frocijon [Frocqo)i] lutor litoreus Nelson and Goldman: Ivpc; [male] adult: Saint 

Simon Islaiid, Ga. (No. 2450, U. S. Natl. Mns.) 

B. Procyon [Procyon] lotor elucus Bangs; male adult: Fort Kissimmee, Fla. (No. 

64b02, U. S". Natl. Mu.s., Biological Surveys collection.) 

124 



North American Fauna 60. Fish and Wildlife Service PLATE 10 





^^-^M^ 



^''M 




Skulls of Procyon, subgenus Procyon. 



125 



.sT0li0''--r)U 10 



PLATE 11 

|Tlii-eo-fourths+ natural size] 

A. Procyon [Procyon] lotor incautus Nelson; tvpe; male adult; Torch Kev, Fla. 

(No. 255060, U. S. Natl. Mus., Biological Surveys collection.) 

B. Proci/on [Procijon] uiat/nardi Bangs; male adult; New Providence Island, 

Bahamas. (No. 121905, U. S. Natl. Mus.) 

126 



North American Fauna 60, Fish and Wildlife Service plate 11 




'm-.H 



t 



-J59i 





Skulls of Piocyon, subgenus Frocijon. 



127 



PLATK 12 

[About three-fourths natural size] 

Procijon [Procf/on] lator excelsus Nelson and Goldman; type; male adult; Owy- 
hee River, Oreg., 10 miles west of Fairylawn, Idaho. (No. 236214, U. S. 
Natl. Mus., Biological Surveys collection.) Distinguished by very large 
size. 

Procyon [Pi-oci/on] lotor -psora Gray; male adult; Nicasio, Calif. (No. 187936, 
U.S. Natl. Mus., Merriam collection.) 

128 



NORTH AMERICAN FAUNA 60, FlSH AND WILDLIFE SERVICE PLATE 12 






# * 







Skulls of Procijon, subgenus Pr 



ocyon. 



129 



PLATE 13 

[About three-foui'ths natural size] 

A. Proci/on [Froci/o/i] lotor hernandezii Wagler; male adult; Tlalpam, Vallev of 

Mexico, Mexico. (No. 51151, U. S. Natl. Mas., Biological Surveys collec- 
tion.) 

B. Procyon [Procyon] lotor putiiilus Miller; male adult; Porto Bello, Panama. (No. 

171484, U. S. Natl. Mus., Biological Surveys collection.) 

130 



NORTH American Fauna 60, Fish and Wildlife Service PLATE 13 



^ 




•^**# 
;§»• 



Cji 



"-sr 



."i ■^fS^ 



.;sJ£»* 



'*Hi 







# 



Skulls of Procijon, subgenus Procyon. 



131 



PLATE 14 

IThree-fourths natural size] 

A. Procyon [Procyo/i] pyginaeus Merriam; type; male young adult ; Cozumel Island, 

Yucatan, Mexico. (No. 108511, U. S. Natl. Mus., Biological Surveys 
collection.) 

B. Procyon [Procyon] insularis insularis Merriam; male adult; Maria Madre 

Island, Nayarit, Mexico. (No. 88978, U. S. Natl. Mus., Biological Surveys 
collection.) 

132 



North American Fauna 60, Fish and Wildlife Service PLATE 14 



,j^ ***"■'' 



■f% 



i^- 













^ 



iS.^ 



C^i 





Skulls of Procyon, subgenus Procyon. 



133 



PLATE 15 

[Seven-tenths natural size] 

A. Proci/on [Procyon] lotor lotor (Linnaeus); [male] adult; Sing Sinn, X. Y. (No. 

12'9146, U. S. Natl. Mus., Biological Surveys collection.) 

B. Procyon [Procyon] lotor hirtus Nelson and Goldman; male adult; Elk River, 

Minn. (No. 187926, U. S. Natl. Mus., Merriam collection.) 

134 



North American Fauna 60, Fish and Wildlife Service PLATE 15 




^^^ 



■■\y 




Skulls of Frocyon, subgenus Procyon. 



135 



PLATE 16 

[About (linT-fourths natural sizf] 

A. Procyon [Proctjon] lotor litoreus Nelson and Goldman; type; [male] adult; 

Saint Simon Island, Ga. (No. 2450, U. S. Natl. Mus.) "Note heavy denti- 
tion. 

B. Proci/on [Proci/on] lotor elucas Bangs; male adult; Fort Kissimmee, Fla. (No. 

64002, U. S". Natl. Mus., Biological Surveys collection.) 

136 



North American Fauna 60. Fish and Wildlife Service PLATE 16 




^^' 




§" 'k ■■/ 






't; 



fit 










^9sSLL^^ 





Skulls of Procijon, subgenus Procyon. 



137 



PLATE 17 

'.bout tlirce-fourths natural size] 



A. Frocyon [Procyon] lotor incuutiis Nelson; tvpe; male adult; Torch Key Fla 

(No. 255060, U. S. Natl. Mus., Biological Surveys collection.) 

B. Procyon [Procyon] maynardi Bangs; male adult; New Providence Island, 

Bahamas. (No. 121905, U. S. Natl. Mus.) 

138 



NORTH AMERICAN FAUNA 60, FiSH AND WILDLIFE SERVICE PLATE 17 



r 






f^< 



7' • 



■>■' 



Skulls of Frocyon, subgenus Prociion. 



139 



PLATE 18 

[About three-fourths natural size] 

A. ProcyoN [Proctioii] lotor excelsus Nelson and Goldman; type; male adult : Owyhee 

River, Oreg., 10 miles west of Fairylawn, Idaho. (No. 236214, U. S. Natl. 
Mus., Biological Surveys collection.) Distinguished by very large size. 

B. Procijon [Procyon] lotor psora Gray; male adult: Nicasio, Calif. (No. 187936, 

U. S. Natl. Mus., Merriam collection.) 

140 



NORTH AMERICAN FAUNA 60, FlSH AND WILDLIFE SERVICE PLATE 18 







^>. 







Skulls of Procyon, subgenus Procyon. 



141 



S7C119°— 50 11 



PLATE 19- 

[Five-sixths natui'al size] 



A. Procyon [Procyon] lotor hernandezii Wagler; male adult; Tlalpain, Valley of 

Mexico, Mexico. (No. 51151, U. S. Natl. Mus., Biological Surveys collec- 
tion.) 

B. Procyon [Procyon] lotor pumilus Miller; male adult ; Porto Bello, Panama. (No. 

171484, U. S. Natl. Mus., Biological Surveys collection.) 



142 



North American Fauna 60. Fish and Wildlife Service PLATE 19 







>? 





iT-^ 



,- i.* f*^ '>^ 



^ 



r 



jsit^rA. 







/ 



10- 




1/ 



...^,^ 




Skulls of Proci/on, suljgeims Procyon. 



143 



PLATE 20 

[About two-thirds natui'al size] 

Procyon [Procyon] pi/gmaeus Merriam ; tvpe; male voung adult ; Cozumel Island, 
Yucatan, Mexico! (No. 108511, U. S." Natl. Mus., Biological Surveys collec- 
tion.) 

Procyon [Procyon] insularis insularis Merriam; male adult; Maria Madre 
Island, Nayarit, Mexico. (No. 88978, U. S. Natl. Mus., Biological Surveys 
collection.) 

144 



North American Fauna 60. Fish and Wildlife Service PLATE 20 



■ i 




'-^ i 



'"> >.| ., 



r: 



>: 



•SS^'jr-' 



t 




^ 



* 







Skulls of Frocijun, subgenus Procyon. 



145 



PLATl'l 21 

[.Six-si'vcnths naliiml size] 

Procyon [Proci/nn] lotor eluais Baiig.s; male adult: Fort Kissinunee, P'la. (No. 
64002, U. S. Natl. Mus., Biological Surveys collection.) Note larger size 
and greater angularity of male. 

Prociion [Procyon] lotor elucus Bangs: female adult; Fort Kissimmee, Fla. 
(No. 64012, U. S. Natl. Mus., Biological Surveys collection.) Note smaller 
size and lesser angularity of female. 

Proci/o)! [Prort/on] lotor lotor (Linnaeus): female adult; Schroon Lake, N. Y. 
(No. ilSSDvO, r. S. Natl. Mus., liiological Surveys collection.) Lateral view 
of tyi:)ical mandibular ramus. 

146 



North American fauna 60, Fish and Wildlife Service PLATE 21 





c >v5^'r 



Skulls, including a mandiliular ramus, of Procyon, subgenus J'roryon. 

147 



PLATE 22 

[ Vbout flvt'-sixths natural size] 

A. Procyon [Euprocyon] cancrivorus panamensis Goldman; type; female adult; 

Gatun, Canal Zone, Panama. (No. 171669, U. S. Natl. Mus., Biological 
Surveys collection.) Upper surface of cranium. 

B. Procyon [Euprocyon] cancrivorus panamensis Goldman; tvpe; female adult; 

Gatun, Canal Zone, Panama. (No. 171669, U. S. Natl. Mus., Biological 
Surveys collection.) Under surface of cranium. Note molar crowns with 
rounded cusps adai)ted for crushing food; and compare with more trenchant 
cusps in subgenus Procyon. 

148 



North American Fauna 60. Fish and Wildlife Service PLATE 22 













Skulls of Procijon, subgenus Eaprocyon. 



149 



INDEX 

[Pago numbers in boldface rcftT to principal entries; those in italic to synonyms 



abru])t.a, Ursus cauda, 3. 
Ailuropoda, 1. 
Ailurus, 1. 

Alabama raccoon, 38. 
albina, Procyon lotor, 5. 
aiimilatiis, Procyon, 33, 35. 
aiispicatus, Procyon lotor, 22, 24, 2S, 44, 
46, 47. 48. 

Haliania raccoon, 75. 
Paja California raccoon, 62. 
Barl)ados raccoon, 79. 
Bassaricyon, 1, 26. 
lirachyurus, Procyon, 4. 
breeding, 13. 

California raccoon, 56. 

calif ornicns, Procyon lotor, 56, 57. 

Canij^eche raccoon, 65. 

Cani])siurus, 25. 

cancrivonis, Euprocyon, 16. 

Procyon, 16, 31, 80, 81. 

Procyon cancrivorns, 82, 83. 

Ursus, 4, 25, 81. 
castaneus, Procyon hernandezi, 5. 
coastal marsh raccoon, 85. 
Colorado Desert raccoon, 54. 
Costa Rican raccoon, 69. 
Cozumel Island raccoon, 76. 
crab-eating raccoon, 4. 
crassidens, Procyon lotor, 24, 28, 66, 67, 

68, 69, 71. 
Cynodictis, 2. 

dickeyi, Procyon lotor, IS, 24, 28, 66, 
67, 69, 70. 

eastern raccoon, 33. 

economic status, 14. 

elongata, Ursus cauda, 3, 6. 

elucus, Procyon lotor, 22, 24, 27, 38, 39, 

40, 42, 44, 46, 49, 113, 125, 137, 147. 
Euprocyon, 1, 2, 3, 4, 25, 27, 28, 29, 80. 

cancrivorus, 16. 

cancrivorus panamensis, 28, 82. 



excelsus, Procyon lotor, 20, 24, 28, 33, 
56, 58, 59, 60, 86, 117, 129, 141. 

flavidus, Procyon lotor, 5. 

Florida raccoon, 42. 

food, 7. 

fusca, Procyon brachyurus, 5. 

fuscipes, Procyon lotor, 24, 28, 37, 38, 
39, 49, .52, 64, 65, 84. 

general activities, 7. 

gloveralleni, Procyon, 22, 24, 28, 79. 

grinnelli, Procyon lotor, 24, 2S, 54, 55, 

62. 
Guadeloupe Island raccoon, 77. 
gularis, Procyon, 33, 35. 

hernandezii, Procyon, 64. 

hernandezii, Procyon lotor, 22, 24, 28, 

.50, .52, 53, 64, 66, 67, 72, 74, 119, 

131, 143. 
hit)ernation, 14. 

Hilton Head Island raccoon, 41. 
hirtus, Procyon lotor, 22, 24, 27. 33, 34, 

35, 37, 50, 52, 111, 123, 135. 
hudsonicus, Procyon, 5. 

incautus, Procyon lotor, 24, 28, 44, 46, 

47, 48, 75, 115, 127, 139. 
ines])eratus, Procyon lotor, 24, 28, 43, 

44, 46, 47, 48. 
instincts, 9. 
insularis, Procyon, 20, 71. 

Procyon insularis, 24, 28, 72,74, 121, 
133, 145. 
Isthmian raccoon, 70. 

key to suligenera, 27. 

to species and subspecies of svibgenus 
Procyon, 29. 
Key Vaca raccoon, 47. 

litoreus, Procyon lotor, 22, 24, 27, 33, 
34, 35, 40, 41, 42, 43, 80, 85, 86, 
109, 111, 113, 123, 12.5, 135, 137, 147. 

longevity, 7. 

Lotor, 25. 

151 



152 NORTH AMERICAN FAUNA GO, FISH AND WILDLIFE SERVICE 



lotor, Meles, 33, 35. 

Procyon, 1, 31, 33, 52. 

Procyon lotor, 21, 24, 27, 33, 35, 38, 
39, 40, 41, 43, 56, 57, 75. 

Ursus, 4, 6, 25, 83. 
Lotor vulgaris, 33, 35. 

Mamprocyonus, 25. 

Mapachiu, 82. 

Maria Madre Island raccoon, 72. 

Maria Magdalena Island raccoon, 73. 

marinus, Procyon lotor, 24, 27, 42, 43, 

44, 46, 47, 48. 
maritimus, Procyon lotor, 24, 36, 85. 
Matecumbe Key raccoon, 46. 
maynardi, Procyon, 24, 28, 75. 78. 79, 

115, 127, 139. 
megalodous, Procyon lotor, 24, 39, 51, 

84. 
melanus, Procyon lotor, 5. 
Meles lotor, 33, 35. 
Mexican Plateau raccoon, 64. 
Mexican raccoon, 52. 
mexicana, Procyon hernandezii, 52. 
mexicanus, Procyon lotor, 24, 28, 50, 

52, 54, 55, 63, 64, 65, 72, 74, 85, 86. 
minor, Procyon, 24, 28, 75, 77, 79. 
Mississippi Delta raccoon, 84. 
molt, 19. 

nanus, Procyon, 3. 
Nasua, 1. 
Nasuella, 1. 
nivea, Procyon, 4. 
nomenclature, 3. 

obscurus, Procyon, 5. 
ochraceus, Procyon lotor, 54, 55. 

Pacific Northwest raccoon, 58. 
pacifica, Procyon lotor, 28. 

Procyon psora, 58. 
pacificus, Procyon lotor, 24, 56, 57, 58, 

59, 60, 61, 86. 
pallidus, Procyon, 54. 

Procyon lotor, 20, 24, 28, 52, 54, 56, 
57^ 62, 63. 
Panama crab-eating raccoon, 82. 
panamensis, Euprocyon cancrivorus, 28, 
82. 

Procyon cancrivorus, 28, 81, 82, 109, 
149. 
pelage, 19. 
Phlaocyon, 2. 
Potos, 1, 27. 



])riscus, Procyon, 3. 
Procyon, 25, 27, 28, 29. ' 
annulatus, 33, 35. 
l)rachyurus, 4. 
brachyurus fusca, 5. 
cancrivorus, 16, 31, 80, 81. 
cancrivorus cancrivorus, 82, 83. 
cancrivorus panamensis, 28, 81, 82, 

109, 149. 
cancrivorus proteus, 82, 83. 
gloveralleni, 22, 24, 28, 79. 
gularis, 33, 35. 
hernandezi castaneus, 5. 
hernandezli, 6Ji. 
hernandezii mexicana, 52. 
hudsonicus, 5. 
insularis, 20, 71. 
insularis insularis, 24, 28, 72. 74, 121, 

133, 145. 
insularis vicinus, 24, 28, 73. 
lotor, 1, 31, 32, 33, 52. 
lotor all)ina, 5. 
lotor auspicatus, 22, 24, 28, 44, 46, 

47, 48. 
lotor calif ornicus, 56, 57. 
lotor crassidens, 24, 28, 66, 67, 68, 

69, 71. 
lotor dickeyi, 18, 24, 28, 66, 67, 69, 70. 
lotor elucvis, 22, 24, 27, 38, 39, 40, 42, 

44, 46, 49, 113, 125, 137, 147. 
lotor excelsus, 20, 24, 28, 33, 56, 58, 

59, 60, 86, 117, 129, 141. 
lotor flavidus, 5. 
lotor fuscipes, 24, 28, 37, 38, 39, 49. 

52, 64, 65, 84. 

lotor grinnelli, 24, 28, 54, 55, 62. 
lotor hernandezii, 22, 24, 28, 50, 52, 

53, 64, 66, 67, 72, 74, 119, 131, 143. 
lotor hirtus, 22, 24, 27, 33, 34, 35, 37, 

50, 52, 111, 123, 135. 
lotor incautus, 24, 28, 44, 46, 47, 48, 

75, 115, 127, 139. 
lotor inesperatus, 24, 28, 43, 44, 46, 

47, 48. 
lotor insularis, 72. 
lotor litoreus, 22, 24, 27, 33, 34, 35, 

40, 41, 42, 43, 80, 113, 125, 137 
lotor lotor, 21, 24, 27, 33, 35, 37, 38, 

39, 40, 41, 43, 56, 57, 75, 85, 86, 

109, 111, 123, 135, 147. 
lotor marinus, 24, 27, 42, 43, 44. 46, 

47, 48. 
lotor maritimus, 24, 36, 85. 
lotor megalodous, 24, 39, 51, 84. 



INDEX 



153 



Procyon — Continued 
lotor melanus, 5. 
lotor mexicanus, 24, 28, 50, 52, 54, 

55, 63, 64, 65, 72, 74, 85, 86. 
lotor ocbraceus, 51^, 55. 
lotor pacificus, 24, 28, 56, 57, 58, 59, 

60, 61, 86. 
lotor imllidus, 20, 24, 28, 52, 54, 56, 

57, 62, 63. 
lotor psora, 13, 24, 28, 54, 56, 58, 60, 

62, 63, 86, 117, 129, 141. 
lotor pumiliis, 24, 28, 69, 70, 80, 83, 

118, 131, 143. 
lotor rufeseens, 5. 
lotor shufeldti, 24, 28, 64, 65, 67, 68, 

70, 76. 
lotor solutus, 24, 27, 33, 34, 35, 40, 41. 
lotor vancouverensis, 24, 28, 58, 61. 
lotor varius, 24, 27, 33, 34, 35, 38, 42, 

43, 50, 84. 
mayuardi, 24, 28, 75, 78, 79, 115, 127, 

139. 
minor, 24, 28, 75, 77, 79. 
nanus, 3. 
nivea, 4. 
obscunis, 5. 
pallidus, 54* 
priscus, 3. 
proteus, 58, 59, 
psora, 56. 
psora pacifica, 58. 
pumilus, 70. 
pygmaeus, 20, 24, 28, 76, 121, 133, 

145. 
simus, 3. 
proteus, Procyon, 58, 59. 

Procyon cancrivorus, 82, 83. 
psora, Procyon, 56. 

Procyon lotor, 13, 24, 28, 54, 56, 58, 

60, 62, 63, 86, 117, 129, 141. 
pumilus, Procyon, 70. 

Procyon lotor, 24, 28, 69, 70, 80, 83, 

118, 131, 143. 
pygmaeus, Procyon, 20, 24, 28, 76, 121, 
133, 145. 

raccoon, Alabama, 38. 
Bahama, 75. 
Baja California, 62. 
Barbados, 79. 
California, 56. 
Campeche, 65. 
coastal marsh, 85. 
Colorado Desert, 54. 
Costa Rican, 69. 



ra ccoon — continued 

Cozumel Island, 76. 

crab-eating, 4. 

eastern, 33. 

Florida, 42. 

(Juadeloupe Island, 77. 

Hilton Head Island, 41. 

Isthmian, 70. 

Key Vaca, 47. 

Maria Madre Island, 72, 

Maria Magdalena Island, 73. 

Matecumbe Key, 46. 

Mexican, 52. 

Mexican Plateau, 64. 

Mississippi Delta, 84. 

Pacific Northwest, 58. 

Panama ci-ab-eating, 82. 

Saint Simon Island, 40. 

Salvador, 67. 

Snake River Valley, 60. 

Ten Thousand Islands, 44. 

Texas, 49. 

Torch Key, 48. 

Upper Mississippi Valley, 37. 

Vancouver Island, 61. 
rufescens, Procyon lotor, 5. 

Saint Simon Island raccoon, 40. 

Salvador raccoon, 67. 

senses, 9. 

shufeldti, Procyon lotor, 24, 28, 64, 65, 

67, 68, 70, 76. 
simus, Procyon, 3. 
Snake River Valley raccoon, 60. 
solutus, Procyon lotor, 24, 27, 33, 34, 

35, 40, 41. 

Ten Thousand Islands raccoon, 44. 
Texas raccoon, 49. 
Torch Key raccoon, 48. 

Upper Mississippi Valley raccoon, 37. 
Ursus cancrivorus, 4, 25, 81. 

Cauda abrupta, 3. 

Cauda elongata, 3, 6. 

lotor, 4, 6, 25, 33. 

Vancouver Island raccoon, 61. 
vancouverensis, Procyon lotor, 24, 28, 

58, 61. 
varius, Procyon lotor, 24, 27, 33, 34, 

35, 38, 42, 43, 50, 84. 
vicinus, Procyon insularis, 24, 28, 73. 
vulgaris, Lotor, 33, 35. 



weights, 22. 



o 



FAUNA OF 
THE ALEUTIAN ISLANDS 
AND ALASKA PENINSULA 



WITH NOTES ON 

INVERTEBRATES AND FISHES 

COLLECTED IN THE ALEUTIANS. 

1936-38 







NUMBER 61 



UNITED STATES 

DEPARTMENT OF THE INTERIOR 

FISH AND WILDLIFE SERVICE 



FAUNA OF 



^A 



UNITED STATES DEPARTMENT OF THE INTERIOR 
FISH AND WILDLIFE SERVICE 

NORTH AMERICAN FAUNA NO. 61 

Fauna of the Aleutian Islands and Alaska Peninsula 

CORRECTION 

A report on so great an area as the Aleutian Islands and the 
Alaska Peninsula, to be comprehensive, must include data col- 
lected by many persons. While I was preparing this report, 
at a time when Ira N. Gabrielson and Frederick C. Lincoln 
were preparing their comprehensive work, Birds of Alaska, 
Dr. Gabrielson kindly lent me his field notes of Alaska trips. 
With his own notes Dr. Gabrielson also suppHed field notes of 
Frank Beals, who was in the Aleutians in the 1940's for the Fish 
and Wildhfe Service, and a few notes of Douglas Gray, who 
made some fine observations in the Aleutians. 

Immediately after issue of this Fauna, Dr. Gabrielson noted 
that several observations were attributed to his notes whereas 
they should have been credited to Frank Beals; somehow, in the 
compilation of many observations the original sources of the 
field notes were confused. Dr. Gabrielson points out that his 
field work in the areas concerned covered the following periods: 
June and July of 1940 ; late September and early October of 1941 ; 
August of 1943; late July to mid-August of 1945; and June, 
July, and August of 1946. Where records outside these periods 
are attributed to Gabrielson's field notes, they should be cred- 
ited to Frank Beals. 

In correcting this misplaced credit, I join Dr. Gabrielson in 
thus giving recognition to the splendid biological notes made 
by Beals (as well as the excellent photographs of his, which 
I have seen). 

Boston Public Library m tat 

c • * J ^ Olaus J. MURIE 

February 1960 i^'ipenntendent of Documents 

DEPOSITORY 



•;r/, 



FAUNA OF 
THE ALEUTIAN ISLANDS 
AND ALASKA PENINSULA 

By Olaus J. Miirie, Biologist 



INVERTEBRATES AND FISHES 

COLLECTED IN THE ALEUTIANS, 

1936-38 



By Victor B. Sclieffer, Biologist 









FAUNA 



NUMBER 61 



UNITED STATES DEPARTMENT OF THE INTERIOR 

Fred A. Seatoii, Secretary 

FISH AND WILDLIFE SERVICE 
Arnie J. Suomela, Commissioner 



Boston Public Library 
Superintendent of Documents 



JAN 18 1960 



DEPOSITORY 




PIBLISHKD BY V. S. FISH AND WILDLIFE SERVICE • WASHINGTON • 1959 
PRINTED AT V. S. GOVERNMENT PRINTING OFFICE, WASHINGTON 



For sale by the Superintendent of Documents, I'. S. Government Printing Office, 
\^'ashington 23, D. C. Price $1.23 cent?. 



CONTENTS 

FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA 
PENINSULA, by Olaus J. Murie 

Page 

Foreword ^iii 

Introduction l 

Geography and geolog-y 3 

Climate 10 

Environment and biotic distribution 11 

Geographic and geologic influences 12 

Asiatic immigrants 12 

Bering Sea avifauna 14 

Fauna of wider northern distribution . , 15 

Southern and southeastern birds 16 

Environmental influence 17 

Ecological classification 18 

Vegetation 22 

Birds 27 

Family Gaviidae 28 

Gavia uhdio-, Common loon 28 

Gavia adaynsii, Yellow-billed loon 29 

Gavia a)cUca, Arctic loon 29 

Gavia stellata, Red-thioated loon 31 

Family Podicipedidae 32 

Podiceps grisegena, Red-necked grebe 32 

Podiceps a}(7'itHs, Horned grebe 33 

Family Diomedeidae 34 

Diomedca nigripes, Black-footed albatross 34 

Diomedea albati'us, Short-tailed albatross 36 

Diomedea immutahilis, Laysan albatross 39 

Family Procellariidae 41 

Puffinns tenuirostris, Slender-billed shearwater 41 

Puffinus gnseiis, Sooty shearwater 44 

Pterodroma inexpectata, Scaled petrel 44 

Pterodroma cookii, Cook's petrel 45 

Fulmarus glaciali!^, Fulmar 45 

Family Hydrobatidae 48 

Oceanodvoma fnrcata, Fork-tailed petrel 48 

Oceanodroma leiworhoa, Leach's petrel 51 

Family Phalacrocoracidae 52 

Phalacrocorax au)-it}is, Double-crested cormorant 52 

Phalacrocorax pelagicus, Pelagic cormorant 55 

Phalacrocorax ^rile, Red-faced cormorant 57 

Family Ardeidae 59 

Ardea hcrodias, Great blue heron 59 

iii 



Page 

Family Anatidae 59 

Olor columbiamis, Whistling swan 59 

Olor buccinator, Trumpeter swan 61 

Branta cayiadensis, Canada goose 61 

Branta nigricans. Black brant 67 

Philacte cayiagica, Emperor goose 69 

Anse7- albifrons, White-fronted goose 73 

Chen hyperborea. Snow goose 74 

Anas platyrhynchos, Mallard 74 

Atias strepera, Gadwall 76 

Anas acuta. Pintail 77 

Anas falcata, Falcated teal 78 

Anas crecca. Common teal 79 

Arias Caroline tisis, Green-winged teal 80 

Mareca penelope, European widgeon 82 

Ma/reca americana, American widgeon 82 

Spatula clypeata, Shoveler 83 

Aythya americana, Redhead 83 

Aythya valisineria, Canvasback 84 

Aythija maiila, Greater scaup 84 

Aythya affinis, Lesser scaup 86 

Aythya fuligula, Tufted duck 86 

Bucephala clangula. Common goldeneye 86 

Bucephula islandica, Barrow's goldeneye 87 

Bucephala albeola, Bufflehead 88 

Clangula hyemalis, Oldsquaw 89 

Histrionicus histrionicus, Harlequin duck 90 

Polysticta stelleri, Steller's eider 92 

Somateria mollissima, Common eider 94 

Sotnateria spectabilis. King eider 97 

Lampronetta fischeri. Spectacled eider 98 

Melanitta deglandi, White-winged scoter 99 

Melanitta perspicillata, Surf scoter 101 

Oidemia nigra, Common scoter 102 

Mergus merganser, Common merganser 104 

Mergus serrator, Red-breasted merganser 105 

Family Accipitridae 106 

Accipiter gentilis. Goshawk 106 

Accipiter striatus. Sharp-shinned hawk 107 

Buteo lagopus. Rough-legged hawk 107 

Aquila chrysaetos. Golden eagle 109 

Haliaeetus albicilla. Gray sea eagle 110 

Haliaeetus leucocephalus, Bald eagle Ill 

Haliaeetus pelagicus, Steller's sea eagle 117 

Circus cyaneus. Marsh hawk 117 

Pandion haliaetus, Osprey 118 

Family Falconidae 118 

Falco nisticolus, Gyrfalcon 118 

Falco peregrinus, Peregrine falcon 119 

Falco columbarius. Pigeon hawk 120 

Falco sparverius. Sparrow hawk 121 

Family Tetraonidae 121 

Canachites canadensis, Spruce grouse 121 



IV 



Page 

Lagopus lagopus, Willow ptarmigan 122 

Lagopiis miitns. Rock ptarmigan 123 

Family Gruidae 129 

Grus canadensis, Sandhill crane 129 

Family Rallidae 130 

Fulica ainericana, American coot 130 

Family Haematopodidae 130 

Haematopus hachmani, Black oystercatcher 130 

Family Charadriidae 132 

Charadrius dtibius, Little ringed plover 132 

Charadrius semipalmatus, Semipalmated plover 132 

Phivialis dominica, American golden plover 133 

Squatarola squatarola, Black-bellied plover 134 

Aphriza virgata, Surf bird , 135 

Arenaria interpres, Ruddy turnstone 135 

Areyiaria melanocephala, Black turnstone 136 

Family Scolopacidae 137 

Capella gallinago, Common snipe 137 

Numenius phaeopus, Whimbrel 138 

Numenius tahiticnsis, Bristle-thighed curlew 138 

Actitis viacularia, Spotted sandpiper 138 

Tringa glareola, Wood sandpiper 139 

Heteroscelns incanum, Wandering tattler 139 

Totanus ruelanoleucus, Greater yellowlegs 140 

Totanus flavipes, Lesser yellowlegs 141 

Calidris canutus, Knot 141 

Erolia ptilocnemis, Rock sandpiper 141 

Erolia acu^niyiata, Sharp-tailed sandpiper 146 

Erolia melanotos, Pectoral sandpiper 146 

Erolia bairdii, Baird's sandpiper 147 

Erolia minutilla, Least sandpiper 147 

Erolia alpina, Dunlin 150 

Limnodromus griseiis, Short-billed dowitcher 150 

Ereunetes pusillus, Semipalmated sandpiper 152 

Ereunetes maun, Western sandpiper 152 

Limosa fedoa, Marbled godwit 153 

Limosa lapponica, Bar-tailed godwit 153 

Limosa haemastica, Hudsonian godwit 154 

Crocethia alba, Sanderling 154 

Family Phalaropodidae 155 

Phalaropus fuliearins. Red phalarope 155 

Lobipes lobatus, Northern phalarope 156 

Family Stercorariidae 157 

Stercorarius pomarinus, Pomarine jaeger 157 

Stercorarius parasiticus, Parasitic jaeger 159 

Stercorarius longicaudus. Long-tailed jaeger 161 

Family Laridae 162 

Larus hyperboreus, Glaucous gull 162 

Larus glaucescens, Glaucous-winged gull 165 

Larus schistisagus, Slaty-backed gull 171 

Lams argentatus, Herring gull 171 

Larus delaivarensis, Ring-billed gull 172 

Larus canus, Mew gull 172 

V 



Page 

Lams Philadelphia, Bonaparte's gull 174 

La7-us ridibundus, Black-headed gull 175 

Rissa tridactyla, Black-legged kittiwake 175 

Rissa hrevirostris, Red-legged kittiwake 176 

Xema sabini, Sabine's gull 178 

Steryia hirundo, Common tern 178 

Sterna paradisaea, Arctic tern 179 

Sterna aleutica, Aleutian tern 180 

Family Alcidae 182 

Uria aalge, Common murre 182 

Uria lomvia, Thick-billed murre 182 

Cepphus columba, Pigeon guillemot 186 

Brachyramphus marmoratum, Marbled murrelet 187 

Brachyramphus brevirostre, Kittlitz's murrelet 188 

Synthliboratnphus ayitiquum, Ancient murrelet 189 

Ptychoramphus aleutica, Cassin's auklet 191 

Cyclorrhynchus psittacida, Parakeet auklet 193 

Aethia cnstatella. Crested auklet 194 

Aethia pusilla, Least auklet 197 

Aethia pygmaea. Whiskered auklet 200 

Cerorhdnca nionocerata, Rhinoceros auklet 202 

Fratercula cornicnlata, Horned puflfin 202 

Lunda cirrhata. Tufted puffin 204 

Family Cuculidae 205 

Cucubis saturatns, Oriental cuckoo 205 

Family Strigidae 205 

Bubo virginianus. Horned owl 205 

Nyceta scandiaca. Snowy owl 206 

Surnia ulula. Hawk owl 207 

Asio flammeus, Short-eared owl 207 

Aegolius funereus. Boreal owl 209 

Family Trochilidae 210 

Selasphorus rufus, Rufous hummingbird 210 

Family Alcedinidae 210 

Megaceryle alcyon, Belted kingfisher 210 

Family Picidae 211 

Dendrocopos pubescens, Downy woodpecker 211 

Picoides arcticus, Black-backed three-toed woodpecker .... 211 

Pico'ides tridactylus. Northern three-toed woodpecker .... 212 

Family Tyrannidae 212 

Sayomis saya, Say's phoebe 212 

Family Alaudidae 212 

Eremophdla alpestris. Horned lark 212 

Family Hirundinidae 212 

Tachycineta thalassina, Violet-green swallow 212 

Iridoprocne bicolor. Tree swallow 213 

Riparia riparia, Bank swallow 213 

Hinindo rustica, Barn swallow 214 

Family Corvidae 214 

Perisoreus canadensis, Gray jay 214 

Pica pica, Black-billed magpie 215 



VI 



Page 

Corvus corax, Common raven 216 

Corvus caiiriuus, Northwestei'n crow 217 

Nucifraga columhiana, Clark's nutcracker 217 

Family Paridae 217 

Pariis atricapillus, Black-capped chickadee 217 

Parus hndsoniciift, Boreal chickadee 218 

Family Certhiidae 219 

Certhia fatniliaris, Brown creeper 219 

Family Cinclidae 220 

Cinchis mexicaniis, Dipper 220 

Family Troglodytidae 221 

Troglodytes troglodytes, Winter wren 221 

Family Turdidae 225 

Turdus migratorius, Robin 225 

Ixoreus yiaevius, Varied thrush 225 

Hylocichla guttata, Hermit thrush 226 

Hylocichla ustulata, Swainson's thrush 228 

Hylocichla mhiima, Gray-cheeked thrush 228 

Luscinia calliope, Siberian ruby throat 228 

Family Sylviidae 229 

Phylloscopus borealis, Arctic warbler 229 

Regulus satrapa, Golden-crowned kinglet 229 

Regnhis calendula, Ruby-crowned kinglet 229 

Family Motacillidae 230 

Motacilla alba. White wagtail 230 

Motacilla flava, Yellow wagtail 230 

Anthus spinoletta. Water pipit 231 

Anth}(s ceriunus, Red-throated pipit 233 

Family Laniidae 233 

Lanius exeubitor, Northern shrike 233 

Family Parulidae 234 

Vermivora celata, Orange-crowned warbler 234 

Dendroica petechia, Yellow warbler 234 

Dendroica coronata. Myrtle warbler 235 

Dendroica striata, Blackpoll warbler 235 

Seiurus noveboracensis, Northern water thrush 236 

Wilsonia pussila, Wilson's warbler 236 

Family Icteridae 236 

Euphagus carolinus, Rusty blackbird 236 

Family Fringillidae 237 

Pinicola enucleator, Pine grosbeak 237 

Leucosticte tephrocotis, Gray-crowned rosy finch 237 

Acanthis hornem-anni, Hoary redpoll 240 

Acanthis flammea. Common redpoll 240 

Spinus pinus, Pine siskin 242 

Loxia curvirostra. Red crossbill 242 

Loxia lencoptera. White-winged crossbill 242 

Passer cuius sandwichen^is, Savannah sparrow 243 

J unco hyernalis. Slate-colored junco 246 

Junco oregayius, Oregon junco 246 

Spizella arborea. Tree sparrow 247 

Zonotrichia leucophrys. White-crowned sparrow 247 

Zonotrichia atricapiUa, Golden-crowned sparrow 248 

vii 



Page 

Passerella iliaca, Fox sparrow 249 

Melospiza Uncolnii, Lincoln's sparrow 254 

Melospiza melodia, Song sparrow 254 

CalcariuH lapponicus, Lapland long-spur 257 

Plectrophenax nivalis. Snow bunting 258 

Plectrophenax hyperboreiis, McKay's bunting 260 

EmbeHza nisfica, Rustic bunting 260 

Mammals 262 

Family Soricidae 262 

Sorex cinereus. Cinereous shrew 262 

Sorex Uindrensis, Tundra saddle-backed shrew 263 

Sorex hydrodromus, Unalaska saddle-backed shrew 263 

Sorex obscurus, Dusky shrew 265 

Microsorex hoyi, Pigmy shrew 266 

Family Vespertilionidae 266 

Myotis lucifugus, Little brown bat 266 

Family Ursidae 266 

Euarctos americaniis, Black bear 266 

Ui'sus arctos, Brown bear 267 

Thalarctos mainti^mis, Polar bear 274 

Family Procyonidae 274 

Procyon lotor, Raccoon 274 

Family Mustelidae 275 

Maries americana, Marten 275 

Mustela erminea, Weasel 275 

Mustela rixosa, Least weasel 276 

Mustela vision, Mink 276 

Gulo liiscus, Wolverine 277 

Lutra canadensis, Otter 278 

Enhydra lutris. Sea otter 278 

Family Canidae 287 

Vulpes fiilva, Red fox 287 

Alopex lagopus, Blue fox 292 

Canis lupus, Wolf 304 

Family Felidae 305 

Lynx canadensis, Canada lynx 305 

Family Otariidae 305 

Eumetopias jubata, Steller sea lion 305 

Callorhinus lirsinus, Northern fur seal 306 

Family Phocidae 307 

Phoca vitulina. Harbor seal 307 

Pusa hispida. Ringed seal 309 

Pagophilus groenlandicus. Harp seal 309 

Histriophoca fasciata. Ribbon seal 310 

Erignathus barbatus, Bearded seal 310 

Family Odobenidae 311 

Odobenus rosmarus, Walrus 311 

Family Sciuridae 314 

Marmota caligata. Hoary marmot 314 

Citellus parryii. Ground squirrel 314 

Citellus kodiacensis. Ground squirrel 316 

Tamiasciurus hudsonicus. Red squirred 316 

viii 



Pasre 

Family Castoridae 317 

Castor canadensis, Beaver 317 

Family Cricetidae 317 

Synaptomys borealis, Lemming mouse 317 

Letnmns trimucronatiis, Lemming- 318 

Dicrostonyx groenlandicus. Collared lemming 318 

Clethrionomys riitilus, Red-backed mouse 320 

Microtus oeconomus, Meadow mouse 320 

Microtus pennsylvanicKs, Meadow mouse 324 

Ondatra zibethicus, Muskrat 324 

Family Muridae 324 

Mus musciilus, House mouse 324 

Rattus norvegicus, House rat 325 

Family Zapodidae 326 

Zapus hudsonius, Jumping mouse 326 

Family Erethizontidae 327 

Erethizon dorsatnm, American porcupine 327 

Family Ochotonidae 327 

Ochotona coUaris, Collared pika 327 

Family Leporidae 327 

Lcpus americaniis, Varying hare 327 

Lcpus otiius, Arctic hare 328 

Family Cervidae 328 

Cervus canadensis, Elk (Wapiti) 328 

Odocoilexis hemiomis, Black-tailed deer 328 

Alces alces. Moose 329 

Rangifer arcticus. Barren ground caribou 329 

Rangifer sp., Reindeer 331 

Family Bovidae 332 

Ovis dalli, Dall sheep (White sheep) 332 

Family Hydrodamalidae 332 

Hydrodamalis gigas, Steller sea cow 332 

Family Balaenidae 333 

Eubalaena siebokli, Pacific right whale 333 

Balaena mysticetiis, Bowhead whale 333 

Family Eschrichtidae 334 

Eschrichti)is ghaicus, Gray whale 334 

Family Balaenopteridae 334 

Balaenoptera physalus. Finback whale 334 

Balaenoptera borealis, Sei whale 334 

Sibbaldus musculus, Blue whale 335 

Megaptera yiovaeangliae, Humpback whale 335 

Family Physeteridae : . . 335 

Physeter catodon, Sperm whale 335 

Family Delphinidae 335 

Gi-atnpus reciipinna. Pacific killer whale 335 

Globicephala scammonii, Pacific blackfish 337 

Lissodelphis borealis, Right-whale porpoise 337 

Lagenorhynch^is obliqnidcns. Pacific striped porpoise 337 

Phocoena vomerina, Pacific harbor porpoise 337 

Phocoenoides dalli, Dall porpoise 338 

Family Monodontidae 338 

Delphinapterus leucas. White whale (Beluga) 338 

ix 



Page 

Famib'- Ziphiidae 339 

1 t7-adius hairdii, Baird beaked whale 339 

Mesoplodon stejnegeri, Stejneger beaked whale 339 

Ziphius cavirostri.^, Cuvier beaked whale 339 

References 34o 



ILLUSTRATIONS 

Figure Page 

Frontispiece: The Brown Bear in the Aleutians xiv 

1. Aleutian Islands 4 

2. Semichi Islands 6 

3. Sketch elevation of Agattu Island 6 

4. Sketch elevation of Buldir Island 6 

5. Sketch elevation of several Aleutian Islands 6 

6. Sketch elevation of three Aleutian Islands 7 

7. Little Sitkin Island 7 

8. Sketch elevation of Rat Island 7 

9. West end of Rat Island 7 

10. West end of Rat Island 8 

11. Southeast end of Rat Island 8 

12. Sketch elevation of Semisopochnoi Island 8 

13. Sketch elevation of West Unalaga Island 8 

14. Sketch elevation of Ilak Island 9 

15. Sketch elevation of Kavalga Island 9 

16. Sadatanak I^^land 9 

17. Sag-chudak Island 9 

18. Sketch elevation of Bobrof Island 9 

19. Anag-aksik Island 9 

20. Sketch elevations of Kasatochi Island and Koniuji Island 9 

21. Seg:uam Island 10 

22. Sketch elevation of Ananiuliak Island 10 

23. Mounds on Kavalga Island 26 

24. Red-faced cormorant 57 

25. Rough-legged hawk 108 

26. Black oystercatcher 131 

27. Aleutian rock sandpiper 143 

28. Least sandpiper 148 

29. Least sandpipers 149 

30. Glaucous-wing-ed g'ulls 166 

31. Black-legged kittiwakes 176 

32. Colony of Pallas's thick-billed murres on Bogoslof Island 184 

33. Pallas's thick-billed murres 185 

34. Kittlitz's murrelet 189 

35. Ancient murrelet 190 

36. Crested auklets 196 

37. Least auklets 197 

38. Least auklet 199 

39. Horned puffins 203 

40. Tufted puffins 204 

41. Aleutian song- sparrow 255 

42. Sea otter 279 

43. Blue fox 294 



NVERTEBRATES AND FISHES COLLECTED IN THE 
ALEUTIANS, 1936-38, by Victor B. Scheffer 

Page 

Introduction 365 

Marine algae 367 

Marine invertebrates 370 

Sponges 370 

Coelenterates 370 

Hydroids 370 

Jellyfishes 370 

Flatworms 371 

RoundwoiTns 371 

Nemertean worms 371 

Brachiopods 371 

Annelid worms 371 

Echinoderms 372 

Brittle stars 372 

Starfishes 372 

Sea urchins 373 

Sea cucumbers 375 

Crustaceans 375 

Copepods 375 

Barnacles 376 

Amphipods 377 

Isopods 379 

Shrimps 379 

Hermit crabs 379 

Anomuran crabs 381 

Other crabs 381 

Mollusks 381 

Bivalves 381 

Snails and sea slugs 383 

Chitons 385 

Devilfishes 386 

Fresh-water invertebrates 387 

Crustaceans 391 

Cladocerans 391 

Copepods 391 

Ostracods 391 

Mollusks 391 

Land invertebrates 392 

Mollusks 392 

Beetles 392 

Bird lice 393 

Diptera 393 

Spiders 393 

Fishes 395 

Literature cited 406 



ILLUSTRATIONS 

FiRure Pagf 

1. Fucus, a brown seaweed 368 

2. Calcareous algae of the Lithothamnion group 368 

3. The 5-rayed starfish Asterias amurensis 373 ' 

4. Twenty-rayed starfish, Pycnopodia helianthoides 374 

5. Green sea urchin Strongylocentrotiis drobachiensis 375 

6. Rock barnacles, Balanns sp 376 

7. Two species of barnacles 377 

8. Parasitic amphipod Paracyamus boopis 378 

9. Common crab. Cancer tnagister 380 

10. King crab, Paralithodes sp 380 

11. Mussels, Mytilus edulis 382 

12. Limpets, Acynaea sp 384 

13. Periwinkles, Littorina sp 385 

14. Fresh-water pool, type 1 (small and clear) 387 

15. Fresh-water pool, type 2 (small and weedy) 388 

16. Fresh-water pool, type 3 (large and barren) 389 

17. Alaska cod, Gadiis macrocephaltis 396 

18. Red sculpin, Hemilepidotns hemilepidotus 397 

19. Irish lord, Hemilepidotus jordani 398 

20. Halibut, Hippoglossus stenolepis 399 

21. Pogie, Lebius superciliosus 400 

22. Pink salmon, Oncorhynchns gorbuscha 401 

23. Native boy netting sockeye salmon 402 

24. Red or sockeye salmon, Oyicorhynchiis nerka 402 

25. Atka mackerel, Pleurogrammus monopterygixis 403 

26. Silver hake, Tneragra chalcogranuna 404 



Foreword 



This report is based on a biological survey of most of the 
Aleutian Islands and the Alaska Peninsula in 1936 and 1937. 
The report was largely prepared soon after the survey, but 
for various reasons it has not been practical to publish it until 
now. Even in manuscript form, this material has been con- 
sulted frequently, and it is issued now in the North American 
Fauna series so as to make more accessible information on one 
of North America's most significant biogeographic regions. 

While the report was being readied for publication, the 
fifth edition of the Check-List of North American Birds ap- 
peared (American Ornithologists' Union 1957). Throughout 
the report, scientific and common names of birds have been 
made to conform to the new Check-List, but generally refer- 
ences to "the A. 0. U. Check-List," without specification, are 
to the fourth (1931) edition. Scientific names of mammals 
have been made to conform in general to the List of North 
American Recent Mammals (Miller and Kellogg 1955) ; com- 
mon names of mammals for the most part follow Hall (1957) . 
The Pinnipeds conform to the nomenclature of Scheffer 
(1958). 

No attempt has been made to include references to all 
recent publications on the Aleutian and Alaskan fauna ; 
references included are those from which data were obtained 
for this report. 

0. J. MURIE 

May 1959 




The Broivyi Bear in the Aleutians. Carlisle Island rising above the fog 



FAUNA OF THE ALEUTIAN ISLANDS 
AND ALASKA PENINSULA 

By Olaus J. Murie, Biologist 



Introduction 



The Aleutian Islands, treeless, fog-bound, and volcanic, extend 
westward from the tip of the Alaska Peninsula for about 1,100 
miles to Attu, which is less than 600 miles from the Kamchatka 
Peninsula of Asia. The Aleutian Islands Wildlife Reserva- 
tion, now a National Wildlife Refuge, was established on this 
chain in 1913. This reservation embraces the islands of the Aleu- 
tian chain between the North Pacific Ocean and the Bering Sea. 
These islands were set apart as a preserve and breeding ground 
for native birds, for the propagation of reindeer and fur-bearing 
animals, and for the encouragement and development of fisheries. 

In 1920, the United States Bureau of Biological Survey ^ was 
given the responsibility of enforcement of the Alaska fur laws 
and administration of the blue-fox industry in the Aleutians. As 
time went on, it became apparent that proper supervision of this 
important wildlife refuge would necessitate an extensive inven- 
tory of the resources of these islands. 

In 1936, assisted by Cecil S. Williams of the Bureau of Biologi- 
cal Survey, I was assigned to make the necessary investiga- 
tions. The motorship Brown Bear was placed at our disposal, 
and H. Douglas Gray and Homer Jewell, both of the Alaska Game 
Commission, joined us at Juneau. A second season was required 
for the work, and, in 1937, Victor B. Scheffer, John H. Steenis, 
H. Douglas Gray, and I made up the scientific party. During 
these two seasons we visited every Aleutian island of any size, as 
well as many islands south of the Alaska Peninsula and several 
points on the Peninsula, including Bristol Bay and the Nelson 
Island region of the Bering Sea coast. In 1938, Scheffer returned 



1 Now a part of the U. S. Fish and Wildlife Service. 



2 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

with the Brown Beai- for another season's work. He made limitec 
studies of the lesser forms of animal life that inhabit the sub 
arctic waters of the North Pacific Ocean and the Bering Sea an<( 
those that occupy the shores and slopes of the islands. 

Our work, and the work of Scheffer, expanded upon informai 
tion obtained by biologists who visited the area late in the lOth 
century and early in the 20th century. In 1902, W. H. Osgood 
for the Bureau of Biological Survey, conducted an expeditioi 
to the base of the Alaska Peninsula. Results of his field worl 
were published as "A Biological Reconnaissance of the Base o: 
the Alaska Peninsula" (North American Fauna Series No. 24 
1904). In 1911, Alexander Wetmore and A. C. Bent investigatec 
the western end of the Alaska Peninsula and some of the Aleutiai> 
Islands (Wetmore's field report was never published). In thd 
summer of 1925, assisted by Fur Warden Donald H. Stevensonc 
I was assigned to field work at the western end of the Alaska 
Peninsula. Additional investigators who visited the Aleutians 
include Lucien M. Turner and William H. Dall (in the 19th cen 
tury), and Ira N. Gabrielson (in the 20th century). 

In the present report, references are made to all individuals 
who are known to have contributed to the knowledge of the fauns 
of the Aleutian Islands. These individual contributions total intc 
a considerable volume of data that have been of inestimable helpi 
in evaluating the Aleutian fauna. In view of this assemblage oli 
data, and for a better understanding of the fauna of this pari* 
of Alaska, the present report embraces all of the Alaska Peninsula 
and the Aleutian Islands. 

In compiling the material presented here, and in gathering the 
field data, I am indebted to my colleagues in the field on all three^ 
expeditions — 1925, 1936, and 1937. These colleagues, already 
mentioned — Stevenson, Williams, Scheffer, Steenis, Gray, and 
Jewell — are men whose zeal for research and loyalty to the joint 
undertaking must ensure success of an expedition. John Selle- 
vold, veteran seaman and captain of the Broion Bear, went beyond 
the requirements of his duty to help us in many ways. 

John W. Aldrich and Allen J. Duvall, both of the U. S. Fish 
and Wildlife Service, have been especially helpful with advice , 
and assistance in working with specimens. Herbert Friedmann,ij 
of the National Museum, has also helped considerably, and Ira N 
Gabrielson, who has made many trips to the Aleutian district,] 
has been especially generous with his field notes. Many others,^ 
both in Washington and in the field, assisted in many ways. 

Also, I must pay tribute to those original inhabitants of tnej 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 3 

A.leutian3, the Aleuts, who as a race have suffered many vicissi- 
:udes through earlier contacts with white men. Those with whom 
i-ve associated were eager to help with information. It is with 
special affection that I recall the friendly cooperation of Mike 
Hodikoff, Chief of Attu village, who was ready to do anything 
to further the work of our expeditions and to add to our knowl- 
?dge. He, with his village, was captured by Japanese invasion 
forces during World War II ; there is no knowledge of his fate. 

GEOGRAPHY AND GEOLOGY 

The Alaska Peninsula and Aleutian Islands (see fig. 1) form 
a great arc that swings across the northern seas for about 1,500 
miles, almost to Siberia. The Aleutian chain alone is about 1,100 
miles long. This arc, together with the Commander Islands, forms 
a barrier that separates the Bering Sea from the North Pacific 
Ocean. The Alaska Peninsula extends southwestward from 
about latitude 59° N., and Amatignak Island, the southernmost 
of the Aleutians, lies nearly as far south as latitude 51° N. — the 
same latitude as the north end of Vancouver Island. 

The north shore of the Alaska Peninsula shelves off gradually 
into the shallow waters of Bering Sea, forming a low coastal 
plain with a comparatively even coastline. However, farther in- 
land the land rises to the rugged volcanic Aleutian Range, which 
runs the length of the Peninsula, and, on the south side, breaks 
off into the deeper water of the North Pacific. Accordingly, the 
south shore is irregular and rugged with bays and headlands and 
offshore rocks and is fringed by offshore islands — notably the 
Kodiak-Afognak, Semidi, Shumagin, and Sanak Island groups, 
as well as a number of smaller ones. 

The eastern Aleutians retain some of the characteristics of the 
Alaska Peninsula. This is most pronounced on Unimak Island, 
which has a low coastal plain, lagoons, and rugged interior moun- 
tains that extend southward to the Pacific Ocean. In fact, Unimak 
Island is separated from the Peninsula by only a narrow strait. 

Numerous eruptions have been recorded since the discovery 
of these islands, and the Aleutian chain proper consists of over 
70 named islands, some small, others large ; Unimak is about 70 
miles long. The chain is irregular and is bordered on the north 
and south sides by deep oceanic troughs. In other words, the 
soutli border of the shallow Bering Sea bottom, which is virtually 
a continental shelf, veers off northwestward so as to leave deep 
waters north of the Aleutian chain. As Stephen R. Capps (1934, 
p. 143) has stated. 



NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 




FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 5 

A line of soundings taken by the fathometer on the Gannet in 1932, ex- 
ending along an irregular course from a point north of Amukta Pass to 
Vttu Island, at varying distances from the intervening islands, shows 
hat north of the islands the 1,000 fathom line lies close to the island 
"estoon and that at a distance of 50 miles or more from them there is a 
■emarkably smooth-floored depression at a depth of 2,000 to 2,200 fathoms. 
The shape of this depression between the islands and the continental mass, 
vhich includes much of Bering Sea, is not known, but it is significant 
.hat the island arc rises as a sharp ridge separating deeps of 2,000 fathoms 
)r more both to the north and south. 

The volcanic nature of this region is well known. Capps (1934, 
D. 142) says. 

Throughout the Alaska Peninsula the volcanoes have broken out through 
)lder sedimentary or igneous rocks, by which they are now flanked. In 
;he Aleutian Islands there are few if any exposures of the basement 
•ocks, and the islands are largely constructional, having been built up 
and above sea level by the accumulation of lavas and volcanic fragmental 
naterial ejected from below. 

Many volcanoes along this remarkable arc are still in an active 
5tate. The eruption of Katmai Volcano, on the Alaska Peninsula, 
n 1912 was one of the great volcanic spectacles of modern times 
(see Griggs 1922). The activities of Bogoslof Island and Mount 
Shishaldin on Unimak Island are well known, and in 1930 there 
tvas an eruption on Gareloi Island. On our visit there in 1937 
\xe examined some of the small craters, from which were issuing 
5team and other gases, and we noted many lava bombs on the 
ower slopes. We found several typical hot thermal springs that 
*vere rimmed with algae. On Kagamil Island, noisy steam jets 
ssued from a rocky bluff, and rumblings could be heard under 
:he boulder beach. After our return from the expedition of 
1937, we learned that there had been an eruption on Yunaska 
Island while we had been exploring other areas. Many of the 
mountains have plumes of steam issuing from the top. Mount 
Cleveland, on Chuginadak Island, erupted in 1944, and Okmok 
and Umnak Islands erupted in 1945. 

As would be expected, most of the islands are mountainous. 
There are a few relatively flat islands, such as Amchitka, Agattu, 
and Semichi. However, there is a low mountain range along 
3ne side of Agattu, and there is a small mountain at one end of 
A.laid Island, in the Semichis. Most of the larger islands have 
lakes and streams, and several, such as Amchitka, Agattu, and 
the Semichis, are dotted with lakes. In keeping with their 
volcanic origin, some of these islands liave notable lava beds that 
furnish nesting crevices for petrels and auklets, as on Amukta 
and Gareloi. Other islands, notably Ogliuga and part of Kavalga, 



6 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

have been covered with volcanic ash in recent times. 'The shore- j 
lines are irregular, with offshore islets, rocks, and undersurfac€i 
reefs, and there are boulder beaches, sandy beaches, and abrupt 
cliffs in great variety. 

The accompanying field sketches, (see figs. 2-22) showing a few 
of the islands in profile, suggest the variety of configuration. 




Figure 2. — Semichi Islands from mountain on Alaid Island (June 1937). 
Note that a narrow spit connects Alaid, in foreground, with the middle* 
island ; Shemya, the easternmost, is in the distance. 



Figure 3. — Sketch elevation of Agattu Island, seen from west end of Alaid 
Island, looking southwesterly. 




Figure 4. — Sketch elevation of Buldir Island, looking southeast. 



AMCHITKA 
(WEST end) 



KISKA 
(N. END) 



CHUGUL DAVIDOF 



LITTLE 
SITKIN 



Figure 5. — Sketch elevation of several Aleutian Islands, looking west. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 7 




'IGURE 6.— Sketch elevation of three Aleutian Islands from Gunner Cove on 
Rat Island, looking northerly. 




'IGURE 7. — Little Sitkin Island from Gunner Cove on Rat Island, looking 

northeasterly. 



'IGURE 8. — Sketch elevation of Rat Island from southeast end of Khvostof 
Island, looking southerly. 







'IGURE 9. — West end of Rat Island (July 1937). Kiska Island in distance. 



8 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 




Figure 10. — West end of Rat Island (July 1937) from beach on south shore 




Figure 11. — Southeast end of Rat Island (June 1937). 



Figure 12. — Sketch elevation of Semisopochnoi Island seen from west end of 
Amchitka, looking northeasterly. Low fog bank on horizon. 



.^ m. 



Figure 13. — Sketch elevation of West Unalaga Island, looking westward. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 9 



Figure 14. — Sketch elevation of Ilak Island, looking- southerly. 



'IGURE 15. — Sketch elevation of Kavalfi:a Island from West Unalg-a Island, 

looking easterly. 



Figure 16. — Sadatanak Island looking easterly. 



^iSSisS^^JMBSm^lllim^^^^^^ 



Figure 17. — Sagchudak Island looking easterly. 




Figure 18. — Sketch elevation of Bobrof Island, looking southwesterly. 






Figure 19. — Anagaksik Island, looking southeasterly. 




'igure 20.— Sketch elevations of Kasatochi Island and Koniuji Island, 

looking west. 



10 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 







ii'"Hiiiiii 



u , ' . . ii M i "" iMMi '."',": '.'ij|»u'^> ...1 M..;,7' .J^i>)ffm> 



Figure 21. — Seguam Island, looking south-southwest. 



Figure 22. — Sketch elevation of Ananiuliak Island, looking southerly. Umn? 

Island in background. 



CLIMATE 

The Aleutian Islands and Alaska Peninsula are south of tl 
severe low winter temperatures of interior and northern Alask, 
and the surrounding waters are generally free of sea ice. ^^ 
characterize winter conditions briefly: temperatures go well b 
low freezing, fresh-water ponds freeze over at times, and sno 
sometimes piles knee deep. But the snow is likely to be w 
and slushy, and there will be some bare ground. At high( 
elevations, hjwever, snow is heavier, and the higher mountaii 
are snowcapped in winter. 

A few temperature records from Turner (1886), with notes C( 
clear days, are of interest: 



Month 



Temperature (°F.) 



Mean 



Maximum 



Minimum 



Number of- 



Clear days 



UN ALASKA, 1878-79 



September 
October. __ 
November 
December. 
January _ _ 
Februarj-. 

March 

April 

May 

June 

July 

August 



48.02 
40.77 
33,50 
35.12 
33.97 
29.25 
32.16 
33.07 



55 

49 
48 
45 
48 
44 
49 
52 



ATKA, 1879 



39 . 90 
42.08 
48.96 
50.31 



65 
64 
65 
69 



36 
26 
21 
19 
20 
7 
15 
21 



30 
30 
38 
45 



Fair day 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 11 

ATTLl, 18S0-81 



July 

August 

September 

October 

November 
December. 
January. - 
Fel)ruary. 

March 

April 

Mav 



52.35 


CO 


51 5f) 


6() 


47.75 


58 


41.12 


49 


35 45 


4(i 


33 i)l 


44 


31.17 


42 


31.!)5 


41 


29.02 


41 


36.70 


52 


39.55 


49 



42 
38 
36 
30 
25 
22 
17 
17 
11 
20 
31 



Sutton and Wilson (1946) observed birds on Attu Island from 
ebruary 20 to March 18, 1945. They report, 

he air temperatures at sea level did not vary much from freezing as 
rule. During the daylight hours it sank somewhat below 32° F. on 20 
' the 27 days, climbed as high as 38" during the day on March 4, sank 
5 low as 15° during the night on March 15, and averaged 31°. On March 
i the greatest temperature variation (15° to 31°), as well as the lowest 
mperature, was recorded. The general aspect was wintry: the sky over- 
ist, the wind raw, the sea turbulent. Highlands and lowlands alike were 
)vered with snow. Along the shore, tufts of rank grass and coarse stalks 
' wild parsnip protruded from the drifts, and boulders, turfy mounds and 
irrow gray beaches were always bare. Elsewhere, save for an occasional 
iff or exposed slope, everything was white. 

A striking feature of the Aleutian climate is the prevalence 
f foggj' or cloudy weather, the abundance of rain in summer, 
nd the frequent violent winds that arise suddenly and un- 
^pectedly. On western Alaska Peninsula, in 1925, we built a 
indbreak of alder brush to protect our tent, and, on the beach, 
^ht gravel occasionally would be blown into our faces. Briefly, 
len, one might say that although the temperature is mild — 
9ither very low in winter nor very high in summer — there 

a minimum of sunshine and a maximum of fog, rain, and 
orm. 



ENVIRONMENT AND BIOTIC DISTRIBUTION 

The Alaska Peninsula and Aleutian Islands, stretching as a 
nd bridge between two continents, present a most interesting 
stribution of plant and animal life. There are, of course, a 
imber of physical facts that bear on the distribution of animals 
id plants — including the location of the area with relation to 
at of other significant areas, the geologic history, the physio- 
'aphic conformation of the land, the ocean currents, and the 
mperature, humidity, and other climatic influences. 



12 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

GEOGRAPHIC AND GEOLOGIC INFLUENCES 

The Aleutian district lies within the Boreal region, and it mj 
be identified as the southern fringe of this great circumpoli 
area throughout which life has so much in common. It 
significant that the Alaska Peninsula and Aleutian Islands li 
in a sense, almost isolated from the mainland of Alaska ai 
extend westward a tremendous distance toward Kamchatk 
thus, in some respects, serving as a "bridge" between Asia ai 
North America. It is also significant that they are near tl 
other intercontinental bridge at Bering Strait, which is recognizi 
as having an important influence on our biota. 

The Aleutian Islands and Alaska Peninsula are of comparative 
recent geologic origin. Volcanic activity is still prevalent, ai 
changes in the surface of the land are still taking place. F 
this reason, and for reasons mentioned later, the area has 
new and changing environment that has not yet been ful 
occupied by flora or fauna to the extent of its potential capacit 
Thus, the area presents an opportunity to see immigration st 
taking place and to note the changes imposed on the newcome 
by an unusual environment. 

As the following sections show, the Aleutian biota is drawii 
its members from several directions. As would be expected, sor 
have come directly from Asia ; some have come from the nor 
on the Alaskan side; others have come from the southeast aloi 
the Pacific coast; and still others are part of the fauna th 
appears to have developed in the Bering Sea region — an ar 
roughly bounded by Siberia, mainland Alaska, and the Aleuti; 
Islands. Many others are drawn from biotic populations th 
at present are so widely distributed in the Palaearctic regi( 
that it is impossible to judge the direction from which th 
entered the Aleutian area. Following, are some of these grou 
that have contributed to the Aleutian biota : 

ASIATIC IMMIGRANTS 

BIRDS 

SCIENTIFIC NAME COMMON NAME 

Branta nigricans Black br; 

Mareca penelope European widge 

Anas crecca ssp Common td 

Haliaetus albicilla White-tailed sea eajli 

Haliaetus pclagicus Steller's sea eaj 1 

Faico rusticolus uralensis Asiatic gyrfalc \ 

Charadriiis dubiiis curonicus Little ringed plo\ i 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 13 

haradrius semipabnatus Semipalmated plover 

luvialis dominica, fulva American golden plover 

renaria interpres interpres Ruddy turnstone 

ringa glareola Wood sandpiper 

rolia acuminata Sharp-tailed sandpiper 

ams hyperboretts hyperboreus Glaucous gull 

arus schistisagiis Slaty-backed gull 

anis argentatus vegae Herring gull 

arus ridibundus sibiricus Black-headed gull 

terna aleutica Aleutian tern 

ethia pygmaea Whiskered auklet 

uculus saturatus hursfieldi Oriental cuckoo 

'iscinia calliope camtschatkensis Siberian rubythroat 

roglodytes troglodytes ssp. (in part) Winter wrens 

otacilla alba lugens White wagtail 

mbenza i-ustica latifascia Rustic bunting 

MAMMALS 

SCIENTIFIC NAME COMMON NAME 

lopex lagojnis Blue fox 

rsiis arctos gyas Brown bear 

rsus arctos middendorffi Brown bear 

Some of these are only occasional visitors, such as the two 
igles mentioned, and Larus schistisagus (slaty-backed gull), 
arus ridibundus sibiricus (black-headed gull), Cuculus saturatus 
orsfieldi (oriental cockoo), and some others. Some have become 
;tablished in the Aleutians, such as Anas crecca (common teal), 
ethia pygmaea (whiskered auklet), and Sterna aleutica (Aleu- 
an tern), and Alopex lagopus (blue fox). Others have reached 
le Alaskan coast in general, including the Aleutian district, but 
>t necessarily by the Aleutian route, such as Falco nisticolus 
''alensis (Asiatic gyrfalcon), Pluvialis dominica fulva (Ameri- 
m golden plover), and the big brown bears. Some, such as 
haradrius seniipalmatus (semipalmated plover) and Branta 
igricans (black brant), have extended eastward considerably 
?yond the Alaskan Peninsula but show greater affinity with 
siatic populations than with those farther east in North Ameri- 
i. In the case of the winter wrens. Troglodytes troglodytes, the 
•igin appears to have been from Asia and from the southeast. 
if course, the bears came by the more remote northern route. 
Plants, too, have begun the long traverse over from Asia. 
L the case of plants which occur widely on both sides of Bering 
Lrait, and which have become established all the way through 
je Aleutian chain, it is difficult to know the direction from 
hich their immigration took place. There are some plants that, 
cording to Hulten's distribution maps (1937a), have obtained 



14 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

a foothold in the western Aleutians but have not been observed 
the east, even though some of them also occur on the Alaskj 
mainland. Concerning the plant distribution, Hulten (1937a, 
44) stated, 

If Commander Islands and westernmost Alaska Penin. are included, as 
this flora, 92 America species reach that area, but 47 of them do not 
further westward than to Umnak. 49 species reach the Aleutians frc 
the west but not other parts of southern Alaska. 40 of them do not rea. 
further eastwards than to the westernmost g'roup of the Aleutians. 

Some of the prominent Asiatic forms that we observed in tl 
westernmost Aleutians are Cirsium kamtschaticum, Veratru 
album oxysepalum, Cacalia aricuhxta, and Sorbus sambucifolii 
These are confined to the Near Islands, though some are thoug 
to have reached as far east as Buldir. 

Hulten says further, "The flora of the middle Aleutians 
very depauperated, probably due to the relatively short tin 
elapsed since the glacial period, when most of their plants we 
exterminated." 

Only the more obvious Asiatic elements are mentioned hen 
Other animal and plant forms probably originated in Siber' 
at a more remote time. 

BERING SEA AVIFAUNA 

The following birds represent a group largely confined to t; 
coastal parts of Bering Sea, although some of them range farth 
north or south. They appear to be characteristic of all shor 

of the Bering Sea, instead of the Siberian side exclusively. 

I 

SCIENTIFIC NAME COMMON NAME 

Phalacrocorax pelagiciis pelagicus Pelagic cormorai 

Phalacrocorax urile Red-faced cormoral 

Branta canadensis minima Canada goc^ 

Philacte eanagica Emperor goc 

Anas crecca nimia Common t( 

Polysticta stelleri Steller's eic 

Arenaria melanocephala Black turnstc 

Numenius tahitiensis Bristle-thighed curl 

Erolia ptilocnemis ssp Rock sandpii 

Limosa lapponica baiieri Bar- tailed god\ 

Rissa tndactyla pollicaris Black-legged kittiwai 

Rissa brevirostris Red-legged kittiwai 

Xema sabini woznesenskii Sabine's g< 

Uria lomvia arra Thick-billed mui( 

Brachyramphus brevirostre Kittlitz's murre( 

Cyclorrhynchus psittacula Parakeet auki 

Aethia crwtatella, , , Crested auW 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 15 

ethia pusilla Least auklet 

'ratercula coryiiculata Horned puffin 

'hylloscopus borealis kennicotti Arctic warbler 

lotacilla fiava tschutschensis Yellow wagtail 

lectrophenux hyperhoreus McKay's bunting 

AUNA OF WIDER NORTHERN DISTRIBUTION 

BIRDS 

SCIENTIFIC NAME COMMON NAME 

avia arctica pacifica Arctic loon 

avia stellata Red-throated loon 

lor columbianus Whistling swan 

ranta canadensis leucopareia Canada goose 

nse7- albifroyis frontalis White-fronted goose 

langula hyemalis Oldsquaw 

omateria inollissima v. nigra Common eider 

uteo lagopus s-johannis Rough-legged hawk 

alcQ rusticolus obsoleteus Gyrfalcon 

agopus lagopus ssp Willow ptarmigan 

agopus mutus ssp Rock ptarmigan 

rolia alpina pacifica Dunlin 

reunetus sp Sandpipers 

halaropus fulicarius Red phalarope 

obipes lobatus Northern phalarope 

^.ercorarius sp Jaegers 

arus hyperboreus barrovianus Glaucous gull 

tema paradisaea Arctic tern 

%rus atncapillus turneri Black-capped chickadee 

df-us hudsonicus hudsonicus Boreal chickadee 

Urdus ynigratorius migratorius American robin 

ylocichla minima fninima Gray-cheeked thrush 

"'''*^''' sp Redpolls 

inco hyemalis hyemalis Slate-colored junco 

isserella iliaca zaboria jTqx sparrow 

ilcarius lapponicus alascensis Lapland longspur 

ectrophenax nivalis ssp Snow bunting 

MAMMALS 

SCIENTIFIC NAME COMMON NAME 

>rex tundrensis Tundra saddle-backed shrew 

^^""« sp Ground squirrels 

icrostonyx sp Collared lemmings 

icrotus oeconomus ssp Meadow mice 

•pus othiis poadromus Arctic hare 

mgifer arcticus grayiti Barren ground caribou 

'Iphinaptenis leiicas Beluga 

These are some of the more northern birds and mammals whose 
stribution with relation to the Alaska Peninsula is such that 



16 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

they probably immigrated southward or southwestward to t 
Aleutian district. There are, of course, a great many oth€ 
of northerly distribution whose general range is such that t 
route of the population movement is uncertain. In the gro 
here listed it will be seen that the Aleutian district has dra^ 
heavily from the fauna that characterizes the northern porti 
of the continent from northern Alaska across to Hudson B£ 
Lemmings, Arctic hares, the jaegers, Sabine's gull, and othe 
came straight down the Bering Sea coastal strip. 

It will be noted that not all of the birds just listed actua. 
nest in the Aleutian district. 



SOUTHERN AND SOUTHEASTERN AVIFAUNA 

SCIENTIFIC NAME COMMON NAME 

Phalacrocorax auritus cincinatus Double-crested cormorf 

Olor buccinator Trumpeter sv( 

Anas strepera Gadw 

Larus glaucescens Glaucous-winged g\ 

Brachyramphus fnarmoratum marmoratuyn Marbled murre 

Synthliboramphus antiquu^n Ancient murrfi 

Ptychoramphus aleutica Cassin's auW 

Cerorhinca monocerata Rhinoceros auW 

Megaco-yle alcyon caurina Belted kingfisi 

Troglodytes troglodytes ssp. (in part) Winter wr 

Ixoreus naevius Varied thri 

Vermivora celata lutescens Orange-crowned warfc 

Pinicola enucleator flamnuila Pine grosbc 

Leucosticte tephrocotis ssp Gray-crowned rosy fii 

Loxia curvirostris sitkensis Red cross 

Passerculns sandwiohensis ssp Savannah span' 

Passerella iliaca ssp Fox span 

Melospiza melodia ssp Song sparr 

Some of these listings give us a clear demonstration of 1 1 
route of influx into the Aleutian district, by way of closely relal i 
subspecies in a series extending along the coastal strip of southe : 
and southeastern Alaska. Such examples are the song sparrov 
fox sparrows, and winter wrens in particular. The fox sparro ' 
present an interesting distributional picture. It is the dark coas 
unalaschcensis group that has worked along the coast and fui 
occupied the suitable habitats as far as the eastern Aleutia i 
But the bright-colored eastern type has come down from 11 
northeast and, at the base of Alaska Peninsula, this type has m^ ( 
contact with the coastal forms. 

Naturally, there could be error in the interpretation of faun 
immigration just cited, because complexities may have interverE 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 17 

nee the territory in question was "opened for settlement" ; how- 
er, the conclusions submitted are based on strong probability at 
ast. 

It will be seen that the coastal mountain masses of southern 
laska and the Alaska Range form a barrier. Although this is 
»t an absolute barrier, presumably it is enough of an obstacle 
at the way of least resistance would be north and west along 
e coast for some species. Similarly, there is an easy avenue 
uthward along the open Bering Sea coast for tundra-loving 
rms. And the Aleutian chain, reaching out close to Siberia, 
an inviting route, 

nIvironmental influence 

There are some striking environmental influences operative in 
e Aleutian district. We know, of course, that humid regions 
id to produce dark pigmentation, and this fact holds true for 
is area. The rosy finches reach their darker hues in the Aleu- 
Ln area, with the darkest in the Pribilofs, The fox sparrows 
ow the same tendency, exceeded in dark tones only by the 
pulations of the excessively humid Pacific rain-forest zone that 
tends from the coast and islands of southeastern Alaska, south- 
ird to the northwest coast of the United States, Except for 
3 aberrant yellow types in the middle Aleutians, the darkest 
:k ptarmigans are found in the Aleutian area, especially on 
tu and the Commanders. Here, parasitic jaegers are, and the 
ctic foxes are, almost entirely in the dark-color phase. In 
imitive times, silver foxes were unusually plentiful somewhere 
this district, judging by the cargoes of the first Russian traders, 
e lemmings, Dicrostonyx, of Unalaska and Umnak do not ac- 
ire a white pelage in winter. 

This is also a region of giantism. Note the huge size of the 
ig sparrows, Savannah sparrows, and rosy finches, which, as 
lera, reach their greatest size in the Aleutians and Commander 
ands. Here, the Aleutian winter wrens, as a group, have de- 
oped unusually long bills. Here, too, we may include the Alaska 
)wn bear, which achieves its greatest size on the Alaska Penin- 
a, Unimak Island, and Kodiak Island. 

Marine biologists have found that in many instances the in- 
•tebrate subspecies in the northern Pacific waters, and even 
ther north, are strikingly larger than forms of the same 
cies farther to the south. This invites interesting speculation, 
pointed out later, the Aleutian waters are unusually rich in 
nkton, and there is an abundant and varied marine inverte- 



18 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

brate fauna. An exceptionally nutritious marine food sour 
should influence the vigor and size of the terrestrial vertebrat 
of that region. 

The song sparrow's habitat in the Aleutians is the beach, ai ( 
it is probable that its food is largely of marine origin — the smi, 
beach crustaceans, for example. Some other land birds, such ;i 
winter wrens and pipits, feed to some extent on the beach. Tl! 
blue foxes feed chiefly on marine life. 

The case of the Alaska brown bears is not so clear, though f(t 
a part of the summer they comb the beaches and live extensive 
on salmon, which are nourished in the sea. One wonders, alsi 
if a certain type of food may, with other factors, help to encouraj. 
melanism (as in the jaeger), or darker shades of color, as in son j 
of the other birds. It is generally accepted that a humid habit i 
produces dark coloration. It is not certain that this tendenci/ 
as well as melanism, is encouraged by rich food. 

This is, of course, pure speculation, yet the significance of ; 
food chain from the sea to the higher vertebrates on the adjacei 
land may be worthy of earnest study. There are many birds th i 
have not responded to environmental influence. The Aleutian soii] 
sparrow has not developed dark pigmentation to an unusual d^ 
gree. The northern form of the fork-tailed petrel, though avera; : 
ing larger in the Aleutians, apparently is paler than those i 
southeastern Alaska. Also, it must be considered that the ii 
terior Alaska and Yukon caribou, as well as the Alaskan moos; 
which have no direct connection with the sea, are the largest (j 
this continent. But these examples suggest that there is somE 
thing in the environment — favorable food, humidity, or otb 
stimuli— that tends to produce dark pigmentation and large si2 ; 
This is an important challenge to future investigation ai < 
understanding. 

ECOLOGICAL CLASSIFICATION 

By the usual standards of life-zone allocation, the Alaska Pej 
insula and Aleutian Islands would fall chiefly in the Arctic Zors 
A part of the Kodiak-Af ognak Island group supports tree growt ' 
and forests encroach on the base of Alaska Peninsula to tii 
vicinity of Mount Katmai. Therefore, these locations would ma: 
the limit of the Hudsonian Zone. However, we find the life-zoM 
classification here to be far from simple. There are probably 
number of physiographic and oceanic reasons for this situatiof 

There are serious difficulties in the interpretation of life zonk 
in the Aleutians that should be considered. The lack of tre4, 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 19 

resumably would indicate some form of Arctic or Alpine Life 
one. So far as latitude is concerned, the southernmost island of 
le Aleutian chain, Amatignak, lies not far north of 51° N,, 
^hich is the latitude of heavily timbered, coastal British Columbia 
1 the vicinity of Vancouver Island ; however, the treeless Aleu- 
ians lie hundreds of miles south of the tree limit in the Brooks 
lange of interior Alaska. Some of the lowest temperatures in 
ilaska are recorded from the timbered interior, while the 
emperatures in the Aleutians are uniformly higher in winter, 
nd the adjacent seas are not frozen over. From the standpoint 
f vegetation growth, summer tempoi^tures are probably of 
reater significance than winter temperatures, and probably do 
jot show so great a variation. Certainly the temperatures aver- 
age much lower and have a lower maximum in summer than 
emperatures found in the forested continental areas. 

Wind is another factor that generally accompanies treelessness 
t high altitudes and latitudes. There is a treeless coastal strip 
I ordering the Bering Sea, with few interruptions, from Alaska 
peninsula to Bering Strait, continuing around to the treeless 
i^rctic coast. This coastal area is characterized by strong winds, 
is contrasted with the comparative stillness of the interior. We 
I now the effect of wind on tree growth at timberline in moun- 
jains. In the Aleutians, I found many instances where the wind 
lad scoured out the soil, exposing the roots of such ground- 
jiUgging plants as crowberry and dwarf willow. If wind is one 
f the factors that establish the edge of forests, it is operative to 
j,n unusual degree in the Aleutians. 

! Forest growth is another important factor to be considered in 
(he Aleutian district. Attention is invited to the series of pub- 
ications on Alaskan flora by Robert F. Griggs (see bibliog- 
aphy) — particularly his 1934 report on the edge of the forest, 
'n which he has assembled numerous data to show that the edge 
if the forest has been advancing in Alaska. This was particu- 
larly evident on Kodiak Island and in the Katmai region, where 
l)r. Griggs worked intensively. According to Griggs' studies, we 
nay reason that, since the last glaciation, climate or a combina- 
ion of other factors has been gradually improving the area 
oward suitability for forest growth. The forest, however, has 
lot been able to migrate fast enough to keep pace with favorable 
limatic conditions and has not reached its potential limit. 

Where, then, is the limit of the potential climax forest growth? 
Vt the end of the Alaska Peninsula? Farther west? On Mer- 
•iam's life zone map, the Hudsonian Zone is shown extending the 



20 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

length of Alaska Peninsula. Spruce trees that were planted c 
Unalaska Island grew, but did not spread. It may be significai 
that tall willows, alders, and tall salmonberry have extendt^ 
westward as far as Unimak Island in heavy thicket form. Her 
salmonberry thickets are similar to those seen in southeastei 
7\.laska. These facts may be indicative of a territory ripe for 
forest. And such an advance line, based on climatic limitatioi 
rather than on the present position of the forest edge, may 1: 
considered to be the boundary of the Hudsonian Life Zone. T? 
biome concept, to rely strictly on the climax end product, woul: 
have the same difficulty here, and published maps of the Tundri 
Biome and the Tundra-Coniferous Forest Ecotone for this are: 
would simply substitute these terms for Arctic and Hudsonia: 
Zones. There is the same potential boundary difficulty. 

Granted that in the Boreal Zone, at least, tree growth is d ■ 
rectly affected by the climatic factors usually associated with th : 
life-zone concept, to what extent is the rest of the biota affecte ] 
by the same influences? To what extent is it influenced by the 
mere presence of trees? It is reasonable to believe that the wood; 
plants that comprise the understory of the Alaskan forest an, 
to a large extent, dependent on association with trees. Som: 
forms, such as blueberries, often extend from open country inti 
scattered forest. But there is a plant association that coincide! 
with forest growth. 

Similarly, there is a fauna that has become specialized fo 
forest habitat — woodpeckers, certain grouse, certain warblers 
jays, squirrels, black bear, and many others. These appear to bi 
limited by the mere presence of trees. There is good reason tl 
believe that wapiti and other deer would have a much mon 
northern distribution if it were not for the physical barrier o< 
deep snow in winter. On the other hand, the red-backed mousq 
the hermit thrush, and the chickadee have inhabited the lengtl 
of Alaska Peninsula. It is possible to assume that these mobil 
woodland forms simply would not wait for the slow-moving fori 
est and thus have adapted themselves to more-open habitat! 
than is nonnal for the species. Also, this would imply a lessi 
specialized response to habitat than some of the other forest 
species, as well as a greater sensitivity to direct climatic stimuli 
Birds and mammals are more or less adaptive and vary between 
wide extremes in tolerance of adverse elements in their environ i 
ment. However, there is a strong tendency for the majority 0:1 
any population to be associated with the distribution of certair: 
major vegetation types. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 21 

One cannot escape the conviction that if we grant a certain 
degree of cHmatic influence on distribution of vegetation to cause 
it to fall into broad life zones, many of the birds and mammals 
that have become adapted to vegetation types will also tend to 
fall into these same life zones. These birds and mammals may be 
affected to a lesser extent by the life-zone climatic influences than 
by the indirect eff"ects of these influences — the vegetation type of 
the habitat. 

It should be kept in mind that, in boreal regions, biotic units 
are not so clearly defined as in desert or semidesert areas. Griggs 
(1934c), writing on Arctic vegetation, says, 

In short every feature of Arctic vegetation, the anomalies in the geographi- 
cal distribution of arctic species, the occurrence of many species in all 
sorts of habitats, and their apparent indifference to the diverse conditions 
thereof, the lack of definiteness to the composition of the plant cover 
in any particular habitat, the physical instability of the ground itself, the 
general ruderal character of arctic vegetation, the lai'ge number of our 
weeds which are native to the arctic — all these testify to an instability 
in arctic vegetation very different from the relatively stable plant forma- 
tions of the temperate zone. 

He states further that — 

First, combined with the demonstrated active migration of the Alaskan 
forest into the arctic, it gives definite support to the supposition that vegeta- 
tion there has not yet recovered from the glacial period but is still in process 
of active readjustment. 

This statement is applicable to the fauna as well, especially 
in the Aleutian district. Native rodents have only begun to en- 
croach on the Aleutian Islands. Savannah sparrows have gone 
only part way. Song sparrows have reached Attn, but fox spar- 
rows have gone only as far as Unimak. Foxes had started to 
enter the Aleutian chain from Alaska, as well as from Siberia, 
before man intentionally affected their distribution. 

Minute organisms that thrive unusually well in the cold waters 
of the northern seas have set up a food chain that developed a 
rich marine biota. This accounts for the presence of the fish, 
pinnipeds, whales, and sea otters that once inhabited these wa- 
ters so abundantly. Given such a good supply of food, with an 
abundance of ideal cliffs and lava beds and boulder beaches for 
nesting sites, it is logical that the present swarming seabird 
colonies have assembled in the Aleutians. 

There is much of the Arctic element in the Aleutians. In- 
deed, the Arctic and Alpine merge on these islands. The moun- 
tain-loving rosy finches and the Arctic snow bunting nest 
practically side by side, close to sea level. Alpine vegetation types 



22 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

are not far above the level of the sea. But if we apply the terij 
"Arctic" here, it must denote the "Low Arctic." 

The Aleutian district is unique. For animal life, it combimj 
favorable climatic factors and unusual food resources. It is 
focal point to which animal life has been coming from norti 
and south, and east and west, and it is a melting pot for faunj 
elements from two continents that have not yet reached equilil 
rium. It is necessary to keep in mind the fluid nature of th. 
Aleutian biota in arriving at any system of zonal delineation. 

From a purely descriptive standpoint, the fauna of the Alei 
tian district stands apart, and it may well merit distinction a 
the "Aleutian Fauna." There may be good reason to consider ij 
as a unit of a more comprehensive Bering Sea fauna. I 

1 

VEGETATION 

In 1937, Eric Hulten published (in Stockholm, Sweden) "Flor; 
of the Aleutian Islands and Westernmost Alaska Peninsula wit' 
notes on the flora of Commander Islands." The same author ha^ 
also published "Flora of Alaska and Yukon," in 10 parts, issuer 
from 1941 to 1950. This work covers the botany of the Aleutiai 
district so thoroughly that no detailed account of the vegetatioi 
need be attempted here, except for mention of some prominen 
plant associations and their distribution. 

The first consideration is the distribution of forests. Th 
spruce-forest edge is found midway on Kodiak Island and i] 
the general vicinity of Becharof Lake on Alaska Peninsula. W 
find elements of the flora, as well as some of the birds, converg 
ing on the base of Alaska Peninsula from two directions. Fror 
the east, the Sitka spruce (Picea sitchensis) of southern Alask; 
has made its way to Kodiak Island at the base of Alaska Penin 
sula, out to the region about Becharof Lake, and now it consti 
tutes the principal forest growth in this area. The status of thi 
white spruce (Picea glauca) is less certain, but this interior! 
Alaska tree has come down from the north to at least as far as 
Bristol Bay, near Nushagak, and it may be considered to havi 
barely reached the border of Alaska Peninsula, inland from thi 
coast. The birch (Betula kenaica) is associated with the conif 
erous growth in all this forested area. 

With the exception of this meager forest, in all lowland por 
tions of Alaska Peninsula and Unimak Island, and to some exten' 
as far west as Unalaska, tall vegetation is in the form of shrul: 
thickets — dwarf birch (Betula nana exilis), willow, and alder 
Alder (Alnus crispa sinuata) is particularly prevalent and forms 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 23 

heavy thickets. Hulten described a new foiTn (Alnus crispa 
laciniata) from Kodiak Island, and mentions Alnus incana as 
occurring in the Katmai district. Shrubby salmonberry (Rubus 
spectabilis) is found in suitable places along the Alaska Penin- 
sula, and in the eastern Aleutians it is found as far as Unalaska 
Island. 

Beyond Unalaska, the vegetation is of the low type ; the willows 
are of the dwarf species, close to the ground, and we find no 
appreciable high-shrub growth until at the very end, on Attn 
Island. Attu Island possesses moderate shrubby and tall plants — 
largely those with Siberian affinities. Hulten (1937a) states: 

In the westernmost Aleutians, on Attu I., are found fragments of a 
high-grown vegetation similar to that growing in the upper subalpine 
belt on the open spots between the Alnus shrubs in Kamtchatka and 
along the Kamtchatka west coast. It is largely built up of Asiatic elements, 
which occur only on the westernmost islands, such as Cirsiiim kamtschaticum , 
Veratrum oxysepalum, Cacalia auriculata, Senecio palustris, and Sorbus 
sambucifolia, but it also includes elements occurring all over our area, such 
as Geranhan erianthuni, Streptojnis amplexifolius, Calamagrostis Langsdorffii 
and others. 

Some plant communities may be distinguished readily. Through- 
out all the coastal areas of southwestern Alaska the sandy beaches 
are bordered with a rank growth of wild rye. In the Aleutian 
district, other members of the Ehjyyius arenarius, or wild rye, 
association are Senecio pseudoarnican, (a groundsel), Lathyrus 
maritimus (beach pea), Honckenya peploides, and Mertensia 
ma7'itima (sea bluebell). Within this association we found the 
low-to-ground Honckenya peploides generally pushing out near- 
est the water. In many places the leafy, bulky Senecio pseu- 
doaimica formed vigorous patches that virtually left no room 
for other plants. The Aleuts used the tall, coarse beach rye, 
Elymus arenarius, for weaving the exquisite "Attu" baskets. 

Near the beach, but clinging to rocky sites, is Potentilla villosa, 
a herbaceous cinquefoil, which is associated with other plants. It 
is separate from the wild rye, or Elymus, association, though it 
is close to the tide, because its habitat is rock, not sand. 

Behind this beach-line association, on a somewhat drier area 
farther from the tide, was another zone of miscellaneous grasses, 
with some other plants. Here, we noted a dense stand of Poa, 
(blue grass), Calamagrostis (brown top), Bromus (brome), and 
other grasses that we did not observe closely ; however, we noted 
the demarcation between outer beach Elymus association and the 
adjacent inner zone of other grasses. 

The dividing line was not always located by a given distance 



24 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

from the beach. I recall a striking instance where a sloping banii 
arose from the beach to a height of about 30 feet. Elymus, ex- 
posed to the sands of the sea, clung to the open face of this 
slope to the crest. At the exact point where the ground levelled] 
off toward the interior, the other, more inland, grass formation 
began with a dense growth. The plants of this inner group bor- 
dering the Elymus association are by no means confined to the 
vicinity of the beach; instead, they become diffused among othei 
plants farther inland. I 

Farther in the interior, and at higher elevations, we find what 
Hulten refers to as a "mosaic" of Alpine heath and meadow. 
Meadow formations have an abundance of Car ex (sedge), to- 
gether with many other species, though sedges occur elsewhere' 
as well. In these meadows are found Artemisia unalaschensis 
(a herbaceous sage), Epilobium angustifolium (fireweed), 
Calamagrostis larulsdorffl (a brown top). Geranium erianthum 
(geranium), Anaphalis margaritacea (pearly everlasting), Aconi- 
tum kam.tschaticu7n (aconite). Polygonum viviyarum (viviparous 
knotweed), T7ientalis (star flower), Bromus aleuticus (brome), 
Castilleja unalaschensis (paint brush), Arnica chamissonis 
(arnica), and Aster peregrinus (aster). Such a meadow asso- 
ciation, as defined by Hulten, is more characteristic of the east- 
ern Aleutians. Prominent patches of the characteristic cottons 
grass, Eriophorum, and Ranunculus (bitterroot) , were found im 
many wet areas. Here and there, were found Geum (avens), 
Caltha (marsh marigold), Habenaria (rein orchis), Lupinusi 
(lupine), Geranium (geranium), and a botanical list too long tO' 
enumerate. 

In the more exposed situations above the meadows, scattered 
in accordance with the character of the terrain, are the heaths. 
Here, are lichens, mosses, crowberry (Empetrum nigrum), and 
cranberry, (Vaccinium uliginosum). Numerous other plants are 
distributed rather indiscriminately. The showy anemone (Anem- 
one narcissi flora) , so prominent when in bloom, is very common. 

Mention should be made of Heracleum lanatum (cow parsnip) 
and Coelopleurum gmelini (seacoast angelica). These robust 
plants grow throughout the Aleutian district, apparently where 
soil is rich. They are particularly conspicuous, together with 
other plants, on old Aleut village sites where the soil has been 
enriched by refuse from human habitation. Such village sites, 
seen at a distance, were recognizable by the deep-green, heavy 
vegetation. 

On some occasions we would note a particularly green high 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 25 

mountain slope where we would find a colony of auklets nesting 
among rock crevices. We came to the tentative conclusion that 
vegetation grew more luxuriantly on the site of such bird colonies 
as a result of fertilization by bird guano and waste food. This 
vegetation was not necessarily of the same species as those grow- 
ing on the Aleut village sites ; however, the reasons for its pres- 
ence in the two instances may have been related. 

We did not have opportunity to study in detail the recovery of 
vegetation on islands recently covered by volcanic ash, as Griggs 
(1936) has done at Katmai and at Kodiak. However, little flat 
Ogliuga Island would furnish such an opportunity. In 1930, 
there was an eruption on Gareloi, and the ash from the erup- 
tion covered Ogliuga. At the time of our visits in 1936 and 1937, 
vegetation was just beginning to recover. Tall vigorous clumps 
of coarse sedges, Carex, and some Juncus, had pushed up through 
the ash here and there. These clumps had caught some of the 
drifting ash driven by the wind, had pushed up higher to clear 
the ash, and in turn had caught more wind-driven ash, until 
mounds had been created which were similar to sand dunes. 

In the north are found the so-called bird mounds, whose origin 
has caused much speculation. One theory is that birds such as 
gulls and jaegers, repeatedly alighting on a small prominence, 
fertilize the spot, thus causing exuberant vegetative growth — a 
process that continues until a tall mound is formed. 

On nearby Kavalga Island, I found that a part of the area 
nearest to Ogliuga evidently also had been in the path of an ash 
fall from a volcanic eruption, probably not so heavy a fall as that 
which covered Ogliuga. 

Some typical "bird mounds" on Kavalga were carefully dis- 
sected, with the result shown in the accompanying diagram. In 
figure 23, parts a and b, two such mounds show (by dark spots) 
the wearing away, or undercutting, by wind erosion. Also, note 
the wind erosion on the side in the diagrammatic section of an- 
other mound, as shown in part c. 

Part c shows, in cross section, the layers of materials in one 
of these bird mounds. Note that the first layer under the vegeta- 
tion consists of lava sand, or ash. Beneath the first layer are the 
alternating layers, in increasing width toward the center, of black 
soil and rotted moss. This was, of course, a fairly crude field 
examination, with no opportunity for more precise analysis of 
materials. But the drifting volcanic ash on nearby Ogliuga, pil- 
ing up in mounds around the pioneering clumps of vegetation, 
suggested a process that may also have operated on Kavalga 



26 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

./■.'JM'-.''.. 



wk 




b 



!■ 



^|i 
//^'w 



Figure 23. — Mounds on Kavalga Island (July 1937). a and b, undisturbed 
mounds, c, cross-section of a mound: 1, wind erosion to black soil; 2, 
vegetation on surface; 3, lava sand 21/4 inches; 4, black earth % inch; 
5, rotted moss 1^/4 inches; 6, black earth V4. inch; 7, rotted moss 2 inches; 
8, black earth 1 inch; 9, rotted moss 21/2 inches; 10, black earth 41/2 inches; 
11, rotted moss 12 inches plus. 



to initiate the formation of the so-called bird mounds. Possibly, 
the creation by the wind of these miniature dunes, together with 
the perching of birds thereon, are both involved in the formation 
of these mounds. 

Wind erosion is very severe on some exposures. In places, the 
wind had eroded the soil in troughs, undermining the vegetative 
turf to form a crude type of terracing. The woody roots of crow- 
berry had been exposed and were already supporting a thin coat- 
ing of lichens. With such constant wind action, one wonders how 
the vegetation became established in the first place. As shown in 
part c, wind erosion apparently had affected only the outer layer. 

Marine vegetation is well represented by the kelp beds, which 
consist of a considerable variety of seaweeds that are prevalent 
throughout the Aleutian district. The kelp is, of course, the 
habitat of numerous marine organisms, and during the summer 
it furnishes a favorite habitat for the sea otter. These kelp beds 
disappear in the winter. 

The oceanic climate of this region, the high humidity and pre- 
cipitation, and the prevalence of strong winds have combined to 
shape the vegetative complex that we find in the Aleutian dis- 
trict. In turn, this complex, together with climatic conditions, 
topography, and the rich marine fauna, has influenced the compo- 
sition of the indigenous fauna. 



Bird 



An effort was made to ascertain the Aleut names for birds and 
nammals. There are difficulties in such an undertaking, because 
)ne must be certain that both investigator and native informant 
are talking about the same bird. To make sure of this, a de- 
5cription of the bird and its calls and habits was supplemented 
,vith a colored illustration by Allan Brooks, which was obtained 
Tom the National Geographic Magazme, and in numerous in- 
stances actual specimens were used for identification. In spite of 
jill these precautions, it was necessary to guard against confu- 
l^ion in the minds of the natives because not all of them know 
i;heir birds perfectly. This is particularly true of the more east- 
ern communities, which are farthest removed from a primitive 
itvay of life. The most accurate information was obtained from 
|:he Attn people living at the extreme western end of the island 
l?hain. 

j There also is difficulty both in accurately hearing names spoken 
!3y natives, and in ivriting them adequately. Not being familiar 
jkvith the technique of the ethnologist, I have used the English 
jilphabet to represent the sounds of Aleut words as closely as 
loossible. The endings of Aleut words, or syllables, are also a 
oroblem, because they are very soft, often somewhere between 
h and ch, and sometimes have a soft r sound included. Final ch, 
as used here, is the same as in the German Buck. R is guttural, 
[glided over, and sometimes is accompanied by an h to emphasize 
i:his quality. /, as in "it." E, as in "let," unless marked long. A, 
'is in "Ah." 

; There are at least three Aleut dialects, which are indicated 
jiere as Attn, Atka, and Unalaska. When available, names from 
Alaska Peninsula, recorded by Wetmore or others, are included. 
Stejneger's names from the Commander Islands are also given 
(most of these names are Russian, but some are Aleut). Jochel- 
5on has listed a few names, but usually he did not designate the 
dialect or the exact species. Some of his names cannot be 
dentified; however, only names that are generally accepted are 
ased here. A few names in Russian and Chukchi, from the 
nainland of Siberia, are also given. 

27 



28 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Without doubt, the Aleut language will disappear, and it ii 
worthwhile to record the names that these people applied to th( 
species in their native fauna. Too often, the professional ethnolo- 
gist obtains only the obvious and generalized terms applied to £ 
fauna; however, primitive societies clearly distinguish various 
species, almost as precisely as the scientist. 



Family GAVIIDAE 
Gav'ia immer: Common Loon 

Attu: Kah-goo-gich 
Atka: Kri-giich 
Qigux (Jochelson) 

The common loon, often observed on salt water in winter andi 
in migration, usually is found nesting in interior lakes and pondsi 
rather than in coastal marshes, but it also breeds throughout the^ 
Aleutian district. In this connection it is interesting to note that 
it does not occur in the Pribilofs. 

Bones of this loon have been identified from Kodiak Island and^ 
from Little Kiska (Friedmann, 1935, 1937). A specimen was 
taken by Bretherton on Kodiak (1896), and the bird was re- 
ported by Chapman at Seldovia (June 30, 1903). Ball (1873) 
reports a "ColymMis torquatus" at Simeonof Island, in the 
Shumagins, on September 2, 1873, and further reports (1874) 
that it breeds on Kiska and is abundant on Amchitka. On July 
23, 1925, I noted a pair of common loons, probably nesting, on a 
pond near Izembek Bay, near the west end of Alaska Peninsula. 

In 1936, our party saw one of these birds (probably a migrant) 
on May 11, near Ushagat Island of the Barren Islands group, and 
on May 23, in Nushagak Bay, we saw six or more. The greatest 
number of these loons was found on Adak Island, though we also 
saw them on Amchitka, Kanaga, and Kiska. In 1937, at least 3 
pairs were found on Agattu, and on June 17 of that year we 
found at least 2 pairs on Semichi Islands, each with 2 small downy 
young. In this instance, when we disturbed the adults, a glaucous- 
winged gull swooped down and carried off one of the young. 

Dall reported that the common loon does not winter in the 
Aleutians, and Mike Hodikoff, chief of Attu Village, stated that 
it arrives at Attu Island in April and departs in October. How- 
ever, during the years 1940 and 1946, Gabrielson found these 
loons on various islands as far west as Atka in midwinter, and 
in early spring and summer they were "common" or "plentiful' 
in numerous localities throughout the entire Aleutian district. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 29 

Also, Cahn (1947) found this loon "not uncommon in winter in 
Captains and Makushin Bays" of Unalaska Island, and he noted 
it also in Iliuliuk Bay; the latest date was March 3. 

Gavia adamsii: Yellow-billed Loon 

Russian, Commander Islands: Bolschoj gagara (Stejneger) 
Russian, latitudes of Yana : Gagara 

Bolschaja gagara (Pleske) 
Chukchi: Uvanketsjouku (Palmen) 

It is extremely rare in this district. Herbert Friedmann (1934, 
1935, 1937) has recorded bones of the yellow-billed loon from 
middens on Kodiak Island, on Amaknak (near Unalaska), and 
on Little Kiska Island in the western Aleutians. A specimen was 
collected at Kodiak by Bischoff in 1868, and Fisher obtained an 
adult male in 1881. 

We did not observe this species on our expeditions, but the 
-"hief of Attn seemed to recognize pictures of the bird and said 
it occurs in his home area occasionally. Stejneger (1885, 1887) 
considered it to oe a rare winter visitor in the Commander Is- 
lands, where he obtained one specimen and saw another. The 
bird he obtained was found on glare ice. unable to rise; evidently, 
it had mistaken the ice for water. 

According to published accounts, this loon migrates along the 
Alaskan coast, from southeastern Alaska, west and north through 
Bering Strait. Presumably, the fall migration is the reverse of 
this. Several specimens are recorded from the Pribilofs (in May 
and August) as transients. In the spring of 1924, I obtained a 
specimen from an Eskimo at Hooper Bay and was informed that 
these loons pass that point in migration. It is likely, however, 
that the yellow-billed loon migration is not confined to the 
Alaskan coastline. In the autumn of 1924, several natives along 
the Koyukuk River in interior Alaska assured me that the yellow- 
billed loon passes through there in migration, though it does not 
nest there. They seemed well acquainted with the species as it 
was described to them, having particularly noted the light-colored 
bill. Therefore, the yellow-billed loon, nesting in the far north, is 
widely scattered in migration and occurs as a transient in the 
Aleutian, Commander, and Pribilof Islands. 

Gov/o arcfica: Arctic Loon 
Gavia arcfica pacifica 

Russian: Gagara 

We could obtain no Aleut name for this species. The Russsian 
name for loon in the general sense seems to have been adopted 



30 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

by the Aleuts, so that we find Bretherton (1896) and Turner 
reporting different forms of this word as the Aleut name for loon 
in general, and Nelson applying it to the red-throated loon. 

The Arctic loon is widely distributed, nesting commonly on 
parts of the coastal plains of Bering Sea and the Arctic Ocean, as 
well as in many inland localities. It is quite common on the east- 
ern portion of Alaska Peninsula, but it becomes exceedingly rare 
to the westward, as the following records show, 

Bretherton and Bent report them nesting on Kodiak Island, 
and Cahalane (1943) found them to be common in the Kodiak- 
Afognak group in 1941. Friedmann (1935) records bones at 
various levels in archeological diggings on Kodiak, this indicating 
a regular occurrence over a long period. 

Writing of his observations on the Alaska Peninsula in 1940, 
Cahalane (1944) says. 

We found this species common on the Naknek River September 3, and at 
Brooks Lake September 9. . . . On the western shore of Shelikof Strait, I 
recorded loons as "common" between Amalik and Katmai Bays, October 4, 
and "abundant" on the following day in Amalik and Kinak Bays and Geo- 
graphic Harbor. 

He also observed them off Cape Nushagak, October 7, but he 
adds, "They were absent from the interior of the Alaska Penin- 
sula, even where suitable habitats existed." 

These observations were made chiefly in the migration period, 
when Arctic loons are strikingly abundant along the Alaskan 
coast. On May 18, 1937, as we were approaching Valdez, Pacific 
loons were scattered widely over the water of the fjord. We 
counted at least 75 at one time. One loose flock contained 50 loons. 

In 1940, Gabrielson observed 30 or more pairs, as well as scat- 
tered individuals, on Kvichak River, July 23, and he noted 2 of 
these birds at the upper end of Iliamna Lake on July 24. On July 
7, 1946, he noted a loon at Port Moller. 

Jaques (1930) found them to be common near Port Moller, 
June 1-23, 1928. On May 29, 1936, I noted at least eight pairs, 
apparently preparing to nest, among the ponds bordering the 
lower reaches of Ugashik River, but they are scarce at the west 
end of the peninsula. 

Farther west, these loons are less numerous. Among the Aleu- 
tians proper, we did not identify a single Arctic loon during two 
seasons of extensive field work and a third season of hasty recon- 
naissance. The chief of Attn Village did not recognize pictures 
of the bird and declared that no such bird occurs there. Donald 
H. Stevenson, former warden in the Aleutian Islands National 
Wildlife Refuge, reported them as "not common." His only spe- 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 31 

cific record is the mention of two birds that he saw at Unalaska 
Harbor, October 15, 1920, which he thought were of this species. 

Austin H. Clark (1910) mentions only one bird, which was 
seen on a lake on Agattu Island in 1906. 

Turner, however, reported this loon, under the name of Uri- 
nator pacificus, as a common lareeding bird in the Near Islands 
(1885), and, later (1886), he specifically reported one at Attu 
in the winter of 1880-81 and said that they nested commonly 
on Semichi Islands. Using the name Urinator ai^cticus, he said 
that this species was to be found among the Aleutians at any 
time of year, and he mentions seeing one at Amchitka Island 
in June. 

These reports of Turner are rather surprising, and certainly 
they are not in accord with more recent findings. We had abun- 
dant opportunity to examine Agattu, Semichi, and Amchitka Is- 
lands, which were specifically mentioned by Turner, and though 
we found the common loon and red-throated loon, we did not see 
an Arctic loon. Stejneger did not record it for the Commander 
Islands, and it has not been recorded for the Pribilofs. 

Gavia arctica viridigularis is known to be an occasional Old 
World straggler from Siberia to Alaska, and it has been recorded 
on the Pribilofs. Turner recorded two forms for the Aleutians, 
therefore it might be expected that viridigularis has occurred 
among those islands. However, in view of the confusion that 
has existed over the identity of the American forms of this loon, 
and because of its complete absence from the Aleutians, in recent 
years at least, a reported occurrence should be well authenti- 
cated before being accepted, 

Gavia stellafa: Red-throated Loon 

Attu: Ka-ka-dra-cha or Ka-da-dra-ka 

Atka: Ka-kach 

Russian, Commander Islands: Gargara (Stejneger) 

Russian, latitude of Yana: Gagara (Birula) ; Malaja gagara (Birula) 

Chukchi: Jouku (Palmen) 

As previously mentioned, the Russian word "gagara" is used 
by natives in various parts of coastal western Alaska. This, and 
the Aleut names, are imitations of the call of this loon. 

The red-throated loon is the most abundant and widespread 
loon in the North, especially on coastal areas, and it occurs on 
both shores of Bering Sea. On the basal portion of Alaska Penin- 
sula it appears to be less abundant. Neither Gabrielson nor 
Cahalane reported seeing it there, although they observed the 



32 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Pacific loon. We did not observe them there on our expeditions 
Osgood (1904) records a pair on Chulitna River, August 12 
1902, and he observed a few others "at comparatively long in- 
tervals" on the Chulitna and Kakhtul Rivers; however, he add{ 
that "they were far exceeded in numbers by the Pacific loon.' 
McKay had collected specimens at Nushagak, and Friedmann haj 
recorded the bird from Kodiak Island. 

At the western end of Alaska Peninsula, however, red-throatec 
loons were abundant. They were noted in some numbers on Izem- 
bek Bay in 1925. Turner (1886) found them to be abundant ir 
the Aleutians and records them nesting on Atka, Semichi, anc 
Agattu. We found these loons to be plentiful on Semichi, Agattu 
and Amchitka, and we noted them on Sanak, Adak, Tanaga 
Kiska, and Attu. Gabrielson records them on Attn, Amchitka 
and Izembek Bay. Friedmann and Cahn also recorded the bird 
from Unalaska. They are present on all islands that bear suit- 
able nesting ponds, and many of these red-throated loons spend 
the winter in the Aleutians. 

Stejneger and Hartert report this loon as "abundant" and 
breeding "frequently" in the Commander Islands. Clark also 
noted a pair on Bering Island in 1906. 

Family PODICIPEDIDAE 

Pod'iceps gr'isegena: Red-necked Grebe 
Podiceps grisegena holbolUi 

Friedmann (1935) records a bone, as well as several skins, 
from Kodiak Island. Cahalane (1943) recorded these birds as 
numerous in Uyak Bay and recorded a few in Kupreanof Strait. 
He also observed 2 birds on Brooks Lake, in the Katmai region,, 
on September 9, and he observed 4 or 5 on the lower Naknek 
River on September 28. He stated, "On Shelikof Strait, the 
species was abundant between Katmai and Kinak Bays on Oc- 
tober 4 and 5, and off Cape Nukshak on the 7th." 

On May 29, 1936, we found one of these birds in a pond, ap- 
parently nesting, near lower Ugashik River, and another was 
swimming in the river. 

A little farther west, in ponds near Port Moller, Jaques (1930), 
reports several, June 4 and 20, 1928, and Gianini (1917) observed 
several at Stepovak Bay in June 1916. 

On April 28, 1925, I obtained a specimen at False Pass, at 
the extreme tip of Alaska Peninsula, and on July 21, 1925, I 
found an adult with two young in a pond near Moffet Cove, at 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 33 

the east end of Izembek Bay, thus positively establishing the 
species as a nesting bird that far west. The bird was heard call- 
ing in the evenings. 

In 1942, Gabrielson observed this grebe at Cold Bay, and on 
October 20, 1943, he obtained a specimen at Kodiak and obtained 
two more in September and October 1946. 

There are a number of records for Unalaska. Turner mentions 
two specimens from there. There is a specimen in the Natioixal 
Museum that was taken by Dall at Unalaska, December 14, 1871, 
and Donald H. Stevenson inforaied me that this grebe occurred 
on the salt water at Unalaska, chiefly in the fall. Laing (1925) 
also reports it at Unalaska and at Atka. 

More recently, Cahn (1947) reported this grebe as not un- 
common at Atka in November, December, and January. 

There are a few records of the red-necked grebe west of 
Unalaska, though we have no positive data on nesting. Taber 
(1946) observed a flock of about 50 at Adak Island on November 
25 and observed another large group December 16 — this group 
disappeared by December 25, These sightings were on the 
salt water of Bering Sea. On June 18, 1936, we observed a 
pair as they arose from the salt water near the northeast shore 
of Seguam Island. We have no record of its occurrence west 
of Adak, but Stejneger (1885) described it as a rare straggler 
in the Commander Islands, where he obtained a specimen. Hartert 
also (1920) considered it a straggler in the Commander Islands, 
where he obtained three specimens. 

Podiceps aurifus: Horned Grebe 

This little grebe is found sparingly in the Aleutian district, 
and there is no evidence that it nests there. Friedmann (1935) 
found osseous remains in a collection of bones from Kodiak 
Island and lists nine specimens taken there, most of which were 
taken in the winter months. On October 1, 1940, Cahalane (1943) 
recorded several grebes in Viekoda and Terror Bays, Kodiak 
Island. Referring to the Katmai region of the Alaska Peninsula, 
he reports one horned grebe on Brooks Lake, September 9, 1940; 
he found them scarce west of the Aleutian Range. On the east 
side, however, he found them abundant and observed "great 
numbers" in early October along the Shelikof Strait coast of 
Katmai National Monument, as well as in most of the inlets from 
Katmai to Kinak Bay. Osgood (1904) recorded several small 
grebes, assumed to be this species, at Becharof Lake, October 



34 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

6-7, 1902, and McKay obtained a specimen at Nushagak, June] 
21, 1881. 

Gabrielson noted two specimens at King Cove, March 25,J 
1942. He obtained a specimen at Kodiak in October 1944 and 
obtained three more in 1946. 

On our expeditions we saw no horned grebes in the Aleutian] 
Islands, but we observed several at Port Chatham, Kenai Penin- 
sula, on May 6, 1936, and observed two at Ushagat Island, of the 
Barren Islands group, 5 days later. 

However, there are a few records for the Aleutian chain.! 
Laing (1925) saw about a dozen horned grebes at Unalaska,] 
presumably in early spring judging from his itinerary. Bailey I 
(1925) records two specimens taken by Hendee at Unalaska, 
September 21, 1922. Cahn (1947) reports on this bird at Un- 
alaska: ''Seen sparingly in any of the bays during December 
and January, always solitary and rather shy. December 2, 1943, 
and February 21, 1946 are the extreme dates of record." Turner; 
(1886) reports seeing a grebe at Attn that he suspected wasi 
auritus, but states that he did not observe it in the Aleutians in 
the summer, "and at no time to the westward of Unalaskai 
Island." However, on July 1, 1946, Gabrielson noted one hornedi 
grebe on Amukta. Taber (1946) noted them in small numbersi 
on the salt-water lagoons at Adak Island throughout the period 
of his observations and mentions specifically the dates November 
22 and January 9. 

The bird noted by Turner at Attu might well have beeni 
auritus, because Stejneger (1885) obtained a skeleton of one ini 
the Commander Islands, though he considered it rare. Hartert 
(1920) also records two horned grebes wintering on the Com-i 
mander Islands. 

There is no evidence that the horned grebe nests in the Aleutian 
district, but, according to these records of its occurrence, it 
evidently winters in those waters. 



Family DIOMEDEIDAE 
Diomedea n'igripes: Black-footed Alba+ross 

Attu: A-la-gri-gich or Ah-la-gri-gich 
Atka: A-ga-lig-ahh or Ah-ga-lig-ach 
Agligax (Jochelson) 

This is the albatross common in the North Pacific during our 
summer season, at least from May to October. The earliest dates 
that I have noted this bird were April 20-23, 1925, in the open 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 35 

sea from Ketchikan to the western part of Alaska Peninsula. 
The black-footed albatross occurs commonly in the Gulf of Alaska ; 
it was observed in the vicinity of Kodiak, and frequently along 
the Alaska Peninsula and the Aleutian chain, though it rarely 
was observed near land. This albatross also was seen in the 
waters of Bering Sea, just north of Alaska Peninsula and the 
Aleutians. We found these birds particulary plentiful at the 
Petrel Banks, north of Semisopochnoi Island. Laing (1925) re- 
ports them "about 100 miles north of Kyska Island." Though 
these birds were usually far offshore, there are two places where 
they came near land — Seguam and Kiska Islands. 

Nelson (1887) was of the opinion that the northern limit of 
distribution was considerably south of the Aleutians and quotes 
T. H. Bean to the effect that latitude 51° marked the northern 
limit. Turner (1886), on the other hand, stated that the species 

is quite a common bird in some localities north of the Aleutian Islands. 
In Bristol Bay in June. 1878, I saw numbers of them in the vicinity of 
Cape Newenhani . . . Toward the western Aleutian Islands they are not 
common but are frequently met. 

It is not certain that this albatross ranged beyond latitude 51° 
N, in the past, nevertheless it does so today, and, at least in the 
Bristol Bay region, it reaches nearly to latitude 58° N. Further 
consideration of albatross distribution will be found in the dis- 
cussion of the short-tailed albatross. 

An interesting story was told to me by several old natives at 
Atka Island. They insisted that albatrosses used to nest in small 
numbers on Bobrof Island, on top of the mountain, in ivinter. 
Judging by the descriptive gestures of one informant, icicles 
formed on their beaks while the birds were incubating. This is 
indeed a strange legend. It seems unbelievable that any of these 
albatrosses could have tried to nest in the Aleutians, but at least 
these Aleuts were talking about the proper nesting season. 

If I correctly understood the information given me by the 
Aleuts, they do not distinguish two species of albatrosses in the 
islands, possibly assuming that nigripes is the young of albatrus, 
in which case this story might more properly apply to albatnis. 

A female D. nigripes collected June 17, 1936, near Seguam 
Island weighed 614 pounds and had a wingspread of 85 inches. 

The black-footed albatross is said to feed on whatever be- 
comes available at the surface of the water, including refuse 
from ships — this is true for our experience in the Aleutians. 
On one occasion, a dozen were following our ship; they were 
attracted by fish offal thrown overboard by the sailors, who 



36 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

were cleaning codfish, At times, we saw the albatrosses, ac- 
companied by shearwaters and auklets, feeding in tide rips. 

Cottam and Knappen (1939) reported on two stomachs of the 
black-footed albatross from the Aleutians. One stomach contained 
fish (mostly Scorpaenidae) , 35 percent; remains of 6 or more 
squids, 55 percent; sea urchin, 2 percent; brown algae, 8 percent. 
The other stomach contained fish (mostly Scorpaenidae) 92 per- 
cent; Gammaridae, 1 percent; brown algae, 7 percent. 

In 1941, Loye Miller (1942) found, in the course of experi- 
mental food studies off the coast of southern California, that 
"the greatest gooney appeal was provided by bacon drippings 
which had congealed to semisolid state." For better handling 
of this bait material, it was mixed with puffed rice before 
cooling. He remarked further that "Bacon grease seems to throw 
the birds almost into a frenzy. Some of them rushed right up 
under the overhang of the poop." 

Miller summed up his findings as follows: 

The most attractive bait discovered is animal fat. Bacon fat was superior 
to beef suet. The semisolid gelatin settling out from roast beef drippings 
was of no interest at all and was neglected after the first taste. Taste 
buds in the tip of the bill appear to be highly sensitive and discriminative. 
The turpentine-linseed flux of paints used in marking is very repugnant 
and seemed to be recognizable by odor before actual contact was made. 
I was repeatedly impressed by their seeming acuity of olfactory perception. 

He noted, further, that in subsurface feeding the albatross would 
tip up, or actually submerge to a depth of at least 2 feet with 
wings partially spread. This suggests a trait similar to that of 
the slender-billed shearwater, though the latter is capable of 
descending to a depth of many fathoms, 

Diomedea albafrui>: Short-tailed Albatross 

Kodiak : Kay-mah-rye-erk (Nelson) 

As mentioned above, the Aleuts apparently do not have separate 
names for the two species of albatross. At least one Aleut 
identified albatrus as the adult bird, nigripes the young. Thus, 
the Aleut nomenclature is confusing, and the names already 
given for the black-footed albatross might apply equally well to 
the short-tailed albatross. 

We are concerned here with what appears to be an extinct 
bird. We had thought that a few remained in the Aleutiar 
district, but when the one specimen we collected in 1937 proved 
to be imniutabilis, serious doubt was thrown on the possibility 
that any of the light-colored birds were albatnis. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 37 

Bering Sea appeared to be the particular domain of the short- 
tailed albatross in summer. Nelson (1887) defined its summer 
range from 50° N. latitude northward through Bering Sea as 
far as Bering Strait. He reported them in the Aleutians and 
quotes T. H. Bean as having found them around the Gulf of 
Alaska, but he considered the mouth of Cook Inlet and the 
vicinity of the Barren Islands as their favorite resort. Nelson 
"found them very common between the islands east of Unalaska" 
during May 1877. Turner also found them plentiful among the 
Aleutians, as well as at Cape Newenham in the Bristol Bay 
region. Friedmann, who has examined bones unearthed from 
ancient village sites on Kodiak, Amaknak, Unalaska, Little Kiska, 
Atka, and Attu Islands, found numerous remains of this alba- 
tross, but he found no remains of nigripes. They are reported to 
have been abundant in the vicinity of the Pribilofs when whalers 
were active there, and they became scarce when whaling was 
abandoned. 

Austin H. Clark (1910), writing of his expedition in 1906, 
reported that — 

We first saw this species about 100 miles east of Unalaska on the day 
before our arrival at Dutch Harbor. On the next day, two were seen near 
the Aleutian chain, one of them within five miles of the islands. Two 
more were seen between Attu and Copper Island, on June 12; on the 20th 
one was observed about 20 miles off the Kamchatka coast, and the next 
day another in the Okhotsk Sea, near the mouth of the Aangan River. 
On October 1 this species was very common about the southern end of the 
Kurils, on both the inside and outside of the chain. 

Clark believed that the birds were more abundant than these 
notes indicate, because they are very shy and not readily observed. 

Stejneger (1885) reported that the species is not a rare visitor 
to the Commander Islands, and he, too, considered them "re- 
markably shyer than D. nigripes." 

Nelson (1887) also considered them shy, though "natives 
of Alexandrovak sometimes spear them from their kayaks." 

According to Otto Geist (in Murie 1936), in earlier days, 
near St. Lawrence Island, these birds "... were often caught 
on the pack ice near the island. This was often easy because 
the birds were very fat and could hardly make their way in the 
air." 

Today, the short-tailed albatross is rare, or extinct. Although 
Nelson had reported it as common in Bering Strait and noted it 
at St. Lawrence Island, in 1887, Otto Geist, in the course of 
archeological work on St. Lawrence Island from 1926 to 193'5, 
did not see this bird. However, bones were found in excavations, 



38 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

and natives stated that it had been present in considerable num- 
ber^ at one time (Murie 1936). Bent (19^2) reports that he 
did not observe this bird during a cruise along the Aleutian chain 
in June 1911. Laing (1925) obsei-ved two whitish albatrosses at 
a distance in the Kuriles in 1924, but he saw no others during two 
crossings of the North Pacific. In the course of many voyages 
across the Gulf of Alaska after 1920, I never saw a short- 
tailed albatross. In 1936 and 1937, we cruised about the Barren 
Islands several times and saw none, although this had been 
considered to be a favorite area for them by T. H. Bean. 

It appears, then, that at one time the short-tailed albatross was 
plentiful in the Aleutian district and Bering Sea region in general, 
but that the population had suffered a drastic reduction in 
numbers, probably about 1900 or a little later. 

Austin (1949) has indicated that Japanese fishei*men and plume 
hunters were responsible for the destruction of this species on its 
nesting grounds. But it seems that the decline began long ago. 
Did the plume trade affect this species, as it affected the Laysan 
albatross ? Whatever the facts might be, the concentrated nesting 
of a species on one or on a few small islands constitutes a serious 
hazard to its perpetuation. 

There is a puzzling problem in distribution revealed by Fried- 
mann's work on bones found in ancient village sites. All bones 
found on the Aleutian Islands proved to be those of D. albatrus, 
and not those of D. nigripes. This indicates that in earlier times 
D. alhatrus was the common bird of the region and that D. 
nigripes was scarce or absent, at least close to the coast. Even 
at Kodiak Island, though there were some bones of D. nigripes, 
Friedmann found numerous bones of D. albatrus, thus confirming 
early reports of this bird's abundance in those more easterly 
waters. 

Friedmann's findings from midden material, therefore, lend 
some support to Bean's designation of 51° N. latitude as the 
northern limit of range of D. nigripes, even though Turner states 
that he saw this bird as far north as Bristol Bay. Otherwise, 
at least a few bones of this bird would have appeared in middens. 

In his study of the distribution of these two forms off the 
California coast, Loye Miller (1940) did not find nigripes in 
channel waters near the coast, but found them farther out; how- 
ever, there is evidence that albatrus did occupy the channel waters 
chiefly. He quotes Willett to that effect, and remarks: "The 
two birds seem to have divided the territory between them, as 
it were." He states, "I have taken from the channel Indian 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 39 

mounds great number of their [D. aJbatrms] bones, but never 
any of D. nigripes." 

Here, we have a parallel with the situation in the Aleutians. 
In both areas (California and the Aleutian district), only the 
short-tailed albatross occurs in middens, except for some over- 
lapping of the two species at Kodiak. Is it possible that in early 
times D. alhaUnis was the species that came closer to land and 
was therefore more available? Apparently, in the north, too, 
these two species had "divided the territory between them." 

We need to know more about the whitish birds being seen today 
in the Aleutian district. Are some of these birds the remnants 
of the vanishing short-tailed albatross, or are they all Laysan 
albatrosses? If they are the latter, will that species eventually 
take over the range of albatms ? 

Turner, in writing of the short-tailed albatross, mentions two 
birds that were killed at Attn in the latter part of March, 1881. 
He claimed that "this species passes the winter in this locality 
and may be found, during very severe weather, about the 
western end of Attu." And he believed that it nested somewhere 
in that region, which also was the belief of the Attu chief at the 
time of our visit there. 

Nelson mentions a specimen, obtained by Dr. Bean, that had a 
wing spread of 88 inches. 

Diomedea immufabilis: Laysan Albatross 

We had not suspected that this bird occupied the Aleutian 
district until a specimen, collected near Ulak island, July 31, 1937, 
eventually proved to be of this species. On the field trip, we had 
assumed this specimen to be D. albatrus, and on the basis of that 
assumption we had recorded our observations on white albatrosses 
as albatrus. However, because Oliver Austin has pointed out the 
extreme scarcity of specimens ©r certain records of the short- 
tailed albatross in recent years, and suggests its probable extinc- 
tion, there must remain doubt about the records on our expedition. 
This, of course, throws much doubt on the possible existence of 
albatrus in the Aleutians today, but I shall record our observations 
for what they are worth, keeping in mind that the species ob- 
served was in doubt in each instance. 

In these seasons of field work, we occasionally saw white- 
colored albatrosses throughout the Aleutian islands. In 1936, we 
saw one between Seguam and Chagulak Islands on June 21. On 
July 31, we saw another sitting on the water between Buldir and 



40 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Kiska Islands, and, later in the evening, we saw one nearer to 
Kiska Island. 

In 1937, we saw them oftener: On May 31, 1 north of Seguam; 
on June 2, at least 3 west of Atka; on June 3, several near the 
Petrel Banks and 2 between Kiska and Segula ; on June 17, 1 east 
of Semichi Islands; on June 18, 1 east of Buldir; and, on July 
31, 1 specimen was taken at Ulak Island, 178° W. longitude. 

In 1938, Scheffer reported the following: On August 11, 4 
were seen south of Atka, 3 of these in 1 group ; August 14, 1 was 
seen south of Khwostof; August 17, 1 was seen northwest of 
Semichi ; August 25, 1 was seen south of Tanaga ; and, on Septem- 
ber 23, 1 was seen in the Gulf of Alaska off Cape Hinchinbrook. 

On June 9, 1940, Gabrielson observed a white albatross fol- 
lowing the ship in the Gulf of Alaska. In 1941, he saw the 
following: February 3, 1 was sighted near Amchitka, and on 
February 7, 2 were seen near Amchitka Pass; June 24, 1 was 
seen at Tanaga Bay ; June 27, 1 was seen near Amchitka ; and on 
July 22, 2 were seen near Attu. 

Were all these birds the Laysan albatross, the same as the 
one specimen collected, or were some, or most, of them albatr-us? 

The specimen collected weighed 4 pounds 6 ounces. Its length 
was 301/2 inches, and wingspread was 77%, inches. 

During 1947, 1948, and 1949, on crusies for the Fish and 
Wildlife Serice in North Pacific waters, Karl W. Kenyon (1950) 
made noteworthy observations on albatross distribution. Alert 
to the confusion in sight identifications of light-colored birds, he 
gave close attention to field characteristics. 

On October 13, 1948, Kenyon saw a Laysan albatross about 230 
miles east of Kodiak, and Captain Carlson said that during 5 
round trips to the Pribilofs he often saw 1 or 2 of the white 
albatrosses with the black wings and back at about this same 
place while crossing the Gulf between Kodiak and Cape Spencer 
but not in the Bering Sea. 

Kenyon records a number of other sightings in the North 
Pacific, but south of the Aleutian chain. He received parts sal- 
vaged by Elmer C. Hanson from two dead birds at the Army 
air base at Amchitka, June 5, 1948, which were sufficient for 
identification as Laysan albatrosses. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 41 

Family PROCELLARIIDAE 
Puffinus tenuirosfris: Slender-billed Shearwater 

Attu: A-la-mach 
Kussian, Commander Islands: "Tschornij Glupisch" (Stejneger) 

In summer, the slender-billed shearwater is distributed widely- 
over the North Pacific and throughout the Bering Sea. On the 
Gulf of Alaska, we observed them often, singly or in small groups. 
Some of the birds that we thought to be slender-billed shearwaters 
may have been the sooty shearwater, P. griseus. 

At Kodiak Island, however, P. tenuirostris has been identified, 
and we obtained a specimen there in 1936. Shearwaters, believed 
to be this form, were noted May 10, 1936, near the entrance to 
Cook Inlet; one was noted near Barren Islands on May 11; they 
were numerous between Sutwik Island and Cape Kumlik on May 
14, and there was a flock at the entrance to Chignik Bay. On 
May 15, a few were seen near Nagai Island, in the Shumagins; 
on August 29, some were noted near Simeonof Island in this 
group, and some were noted between that point and the main- 
land; next day, between Kupreanof Harbor and Chignik, more 
were sighted. Again, on September 1, we passed through dense 
masses of shearwaters north of Karluk, in Shelikof Strait. These 
birds occur also on the north side of Alaska Peninsula and were 
seen as far east as the entrance to Bristol Bay. 

Slender-billed shearwaters occur all through the Aleutian 
Islands, with their center of abundance apparently at the eastern 
end of the chain, among the Fox Islands. Unimak Pass is a 
favorite feeding place, with large concentrations also observed 
in other places as far as the western end of Umnak Island. 

Many published accounts describe the hordes of shearwaters 
observed at various times. Arnold (1948), during an hour and a 
half, June 9, 1944, recorded 160,000 shearwaters in Unimak Pass. 
Probably the most outstanding was the flock noted by Scheffer 
in Umnak Pass on September 3, 1938. He says, in his field report: 

In the Pass we saw the greatest concentration of shearwaters that we have 
ever seen in the Aleutians. Captain Sellevold remarked that it was the 
greatest in his experience. The Pass is 3 miles wide. We estimated that 
the raft of birds extended for 25 miles by 2 miles wide, or an area of 50 
square miles. From 5:30 a. m. to 8:00 a. m. the ship passed through 
dense masses of the birds, about half of them on the water and half 
flying back and forth ... At 5 p. m. the birds had thinned out by more 
than half. 

Apparently this bird is much less abundant to the westward, 
though in 1941 Gabrielson saw "thousands" at Attu and "several 



42 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

thousands" at Atka. Stejneger (1885) considered it rather scarce 
in the Commander Islands. It appears to be relatively scarce in 
Bering Sea north of the Aleutian chain, though it does range far 
northward. In 1944, Gabrielson saw "thousands" between St. 
Matthew and Nunivak Islands and a "scattering of birds" be- 
tween St. Matthew and Adak. Nelson (1887) mentions a specimen 
obtained by Dall in Kotzebue Sound in 1899 and adds, "just 
northwest of the straits, the last of August, 1881, quite a number 
of dark-plumaged birds v/ere seen, with many Rodgers's fulmars, 
which appeared to differ m size and appearance from the latter, 
and which I am inclined to think belonged to this species," Nelson 
records the Eskimo name for the slender-billed shearwater in 
Kotzebue Sound, thus showing that it must habitually occur 
there. A number of specimens have been taken at St. Lawrence 
Island in recent years (Murie 1936). The Eskimos of this 
island have certain taboos in connection with eating the bird. 
A specimen is recorded from the Pribilofs. 

According to Bent (1922), the northward migration is mainly 
in the western Pacific-, and the southward migration is in the 
eastern Pacific. He says they occur in the Okhotsk Sea, as well 
as in Bering Sea. 

Slender-billed shearwaters will feed on refuse from a ship's 
galley. On August 10, 1937, three of these birds came up to 
the stern of our ship as we drifted in the fog. They readily 
ate bits of beef and fish thrown overboard. They would dive for 
pieces that sank, keeping wings half opened, but propelling them- 
selves entirely by their feet under water. Apparently they can 
go to a considerable depth ; they would sometimes pursue a 
baited fishhook thrown in for cod and remain underneath for a 
considerable length of time. 

On two occasions we watched a shearwater pursue an injured 
codfish that had been discarded by the sailors as being wormy; 
as the fish wiggled feebly through the water, the bird pursued 
it and snipped out pieces of the gills from the still-living fish. 

But probably the most important food of the shearwater con- 
sists of crustaceans and other small invertebrates. On numerous 
occasions they joined with fulmars and auklets in the turbulent 
tide rips, where all were busy feeding. Sample tows taken in 
such places revealed a greater quantity of plankton than in 
adjacent areas unoccupied by birds. In this connection, it may be 
significant that the center of abundance of shearwaters in the 
Aleutians today coincides fairly well with localities where whales 
were once particularly abundant — in the Fox Island group. Per- 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 43 

haps both birds and whales were attracted by the swarming in- 
vertebrate life. 

Cottam and Knappen (1939) reported on 10 stomachs of this 
species that were collected in Alaska. They tabulated the total 
contents as follows : 

Amphipods, 13%; schizopods, 15%; undetermined crustaceans, 20.8%; 
squid (Loligo sp.) 16.1% ; undetermined marine invertebrate flesh, possibly 
squid, 29.4% ; and fish, 5.7%. Gravel was found in each stomach and 
averaged only slightly less than 40% of the contents. 

In their summer range, the only foe of the shearwater that we 
could discover is the northern bald eagle. The shearwater fur- 
nishes one of the principal items of the eagle's diet, probably 
because of its great numbers. These birds also perish in storms, 
perhaps the storm mortality rate is higher than that caused 
by natural enemies. Early in September 1937 a heavy rain- 
storm was raging for several days about Unimak Island. While 
the storm was still at its height, we began to find dead shear- 
waters on a strip of beach. They finally totaled over 30 ; one 
bird was still alive. The birds from this one sample beach 
may have been representative of the destruction of birds over 
a large area. In 1925, I frequently found dead shearwaters on the 
beaches of Alaska Peninsula. 

On May 31, 1936, as we approached Cave Point on Unimak 
Island, a lone shearwater was sitting on the water. C. S. Williams 
shot it for a specimen, and it was found to be poor in flesh. On 
June 30, 1937, Scheffer shot one that was in a similar condition 
near Rat Island. As it sat on the water, we noticed that it 
appeared to be "dumpy." Grinnell (1900) says, "On July 4th, 
1899, I secured a single speciment about 4 miles off Cape Blossom. 
It was resting on the water not far from a small icefloe and was 
in an emaciated condition." 

Certainly, there is a mortality element of some sort operating 
on the shearwater, as on other sea birds. 

Occasionally, shearwaters come close to land. At least 1 was 
seen in Nelson Lagoon, and in 1925 they were noticed at the 
entrances to Izembek Bay, on the Alaska Peninsula, and at least 
1 flew over the shallow bay. Some of the large flocks noted in the 
turbulent waters of various island passes are not far from the 
rocky points. 

Chief Hodikoff, of Attu Island, declared that shearwaters 
formerly nested abundantly on Agattu Island and that a few 
were seen on Semichi Island in 1932. He said there were "not 
many" now since the introduction of blue foxes. He stated that 



44 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

they nested "on level ground, some in clumps of grass" and that 
as soon as the young were able to fly they left their nesting 
grounds. 

It is interesting to note that, about 1879, Turner was on 
Amchitka Island with some Attn natives, when they picked up 
a dead shearwater. The natives told him that the birds "breed 
plentifully in the Semichi Islands." Though the breeding range 
of shearwaters is now well established, perhaps we should not 
entirely ignore native information of this kind. 

PuffJnus gr'iseus: Sooty Shearwater 

According to the fourth edition of the Check List of NorthI 
American Birds, of the American Ornithologists' Union, thei 
sooty shearwater occurs in the Aleutian and Kurile Islands. Oni 
all of our expeditions, we scrutinized flocks of shearwaters in am 
attempt to identify this species among the predominent slender- 
billed shearwaters. Although we thought that we could seei 
differences in some instances, positive identification was doubtful.l 
But, in the series of specimens of shearwaters collected in thei 
Aleutian district, a single specimen proved to be P. griseus. 

Nichols (1927), speaking of his voyage between Seattle and 
the Aleutian Islands, says : 

Of the sooty shearwater I have no satisfactory identification, but am of thei 
impression that it replaced the generally common slender bill farthest offi 
shore at a point midway between the islands and the coast, and to somei 
extent at least on the east side of the Gulf of Alaska. 

Pferodroma inexpecfafa: Scaled Petrel 

Attu: Le-vi-dre-che 

This petrel has a wide range, but we have little informa- 
tion on it in the southern Alaskan waters. There is a record for 
Kodiak Island, a specimen collected by Fisher, June 11, 1882, that 
served as the type of Ridgway's Aestrelata fisheri. Wetmore 
collected a specimen at the Alaska Peninsula, August 6, 1911, 
and while crossing from Cape Muzon to Unimak Pass he ob- 
served a number of birds that appeared to be of this form. On 
the same expedition, A. C. Bent also observed the bird in the 
North Pacific, while sailing to the Aleutian Islands, and Rollo 
H. Beck, who was a member of the expedition, took a specimen 
at Kiska Island on June 17. 

Nichols (1927), speaking of seeing this petrel on his trip 
from Seattle to Nome, Alaska, says : 

Seen in the Pacific on 1 day only, August 5, when midway between the 
islands and the west coast, noon position 53° 36' N., 145° 37' W. They were 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 45 

frequent all day, singly and in small groups, a conservative estimate for 
the total number, 30. 

We did not see this bird on our expeditions, but at Attu Island 
the native chief insisted that there was a third petrel, calling it 
le-vi-dre-che. He said that it is gray in color — if we understood 
him correctly — but that it is distinct from the forked-tailed and 
the Leach's petrels, with which he was also familiar and for 
which he had names. 

The chief provided some native guides, and we visited two 
small islets. Cooper and Gibson Islands, which are adjacent to 
Attu. We searched diligently and hopefully, but we found only 
the other two species. Inasmuch as it is known to breed in New 
Zealand, and because the egg dates are "December 24 to January 
7" according to Bent, one would not expect to find it nesting in 
the Aleutians. But in view of the Attu chief's confidence in the 
matter, there was a possibility that the bird may enter burrows 
to roost. 

Pferodroma coo/://; Cook's Petrel 
Pferodroma cook'ii or'ienfalis 

A. W. Anthony (1934, p. 77) recorded a specimen of this petrel, 
postively identified, which was taken at Adak Island by members 
of the crew of the U. S. S. Kingfisher. It was 1 of 2 such birds 
that came aboard the ship, and the specimen saved had been 
sent to the San Diego Zoo in California, where it was placed in 
the collection of Louis B. Bishop. 

Fulmarus glacialis: Fulmar 
Fulmarus glacialis rodgersii 

Attu : Kil-u-gho6-kin 

Atka: Ah-ga-luch 

Probably Russian Commander Islands: Glupisch (Stejneger) 

The Atka name for fulmar should not be confused with their 
name for killer whale, which was recorded as A-ga-loh, with 
shorter syllables. 

Fulmars are common in the North Pacific and Bering Sea 
region, though in varying numbers. They are seen on the Gulf 
of Alaska, in small groups, in company with the black-footed 
albatross and shearwaters. We found them in Shelikof Strait, 
between Kodiak-Afognak Islands and the mainland, and farther 
west near the Shumagins. A few were seen north of the Alaska 
Peninsula, east to Bristol Bay, and north to Nunivak and Nelson 
Islands. They were more abundant among the eastern Aleutians, 



46 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

and they are fairly prevalent throughout the chain, some having 
been noted at Wrangell Cape on the west side of Attu Island. 

Arnold (1948) recorded a concentration of about 38,000 fulmars 
in Unimak Pass on June 9, 1944. Throughout the Aleutian 
district, the dark color phase predominates, though some con- 
centrations of the light phase were encountered. On May 21, 
1936, Cecil Williams estimated that 50 percent of the fulmars 
seen in Unimak Pass were light colored, and that most of those 
sighted on the north side of Unimak Island were whitish, though 
there were entire bands of dark ones. 

It is interesting to recall that Nichols (1927) found "the dark 
phase to light phase about as 99 to 1" in Shelikof Strait; else- 
where in the Pacific, it was about 9 to 1 ; near Unimak Pass, 
the pale and dark birds "were in .about equal numbers ;" and in 
Bering Sea, they were almost all pale. 

Nesting 

Nesting fulmar colonies have been known in the Bering Sea 
region, on the Pribilofs, where light-colored birds are in the 
majority; on the Siberian coast opposite St. Lawrence Island; 
and at Copper Island, where light-colored birds are rare. Nesting 
places in the Aleutians had been suspected, but they had not been 
found. It was gratifying, therefore, to find several such colonies 
in 1936 and 1937, and in 1940 Gabrielson found other colonies 
east of the Aleutian chain. 

In 1940 Dr. Gabrielson learned that Sea Otter Island, near 
Afognak, supports a colony of fulmars. This nesting colony would 
account for the fulmar flocks so often observed in Shelikof 
Strait. In the same year, on June 18, he found nesting colonies 
in the Semidi Islands, and says, "The enormous concentrations of 
Fulmars, for example, was a great surprise to us, as the colony 
apparently has developed since the previous exploration." In 
the Semidi group, he found the fulmars in "huge colonies" on 
Aghik and Choweit Islands, and there were fulmars also on 
Kateekuk, Anowik, Kiliktagik, and Suklik Islands. They were 
mostly of the dark color phase, with an occasional light-colored 
one. 

In the Aleutians proper, we had found at least four nesting 
places. Among these, Chagulak Island is outstanding. As we 
approached this island on June 15, 1936, it loomed as a peaked 
mountain top rising sheer from the water. It affords nesting 
sites for a variety of sea birds, and a swarm of fulmars swirled 
above its top and milled about its slopes. On the cliffs of the 
south side there were small recesses in the red crumbly rock 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 47 

strata. Each of these recesses was occupied by a fulmar, nesting 
with a single egg. They nested also on other types of ledges 
and on the grassy slopes all the way to the snow line. 

On the same day, we found another large fulmar colony on the 
neighboring island of Amukta. These two islands have the largest 
and the principal nesting colonies of fulmars in the Aleutian 
chain. 

In 1937, we found fulmars nesting in considerable numbers on 
the reddish cliffs of Segula Island (also known as Chugul), 
though this group was not nearly so large as those on Amukta 
and Chagulak. 

Another nesting place is Gareloi Island. Natives had assured 
us that fulmars nest there, though we did not find the birds on 
the first trip. In 1937, however, we found them in limited num- 
bers on the south side of the island. According to the natives, 
their numbers had been decimated, at least on all accessible ledges, 
since the introduction of the blue foxes. The volcanic eruption 
of 1930 also disturbed them; however, if the foxes were re- 
moved, this colony should increase. 

Natives told us that the fulmars nest on outlying rocks at 
Unalga Island, southwest of Gareloi, but we found none there in 
August. They are also reported to nest on Agattu, but we 
failed to find them. 

It is highly probable that a nesting colony will eventually be 
discovered in the general vicinity of Unimak Pass, perhaps on 
some isolated cliff or islet at Unimak Island, because fulmars are 
common in that area. A full schedule and bad weather prevented 
us from exploring that part of the Aleutian chain as thoroughly 
as we wished. 

Food Habits 

We had assumed that fulmars feed to a large extent on plank- 
ton, for we often found them congregated in tide rips, busily 
feeding. And Arnold (1948), observing a huge concentration 
of fulmars and shearwaters in Unimak Pass, found that they — 

evidently were feeding on a type of reddish-orange water life. On occason, 
when one of the birds was hard-pressed to leave the area in the immediate 
vicinity of the ship, it would turn its head down and to one side and 
regurgitate a reddish-orange liquid substance. 

However, records show a great variety in fulmar diet, including 
refuse from ships. 

A. W. Anthony (1895) has pointed out an interesting feeding 
habit of the fulmar off the California coast. Speaking of a large 
jellyfish that is abundant along that coast, he says. 



48 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

I have often seen a fulmar sitting on the water by the side of a jellj 
fish, part of which it had eaten, so filled that it could scarcely move oul 
of the way of the boat. Specimens shot while these Medusae are commor 
I have always found with the stomach filled with these alone, and a hali 
pint of the slimy mass will often run from their mouths when lifted froir 
the water by their feet. 

I think the fulmars enjoy a monopoly of this diet, for I have nevei 
seen other species eating it, nor will gulls, nor any sea birds that I have 
observed, pay any attention to a fulmar that is eating a jelly fish though 
they all claim their share if the food is of a kind that they care for. 

The abundance of the fulmars off this coast would seem to have some 
relation to the abundance of Medusae, since the winter of 1893-94 was 
noted for the almost if not entire absence of fulmars as well as jelly fish 
until some time in late February or March, when both jelly fish and fulmars 
appeared in small numbers. 

This is quoted at some length because it reveals an important 
habit of the fulmar, which also was noted among the Aleutians, 
where a large brown jellyfish, Cyanea capillata, often proved an; 
attraction to fulmars. 

Mortality Factors 

Aside from the danger from foxes on accessible nesting sites, 
the only other natural enemy on which we have information is 
the northern bald eagle. The fulmar appears to furnish an 
important item in the eagle's diet throughout the Aleutian Islands 
as a whole, though murres and other species may dominate the^ 
diet of individual eagle pairs. The drain on the fulmar population 
by eagles could not be significant, in view of the great variety 
of birds on which the eagle preys. More important are man's 
activities, such as the raising of blue foxes. Dead fulmars are 
found on beaches, but, at present, it is hard to estimate the 
results of storms or disease. At any rate, we have several large 
flourishing colonies of fulmars, and those that have been depleted 
should increase again owing to the protection now being given. 

Family HYDROBATIDAE 

Oceanodroma furcafa: Fork-tailed Petrel 
Oceanodroma furcafa furcafa 

Attn : A -la-via- go Ke-kech 

Atka :Ki-ki-tich-noch 

Russian, Commander Islands: Sturmofka (Stejneger) The Atka name is 
applied to both this petrel and to Leach's petrel. 

In 1939, Grinnell and Test separated the forked-tailed petrel 
into two races, designating the southern form O.f. plumbea, 
whose range is said to extend northward to "the Alexander 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 49 

Archipelago, just short of Cross Sound." Sitka birds proved to be 
intermediate, but closer to plumbea. 

In looking over material from the Aleutian district, it is obvious 
that there is variation in the characters used by Grinnell and 
Test — size and color (light or dark) — and many Aleutian speci- 
mens are puzzling in this respect. A specimen from Kodiak is 
similar to the Aleutian group. One from Ugashik, on the north 
side of Alaska Peninsula, obviously is furcata. This also is true 
for two from Nushagak. But a series from Belkofski, on the 
Alaska Peninsula, are darker than other Aleutian specimens — 
fully as dark as a series from Forrester Island, Stephen's Passage, 
Sitka, and Icy Strait — though the Belkofski petrels are larger. 
Since we are dealing with average characters, it is clear that 
the birds from the Aleutian district, from Kodiak and Nushagak 
west to Attu Island, should be called O.f. furcata. 

It is of interest to note that birds from the Commander Islands 
and Kamchatka are paler and (on the average) larger than 
those from the Aleutian Islands. A few from the Aleutians are 
squally large and pale, and one from as far east as Nushagak 
is identical with many of the Kamchatka birds. These birds bear 
Dut Grinnell and Test's statement of an increase in size and a 
3olor transition from dark to pale, in the populations from south 
to north and west. Probably, we should consider the Siberian 
3irds as the culmination of this trend toward larger size and 
paler coloration, and, for the present at least, we should class 
:hem with furcata of the Aleutian district. 

The forked-tailed petrel ranges widely over the North Pacific 
md Bering Sea and is the dominant species among petrels there. 
From May 29 to June 4, 1911, Wetmore found these birds common 
)n the Gulf of Alaska. Friedmann (1935) records several speci- 
mens and eggs from Kodiak. Specimens have been taken at 
N'ushagak by Hanna and Johnson and have been taken at Ugashik 
3y McKay. We observed them in the Shumagin Islands and 
'ound them to be abundant throughout the Aleutian chain. 
?tejneger (1887) found them nesting in the Commander Islands. 

Nesting of this species in Bering Sea proper has not been re- 
Dorted, though the bird occurs far northward. Nelson (1887) 
'ound the birds off Nunivak Island in June 1877, but he speaks 
)f them chiefly as autumnal visitors, as far north as St. Lawrence 
[sland, Bering Strait, and Plover Bay, Siberia. Two specimens 
vere secured from Kotzebue Sound. According to Nelson, they 
ingered in Bering Sea even after the formation of ice, and the 
Eskimos told him that they were captured on the ice, near air 



50 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

holes, in a weakened condition. He saj^s that they were founc 
on the lower Yukon, and that one was found about 75 miles 
up the Tanana River, near an air hole in the ice, late in November, 
These petrels are also recorded from the Pribilofs (Preble anc 
McAtee 1923). Cahn (1947) reports from Unalaska: "Seen 
abundantly in the fall and winter far out in Bering Sea." 

The Aleutians must be considered to be the stronghold of thisi 
petrel. We found them on most of the islands. Experience taughti 
us that wherever we found petrel wings left by blue foxes, ori 
petrel remains in fox droppings, it was safe to assume that the 
birds nested on the island. 

The following islands were specifically noted as being nesting 
places for these petrels: Sanak group. Egg Island (in Akutanl 
Pass), Uliaga, Kagamil, Chuginadak, Herbert, Yunaska, ChagulakJ 
Amukta, Amlia (reported by natives), Atka (on Korovin 
Volcano, reported by L. M. Turner), Salt (until destroyed by 
fox raising), Kasatochi, Igitkin, Ulak, Bobrof, Tanaga, Ilak (re- 
ported by natives, but now destroyed by blue foxes), Gareloi, 
Semisopochnoi, Little Sitkin, Chugul (reported by natives), Kiska, 
Buldir, Agattu, and Attu, It is almost certain that the birds nest 
also on most of the other islands — on the Shumagins, and probably 
on other islands off the Alaska Peninsula. This list serves to 
show the uniform distribution of these birds. 

It is well known that the forked-tailed petrel nests underground.^ 
It may make its own burrow, often as a side tunnel from the 
wall of the tufted puffin's burrow, or it may nest in natural 
cavities, such as those found in lava beds. Amukta Island fur- 
nishes a typical example, where the moss-covered lava formation 
had neat round holes through the vegetative crust leading in to 
irregular cavities beneath. 

Food Habits 

The forked-tailed petrel is said to skim the oil from the surface 
of the water near a wounded seal or whale, but we do not have 
extensive data on its food habits. Preble and McAtee (1923) 
record one stomach from the Pribilofs that contained a few 
fish bones. Scheffer made interesting observations at Kagamil 
Island on August 30, 1938, when about 25 forked-tailed petrels 
and 3 Leach's petrels were picked up on the deck of the ship. 
Six piles of regurgitated material on the deck contained broken 
remains of small fish — the largest was about the size of a man's 
little finger 

Mortality Factors 

The principal enemy of the petrel, so far as our observations) 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 51 

go, is the introduced blue fox. Petrels appeared to be especially 
palatable to these carnivores, or perhaps they are easy prey at 
their burrows. On Salt Island the species had been entirely elimi- 
nated by foxes, and this appeared to be true of Ilak, also. 

Petrels are attracted by ship's lights at night. They flock 
around the ship, chirping and chattering incessantly, striking 
the rigging and fluttering about the deck. Often, they get into 
staterooms, the galley, or other portions of the ship, and some- 
times in the excitement an egg or two may be dropped on the 
deck. Joseph Mailliard (1898), writing of the petrels on St. 
Lazaria Island, Sitka Bay, quotes Grinnell as saying that it was 
impossible to keep a fire alight in the middle of the night be- 
cause the petrels flew into it in such numbers that they ex- 
tinguished it. 

Oceonodroma leucorhoa: Leach's Petrel 

Oceanodroma leucorhoa leucorhoa 

Attn : Ke-Kech 

Atka : Ki-ki-tich-noch 

Russian, Commander Islands: Malinka tschornaja sUirmofka (Stejneger) 

The Russian name given by Stejneger means a small black 
petrel. Leach's petrel has a more southern distribution than O. 
furcata and does not range far into Bering Sea. Though it 
nests throughout the Aleutian chain and on Copper Island, ac- 
cording to Stejneger, it is much less abundant than the other 
species. Farther south, it becomes much more abundant. Bent 
(1922) states that on St. Lazaria Island, at Sitka Bay of Baranof 
Island, Grinnell and Mailliard estimated that 0. leucorhoa out- 
numbered O. furcata four to one. They also are extremely nu- 
merous on Forrester Island. 

On the Aleutians we found this ratio reversed. Among 
the remains of petrels left by foxes, those of O. leucorhoa were 
scarce. The petrel colonies of any size were O. furcata. When 
flocks of petrels fluttered about the ship's lights at night they 
were usually furcata, though sometimes there were a few leu- 
corhoa. It is interesting to note that in 1941 Gabrielson found 
Leach's petrel outnumbering furcata at Kasatochi Island. There 
may have been a nesting colony nearby. 

Thus, we find that these two species more or less inteiTningle 
in their ranges. But 0. furcata has a more northerly center of 
abundance, and ranges farther north, while 0. leucorhoa is more 
concentrated somewhat farther south, and does not reach far 
into Bering Sea. 



52 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

The natives declared that these petrels (probably referring to 
both species) nest "everywhere," but the following islands are 
the only ones for which we have precise evidence of nesting of 
Leach's petrel: Sanak Island group. Egg Island (McGregor, 
1906), Amchitka (reported by Dall), Davidof, Kiska, Buldir, 
Agattu, and Attu. Dr. Gabrielson noted evidence of their presence 
on Amatuli Island, of the Barren Islands group, June 13, 1940. 
They were especially numerous on Buldir Island, where we found 
the greatest number of nesting burrows. No doubt they occur 
on a great many other islands, as the natives intimated, but 
probably in such small numbers that they are not easily detected. 

On Buldir Island, where we had the best opportunity for 
examination, their burrows were found in the sandy banks above 
the beach and along a stream, as well as on high grassy slopes, 
well up on the higher part of the mountain. The burrows extended 
about arm's length, often with a very small entrance so that it 
was difficult to insert the hand, but generally there was an en- 
larged chamber at the end. In sandy soil, the burrows were 
larger in diameter; in sod, they were much smaller and the en- 
trance was more obscure. There was a flat nest of dried grass in 
the end chamber. Usually, there was a single bird in the nest, 
but in at least 1 burrow there were 2 birds. 

Food Habits 

Leach's petrels have been observed following whales for food 
fragments, and they have been seen picking fish refuse in the 
vicinity of fishing boats. We obtained no additional data on their 
food habits. 

Mortality Factors 

Foxes prey on these birds where nesting colonies are available. 

Family PHALACROCORACIDAE 

Phalacrocorax aur'ifus: Double-cres+ed Cormorant 
Phalacrocorax aur'ifus cinc'inatus 

Attu: Ki'ich-tirch 
Unalaska: T'chung-ahh 

Of the three species of cormorants nesting throughout the 
Alaska Peninsula and Aleutian Islands, the double-crested is 
much less common than the pelagic or the red-faced. It nests 
on Kodiak Island. Cahalane (1943) reported it to be abundant in 
the Kodiak-Afognak group, and Gabrielson noted a few at Whale 
Island and four in Uyak Bay. Probably it nests in the Barren 
Islands also. Several were seen at Ushagat Island of this group. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 53 

on May 11, 1936. On May 7, 1936, we had seen several of these 
birds on Chisik Island, in Tuxedni Bay, Cook Inlet, obviously 
preparing to nest. C. J. Rhode noted a colony of about 50 
cormorants on islands of Skilak Lake, Kenai Peninsula. Identifica- 
tion was uncertain, but it is probable that these were double- 
crested cormorants. 

Osgood (1904) recorded this cormorant nesting on islands of 
Iliamna Lake. He saw them flying up and down Nogheling 
River, and several were seen on Lake Clark (one specimen taken) ; 
however, he believed few, if any, nested on that lake. He states 
that from the Mulchatna River near the mouth of the Tikchick to 
Nushagak these cormorants were seen daily, but not in great 
numbers. Again, he saw several on Becharof Lake, October 4 to 7, 
1902. 

Cahalane (1944) observed coraiorants, "presumably all of this 
species" (double-crested), rather commonly in the Katmai region 
of the Alaska Peninsula in 1940, and he reported them in 
September on Naknek River, on Naknek Lake, at the south shore 
of Iliuk Arm, at the mouth of Savanoski River, on Brooks River, 
and on Brooks Lake. By September 27 and 28, they were 
relatively scarce on Naknek Lake and River. On the Pacific side, 
he reported them to be much more numerous and recorded them 
as "common to abundant" in the bays of the Katmai coast, 
October 4 to 7. 

On July 23, 1940, Gabrielson noted 1 or 2 double-crested 
cormorants on the Kvichak River, and on July 25, he reported 
2 more on Iliamna Lake. In the same season, he also noted 
them at the Semidi Islands. 

On August 29, 1936, we saw at least two double-crested cor- 
morants near Simeonof Island, in the Shumagin group. They 
probably nest among those islands. On May 16 and 17, 1928, 
F. L. Jaques (1930) saw cormorants, which he thought to be this 
species, near the Shumagins and near Belkofski. 

The largest colony of this connorant was found by the writer 
in 1925, in Isanotski Strait, at the extreme tip of Alaska Penin- 
sula. On the larger of the two Isanotski Islands, which was 
not over 150 yards long, there were at least 25 nests in a close 
group on a grassy slope. On July 27 the nests contained from 
2 to 5 eggs, but usually there were 4 or 5. A specimen was 
collected. It is interesting to note that Beals and Longworth, 
on June 10, 1941, 16 years later, stated in their field report that 
"Small colony of 50 or more birds nesting on the most northerly 
of the two Isanotski Islands." 



54 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Donald H. Stevenson, who accompanied me' in 1925, described 
cormorants nesting on a small island in Swanson Lagoon, on 
the north part of Unimak Island. His description, and the 
location of the nesting site, leaves little doubt that a colony of 
double-crested cormorants were nesting in Swanson Lagoon at 
that time. 

Among the eastern Aleutians we found several nesting groups, 
though they were small in number. On July 16, 1936, we observed 
five of these cormorants perched on low rocks at one of the 
little islands (which we designated "Puffin Island") in Trident 
Bay, Akun Island. On June 7, we had seen about a dozen at the 
west end of Umnak Island; a specimen was taken here. Nesting 
was not actually observed in these instances. But on June 8, 
we found several of these cormorants nesting on Kagamil Island. 
Some also were found on Uliaga Island near by, and two small 
colonies were found on Carlisle Island. Seven nests, and a 
number of birds, were located on Herbert Island. The nests 
among the Islands of the Four Mountains were on ledges of 
sheer cliffs and in the walls of high caverns (sometimes very 
high), which was in great contrast to the nesting on the low 
Isanotski Islands, observed in 1925. In some cases, these cor- 
morants were nesting in close proximity to red-faced and pelagic 
coiTnorants. 

We observed no double-crested cormorants west of these islands. 
The natives of Atka assured us that this cormorant does not 
occur in that part of the Aleutians, We are fairly confident that 
today this species does not nest west of the Islands of the Four 
Mountains. Yet, the Chief of Attu appeared to be familiar 
with this bird; he gave us the native name and declared that 
formerly it was abundant, though it has become scarce in recent 
years. Austin H. Clark (1910), writing of his expedition of 
1906, said "I have a note of a few [double-crested cormorants] 
being seen in Unalga Pass near Unalaska, and I found them 
at Atka, Attu, and Agattu." Turner (1885) also, writing of the 
Near Islands, reported double-crested cormorants to be abundant, 
resident, and breeding. 

Clearly there has been a drastic change in distribution of 
this species since about 1906, (the time of Clark's notes). The 
cause of this restriction of range has not been determined. 

The species P. auritus as a whole is quite versatile in nesting 
habits. Many of the prominent nesting sites in the northern 
part of the continental range are on low islands. Elsewhere, 
cormorants nest in trees and on cliffs as well as on low islands. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 55 

Therefore, there must be adaptability in the species. The in- 
troduction of blue foxes on a large scale in the Aleutians dis- 
couraged the low-ground type of nesting; therefore, the birds 
nested in the cliffs in greater numbers. This change in nesting 
locale of the double-crested cormorant may have resulted in 
competition with the more agile red-faced and pelagic cormorants. 
It may be pointed out that in the absence of blue foxes, the 
double-crested cormorant would nest on low flat ground, away 
from cliffs — a habitat not usually desired by the other two 
species. What the human factor might have been in the 
ecological picture is hard to say, but, under conditions prevailing 
in recent years, cormorants could hardly succeed in nesting except 
on well-protected cliffs. 

Whatever the factors, it is a fact that the double-crested cor- 
morant has virtually disappeared from the Aleutians west of 
the group known as the Islands of the Four Mountains. 

Phalacrocorax pelag'icus: Pelagic Cormorant 
Phalacrocorax pelag'icus pelag'icus 

Attu: Kri-li-ti-kch or Kri-li-ti-kich; TiV-i-toch (1-year-old young) 
Atka: AgW-i-uh (possibly referring to any cormorant species) 

Agayi'ix (Jochelson) 
Russian, Commander Islands: Malmky Uril (Stejneger) 

This is the most abundant cormorant in the North Pacific and 
Bering Sea region. We found them in the Barren Islands, the Ko- 
diak-Afognak group, Chisik Island in Cook Inlet, Sutwik Island, 
Chignik Bay, Shumagin Islands, and along the north side of 
Alaska Peninsula to Bristol Bay. 

Hine (1919) said, "Colonies of this cormorant nested on the 
shelves of the sea wall along Katmai and Kashvik bays during 
the 1919 season." 

Osgood (1904) found these birds on Becharof Lake. Gabriel- 
son found them to be common in the Semidi Islands. 

We found the pelagic cormorant to be numerous throughout the 
Aleutian Islands, and they are common in the Commander Islands 
(Stejneger 1885). This bird has a more northerly distribution 
than other species, for it occurs on both sides of Bering Sea and 
as far as the Arctic coast. 

Ordinarily, this species was the more common in the waters 
about the Aleutian Islands, though among the birds actually 
nesting P. urile outnumbered P. pelagicus. There appeared to be 
a considerable number of nonbreeding P. pelagicus, in subadult 
plumages. In some cases we found no nests, though the birds 
were present in considerable numbers. 



56 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Usually, the nests were placed on ledges of steep cliffs, thougl- 
sometimes they were on overhanging walls of caverns well ou1 
of reach of the surging water. Only once was a differeni 
nesting site noted. In 1925, I found an unusual nesting situatior 
near Izembek Bay at the west end of Alaska Peninsula. On Gler 
Island, at this bay, on May 20, there was a considerable colonj 
of P. p. pelagicus in a compact group on a low point. Some oi 
the birds had white flank patches at this time. On June 1, Steven- 
son reported that the birds were sitting on nests. On June 26, 
about 30 nests were counted, but there were no eggs. On July 27, 
the cormorants were still plentiful at Glen Island, but there had 
been no success in nesting. 

The interesting fact in this instance is the unusual nesting 
site, which was a low sandy point above tidewater. While there 
had been a few birds in breeding plumage, the majority seemed 
to be immature birds. Amak Island, with immense cliffs occupied 
by large numbers of P. urile and only a few P. p. pelagicus, is 
about 12 to 14 miles out to sea. It was not determined whether 
this was an abortive attempt at nesting on Glen Island by cor- 
morants crowded off Amak Island, or whether the birds were 
immature. Throughout the Aleutian chain, both P. urile and P. p. 
pelagicus are found nesting on the same cliffs. 

While the pelagic cormorant is a salt-water bird almost ex- 
clusively, Osgood found it on Becharof Lake, as noted previously, 
and natives of Atka Island said these birds will go to the lakes of 
Amchitka Island in winter. 

Cahn (1947) reports them at Unalaska as "abundant every- 
where along the rocky shores from September to May," and 
Taber (1946) says the species was present at Adak in winter, 
where they continuously lived in salt water, never in fresh-water 
lakes. Sutton and Wilson (1946) found them at Attu in the 
summer and in the winter. 

Food Habits 

The food of this cormorant is assumed to be fish, but, according 
to Preble and McAfee (1923), a considerable percentage consists 
of various Crustacea, at least in the Pribilofs. Sutton and Wilson, 
at Attu, obtained a specimen on February 28, and report: "Its 
stomach and crop were packed with small sculpins which it had 
caught in water about 15 feet deep along the west side of Casco 
Cove." 



I 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 57 

Pbalacrocorax urile: Red-faced Cormorant 

Attu: Ing-a-tohh or Ing-a-torh 
Atka: Ing-a-tohh 
Russian, Commander Islands: Bolschoj Uril (Stejneger) 

Walrus Island, in Bering Sea, has been considered the chief 
nesting place of the red-faced cormorant in Alaskan waters. It 
was a surprise, therefore, to find that the red-faced cormorant 
is the dominant nesting cormorant throughout the Aleutian chain. 




*/ ■ *.' .'' V ^*M 



Figure 24. — Red-faced cormorant. 



58 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

On May 16, 1936, while anchored at Unga, in the Shumagin 
Islands, we discovered a good-sized colony of red-faced cor- 
morants on the precipitous cliffs of a rocky point. There were 
about 300 birds beginning to nest. Some of them were carrying 
nesting material; many had no nest at all. In our experience, 
this is the easternmost colony of this species. 

In 1925, I found a colony on Amak Island, a small conelike 
island off the west end of Alaska Peninsula in Bering Sea. It 
was estimated that there were between 4,000 and 5,000 birds 
nesting on the high cliffs — by far the largest colony known south 
of the Pribilofs. When visited again in 1936, these birds were 
still nesting on the same cliffs, many of them carrying nesting 
material (on May 31). On May 22, 1928, Jaques (1930) ob- 
served this species near Port Moller. 

Bogoslof Island is also occupied by .od-faced cormorants, 
mostly in the form of individuals scattered among a large number 
of murres. 

While most of the Aleutian colonies of the red-faced cormorants 
are very small, there are a few sizable concentrations. On Adokt 
Island, one of the Baby Islands group in Unimak Pass, we esti- 
mated 500 nests. There were also large concentrations on certain 
islets in the Bay of Islands of Adak Island. But in many instances 
there were small groups, sometimes six or less. 

Following are the islands, east to west, on which the red- 
faced cormorant was found nesting: Unga (Shumagins), Amak, 
Adokt and Excelsior of the Baby Islands group. Egg (probably), 
Poa, Tangik, Bogoslof, Ananiuliak near Umnak (not certain), 
Uliaga, Kagamil, Carlisle, Herbert, Yunaska, Chagulak, Amukta, 
Seguam, Ulak, Kasatochi, Igitkin (probably), Adak, Gareloi, 
West Unalga, Semisopochnoi, Amchitka, Little Sitkin, Davidof, 
Kwhostof, Kiska, Agattu, Semichi, and Attn. 

These birds were identified at other islands, though nests were 
not actually obseiwed. This gives the red-faced cormorant a 
fairly uniform distribution as a nesting bird from Unga and 
Amak Islands, and Port Moller, all the way to Attn, and they 
are known to nest still farther west, in the Commander Islands. 
The red-faced cormorant also winters in the Aleutian waters. 

Mortality Factors 

Birds which prey on the cormorants are the bald eagle, pere- 
grine falcon, and glaucous-winged gull. In a study of the food 
habits of the bald eagle in the Aleutians it was found that 
cormorants had been taken for food frequently; however, the 
species of cormorants was not determined. It seemed likely 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 59 

that most of the remains that were examined were those of the 
pelagic cormorant, but no doubt the red-faced cormorant would 
be taken also where it is available. We found no remains of the 
double-crested species, but that form is relatively scarce. 

On Amak Island, several carcasses of red-faced cormorants, 
probably killed by peregrine falcons that nested nearby, were 
found beneath the nesting ledges. Falcons are indicated as the 
predator because eagles would have carried their prey away to 
their feeding places. 

Glaucous-winged gulls persistently seek the cormorant's eggs, 
and they are most successful when the parents are frightened 
off the nests, thus exposing the eggs to predation. 

Family ARDEIDAE 

Ardea herod'ias: Great Blue Heron 
Ardea herodias fann'ini 

The heron seldom enters the territory with which we are 
concerned. Osgood (1901) reports, "A great blue heron was 
seen at Hope by E. Heller." In the same general vicinity, May 8, 
1936, the first mate of our ship reported seeing a heron early 
in the morning, as we were approaching Anchorage. It was 
recorded at Portage Bay (Nelson, 1887). So far as we know, 
then, this heron reaches the western part of its range at about 
the head of Cook Inlet. 

Family ANATIDAE 
Olor columbianus: Whistling Swan 

Attu : Kon-kirch 

Qumqix (Jochelson — dialect not given) 
Atka: Ko-kin-yeh (or ko-kin-e-rech ?) 

Whistling swans nest on Kodiak Island and in suitable areas 
along the Alaska Peninsula. Osgood (1904) specifically mentions 
Swan Lake, Chulitna River, and "upper waters of the Nushagak 
system, and near the Ugaguk River and Becharof Lake." 

Einarsen (1922) observed a swan with four young near 
Ugashik on June 26, 1922, and Jaques (1930) observed eight 
swans near Port Moller from May 24 to June 14, 1928. 

While in the Katmai region in September 1940, Cahalane 
(1944) observed swans "from Kwichak River to Naknek," on 
tundra pools, on Lake Grosvenor, mouth of Savanoski River, 
between Iliuk Arm and Mount Katolinat and above New Savanoski 
Village, 



60 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

On July 19, 1940, Gabrielson noted three pairs of swans be- 
tween Naknek and Brooks Lake, and on July 21, while flying 
over the country from Becharof Lake to Egagik, by way of Ruth, 
Ugashik, and Mother Goose Lakes, he saw numerous pairs of 
swans with 1 to 5 young among the many tundra pools and lakes 
of this area. 

According to local residents (1936) , swans nest on small islands 
in ponds near Ugashik River, and up the river from Nelson 
Lagoon, At Chignik I was informed that swans nest in Black 
Lake, the "second lake up Chignik River." Gabrielson was told 
that they nest in the King Cove-Cold Bay area. 

In 1925, though none were found nesting, a swan was seen 
flying over Hazen Point in Izembek Bay on June 13, and on 
July 23 there was a group of three in a lake near the sand dunes 
there. In the same year, on April 29 and on several subsequent 
dates, two were seen at Urilia Bay, on Unimak Island, and a 
trapper said that he saw a few swans in that locality each year. 

More recently, we have precise information that swans nest 
on Unimak Island, for in 1936 we obtained an egg, which, we were 
told, had rolled out of a nest on Ikatan Flats. The following year 
we learned that a pair had returned to the same flats. 

In 1941, Beals and Longworth noted several swans at Unimak 
Island, and they reported that on August 31 a trapper observed 
3 pairs near Swanson Lagoon — eacli pair with 2 young. It is 
also reported that a pair nested on Ikatan Flats in 1940. 

Chase Littlejohn (manuscript notes) says, "Only a few seen 
at Morzhovoi Bay, where I know at least one pair reared their 
young in 1879. I found them with their parents in a lake still 
unable to fly on August 29." 

Swans are not known to nest west of Unimak Island. 

Dall (1874) reports the killing of three swans at Sanak Island 
in September 1872 by a sea-otter hunter, who said they were 
not uncommon there in the fall. 

Apparently, swans have not been considered a part of the 
fauna of the Aleutian chain proper. Dall stated that they did not 
occur there, though Turner said a few wintered on Attu Island. 
It is possible that conditions have changed, for there is. ample 
evidence that swans occur on many of the islands — at least in 
winter. At Atka Island, the natives assured us that swans winter 
in the Aleutians, and they specifically mentioned Kanaga and 
Amchitka, where swans had been observed on the lakes. On 
Amchitka, we found swan remains among the native buildings. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 61 

and we learned that about 20 had been killed there the previous 
winter. 

We also found swan wings on Semisopochnoi Island, thus show- 
ing that some had been killed there in winter by native fox 
trappers. Friedmann (1937) found four swan bones in midden 
material from Little Kiska. 

The native chief on Attn Island assured us that swans winter 
abundantly among the lakes on the south side of that island, 
occurring in flocks of "eight, ten, to twenty-six." In 1924, the 
swans were said to have walked among the houses of the village, 
and, in 1932, "hundreds and hundreds" were seen among the 
lakes on the south side. Now, they occur only in small numbers. 
According to the natives, the swans arrive at Attn Island late 
in October, and they leave late in April. 

Mortality Factors 

It is probable that man has been the greatest enemy of the 
swan, for, under ordinary circumstances, the swan probably 
is able to protect itself against natural enemies. A trapper at 
Port Moller, on the Alaska Peninsula, told me that he had 
seen a swan defend itself against a red fox, and he doubted 
whether foxes were much of a hazard. 

Olor buccinator: Trumpeter Swan 

Quoting Friedmann (1937) on Kodiak Island: "A synsacrum 
and 2 tarsometatarsi were found in the superficial levels and 
another tarsometatarsus in the intermediate depths in 1935; in 
1936 a metacarpal and the head of a humerus were collected." 

There are no other records. 

Branfa canadensis: Canada Goose 
Branfa canadensis leucopareia 
Branfa canadensis minima 

Attu : Legch 

Atka: Luch or lug-ach, or lagix (Jochelson) 

Resident whites: layid geese 

The white-cheeked geese were formerly common migrants 
throughout the Aleutian Islands area and nested on many of 
the islands. These populations now (1936, 1937, and 1938) have 
been universally reduced. 

The forms of the white-cheeked groups of geese that nest in 
the Aleutian district is a question that has led to endless confusion. 
Our latest findings show that leucopareia and minima are so in- 
extricably associated throughout the Aleutians that it is desir- 



62 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

able to discuss them together. As far as we were able to learn, 
the Aleuts have only one name for this general type of goose. 
However, the Eskimos at Hooper Bay distinguish between these 
two forms, and they have a distinctive name for each form. 

It should be noted here that in much of the previous work with 
these birds, the name hutchinsi was used to identify the form 
that we now call leucopareia (A.O.U. Check List) ; and this 
change has resulted in considerable confusion and misunderstand- 
ing of the literature. 

It is certain that 2 forms of the white-cheeked geese nest in the 
Aleutians, but there is a question about the taxonomic rank to 
accord these 2 forms. As to considering them races of the 
same species, I agree with Bent (1925) that "Both the cackling 
goose [minima] and the Hutchins goose [leucopareia'] are said to 
breed on the Aleutian Islands, but it seems hardly likely that these 
two subspecies should occupy the same breeding range." 

The situation we find here supports Taverner's conclusion 
(1931) and the findings of Aldrich (1946) that we have two 
species. Aldrich has proposed that the smaller species includes 
three subspecies: true hutchinsii (not leucopareia), minima, and 
asiatica, and that B. canadensis includes the other subspecies of 
this group. On June 23, 1911, a female was collected on Attu 
Island by R. H. Beck, which Bent (1925) recorded as minima. On 
June 13, 1937, John H. Steenis collected a male goose of this 
group on Agattu Island. These specimens were studied by Aldrich, 
and he agreed that the Attu specimen was true minima, and that 
the one from Agattu was equally typical of leucopareia. 

At Hooper Bay (south of Yukon Delta), we found the Alaskan 
cackling goose {minima) nesting nearest the sea, while the lesser 
Canada goose {leucopareia) nested farther inland, though the 
two nesting ranges were adjacent. Two groups of Eskimos, an 
inland group and a coastal group, with slightly differing dialects, 
both recognized these two species of geese as different and had 
a name for each. With two geese populations nesting in such 
close proximity, without space for "intergrades," it would be 
illogical to consider them subspecies, aside from the facts shown 
by examination of characters. In the Aleutian district, these two 
species occupy ranges similar to the kinds in the Hooper Bay 
district. 

Former numbers — Turner found "thousands" of geese on the 
Near Islands, of which Agattu and Semichi were the chief breed- 
ing grounds. They nested on Unaska, Amlia, Atka, Adak, 
Kanaga, Tanaga, Kiska, Buldir, Semichi, and Agattu. On some 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 63 

of these islands, the foxes had forced the birds to nest on offshore 
islets, and on Attn the natives hunted them extensively and do- 
mesticated them, clipping- the wings of young birds. Jochelson 
(1933) says: "Some of them breed on the Four Mountain Is- 
lands." 

Bill Dirks, Atka chief, mentioned as former nesting grounds: 
Tanadak, Unak, and Tanaklak (all near Great Sitkin), as well as 
Amchitka, Ulak, Tanadak (the one near Kavalga), and Kiska. 
He also stated that at one time there had been a native village 
on Buldir, and that the villagers used to pinion young geese 
to prevent them from migrating in the fall so that they would 
be available later in winter. Dirks recalled that his father once 
obtained 50 goslings on Buldir, and brought them to Atka, where 
he fattened them for food. Nelson (1887) saw a flock of do- 
mesticated geese at Unalaska, which had been obtained in the 
western Aleutians. 

We must include Attn in the breeding range, for it was on that 
island that Beck collected the nesting goose examined by Aldrich 
and identified as minima. Evidently a few geese have been able 
to nest in spite of foxes, and in primitive times undoubtedly a 
great many nested there. 

As late as 1911, Wetmore reported at Kiska "Two flocks of 
rather good-sized geese were seen flying over high up June 18. 
One of the officers reported seeing two on an inland lake. None 
were taken." And, again at Atka, he reported, "a flock of geese 
seen flying high up June 13." 

Austin H. Clark (1910) has presented a striking picture of 
geese in abundance : 

This goose is the most abundant bird on Agattu, where it breeds by 
thousands. When we approached the shore we saw a number of geese fly- 
ing about the cliffs and bluffs, and soaring in circles high in air. On landing 
I walked up the beach to the left and soon came to a small stream which 
enters the sea through a gap in the high bluffs, when I saw fifty or more 
of these birds along the bank preening their feathers. From this point I 
walked inland over the rough pasture-like country toward a lake where 
this stream rises. Geese were seen on all sides in great abundance, walking 
about the grassy hillsides in companies of six or eight to a dozen, or flying 
about from one place to another. 

Migration 

As would be expected, in the days when the lesser Canada 
goose and the Alaskan cackling goose flourished there was an east 
and a west migration along the Aleutian chain. In 1925, Donald 
Stevenson, former reservation warden, said that geese from the 
western Aleutians came eastward in the fall to join the throngs 



64 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

concentrated about Isanotski Strait. Atka natives said that geese 
passed eastward at Isanotski in August. 

Chief Ermeloff, of Umnak, said that geese passed there "in the 
fall." Nick Kristensen, who has lived many years on Unimak 
Island, said geese arrived at Urilia Bay before they reached St. 
Catherine Cove, and he wondered, because Urilia Bay lies west 
of St. Catherine Cove, if this meant they "came from Siberia 
somewhere." 

Jochelson (1933) says: "In April it flies to the west, in 
October to the east, resting on the islands." 

It is evident that there was an annual fall migration eastward 
along the Aleutians. When the Aleutian birds arrived at the 
west end of the Alaska Peninsula, they undoubtedly joined the 
throngs of cackling geese that came down from the north. 

On August 14, 1936, we noted six cackling geese' flying south- 
ward over Nunivak Island. We were told that they linger a 
bit on the south side of Nunivak Island before continuing farther 
south. According to local information, they generally arrive at 
Unimak and the Alaska Peninsula about September 1, but they 
do not become numerous until 1 or 2 weeks later. Then, they 
assemble in surprising numbers and congregate at Urilia Bay, 
Swanson Lagoon, and St. Catherine Cove, all on Unimak Island, 
and at Izembek Bay, head of Morzhovoi Bay, Nelson Lagoon, 
and Port Moller on the Peninsula. In 1942, Gabrielson reported 
the first fall migrant at Izembek Bay as early as "late in July." 

In 1925, accounts of the coming of the geese in "countless 
thousands" and "millions" testified to unusual concentrations, and 
it is safe to say that this area is the prinicpal gathering place for 
geese nesting along the shore of Bering Sea northward, as 
well as those from the Aleutians proper. The emperor goose and 
the 2 forms of the Canada goose all assemble here — of the two, 
the Canada geese are in the majority. 

This area seems to be a place where the geese can fatten in 
the fall before continuing to their wintering grounds. They 
are said to feed to some extent on eelgrass; yninima and leuco- 
pareia feed mostly on crowberry (Empetrum nigrum) and other 
berries and spend so much time on the slopes seeking these foods 
that they are known locally as "land geese" — distinguishing them 
from the "beach goose," which is the local name for the emperor 
goose. 

The geese become very fat and leave for the south about 
November 1, though according to some reports it is as late as 
November 15 or 20. Probably, the earlier date is the more 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 65 

usual one. In 1942, according to Gabrielson, the geese departed 
rather suddenly, eastward, on November 20. 

This situation is quite comparable to that on the other side 
of the continent at the head of James Bay, a southern extension 
of Hudson Bay, where the blue geese spend more than 2 months 
fattening, and then continue south about November 1. 

As the lesser Canada geese and the Alaskan cackling geese 
move south, they are noted in many other places, such as 
Metrofane and Mallard Bays in the Chignik area, at Simeonof 
Island in the Shumagins, and the Sanak Island group. Chase 
Littlejohn (manuscript notes) said: "A large number are seen 
annually at Sanakh in the fall where they remain for a short time 
at this season; they are very fat and toothsome . . . They are 
also numerous on the peninsula where they feed entirely on 
berries." 

Our information on the white-cheeked group of geese for 
the more eastern parts of the Alaska Peninsula is, at this time, 
not as complete as the information that we have for other parts 
of this group's range. Osgood (1904) reported a flock of the 
birds at the mouth of the Chulitna River on August 5, 1902. 
Others were seen later on the Mulchatna and were seen between 
the Mulchatna and Nushagak. On July 6, 1925, I saw a pair of 
geese, not specifically identified, on the tide flats of Izembek Bay ; 
it is possible that they were nesting birds. In August 1911, Wet- 
more repeatedly saw "a small goose" on the marshes back of Thin 
Point. On July 28, 1911, he saw another at Morzhovoi Bay and 
saw three more on July 30. All of these, he provisionally identi- 
fied as cackling geese. 

The spring migration is much less noticeable, no doubt be- 
cause the birds are intent on reaching the nesting grounds and 
therefore do not gather in large concentrations, and also because 
their numbers have been greatly reduced since the previous 
autumn. Residents at False Pass were undecided whether the 
geese pass through there in the spring. We were told that they 
also pass through the Chignik area, and at Simeonof Island 
in the Shumagins, and at Sanak Island farther west. At Sanak, 
we learned that the geese gather on the water enclosed by Sanak, 
Elma, and Caton Islands, though they do not linger there in the 
fall. This suggests that in the spring they have completed a 
lengthy flight over the ocean, thus needing both food and rest. 
Chase Littlejohn, writing of the migration at Sanak in 1887-88, 
said, "They used to stop here on their way north a few years ago, 
but they rarely if ever do now, for what reason I do not know." 



66 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Evidently, the geese have resumed the practice in view of our 
information for more recent years. 

Jaques (1930) reported that "Three flocks of what were 
probably cackling geese were migrating to the southwest May 
16, inside the Shumagin Islands." They may have been headed 
for the Aleutians, judging by the direction they were taking. 

It is evident, from information at hand, that the spring migra- 
tion took place in April and part of May, but it was not so 
spectacular as the fall migration. 

Nesting Habits 

Agattu, in the Near Islands group, is the most favorable for 
geese. Most of the island is a lowland, liberally dotted with lakes. 
This makes it easy to understand why such islands as Semichi, 
Amchitka, Tanaga, and Kanaga were at one time a goose para- 
dise — all of them have extensive lowlands with lakes. 

There is another type of nesting habitat which is typified by 
Buldir Island — a domelike island rising sheer from the sea. 
Buldir possesses beaches and a small grassy valley cut by a 
stream. In this valley, where the grasses and sedges are heavy 
and rank, there were no geese. High on the mountain there are 
little depressions, benches, and valleys, which are cut by water 
courses. In this terrain, where the grasses and sedges are short 
and tender, there were geese — even though there is fog much of 
the time. So, on Buldir, the geese apparently have found an en- 
vironment that is suited to them. 

It is interesting to note that these geese do not hesitate to take 
to salt water. One, with two downy young, was seen in a bay at 
Agattu, and another was seen in the water near Chagulak, an 
island at Amukta Pass. The presence of a goose at Chagulak 
suggests another high-mountain habitat, because that island is 
extremely precipitous. 

Present numbers — We have just enumerated the early accounts 
of "thousands" of geese, including Turner's "thousands" in the 
Near Islands, and Clarke's tale of abundance on Agattu. Today, 
the Aleutian district presents a striking example of the rapid de- 
cline of a species; the general opinion is that the fall concentra- 
tions in the False Pass area have greatly declined, apparently 
involving to some extent the geese from the more northerly nest- 
ing grounds. 

We were surprised to find no sign of these geese on the lake- 
dotted flats around the lower part of the Ugashik River, and in 
1937 we observed only a few pairs of geese on Agattu Island — 
probably less than 6 pairs in 4 days of traveling over the island. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 67 

One pair had 2 young, and another had 5 young. In the Semichis, 
we found feathers and a few droppings on Alaid Island. On 
June 15, 1936, the captain and the mate of our ship saw a 
"small goose" of the canadensis type near the shore of Chagulak 
Island, and we found signs of geese on Buldir. However, they 
had disappeared on most of the islands, and our total observa- 
tions indicated that only a few pairs remained in the Aleutians. 
In fact, these geese are so scarce that the migration is no longer 
noticeable, and some of the younger Aleuts didn't seem to know 
about it. When the remaining geese that go to the Aleutians 
are killed, it will be a long time for a migration to become rees- 
tablished, and consequently an extensive habitat for minima and 
leucopareia will lie vacant. 

Causes for decline — The natives, as well as several writers, 
have assumed that the disappearance of these geese from many 
islands was due to the introduction of blue foxes. Undoubtedly, 
this is true, yet on Buidir where there are no foxes, the geese 
are not plentiful. Undoubtedly, another important cause for 
their decline is increased hunting along the migration route and 
on the wintering grounds in the south. 

Administrative action has already been taken to free certain 
favorable islands, including Agattu, from foxes. Further, to 
preserve these geese, it remains for sportsmen to protect the 
birds on the wintering grounds. With such a combination of 
protection, it is still possible to prevent these geese from losing 
their present tenuous hold in the Aleutians, and perhaps it would 
be possible for them to build up to a point where they will be 
safe from extinction. 

Branfa nigricans: Black Brant 

Attu: Agru-ge la-ghe 

Nelson Island Eskimo: NiiV:-hla-ra-nak 

Hooper Bay Eskimo: Nuk-lu-nnK- 

Hooper Bay, a more inland dialect: NuJ:-Iu-gu-nuk 

Nelson (Eskimo dialect) : Luk-hhig-u-huk 

Russian, Yana district: Njemok (Pleske) 

Chukchi: Ncdljuitti (Palmen) 

The black brant is only a migrant in the Aleutian district, but 
it occurs in considerable numbers. In 1936, we w^ere told at 
Port Moller that the brant appear there in April, and we re- 
ceived the same information for the Chignik area. We had seen 
them on northward migration near Seymour Narrows, British 
Columbia, on April 24, and on Queen Charlotte Sound on April 



68 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

25. Donald Stevenson, in 1925, said that he had seen them at 
King Cove "late in April." 

Apparently, the many bays at the western end of Alaska 
Peninsula are favorite gathering places for black brant in migra- 
tion. In 1925, Stevenson and I observed them on Izembek Bay, 
where they were present on May 20 in small flocks, on the water 
and flying from point to point. However, some flocks contained 
as many as 200 birds, and about 5,000 black brant were estimated 
for the entire bay. The following day, at the east end of the 
bay, there were only a few groups. 

Stevenson arrived at this bay on June 2 and found the brant 
to be plentiful. As he passed Applegate Cove, he saw a "swarm" 
of brant up the bay, rising and settling in a funnel-shaped mass. 
There were other groups of 50 to 75 brant flying up the bay, 
some of these joining the large flock. The following day he 
saw more of them, and, in each instance, they seemed inclined 
to move in a northeasterly direction. 

On June 16, we saw a small flock and a single bird; next day 
we saw several small flocks near the outer sand islands. This 
was their last appearance. 

In 1943, Gabrielson found black brant on the Sanak Islands on 
April 30, and the next day, at King Cove, he saw 100, or more, 
heading toward Cold Bay. In 1944, residents at Port Moller 
reported the first spring flight on April 26. 

A. C. Bent (1925) quotes Chase Littlejohn as saying that the 
brant move westward along the Alaska Peninsula, 1 or 2 miles 
offshore, turn into Morzhovoi Bay, and thence go into the Bering 
Sea. This probably outlines the spring migration fairly ac- 
curately. 

While we were at Nunivak Island on August 14, 1936, black 
brant had arrived from the north. Eskimo said that these brant 
remain on the inland lakes of that island for about 2 months, 
or until sometime in October, before continuing south. In the 
meantime, many others have gone farther south, because at Port 
Moller, on August 29, the residents said that the brant were due 
at that time and that they would remain there until about 
November 1, before continuing south. They also return to Izembek 
Bay during their migration. 

Dall reported that black brant were nesting on some of the 
western Aleutians, but Nelson was undoubtedly correct when he 
assumed that these birds must have been small geese of the 
canadensis group. Friedmann (1937) records the following re- 
mains from native middens: One skull and 2 sterna from Little 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 69 

Kiska; 1 humerus from Atka; and 4 humeri from Attu. In 1936, 
the Attu natives told us that black brant appear there occasionally 
in the fall, sometimes in company with the emperor goose. 

Stejneger (1885) reported the species occuring sparingly in 
the Commander Islands in migration. 

Philacfe canagica: Emperor Goose 

Attu: Il-d-ghir-bch 
Atka: Kd-ghu-miing 

Qdmgdn (Jochelson) 

The emperor goose apparently does not commonly nest in 
the Aleutian Islands, nor on the Alaska Peninsula, but at least 
one record of nesting was established. During June 1925, a 
Bureau of Fisheries boat had stopped for a short time at Amak 
Island, on the way to Port Moller. The pilot informed me that 
during that stop at least three pairs of emperor geese were seen. 
On July 10, 1925, during a visit to Amak, I found the remains of 
a young emperor goose in a bald eagle nest. The feet, stomach, 
and numerous pinfeathers were present in the nest, and were 
collected. This appears to be the southernmost nesting locality. 

The Aleutian district is certainly the prmcipal wintering place 
for emperor geese. We noted evidence of such occurrence and 
obtained statements of natives and others who were familiar 
with specific localities, and in 1941 and 1942 Gabrielson noted 
them as plentiful at a number of the islands he visited in the 
winter months. They are reported as spending at least a part 
of the winter as far east as Port Moller, on the north side of the 
peninsula, leaving when the ice formed but returning when the 
water opened again. Some of these geese winter at Urilia Bay 
on Unimak Island and on Izembek Bay; a few geese winter near 
Chignik on the south side of Alaska Peninsula, and some of them 
winter at Simeonof Island in the Shumagins. A banded bird was 
recorded at King Cove in the fall of 1926. 

Turner (1886) makes the sweeping statement that these birds 
winter on the south side of Alaska Peninsula and on offshore 
islands as far east as Cook Inlet. Friedmann recorded bones of 
this goose in all layers of Kodiak middens. Today, they are 
less numerous along those shores, possibly because of the advent 
of white men and an increased kill resulting from modern 
weapons. 

Emperor geese are known to winter in some numbers in the 
Sanak group. We found recent remains at Unalaska, June 3, 
1936, and on Bogoslof Island, June 5, 1936. Eyerdam (1936) 



70 NORTH AMERICAN FAUNA 61, FISH A.ND WILDLIFE SERVICE 

obtained two specimens at Unalaska on June 20 and on August 
7, 1932. These geese are known to winter on the following 
islands: Unimak, Unalaska, Sanak, Umnak, Amukta, Seguam, 
Atka, Adak, Tanaga, Kanaga (abundant), Amchitka, Ulak (longi- 
tude 178^ W.), Ogliuga, Kavalga, Semisopochnoi, Kiska, and 
Attu. The chief of Attu declared that they were in that locality in 
"millions." These are the islands on which we have specific 
information. Undoubtedly, emperor geese occur on many, if not 
all, of the other islands ; almost certainly they occur on Agattu 
and Semichi, for example. 

As may be expected, there are many records of. winter oc- 
currences farther south, in Washington, Oregon, and California. 
These records are numerous enough to suggest that some strag- 
glers find their way into those southern localities quite regularly ; 
however, the regular wintering area is confined to portions of 
Alaska Peninsula, the Shumagin and Sanak Islands, and the 
Aleutian chain. Apparently, they are rare on the west side of 
Bering Sea during the winter. Stejneger (1887) records two 
specimens taken at Bering Island, April 6, 1886. 

The spring migration varies according to the locality and 
the age class. Natives declare that emperor geese leave Attu 
Island in April; Turner (1886) gave the date as the "latter 
part of March." He also stated that after the middle of April 
considerable numbers of geese begin to arrive on the north side 
of Alaska Peninsula, particularly in the neighborhood of Ugashik. 

In 1924, I observed the spring migration at the nesting grounds 
at Hooper Bay. The first migratory wave began about the 
middle of May and continued to the end of the month. There 
was another notable flight about June 5 and 6, which appeared to 
end the migration of breeding birds. Nesting had begun at that 
time. 

A second distinct migration at Hooper Bay took place from 
June 21 to July 1. These were immature birds, probably all 
nonbreeders. 

It was my good fortune to observe the other end of such 
migration in 1925, at Izembek Bay and Unimak Island. On 
April 29, 1925, and for several days following, flocks of emperor 
geese were noted at Urilia Bay, on the north side of Unimak 
Island, many of them flying northeastward. On May 17, a 
flock of 250 was seen standing on an exposed sand bar in St. 
Catherine Cove. On May 20, they were common in Izembek Bay, 
and Donald Stevenson noted a flock of 300 there on June 2. We 
saw a similar-sized flock on June 8, at Mofl'et Cove, where they 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 71 

were noted throughout June in diminishing numbers. The last 
flock was seen on July 7. The time of gradual disappearance 
on Alaska Peninsula corresponds very well with the time of 
the late migration noted at Hooper Bay the previous year. The 
lingering flocks in Izembek Bay were mostly immature birds. 
One bird, which was collected in adult plumage, proved to be a 
nonbreeder. 

A few late occurrences were noted farther west. C. S. Williams 
noted a group of about six emperor geese on Uliaga Island 
on June 8, 1936 ; and a bald eagle's nest on Kavalga Island con- 
tained remains that were fresh enough to indicate a kill in July. 

Apparently, there is an eastward movement of emperor geese 
along the Aleutian chain, and a consequent "piling up" at favorite 
locations on the Alaska Peninsula, until the northern flights are 
well under way. 

The exact reversal of this process occurs in the fall. Some 
time early in September, the emperor geese begin to arrive from 
the north in the vicinity of Izembek Bay. And, according to the 
enthusiastic accounts of local residents, these emperor geese are 
almost as numerous as the cackling geese before the latter de- 
clined in numbers. At Port Moller, emperors are said to arrive 
as early as the latter part of August. They congregate on 
Nelson Lagoon, Izembek Bay, head of Morzhovoi Bay, locally 
in Isanotski Strait, St. Catherine Cove, Swanson Lagoon, and 
Urilia Bay. Most of these geese move westward some time in 
November. Incidentally, Swarth (1934) states that emperor 
geese were present on Nunivak Island, to the north, as late as 
October 29, 1927. The Attn chief said that they arrive at that 
westernmost point in the Aleutians late in October. 

Apparently, in fall migration the immature birds again lag 
behind their elders. According to Swarth, the first arrivals on 
Nunivak Island, observed by Cyril G. Harrold, August 20 to the 
middle of September, were white-headed adults. "On September 
15 the first young birds (dusky headed) were seen and they 
were common thereafter." 

Food Habits 

It is well known that the emperor guose is largely a beach 
feeder; in fact, it has earned the local name "beach goose." Yet, 
it is reported as occasionaly feeding on the berries of the 
tundra, notably Empefrum nigrum. Swarth (1934) sums it up 
thus. 

The emperor geese fed mostly upon the sea shore, but occasional flocks 
were encountered on the tundra, feeding upon berries. The one adult male 



72 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

of the series had its face stained and the throat and entire intestinal tract 
dyed blue from a diet of berries. 

In the spring of 1925, these birds were feeding at low tide on 
tide flats in Izembek Bay. The tide is about an hour later at the 
head of Izembek Bay than at the entrance; the same situation 
exists between the two sides of the long Hazen Point. The 
emperor geese were well aware of this, and when their feeding 
grounds were flooded by the incoming tide they simply flew up 
to Hazen Point, crossed over a few hundred yards to the east side, 
where the flats were still exposed, and continued feeding. The 
narrower parts of this point were favorite flyways. In this area, 
the principal food was thought to be eel grass. On June 14, at 
the margin of a pond, it was noted that the grass was grazed 
off short ; the area was trampled and was littered with droppings. 
However, the stomach of an immature bird found in a bald eagle's 
nest on Amak Island on July 10 contained remains of small crabs. 

Emperor geese are often reported as feeding on some kind of 
kelp in winter. At Kanaga Island, we were told that they feed 
on kelp and the green shoots of Elymus, which, even in winter, 
may be found under the dead vegetation. One informant stated 
that the geese probed into the ground and pulled out the horizon- 
tal rhyzomes of Equisetum. We had noted droppings on Ogliuga 
Island consisting of the herbaceous parts of Equisetum arvense; 
but these droppings could not be positively identified. 

Several observers in the Aleutians reported that emperor geese 
feed extensively on green "sea lettuce," as well as Fucus, and the 
"exposed roots" of Elymus. 

Chase Littlejohn, apparently referring to Sanak Island and 
Morzhovoi Bay, says: "Here they live almost entirely on a 
bright green seaweed, locally known as sea lettuce, but at times 
eating small mussels." 

Cottam and Knappen (1939) have presented a comprehensive 
statement on the food habits of the emperor goose, based on 
analyses of 35 stomachs. Few, if any, of these stomachs were 
obtained in the Aleutian Islands, yet the data agree fairly well 
with observations made in this area. Their findings (based on 
the contents of 33 stomachs) show 91.58 percent vegetable matter, 
and 8.42 percent animal matter. Their findings are further sum- 
marized as follows: Algae, 30.73 percent; eel grass and other 
pond weeds, 13.91 percent; grasses and sedges, 24.94 percent; 
undetermined and miscellaneous plant fiber, 22 percent; bivalve 
mollusks (Pelecypoda), 3.66 percent; crabs and other crustaceans, 
2.18 percent; rodents and fishes, 1.76 percent; and miscellaneous 
animal life, 0.82 percent. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 73 
Mortality Factors 

At Sanak Island, a resident declared that one winter he 
found 15 dead emperor geese on the beach. Although he thought 
that the deaths were caused by the frozen condition of the fresh- 
water creeks, the precise mortality factor here must remain 
unknown. 

Among the natural enemies of the emperor goose is the bald 
eagle. However, there is no evidence that the eagle materially 
affects the goose population. 

Anser albifrons: Whl+e-fronted goose 

Anser albifrons frontalis 

Attu : Kog-a-la-gich 

Russian, latitudes of the Yana: Kasorka (Pleske) 

Bones of white-fronted geese are recorded by Friedmann in 
middens on Kodiak, Amaknak, Little Kiska, and Attu Islands. 
The Attu natives informed us that they have seen these geese 
in September; but they stated that the sightings are rare and 
that these geese do not winter there. Ste.ineger (1887) stated 
that occasionally these geese visit Bering Island in spring migra- 
tion. Turner did not observe this species in the Aleutian Islands. 
In 1925, I learned of a trapper at False Pass who had a white- 
fronted goose in captivity; he had caught the goose at St. Cath- 
erine Cove during the previous autumn. Residents of the area 
stated that this goose is very scarce around the west end of 
Alaska Peninsula. 

The white-fronted goose is a rare migrant in the Aleutian 
chain; therefore, Turner (1886) no doubt was partly right when 
he said, "They probably never visit the islands lying west of the 
mainland, as that region does not contain their particular food 
in sufficient quantity to induce them to visit it." His further 
statement that the Russians at St. Michaels referred to it as the 
^un-dri-na goose, or lowland goose, is further explanation of its 
scarcity in the Aleutians, where most of the land is rugged. 
Farther east along the Alaska Peninsula, however, suitable 
ground is available, and we found nesting birds on the tide flats 
at Ugashik River. On May 27-29, 1936, at least six pairs were 
noted in that area. 

Osgood (1904) records that he saw these birds at the base of 
Alaska Peninsula in 1902 ; he frequently saw them on the Chulitna 
River in early August, saw one on the Mulchatna River on 
September 3, and on the trip from the Mulchatna River to 
Nushagak he saw a considerable number of these birds each 
day. 



74 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

On July 23, 1940, Gabrielson observed three pairs of white- 
fronted geese along Kvichak River, above Naknek, and he was 
informed by natives that this is the common nesting goose at 
the base of Alaska Peninsula. He estimated that he saw 500 
birds along the Chulitna River on July 26. 

To sum up, the white-fronted goose nests on the eastern por- 
tions of Alaska Peninsula, at least as far west as Ugashik 
River; farther westward throughout the Aleutian district it is 
only a rare visitor, 

Chen hyperborea: Snow Goose 

On October 1, 1942, Gabrielson noted two snow geese with a 
flock of Canada-type geese at Cold Bay. Again, on October 20, 
1944, he saw 4 large flocks flying over Olga Bay at Kodiak Island, 
and, on the same day, he stated that 1,000 to 1,500 birds settled 
near Kodiak village, where several were shot by the townspeople. 
He comments: "They are seldom seen here, though more fre- 
quently at the south end of the island." 

This is the only information available for this goose. There 
are no records concerning the area to the west. 

Anas plafyrbynchos: Mallard 
Anas platyrhynchos plafyrhynchos 

Attu : Argh'-ich 

Atka: Ag-ich (apparently the same word in both dialects) 

Russian, Commander Islands: Selesenn (Stejneger) 

The mallard is widespread throughout the length of the Alaska 
Peninsula and Aleutian Islands, both as a breeding species and as 
a winter resident. Stejneger (1887) reported also that it was 
"resident, breeding numerously in Bering Island; comparatively 
rare on Copper Island." In 1886, Turner reported that the mallard 
was plentiful in the Aleutians in winter, and stated that it 
breeds sparingly on Agattu and Semichi Islands and that a few 
pairs were seen on Amchitka Island in the latter part of May 
1881 — which indicates nesting. Our expeditions verify this in- 
formation. In 1936, Attu natives stated that they had observed 
these birds nesting near streams, and stated that they winter 
there. The following season, on June 9, which was during the 
nesting season, we saw several mallards along the shore of 
Attu Island. Wilson (1948) observed them at Attu in the 
breeding season when some of them were paired. The last ones 
observed were on August 28. We found a number of mallards 
among the lakes of Agattu Island, and on June 13 we found 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 75 

a female with eight downy young. On June 21, 1937, a pair 
was seen among the lakes on the southeast part of Kiska Island, 
and another pair was seen in a lake at the South Harbor. On 
July 5, we flushed two males and a female from a pond on 
Amchitka Island. The natives of Atka also assured us that 
mallards are found there both summer and winter, which suggests 
nesting. June 20, 1941, Gabrielson saw a female with four 
young at Unalaska Island. The islands mentioned here are the 
principal ones that contain ponds and lakes. However, Calm 
found this bird nesting at Unalaska Island. 

Farther east, we obtained additional nesting data. On May 
7, 1925, I found a nest of 11 eggs at Urilia Bay, Unimak Island. 
On June 6, a nest of 5 fresh eggs was found in a stream valley 
below Aghileen Pinnacles, western Alaska Peninsula, and on June 
23 a nest of 10 eggs was observed on the tide flat at Hazen 
Point, Izembek Bay. In 1936, residents at Port Moller assured 
us that mallards nest around Nelson Lagoon, and in 1928 Jaques 
(1930) found it a ''common breeder in the Port Moller region." 
On May 29, 1936, we saw a single male at Ugashik River. We 
had seen a pair at Chisik Island, Tuxedni Bay, in Cook Inlet 
on May 6, and, on May 9, another pair was observed at Anchor- 
age. According to Osgood (1904), "McKay found the species 
breeding at Nushagak and took a number of specimens there 
in May and June, 1881." Gabrielson noted a few along Kvichak 
River July 23, 1940, including one brood of young. He also noted 
a male in the Barren Islands on June 13. 

Mallards undoubtedly nest on various islands south of the 
Alaska Peninsula. On August 29, 1936, I saw two mallards on 
a pond on Simeonof Island, in the Shumagins, and the local 
rancher said they nest there. On Afognak Island, September 2, 
1936, 14 mallards were seen in a lily pond. These could have 
been migrants, yet mallards undoubtedly nest there because they 
are known to nest on Kodiak, nearby. 

As stated above, mallards winter throughout the territory 
under discussion. Localities where considerable numbers have 
been reported are Unalaska, Kanaga, and Unimak. We were 
told by natives of Unimak that when the bays and lakes freeze 
over, the mallards move to the unfrozen streams in the interior 
of the island and return to the lowlands only when the ice has 
disappeared. 

In the summer and fall of 1936 there was an unusually large 
run of salmon up the streams of Unimak Island ; at that time, 
mallards and other ducks, we were told, assembled there to feed 
on free-floating salmon eggs. 



76 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Anas strepera: Gadwall 

The gadwall has been considered to be "accidental" in Alaska, 
on the strength of two records for the Pribilof Islands. It was, 
therefore, of particular interest to find that this bird nests 
regularlj^ in parts of Alaska. 

On Maj' 16 and 17, 1937, several pairs of gadwalls were 
found on the Copper River Flats, near Cordova, not far from 
the mouth of Eyak River. Evidently, these were paired birds 
that were preparing to nest. 

Alfred M. Bailey (1927) reports a pair at Bartlett Cove, 
Glacier Bay, on August 9, and "felt sure" he had identified a 
band at Holkham Bay on September 25, though the light was poor. 

Cahalane (1943) reports that two gadwalls were shot on Oc- 
tober 2, 1940, at the head of Terror Bay, Kodiak Island. 

Chase Littlejohn says "A few of these ducks were shot by me 
while on their way north in the spring at Dolgoi Island, near 
Belkofski. They were the only ones seen." 

On the north side of Alaska Peninsula, where suitable marsh- 
lands are present, the gadwall is fairly common. On May 27 to 
29, 1936, they were common on the tidal marsh and on numerous 
ponds adjacent to Ugashik River where they were courting and 
preparing to nest. Generally, a female would be seen flying about, 
pursued by two or more males. On May 28, this species was 
recorded as "the principal duck seen," and on May 29 "they and 
the scaups made up most of the duck population." A pair was 
collected for specimens. 

At Port Moller, residents assured us that gadwalls nest plenti- 
fully in the lakes upriver from Nelson Lagoon. 

On May 8, 1925, I observed 4 gadwalls near the shore of a 
lagoon at Urilia Bay, Unimak Island; 2 of these were taken for 
specimens. On May 21, five gadwalls were seen among the ponds 
on Hazen Point in Izembek Bay; gadwalls were seen repeatedly 
as late as July 25. 

Beals and Longworth, in a field report, mention that they saw 
4 gadwalls on Unimak Island, March 19, 1941, 1 of which was 
collected. Local residents considered it to be uncommon. 

Gabrielson reports a male and female on a lake at Izembek 
Bay, June 4, 1942. During the fall and winter periods of 1943 
and 1944, he found them to be common among the Kodiak- 
Afognak Islands. 

Turner (1886) records a specimen taken at Unalaska Island 
in December, 1878, and states that they are "abundant" along the 
Yukon Delta district in summer. Nelson does not mention it. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 77 

however, and we did not see this duck in the Hooper Bay district 
in 1924. 

Taber found 5 males and 4 females at Adak Island during the 
winter of 1945-46. 

Stejneger says (1887), "Reported by Dybowski as taken on 
Bering Island." Hartert (1920) mentions a specimen shot on 
Copper Island on May 13, 1911. 

Thus, we find (as is the case with a number of species that re- 
quire lowland marsh) that this duck nests along the lowlands on 
the north side of Alaska Peninsula, possibly also on Unimak Is- 
land, but it occurs only as a straggler in the Aleutians to the west. 

Anas acuta: Pintail 

Russian, Commander Islands: Vostrochvost (Stejneger) 

This widely distributed bird is not common in the Aleutian 
district, but it does occur here and there throughout the entire 
area. It is known to occur on Kodiak Island (Friedmann 1935; 
Howell 1948), where Gabrielson found it plentiful in fall and 
winter. Cahalane (1944) observed pintails in several localities in 
the Katmai region in the autumn of 1940, but his report implies 
that this species is not plentiful. Gabrielson noted a female on 
Naknek River on July 19, 1940, and on July 23 several females, 
evidently with broods, were noted on Kvichak River. He also 
found it at Unimak, Cold Bay, Izembek Bay, Shumagin Islands, 
and Kodiak-Afognak Islands; they were rather plentiful in the 
last-mentioned localities in fall and winter. Einarsen (1922) 
found pintails nesting near Ugashik in 1922, and Jaques (1930) 
found it to be a common breeding bird around Port Moller in 
June 1928. 

On May 23, 1936, we saw 2 pintails near Dillingham, Bristol 
Bay, and, on May 26, 2 more pintails were seen near Snag Point. 
On May 27 to 29, an occasional pair was seen on the flats near 
Ugashik River, where they evidently were nesting. 

Residents on Unimak Island stated that pintails nest there, and 
this was verified by my observations in 1925. In that year, they 
were first seen at Urilia Bay on April 30. On May 4, Donald 
Stevenson saw 10 males flying about, and on May 17 a pair was 
seen at St. Catherine Cove. Pintails were also present on Izembek 
Bay, and on June 30, near Point Grant, in the midst of nesting 
Arctic terns and Pacific eiders, a nest of eight eggs was found. 
Near Frosty Peak, a female that obviously had eggs, or young, 
nearby was observed. 

Turner did not observe the pintail in the Aleutians during the 



78 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

nesting season, and we found that the natives 'did not recognize 
pictures of the bird, yet the bird assuredly occurs in these is- 
lands. We noted pintails on a lake on Umnak Island May 30, 
1937, and on August 23, 1937, Steenis obtained a specimen there. 
On June 16, 1936, C. S. Williams reported a pair on Amukta Is- 
land. Laing (1925) saw a small flock at Kuluk Bay, Adak Island, 
April 13, 1924. We noted a pair among the lakes at the south- 
east point of Kiska Island on June 5, 1937, and, in the same vicin- 
ity, on June 21 we saw two males and a female. Remains of a 
pintail were found in a bald eagle's nest on the north side of Little 
Sitkin Island, and, on June 17, 1937, a pair was observed on 
Alaid Island, of the Semichi group, which is near the western 
end of the chain. On Attu Island, Wilson (1948) found three 
pairs that he thought to be nesting. 

Undoubtedly, all these records denote nesting throughout the 
Aleutian chain, though successful nesting in recent years may be 
adversely affected by the blue-fox industry. 

Stejneger (1887) says pintails are very numerous on Bering 
Island, but less common on Copper Island. 

Turner did not think pintails wintered in the Aleutians; how- 
ever, he recorded them at Unalaska as late as November. More- 
over, Beals and Longworth (field report) state that pintails are 
plentiful in winter in the vicinity of Unimak Island. On March 1, 
a flock of 25 was recorded; on March 16, 2 were noted; on March 
26, a flock of 23 was noted. Moreover, Taber observed a flock of 
48 that were wintering at Adak, in 1945-46. 

Anas falcafa: Falcated Teal 

Rowland Wilson (1948) reports an unusual observation, in part, 
as follows: 

On May 23 and 24, 1945, Lt. C. L. Stone and I observed a male and female 
of this handsome species, together with two Tufted Ducks and three male 
and four female Greater Scaups, on a little "pothole" pond inland from 
Murder Point [Attu Island]. We had abundant opportunity to watch the 
teals, for they were not shy. On the 24th we saw the male diving several 
times. He went under rather awkwardly, giving us the impression that 
he was not used to such activity. The female did not dive while we watched 
her. . . . Possibly they had been blown in from the west by a recent storm. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 79 

Anas crecca: Common Teal 
Anas crecca nim'ia 

Attu: CheerrJi-ooU (obviously the Russian name) 

Atka: Krech-cheer-tha (derivation from Russian is at least suggested by 
the middle syllable) 
Ataxciyax (Jochelson — probably the true Aleut name) 
Russian, Commander Islands: Tschirok (Stejneger) 

It is now well established that the breeding species of teal 
throughout the Aleutian chain is Anas crecca. During our expedi- 
tions, with only one exception, when a close view of males was 
possible, or when specimens were collected,- the bird proved to be 
the common teal. Beals and Longworth collected a male at Uni- 
mak Island, June 11, 1941, This is the easternmost point for 
which we have a record of this bird. Swarth (1934) records 3 
specimens, 2 males and 1 female, taken on Akutan Island, May 24, 
1927. We found these teals common throughout the Aleutian 
chain, and they are to be found on most of the islands where 
suitable habitat is available. Bent lists a specimen collected by 
Lucien M. Turner on Atka Island, June 28, 1879, and one taken 
by J. Hobart Egbert on Kiska Island, July 14, 1904. He also 
states that in 1911 his party collected "quite a series" of speci- 
mens in the western and central islands, and every male proved 
to be this form. Laing (1925) records two males taken at Adak 
Island, April 13, 1924. On our own expeditions, several specimens 
were taken, including males on Kagalaska, July 4, 1936, and on 
Amchitka, July 24, 1936. 

Gabrielson noted a pair of common teals on Amukta Island, 
June 25, 1940; he saw about a dozen on Amchitka, June 28, and 
saw others at Tanaga, Ogliuga, Atka, Ulak, Kavalga, Segula, and 
Adak. 

These teals are the most abundant fresh-water ducks in the 
Aleutians. Broods of young were seen on the small islands, 
Ogliuga and Skagul, and two broods were seen on Kanaga. On 
July 7, 1937, we found a nest of seven fresh eggs on Amchitka. 
On July 3, 1936, a female with two downy young were seen in a 
shallow grassy pond on Adak Island. The natives said that teals 
nest on Attu, and a male was seen on Agattu, June 15, 1937. On 
August 23, 1937, I counted at least 42 teals on a lake near Nikol- 
ski Village, Umnak Island, and Steenis, on the same day, saw a 
greater number. Pairs were seen on various other islands, and it 
is certain that they nest throughout the length of the Aleutian 
chain. 

Stejneger (1887) reported the European (common) teal as an 



80 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

abundant visitor on Bering Island, but less common on Copper 
Island. The species also occurs in the Pribilof Islands. 

Apparently, while nesting, the teals are less susceptible to the 
predations of the blue fox than are most other waterfowl, though 
the chief of Atka Village declared that they were much more 
numerous in early days. They generally occupy shallow, weedy 
ponds, though they also spend much time on the beaches — some- 
times on boulder beaches. They feed extensively along the beach 
margin and are often found on salt water. This was noted par- 
ticularly on Ogliuga Island. The nest found on Amchitka Island, 
July 7, 1937, was situated in the dense stand of Elymus border- 
ing the ocean beach, and the female repeatedly was observed 
feeding on this open beach. 

Cottam and Knappen (1939) have reported on the contents 
of five stomachs of this species, and say that "three out of four 
birds taken in coastal Alaska had fed almost exclusively on soft- 
bodied crustaceans." The authors felt that the high percentage 
of animal matter (80.2 percent) was probably not typical and 
would not be maintained in a larger series of stomachs. However, 
our observations on the feeding habits of these teals in the Aleu- 
tians are in accord with these findings from the stomach analyses. 

Evidently, the common teal winters in the Aleutians. We were 
assured of this by the natives of Attn and Kanaga, and residents 
of Unimak also stated that teals winter there. Furthermore, 
Donald Stevenson, who spent several winters in the Aleutians, 
furnished positive evidence of it, for in his field reports he said, 
in part (referring to Unalaska Island), 

They were again noted here Nov. 2, 1920, and at intervals in the month 
of November until November 21. Then again here January 7, 1921, to 
January 31, 1921. Being often observed feeding in small pools of salt water 
along the beach after the cold weather had set in and had frozen the fresh 
water streams. . . . Existed in great numbers at Umnak Island, near Otter 
Point November 22, 1920, in small fresh water pond, and in large fresh 
water stream. . . . Observed about five hundred here Dec. 13, also noted here 
Dec. 18, 1920. Specimens taken were in a fine fat condition. 

In 1943, Cahn noted 1 common teal at Unalaska Island on Oc- 
tober 14, and 2 on December 2. Taber noted a flock of 47 at 
Clam Lagoon, Adak Island, from November 1945 to late January 
1946. Sutton and Wilson saw a male at Attn, March 5, 1945. 

Anas caroUnensis: Green-winged Teal 

The common teal occupies the Aleutian Islands, and the green- 
winged teal occupies the Alaska Peninsula. There is some over- 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 81 

lapping in range. On June 5, 1937, Steenis and I observed 3 teals 
at a small pond on Kiska Island — at least 2 were males in bright 
plumage. One was clearly A. crecca, with a plain breast and a 
light-colored scapular streak. The other bird lacked the scapular 
streak, and plainly showed the crescent on the side of the breast. 
We both saw these markings, but we failed to note the markings 
of the third bird. It may have been a female. It is interesting 
to note that both species of teals occur on the Pribilofs. 

Cahn, writing of his observations at Unalaska Island during 
the war years, says of the green-winged teal : "Observed in every 
month of the year except August in four years of observations; 
inhabits the same area as A. crecca, but more common." Taber 
did not record it as being present in winter at Adak. 

Because of the difficulty of identification in the field, and be- 
cause females of the two forms are indistinguishable, even with 
specimens, there is confusion about their ranges and the extent of 
territorial overlapping. Until more collecting is done, and be- 
cause specimens of nimia east of Unimak are lacking, we may 
assume for the present that the birds of Alaska Peninsula are 
chiefly carolinensis. They occur in suitable locations along the 
peninsula. In 1925, they were noted on Unimak Island and 
Izembek Bay. In 1936, we were informed by residents that teals 
nest up the river from Nelson Lagoon, and we saw at least two 
teals at Ugashik River on May 29, 1936. Osgood (1904), how- 
ever, found teals scarce in the interior of the base of Alaska 
Peninsula, and says, 

One old female was seen on the Nogheling River July 21, and no more 
appeared until we neared the coast on the lower Nushagak River. Immense 
flocks were seen in late September in the vicinity of Nushagak. McKay 
obtained several specimens at Nushagak and at Ugashik. 

The National Museum has a male green-winged teal that was 
taken at Nushagak, May 6, 1883, by Paul J. Kojevnikoff. 

Cahalane (1944) has observed the green-winged teal on the 
mainland only once positively : a flock of 12 was seen September 
24, 1940, near the mouth of Savanoski River. 

Gabrielson saw several green-winged teals, obviously with 
broods, up the Kvichak River, July 23, 1940. On April 27, 1942, 
he positively identified 21 of these birds at King Cove, and later 
he saw many more at Cold Bay. 

Green-winged teals occur on Kodiak Island, according to Fried- 
mann (1935) and Howell (1948). Cahalane and Gabrielson 
found them to be numerous in the Kodiak- Afognak area, but there 
are no records for the rest of the territory under discussion. 



82 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Teals winter at Unimak Island, according to residents, and, 
according to Cahn, they winter as far west as Unalaska Island. 



Mareca penelope: European Widqeon 

Russian, Commander Islands: Svistsch or Svistun (Stejneger) 

Four specimens have been collected on the Pribilof Islands, 
and Dall (1873) records a specimen taken at Unalaska, October 
12, 1871. He says it is "not uncommon among the ducks brought 
in by the native hunters of that locality." He considered it to be 
a winter visitor, "migrating about May 1st." 

On June 21, 1937, we found a pair at a lake back from the 
beach at the more southerly harbor on Kiska Island. As the two 
birds flew by, I clearly saw the buffy coloration on the head of 
the male. At that season of the year, a pair suggests nesting. 

Stejneger (1887) records this bird as being a visitor to the 
Commander Islands in migration. 

Mareca americana: American Widgeon 

The Am.erican widgeon, or baldpate, is rare in the Aleutian 
district. On May 27, 1936, 2 or 3 were observed on the flat marshes 
near Ugashik River, and 2 males were seen May 29. We saw none 
to the westward, though Gabrielson records seeing a male and a 
female on a pond near Izembek Bay on June 6, 1942, and saw 
others at Port Moller, July 7, 1946. 

Osgood (1904) mentions specimens taken by McKay at Cape 
Constantine and Ugashik in Septeml)er 1881. 

Hine (1919) observed this duck occasionally in the Katmai 
Region in 1919, and he obtained specimens near the mouth of 
Katmai River. 

Friedmann (1935) records seeing the baldpate at Kodiak, and 
a specimen was taken. He also (1937) reports that bones of this 
duck were found in middens on Little Kiska Island. Gabrielson 
records that the species was "common" in the Kodiak-Afognak 
Islands in the fall and winter months of 1943 and 1944. 

Howell (1948) reports as follows for Kodiak: "Two were seen 
May 31, at Middle Bay, and one on June 16, at Bell's Flats," in 
1944. 

Turner states that it is rarely seen on Attn Island. 

Finally, Stejneger (1887) found a dead bird of this species 
among the sand dunes of Bering Island. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 83 

Spatula clypeafa: Shoveler 

Attu: Koo-chii-tuh or Koo-chu-thoh 

Eussian, Commander Islands: Soksiin (Stejneger^ 

The natives of Attu Island seemed to know this duck and had 
a name for it — if their identification is correct. They recognized 
a picture, agreed on the spoon-shaped beak, and claimed that the 
shoveler nests on Attu Island. 

Stejneger (1887) considered it to be one of the rarer ducks on 
Bering Island, but he thought that it breeds there — hence, it would 
not be surprising to find it among the Near Islands. 

We did not find the shoveler in the Aleutians, but on May 29, 
1936, a male was seen among some other ducks in a pond near 
Ugashik River on the peninsula. Two specimens were taken by 
McKay near Nushagak, on August 14, 1881, and on September 
24, 1882. Cahalane (1944) records 1 bird seen by him, September 
7, 1940, on Brooks River, and Gabrielson observed 2 at Morzhovoi 
Bay, June 21, 1940 — the westernmost point for which we have 
precise record. 

The shoveler is scarce in the Aleutians and Alaska Peninsula, 
and it is comparatively scarce on other parts of the Bering Sea 
coast. The only place where we found them in considerable num- 
bers was in the vicinity of Cordova, on the Copper River flats 
near the mouth of Eyak River. There, on May 16 and 17, 1937, 
we saw many of them engaged in courtship, evidently preparing 
to nest. 

Ayfhya americana: Redhead 

Attu: Ka-ve ini'-much 

The A. 0. U. Check-List states that the redhead is a casual 
visitor on Kodiak Island, Alaska, and Friedmann (1935) men- 
tions a specimen taken there by Rutter. 

On June 16, 1936, I had a glimpse of a pair of ducks, identified 
as redheads, rising from a pond near the beach on Amukta Island. 
Upon arrival at Attu, Chief Hodikoff declared that a few ducks 
(like those in the picture of redheads that we showed him) nest 
on Attu and remain in winter. He gave us the native name, Ka-ve 
(head) im-much (round). He was certain of his identification. 

At the time, we were concerned only with the redhead, but be- 
cause of its similarity to the pochard, which occurs on the Pribi- 
lofs, it is possible that the Aleut chief was really referring to the 
Old World species, Nyroca ferina, and conceivably the birds that 
we noted on Amukta were also of that species. 



84 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Aythya vallsineria: Canvasback 

In 1925, I was told by a trapper that canvasbacks had been seeni 
on Urilia Bay, on Unimak Island. It was thought at the time 
that he had confused the birds with some other canvas-backed^ 
type of duck. But the report is more plausible since Friedmanni 
(1937) referred to this species — five humeri found in old middensi 
at Dutch Harbor. The bird has also been recorded for the Pribi- 
lofs. No other records for canvasbacks were obtained. 

Ayfhya marila: Greater Scaup 
Ayfhya marila nearcfica 

Attu: Han-o ka-ve-too 

Russian, Commander Islands: Tschernik (Stejneger) 

Four specimens of ihis species, which were breeding birds, werei 
collected by Donald Stevenson at Izembek Bay in June 1925. On' 
geographic grounds, also, the scaup of this region should be 
A. marila nearctica, rather than A. affinis. The American greater 
scaup was recorded from Kodiak Island by Friedmann (1935). 
Concerning this bird, Osgood (1904) says, "Scaup ducks, doubt- 
less this species, were seen in small flocks along the Nushagak 
River September 4 to 9. McKay took them in May and July at 
Nushagak and Ugashik." And again, he says (1901), "a flock of 
six scaup ducks were seen on a pond near Tyonek September 17." 

Cahalane records this duck on the Naknek River, where it was 
abundant, on September 28, 1940, and he found it to be common 
on Brooks Lake, September 9, though he did not see it in the more 
interior portions of the Katmai National Monument. He also 
said that they were fairly common in the Kodiak-Afognak area. 

On July 23, 1940, Gabrielson observed four broods of greater 
scaups on the Kvichak River. In later years, he saw them in num- 
bers at Unimak, Atka, Kanaga, Umnak, Unalaska, Amchitka, 
Shumagin, Sanak, and Kodiak-Afognak Islands. 

We saw two greater scaups near Chisik Island, Cook Inlet, May 
7, 1936. On May 27-29, 1936, scaups were common, flying about 
in pairs, near Ugashik River. In June 1928, Jaques found them to 
be common near Port Moller. They were reported to be common 
near Chignik, maintaining their numbers better than other ducks 
in that vicinity. 

In 1925, I found scaups nesting in Izembek Bay. About the 
middle of May of that year, there were small bands in St. 
Catherine Cove, at Unimak Island, swimming on the salt water 
or on the small ponds on the shore, sometimes segregating in 
pairs. In the middle of June, they were particularly common 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 85 

about Hazen Point, and they were seen elsewhere in Izembek 
Bay. On June 20, they were still flying about in pairs. On that 
date, a female was seen standing near a recently constructed nest 
cavity. On June 30, 3 nests were found on small islands near 
Point Grant; 2 of these nests contained 9 and 10 eggs respectively. 
The number of eggs in the third nest was not recorded. On July 
26, a nest of seven eggs was found on a gull island. 

Residents stated that scaups nest on Unimak Island. 

Scaups were noted at intervals throughout the Aleutian chain. 
Four or five were seen near Nikolski, Umnak Island, on May 
30, 1937; 7, mostly males, were seen on Corwin Lake on Atka, 
June 22, 1936 ; several were noted on Amchitka, July 1937 ; a flock 
of 30 was seen on a lake on Kiska, July 26, 1936 (where half a 
dozen were seen on June 4, 1937) ; and several pairs were seen on 
Agattu Island in the middle of June 1937. Steenis observed four 
pairs and a female there, and other members of the party observed 
paired scaups. On June 15, 1937, on Agattu, I found a scooped- 
out nest cavity with a little down and some white breast feathers, 
which I thought to be a scaup nest. Austin H. Clark (1910) 
found this species to be rather common at Attn and Agattu. 

Chase Littlejohn (manuscript notes) says, "Found breeding 
at Sanak, Ukamuk [Chirikof Island], and Morzhovoi Bay, each 
nest contained nine eggs. They congregate in large flocks in 
winter at Sanakh and remain so until spring, when they pair off 
and begin nesting." 

The Attn chief assured us that scaups nest on Attn and winter 
there. On Kanaga Island, also, we were assured that scaups are 
plentiful in winter, and that they become very tame around the 
dock. 

Taber found them wintering at Adak, and for Unalaska Island 
Cahn reports — 

An abundant winter inhabi:-;.t of all the larger bays, in common with the 
Harlequin ducks and white-winged scoters. The greatest numbers occur in 
December and January, and the species disappears entirely in April as a 
rule; May 3, 1946 is the latest recorded date. It returns again a few at a 
time, in September and October, gradually increasing in abundance. 

Sutton and Wilson found scaups wintering at Attn Island. 

At Unimak Island, March 1, 1941, Beals and Longworth ob- 
served two rafts, of at least 1,500 scaups each, on Swanson 
Lagoon, and a trapper assured them that these ducks spend the 
winter there. 

In several localities, mention was made of the scaup's habit of 
assembling near docks. In some cases, at least, fish offal appears 



86 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

to be the attraction. This was definitely reported to be the case 
at False Pass, where the ducks gather at the cannery docks during 
the fishing season. 

Ayfhya affin'is: Lesser Scaup 

Friedmann (1935) lists the lesser scaup in the avifauna of 
Kodiak Island on the basis of osseous remains found in middens. 
We did not identify this duck on our expeditions, and we as- 
sumed that the scaups observed were of the larger form. 

Ayfhya ful'igula: Tufted Duck 

Rowland Wilson (1948) added this species to the list of Aleu- 
tian birds, bearing out our assumption that it could easily occur 
among the western islands. He reports seeing 2 males and 2 fe- 
males in a little pothole, inland from Murder Point on Attu Is- 
land on May 23 and 24, 1945; he watched them for some time, 
and the "tufts of long, loose feathers which streamed down from 
the nape of each male" were noted in detail. 

Bucephala clangula: Common Goldeneye 
Bucephala clangula americana 

Attu: Ha-no sakh-oi-a 

Russian, Commander Islands: Gogol (Stejneger, referring to the closely 

related European form). 
This may be the "whistler" amtdtux, given by Jochelson, for which no dia- 
lect was mentioned. 

Friedmann (1935) lists bones found in Kodiak middens, which 
he assigned to this subspecies on geographic grounds, and he 
mentions two specimens collected there by Fisher. On March 21, 
1924, Laing (1925) observed three of these ducks at Uyak Bay of 
Kodiak Island. 

Gabrielson noted that this duck is plentiful in the Kodiak- 
Afognak area in fall and winter; he found it in the winter at 
Unalaska, Umnak, Kanaga, and Atka, and at King Cove and Cold 
Bay in spring and fall. 

I observed the goldeneyes in 1925 at Unimak Island. On April 
29, 1925, I saw a pair flying over a lagoon at Urilia Bay, and on 
May 1, 1925, I saw a flock of about 10. On May 4, 1925, Donald 
Stevenson reported at least 200 on Peterson Lagoon. Identifica- 
tion could not be positive on all of these instances, but they were 
assumed to be americana on the basis of known distribution. 

Friedmann identified a goldeneye humerus in middens of Dutch 
Harbor, and Laing (1925) observed nine goldeneyes at Unalaska, 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 87 

March 21 and 22, 1924 ; however, those were immature birds, or 
females, and identification of the species was doubtful. Turner, 
also, records this duck for Unalaska in December, and he says 
that it winters there. 

Cahn reports the goldeneye for Unalaska Island, saying. 

Strictly a winter visitor, present in considerable numbers but never in 
large flocks. Goldeneyes drift in by one's and two's in late October (October 
24, 1943), and are common in the larger bays during the period of December 
through February, at which time they disappear far more abruptly than 
they arrive. April 11, 1946, is the latest recorded date; this is unusually 
late. 

Laing observed an unidentified goldeneye at Adak Island, and 
at Attn Island he positively identified two adult males that "were 
found ardently courting in a flock of six." 

Taber found goldeneyes wintering at Adak Island in 1945-46. 

When shown colored illustrations, the Attn chief picked out the 
common goldeneye and said it was plentiful there in winter, 
arriving in November and (he thought) leaving early in March. 

Sutton and Wilson found them wintering at Attu. Stejneger 
(1885) reported that the European common goldeneye occurred 
at the Commander Islands in winter in small numbers. 

Because there are so few specimens, and because racial identi- 
fication cannot be ascertained in the field, it would be possible 
that the Old World form (keeping in mind that it is recorded 
from the Pribilofs) occurs in the western Aleutians and has not 
been detected. In the case of this form, we are leaning heavily 
on assumed geographical distribution. 

Bucephala islandica: Barrow's Goldeneye 

We saw several Barrow's goldeneyes at Seward, May 5, 1936; 
at least 12 at Port Chatham, Kenai Peninsula, on May 6; 2 at 
Chisik Island, Cook Inlet, May 7; and 1 male at Kodiak Island, 
May 13. Friedmann (1935) has given a number of records for 
Kodiak Island, and Gabrielson noted them in winter and fall at 
Unalaska and Kodiak-Afognak Islands. 

With regard to the base of Alaska Peninsula, Osgood (1904) 
reported — 

One was seen on the Nogheling River July 20, and one was killed there 
some days later; another was shot by W. L. Fleming on a small pond near 
the head of Lake Clark July 28. Several immature birds were killed at 
the mouth of the Chulitna River August 4. Rather common at intervals 
along the Chulitna River August 12 to 17; generally seen in family parties 
of 6 to 10. Near Swan Lake a flock of about 15 was seen feeding on a shal- 
low lake in company with a flock of 10 swans. Seen almost daily in pairs 
or small flocks along the Malchatna and upper Nushagak September 3 to 6. 



88 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Cahalane (1944) found this duck "numerous in the upper por- 
tion of the Naknek River, September 4." They were also abundant 
in Amalik, Kaflia, and Kukak Bays on October 5 and 7. Hine 
(1919) considered it to be a common species in the Katmai Bay 
area, and he obtained a specimen there. 

Thus, it would appear that the Barrow's goldeneye is confined 
pretty much to the basal part of Alaska Peninsula, adjacent is- 
land groups, and mainland areas as a breeding bird. The Ameri- 
can common goldeneye, on the other hand, is the form generally 
encountered to the westward, as a migrant. There are no data to 
show that any goldeneye nests west of the more or less tree- 
covered base of Alaska Peninsula. 

Bucephala albeola: Bufflehead 

Attu : Chirr-u-num Sakh-oi-a 
Atka : Mith-i-me-thruh 

The bufflehead occurs sparsely throughout the Aleutian district. 
Friedmann's data (1935) show that this duck is rather common 
on Kodiak Island, and that it nests there. Osgood (1904) reportsi 
that "two specimens were seen at Cold Bay October 17 among f 
some ducks killed on the bay by natives. One was taken at 
Nushagak by McKay, May 2, 1882." 

We noted at least six buffleheads at Port Chatham, Kenai Penin- 
sula, May 6, 1936, At Chignik, we were told that buffleheads are 
seen there in autumn. Gabrielson has seen them at Cold Bay, at 
the Shumagin Islands, and at Sanak Island. 

We found two females in a pond at Ikatan Peninsula, Unimak 
Island, May 19, 1936. Beals and Longworth noted one at False 
Pass, March 7, 1941, and four on Ikatan Peninsula on April 15, 
1941. 

Turner (1888) says that this duck occurs in winter at Unalaska, 
where he obtained specimens, and he adds that they are rare to 
the westward, where they are present only in winter. Gabriel- 
son found them in winter at Unalaska, Atka, Amchitka, and 
Umnak. Over a period of 4 years, Cahn saw only one bufflehead 
at Unalaska Island (on February 22, 1944). 

Ray Clark, storekeeper on Umnak Island, said that butterballs 
(buffleheads) remain there in winter. 

Wetmore (manuscript notes) says that R. H. Beck saw a pair 
of buffleheads in a pond back of Atka village on June 13, 1911; 
Laing (1925) saw "fully thirty-five" in a small lagoon on Adak 
Island on April 11, 1924. And Taber found the species wintering 
at Adak, where there were 32 birds noted in a census on January 
13, 1946. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 89 

The Attu chief stated that his island is within the wintering 
range of the bufflehead, but he insisted that they also nest on 
Attu, "up high." 

Stejneger (1887) reports the bufflehead as "an accidental visi- 
tor during the winter of 1882-83" in the Commander Islands. 

Incidentally, this bird is mentioned in a number of records for 
the Pribilof Islands. 

Clangula byemalis: Oldsquaw 

Attu: Ang-lach 

Atka: A-lang-ach 

Unimak: Alg-nach' 

Russian, Commander Islands: Sofka (Stejneger) 

Russian, Yana region : Savka (Birula) 

Chukchi: Pojgochek, male 

Achak, female (Palmen) 

The oldsquaw is fairly common, especially in migration, and 
has been reported by most writers on southwestern Alaska. It 
occurs on Kodiak, and Osgood (1904) reports it from various 
places at the base of Alaska Peninsula and the Bristol Bay region. 
Einarsen (1922) reports several of these birds nesting near 
Ugashik in 1922. Laing (1924) counted as many as 200 at Dolgoi 
Island, March 23, 1924, and apparently he saw it in many other 
unidentified localities. He collected a specimen at Kodiak, March 
21, 1924. Dall (1873) considered it to be abundant east of 
Unalaska. 

We noted the bird at various points : 2 migrating flocks on the 
Gulf of Alaska, May 2, 1936; 150 birds at Chisik Island, Cook 
Inlet, May 7 ; a flock of 30 in Shelikof Strait, May 13 ; 1 bird in 
Nushagak Bay, May 23; and 2 birds on the flats at Ugashik 
River, May 27. They are said to arrive at Chignik "late in the 
fall." 

The oldsquaw is common in migration along the Alaska Penin- 
sula and adjacent islands, but we were unable to establish nesting 
records. 

They are rather common in the Aleutians at certain seasons, 
especially in winter. In 1925, I found them to be numerous 
about False Pass in the latter part of April and in May. One 
was seen in St. Catherine Cove as late as May 20, but none was 
seen after that date. Beals and Longworth (field report) re- 
corded them at False Pass and neighboring points on January 12, 
13, and 19, 1941, and they observed them daily through March 
and as late as April 10. 

Wetmore (manuscript notes) saw 2 birds at Unalaska Island 



90 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

on June 7 and 11, 1911, and McGregor (1906) collected 1 female 
in worn plumage on July 20, 1901, at Tigalda Island. 

Cahn often found them wintering at Unalaska Island; the| 
earliest date was November 3, 1943, and the latest date was April 
19, 1946. Taber found them wintering at Adak. 

We did not see many oldsquaws in the Aleutians; however, at 
Atka Island we were told by the natives that they formerly had 
nested on that island, but not "since the foxes came." The natives 
said that these birds winter on Atka and Kanaga in large num- 
bers. The Atka chief assured us that these ducks nest on 
Amchitka — he stated that although he had seen the young, he, 
had not seen any nests. 

Kiska Island appears to be one of the favorite localities for 
the oldsquaw. This island was mentioned by Dall as the western 
limit of its range. We saw several birds there as late as June 4 
and 5, 1937, and Wetmore reported them to be fairly common 
near the entrance to Kiska Harbor, June 17 to 21, 1911. We also 
found oldsquaw remains in two bald-eagle nests on that island; 
oldsquaw remains were also found on West Unalga, and in eagle 
nests on Rat and Little Sitkin Islands. 

Dall (1874) said that the oldsquaw was resident as far west 
as Kiska, but that it was not abundant. We learned from thei 
Attu natives that it nests on Agattu and is abundant in the Neari 
Islands in winter. This is substantiated by the report by Suttonr 
and Wilson on Attu. The oldsquaws wintered there, and after 
March 4 they were observed courting. Turner (1886) said that 
few of these birds nested in the Aleutians, but that many of 
them wintered there. In 1887, Stejneger reported oldsquaws 
"breeding numerously on Bering Island." Gabrielson also ob- 
served them wintering as far west as Atka. 

To sum up. Turner's statement (see above) applies very well 
to the Aleutian district as a whole. 

Hisfrionicus hJstrionJcus: Harlequin Duck 

Attu: Kagh'-i-ach 

Atka: Kagh'-a-thi-ga 

Unalaska : Kang-a-rich 

Unimak: Kang-ath'-a-gich 

Russian, Commander Islands: Kameymschka (Stejneger) 

This is the most abundant duck in the Aleutian Islands. We 
found harlequin ducks at practically every island that we visited, 
singly sometimes, generally in small groups, and occasionally 
in larger flocks. It is safe to say that, at one time or another, 
harlequin ducks occur at every island, large or small, from Uni- 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 91 

nak to Attu. Stejneger has reported them to be common in the 
Commander Islands. 

They were also found east of the Aleutians — at Amak Island, 
at Izembek Bay, and at False Pass. In the Shumagin group, we 
observed them at Unga, Nagai, and Simeonof Islands. They 
^vere at King Cove, the Barren Islands, Afognak, Port Chatham 
m Kenai Peninsula, and at Seward. Osgood (1904) observed 
them along the Egegik River and "about the mouths of the larger 
streams that empty into Becharof Lake." He found them to be 
common at Kanatak and Cold Bay, and he mentions specimens 
taken by McKay and Johnson at Igushik and Nushagak. 

Cahalane (1944) reported harlequins in large numbers in the 
general region of Katmai National Monument in the fall of 1940, 
and Hine (1919) considered them to be one of the most common 
ducks in the Katmai Bay area in the summer of 1919. Cahalane 
ialso recorded them as being abundant in the Kodiak-Afognak 
igroup in the fall of 1940, where Gabrielson noted 200 on June 16, 
,1940. W. Sprague Brooks (1915) observed them on April 19, 
1913, at the Semidi Islands, and on April 22, 1913, he saw them 
at King Cove. 

Although these birds occur on the north side of Alaska Penin- 
sula, they are more common on the south side, which is more 
Irugged. Evidently, these birds nest on Alaska Peninsula. On 
I July 19, 1940, Gabrielson noted a pair flying along Kittiwake 
iCreek, between Brooks and Naknek Lakes, and Friedmann (1935) 
[states that Bretherton found them breeding in June on Kodiak 
Island. In the spring of 1925, I often observed two pairs along 
a stream just north of Aghileen Pinnacles, near the western end 
of Alaska Peninsula. Eventually, on June 3, only the males were 
seen; presumably, the females were nesting. 

On July 16, 1911, Wetmore (manuscript notes) observed a fe- 
male and a group of young in King Cove. 

It is difficult to determine the status of the harlequin ducks 
in the Aleutians. The natives insisted that they nest along streams 
and that their nests are very hard to find. In way of substantia- 
tion, we found no nests and no broods of young. However, we 
found these birds on islands that had no suitable nesting streams. 
On the other hand, Austin H. Clark (1910) reported: "It was 
common about Atka, where 1 or 2 were seen inland on a small 
stream ; on Attu and Agattu it was also numerous on the streams 
as well as along the coast." 

Turner (1886) described a deserted nest on Unalaska Island, 
in a hollow formed by two blocks of rock. A native assured him 



92 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

that it was the nest of a harlequin duck. Here, again, our own] 
experience was baffling. Though there were numerous cKffs andj 
many available sites for nesting along the rocky shores, we saw 
no young brood throughout the two summers of observations. ' 
Wetmore, however, had pertinent observations at Kiska Island 
in 1911, when he says (manuscript notes) that | 

some of them were already nesting along the base of a high rocky cliff, asi 
they seemed very anxious while I was along there, those on the water.- 
whistling and swimming in small circles. I saw one or two females slipping 
quietly away from shore ahead of me, but flushed none from the beach itself. 

Beals and Longworth found harlequin ducks wintering at 
Unimak Island, and stated that they nest there. Elsewhere in the 
Aleutians, natives said that they are more numerous in winter 
than in summer. 

Stejneger (1885) found no evidence of nesting in the Com- 
mander Islands, and stated that the natives knew of no nesting. 

From these various observations, it can be concluded that the 
harlequin ducks nest on the Alaska Peninsula, possibly rather 
commonly; that they also nest in numbers unknown in the Aleu- 
tian Islands; that immature birds, various nonbreeders, and 
males gather for the summer in these waters; and that they 
winter there in great numbers. 

We had little opportunity to study food habits, and it must be 
assumed that, in the salt water, it consists of marine inverte- 
brates. The teacher of the native school at Atka informed us 
that in the autumn of 1936, when there was a large run of salmon 
up the streams of Atka Island, harlequin ducks were seen on the 
streams, presumably feeding on salmon eggs. However, we have 
no certain data on this subject. 

PolysHcfa steller'i: S+eller's Eider 

Chukchi: Kataadlin (Palmen) 

This little eider of the Bering Sea region occurs abundantly 
along the Alaska Peninsula and the Aleutian chain at certain 
seasons — particularly in winter. It is recorded as far east as 
Kodiak, where Friedmann (1935) lists many specimens and ob- 
servational records, including some bones from middens. Dall 
(1873) says it was observed in the Shumagins "in March, and in 
the summer months." The same writer (Dall 1874) reports them 
as wintering at Sanak Island, but he considered Unalaska to be 
the center of abundance for this species. He remarked upon the 
irregularity of their occurrence, because he had found Steller's 
eider, together with the Pacific eider, to be numerous at Unalaska 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 93 

in May 1872, however in May 1873 he did not see a single one of 
either species in that locality. 

Osgood (1904) considered the Steller's eider to be common 
about Bristol Bay, and he mentions specimens collected by McKay 
and Johnson at Nushagak and Ugashik. On October 4, 1940, 
Cahalane (1944) observed a group of 6 in Amalik Bay, and he 
saw 6 again (perhaps the same group) the next day. Gabriel- 
son noted 2 males at Morzhovoi Bay June 21, 1940, and collected 
1, which was not in breeding condition. In 1936, we observed 
several sizable flocks in Nelson Lagoon. 

In 1925, I found this duck to be rather common in Izembek Bay. 
On May 17, 1925, there were small groups at St. Catherine Cove, 
Unimak Island, and on May 20 about 200 were seen there, as 
well as several bands offshore in Bering Sea. Several flocks, 
totaling at least 300 birds, were spending the summer in Izembek 
Bay; they used Glen Island, near its entrance, as their home- 
ground. These were immature birds of both sexes, though there 
was an occasional one in adult male plumage. One male in adult 
plumage was collected on June 17. The testes were very small. 
None of the birds were seen on the adjacent marshlands, and 
there was no evidence of nesting. 

Turner (1886) testifies to the presence of the Steller's eider 
among the Aleutians in winter, even to the western end of the 
chain. Stejneger (1887) said that they wintered in the Com- 
mander Islands in "countless numbers," arriving early in Novem- 
ber and remaining until after the middle of May. Friedmann 
(1937) has recorded five humeri of this duck from middens on 
Little Kiska Island. Beals and Longworth observed them often 
in January, March, and April, 1941, and saw them as late as April 
25, at False Pass. 

Although we did not find the Steller's eider nesting, older rec- 
ords furnish rather good evidence of nesting on the Alaska 
Peninsula and Aleutian chain. A. C. Bent (1925) records some 
notes sent to Major Bendire in 1892 by Chase Littlejohn, which in- 
cluded a statement that "a few were nesting at Morzhovoi Bay 
in June." Dall (1873) writes of the pairing of these ducks at 
Unalaska and describes a nest found on Amaknak Island, May 
18, 1872. It contained a single egg. Turner (1886) saw a few 
of these ducks at the western end of Attu Island in July 1880, 
and the natives told him that the species nested sparingly on 
Agattu Island. 

Judging by the information available to us, we must recognize 
the strong probability that at one time the Steller's eider nested 



94 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

on Alaska Peninsula and Aleutian chain (though undoubtedly in 
small numbers), and that it wintered there in great numbers. 
It is also clear that there has been a great diminution in num- 
bers. On our two expeditions in 1936 and 1937, we were in the 
general region early enough to have observed these ducks before 
all of them had left their wintering grounds. We saw very few, 
and those that we saw were on the north side of Alaska Penin- 
sula. We saw none in the Aleutian chain. We found the natives 
of Attn Island — who have had only limited contact with the white 
man — to be well versed in their local fauna, much more so than 
natives farther east. These Attn natives did not recognize pic- 
tures of the Steller's eider and declared that it does not occur 
there, even in winter. They could be mistaken; however, if we 
accept their testimony there must have been a great decline in 
numbers since 1880, when natives told Turner that these birds 
nest "sparingly" on Agattu Island. 

A. C. Bent (1925) considers the principal migration route in the 
fall "southward along the Siberian coast of Bering Sea to their 
winter homes in the Kurile, Commander and Aleutian islands." It is 
probable that the migration along the Siberian coast is the greater 
one, but if the information furnished by the Attu Islanders proves 
correct (and since we know these birds do winter in the eastern 
Aleutians), it is unlikely that the Siberian birds go to the Aleu- 
tians. On the other hand, we now know that there are large 
nesting populations on the American shores — at Hooper Bay and 
Nelson Island — and we have observations pointing out that the 
eastern Aleutians and parts of Alaska Peninsula, even Kodiak 
Island, are the principal concentration points in winter. In view 
of these facts, we must conclude that there is also a southward 
migration down the Alaskan coast of Bering Sea to the eastern 
Aleutians and Alaska Peninsula, and that the majority of the 
birds wintering in the Aleutians nest on the Alaskan coast. 

Somaferia mollissima: Common Eider 
Somo/er/o mollissima v. nigra 

Attu: Kaf-segh'-ich, male 

Chd-is, female 

Ku-ku-toch, young 
Atka: Ka-smn'-ich, adult (sex?) 

Ku-ku-toch, young 

Kasimax (Jochelson — dialect not given) 
Russian (?), Copper Island: Pistrak (Stejneger) 
Chukchi : Kupuken, male 

Emngi, female (Palmen) 

Common eiders were observed at practically eveiy island of the 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 95 

Aleutian chain and are as universally distributed as the western 
harlequin duck, though not so abundant. Apparently, they are 
not plentiful on the Commander Islands, for Stejneger (1887) 
says that they breed in "very limited numbers in a few places 
on Copper Island, only occasionally flying over to Bering Island, 
round the shore of which a few may be seen in winter." 

In the Aleutians, we saw these birds in small groups, generally 
paired, and they nest, or try to nest, throughout the Aleutian 
chain. Blue-fox raising has seriously interfered with nesting, on 
certain islands. 

Nesting of common eiders was determined for the following 
islands: 

Attn — quite a number nesting on rocky islets in Massacre Bay. 

Agattu — preparing to nest. 

Semichi — nesting on islets in a lake. 

Buldir — nesting on the beach (no foxes present). 

Chugul, Little Kiska, Kiska — nesting reported by Wetmore in 
1911. 

Amchitka — nesting on offshore rocks and preparing to nest on 
beach. 

Ogliuga — plentiful, many young birds seen. 

Aiktak, Kavalga, Ulak, and Tanaga — nesting reported by Gab- 
rielson. 

Little Tanaga, Kanaga, Adak, Aso, Igitkin, Salt, Atka, Chu- 
ginadak. 

Baby Islands — nesting on Adokt and Excelsior (no foxes pres- 
ent). 

These are the nestings actually observed. The birds were ob- 
served at many other islands, where they were probably nesting. 
Were it not for the predations of introduced blue foxes, they 
undoubtedly would nest on practically all islands. 

In 1925, I found nesting groups in Izembek Bay, Alaska Penin- 
sula (particularly on Glen Island and islets near Point Grant), 
as well as on a gull island far out in the bay. On May 22, 1936, 
we found flocks of common eiders in Nelson Lagoon, and in one 
place I counted 111 males on the beach. Residents said that they 
nest abundantly on some grass-covered sand islands there. Os- 
good (1901) mentions a young bird and a set of eggs secured 
by T. H. Bean in July 1880 at Chugachik Bay (Kachemak Bay). 
In 1936, we saw them in Ugashik River, but we did not remain 
long enough to determine their nesting status. 

Thus, we have a fairly accurate and continuous record of 
nesting from Bristol Bay westward to Attu Island. 



96 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

On the south side of Alaska Peninsula, our observations arej 
more scattered. Common eiders are known to nest as far east as I 
Kodiak, and at Chisik Island, in Cook Inlet, we observed at least 
12 pairs in the spring of 1936. The local game warden assured | 
us that they nest on Duck Island nearby, and on May 13, 1937, j 
several common eiders were seen in Icy Straits — the farthest east 
that we had observed this species. Mrs. Frank C. Hibben (1942, 
p. 182) found them nesting in Glacier Bay, the most easterly 
nesting record for southern Alaska. 

Gabrielson observed about 12 common eiders at Kodiak on June 
14, 1940 ; at least 40 birds and 1 nest were seen in the Semidi Is- 
lands on June 18; and a few were seen at Morzhovoi Bay on 
June 21. 

As might be expected, companies of immature birds (nonbreed- 
ers) spend the summer in the waters along Alaska Peninsulai 
and the Aleutians. Furthermore, this is the principal wintering 
ground for the species; they do not venture farther south in any 
great numbers. 

Nesting Habitat 

These eiders utilize a variety of nesting sites. Probably they 
would prefer low islands of gentle slopes (such as the sand is- 
lands of Izembek Bay) , where they can nest in the grass. In such 
places, they nest both on the slopes and on the beach. Similar 
situations may be found in the Aleutians — the beach of Buldir 
Island is an example. There are few places in the Aleutians 
where they can nest with safety on the principal shorelines be- 
cause of the introduced blue fox. We found a few birds nesting 
on the shores at Amchitka and Agattu, but, being adaptable, 
they now seek the grassy tops of offshore rocks and pinnacles, 
or islands in lakes, where they are protected by water. The 
natives assured us that they also nest on ledges of sheer cliffs, 
where foxes are unable to climb. 

Mortality Factors 

The blue fox is probably the most potent predator that the 
eiders face in the Aleutians. In addition to this introduced enemy, 
the northern bald eagle also obtains an occasional eider, but ap- 
parently it does not prey extensively on the species. In a total 
(taken during three seasons) of 466 food items that were identi- 
fied in 32 eagle nests and at a few perching places, only 8 common 
eiders are represented. In one of these instances, the eider had a 
nest within 10 feet of an eagle's perch, and it was to be expected 
that the eagle would eventually seize the bird. It is surprising that 
so few eiders are taken by the eagle, because this duck does not 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 97 

appear to be agile on the wing and is present everywhere. Prob- 
ably the great variety of "sea birds" attract the eagle's attention 
more readily. 

Gulls and ravens are another potent factor in stabilizing the 
eider population, because they prey on the eggs and young. The 
raven is included here solely on the basis of fragmentary ob- 
servations elsewhere. But the glaucous-winged gull was observed 
at times to be active in raiding eider nests. There is an interest- 
ing relationship here that has been noted frequently. Gulls and 
eiders often nest on the same area. Presumably, this should give 
the gulls a better chance to rob the nests of their duck neighbors ; 
however, it does not appear to be that simple, and the situation 
deserves careful study. Assuredly, upon entering such a mixed 
nesting colony, one finds a number of eider nests already rifled; 
yet, many others have not been disturbed. It is noticeable that 
human intrusion, which forces the eiders to leave hurriedly with- 
out covering the eggs, gives a splendid opportunity to the first 
passing gull, and the gulls readily take advantage of it. 

In 1925, in Izembek Bay, I found eiders nesting in the midst 
of a gull colony and found others nesting in a colony of terns. 
All these birds seek the same type of nesting terrain, regard- 
less of neighborly problems. During that season, an effort was 
made to reduce the hazard for nesting eiders by carefully cover- 
ing the disturbed nest with down, just as the bird would have 
done. So far as the results could be observed, this method was 
effective. One will sometimes find gull and eider nests in amaz- 
ingly close proximity, apparently with no detriment to the eider. 

After being hatched, the small duckling still faces danger 
from the gulls. A number of decimated broods were seen, and 
sometimes, as observed at Ogliuga Island, several families then 
join together in a band. 

But in spite of all these nesting hazards, the eiders hold their 
own — they occupy the entire Aleutian district in fair numbers 
and are plentiful enough to utilize whatever nesting sites are 
available to them. 

Somaferia specfabilis: King Eider 

Attu: Sakh'-uch 

Sdkux (Jochelson) 
Russian and Yukat, latitudes of the Yana: Turkan (Pleske) 
Chukchi: Jekadlin (Palmen) 

Information on the king eider is incomplete. We know that it 
spends the winter among the Aleutian Islands, the Shumagins, 
along the Alaska Peninsula, and as far east as Kodiak, where 



98 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Friedmann (1935) has recorded specimens taken and many bones 
found in middens. He also found many bones in middens of 
Dutch Harbor and Little Kiska. In the wintering season, Gabriel- 
son found this eider at various points from Kodiak to Unalaska, 
and Hine (1919) obtained specimens near the mouth of Katmai 
River on June 25, 1919. Though we have relatively few published 
reports of king eiders from the winter range, these ducks must 
occur along the Aleutians and Alaska Peninsula and the ad- 
jacent seas in large numbers, judging by the striking northward 
migration we observed at Hooper Bay in the spring of 1924. 

The Attu chief appeared to recognize this duck; he gave us a 
name for it and stated that a few of them nest at Attu Island 
and that a few winter there. 

Judging by the relatively large number of bones found by 
Herbert Friedmann in the middens at Dutch Harbor and Kodiak, 
and considering the statement of the Attu chief that only "a 
few" winter there, the king eider evidently assembles in the 
greatest numbers among the eastern Aleutians and along the 
Alaska Peninsula. In 1925, I was told by local residents that 
many of these ducks winter at Isanotski Strait and at Wide Bay. 
Beals and Longworth (field report) observed king eiders at 
Isanotski Strait, Ikatan Peninsula, and at neighboring areas at 
intervals from early January to the latter part of May 1941 ; 
their numbers began to diminish in May, and at the end of May 
practically none were left. Four specimens were collected on 
January 13 and 24 and March 6. 

In winter, Cahn found the king eider to be more common than 
the common eider at Unalaska Island, and he says, 

present from early December to early March, usually in small flocks of 
three to six, or solitarily. Dec. 2, 1945, is the earliest record; April 3, 1944, 
the latest. The gizzard of a female found dead contained two specimens of 
the snail Callistoma. 

Though we do not have nesting records for the Aleutians — the 
Attu chief's statement about their nesting on that island may 
properly be questioned — a number of king eiders spend the sum- 
mer near Alaska Peninsula. In 1925, I observed a flock of about 
200 birds (females and immature males) that spent the summer 
at Glen Island in Izembek Bay. 

Lampronetfa fischer'i: Spectacled Eider 

Information on this eider is disappointingly meager for the 
area under discussion. We saw none during the course of our 
expeditions to the Alaska Peninsula and the Aleutians. They are 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 99 

considered to be winter residents there, and A. C. Bent (1925) 
says they occur sparingly east to Sanak Island. Friedmann (1934) 
records a humerus froiri native-village middens on Kodiak Is- 
land. Dall (1873) records it as rare at Unalaska as a winter 
visitor, leaving there in May for northern nesting grounds. 

Surprisingly enough, Turner (1886) says "This species occurs 
among all the Aleutian Islands, where it breeds and is a constant 
resident, but extremely shy." This certainly is not the case 
today. Ball's statement, above, would seem to be more credible, 

Melaniffa deglandi: White-winged Scoter 
Melaniffa deglandi d'lxon'i 

Attu: Tru-pan-ach (obviously of Russian origin) 
Atka : Ta-mu-ghci-luh 

Russian, Commander Islands: Turpan (Stejneger) 

The Kanddgix of Jochelson (dialect not indicated) may possibly refer to 
this duck. 

White-winged scoters have always been common along the 
southern Alaskan coasts in spring migration, and in the course 
of several voyages they have been noted regularly in late April 
and early May along the southeastern Alaskan waters, as well as 
farther west. In 1936, we noted a few at Seward on May 5; 
at least 20 were noted at Port Chatham, Kenai Peninsula, on 
May 6; a few individuals were seen among the Barren Islands, 
May 10 and 11; and several were noted at Kodiak. They were 
common in Kupreanof Strait on May 13. 

In the fall of 1940, Cahalane found that scoters were numerous 
in the Kodiak-Afognak area. Early in September, he found them 
to be abundant in Naknek River, but none were seen by the end 
of September, He says (1944), "On the Pacific side of the area 
scoters were very numerous during the first half of October. 
They were 'abundant to very abundant' along the entire main- 
land coast from Katmai Bay to Point Nukshak." 

On June 16, 1940, Gabrielson noted 100 scoters near Whale 
Island. 

Osgood (1904) observed a flock of six scoters on Neekahweena 
Lake, about halfway up the Chulitna River on August 14, 

Chase Littlejohn, referring to the area between Kodiak Island 
and the west end of Alaska Peninsula in 1887-88, wrote, "Seen 
often during winter, I saw a number of birds at Ukanuk in 
summer where I am sure they breed but for want of time I 
did not succeed in finding their nests," 

In 1936, we found these ducks to be abundant in Nushagak 



100 NORTH AMERICAN FAUNA 6 1 , FISH AND WILDLIFE SERVICE 

Bay on May 26, and we observed several on lower Ugashik River 
on May 27 to 29. 

Jaques (1930) noted them as abundant in southeastern Alaska, 
May 1 to 9, 1928, and saw a few about Port Moller, May 22 to 
June 2, 1928. 

In 1925, I found them about the western end of Alaska 
Peninsula: At King Cove, April 25; plentiful at False Pass, 
April 28; a flock in Bering Sea near St. Catherine Cove (Unimak 
Island), May 17; and a few near Izembek Bay, May 20. As 
late as July 27 a few (possibly nonbreeders) were found along 
the coast in the vicinity of Izembek Bay. 

We did not observe white-winged scoters in the Aleutians 
west of Unimak Island, but Wetmore (manuscript notes) re- 
ported "a great raft" of these birds at Tanaga Island, June 25, 
1911, and he noted small flocks in Kiska Harbor, June 17-21, 
1911 ; Gabrielson noted a few at Akun, July 9, 1941. 

A. C. Bent (1925) suggests that the species may possibly 
breed in the Aleutians, basing his conjecture on these summer 
observations. This is possible, not only in the Aleutians but 
also on the peninsula, especially before the introduction of blue 
foxes on the islands. However, we have no nesting records for 
this entire district. 

According to general information and statements of natives, 
white-winged scoters winter in large numbers in the Aleutians 
and along the Alaska Peninsula. In 1941, Beals and Long\vorth 
(field report) recorded these ducks at intervals from January 
12 to June 12 in the region about eastern Unimak and the 
adjacent Alaska Peninsula, and Gabrielson recorded wintering 
birds from Kodiak to Unimak. 

Cahn (1947) writing of Unalaska Island, says: "An abundant 
fall and winter visitor, especially from December to February." 
And Taber (1946) found a few of these birds wintering at 
Adak. Sutton and Wilson (1946) observed one scoter at Attu, 
March 17, 1945. 

G. H. Mackay in 1891 (quoted in Bent 1925) gave an interesting 
account of a mass migration of white-winged scoters to their 
nesting grounds, as observed in Rhode Island. He stated that 
it generally took place about the middle of May and that the 
daily flight was begun in the afternoon. 

We observed a similar occurrence on the other side of the 
continent when we visited Nushagak Bay in 1936. As we went 
up this bay on May 23, we saw large numbers of white-winged 
scoters assembled there, some of them flying about in pairs. 
The following is quoted from our field report; 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 101 

On the evening of May 26, as we were going back out through Nushagak 
Bay, we observed flock after flock of white-winged scoters flying high in 
the air in goose-like formation, all heading up the bay in a general 'inland' 
direction. Some flocks contained 75 or 100 birds. It appeared that we were 
witnessing a movement, en masse, from a temporary salt-water meeting 
place to the inland nesting grounds. 

Melaniffa persp'icillafa: Surf Scoter 

We observed this bird in considerable numbers in the spring, 
on the way to the Aleutians, along the coast of southeastern 
Alaska. They (apparently all males) were particularly numerous 
on the south side of Millbank Sound on April 25, 1936. One 
large flock arose from the water and strung out for a mile. 
It must have contained at least 1,000 birds. There were other 
smaller flocks. We saw 4 or 5 of these birds at Port Chatham, 
Kenai Peninsula, on May 6, which was the last sighting. 

Surf scoters are known to occur at Kodiak Island. Cahalane 
(1944) says: '*A11 of the surf scoters seen were on the Shelikof 
Strait coast of Katmai National Monument, Oct. 4 to 7. They 
were 'common' in Kinak Bay, but were abundant from Katmai 
to Amalik Bay and in Kaflia and Kukak Bays." Gabrielson also 
observed them at Kodiak in early spring and fall. 

Laing (1925) observed these birds in spring as far west as 
Dolgoi Island, south side of Alaska Peninsula, but he saw none 
west of there. 

On July 23, 1940, Gabrielson noted four old males up the 
Kvichak River, and in winter he saw a few at Unimak. 

Wetmore (manuscript notes) reported, "A small flock of 
scoters, that I took to be this species, was seen June 4 in Lost 
Harbor, on Akun Island, and others were seen June 10 and 11, 
in Chernofski Harbor (Unalaska Island)." He reported none 
west of that point. 

Dall (1873), referring to the surf scoter under the name 
Melanitta velvetina, says: "Killed Oct 27th, 1871, at Unalaska, 
and noticed at intervals there during the winter. It was not 
seen at the Shumagins, though it may occur there. A winter 
visitor." 

Beals and Longworth reported a single male as False Pass 
on March 1, 1941, remarking that they saw this bird on several 
occasions. 

Cahn observed 3 scoters at Captain's Bay, Unalaska Island, 
April 3, 1943, and saw 1 on March 16, 1945 ; Taber saw 1 at Adak, 
December 14, 16, and 23, 1945. 



102 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Austin Clark (1910) reports that "A few were seen at Attu 
and Agattu." 

Turner is the only observer who states that the surf scoter 
is "common among the Aleutian Islands," and is "abundant" in 
winter. He also says that "The surf duck is the svestu'n or 
whistler, of the Russians." There is much confusion in Turner's 
account. While the surf scoter does make a whistling sound with 
its wings, the vocal whistling is so characteristic of the common 
scoter that if any 1 of the 3 scoters should be so designated, 
it should be Oidemia n. americana. If sound of wings is the criter- 
ion, then deglandi is outstanding. Furthermore, Tunier (1886) 
says of 0. n. americana, "The male is noted for the gibbosity of 
pinkish-white near base of bill ; the lower edge of the swelling is 
deep red, gradually blending with the black of the rest of the bill." 
Assuredly, this fits perspicillata and not americana, and testifies 
to Turner's confusion on these species. 

In any case, the surf scoter is comparatively scarce today in the 
Aleutians, 

Oidemia nigra: Common Scoter 
Oidemia nigra americana 

Attu : Hoo-vai-ach 
Atka: Koo-ghang-ach 

Russian (reported at Unimak) : Swiss-toon (No doubt the svestun applied 
by Turner to perspicillata.) 

Laing (1925) observed the common scoter at Kodiak, March 
21, 1924, and Friedmann (1935) has recorded a number of 
specimens from Kcdiak Island, though we do not have nesting 
records from there. Cahalane (1943) noted a small number 
of these birds in Viekoda Bay in the fall of 1940, and he saw 
a larger number in Uyak Bay. He also reported that this scoter 
was numerous in the fall of 1940, along the coast from Katmai 
to Amalik Bay, but he reported that noticeably fewer birds were 
seen north of this area. A few were noted in Kaflia and Kukak 
Bays. 

Osgood (1904) reported a few broods of young on ponds near 
Lake Clark, and he adds "Females with young were also seen 
occasionally along the more sluggish courses of the Chulitna 
River." 

On July 19, 1940, Gabrielson noted adults on Naknek River, 
and he saw a female with three young on a small lake at Egegik. 
On July 23, he noted three broods up Kvichak River and noted 
the species again near Iliamna Lake on July 24 to 26. 

On May 23, 1936, we noted 15 or 20 males among large numbers 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 103 

of white-winged scoters in Nushagak Bay. These probably were 
migrants. On May 29, several flocks were flying about at the 
mouth of Ugashik River, and on the adjacent marshes two pairs 
were discovered among the ponds, the males whistling. Ap- 
parently, these birds were preparing to nest. 

Jaques (1930) observed them "about Moller Bay and on fresh- 
water pools on the tundra. May 23 to June 20 — not common." 
This, too, suggests nesting. 

In 1925, I saw this species at the western end of Alaska Penin- 
sula — a flock of both sexes at King Cove, April 25, and a few 
at False Pass on April 28. On June 13, small bands were flying 
about on Izembek Bay, whistling. By June 20, the birds were 
generally paired; on June 13, a female, taken for a specimen, 
contained a perfectly formed, hard-shelled egg. There can be no 
question about these birds nesting at Izembek Bay, chiefly at 
Ilazen Point. 

At Unimak Island we were told that this duck nests at Swanson 
Lagoon, which would be expected. 

Laing (1925) observed this duck at Dolgoi Bay, March 23, 
and says: "From Unalaska, where twenty-five were seen on 
March 26, the species was present in most of the harbors as 
far as Hitokappu in the southern Kurils, May 7. It was noted 
at Copper Island, Oest, Kamchatka, or Petropavlovsk." This 
statement indicates that it was noted along the Aleutian chain. 

Bishop (1900) recorded a number of these ducks off Unalaska, 
October 5, 1899. 

The Atka natives stated that this scoter winters sparingly in 
the Aleutians, while the Attu natives said that it was abundant 
there in winter. This is also borne out by Gabrielson's observa- 
tions on wintering birds from Kodiak to Atka. 

Beals and Longworth noted common scoters quite often in vari- 
ous places about the east end of Unimak Island from January 19 
to June 12, 1941. 

Cahn (1947) reported for Unalaska Island: "Common in very 
large flocks in all the major bays from December to February, 
inclusive." Taber (1946), writing of Adak for the winter of 
1945-46, states, "This was the most common bird of the area; 
it was seen in groups of 2 to 70 on the salt lagoon and the open 
sea." Sutton and Wilson (1946) found it wintering commonly 
at Attu. 

Bent (1925) records this bird as nesting in the Aleutian 
Islands; this is verified by the A.O.U. Check-List (fourth edition). 
It is possible that both statements are based on Turner's account. 



104 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

But, considering (1) the apparent confusion of the three scoters 
by Turner (shown by his description and misplacement of the 
Russian names) ; (2) that Bent and his party did not observe 
it nesting there in 1911; (3) that we did not find any evidence 
of it nesting there on expeditions throughout the chain, when all 
islands were examined; and (4) that none of the Aleuts re- 
ported it nesting; then we must conclude that nesting of the 
common scoter in the Aleutian Islands as a whole must remain 
in doubt. 

Mergus merganser: Common Merganser 
Mergus merganser amer'icanus 

Attu: Chu-vai-ach, Siss-uch 

Tan-nnm-aJi-tiim sak-oi-a 

Chung-ung-e-koo-loo-ghearch 

Ha-Ka chai-u-too 
Russian, Commander Islands: Bolschoj Krachal (Stejneger) 

Friedmann (1935) records a number of specimens from Kodiak, 
as well as a number of eggs, which he said to be those of the 
common merganser, and he quotes Brethei*ton as saying that 
this duck nests on Kodiak. 

Osgood (1904) had very little information on this merganser 
for the base of the Alaska Peninsula, but he mentions an adult 
male killed at Becharof Lake. 

Cahalane (1944) observed several on the Naknek River on 
September 4, 1940. 

Jaques (1930) found flocks of these ducks (most were males) 
near Port Moller in late May and June, but he saw no sign of 
nesting. 

In 1936, we were informed by residents at Chignik that two 
kinds of mergansers occur there. 

A number of records of occurrence are available for Unalaska, 
probably because it has always been a prominent port where 
vessels put in during voyages through that region. Dall (1873) 
said several specimens were taken there on December 20, 1873, 
and he adds that none were seen in the Shumagins. Turner 
says they winter at Unalaska, but do not breed there. Eyerdam 
(1936a) reports that two birds were collected at Unalaska on 
June 10 and August 6, 1932. 

We saw no common mergansers in the Aleutians. The chief 
of Attu Island, who furnished the series of names for this bird, 
said that a few common mergansers nest there but that they 
are more numerous in winter. 

To sum up, the common merganser occurs sparingly from 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 105 

Kodiak to Attu ; the best evidence of nesting comes from Kodiak 
Island; and (possibly) it nests on Attu Island. We know that 
it is an inland form — more so than M. serrator. 

Mergus serrafor: Red-breasted Merganser 
Mergus serrafor serrator 

Attu: Cruch-ah'-lich 
Atka: A-ga-lai-ahh 

Agldyax (given by Jochelson as applying to two species) 
Russian, Commander Islands: Krakhal (Stejneger) (The Attu name is 
undoubtedly a corruption of the Russian.) 

This is the commoner merganser of the Aleutian district. It 
breeds on Kodiak Island (Friedmann, 1935) , and Cahalane (1943) 
found it generally very abundant in the Kodiak-Afognak group in 
1940. He also observed it in various places in the Katmai region, 
where Hine (1919) also reported it to be common. 

Osgood (1904) found it "exceedingly abundant on all the lakes 
and rivers" visited at the base of Alaska Peninsula, and he 
mentions seeing broods of young on Iliamna, Chulitna, Kakhtul, 
and Nushagak Rivers. He remarks, "From start to finish probably 
more mergansers were seen than any other species of water 
bird, with the exception of the large gulls." 

Gabrielson also noted this duck in 1940 on the rivers tributary 
to Bristol Bay. There were at least 50 broods of young, in all 
ages, on the Kvichak River, July 23. 

This merganser was reported as common at Chignik, and 
Jaques (1930) found it paired on King Salmon Creek, near Port 
Moller after June 11, "possibly breeding." 

On May 26, 1936, we saw two females in Nushagak River at 
Snag Point, and a pair was seen back on the marshes among the 
lakes near Ugashik River, where they probably nest. 

In 1925, I found this merganser nesting about Izembek Bay, 
and, on May 25, 1925, 4 were seen on a mountain stream below 
Aghileen Pinnacles. (On May 4, and on several subsequent days, 
red-breasted mergansers were noted at Urilia Bay, on Unimak 
Island.) On July 5, a nest with six eggs was found on a small 
island near Point Grant, and another nest was found on a little 
island far out in Izembek Bay, in the midst of a colony of 
glacous-winged gulls. Red-breasted mergansers with molting wing 
feathers were seen late in July. 

Chase Littlejohn, in 1887-88, said that this duck breeds at 
Sanak and at Morzhovoi Bay, where they remained all winter. 

McGregor (1906) found three nests on Round Island, Beaver 
Inlet, Unalaska Island, July 4, 1901. On June 3, 1936, we saw 



106 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

six of these mergansers at Unalaska — Wetmore also had observed I 
them here on June 6 and 7, 1911, and had collected a specimen. 

At Unalaska Island, Cahn (1947) found a brood of 9 young | 
in the Makushin Valley swamp, June 23, 1944, and he observed 
a brood of 11 downy young on Coxcomb Lake, July 4, 1945. 

On August 15, 1937, we flushed a female from a grass-topped 
islet off the shore of Amlia Island. We had found 3 pairs oni 
Kiska Island, June 4 and 5, where Wetmore had seen 1 pair 
in June 1911. We found a foot of red-breasted mxerganser in an 
eagle's nest on Buldir. On June 17, we saw a flock of 7 at Semichi 
Islands; 6 were noted on Amchitka Island. Incidentally, Dall 
(1874) had reported that Amchitka was the only place in the 
western Aleutians where this species had been observed. 

In 1936, we noted a flock of seven red-breasted mergansers on 
Corwin Lake, Atka Island, June 22. Several were seen on 
Kanaga, June 29, and eight were seen in a lake on Kiska, July 
26. At Adak Island, July 3, two were seen in Bay of Islands, and 
three or four in Kuluk Bay. June 26-27, 1911, Wetmore found 
them to be fairly common in the small lakes back of Bay Water- 
falls, Adak Island, where he found a brood of nine downy young 
about a week old — he suspected that there was a brood in another 
lake. And on September 3, 1944, Gabrielson found a brood on 
Amchitka, thus definitely establishing a nesting record for that 
part of the Aleutian chain. 

The Attu chief said that these ducks nest on Attu, and Atka 
natives reported them nesting on their island. Turner also re- 
ported them nesting on Atka. 

We can definitely state that the red-breasted merganser nests 
from Kodiak to Attu, and, according to Stejneger (1885), it is 
a very common breeding bird in the Commander Islands. 

Apparently, it winters in the Aleutians also (though perhaps 
in small numbers), because Taber (1946) observed them at 
Adak from December 9, 1945, to January 13, 1946. 



Family ACCIPITRIDAE 

Accipifer gen f His: Goshawk 
Acc'ip'ifer gentilis africapillus 

The goshawk occurs on Kodiak Island, as shown by specimens 
recorded by Friedmann (1935). Harrold saw one on Sitkalidak 
Island, near Kodiak, in May 1927 (Swarth 1934). Howell (1948) 
found a goshawk nest July 9, 1944, located in a 35-foot spruce at 
Middle Bay, Kodiak Island — there was a single young, which- flew 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 107 

from the nest. Osgood (1901) saw goshawks frequently near 
Tyonek, and two immature birds were collected. Osgood (1904) 
again reported a goshawk at Iliamna Pass, July 13, 1902, and 
several immature birds were observed repeatedly at the mouth 
of Chulitna River. Later, he observed the species at Nushagak. 

This sums up normal distribution of the goshawk in the region 
under discussion, though on August 15, 1946, Gabrielson recorded 
one at Dutch Harbor, and, on August 20, he noted another at 
Simeonof Island in the Shumagins. Ordinarily, the goshawk is 
confined to the Kodiak-Afognak area and the base of the Alaska 
Peninsula — the regions that contain the forested areas. 

AcdpHer sfrJafus: Sharp-shinned Hawk 
Accip'iter striatus velox 

Friedmann (1935) records a specimen collected by Bischoff on 
Kodiak Island, March 10, 1869. Osgood (1904) reports seeing 
a sharp-shinned hawk on the Mulchatna River, September 3, 1902 ; 
apparently, these are the only records for the area here con- 
sidered — this bird sharing the forested areas with the goshawk. 
But Swarth (1934) reports a specimen taken on Nunivak Island, 
north of the area here considered, on September 14, 1927. This 
bird was found among the boulders on the shore, far from any 
forest, which is a most unusual occurrence. 

Bufeo lagopus: Rough-legged Hawk 

Buteo lagopus s.johannis 

We observed the rough-legged hawk at Kodiak and Afognak 
Islands. Friedmann (1935) has recorded a number of specimens 
in both light and dark color phases from Kodiak, and he mentions 
Bretherton's statement that this species nests there. Osgood 
(1904) reports a nesting pair on an islet in Lake Clark, and he 
observed one bird near the mouth of Chulitna River and another 
on the lower Nushagak. McKay took a specimen in 1881 on the 
Aleknagik River. 

Cahalane observed these hawks on the wxst side of Alaska 
Peninsula, on Naknek River and Three Forks, in September 1940, 
and, in the same year, Gabrielson noted one at Kodiak, June 14, 
and one at Dillingham, July 17. 

Gianini (1917) observed these hawks nesting in "fair num- 
bers," in Stepovak Bay, in 1917. In 1911, Wetmore found them 
to be fairly common near Frosty Peak, and he noted one at Un- 
alaska. 

In 1925, I found a number of nests on cliffs about Izembek 



108 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Bay and at least 4 nests on Amak Island, and I noted five or si t 
hav/ks on Unimak Island. In 1936, when our party visited Ama f 
Island very briefly, two rough-legged hawks were noted ther[ 
again. They occur also in the Shumagins, because we saw on| 
at Unga Island. 

Rough-legged hawks have been noted by various ornithologist! 
in the Fox Islands group. We saw them on East Unalga, Ur 




Figure 25. — Rough-legged hawk. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 109 

alaska, Umnak, and Ananiuliak (the last is a smaller island off 
the west end of Umnak). Swarth (1934) reports that these 
hawks were seen almost daily on Akutan from May 17 to June 
13, 1927, by Harrold, nesting and in both color phases. He 
also found this species nesting at Unalaska. 

Our own observations, and the published record, show that 
rough-legged hawks nest along the Alaska Peninsula and on 
suitable offshore islands, and westward in the Aleutians as far 
as Ananiuliak Island — but no farther. It is significant that this 
breeding range coincides exactly with the distribution of rodents, 
for no rodents originally occupied the Aleutian Islands west of 
Ananiuliak. Rats and ground squirrels have been introduced on 
a few islands to the westward, but evidently these introductions 
have not yet affected the original distribution of the rough- 
legged hawk. 

Rodents constitute the chief item in the diet of these hawks, 
as was verified by a number of observations. Speaking of the 
area about Frosty Peak, Alaska Peninsula, Wetmore reported 
in 1911: "The thousands of ground squirrels (Citellus) here fur- 
nished them an abundant food supply as the crops of those taken 
testified." 

On Amak Island, in 1925, I found a quantity of mouse fur, 
three Microtus, and the wing of a Savannah sparrow in a rough- 
legged hawk's nest. Microtus amakensis is the only rodent there. 
The stomach of a female hawk collected by Harrold on Akutan 
Island contained two field mice Microtus. 

Stejneger (1885), speaking of Archibuteo kigopus, said that it 
was occasionally seen in the Commander Islands, and he thought 
that it might become established there, because mice had been 
introduced. 



Aquila chrysaefos: Golden Eagle 
Aqu'ila chrysaefos canadensis 

Both Turner and Dall reported the golden eagle to be abundant 
in the Aleutians. Austin H. Clark (1910) reported: "I observed 
this species once on Unalaska and several times on Atka, where 
it appears to be rather common." 

Chase Littlejohn (manuscript notes), speaking of the area 
from the southwest end of Kodiak Island to the end of Alaska 
Peninsula, including adjacent islands, says. 

Saw quite a number of these fine birds but only obtained one, which was 
unavoidably lost to my collection. He was caught in a steel trap. A couple 



110 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

of days before, he had killed and eaten a silver fox whii^h was in a trap. 
It seems he returned to pick up the fragments and was himself caught. 
He measured nine feet from tip to tip. 

The exact locality was not given. 

In 1936, Douglas Gray and C. S. Williams saw an eagle at 
Unalaska, which they described as having considerable white 
on the tail, but with a temiinal dark band. This assuredly 
suggests that the bird was a golden eagle. At Chignik we were 
told that one had been killed there, but we did not see the 
specimen. 

Cahn (1947) writes, "While probably not rare in the higher 
and wilder parts of Unalaska Island, this species is uncommon 
around Dutch Harbor, Two records in four years: June 17, 
1944, over Mt. Ballyhoo, and August 7, 1944, sitting atop a 
mast on a ship anchored at a dock." 

Osgood (1904) refers to a specimen that was supposed to 
have been collected by McKay at Nushagak, but he was unable 
to find it in the National Museum collection. 

Friedmann (1937) found a sternum of this species in midden 
material from Kodiak Island — the only record for that locality. 

Thus, we have quite a number of records (mostly based on 
observations) ; however, authentic specimens are rare. It is a 
little difficult to conceive of the golden eagle as abundant in the 
Aleutians, in view of observations dating back to the time of 
Turner and Dall, but there seems to be ample evidence to conclude 
that at one time the bird was more common that it is today. 
It is now only an occasional straggler in the Aleutian Peninsula 
region. 

Haliaeefus albicilla: Gray Sea Eagle 

Friedmann (1935) records osseous remains of the gray sea 
eagle from village middens on Kodiak Island. Bishop (1900) re- 
ported the first record of this bird for North America — a young 
female that was found dead at Unalaska, October 5, 1899. 
Again, in 1905, he records a specimen that was taken at Van- 
couver Island, March 18, 1898. 

Eyerdam (1936) says, "Several of these birds were seen on 
Unalaska Island on May 25th and May 30th, One was killed 
near Dutch Harbor by a seaman from one of the coast guard 
cutters, who kept the claws, tail and wing feathers for souvenirs." 
It is unfortunate that a specimen was not saved, since it is 
rather remarkable to casually see "several" of a species so rare 
in North America. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 111 

Cahn (1947) reports seeing one of these birds at Dutch Harbor 
on May 16, 1945, and he reports that he watched it under 
favorable circumstances for 10 minutes. Sutton and Wilson, at 
Attu, watched two dark-headed, white-tailed eagles, identified as 
this species, on March 15, 1945. 

We did not see this bird on any of our expeditions. A 
number of times we thought that we had sighted one, but each 
time it proved to be a bald eagle in one of its immature plumages. 
These plumages can be confusing, and we felt that records of 
the gray sea eagle should be based on specimens. 

Haliaeetus leucocephalus: Bald Eagle 
Haliaeetus leucocephalus alascanus 

Attu : Tirrgh-hich 
Atka : Tig-a-lach 

A-waich'-7-ich (immature) 
Alaska Peninsula: Tikh-lukh (Wetmore) 

The bald eagle is commonly distributed throughout the length 
of Alaska Peninsula and adjacent island groups, and the Aleutian 
chain. It is numerous in some places. In the Aleutians, nearly 
every island that we visited had at least 1, often 2 or more, 
pairs, nesting. They are numerous about the larger islands. 
Williams noted 15 eagles in Bay of Islands, Adak Island, July 2, 
1936, and more were found on other parts of the island. On June 
29, we saw several at Kanaga Island. The caretaker of a fox- 
ranching establishment there had killed 14 of these eagles for 
the bounty, and he planned on raiding 20 more nests later. 

For some reason, the bald eagle is scarce in the Near Islands — 
including Attu, Agattu, and Semichi. We observed a single pair 
on Agattu in 1937, but we saw none at Attu or Semichi and the 
natives assured us they were very scarce. However, we found 
a nest on Buldir Island, and from that point eastward bald 
eagles were common. 

Not only do eagles occur along the Alaska Peninsula, they also 
occur on the ofi'shore island groups. In 1940, Gabrielson observed 
them in several places at the base of Alaska Peninsula. At 
Kodiak, in 1936, one merchant erected a sign advertising the 
fact that eagle feet were acceptable as cash (bounty could be 
collected for them). 

Plumage and Other Color Changes 

Too few specimens were handled to obtain precise information 
on plumage changes. A. C. Bent (1937) states that he believes 
the bald eagle assumes the adult plumage in the fourth year. 



112 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE , 

Field observations on numerous immature birds in Alaska were' 
confusing, and we were unable to correlate some plumage patterns 
with age. 

The downy-young plumages are well known and are well de>- 
scribed by Bent. However, the color of beak, eyes, and other 
soft parts is not so well known. A young bird in the dark-down 
stage on Ananiuliak Island had a slate-colored upper mandible, 
the cere was of a similar color, but it was of a little lighter 
shade. The tip of the lower mandible was similar to the upper 
in color, but posteriorly the margin of the gape was flesh color, 
becoming paler posteriorly and shading into a near-yellow at thei 
corner of the mouth. The lores were dull bluish. The iris wasi 
dusky brown, and the pupil was blue. The eyelids were pale 
plumbeous. The feet were a yellowish-clay color, and the claws 
were slaty. 

The flrst-year plumage is dark; as Bent says, "uniformly 
dark 'bone-brown' to 'clove brown' above and below; the flight 
feathers are nearly black, but there is usually a slight sprinkling 
of grayish white in the tail." In the first year, both the bill and 
cere are of a blackish-slate color. The iris is brown, and the 
pupil is black. At this stage, the eyelids are still plumbeous. 

The plumages preceding the final adult stage are hard to 
define. There appears to be much variation, probably over a 2- 
year period. Assuming a 2-year period for the postjuvenal 
phases, the plumage varies in the degree of white mottling. 
The essential feature is a pattern that includes patches of dull- 
white mottling on scapulars and back (which, in flight, show as 
three distinct areas), and light-colored upper tail coverts and 
considerable white in under parts. In one phase of this plumage, 
which must be in the second year, the bill and cere are still 
blackish and the eye is still a rich brown. The preocular area 
is essentially white, the eyelid is plumbeous, and the gape is 
dull yellowish. The feet are yellow. 

A later phase, which possibly may represent the third year, 
still includes the dark bill, with a dull-yellowish hue appearing 
on the lower mandible and the margin of the cere. The eye is 
dull yellow also, and a yellowish tinge is encroaching upon the 
preocular area. The eyelid is gray, and the gape is yellow. There 
is much light speckling on the head, though the head is chiefly 
brownish. The specimen on which this description is based did 
not have the light mottling on upper parts falling into a pattern 
of three light patches, as was seen on many birds; instead, it 
was more scattered. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 113 

In still another phase, which is quite advanced, the head is 
white, speckled with a blackish hue. The beak is a dull-yellowish 
tone — perhaps best designated as tan, somewhat streaked with 
a slaty tone. The lower mandible is bright yellow at the base. 
The cere is a mixture of gray black and yellow. The eye is 
yellow (as in the adult), the eyelid is a brighter yellow, the 
preocular area is pale yellow, and the gape is a rich, bright 
yellow. 

These are the advancing stages in development, the transition 
from dark "soft parts" to the characteristic yellow of the adult, 
but it was not possible to allocate all of these plumages to age 
groups. 

Nesting 

Trees are absent in the area except in a limited portion of 
the base of Alaska Peninsula, therefore nests are placed on cliffs 
or pinnacles, or on low ground. Many nests are inaccessible to 
man by ordinary means of climbing. Frequently, a nest is placed 
on the top of a pinnacle, which sometimes is separated from an 
adjacent cliff by a narrow chasm, and which is surrounded by 
water, at least at high tide. At times, the nest is placed on a 
cliff, where it may be fairly accessible to man. In one case, on 
Buldir Island in 1936, a nest was found on a small rock outcrop 
on a slope, where one could walk to it without climbing. The same 
place was visited the following year; the former nesting site 
was abandoned, and the eagles (probably the sam.e pair) had 
made their nest on the flat grassy valley bottom below. There 
was not even a hummock at the nest location. 

In 1925, on Unimak Island, a nest containing eggs was placed 
on the top of a smooth sand dune. It is interesting to note that 
on June 9, 1941 (16 years later), Beals and Longworth re- 
ported finding an eagle's nest on a sand dune in the same 
locality. As a rule, eagles seek inaccessible locations on cliffs and 
obviously prefer pinnacles. 

Nests are generally built by assembling a layer of dried grasses, 
mosses, and other vegetable debris. Sometimes kelp is used. 
Kelp nests are rimmed with the dried stems of Heracleum and 
Ligusticum, which are the largest material available in lieu of 
twigs from trees. In some cases, however, the eagles use sticks 
from the driftwood on the beach. 

Eagles build various types of nests. The nest on the sand 
dune, already mentioned, consisted of a cavity that was 360 
mm. wide and 130 mm. deep, heavily lined with dry grass, bits 
of moss, and a small amount of dead eelgrass from the beach. 



114 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

A number of large dry stalks of Heracleum lanatum lay around 
the rim, though these were not used in the construction of the 
nest proper. 

Another nest was on a rock mass rising from a slope on Amak 
Island. A few dried plant stems were the only evidence of nest ! 
construction, and the single young sat on a bare spot, well 
trampled, on top of the rock, surrounded by a fringe of green 
grass. 

Another nest on the same island was somewhat similar. It 
was on the grassy top of a high cliff. Two well-feathered young j 
perched in a bare trampled spot about 8 feet long, which wasi 
crescent-shaped because of a hump in the middle of the space. 
There were the usual dry cow parsnip stems around the edge, 
but there was practically no nesting material in the center. 

A third nest on this island was more substantial, consisting of 
dry grass with dry cow parsnip stems around the rim. 

These scanty nests contrast sharply with a nest found at 
Amukta Island, June 16, 1936. This nest, on top of a pinnacle, 
was built of kelp, grasses, and driftwood to a height of 4 feet. 
A nest observed at Kanaga Island, June 29, 1936, was on the 
grassy top of a pinnacle ; it was made mostly of moss and had 
a wide platform rimmed with dry stems of Heracleum and 
Ligusticum and a few driftwood sticks. 

A nest on the grassy top of a columnar rock on the shore of 
Kiska Island was in the form of a bulky mass, consisting mostly of 
kelp. 

Still another nest, on a rocky point of Little Sitkin, was built 
largely of dry stems of Heracleum and Ligusticum and willow 
roots, with a lining of finer vegetation. The willow there is a 
prostrate form, whose roots often are partly exposed by wind 
erosion. 

These examples illustrate the general type and the variations 
of bald eagle nests. Some of the bulky nests resulted from an 
accumulation of material over a long period — a typical example 
was found at Amchitka Island, July 11, 1937. This nest — a 
shallow affair — was made mostly of moss on the grass-topped 
point of a pinnacle rising from the beach. It rested on a mass 
of old sod and soil to a depth of about 6 feet. This accumulation 
was filled with bird bones. Evidently, this accumulation had been 
built up by annual increment of debris left by nesting eagles 
for many seasons. 

Our various expeditions were usually too late in the season 
to observe eggs — there were young in nearly every case. The 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 115 

number of young, in a series of 34 nests, varied from 1 to 3 per 
nest, though in 1941 Beals and Longworth reported a nest with 
4 young. In 1 nest, there was 1 live youngster and 1 partly 
eaten dead youngster; in 2 other nests, there was 1 young and 
1 rotten egg containing an embryo. All of these must be con- 
sidered as having had two fertile eggs originally. On that basis, 
there were 12 nests with 1 young, 17 nests with 2 young or eggs, 
and 5 nests with 3 young. 

In every nest that we observed, the nesting birds were white- 
headed adults. One report, from Cecil Williams in 1936, indicated 
a nesting pair, in immature plumage, on TJliaga Island. 

Food Habits 

I have discussed the food of this eagle in detail in "Food 
habits of the northern bald eagle in the Aleutian Islands, Alaska" 
(Condor, 1940, vol. 42, No. 4, pp. 198-202). The data presented 
were based on examination of 28 nests. In addition to this 
published material, data from 4 other nests are available, com- 
prising 21 more food items. This additional material agrees with 
the published percentages. 

In the Aleutian district, birds constitute the major part of the 
bald eagle's diet — 58.9 percent on the basis of material obtained 
in 1936; 86 percent for 1937. As would be expected, most of 
the birds taken are the so-called sea birds, chiefly shearwaters, 
fulmars, cormorants, glaucous-winged gulls, murres, ancient 
murrelets, paroquet auklets, crested auklets, and horned and 
tufted puffins. Fulmars and shearwaters head the list. Two 
ravens had been eaten. Others taken included : Petrels, kitti- 
wakes, pigeon guillemots, ptarmigan, least auklets, and ducks, 
though none of these are taken in great numbers. Ducks were not 
preyed on extensively, probably because of the abundance of other 
birds, although harlequin ducks, oldsquaws, European teals, 
pintails, common eiders, red-breasted mergansers, and three 
emperor geese were identified in food remains. 

Mammals are not universally available to eagles in this district 
and are seldom found in the diet. Tlie ground squirrel is by far 
the most common mammal captured. Others, which occasionally 
are taken, are the house rat, the field mouse, the blue fox, and, 
possibly, the domestic sheep at Umnak Island. In 1938, Scheff'er 
reported that one of the men in charge of the sheep on Umnak 
Island declared that he had never seen eagles bothering live sheep, 
though they will eat carrion. Another informant, a sheep herder 
at Unalaska, said that eagles will not bother healthy sheep, 
but they will attack dying ones and will feed on dead ones. 



116 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

He had seen both ravens and eagles feeding on carcasses of 
winter-killed sheep, Beals and Longworth, in 1941, reported 
that local residents on Unimak Island believed that the bald 
eagle kills caribou fawns. However, this would need verification. 
It is known, of course, that eagles feed on dead whales and seals. 

It has been thought that bald eagles kill many blue foxes. 
But, according to the evidence we obtained, this is not the case 
in the Aleutian district. The remains of only one fox were found 
in an eagle nest, and these remains could have been carrion 
because we found a few dead foxes on the beaches. To further 
refute this theory, many blue fox families were being raised 
successfully in the vicinity of eagle nests. 

A moderate percentage of fish and invertebrates is eaten by 
the bald eagle. To what extent this eagle feeds on dead or 
spawning salmon on the Alaska Peninsula was not determined. 
In July 1911, at Morzhovoi Bay, Wetmore observed them feeding 
on dog salmon taken from shallow rapids. Edward D. Crabb 
(1923) apparently found fish remains to be prominent in nests 
examined along Alaska Peninsula; there were parts of seven 
Dolly Varden trout in one nest. Edward J. Reimann (1938) 
observed a bald eagle taking a mullet out of the water, reaching 
for it with one foot. Beals and Longworth found two sockeye 
salmon and the head of a sea gull in a nest on Unimak Island, 
June 9, 1941. We did not see bald eagles capture live fish, but 
Atka mackerel were often observed near the surface of the 
water, where an eagle could very easily seize one. 

In the Aleutian chain proper, the main food of the bald eagle 
consists of sea birds. There are some indications that fish of 
various kinds are more prominent in the diet along the Alaska 
Peninsula, where we did less work on this bird. At any rate, 
there is abundant evidence that the eagle is not a serious detri- 
ment to man's interests throughout the Aleutian district. 

Banding 

A number of nestling bald eagles were banded in the Aleutian 
Islands in 1937. Of these, six returns were obtained. All six 
had been banded in June ; 1 on Little Kiska Island, 2 at Little 
Sitkin, and 3 (all in one nest) on Rat Island. The following winter, 
all of these were killed by natives on Attn Island. This shows a 
westward drift of immature eagles, at least in the western part 
of the Aleutian chain. 

These eagles are permanent residents in the Aleutian district, 
summer and winter. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 117 

Haliaeefus pelagicus: Steller's Sea Eagle 

In the course of all our expeditions to the Aleutians, a Steller's 
sea eagle was never observed, though we scrutinized all eagles 
closely for such a possibility. Charles H. Gilbert's specimen from 
Kodiak Island (1922) is the only record of a specimen obtained 
from the Aleutian district; however, more recently, Friedmann 
(1935) has recorded several bones of this species from middens 
on Kodiak Island. G. Dallas Hanna (1919, 1920) has recorded 
a specimen taken in the Pribilofs in December 1917. These are 
the only records for North America based on actual specimens. 
Austin H. Clark (1910) reported seeing one of these eagles near 
Unalaska on May 26, 1906. 

Leonard Stejneger (1885) says of this eagle: "The habitat is 
especially the mainland of Kamschatka, where it is abundant, 
but also all the countries bordering the Okotsk Sea. On Bering 
Island it is only an occasional visitor, being chiefly an inland 
bird preferring the quiet rivers and lakes surrounded by dense 
forests." 

Circus cyaneus: Marsh Hawk 
Circus cyaneus hudsonius 

Friedmann (1935) records a specimen taken on Kodiak Island 
by Bretherton on April 2, 1894. Osgood (1901) reports the 
marsh hawk near Homer and Hope, in the Cook Inlet region, 
and again, in August 1902, he found them at intervals along the 
Kakhtul River and occasionally, all the way to Nushagak. Caha- 
lane (1914) observed 4 marsh hawks in Katmai National Monu- 
ment in September and October 1940; 2 of these were males. 

We did not see any of these hawks west of Kodiak Island 
during three expeditions, nor does Wetmore record any west 
of Kodiak Island in his field report for 1911. But Turner (1886) 
records a flock of 10 of these hawks at Unalaska, and he remarks 
that it is a rare summer visitor to Attn Island ; however, this 
statement is surprising in view of present-day information. Those 
observed at Unalaska must have been a migrant group. But 
Cahn (1947) contributes the valuable information that he ob- 
served a male in Makushin Swamp, Unalaska Island, June 7, 
1943, and saw a female over the swamp at the end of Captain's 
Bay on July 7, 1944. 

Although the marsh hawk may occasionally appear to the 
westward, it certainly prefers the meadows and marshes of the 
more wooded parts of Alaska, including the base of Alaska 
Peninsula and the Kodiak-Afognak group. Along the north side 



118 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

of Alaska Peninsula all the way to the west end, and on Unimal^ 
Island, numerous marshy areas with an abundance of mice and 
birds may be found ; yet, this treeless region generally is avoided 
by the marsh hawk. 

Family PANDIONIDAE 

Pand'ion haUaefus: Osprey 
Pandion hal'iaefus carolinensJs 

This bird has not been recorded from the Kodiak-Afognak 
Islands, but Osgood (1904) reports it to be quite common on 
nearly all watercourses that he has traveled, and he specifically 
mentions the Nogheling, Chulitna, and Kakhtul Rivers — all these 
are north of Lake Iliamna. 

Cahalane (1944) saw 2 American ospreys — 1 at Naknek River, 
September 4, 1940, and the other at the outlet of Brooks Lake, 
September 7. 

In 1940, Gabrielson saw 1 osprey at Wood River Lakes, July 
18; he observed 1 at Brooks Lake, July 19, and he saw another 
near the upper end of Iliamna Lake on July 25, 

There are no records of sighting the American osprey farther 
west, but Stejneger (1885) said that it is an occasional visitor 
in the Commander Islands and that is is very abundant in Kam- 
chatka. This is another bird that does not venture out into the 
treeless areas to nest. 

Family FALCONIDAE 

Faico rusficolus: Gyrfalcon 
Faico rusficolus uralensis 

Attu : Kns-SHm Ah'-ghu-Uch 

The Attu chief described a bird larger than the peregrine 
falcon, and gave us the above name. (If it were different from 
the peregrine falcon, and larger, it could hardly be anything but 
a gyrfalcon.) The chief declared that it nests and winters on 
Attu Island. Austin Clark (Collins et al. 1945, p. 37) says 
"Lieutenant Nelson, an experienced falconer, believes he saw 
gyrfalcons on Kiska, though only one, in the white phase, was 
identified with certainty." 

We did not identify this bird on any of our expeditions, noi 
did Wetmore record it. Nelson (1887), using the name FaJcc 
rusticolus gyr faico, said that it was very common along the 
Bering Sea coast, but less common in the Aleutian Islands. Swarth 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 119 

(1934), using the name Falco rusticolus candlcans, records a 
pair seen on Akutan and several on Unalaska by Cyril G. Harrold. 
Beals and Longworth report — 

March 2, False Pass: 1 falcon, very light, almost grey, on a cliff approxi- 
mately 1500 feet elevation. March 14, False Pass: 2 at elevation of 1800 
feet, color white. March 24, False Pass: 1 with color predominantly white, 
flying over alders back of cannery. May 13, False Pass: 2 almost pure white 
falcons at 1500 feet. 

These are all sight records, and one cannot be sure which 
form of gyrfalcon was represented. But there are several speci- 
mens in the National Museum that are referable to iiralensis: 
Three were taken at Nushagak, September 1, 1881, October 20, 
1881, and December 5, 1882; and a juvenile specimen was ob- 
tained from Herendeen Bay, taken July 15, 1890, by C. H. 
Townsend, and marked by Friedmann as "Prob. uralensis." Fried- 
mann has also recorded 2 from Kodiak Island, 1 taken by Fisher, 
September 18, 1882, and the other (no longer extant) by Panshin 
in 1871. 

On September 21, 1942, Beal obtained a specimen at Cold Bay, 
on Alaska Peninsula. 

Stejneger (1885) listed Falco rusticolus and Falco islandus for 
the Commander Islands. The former, he says, is not uncommon in 
winter — feeding chiefly on "the numerous field mice which now 
infest that island," — and possibly nests there. He states that 
F. islandus breeds there in limited numbers. 

Hartert (1920) records 4 white and 4 dark immature birds 
and 1 white and 3 dark adults from the Commander Islands, all 
taken in winter ; he lists them all under Falco rusticolus canclicans. 

Falco rusficolus obsoletus 

In the National Museum there is a specimen taken by McKay 
at Ugashik in 1881 ; it was identified by Friedmann as obsoletus. 
To what extent this bird occurs in the Aleutian district is un- 
known, nor do we know how many of this form were represented 
in the sight records listed under uralensis. 

Bond (1949) has thrown some doubt on the classification of 
western American gyrfalcons, but I have not had an opportunity 
to evaluate the situation. 

Falco peregrinus: Peregrine Falcon 
Falco peregrinus anatum 

We did not obtain specimens of anatum in the Aleutian district, 
though Friedmann (1935) records a specimen from Kodiak Is- 



120 NORTH AMERICAN FAUNA fil, FISH AND WILDLIFE SERVICE 

land, which he suggests may possibly have been a migrant or 
vagrant. Likewise, the duck hawks reported by Captain Ammann 
and Lieutenant Nelson on Kiska (Clark 1945, p. 36) would have 
to be migrants or vagrants if we are to retain the subspecificj 
status of these forms. There is, of course, the possibility that' 
these birds were wanderers from Siberia (F. p. calidus). 

Faico peregrinus peale'i 

Attu: Ah'-ghu-lich 

Atka: Ah'-ghu-lich 

Commander Islands: Agulekh (Stejneger) 

Russian, Commander Islands: Tschornij Jastrip, black hawk (Stejneger) 

The Aleuts of the Commander Islands speak the language of 
the Aleutian Islands, and, evidently, the name for this falcon 
is the same in all dialects. 

It is pretty well established that the nesting birds of the 
Aleutian chain are pealei, and that this form also nests in the 
Commander Islands. Probably the same form occupies the Alaska 
Peninsula and adjacent island groups. However, there is a specif 
men of F. p. anatum recorded by Friedmann from Kodiak Island,! 
and we did not collect specimens of F. p. pealei east of the Aleutian 
chain. 

We found this falcon to be common throughout the Aleutians. 
It nests on nearly all the islands, usually on high inaccessible 
ledges. It is a resident breeding bird and also winters in the 
Aleutians. 

As would be expected, the peregrine falcon feeds chiefly oni 
birds. Casual observations revealed that the least auklet and 
the crested auklet are taken — one falcon was seen carrying a 
crested auklet. It has been reported as capturing ptarmigan and 
shorebirds, but it is believed that sea birds furnish a large portion 
of its food. 

FaIco columbarius: Pigeon Hawk 
FaIco columbarius bendirei 

Friedmann (1935) records a number of specimens of bendirei 
from Kodiak Island, though, as he points out, Peters states in his 
check-list of the birds of the world that suckleyi is the breeding 
bird of Kodiak Island. The 1931 A.O.U. Check-List considers 
the Kodiak birds to be bendirei. Two specimens were obtained 
from Kodiak by Gabrielson, August 8, 1945. 

Osgood (1904) records specimens from the Nogheling and 
Chulitna Rivers, Nushagak Village, and Aleknagik Lake, and 
he observed others on the Kakhtul and Nushagak Rivers. He also 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 121 

reported them common in the Cook Inlet region — specifically men- 
tioning Hope and Tyonek (1901). 

Cahalane (1944) observed a pigeon hawk on lower Ukak River, 
September 9, 1940, and saw another on Windy Creek, September 
16. 

Hine (1919) also found these little hawks to be common in 
the region of Katmai Bay in the summer of 1919. 

The pigeon hawk is exceedingly rare farther west, but there 
are a few records available. Turner (1886) mentions a specimen 
taken at Unalaska in 1879. Bailey (1926) says "Hendee collected 
one at Unalaska Sept. 25, 1922, and saw another the next day." 

Captain G. A. Amman reported a pigeon hawk, not positively 
identified, on Kiska Island. 

Taber had an opportunity to observe a pigeon hawk rather 
closely at the military establishment on Adak Island on De- 
cember 9, 1945. 

It must be considered rare in the Aleutians, however. 

Hartert (1920) records a female Falco columbarius insignis 
collected on Bering Island, June 10, 1915. 

Falco sparverius: Sparrow Hawk 
Falco sparverius sparverius 

The only record of the sparrow hawk is the statement by 
Dall (1873) that one was killed at Unalaska in the fall of 1871, 
but it was not preserved. There are no other records of this 
species in the entire area under discussion ; therefore. Ball's 
inability to preserve the specimen is unfortunate. 

Family TETRAONIDAE 

Canachifes canadensis: Spruce Grouse 

Osgood (1901, 1904) found spruce grouse to be plentiful in the 
wooded portions of the base of the Alaska Peninsula and the 
Cook Inlet region. Friedmann (1935) refers to a specimen from 
Kodiak Island, which was mentioned by Baird, Brewer, and Ridg- 
way. Cahalane (1944) found this bird to be abundant in the 
spruce forests north of Mount Katolinat, in the fall of 1940, and 
saw evidence of its presence north of Savanoski River. The 
Kodiak Island record had been referred to the Valdez spruce 
grouse, C. c. atratus, by Friedmann, but it is not known what 
the Alaskan Peninsula birds would be referable to. 

This bird could not be expected to occur west of the forested 
portions of Alaska Peninsula. 



122 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Lagopus lagopus: Willow Ptarmigan 
Lagopus lagopus alascensis 

Aleut: Alladek (Wetmore) 

The willow ptarmigan, distributed throughout the Alaska Pen 
insula, is represented by two races, L. I. alascensis and L. I. murieh 
Gabrielson and Lincoln (1949) referred the subspecies on thi] 
Alaska Peninsula proper to alascensis, as distinct from the race; 
on nearby islands. 

Alaska willow ptarmigan were observed at the west end o: 
the Alaska Peninsula in 1925. About the middle of May, th( 
males were strutting and crowing in a lively fashion at Izembel 
Bay. On June 14, very few females were seen. Evidently, thej 
were incubating, because on June 22 I found a nest of nine eggs 
pipped, ready to hatch, and late in July there were broods o] 
young on the marsh at Moffet Bay. 

Concerning the boldness of males at this time, I find the follow- 
ing in my field notes for June 3 : 

The female was sulking among the alder stems on the shore of a ponci 
and I stood on a rise nearby. The male rushed between me and his mate 
growling, puffing out his chest, and elevating his combs. He was a splendic 
bird as he strutted, following his mate as she sneaked along in the brusl 
but keeping out in the open himself, evidently to attract attention awaj 
from the female. I was within 15 feet of him at times. 

On July 3, Donald Stevenson watched a pair of ptarmigan pro- 
testing the approach of a brown bear. The bear had been walk- 
ing across a gentle slope toward the mountains and evidently haci 
disturbed a brood of young birds. Both parents were pretending^ 
to be crippled before the huge intruder. The bear made several 
lunges at the birds, but finally continued on its way. 

Lagopus lagopus mur'ie'i 

Aleut: Alladak (Wetmore) 

This ptarmigan occurs on Kodiak Island, Unga, Nagai, Little^ 
Koniuji, Simeonof, and Popof Islands of the Shumagins, and 
Atka, Unalaska, and Unimak of the Aleutians. 

This willow ptarmigan was described by Gabrielson and 
Lincoln in 1949, as follows: "As compared with L. /. alascensis, 
this race is much redder and darker when skins in comparable 
plumages are compared. L. I. alascensis is buffy; the new race, 
muriei, more reddish and darker, near walnut brown, while 
alexandrae [of Baranof and adjacent islands] is dark brown to 
bister." 

As to distribution, they commented: "Somewhat to our sur- 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 123 

rise, all birds from Morzhovoi Bay, only a few miles across from 
'alse Pass, certainly belonged to alascensis, while those from 
Jnimak Island just as definitely belonged to the island group 
muriei) ." 

Beals and Long^vorth (field report, 1941) reported numerous 
tarmigans on Unimak from February 26 to April 10, in flocks of 
5 to 300 birds. They noted, on March 6, at False Pass as fol- 
)ws: "Large flocks of 300 or more birds each flew about the 
Iders back of the cannery. We saw several flocks of 75 to 100 
irds in Sourdough Flats and vicinity the same day." On March 
4, they reported "ptarmigan by the hundreds" in the valley back 
f False Pass. On March 31, at Ikatan Valley, they saw 3 flocks 
f 100 birds each, and saw numerous groups of 10 to 15 birds. 
)n April 2, at Sourdough Flats, they reported, "Flock after 
lock of 100 to 150 or more each all through this area. The flocks 
ept moving ahead of us until several thousand ptarmigans were 
:athered in one large brood across the valley floor. It looked 
nd sounded like a gigantic chicken ranch." On April 10, at 
:'alse Pass, a flock of "several hundred" were noted ; the males 
/ere "reddish brown about the head and shoulders." 

During field work on Unimak Island in May, 1925, I found 
hese ptarmigan common in the lowlands and on the middle 
lopes of the mountains. On April 30, I saw three males that had 
.cquired much of the brown plumage, but on May 4 the females 
hat I observed were still mostly white, though speckled with 
>rown. On May 5, I saw one male in almost complete summer 
)lumage. 

An interesting incident occurred on May 19, 1925, at St. 
Catherine Cove. I was about ready to leave my cabin, when the 
lattering call of a male willow ptarmigan sounded close by. The 
all was followed by a light patter on the floor of an adjoining 
hed. Before going into the shed to investigate, I glanced out 
he window and saw a peregrine falcon. In the shed, I found a 
ock ptarmigan that ran out through the open door, only to return 
ilmost at once. But my presence proved too much for him, and 
le finally bolted out through the open door and, with lusty crow- 
ng, took flight and disappeared over a rise. By this time, the 
"alcon was some distance away. 

.agopus mufus: Rock Ptarmigan 

The rock ptarmigan occurs on the Alaska Peninsula, on the 
■astern Aleutian Islands as far west as Yunaska, on the middle 
md western Aleutians from Atka Island as far west as Kiska 



124 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

and, on the extreme western end of the chain, Attu Island. It i i 
represented by eight subspecies: neUoni, yunaskensis, atkhensi 
chamberlaini, sanfordi, gabrielsoni, iotonsendi, and evermann 

As reported elsewhere (Murie 1944, p. 122), the rock ptai 
migan of the Aleutian Islands-Alaska Peninsula district fall int 
two groups : A dark, more or less blackish group (in summe 
plumage) , and a yellowish group (also in summer plumage) . Th 
dark group, comprising nelsoni, yunaskensis, and evermanni, 0( 
cupies the Alaska Peninsula and the eastern Aleutians as fa 
west as Yunaska, with the representative subspecies evermani 
on the extreme western end of the chain, Attu Island. The yello'w 
ish group, comprising the other five forms, occupies the middl 
and western Aleutians from Atka Island as far west as Kiska. 

The five so-called yellowish ptarmigan races in the middl 
Aleutians are actually very similar in appearance and are hard t: 
distinguish without a series for comparison purposes. The fa( 
that so many forms can be separated within such a comparativel 
limited area can be explained only by the partial isolation al 
forded by island habitat, though a given race is not necessaril 
confined to a single island, but may occupy a group of island; 

Close knit as these five "yellowish" races are, it is still possibl 
to separate them. The three eastern forms, atkhensis, chamber 
laini, and sanfordi (the most difficult to distinguish one from an 
other), form a group characterized by pale coloration, and, moi; 
particularly, by finer barring in the plumage. The two wester 
forms, gabrielsoni and townsendi, have much heavier barring. 

As Bent has pointed out, middle-Aleutian ptaraiigan occup 
lowland areas, comparatively speaking, in contrast with the higl 
mountain habitat of rock ptarmigan farther east. This does m 
mean that the middle-Aleutian races avoid highlands; they 0(; 
cur on relatively high ground on Atka and Kiska, and elsewhere 
But the terrain of these islands is not particularly rugged, nci 
of the high-mountain type. Amchitka, for instance, is a low ii 
land. It is true, as Bent has said, that these rock ptarmigan li\ 
to a large extent in grassy areas, but it does not necessarily fo 
low that they have responded directly to environment by takin 
on colors that blend with the color of dead grass. This is 
possibility, but at present we do not have sufficient facts for 
conclusive decision. 

Lagopus mutus nelsoni 

Nelson's rock ptarmigan is a dark race that occurs throughoii 
the Alaska Peninsula area, including Kodiak and Afognak in 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 125 

ands, the Shumagins, and other neighboring island groups, and 
v'est in the Aleutians as far as the Islands of the Four Mountains. 

We had suspected that birds from the Islands of the Four 
/[ountains might have developed new characteristics due to iso- 
ation. Ground color of the plumage of birds taken on these is- 
ands in early summei differs in tone from that of evermanni or 
idgwayi (of the Commander Islands) — although there is a slight 
livaceous cast, the predominating shade is yellow-brown. The 
Tound color also differs from that of specimens of nelsoni from 
Jnalaska and farther east, tending to gray rather than to red 
n overall effect. Such differences, if real, are too insignificant 
warrant naming a new form. They can best be referred to a 
light deviation from the usual in nelsoni. There is a similar situa- 
ion at Kodiak and neighboring localities, where nelsoni shows an 
pproach to dixoni (of southeastern Alaska), because the ground 
olor of nelsoni has a slightly grayish cast. 

The females of each Aleutian form cannot readily be distin- 
;uished, but there is a discernible difference between females of 
i.elsoni (which represent the darker forms) and the females 
if the middle Aleutians (which represent the pale forms). On 
emale nelsoni, barring is black, broad, and in contrast ; whereas, 
•n the pale forms, the barring is less sharp, and the bars tend to 
)e discontinuous with a softer effect. 

Spring-plumage changes in Nelson's rock ptarmigan come much 
ater than the changes in willow ptarmigan. A 1-pound speci- 
nen, collected on Ushagat of the Barren Islands group on May 
1, 1936, was still mostly white. On May 7, 1925, I found, on 
Jnimak Island, that the plumage was still nearly all white. On 
day 14, 1925, males were seen with well-speckled plumage, but 
t was a long time before these birds attained full summer dress. 
)n June 6, in the Izembek Bay region on Alaska Peninsula, 
nales still had considerable white in the plumage, but the females 
lad changed completely into summer plumage. A female taken 
m Dolgoi Island, May 24, 1937, contained well-developed eggs. 

Nelson's rock ptarmigans are largely inhabitants of the high 
nountains, though they are not confined to the steeper parts. 
?hey are often found on gentler middle slopes — in this respect, 
hey resemble the rock ptarmigan of interior Alaska. 

agopus mutus yunaskensis 

The Yunaska rock ptarmigan was described by Gabrielson and 
Jncoln (1951) on the basis of a specimen collected by Gabriel- 
on on Yunaska. As might be suspected, although it is grayer 



126 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

than nelsoni, it seems to be more nearly .allied to this darkei 
group to the east. Although its range is thought to be confinec 
to Yunaska, this is not certain, 

Lagopus mufus afkhensis 

Atka: A-gha-de-gach, or A-gha-de-gah 
Agdikax (Jochelson) 

Although native names seem to apply to rock ptarmigan ir 
general, it seems appropriate to apply the Atka dialect name tc 
this form found on Atka Island. Turner's rock ptarmigan maj 
possibly occur on Amlia Island (separated from Atka by onlj 
a narrow pass) , though no specimens were collected on Amlia 
Whether this form occurs eastward as far as Seguam is unknown 
but it is logical to assume that it does. It may be expected thai 
atkhensis also occurs westward to the next group of islands, anc 
that it intergrades with the neighboring form, chamherlaini, oJ 
Adak. 

One is struck by the grayish color of atkhensis, even in flight 
as compared with nelsoni. This color is generally more rufesceni 
than chamherlaini, though both birds have the characteristic 
variegation of gray and rufescent patches. In fact, atkhensis anc 
chamherlaini are hard to distinguish, though, when a good series 
of each is laid out for comparison, the difference can be seen. 

On April 4, 1924, Laing obtained a series of 15 birds on Atka — 
a few of these birds were still in full white plumage. The birds 
were just beginning to molt into summer dress, a change that 
appears to be slightly earlier in atkhensis than in nelsoni on 
Unimak Island. 

Rock ptarmigan have always been abundant on Atka, as manj 
visiting collectors have testified. They are able to maintain theii 
numbers in spite of the blue foxes. It was on Great Sitkin — £ 
neighboring island — that I obtained the only direct evidence oi 
cyclic behaviour among the Aleutian ptarmigan. John Taylor, whc 
had a lease on Great Sitkin to raise blue foxes, said that he hac 
placed 14 foxes on the island in 1934. At that time ptarmigar 
were numerous — "Thousands of them" was the way he expressec 
it. When Taylor returned to Great Sitkin the following year 
ptarmigan were scarce. He did not think that the blue foxes or 
this large island could have been responsible for such a swifl 
and marked decrease. There had been red foxes on the islanc 
before the planting of the blues. This sudden decrease in the 
ptarmigan population appears to have been a case of the char- 
acteristic "die-off" of ptarmigan. Similar fluctuations were no1 
reported for Atka Island. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 127 
Lagopus mufu.s chamberlain} 

Chamberlain's rock ptarmigan is known only from Adak Is- 
land, but it undoubtedly occurs on neighboring islands. It is 
somewhat grayer than atkhensis, and is darker on the top of the 
head and upper neck — an effect, not particularly striking, that 
results from a greater number of black markings. No doubt 
chamherlaini and atkhensis intergrade on some of the intermedi- 
ate islands. 

Laing (1925) found rock ptarmigan on Adak "even more nu- 
merous" than on Atka. He collected five males on Adak, April 13, 
1924, that had started to molt into summer plumage. The date, 
like that for the molt into summer plumage by atkhensis, is un- 
usually early for ptarmigan. 

Lagopus mufus sanfordi 

Sanford's rock ptarmigan is abundant on Kanaga, Tanaga, and 
neighboring islands. Bent (1932), speaking of Tanaga, says: 
"The ptarmigan were tamer and more abundant here than on 
any of the other islands that we visited ; we shot more than 40 
in one afternoon." The two specimens we obtained on Kanaga Is- 
land are referable to sanfordi, showing that this race occupies 
Tanaga and Kanaga, at least. 

As Bent said, Sanford's rock ptarmigan is paler than chamher- 
laini, and is somewhat more ochraceous than either chamherlaini 
or atkhensis. Bent (1932) says: 

Although I described and named this race myself (1912), in honor of my 
friend Dr. Leonard C. Sanford, who cooperated with me in organizing our 
expedition to the Aleutian Islands, I must confess that it is only slightly 
differentiated from the Adak ptarmigan. We all noticed a diiference when 
our birds were collected, and when we laid our series of about 40 specimens 
of sanfordi beside nearly as many of chamberlaini, it was easy to see that 
the Tanaga birds were appreciably paler than the Adak birds. The Tanaga 
birds ai'e therefore the lightest in color of any of the Aleutian ptarmigan, 
and have the finest vermiculations. 

Lagopus mufus gabr'ielsoni 

Gabrielson's rock ptarmigan occurs on Amchitka Island, the 
type locality, as well as on Little Sitkin Island and Rat Island. 
We have no specimens from Semisopochnoi Island, where this 
form may occur also. In 1938, Scheffer obtained, on Amchitka, 
a specimen whose crop was filled with berries of Empetrum 
nigrum, no doubt a favorite food of all these rock ptarmigan. 

In summer plumage, the male gahrielsoni differs from sanfordi 



128 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

in that the ground color is more uniformly dark, and the bars 
are broader and extend farther down the flanks and back; it 
differs from townsendi in that the coarse barring is not restricted 
to the anterior part of the body, and the ground color is paler 
and less ochraceous. 

Lagopus mufus fownsendi 

Differences between toivnsendi and gabrielsoni have just beer 
given. Townsend's rock ptaraiigan is found on Kiska and on 
Little Kiska Islands. We have no specimens to prove that it oc- 
cupies Chugul Island. It is possible, but unlikely, that toivyisendi 
is found on Buldir Island, far to the west ; in any event, we found 
no ptarmigan there on several visits. 

Lagopus mufus evermanni 

Attu: A-ti-ka-tooJi-ach 

Russian, Commander Islands: Kuroptka or Kuro-paschka (Stejneger) 

Russian, Yana River region: Mala Kuropatka (Pleske) 

(The Attu and Russian names undoubtedly refer to all rock ptarmigan.) 

Evermann's rock ptarmigan occupies Attu Island. Apparently, 
ptarmigan have always been scarce on Attu, even before the in- 
troduction of blue foxes. According to Turner (1886) , the natives 
reported ptarmigan on Agattu Island, but we did not obtain 
specimens there on our visit in 1936. 

Comparison of evermanni with 7%dgwayi (of the Commander 
Islands) shows that evermanni, darkest of the rock ptarmigan 
series, is closely related to the Commander Islands form. In 
ridgwayi, the ground color shades from dark buckthorn to hazel 
This ground color varies with different specimens and on differ- 
ent parts of the body; it is heavily overlaid with a close pattern ol 
fine black vermiculation and is spotted with blackish feathers, 
In the Attu evermanni, the ground color suggests buckthorn 
brown, as in ridgivayi; but it is duskier and less rufescent, and 
the black vermiculations are more closely woven and the black: 
feathers are more prevalent. The plumage characteristics in, 
evermanni give the effect of a darker bird than ridgwayi. Com- 
paring evermanni and ridgwayi with nelsoni from the Islands of 
the Four Mountains in the eastern part of the Aleutian chain, 
we find that there is a general resemblance among the three, but 
that ridgwayi and evermanni show the closest affinity, while, in 
ground color, nelsoni tends to be more olivaceous with an abun-' 
dance of Dresden brown. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 129 

Family GRUIDAE 

Grus canadensis: Sandhill Crane 
Grus canadensis canadensis 

This sandhill crane nests on practically the entire length of 
Alaska Peninsula, so it is not surprising that Friedmann (1935) 
includes this species in the avifauna of Kodiak Island also, on the 
basis of a reported specimen as well as on bones found in middens. 

On May 26, 1936, a crane was heard calling on the tundra at 
Snag Point, Nushagak River ; cranes were heard several times on 
the flats about Ugashik River, and, on May 27, one was observed 
flying high in the air. On May 29, there was a pair and a group 
of three, on the tide flats. Curiously enough, 1 bird in this group 
of 3 appeared to be much larger than the others, giving the ap- 
pearance of an adult with 2 immature birds. It is believed that 
cranes nest on the flats along Ugashik River. 

A local trapper said that cranes commonly nest upriver from 
Nelson Lagoon, and Gabrielson received reports of cranes in the 
Cold Bay district. 

In 1925, I found cranes on Unimak Island and adjacent parts 
of Alaska Peninsula, and, on May 1, 1925, two cranes were seen 
at Urilia Bay. On May 21 and 22, 1925, at Moffet Cove, on 
Izembek Bay, a pair, or a group of 3, were noted on several 
occasions, and on June 14, 1925, Donald Stevenson saw 1 bird. 
This bird (seen by Stevenson) was very fearless, and we thought 
that its mate probably was on a nest not far away. On July 18, 
on a marsh in another part of Moffet Cove, a pair of cranes tried 
to decoy us away — evidently they had their young nearby. 

Thus, it is clear that the crane nests in suitable areas along the 
Alaska Peninsula, probably also at Urilia Bay, Unimak Island. 
If it nests at Unimak Island, this point probably is the western 
limit of its breeding range. 

On August 24, 1937, I found the decomposed remains of a 
crane (evidently a migrant straggler) on the beach of Bogoslof 
Island. Turner (1886) states that the natives reported killing 
a crane on Attu in October. In Turner's opinion, this bird was a 
storm-driven straggler. 

But Austin H. Clark (1910) gives us a more significant ob- 
servation when he says — 

On the morning of June 8 while ashore on Agattu Island I encountered a 
pair of these birds, but could not succeed in getting anywhere near them. 
On being flushed, they never flew for any great distance, but always alighted 
far out in the open pasture-like areas, out of reach from any rocks or other 
suitable cover. 



130 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

It is not impossible that cranes nested occasionally on flat, 
lake-strewn Agattu Island. 

Hartert (1920) records two adult specimens, a male and a fe- 
male, from the Commander Islands, and states that they probably 
visit that area from time to time. 



Family RALLIDAE 

Fulica americana: American Coot 
Fulica americana americana 

The only record of this bird was obtained by Gabrielson. On 
December 10, 1943, John Gardner, of False Pass, observed this 
bird in a small stream near his home. It remained for several 
days and appeared to be sick; on December 15, it was killed by a 
dog. The head, wings, and feet were saved for identification. 
Residents of the community had never seen a coot there before. 



Family HAEMATOPODIDAE 
Haemafopus bachmanJ: Black Oysterca+cher 

Attu: He-gich 

Hekh (Turner) 
Atka: Hech 

Hegis (Turner) 
Unalaska: Hekh (Turner) 

Hekli (Clark) 
Russian: Morskoi Ptookh, "Sea Cock" (Turner) 

(The variations in native names are unquestionably due to individual speech mannerisms, 
and represent the same name in all dialects.) 

Speaking of Haematopus osculans, Stejneger (1885) says. 

The Russians of Kamtschatka apply to this bird especially the name 
Ptuschok (pi. "Petuschki") , a chicken, a term used for Simorhynchus 
pygmaeus by the natives of Bering Island, for Leucosticte griseonucha by 
those living on the Prybilof Islands (according to H. W. Elliott), and to 
other birds in different parts of the vast empire where the Russian tongue 
is spoken. 

It would not be surprising, then, to have this name appear, var- 
iously applied, in western Alaska. 

The black oystercatcher breeds commonly from Kodiak Island 
westward for the length of Alaska Peninsula, especially on 
islands. We recorded one at Chignik, but we did not record the 
species on the shore of Alaska Peninsula. On May 11, 1936, one 
was found on Ushagat (Barren Islands), and we observed them 
also on Kodiak Island. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 131 





Figure 26. — Black oystercatcher. 

In 1940, Gabrielson noted 6 of these birds at Whale Island, and 
he saw 15 or 20 in the Semidi Islands. 

On May 16, 1936, we obtained a specimen of the black oyster- 
catcher on Nagai Island, in the Shuinagins. 

Chase Littlejohn (manuscript notes, 1887-88) wrote: "Found 
from Sanakh to Kodiak in limited numbers. Their warning cry 
at the approach of man if heard by the sea otter causes the 
latter to make off at once, for this reason they are much hated 
by otter hunters." Turner also mentions the reactions of sea- 
otter hunters. 

The north side of Alaska Peninsula is not suitable for this bird 
because it lacks a rocky-shore habitat. But the bird nests on Amak 
Island, near the west end of the Peninsula, which is probably 
the eastern limit of its nesting range in Bering Sea. 

In the Aleutian Islands proper, the black oystercatcher occurs 
on nearly every island and is a fairly constant feature of the 
rocky-shore fauna. On Attn Island, however, we saw none, and 
we were assured by the native chief that they do not occur there. 
This probably is true of all the Near Islands. We have no record 
of this bird west of Kiska. Turner points out (1886) that the 
distance between Kiska and the next island, Buldir, may be too 
much of an over-water flight for this bird. Strangely enough, 
in 1885, Turner had reported it as a rare visitor to Attn, occur- 
ring oftener on Semichi and Agattu. 



132 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

In this connection, it is interesting to note Stejneger's remarks 
on Haematopus osculans, of Siberia (1885) : "This bird comes 
only as a rare visitor to the [Commander] islands during the 
migration seasons. This is rather strange, as it inhabits the 
nearest coast of the mainland." 

Perhaps the oystercatchers are merely conservative — lacking 
the exploratory tendency of some other species — and have not yet 
had time to extend their range to the end of the Aleutian chain. 
However, if we believe Turner's report of 1885, rather than his 
report of 1886, the oystercatchers had reached Attu in 1885. 

Apparently, oystercatchers do not exceed a certain popula- 
tion density and are scattered rather thinly along rocky shores. 

Usually, there were only a few pairs on an island (about six), 
although more birds can be found on the larger islands. But 
sometimes, in summer, they gather in loose flocks. On Ogliuga 
Island, August 6, 1936, at least 25 or 30 were seen. On tiny 
Salt Island, off the shore of Atka, on July 8, 1936, a flock of 13 
was noted. 

A nest was found June 28, 1936, on a small islet off Little 
Tanaga Island. The nest was in the grass — merely a shallow 
depression lined with a few bits of barnacle shells — and con- 
tained two eggs. Gabrielson (1941) found a nest on Tanaga 
Island that contained 2 young and 1 pipped egg. 



Family CHARADRIIDAE 

Charadrius dub'ius: Litrle Ringed Plover 
Charadrlus dub'ius curonicus 

The only record of the little ringed plover is the one by Schalow 
(1891, p. 259), for Kodiak Island, which originally was recorded 
as Charadrius alexcindrinus Pallas. Oberholser (1919) concluded 
that this record should be identified under Charadrius dubiu^ 
curonicus, and it was so listed in the 1931 A. 0. U. Check List. 
This record is considered doubtful, and has been dropped from 
the 5th edition of the A. 0. U. Check List. 

Charadrius semipalmafus: Semipalmated Plover 

The semipalmated plover is recorded from Kodiak Island 
(Friedmann 1935) , and we observed two on the beach of Ushagat, 
Barren Islands, May 11, 1936. Howell (1948) found a nest 
with eggs at Kodiak Island, May 31, 1944. It occurs through- 
out the length of Alaska Peninsula. G. D. Hanna collected a 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 133 

specimen, May 23, 1911, at Nushagak. Gianini (1917) found 
them to be common about Stepovak Bay on the south side of 
Alaska Peninsula, where they appeared to be nesting, in May 
and June 1916. Jaques (1930) recorded them on the north side 
near Port Moller. We saw three at False Piss, Unimak Island, 
August 23, 1936. In 1925, I observed the species at False Pass 
and at St. Catherine Cove, May 16 and 17. In the latter part of 
May 1925 they were found again in a valley below Aghileen 
Pinnacles, near Izembek Bay, and at Applegate Cove. At the 
time, it was believed that they were nesting. Wetmore found a 
few of these Kirds at Thin Point, near Cold Bay, in August 1911, 
and he obtained two immature specimens at the east base of 
Frosty Peak on August 6. Beals and Longworth, reporting on 
False Pass, in 1941, noted one on a gravel bar of an old stream 
bed May 9, and remarked that "2 are seen on this gravel bar 
every time we pass. They were not observed after the 25th of 
May." In 1940, Gabrielson also noted three of these birds at 
Morzhovoi Bay. Donald Stevenson obtained a specimen on Uni- 
mak Island, May 25, 1922, and made this notation: "Arrived 
about May 1st. Rather common along glacial stream beds. 
Breeds." 

McGregor (1906) obtained a male and a female in English 
Bay, Unalaska Island, May 27, 1901, and he obtained two imma- 
ture birds on Unimak Island, August 14. 

In view of all these observations, the evidence is rather con- 
clusive that the semipalmated plover nests as far west as Unalaska 
Island. 

PluviaUs dominica: American Golden Plover 

Pluvia'is dominica fulva 

Attu : Svegch 

Smix (Jochelson) (Probably refers to this bird; no dialect given) 

Osgood (1904) says of this species at the base of Alaska Penin- 
sula: 

A few small flocks were seen on the tide marshes and along the mud flats 
about Nushagak September 12 to 26. Several were seen at Igagik and 
others occasionally along the Ugaguk River, as far up as the mouth of 
Becharof Lake. Specimens were taken at Nushagak by McKay in June, 1881. 

Hine (1919) observed these birds at Kashvik Bay in 1919, and 
he collected a specimen on August 24. 

Friedmann (1935) records a number of specimens and obser- 
vations of this bird at Kodiak, where it is no doubt a regular 
migrant. 



134 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Farther westward, records are available all the way to Attu. 
On May 16, 1925, Donald Stevenson saw a bird at False Pass, 
which he thought was the golden plover. Eyerdam (1936) saysl 
"Frequently seen and collected at Unalaska and Unimak Island." 
Dall (1873) reports a specimen of '^Charadrius virginicus, Borck" 
taken June 22, 1872, at Popof Island, in the Shumagins. Turner 
(1886) observed a golden plover at Sanak in July 1878, and on 
May 17, 1879, at Atka Island, he identified the plucked body of a^ 
golden plover. Again, in the early part of October 1880, he saw^ 
two golden plovers on the beach at Massacre Bay, on the south' 
side of Attu Island. ,,^^ 

Gabrielson obtained specimens at Cold Bay and in the Shu- 
magins, in 1943 and 1944 respectively. 

On June 3, 1937, we observed a golden plover circling over thei 
stormy sea between Segula and Semisopochnoi Islands. 

The chief of Attu village declared that he knew of the golden| 
plover; he recognized a colored picture of it, gave us the Aleut' 
name, and referred to it as the "gold snipe." He insisted that 
this plover nests commonly on Attu Island, and that it remains 
until October. 

The reported nesting on Attu requires verification, but it is 
safe to say that the golden plover may appear anywhere — as ai 
migrant or nonbreeder, at least, from Kodiak Island to Attui 
Island, though it does not appear to have been observed in great 
numbers. 

Stejneger (1885) remarked that "The individuals of fulvusi 
breeding in America migrate in winter along the Asiatic coasts, i 
thus giving evidence of the way in which the species once im- 
migrated into Alaska." The records here given, however, are 
proof of a migration along the Alaskan coast. Conover (1945) 
has shown that both the American and Asiatic forms occur in 
Alaska, and that fulva predominates on the Bering Sea coast ot 
Alaska. Thus, both forms could occur in the Aleutian district. 

Squafarola squafarola: Black-bellied Plover 

Friedmann (1935) says of the black-bellied plover, "The only.! 
Kodiak record I have found is a specimen referred to by Salvim 
and Godman in their description of this species in their great! 
work on Central American birds." 

Osgood (1904) says "Two black-bellied plover were collectedi 
by McKay at Nushagak Aug. 8 to 14, 1881." 

Turner (1886) says "They occasionally occur in the spring 
migrations on the Aleutian Islands, the more abundantly on the 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 135 

western islands than those in the vicinity of Unalaska. I saw 
several on Sanakh Island in the spring of 1878, and also in late 
August of 1879." 

Stejneger (1885) says that they occur on the Commander Is- 
lands in fall migration only. 

We saw none of these birds on any of our expeditions. 

Apbriia virgafa: Surfbird 

Turner (1886) says of the surfbird "At Sannakh Island in 
1878, and at Kodiak in 1881, I saw several individuals of this 
species, but under circumstances which rendered it an impossi- 
bility to collect them." 

Friedmann (1934) records a specimen to the northward at 
Goodnews Bay on the Bering Sea coasts taken August 12, 1933. 

The Attn chief, who is well versed in his native avifauna, did 
not recognize a picture of this bird. -'^' 

Arenar'ia inferpres: Ruddy Turnstone 
Arenaria inferpres inferpres 

Commander Islands (native) : Kidmalgikh (Stejneger) 

Russian, Commander Islands: Kasnonogoj Kidik, i.e., red-legged sand 

snipe (Stejneger) 

A series of specimens was available for study : 2 from Nusha- 
gak, 1 from King's Cove, 2 from Unimak Island, 1 from Unalaska, 
2 from Umnak Island, 1 from Ogliuga Island, and 1 from "Aleu- 
tians." In addition to these (which we examined), McKay ob- 
tained a turnstone at Nushagak, August 12, 1881. 

We carefully compared the above-mentioned specimens with 
series of A. i. morineUa from eastern localities and with speci- 
mens of A. i. interpres. The relationship between these two forms 
did not seem to justify the insertion of an intermediate sub- 
species, such as A. i. oaJmensis, as has been proposed. More- 
over, the present series from the Aleutian district agrees with 
the characters of A. i. interpres. One specimen. No. 118845 of the 
U. S. National Museum, taken by William Palmer, at Unalaska, 
May 19, 1890, is much like morineUa and perhaps could pass for 
that race, especially because of the coloration of the head. But, 
when the extensive black on upper parts and the restricted brown 
areas and paleness on the wings is considered, it seems best to 
refer it to interpres. 

On May 22, 1936, we observed a small flock of turnstones, 
believed to be of this species, at Nelson Lagoon. On August 20, 
2 or 3 ruddy turnstones were seen at Port Moller, where they 
wei'e feeding on the beach with Aleutian sandpipers. 



136 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

In 1925, I took three specimens of ruddy, turnstones at St. 
Catherine Cove, Unimak Island — others were seen, including one 
at False Pass, 

Wetmore observed them at King's Cove in August 1911; Mc- 
Gregor (1906) noted them on Unimak Island, August 14, 1901, 
and he obtained two specimens on Amaknak Island, August 17. 

Laing (1925) found 10 of these birds at Unalaska on August 8, 
1924, and collected 3. Cahn (1947) saw one ruddy turnstone, in 
company with other sandpipers, at Summer Bay, Unalaska Is- 
land, July 18, 1944. 

Gabrielson found small groups of ruddy turnstones at Amchitka 
in September 1944. 

Littlejohn (1887-88) wrote "Plentiful in the fall at Sanakh 
where some remain during winter. They are very fat and tooth- 
some. Also numerous at Morzhovoi Bay." 

We found small flocks at Ogliuga and Skagul Islands, on July 
23 and August 5, 1936, and the following year they were seen 
again at the same place on July 27 and on August 4. One speci- 
men was taken. On June 5, 1937, we saw 1 on Kiska Island; on 
July 31, w-e saw 1 on West Unalga; and on August 2, we saw 12 
on Ilak Island. 

Turner (1886) says "The turnstone is of more frequent occur- 
rence in the region about the shores of Bristol Bay, the Alaska 
Peninsula, and the Aleutian Islands ; perhaps more common on the 
western islands of that chain than to the eastward. I saw indi- 
viduals at Attn, Amchitka, Atkha, and in the vicinity of Belkov- 
sky village." And he adds : "They do not arrive on the Aleutian 
Islands until the middle of May, and none were observed any- 
where after the 1st, of October." 

Nesting throughout this region was not established. Stejneger 
(1885) states, concerning the Commander Islands, that they are 
at least migrants, and that possibly some of them breed. 

Arenaria melanocephala: Black Turnstone 

Bretherton (1896) found the black turnstone breeding on 
Kodiak Island, and Friedmann (1935) lists a number of other 
records for that island. Osgood (1904) collected one bl5.ck turn- 
stone at Lake Clark, base of Alaska Peninsula, July 23, 1902, 
and observed others at Nushagak. He also mentions specimens 
taken in June, July, and August, at and near Nushagak and 
Ugashik, by McKay and Johnson. 

During August, Hine (1919) found these birds at Kashvik Bay 
in increasing numbers ; by August 25, they were one of the most 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 137 

abundant shorebirds, being observed in flocks of at least 100. 
Specimens were taken. Evidently, this is in the migration route. 

July 23, 1940, Gabrielson found this turnstone to be common 
along Kvichak River, above Naknek. 

We frequently saw the black turnstone on the tide flats at 
Ugashik River, May 27 to 29, 1936. One day, I noted 8 pairs, 
and found a deserted nest containing 3 eggs. Evidently, the birds 
were on their nesting grounds, which were confined to the tide 
flats rather than to the somewhat higher mossy areas farther 
back. 

Littlejohn (notes) wrote "Saw one flock in the spring at 
Sanakh. Tried hard to obtain a specimen but failed. They were 
very wild." 

Turner (1886) saw one of these turnstones at Belkovsky, south 
side of Alaska Peninsula, in the early part of August 1881. He 
says that they were reported to be plentiful on Unga and Sanak 
Islands, where natives claimed this bird interfered with hunting 
of marine mammals by making its characteristic outcries. The 
natives had stated that the black turnstone is not found on 
"Unalashka and other islands west of the mainland." 

Family SCOLOPACIDAE 

Capella gallinago: Common Snipe 
Capella gallinago delicafa 

Attu: Goo-lech' -arch (?) 

The Attu chief insisted that he recognized a picture of a Wil- 
son's snipe and gave us the native name, adding that the bird 
nests on Attu as well as on other islands. Since this is at vari- 
ance with all other information, one must seriously question it. 
There is the possibility that the chief was referring to an allied 
form from Siberia, which resembles the Wilson's snipe, and 
which may occur sometimes in the Near Islands. 

On May 12, 1936, a Wilson's snipe Mas performing high in the 
air over Kodiak Island, evidently on its nesting ground. Again, 
on May 25 and 26, several of these snipe were performing at 
Snag Point, Nushagak River. Osgood observed this species at 
various parts of the base of Alaska Peninsula, and, he records a 
specimen taken by McKay, April 25, 1882 (1904). Hanna also 
obtained a specimen at Nushagak, May 16, 1911. 

Cahalane (1944) observed the common snipe in several places 
within the Katmai National Monument in 1940, and on July 17, 
1940, Gabrielson saw two snipe at Dillingham. 



138 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Jaques (1930) found these birds in the Port Moller region in 
June, and Bent (1927) includes the Shumagin Islands in the 
breeding range. 

In 1925, I noted one common snipe at Urilia Bay, Unimak Is- 
land, on May 3, and another was heard several times at Moffet 
Cove, Izembek Bay, on July 22. Undoubtedly these were nesting 
birds, so there is good evidence that the nesting range reaches 
westward at least as far as the Shumagins and Unimak Island. 

Numenius phaeopus: Whimbrel 
Numenius phaeopus hudson'icus 

The occurrence of curlews or whimbrels was rather sketchy 
and none was found breeding. Osgood (1904) reported three 
specimens collected by McKay at Nushagak in August 1881. 
Cahalane observed a flock of seven flying in an easterly direction 
about 5 miles above Naknek village, on Naknek River, September 
2, 1940. 

On July 23, 1925, I observed a flock of six curlews flying over 
the marsh at Moffet Cove, Izembek Bay. On June 5, 1937, 2 
curlews were seen at Kiska Island in company with 16 Pacific 
godwits and a ruddy turnstone. Again, on July 30, a curlew was 
seen on Kavalga Island. These were thought to be phaeopus, but 
specimens were not taken, and it is possible that some, or all, 
were tahitiensis. Stejneger (1885) reports the eastern whimbrel 
as a migrant on Bering Island. 

Numenius fah'if'iensis: Bris+le-thighed Curlew 

On July 23, 1940, Gabrielson recorded in his field notes, for the 
Kvichak River, above Naknek, "Flock of 20 flew over. Dufresne 
has seen as many as 200 in the past 3 days around Naknek." 

This is the only record we have, but, in 1924, we had observed 
migrating flocks of immature birds at Hooper Bay, and it is 
logical that bristle-thighed curlews should pass over the basal 
part of Alaska Peninsula in migration. 

Actifis macularia: Spotted Sandpiper 

Friedmann (1935) lists the spotted sandpiper in the Kodiak 
avifauna on the basis of four specimens collected by Wosnes- 
sensky during 1842-43. Speaking of the base of Alaska Penin- 
sula, Osgood (1904) says — 

When we arrived at Lakes Iliamna and Clark, in the latter part of July, 
the majority of the spotted sandpipers, which doubtless breed in the region, 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 139 

had migrated, and only scattering stragglers remained. One small flock 
3f 8 or 10 hornotines was seen nervously flitting from point to point along 
the gravelly beaches of Lake Clark July 25. Some days later a few belated 
individuals were found along the lower part of the Chulitna River. Prac- 
ticably all were gone before August 10. 

We found none of these birds farther west. 

Tr'inga glareola: Wood Sandpiper 

There is a single record of this bird for Sanak Island — a speci- 
men taken by Chase Littlejohn on May 27, 1894 (Littlejohn, 
1904). The bird was found among some Aleutian sandpipers, 
and another, thought to be of this same species, was seen. 

Stejneger (1885) reported it rather common and breeding in 
the Commander Islands. 

Heferoscelus incanum: Wandering Tattler 

Russian, Commander Islands: Tschornij Kidik (Stejneger) 

A wandering tattler was seen on Kodiak Island, May 12, 1936, 
and 6 or 7 were seen on the beach at Karluk, Kodiak Island, 
September 1. Hine (1919) collected two specimens at Katmai 
Bay in 1919. Gabrielson noted a wandering tattler near Iliamna 
Lake on July 24 and 26, 1940, and he noted the species at Cold 
Bay, King Cove (with specimens), and Kodiak, as well as at 
Dutch Harbor and Amchitka. We obtained a specimen on Nagai 
Island, Shumagin group. May 16, 1936, and we saw one at False 
Pass, August 23. Scheffer saw one on Sanak Island, August 28, 
1937. Nelson (1887) had seen one on Sanak Island, May 15, 
1877. 

The wandering tattler has frequently been reported in the east- 
ern Aleutians. Bishop (1900) obtained 2 at Unalaska, October 5, 
1899; McGregor (1906) recorded 2 specimens from English 
Bay, Unalaska Island, June 2, 1901; Laing (1925) saw 4 at 
Unalaska, where Turner also recorded 1; and Swarth (1934) 
reports 6 at Akutan, which includes 3 specimens taken. 

On July 16, 1911, Wetmore obtained a breeding female at King 
Cove, Alaska Peninsula, and said he judged that she had young 
in the vicinity. He found the birds to be common there in August. 

In 1925, I observed wandering tattlers at False Pass and 
Izembek Bay. On May 21, there was a pair on the beach, calling 
and perching on various boulders. On July 19 and 23, there was 
a pair and a single bird on a small gravelly stream flowing out of 
the marsh at Moffet Cove. August 8 and 9, there were several 
on the gravelly stream at False Pass, and, the next day, five 



140 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

were collected on the rocky beach at Ikatan Peninsula. Thes(i 
last-mentioned were extremely fat. Local residents said thai 
these birds occur on streams in the vicinity of Becharof Lake. 

Undoubtedly, the wandering tattler nests along the gravel 
bordered streams in this region. It was on such a habitat tha 
Adolph Murie and I found a nest in Mount McKinley Nationaj 
Park in 1923, and, considering the available evidence, it is monj 
than likely that the wandering tattler nests along the Alaska | 
Peninsula, on Kodiak Island, and probably on other suitable j 
adjacent islands. It is possible that it nests on many of the I 
Aleutian Islands farther west also. Turner (1886) said: "Amongj 
the Aleutian Islands it was observed once on Unalaska, severaj 
on Atkha, and twice on Attu." Clark (1910) reported it a1 
Unalaska, Agattu, and Attu Islands, but it was not common 
Scheffer saw one at Atka, June 1, 1937. We also noted one or 
Kiska Island, June 5, 1937, and obtained a specimen on Herberl 
Island, August 22. 

Stejneger (1885) reported this bird to be common in the Com-; 
mander Islands, and he suspected that it nested there. While 
visiting those islands, he also obtained a specimen of Meter osceluf 
brevipes. It is possible that some of our sight records in the 
Aleutians represent the latter form. It can be expected in the 
Aleutians, for it has been found on the Pribilofs. 

Tofanus melanoleucus: Greater Yellowlegs 

Osgood (1904) described a pair that evidently was nesting all 
a small pond on the portage trail between Lake Clark and Lak€( 
Iliamna. He found the species again at Swan Lake and Mulchatna^ 
River, and he mentions two specimens taken by McKay at 
Nushagak, August 14 to 28, 1881. Hine reported that it com- 
monly nested along the shore of Katmai Bay (1919). We ob- 
served two of these birds at Anchorage in 1936, and we were 
informed by local people that the species nests there. Bretherton 
said that it occurs on Kodiak Island and that it probably breeds 
there. Howell reports seemg two birds at Kodiak on May 9, 1944. 
In 1940, Gabrielson found this bird to be common on Alaska 
Peninsula; he obtained a specimen, and saw others, at Cold Bay 
in September 1942. 

We observed three greater yellowlegs on the beach at Port 
Moller, August 20, 1936. On July 7, 1925, I observed one of these 
birds circling about on Amak Island, and from July 18 to 24 
they were common on the marsh at Moffet Cove, Izembek Bay. 

Evidently, this bird nests at the base of Alaska Peninsula, 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 141 

and it is possible that it nests as far west as the end; however, 
this is not certain, 

Totanus flavipes: Less3r Yellowlegs 

We did not encounter this species on our expeditions to the 
Aleutian Islands, and records are few. Friedmann (1935) men- 
tions a specimen (not available) collected by Bischoff on Kodiak 
Island; its occurrence at Karluk River was reported by Bean 
in 1889. Cahalane reported (1943) that he saw "considerable 
numbers of these birds September [1940] on mud flats exposed 
by falling tide on the Naknek River below the rapids." 

Calidris canufus: Knot 

This species is mentioned here on the strength of Turner's re- 
mark (1886) : "I have not observed this bird west of Ugasik, 
on the eastern end of Aliaska, where it was quite plentiful in the 
latter part of June 1878." Presumably, it migrates through the 
Aleutian district, but we do not know the subspecies that are 
involved. 

Erolia pfilocnemis: Rock Sandpiper 
Erolia pfilocnemis pfilocnemis 

Using material that is available in the U. S. National Museum, 
ptilocnemis, couesi, quarta, and maritima were carefully com- 
pared. The last-named species appeared to be more stable in 
characters than the forms from Bering Sea. In some instances 
there was a close similarity, shown, for instance, between cer- 
tain specimens of couesi from the Aleutians and specimens of 
maritima; winter plumages are quite similar. There seemed good 
reason to include them all as forms of one species — maritima. 
However, Conover (1944) studied a much greater series — more 
than 500 specimens — and concluded that two basic species exist. 
His conclusion is followed here. 

E. p. ptilocnemis is larger than the other Bering Sea forms, 
and it is paler, both in summer and winter plumages. Compared 
with couesi, there is more tan color in the plumage of the back 
(less of the rusty brown and less of the black admixture). Even 
the primaries and tail are of a lighter color. 

In immature plumage, the feathers of the back are dark and 
narrowly edged with rusty brown in a smooth regular pattern, 
thus being distinguished from the broadly edged feathers of the 
adult at that time of year. In this immature plumage, the differ- 



142 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

ence in color of upper parts between ptilocnemis and couesi is not 
striking. The under parts of ptilocnemis are much paler with a 
pale buffy and gray area across the breast, the throat is nearly 
white and finely spotted, and the streaks on the breast and up- 
per flanks are narrow and pale. In contrast, the under parts of 
couesi in the same plumage are heavily and boldly streaked and 
spotted, thus giving the bird a darker appearance. In the winter 
plumage also, ptilocnemis is markedly paler than couesi. 

In all races, the measurements of wing and exposed culmen 
average greater in the female than in the male. Measurements, 
in millimeters, of 13 males and 19 females of ptilocnemis are as 
follows : 

Males: wing, 118 to 132 (125.6); exposed culmen, 27 to 32 
(29.3) 

Females: wing, 125 to, 136 (129.5) ; exposed culmen, 29.5 to 
37.5 (33.4) , . ,: 

The Pribilof sandpiper nests on St. Matthew and the Pribilof 
Islands, but, as would be expected, it occurs on Alaska Peninsula 
and the Aleutian Islands in migration. Probably, it winters in 
this area to some extent. At any rate, among the specimens ex- 
amined there are at least four from the Bristol Bay region that 
are referable to ptilocneynis. One of these, a female, was taken by 
C. L. McKay, at Point Etolin, April 8, 1883. Three others were 
collected by J. W. Johnson, at Nushagak, April 1 and April 18, 
1885. McGregor (1906) records that this species was collected 
on Unimak Island, August 14, 1901, and on Tigalda Island, Au- 
gust 5, 1901. 

Erolia pfilocnemis couesi 

Attu : Too-loo-goo-yuch 

Atka: Chu-Iich'-tah 

Alaska Peninsula: Tsoo-gooch (Wetmore) 

Russian, Commander Islands: Lajdinij kulik (Stejneger) 

In measurements, couesi is quite comparable to maritima and 
quarta, but all three are definitely smaller than ptilocnemis. The 
Aleutian sandpiper is decidedly darker than the Pribilof sand- 
piper — the markings on the under parts are bolder and heavier; 
the upper parts contain more black and a greater proportion of 
rusty brown. In this respect, couesi approaches quarta. 

Measurements, in millimeters, of 29 males and 24 females of 
couesi are as follows : 

Males: wing, 110 to 123 (117.1); exposed culmen, 25 to 34 
(27) 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 143 

Females: wing, 113.5 to 127 (120.5) ; exposed culmen, 27 to 33 
(30.7) 

The Aleutian sandpiper nests throughout the Aleutian Islands, 
where it is the common shorebird ; it also nests along the Alaska 
Peninsula and adjacent islands — at least as far east as Port 
Moller (Jaques 1930), and undoubtedly all the way to the base 
of the Peninsula. Hme (1919) observed it at Katmai Bay in 
1919. At least two specimens in immature plumage were taken 
by Johnson, at Nushagak, July 11 and 18, 1884, and another was 
taken April 18, 1885 — all these specimens appear to be couesi. 
There may be some question in regard to the breeding status of 
this bird on Kodiak Island ; however, it winters there. 




FiGUBE 27. — Aleutian rock sandpiper. 



The winter range includes all of the Alaska Peninsula-Aleutian 
district. 

The Nesting Period 

Extensive observations on the nesting of the Aleutian sand- 
piper were possible in 1925, when I spent a season on Unimak 
Island and the adjacent part of Alaska Peninsula. On April 29, 
flocks of Aleutian sandpipers (as many as 20 birds) were feed- 
ing along the lagoon at Urilia Bay. The first mated pairs were 
noted on May 3 ; these mated birds had left the shorelines and 
were nesting on the mossy tundra. By May 7, they had become 



144 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

more plentiful, and mating was in full swing. At this time, : 
flocks were still common on the beaches — several flocks were I 
noted at St. Catherine Cove on May 17. A flock of 150 to 200 
birds, feeding on the tide flat, occasionally would rise, maneuver, 
wheel, and turn in the air (in characteristic sandpiper fashion), 
then settle back on the beach. Thereupon, a great babel of 
chattering would arise, as they all dabbled busily in the wet sand 
and mud. 

On May 18, a single bird was collected on the beach. It proved 
to be a female with an egg almost ready for the shell. The next 
day, a flock of 400 was seen. The significance of seeing these 
large flocks at the same time that others were nesting is hard 
to determine. They must have been nonbreeders or late nesters. 

On May 23, these sandpipers were common on the higher 
tundra back of Moffet Cove, Izembek Bay trilling and calling, 
evidently nesting or still making preparations. Some had ob- 
viously selected the nesting place or had eggs. By May 28, egg 
laying was definitely under way. 

A nest containing four eggs, found June 5, was a cavity in the 
ground lined with a few tiny leaves — diameter was 100 mm.; 
depth was 53 mm. 

These nesting habits were verified on later expeditions (in 
1936 and 1937) throughout the Aleutian chain. Some sandpipers 
nested close to tidewater, others nested back in the hills — some- 
times a considerable distance from a body of water. On June 1, 
1937, on Atka Island, I found 2 nests, each containing 4 eggs. 
They were shallow depressions in a mass of low vegetation, 
lined with bits of lichens, straws, and dwarf-willow leaves. 

Another nest, with four eggs, was found June 4, high up on 
Kiska Island. It was a depression in the moss beside a rock ; the 
cavity was 3 by 4 inches wide, and li/o inches deep. 

On June 22, 1936, on Atka Island, I found a dead, newly 
hatched young. On June 22, 1937, Scheff'er found a brood of 4 
recently hatched young on Little Kiska Island. Another brood 
of 4, several days old, was found on Little Sitkin Island on June 
27 ; and, on June 29, a brood of 3 was found on Rat Island. 

I heard the mating song of the Aleutian sandpiper at Izembek 
Bay in 1925. Quoting from my field report, the song suggested — 

the droning trill of toads, varied by a repetition of "per-deerrrr, per-deerrrr" 
. . . very much like the red-backed sandpiper's call, but shorter. Later on, 
vi'hen frightened from their nests, they had a variety of alarm calls. As they 
flew away, they would call "Ka-deer, ka-deer, ka-deer," similar to the notes 
of mating time, but shorter and sharper, and they also uttered a very rapid 
"uh-uh-uh-uh-uh-uh-uh." 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 145 

Wetmore, in his field report for 1911, says: "The males have 
a trilling note, almost a whinny that is hard to describe. Also a 
quick musical whistled turdle turdle, on the Carolina wren order." 

The first signs of flocking were noted early in July. On July 5, 
1937, a group of 3 adults was observed flying along the beach on 
Amchitka Island, and, from July 10 to July 20, 5 or 6 were 
seen in groups several times. On July 24, 1925, two immature 
birds were collected at Izembek Bay, and several flocks were seen. 
On July 29, 1937, flocks of 40 or more were seen on Ogliuga Is- 
land ; after July 29, they generally were seen in flocks. 

Erolia pfilocnemis quarfa 

Russian, Commander Islands: Lajdinij kulik (Stejneger) 

Ernst Hartert (1920) described quarta from the Commander 
Islands and said — 

The purple sandpiper of the Commander Islands differs from E.m.couesi 
from Alaska and the Aleutian Islands as follows: 'In the winter plumage 
the foreneck and jugulum are darker slate-colour and less mixed with white. 
In the full summer plumage the edges to the feathers of the upperside are 
much wider and of a brighter ferruginous, so that the upperside looks quite 
rust-red, with mostly concealed black centers to the feathers. The wings 
measure 121-127, in one female even 130 mm.' 

In the series from the Commander Islands (in the U. S. Na- 
tion Museum), 6 males and 5 females measure, in millimeters, as 
follows : 

Males: wing, 117-129 (121) ; exposed culmen, 25.5-28.5 (27). 

Females: wing, 120-126.5 (122.8); exposed culmen, 27.5-33 
(29.5) 
These measurements easily fall within the size range of couesi. 

While it is true that quarta is essentially a Siberian form, there 
are a number of records for Alaska. A. C. Bent (1927) reported 
these birds, at least one of which was a breeding bird, from 
Attn Island. Two specimens in the U. S. National Museum, Nos. 
131763 and 131764 (probably the ones mentioned by Bent), as- 
suredly are qimrta. Another Attu specimen. No. 201468, is very 
similar to the less brightly colored specimens from the Com- 
mander Islands. There is another specimen, No. 298506, from 
Izembek Bay, Alaska Peninsula, that is very similar to quarta 
and is practically identical with a specimen from St. Lawrence 
Island, No. 165056. Another specimen, No. 230608, from Morzho- 
voi Bay, has the coloration of quarta. Moreover, Bailey (1943) 
records two specimens from Cape Prince of Wales, taken June 6, 
1922, that were identified as quarta. 



146 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Four specimens were collected by F. L. Beals on Unimak Is- 
land in January 1941. One of these is very dark, thus agreeing 
with the description of quarta in winter plumage, and another is 
nearly as dark as quarta. These specimens have not been identi- 
fied definitely. 

It should be pointed out that in the series from Bering Island 
(the type locality), there are several specimens that lack the 
extreme of bright ruf escence which characterizes quarta ; in fact, 
these specimens are very similar to average couesi. One speci- 
men from Bering Island, No. 89037, is as pale as some ptilocnemis. 

In the light of this circumstance, it is diflScult to evaluate the 
Alaskan records. Are these stragglers of quarta, or are they ex- 
tremes in variation within the population of couesil Until more 
Siberian material is obtained, and until a more extensive knowl- 
edge of quarta is at hand, it may be best to accept our records 
as stragglers of the Old World forai. 

Since the above studies were made, Conover (1944) reviewed 
the group and referred the mainland birds north of Alaska 
Peninsula to tschuktschorum. 

Erolia acuminata: Sharp-tailed Sandpiper 

Specimens of this sandpiper have been obtained in various 
parts of Alaska, including St. Lawrence, St. Michael, and Nuni- 
vak Islands, the Pribilofs and the Russian-held Commander Is- 
lands. We saw none of these birds on our expeditions to the 
Aleutian Islands, but Bailey (1925) reported the capture of a 
specimen (a young of that year) by Hendee, on Unalaska Island, 
on September 27; and Bishop (1900) obtained a specimen at 
Unalaska, on October 5, 1899. Undoubtedly, this species occurs 
in the Aleutian district during migration more often than is 
shown by published records. 

Erolia melanofos: Pectoral Sandpiper 

The pectoral sandpiper proved to be exceedingly rare. Osgood 
(1904) says "One was taken by Johnson at Nushagak October 15, 
1884. The species was not seen by our party." Gabrielson saw 
three of these birds up the Kvichak River, July 23, 1940. 

On July 23, 1925, I observed two birds in the grassy marsh 
at Moflfet Cove, Izembek Bay, which were believed to be imma- 
ture pectoral sandpipers, but, unfortunately, specimens were not 
obtained. The Alaska Peninsula should be in the migration route. 

Bishop (1900) obtained a specimen at Unalaska October 5, 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 147 

1899; Turner (1886) reports taking 3 specimens on Attu Is- 
land; and Hartert (1920) reports taking 2 specimens from Ber- 
ing Island. 

Eventually, this bird may be found nesting on some of the fa- 
vorable habitats on the north side of Alaska Peninsula, such as 
those near Ugashik River, but at present the nearest known 
nesting locality, reported by Friedmann, is at Goodnews Bay 
(1933). 

Erolia bairdil: Baird's Sandpiper 

Friedmann (1935) records a number of specimens from Kodiak 
Island ; only one of these specimens is now available for verifica- 
tion. This specimen was taken by Townsend, August 15, 1888. 

According to Nelson (1887), Dall recorded Baird's sandpiper 
from Kodiak and from Amak Island, north of Alaska Peninsula, 
but there are no specimens to support these records. 

More recently, August 7, 1945, Gabrielson obtained a specimen 
at Wide Bay, Alaska Peninsula. Furthermore, he recorded them 
at Togalak Island, August 5, 1941 ; at Unalaska, Adak, Amchitka, 
Shemya, Agattu, and Kodiak in 1943; and at Amchitka, Adak, 
and Kodiak in 1944. These records reveal that this bird is more 
numerous in ^^he Aleutian district than was formerly supposed. 

Erolia minufilla: Least Sandpiper 

Attu : Kre-a-via-ghre — choo ( ?) 

The chief of Attu village said that he recognized a colored pic- 
ture of the least sandpiper, and he gave us the native name for 
it. But because the lack of striking markings makes identifica- 
tion difficult, and because we have no records for the western 
Aleutians, the chief's statement needs verification. 

During our brief stops at Kodiak Island we did not see this 
bird, but Friedmann (1935) records 6 adults and 9 downy young 
from Kodiak in the Thayer collection. We observed least sand- 
pipers at Port Chatham, Kenai Peninsula, May 6, 1936, and we 
observed it again on Ushagat Island, Barren Islands, May 10, 
where two specimens were taken. Several of these birds were 
noted at Chignik on May 14, and, on May 24, 1937, a pair was 
seen on Dolgoi Island. 

Hine (1919) observed a few least sandpipers, and took a speci- 
men, near the mouth of Katmai River, July 23, 1919. 

Dall (1873) reported it to be rather abundant along the beaches 
of Popof Island, in the Shumagins, June 20, 1872, and he obtained 
specimens at that time. 



148 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 




Figure 28. — Least sandpiper. 



On May 25, 1936, six, or more, least sandpipers were found in 
the marshy vegetation at Snag Point, Nushagak River, where 
they were evidently nesting; the following day, a male, with in- 
cubation patches, was collected. The flight song was heard here 
also. 

Jaques (1930) reported that after May 25 this bird was abun- 
dant about Port Moller in the vicinity of tundra pools. In 1911, 
Wetmore observed the species in August at King Cove near 
Thin Point. Late in July, he saw them at Morzhovoi Bay under 
circumstances that suggested they had just finished nesting. He 
also mentions seeing them on August 25 between King Cove and 
Little Koniuji Island, and on August 26 he saw them off Chignik 
Bay. Gabrielson, on June 21, 1940, found 6 or 8 in a high meadow 
at Frosty Peak, and he took specimens there and at Unalaska, 
Alaska Peninsula, and the Shumagins. Gianini (1917) reported 
them to be common and breeding at Stepovak Bay, where he 
found a nest with four eggs. 

In May 1925, I observed these sandpipers about Urilia Bay and 
St. Catherine Cove, Unimak Island, where they were common by 
May 19. At Hazen Point, Izembok Bay, a pair was seen on May 
21, and, on May 29, birds were observed going through their mat- 
ing performance in the valley below Aghileen Pinnacles. Evi- 
dently these birds were nesting in the marshy valley bottom. 
Least sandpipers were found nesting commonly at Hazen Point, 
where a nest was found on June 22. The nest consisted of a slight 
cavity in the matted vegetation, with a few small round leaves 
in the bottom, and it contained four well-incubated eggs. On June 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 149 

10, a small flock, probably nonbreeders, was noted at Hazen 
Point. During July, this sandpiper was common near Frosty 
Peak and the islands near Point Grant, and, on July 24, a number 
Df flocks, probably immature birds, were feeding on the tide flats. 

Chase Littlejohn (notes) wrote that he "Found [it] breed- 
ing from Kodiak to Sanakh, but not in great numbers, a few 
remain during winter." 

McGregor (1906) obtained a specimen on Amaknak Island, 
May 17, 1901, and obtained another on Tigalda, August 5. Eyer- 
dam (1936) obtained a specimen at Unalaska on May 17, 1932, 
and Gabrielson collected one there on July 4, 1946. Swarth 
(1934) records two specimens taken on Akutan Island on May 19 
and 31, by Cyril G. Harrold, who had remarked that "Several 
pairs were observed on the flats on Akutan Island. The male 
has a strange flight song consisting of a repetition of several low 
notes uttered while the bird is alternately gliding and hovering." 

On May 30, 1937, a pair of least sandpipers was seen by our 
party at Nikolski Village, Umnak Island. 

We have no records beyond Umnak, but the data indicate that 
the least sandpiper nests as far west as Akutan — very probably as 
far as Umnak. 







Figure 29. — Least sandpipers. 



150 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Erolia alpina: Dunlin 
Erolia alpina pacifica 

The red-backed dunlin, or sandpiper, occurs in some localitie 
on the Alaska Peninsula. Osgood (1904) observed several flock 
flying up and down the Egegik River on September 29. He men 
tions several specimens taken by McKay at Ugashik in May an( 
July 1881. G. D. Hanna obtained three specimens at Nushagal 
on May 31, 1911, and Hine obtained a specimen near the moutl 
of Katmai River, August 23, 1919. 

The specimens taken by McKay suggest nesting. Certainly 
in 1936 we found good evidence of nesting at Ugashik River 
these sandpipers were common on the tide flats on May 27 an( 
29. They were paired and evidently breeding. One was obvioush 
flushed from a nest, though the nest was not found. 

On April 29, 1925, I saw a red-backed dunlin feeding on th< 
shore of a lagoon at Urilia Bay, Unimak Island, in company witl 
some Aleutian sandpipers. This may have been a migrant be 
cause none were found nesting on the marshes about Izembel 
Bay. The dunlin's westernmost nesting locality on Alaska Penin- 
sula is, so far as we know, the tidal marshes about Ugashil 
River. 

Taber (1946) noted a few red-backed dunlins wintering or 
Adak Island. 

Stejneger (1885) reported this bird as a migrant in the Com- 
mander Islands. 

LJmnodromus grheus: Short-billed Dowitcher 
L'imnodromus griseus caurinus 

A series of specimens from various parts of the Alaska Penin- 
sula is available, and comparison of these birds with those in 
other series from differing localities brings up the question ol 
the subspecific status among the dowitchers. Specimens from 
Point Barrow, St. Michael, Hooper Bay, Fort Yukon, Nushagak, 
Ugashik, and the west end of Alaska Peninsula were examined 
and compared with numerous specimens from eastern localities. 

At the time that these comparisons were made, it appeared that 
the Alaska Peninsula birds should properly be included with 
scolopaceus. Since then, Pitelka (1950) has studied this genus 
intensively with nearly 3,000 specimens. On the basis of this 
study, he concluded that scolopaceus and griseus are distinct 
species, and that griseus includes three forms — griseus, hender- 
soni, and a new subspecies, caurinus. Previously, Aldrich (1948) 
had concluded that intergradation between populations could be 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 151 

lemonstrated among North American dowitchers and, therefore, 
)nly one species was involved. 

The designation of L. g. caurinus as the breeding form for 
;oiithern Alaska would tend to solve some of the classification 
Droblems of Alaska Peninsula specimens. However, it still seems 
difficult to visualize specific status for scolopaceus, as proposed by 
:^itelka. As one example, a female from Nushagak, with the spot- 
ing of the underparts characteristic of the griseus group, was 
nated with a male, that was heavily barred on the underparts, 
ypical of scolopaceus. Should we consider this to be a case of hy- 
)ridization between two ordinarily isolated species, or should it be 
considered a case of intergradation between two races of the same 
species? A parallel situation exists in the case of the fox spar- 
rows at the base of Alaska Peninsula. 

A specimen from Ugashik River, Alaska Peninsula, was com- 
pared with one from La Saline, Athabaska River, which, pre- 
sumably, is the range of the proposed race hendersoni. Both are 
nales — the Canadian specimen was taken May 12, 1920, and the 
Alaskan specimen was taken May 27, 1936, from a mated pair, 
rhese two specimens are almost identical. The longer bill is on 
he Canadian bird, 60.5 mm., while the bill of the Alaskan bird 
s 54.5 mm. The wing of the Canadian bird is smaller than that 
)f the Alaskan bird (144 mm. and 147 mm.). Both birds are 
leep buff, with very little spotting, the round spots occurring on 
;he sides of the breast and on the flanks. The Alaskan bird has 
I little more white on the belly than the Canadian bird, though 
he latter has a pale, noticeably whitish edging on the feathers 
)f the under parts. On the upper parts, the Canadian bird is 
somewhat darker buff than the Alaskan one. If a mixed series 
3f these birds were laid, it would be most difficult to separate 
them. 

Another specimen from Nushagak River is mostly white un- 
ierneath and rather heavily spotted. This is a female; the wing 
measures 145.5 mm., and the bill measures 62 mm. Neither of 
[hese two Alaskan specimens has barring on the side of the 
Dreast. Without knowledge of the locality, one would place these 
;wo, both breeding birds, with the Canadian group ; however, other 
3irds from Nushagak show plumage associated with typical 
scolopacens. Indeed, most significant of all, the female from 
Nushagak, lacking the bars, was mated with a male that was 
heavily barred. Other birds from the Bering Sea coast vary 
greatly in degree of spotting, in amount of barring, in amount 
of white underneath, and in shade of solid buff color. Length of 
bill also varies greatly — even within each sex group. 



152 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

In view of so much variation, obvious in sny series from 
given locality, and because of the extreme overlapping show 
here, it would seem that subspecific variation best expresses th 
nature of the forms. 

Friedmann (1935) reports one record for Kodiak Island, "tw 
specimens collected by Wosnessensky in 1842-1843, now in th 
Zoological Museum of the Academy of Sciences at Leningrad." 

As mentioned above, we found several breeding birds on a lo^ 
marshy area near Snag Point, Nushagak River, in 1936, and 
female, collected there on May 25, contained two eggs almos 
ready for deposition of the shell. Several pairs M^ere seen on th 
tide marshes at Ugashik River, May 27 and 29. 

Jaques (1930) observed several of these birds, and collectei 
one at Port Moller, June 12, 1928. 

At Izembek Bay, near the west end of Alaska Peninsula, Donal( 
Stevenson collected an immature female, July 2, 1925, and 
obtained another immature female there, July 24, 1925. Thes( 
birds possibly could have been migrants, but we believe the: 
were on their nesting grounds. The locality is excellent habita 
for this bird. 

Cecil Williams, a member of our party in 1936, reported seeing 
a long-billed dowitcher on Bogoslof Island, June 5, at the little 
"sulphur lake." On such a barren island, this sighting is a mosl 
surprising occurrence. 

Ereunefes puslllus: Semipalmated Sandpiper 

Eyerdam (1936) reports that this bird was collected at Una- 
laska and Unimak— the only report of this species for the Aleu- 
tion district. I have not had an opportunity to see these speci- 
mens. 

Ereunefes maun: Western Sandpiper 

Friedmann (1935) mentions that specimens were collected on 
Kodiak by Bischoff, August 10 to 15, 1868. These are the only 
positive records based on specimens. This bird should occur 
there in migration more commonly than these meager records 
show. On May 10, 1936, C. S. Williams obtained a specimen on 
Ushagat, one of the Barren Islands, which are not far from 
Kodiak. Osgood (1904) mentions two specimens collected by 
McKay at Nushagak, and Hine (1919) observed them commonly 
in the Katmai-Kashvik Bay area, where he collected specimens. 
Wetmore found these birds to be common near Thin Point, on 
the Alaska Peninsula, August 3 to 13, 1911, and back of King 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 153 

Cove, July 12 to 20, 1911. Eyerdam (1936) reported that he 
collected specimens at Unalaska and Unimak, and Gabrielson 
obtained a specimen at Cold Bay, July 20, 1942. 

Turner (1886) stated that "This sandpiper is abundant in all 
the Aleutians. At Atkah and Amchitka it is extremely abundant." 
Turner's observations must have been made during certain mi- 
gration periods. Certainly, his statement does not fit present- 
day conditions, because, except for Eyerdam's records, no one 
else appears to have seen these birds in the Aleutians. 

Limosa fedoa: Marbled Godwit 

Osgood (1904) says of this species that "Two immature speci- 
mens of the marbled godwit were taken by McKay at Ugashik 
July 16-18, 1881." 

This is 1 of the 3 unusual records of this species for Alaska. 

Limosa lapponica: Bar-tailed Godwit 
Limosa lapponica baueri 

Attu: Mi-ii-keegh 
Atka: Chu-ee-gech 

Dall obtained a specimen on an islet in Akutan Pass, June 2, 
1872, and he noted it at Unalaska, June 9. He stated that it 
breeds there. Nelson (1887) said "On May 26, 1877, while I was 
at Unalaska, a native brought in half a dozen of these birds, and 
on June 3 I obtained three others from the sandy beach of a 
small inner bay." He said that they appeared to be migrating. 

Cahn observed one of these birds near Unalaska Island, on the 
beach of Hog Island, May 21, 1946. 

Turner (1886) said that— 

This godwit is found on the Aleutian Islands in the latter part of May as it 
is on its way to the northward. On Atkah Island I obtained three specimens. 
They were on the sandy beach of the west side of Nazan Bay. They re- 
main but a few days, and are probably stragglers from the m^in body of 
their kind. 

At Amchitka I saw four of this species on May 24, 1881. They were in 
Constantine Harbor of this Island. 

I do not think they breed on any of the Aleutian Islands. 

Joseph Grinnell (1910) has also recorded two specimens taken 
at Unalaska by C. L. Hall on May 29 and June 4, 1894. 

Donald Stevenson obtained a male bird on Unimak Island, 
June 3, 1922, and noted that "A few observed, this one only 
taken. Was very thin and weak. Sex organs swollen." 



154 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Chase Littlejohn (manuscript notes for 1887-88) noted that- 
Many of these seen in the spring going north at Morzhovoi Bay but thej | 
do not stop, they take almost the same route as the black brant but do not! 
bother about flying around the sand bar. As they are not seen in the fall! 
they must take some other route. 

A. C. Bent (1927) said— 

On its spring migration the Pacific godwit passes through the Aleutian 
Islands and the Pribilof Islands on its way to its breeding grounds in 
northwestern Alaska. I saw two birds on Atka Island on June 13, 1911, 
probably belated migrants; it has been said to breed near Unalaska, but 
this seems hardly likely. 

On our own expeditions, we met with this bird only once. On 
June 5, 1937, we found a flock of 16 Pacific godwits, 2 Hudsonian 
curlews, and a European turnstone at the south end of Kiskai 
Island. Two specimens of the godwit were taken. 

We have no records for the Alaska Peninsula, and we have no 
proof of nesting in the Aleutians. According to Stejneger (1885), 
this bird is a regular migrant in the Commander Islands. 

Limosa haemasfica: Hudsonian Godwit 

Osgood (1901) wrote "Nine specimens were taken by Bischoff 
at Fort Kenai. At least two of these are still in the National 
Museum — one an adult in breeding plumage, the other in fall 
plumage." 

A. C. Bent (1927) wrote that "It has been reported from 
Alaska (Kenai, Nulato, Ugashik, mouth of the Yukon River, and 
Point Barrow) ." 

These observations show that this godwit rarely appeared near 
the base of Alaska Peninsula. 

Crocefh'ia alba: Sanderling 

Chase Littlejohn (manuscript notes for 1887-88) says "Only 
three seen during my stay, and these were seen during very cold 
weather. Twice alone and once with Aleutian sandpipers." He 
does not mention localities here, but his observations covered the 
general region from Kodiak to Sanak Island. 

On February 7, 1941, F. L. Beals obtained a male specimen 
on Amchitka Island. 

Stejneger (1885) reports the sanderling to be a rare migrant 
in the Commander Islands. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 155 

Family PHALAROPODIDAE 
^halaropus fulicarius: Red Phalarope 

Attu : A-clii-li-rhd-uch 

Russian, Yana River region: Plavounetz, more often Petouschok (Pleske) 

We observed flocks of red phalarope in spring migration as well 
as later in the summer, when some of them may have been re- 
suming from the north. On May 22, 1937, several flocks were 
seen in Shelikof Strait, many in the red plumage, and the next 
.lay, they were common all the way between Sutwik Island and 
the Shumagins. These were chiefly in the red plumage. On Au- 
gust 26, as we approached East Unalga Island from Unalaska, 
flocks were seen, this time in whitish winter plumage. On the 
evening of May 21, 1936, while passing off"shore from Unimak Is- 
land in Bering Sea, we saw bands of red phalaropes, totaling 
nearly 100, flying over the water. On July 15, over 100 were fly- 
ing near the Baby Islands in Akutan Pass, and near Rootok Is- 
land, Next day, more of these birds were seen near Rootok Island. 
On two occasions, they were seen feeding along a line of dead kelp. 

Cahalane (1943) says "N. J. Benson told me that in August 
1940 he had seen a flock of 'at least five thousand' of the 'whale 
birds' in Shelikof Strait." 

Turner (1886) wrote that he "saw but few of these birds at 
Nushagak. At the mouth of Ugasik River, and the low grounds 
surrounding it, I saw hundreds of these birds." 

Jaques (1930) says that the red phalarope was "First seen 
near the Shumagin Islands May 15 and 16, at Moller Bay, and 
throughout Bering Sea on the northward voyage." And again, 
"Only one bird (at Port Moller) was seen on or near the shore." 

There is a strong probability that a few red phalaropes nest 
on parts of Alaska Peninsula. Turner's observations at Nushagak 
and Ugashik, and the bird noted by Jaques at Port Moller, 
suggest nesting, because these are birds of the open sea when 
on migration. Furthermore, on May 25, 1925, I found a female 
along the stream flowing northwesterward from Aghileen Pin- 
nacles, on the north side of Alaska Peninsula, and on May 29 
another female was flushed from a pond in the upper end of the 
same valley. On June 22 Donald Stevenson shot a female at 
Hazen Point; he thought that this bird had incubation patches. 

Nelson (1887) says "It is an abundant summer visitant on 
the Near Islands, .and breeds abundantly on some of the Com- 
mander group." 

Turner, on the other hand, (1886) says "I have no record 



156 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

of their occurrence in the Aleutian Islands. They may occasionally 
occur there with other species." 

Stejneger (1885) merely reports a flock seen at sea, near the 
Commander Islands, Hartert (1920) records five specimens taken 
on the Commander Islands, and he remarks that the late dates, 
May 25 to June 16, suggest breeding. 

Clark (1910) reports that when he approached Unalaska 
"thousands of these birds were seen, mostly in flocks of from 
fifty to a hundred or more, but many singly or in small com- 
panies." 

The chief of Attu Village said that the red phalarope does 
not nest in the Aleutians, but he stated that it is plentiful there 
in winter. This probably is true, though the above data gives 
good evidence of nesting along the Alaska Peninsula. 

Lobipes lobafus: Northern Phalarope 

Attu: Chirr-teg-ech 

Chimt-khukh (according to Turner) 
Atka: Chir-riz-ing-ah 

Large numbers of the northern phalarope migrate along the 
southern Alaskan coast. On May 8, 1937, while passing through 
Snow Pass in southeastern Alaska, we enjoyed the impressive 
spectacle of several thousands of northern phalaropes resting on 
the water. There was much dead kelp, which apparently afforded 
good feeding. On May 16 and 17, northern phalaropes were 
abundant on the tide flats at Eyak River, near Cordova, and local 
residents declared that they nest there. On May 11, 1936, we 
observed a small group between the Barren Islands and Afognak 
Island, and on May 13 we saw a flock of about 25 in Kupreanof 
Strait as well as smaller groups near Kodiak Island. All of these 
flocks were seen over open water, where they sometimes alighted 
and swam about. 

Cahalane (1944) observed two northern phalaropes in the 
Katmai region in September 1940, and Hine (1919) noted them 
on a number of occasions near the mouth of Katmai River where 
he obtained specimens. 

Littlejohn wrote : "Seen often at sea in large flocks and found 
nesting at Kodiak and Sanakh in April 1888." 

On May 25, 1936, two were seen near a pond on the tide flat 
at Snag Point, Nushagak River, evidently preparing to nest, and, 
on May 27 and 29, on the tide flats at Ugashik River, many more 
seemed to be preparing to nest. Some were seen in small flocks, 
others in twos and threes. 

Jaques (1930) found this bird "abundant about the Port 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 157 

/Toller region in all sorts of pools on the tundra after June 1 
ntil our departure on June 22." 

Gianini (1917) noted the species at Stepovak Bay as "one of 
he most common and interesting of the smaller birds. Every 
lond had a pair or more ... I found no nests nor saw any young, 
et these birds breed there." 

In 1911, Wetmore found these phalaropes evidently breeding 
n the Morzhovoi Bay region. 

In 1925, I observed many northern phalaropes, obviously 
ireeding, in the wet valley bottom below Aghileen Pinnacles, on 
lazen Point, and on the marshes at Moffet Cove. Two males 
hat were collected June 15 had incubation patches, and, on July 
.9, Donald Stevenson saw a young bird. 

Turner (1886) says: "Hundreds of them were seen on the 
ow grounds on the northern side of Alaska." 

The northern phalarope also nests on many of the Aleutian 
slands. We found them on Unimak, Unalaska, Atka, Little 
?anaga, Adak, Amchitka, Ogliuga, Little Sitkin, Kiska, Little 
Ciska, Buldir, Semichi, and Agattu. Swarth (1934) reports a 
)air taken on Akutan. 

Wetmore found them nesting on Adak, Tanaga, and Kiska, and 
le believed that they nested on Atka. 

Turner (1886) says that they are abundant on the western 
slands in the Aleutian chain, and he adds that many of them 
)reed on Atka, Amchitka, Semichi, and Agattu. 

On Buldir Island, we were much interested to find two of these 
)irds high on the mountain, in the area occupied by nesting 
leese. 

Stejneger reported the northern phalarope to be a common 
)reeding bird in the Commander Islands. 



Family STERCORARIIDAE 
Ifercorar'ius pomarinus: Pomarine Jaeger 

Russian, Yana and Indigirka regions: Terbei (Pleske) 
Chukchi: Aunuklinuadr-ukanodliyi (Palmen) 

According to Pleske, the Russian name "Terbei" applies to 
jaegers in general. He states that in northern Siberia, people 
3f various languages use one name for all jaegers, adding "large" 
3r "small" for the different kinds. Similarly, among some 
Eskimos I found that the same name was applied to parasiticus 
and longicaiulu-s. 



158 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE ! 

The pomarine jaeger proved to be a rare bird in the AleutiaiJ 
district, and there was no evidence of nesting. I 

On the evening of May 21, 1936, several miles off Urilia Baj, 
Unimak Island, 3 pomarine jaegers passed the ship — 2 wer s 
together, followed by a single bird that Cecil Williams identifieii 
at close range. On May 22, another pomarine jaeger was seeii 
offshore from Nelson Lagoon, Alaska Peninsula, and several 
others were seen farther east later in the day. On two occasions, 
we noticed a jaeger trying to rob an Arctic tern. On May 23 
three or four jaegers were seen in outer Nushagak Bay. Farther 
west, on July 4, three jaegers were observed between LittL 
Tanaga and Kagalaska Islands. They probably were pomarinus 
because they were large and were light underneath; however' 
positive identification was impossiblq. Another was seen at Wes 
Unalga Island on August 3. 

Pomarine jaegers were seen again in 1937. One was seei 
near Resurrection Bay on May 20; 1 was observed near th 
Shumagins, May 23, and several were seen near Deer Island 
May 24; 1 was seen near Unimak Island, May 25; and 1 wa 
sighted west of Kiska Island, June 6. One June 17, the captain o 
our ship counted 7 of these birds off Semichi Islands — I verifie( 
4 of them. Later in the day, another was seen at sea, farthe 
eastward. On August 19, at Cape Cheerful, near Unalaska, ther 
were quite a number of these jaegers among the shearwaters 
and on August 24 one was seen near Bogoslof, among gulls anc 
shearwaters. 

Austin H. Clark (1910) saw a pomarine jaeger at Bower's Ban] 
in Bering Sea on June 3 — this is the only one that he noted. 

Pomarine jaegers were always found either at sea or weJ 
offshore, they never were seen on land. Apparently, the water 
about the Alaska Peninsula and the Aleutians afford excellen 
foraging for nonnesting individuals. Jaegers probably parasitize 
gulls and shearwaters in this area, though no doubt they are als^ 
capable of feeding directly from the water, where marine organ 
isms are so abundant. 

In 1924, we found the pomarine jaeger nesting commonly a 
Hooper Bay, and no doubt the nonbreeding individuals would b 
attracted to the Aleutian area, which must lie in their migratioi 
path. 

Hartert (1920) records two specimens taken on the Commande: 
Islands. 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 159 

Sfercorar'ius parasiticus: Parasitic Jaeger 

Attu: Klu-pa-soch 

Atka: Ke-uch 

Russian, Commander Islands: Rasbojn.ik (Stejneger) 

Chukchi: UacW Ukangodlin (Palmen) 

The parasitic jaeger nests on Kodiak Island (Friedmann 1935; 
Bent 1921), throughout the Alaska Peninsula, and along the 
Aleutian chain. On May 23, 1936, 2 of these birds were seen in 
outer Nushagak Bay; on May 25, 3 were seen, and, the next day, 
2 were observed over the marshy tide iiats at Snag Point, 
Nushagak River. On May 27 and 29, a number of these birds 
were seen flying about over the tide flats at Ugashik River — 
both the light- and dark-color phases v/ere noted. C. S. Williams 
collected one in the light-color phase. On August 26, there were 
five (all of which were in the light-colored phase) over the marsh 
at Sand Point, Popof Island. 

On June 18, 1940, Gabrielson saw a pair at the Semidi Islands, 
and on July 17, he saw three birds at Dillingham. On July 19, 
he saw 12 parasitic jaegers on Naknek River. 

Jaques (1930) says the parasitic jaeger was "Common along 
shore and over the tundra north of Port Moller, where it was 
breeding in June." 

In 1925, I found these jaegers to be numerous about Izembek 
Bay during the nesting season, and I suspected that they were 
nesting, though proof was lacking. Wetmore reported these 
birds "tolerably common" about Morzhovoi Bay, and Gianini 
(1917) found them quite common about Stepovak Bay. 

We saw this bird frequently throughout the Aleutian chain. 
They were in pairs, on characteristic tundra habitat, and they 
acted in a manner typical of nesting. Unfortunately, we had no 
time to hunt for nests. 

On June 12, 1936, 4 pairs of jaegers were found on Chuginadak 
Island ; June 14, 1 was seen on Herbert ; June 18, a pair was seen 
on Seguam; June 29, 3 were seen on Kanaga; July 23, 2 were 
seen on Ogliuga; July 26, several were observed on Kiska; July 
30, 3 were sighted on Attu; July 31, at least 21 were seen high 
up on Buldir ; August 4, 5 were seen on Kavalga and 3 were seen 
on Skagul Island. 

In 1937, they were noted again : June 4 and 5, 6 were seen on 
Kiska ; June 7 and 10, 4 were seen on Attu ; June 11 to 15, common 
on Agattu; June 17, 6 were seen on Semichi; June 18, abundant 
on Buldir; June 21, common on Kiska; June 22, a pair was 
seen on Little Kiska; June 23, 3 were seen on Chugul; June 27, 
a pair was seen on Little Sitkin; June 30, at least 2 pairs were 



160 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

seen on Rat Island; July 4, 3 were seen on Semisopochnoi ; July 
5 and 11, about 7 were sighted on Amchitka; July 29, at least 
4 were seen on Ogliuga; July 30, three or four were observed on 
Kavalga; July 31, 2 were seen on West Unalga; and on August 2, 
2 were seen on Ilak. 

On the Alaska Peninsula, the black-color phase of the parasitic 
jaeger is particularly common, though the light phase probably 
predominates. In the Shumagins, all five birds seen were light 
colored. 

Among the Aleutian Islands, however, the light-color phase 
is a rarity. More than 100 parasitic jaegers were recorded, and, 
of this number, only 4 were specifically mentioned in our field 
notes as being light colored; nearly all the rest were mentioned 
as being definitely dark. Possibly in no other area is the parasitic 
jaeger population so uniformly dark. 

Stejneger (1885) says of the color phase that "On the Com- 
mander Islands the dark form is the most common. A few only 
with white lower surface were seen and one secured." 

Hartert (1920) obtained 4 adults in the Commander Islands 
with white underparts, and he obtained 3 of the dark phase. 

Dall (1874) noted the same tendency, believing, however, that 
the dark color was in the immature plumage ; this becomes obvious 
when he says "nor have we ever obtained any in completely 
adult plumage. All our specimens are of a nearly uniform dark 
slate color." 

Bent (1921) has suggested that the dark color phase may be 
a distinct species. That appears doubtful, however. Bent quotes 
Grinnell as saying that he found a light and a dark bird mated. 
On two occasions in the Aleutians we observed trios, one of 
which was white. Unless we can show that normally the two 
color types keep segregated in breeding, with only an occasional 
crossbreeding that may be construed as hybridization, it will be 
better to consider that they are color phases. 

A dark-color phase, becoming dominant or very prominent 
locally, is known among other animal species — for example, in 
the case of the marmot in parts of the Rocky Mountains, the 
ground squirrel in eastern Alaska, and the Arctic fox in the 
Aleutians. 

Food Habifs 

The name of this bird suggests its food habits. It is known 
to rob gulls and terns of their food. The Arctic tern and European 
turnstone were seen pursuing parasitic jaegers, evidently rec- 
ognizing them as foes. On Alaska Peninsula, there was evidence 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 161 

hat these jaegers were seeking fragments of salmon left on 
he banks by brown bears. On Buldir Island, where parasitic 
aegers were so numerous, a colony of nesting glaucous-winged 
rulls probably furnished a food supply for the jaegers. 

On Agattu Island, 24 pellets were obtained ; these pellets con- 
;ained the remains of 17 forked-tailed petrels, 3 murres, and 5 
midentified birds. Of the last mentioned, 2 were possibly paroquet 
luklets and 1 was a small, sparrow-sized bird. 

Some of these items, especially the murres, were undoubtedly 
carrion. A nesting colony of forked-tailed petrels, near the 
perch where the pellets were obtained obviously was the source 
Df the items found in the pellets — however, the method of capture 
was not ascertained. It is, of course, possible that the jaegers 
found parts of petrel carcasses left by blue foxes. 

Certain observations suggest that the parasitic jaeger is 
not solely a carrion eater and robber, but that it hunts part of the 
time in the manner of a hawk. On Semisopochnoi Island, Douglas 
Gray and I sat on a slope in the midst of a least auklet colony 
and watched the performance of a parasitic jaeger. For over an 
hour, we watched the bird repeatedly pursue these little auklets 
as the flocks came in from the sea. It did not stoop from a 
height, but it would single out a bird and follow it as swiftly 
as possible on the level or at various angles, in irregular flight. 
There are two possible interpretations. It may have been trying 
to capture an auklet, or it may have been trying to make it 
disgorge. So far as we could see, in spite of its persistence, it 
did not succeed in either purpose. 

Stejneger (1885) says, of the Commander Islands: "In the 
autumn they seem to feed to a great extent on the berries of 
Empetrum nignim, and their excreta at that time are colored 
dark blue." 

Sfercorar'ius longJcaudus: Long-tailed Jaeger 

Chukchi: A7ikakenuadl'-nkangodlin (Palmen) 

The long-tailed jaeger is rare in the Aleutian district. Fried- 
mann (1935) records a few bones found in middens on Kodiak 
Island. Osgood (1904) reports one on Iliamna Lake, July 16, 
1902, and he records specimens taken by McKay at Nushagak and 
Ugashik in July and August 1881. 

On July 17, Gabrielson recorded a long-tailed jaeger at Dilling- 
ham; and, on July 19, he noted three on Naknek River. 

Gianini (1917) is the only observer who has reported these 
jaegers to be common; his observations were made at Stepovak 



162 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Bay, where he collected a specimen. Apparently, Wetmore did 
not observe them on Alaska Peninsula in 1911, and I did no1 
positively identify this jaeger at Izembek Bay in 1925. 

Turner (1886) says "The Long-tailed Jaeger is rarely seer 
on the Eastern Aleutian Islands. I saw one on Sannakh Island ir 
July, 1878. I saw a few at Atkah Island in 1879, and two a1 
Attn Island in 1880 . . . This species is reported to breed at the 
Semichi Islands." 

Friedmann (1934) reports a number of bones of this jaegei 
found in middens on Amaknak Island. 

Stevenson obtained a specimen on Ilak Island, September 8, 
1921. 

We observed it only once in the Aleutians. On June 13, 1937, 
I watched a bird in the normal light-colored plumage, flying about 
with three parasitic jaegers. 

Clark (1910) reports seeing one on Bower's Bank, Bering Sea. 

Hartert (1920) records two specimens from Commander Is- 
lands. Stejneger (1885) stated that they do not nest there. 



Family LARIDAE 

Larus hyperboreus: Glaucous Gull 
Larus hyperboreus hyperboreus 

Russian, Murman coast: Kluscha (Pleske) 
Chukchi: Yttak, tchikerga (Palmen) 

Though the glaucous gull normally nests north of the Aleutiani 
district, it reaches this area in considerable numbers in winter! 
and in migration to more southern localities. As Friedmann' 
suggested (1935), it is practically certain that Turner was in 
error when he reported "countless thousands" of these gulls on 
cliffs at Kodiak. Friedmann records several bones found in 
middens on Kodiak, adding, "Macoun mentions a bird in the Hen- 
shaw collection, and 3 eggs taken in June, 1880, now in thei 
Mailliard collection, [which] are the only other records I have 
found." 

While these are recorded under the name Larus h. hyperboreus,\ 
they could possibly refer to L. h. harrovianus. We do, however, 
have at least one undoubted specimen of L. h. hyperboreus — a 
male taken at Unalaska Island by Wetmore on June 9, 1911. 
Though this specimen was listed by Oberholser (1918) under 
L. h. harrovianus, examination of the speciman shows that it 
has the massive beak that characterizes L. h. hyperboreus, the 
measurements being greater than in barrovmnus. Since it is 



FAUNA OF THE ALEUTIAN ISLANDS AND ALASKA PENINSULA 163 

[nown that the eastern glaucous gull nests on St. Matthew and 
/Valrus Islands, we would expect it to visit the Aleutian district 
it times, though the western glaucous gull would be most pre- 
valent. Oberholser (1918) also mentions Diomede Islands for 
;his form. 

.arus hyperboreus barrovianus 

Though the status of barrovianus has been belabored repeatedly 
3y able ornithologists, certain specimens obtained in the Aleutians 
nduced me to examine the whole question again. More than 200 
specimens were examined in the U. S. National Museum and in 
;he American Museum of Natural History in New York. 

It is useless to deny the difficulties in recognizing barrovianus 
is a distinct form. In studying series from a given locality, one 
is confronted with specimens that do not fit a given description. 
Gulls are variable, and one must be cautious in arriving at con- 
:lusions. On the other hand, if one is careful to give due weight 
to breeding territory, and to allow for migration to explain cer- 
tain irregularities, many of these difficulties disappear. 

All gulls of the species hyperboreus are pale mantled, but true 
hyperboreus is noticeably paler than barrovianus. Furthermore, 
hyperboreus is definitely larger and has a decidedly larger and 
more massive beak. Listed measurements do not adequately ex- 
press the difference. Depth of bill of the two forms overlaps, or 
meets, at about 23 mm., though most of them are above, or below, 
this figure, and a difi'erence of even 2 mm. makes a considerable 
diff'erence in appearance. 

A good series of specimens from Point Barrow and the east 
shore of Bering Sea are remarkably uniform in the characters 
assigned to barrovianus — the darker mantle, the smaller size, 
and especially the smaller bill. Available specimens from eastern 
North America are confusing, but it is notable that when winter 
specimens are eliminated, and apparent breeding birds are used, 
they fall more generally into the group of hyperboreus. This was 
especially true of Greenland, where a good series of breeding 
birds presented a clear picture of Larus h. hyperboreus, diS here 
described. 

The confusing aspect of the distribution of these two forms is 
the considerable number of small-sized birds found along the 
Atlantic coast in winter, which apparently agree with barro- 
vianus, but which are far from the type locality. Possibly we 
should expect this. Oberholser gave the breeding range as 
extending along the Arctic coast as far east as "the territories of 



164 NORTH AMERICAN FAUNA 61, FISH AND WILDLIFE SERVICE 

Yukon and western Mackenzie." It is entirely possible that the 
breeding range extends much farther east. Among specimens 
examined, L. h. hyperboreus was found to the westward across 
northern Eurasia, Greenland, and northeastern North America! 
as far west as Baffin and Ellesmere Islands. In any case, it may 
be expected that many of the birds can find their way from 
Arctic Canada to the Atlantic coast in winter. The gulls are far- 
ranging birds. Steller's eider has been recorded from the Gulf 
of St. Lawrence (Fisher 1900). 

It seems logical to consider Larus h. hyperboreus as breeding 
throughout the Arctic regions of Siberia and Europe, traveling as 
far west as Baffin and Ellesmere Islands (and probably neighbor- 
ing areas) and, from the west, traveling eastward to the Beringi 
Sea coast of Siberia. In Bering Sea, the birds obviously have' 
come eastward as far as St. Matthew and Walrus Islands. There 
is a specimen from St. Matthew taken