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HARVARD UNIVERSITY

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Museum of Comparative Zoology

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MAMMALS OF WASHINGTON

BY

WALTER W. DALQUEST

University of Kansas Publications
Museum of Natural History

Vol. 2, pp. 1-444, 140 figures in text
April 9, 1948

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MAR -8 1350

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UNIVERSITY OF KANSAS

LAWRENCE
1948

MAMMALS OF WASHINGTON

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Mounl l; linn i from [ndian Henry's Bunting Ground, July. 1934. (.Fish and
Wildlife Service photo by Vich.i IV SdiH'tVr. \.> S64.)

MAMMALS OF WASHINGTON

j

BY

WALTER W. DALQUEST

(Contribution from the Museum of Natural History. University of Kansas)

University of Kansas Publications
Museum of Natural History

Volume 2, pp. 1-444, 140 figures in text

April 9, 1948

toUS. MP. ZOOL
ilBRW

IHAR -8 1950

UNIVERSITY

UNIVERSITY OF KANSAS

Lawrence

1948

(7)

University of Kansas Publk vtions, Mi seum of Natural Historv
Editors: E. Raymond Hull and Donald F. HotTmeistcr

Volume 2, pp. 1-444. 140 figures in text
April 9, 1948

University of Kansas
Lawrence, Kansas

Printed by

Ferd Voiland Jr., State Printer

Topeka, Kansas

1948

?l-4!)!i:{

(S)

CONTENTS

PAGE

Introduction 13

Physiographic Provinces of the State 16

Distributional Areas 20

Climate and Vegetation 25

Life-zones and Ecology 32

Geologic History of Washington 46

The Faunas 52

Speculation as to Emigrational History of the Mammals. . 54
Speculation as to the Later Distributional History of the

Mammals 68

Explanation of Treatment 110

Check List of Mammals 112

Accounts of Species and Subspecies 121

Addenda 416

Bibliography 417

Index 430

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ILLUSTRATIONS

Topography

1 [(JURE PAGE

Mount Rainier from Indian Henrys frontispiece

1. Cascade Mountains at Canadian Boundary 17

2. Columbia River one mile south of Kellers Ferry 19

4. Cascade Mountains in Chelan National Forest 21

5. Mount Rainier. Yakima Park 22

6. Columbia River in Stevens County 23

7. Blue Mountains, Washington 24

8. North Side of Mount Rainier 26

9. Mount Rainier : Cowlitz Chimneys 28

10. A "pothole" crowded by drifting sand 30

12. Arctic-Alpine Life-zone on Mount Rainier 33

13. Humid subdivision of the Transition Life-zone 34

14. Timbered, arid subdivision of the Transition Life-zone 85

15. Upper Sonoran Life-zone 37

16. Canadian Life-zone 38

18. McDowell Lake, Little Pend Oreilh Wildlife Refuge 50

19. Pend Oreille River near Newport 59

20. Rocky bluff along north bank of the Columbia River 65

Mammals

FIGURE PAGE

21. Gibbs shrew-mole 122

23. Coast mole and Townsend mole 125

25. Coast mole 128

41. Long-eared bat: female with young 161

42. Boulder Cave : habitat of long-can d bats 162

48. Female black bear and two cub- ■. 171

49. Black bear in "hibernation" 173

54. Fisher 187

59. Wolverine 202

60. Wolverine: dried pelt 203

62. River otter 208

66. Badger 220

69. Coyote 226

72. Cougar oi mountain lion : skin 234

73. Cougar or mountain lion : pelts 235

75. Canadian lynx 238

7(i. Trapper's catch of nun Canadian lynx 239

7s Bobcat 241

82. Townsi nd chipmunk 260

87. Golden-mantled ground squirrel 282

vi. Feeding si .i f Honda- -quirre] 287

91. Northern flying squirrel 292

95. Northern pockel gopher 302

96. Gianl mounds formed by pockel gophers 304

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Illustrations 11

Figure page

97. Food cache of northern pocket gopher 306

99. Beaver 315

100. Beaver lodge and pond 316

101 . Cottonwood pole carved by beaver 317

102. Road flooded by beavers 320

106. Deer mouse 327

112. Pennsylvania meadow mouse 345

1 15. Runways of Townsend meadow mire 350

119. Muskrat 360

121. Mountain beaver 366

123. Big jumping mouse in hibernation 370

130. Elk 392

131. Group of elk 393

132. White-tailed deer 395

133. White-tailed deer: fawn 396

134. Left antler of white-tailed deer 397

135. Antlers of white-tailed deer 398

136. Mule deer 400

137. Black-tailed deer 401

138. Mountain goat 407

139. Group of mountain goats 408

Distribution" Maps

3. Mammalian distributional areas 20

11 . Life-zones of Washington 32

17. Extent of Vashon-Wisconsin ice 48

22. Gibbs shrew-mole 123

24. Townsend mole 127

26. Coast mole 129

27. Cinereous shrew 1 33

28. Merriam shrew and Trowbridge shrew 134

29. Wandering shrew 1 37

30. Dusky shrew 140

31 . Mountain water shrew 141

32. Bendire water shrew and pigmy shrew 143

33. Big myotis 147

34. Yuma myotis 149

35. Fringe-tailed myotis and Keen myotis 150

36. Long-eared myotis 152

37. Hairy-winged myotis 155

38. California myotis 156

39. Small-footed myotis 157

40. Silver-haired bat 160

43. Long-eared bat 163

44. Western pipistrelle 164

45. Big-brown bat 1 66

46. Hoary bat 168

47. Pallid bat 170

50. Black bear 175

51 . Grizzly bears 1 77

12 University of Kansas Pvbls., Mrs. Nat. Hist.

Figure page

52. Raccoon 181

53. Western marten 185

55. Fisher 188

56. Ermine 192

57. Long-tailed weasel 196

58. Mink 199

61. Wolverine 204

63. River otter 209

64. Civet cat 214

65. Striped skunk 218

67. Badger 221

68. Red fox 225

70. Coyote 230

71. Wolf 232

74. Cougar 236

77. Canadian lynx 240

79. Bobcat 243

80. Least chipmunk 252

81. Yellow-pine chipmunk 256

83. Townsend chipmunk 261

84A. Marmots 266

84B. Townsend and Washington ground squirrels 268

85. Columbian and Beechey ground squirrels 274

86. Golden-mantled ground squirrels 280

88. Western gray squirrel 284

90. Red and Douglas squirrels 289

92. Northern flying squirrel 294

93. < treat Basin pocket- mouse 298

94. Ord kangaroo rat 300

98. Northern pocket gopher 308

103. Beaver 321

104. Northern gras>hopper mouse 323

10.5. Western harvest mouse 325

107. Deer mouse 329

108. Bushy-tailed wood rat 335

109. Northern lemming mouse 337

110. Beather vole 339

111. ( rapper and California red-backed mic( 342

113. Pennsylvania meadow mouse 346

111. Montane and Townsend meadow mice 348

116. Long-tailed meadow mouse 352

117. Water rat 350

118. Creeping mouse 357

120. Muskrat 362

122. Mountain beaver 368

124. Big jumping mou.se 372

125. Porcupine 375

126. Pika 378

127. Snowshoe rabbit 383

128. Black-tailed jack rabbit 386

129. Nuttall cottontail 388

INTRODUCTION

Mammals of Washington are of especial interest to the naturalist
because many of them are recent immigrants; much of Washington
was buried under thick glacial ice until relatively recently and
many of the mammals, therefore, have inhabited the area only since
the ice disappeared. The evolution or development of certain sub-
species, in Washington, has certainly occurred within the last few
thousand years. To be able thus to date such evolutionary changes
as have occurred is of course a matter both of importance and in-
terest to zoologists. The evolutionary changes in several species are
relatively great. In color, for example, the bobcat in the humid
coastal area of western AVashington is notable for its dark color-
ation, whereas in the more arid area of southeastern Washington
it is remarkably pale and of a different subspecies. Within the
limits of the state of Washington, elevations ranging from sea level
to more than 14,000 feet occur. Since different elevations have their
characteristic mammals, more kinds are found in Washington than
in other areas of corresponding size that lack such topographic di-
versity. Expressed in terms of the life-zone concept, Washington
includes faunas ranging from the Upper Sonoran Life-zone to those
of the Arctic-Alpine Life-zone.

The basis for a study of the mammals of Washington was laid
in 1929 by W. P. Taylor and W. T. Shaw in the "Provisional List
of the Land Mammals of the state of Washington." Bailey's "Mam-
mals and Life Zones of Oregon'' and "The Recent Mammals of
Idaho" by W. B. Davis deal with the habits and distribution of
mammals in the areas bordering AVashington on the south and east,
and were very useful in the organization of the present report.

The study was first planned from the taxonomic and ecologic
point of view. Such a study, of necessity, involves the classification
and distribution of the forms concerned. Classification has required
more work than any other part of the study and has been, in a
sense, the nucleus of the study. Nevertheless, as the report began
to take form it was recognized that the part dealing with classifi-
cation and other purely technical aspects of the paper probably
would be uninteresting to the average reader. Therefore it was felt
that a greater impetus to the study of the mammals of Washing-
ton would be given by reducing the taxonomic accounts to the mini-
mum and dealing principally with the problems of distribution.

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14 University of Kansas Publs., Mrs. Nat. Hist.

As originally planned, the present report was to be of joint au-
thorship by Dr. Victor B. Seheffer of the United States Fish and
Wildlife Service, Seattle, and the writer. The press of other work
prevented Dr. Seheffer from devoting as much time as he had
planned to the project. He has, however, contributed his field
notes, specimens, and photographs, and in many other ways as-
sisted in the project.

Field work on mammals of Washington was carried out by the
writer from 1936 to 1940 but a decision to prepare a complete re-
port was not reached until 1938. Intensive field work was done be-
tween 1938 and 1940. In August, 1941, the author took up resi-
dence at Berkeley, California. Drafting of the manuscript was be-
gun at that time as a student under Professor E. Raymond Hall.
War conditions and the press of other work delayed completion.
Subsequently, the manuscript was put in final form at the University
of Kansas.

Many persons in addition to Dr. Seheffer have given assistance
in the course of this work. Dr. E. Raymond Hall, in particular,
encouraged the project and gave assistance in various ways includ-
ing critical attention to the manuscript. I am indebted also to Dr.
Trevor Kincaid and Mrs. Martha Flahaut of the University of
Washington, Dr. Seth B. Benson and Dr. Alden H. Miller of the
University of California, Dr. H. H. T. Jackson of the U. S. Fish
and AVildlife Service, Dr. George E. Hudson of the Charles R. Con-
ner Museum, Mr. Burton Lauckhart of the State of Washington
Department of Game, and Mr. Ernest Booth of Walla Walla Col-
lege. Mrs. Peggy B. Dalquest typed and edited the several pre-
liminary drafts of the manuscript and aided in the laboratory and
field work. Thanks are due also to many others, including game
protectors, hunters and trappers, who have given assistance. The
names of some of them are mentioned in the following pages.

Approximately ten thousand specimens of mammals were used.
In decreasing order, according to the number of specimens studied
from Washington, the following collections are to be mentioned:
materials obtained principally from southern Washington in the
years 1939 to 1942 as a result of the interest of Miss Annie M.
Alexander and Dr. E. Raymond Hall; these materials are in the
University of California Museum of Vertebrate Zoology. The
writer'- own collection which at one time numbered 2,500 specimens
was the second source. The residue, the part not destroyed by fire
at the writer's home in the spring of 1942, in Seattle, now is in the

Introduction 15

Museum of Vertebrate Zoology at the University of California
2nd the Museum of Natural History at the University of Kansas.
The other collections are those of the Biological Surveys of the
United States Fish and Wildlife Service, the Washington State
Museum at Seattle, the Charles R. Conner Museum at Washington
State College, and the Museum of Natural History at the University
of Kansas. In the latter collection are some materials obtained
nearly half a century ago by the late L. L. Dyche, some recently
taken specimens added by reason of the provision for work of this
kind by the University of Kansas Endowment Association and, as
noted above, a part of the author's original collection. Selected
specimens from several other collections have been used and these
are indicated in the text when particular reference is made to the
specimens. Most of the specimens studied were conventional study
skins with skulls. In some instances skeletons, skins alone, skulls
alone, or entire animals preserved in alcohol have been used.

16 University of Kansas Publs., Mus. Nat. Hist.

PHYSIOGRAPHIC PROVINCES OF THE STATE

The state of Washington was divided into seven physiographic
provinces by Culver (1936). Culver points out that the physiog-
raphy, though complicated in detail, is basically simple.

The state, including Puget Sound and other inland waters, is
nearly rectangular in shape and is 69,127 square miles in area. Its
western boundary is the Pacific Ocean. Politically, it is bounded
on the north by the United States-Canadian Boundary (49° north
lat.) , on the east by the state of Idaho, and on the south by the state
of Oregon.

The Cascade Mountain Range, or Cascade Mountains Province,
runs from the northern to the southern boundary and divides the
state into two sections, of which the eastern is slightly the larger.
The mountain range trends approximately 10° east of north and
continues uninterruptedly into British Columbia, but on the south
the Columbia River separates the Washington Cascades from the
Cascades of Oregon. Near the northern border of the state the
range is wide, extending from the Mount Baker Range on the west
to Mount Chopaka, 100 miles to the east. In the central part of
the state it is more compact, being some 60 miles wide in the vicinity
of Mount Rainier. Farther south it expands to approximately 100
miles.

The Cascades of Washington possess five great volcanic cones.
These are Mount Baker (10,750 feet elevation) on the north, Glacier
Peak (10,436 feet) in the north-central part, Mount Rainier (14,408
feet) in the central area, and Mount Adams (12,326 feet) and
Mount St. Helens (9,697 feet) on the south. Excluding these vol-
canic peaks, the crests of the Cascades have a relatively uniform
level descending from an average of 8,000 feet at the north to 5,000
feet at the south. Ranges jutting southeastwardly from the north-
central Cascades parallel elongate intermontane valleys. These
include the Entiat and Wenatchee mountains. The latter range
reaches the Columbia River and forms an important hairier to
mammalian movements.

There is an extensive area of anticlinal ridges extending from the
southern Cascades to the Columbia River. This area includes the
Horse Heaven and other hills. Most of it is drained by the Yakima
River and is termed, in this report, the Yakima Valley Area.

Physiographic Provinces

]7

The Puget Sound Trough, or Puget Sound Province, is immedi-
ately west of and parallel to the Cascade Mountains. It is part of
a structural downwarp that extends southward into Oregon. Most
of the area is below 1,000 feet elevation, and much of the northern
part is below sea level and therefore flooded by the marine waters

Fig. 1. Cascade Mountains at Canadian boundary, looking west along
boundary trail. Monument 100 in foreground, headwaters of Ashnola River in
broad valley at ri«;ht. (Forest Service photo. No. 4328.)

of Puget Sound. The most prominent feature of this area is Puget
Sound. This is a glacially-carved and drowned river valley, studded
with islands, peninsulas, fjords and bays that all possess a general
north-south orientation resulting from the direction of ice movement.
Puget Sound is connected with the Pacific Ocean by the Strait of
Juan De Fuca, a wide channel separating the state of Washington
and Vancouver Island.
2—4993

18 University of Kansas Plbls., Mis. Nat. Hist.

The San Juan Inlands represent the glaciated remnants of moun-
tains that, in pre-glacial time, may have connected the mountains
on Vancouver Island with the Cascades of Washington. The San
Juan Islands lie at the junction of Puget Sound, the Strait of
Georgia, and the Strait of Juan De Fuca. As a result of a boundary
dispute and subsequent arbitration, the islands were apportioned.
on the basis of the deepest channel separating them, between Canada
and the United States. The American portion includes more than
400 islands. These vary in size from mere rocks above high tide
to Orcas Island, 60 square miles in area.

The Olympic Peninsula, or Olympic Province, lies between Puget
Sound and the Pacific Ocean. The Strait of Juan De Fuca separates
this peninsula from Vancouver Island on the north. In the south
the valley of the Chehalis River is a convenient boundary for the
province. The central portion of the peninsula is occupied by the
Olympic Mountain Range. This range is nearly oblong in shape,
measuring some 70 miles east to west by 45 miles north to south.
The mountains are extremely rough and jagged. They rise from
sea level to above 6,000 feet. The highest peak. Mount Olympus,
is 8.150 feet in elevation.

South of the Olympic Province and west of the Puget Sound
Trough is an area of low, rough hills. Culver called it the Willapa
Hills Province.

The northern third of the land east of the Cascade Mountains,
or northeastern Washington, is termed the Okanogan Highland
Province by Culver. Its southern boundary is set at the east-
west flow of the Spokane and Columbia rivers. The outstanding
physiographic feature of this area is its division into north-south
trending areas of lowland with intervening highlands and mountain
ranges. The rivers are, from east to west, the Clark Fork, Colville,
Columbia. Kettle, San Poil and Okanogan. Not .ill intervening
highlands are separately designated as mountains. Among these
named are the Pend Oreille, Huckleberry, Kettle River, and
Okanogan ranges.

The part of eastern Washington south of the Okanogan Highland
Province, save the extreme southeastern corner of the state, consti-
tutes the Columbia Lava Province. This is an extensive, relatively
level plateau that lies mainly below 2,000 feet elevation. The
plateau consists of gently folded lava flows that reach a depth of
f. 1)00 feet in some places (Russell. 1893) and slope inward from the

Physiographic Provinces

19

east, north, and, in part, the west (Flint, 1938). These horizontal
layers of basalt are extremely resistant to erosion by other than
large rivers. Two great gashes cross the Plateau diagonally from
the northeast to the southwest; these are Moses Coulee and the
Grand Coulee. These old coulees are the former vallevs of the

^mm^

Hi

Fig. 2. Columbia River one mile west of Kellers Feny, Washington, ele-
vation 1,060 feet, April 16, 1940. (Fish and Wildlife Service photo bv Victor
B. Scheffer, No. 933.)

Columbia River, and were formed at the time when the course of
the river was successively blocked by the advance of Pleistocene ice.
The Snake River crosses the southern edge of the Columbia Lava
Province and separates the plateau proper from an area of similar
land to the southward.

The Blue Mountains Province is an area of relatively small ex-
tent in the extreme southeastern corner of the state of Washington.
There, the province concerned constitutes, as it were, a northward
extension of the Blue Mountains of Oregon. The mountains rise to
only 5,000 feet elevation in the Washington part of the Blue Moun-
tains Province.

20

University of Kansas Pvbls., Mis. Nat. Hist.

DISTRIBUTIONAL AREAS

The physiographic provinces are areas of land form. The form
of the land has a considerable effect on the temperature, humidity.
drainage, weathering, soil, and other non-organic features that com-
bine to produce the various life-zones and influence the distribu-
tion of mammals. One might therefore expect a close correlation
of mammalian distributional areas with physiographic provinces.
Although there is a correlation, it is not exact because the dis-
tribution of mammals is influenced also bv certain other factors.

I'H;. 3. Mammalian distributional areas of Washington. A. Western Wash-
ington. B. Cascade Mountains. ('. Northeastern Washington. 1). Blue
Mountains. E. Southeastern Washington. F. Yakima Valley. ('<. Colum-
bian Plateau.

Among these are historical factors and isolation by geographic bar-
riers.

The Cascade Mountains Province of Culver includes the Yakima
Valley Area. This province contains two completely different mam-
malian distributional areas. The higher mountains possess a boreal,
alpine fauna; the Cascade Range itself i- called the Cascade Area

Distributional Areas

21

in this report. The Yakima Valley Area possesses a desert fauna
derived from the desert of eastern Oregon.

The land west of the Cascades is separated into three physio-
graphic provinces, the Puget Sound, Willapa Hills, and Olympic
Mountains Province. The differences between the mammalian
faunas of the Puget Sound and Willapa Hills provinces are slight.
The Olympic Mountains possess a few species not found in the
lower areas. The similarities of the faunas of the three provinces
far outnumber their differences, and it seems best to consider them
subdivisions of one distributional area.

Fig. 4. Cascade Mountains in Chelan National Forest, looking southwest at
Straight Ridge; Cataract Creek (Methow watershed) at left. (Forest Service
photo, No. 4260.)

The Okanogan Highland Province extends, from a physiographic
point of view, west of the Okanogan River Valley. This valley.
however, is a fairly efficient barrier to mammals. Thus the part of
the state east of the Okanogan Valley and north of the east-west
flow of the Spokane and Columbia rivers may be called the North-
eastern Washington Distributional Area.

22

University of Kansas Ptjbls., Mrs. Nat. Hist.

The Columbia Lava Province includes the land both north and
south of the Snake River. Since the Snake River serves as a barrier
to some species, it seems better to term the area north of the Snake
River the Columbian Plateau Area and that to the south the
Southeastern Washington Distributional Area.

The Blue Mountains Province and the Blue Mountains Distri-
butional Area are the same.

Fio. 5. Yakima Park (or Sunrise Park), elevation 6.000 feet. Mount Rainier.
August 29. 1932. (Photo by 116th Photo Section, Washington National Guard,
No. 014-36A-116.)

DlSTRlBLTIOXAL AREAS

23

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v *v

' 2i_*?- *.-F i>:

..<** * '■" <-«r

* "♦*

♦.*

Fig. 6. Columbia River at Hunters Ferry, Stevens County, Washington,
April 15, 1940. (Fish and Wildlife Service photo bv Victor B. Scheffer, No.
932.)

24

University of Kansas Publs., Mus. Nat. Hist.

Fig. 7. Blue Mountains, Umatilla National Forest, Washington, looking
north-northeast across Al Williams Ridge to Tucannon River; 1933. (Forest
Service photo, No. 4437.)

Physiographic Provinces

Cascade Mountains

Puget Sound

W ilia pa Hills [

Olympic Mountains J

Okanogan Highlands
Columbia Lava
Blue Mountains

Distributional Areas

Cascade Mountains
Yakima Valley

Western Washington

Northeastern Washington

Columbian Plateau
Southeastern Washington

Blue Mountains

Climate and Vegetation 25

CLIMATE AND VEGETATION

The life-zone theory of plant and animal distribution was proposed by
Merriam (1892). Merriam's life-zones have been severely criticized by many
authors, especially because an error was made in computing some of the data
on temperature. However, zonation of vegetation and animals is obvious in
Washington, and the life-zone concept has been employed in Washington by
numerous botanists and zoologists. Among them are: Piper (1906), Taylor
and Shaw (1927), Jones (1936, 1938) and St. John (1937).

The higher parts of the Cascade Mountains are in the Arctic-alpine Life-
zone. This is the area of wind-swept ridges, living glaciers, and permanent
snow fields.

Trees are absent but a few shrubs are present; these include: Juniperus
sibirica, Salix cascad-ensis, Salix nivalis, Gaultheria humijusa, Empetrum
nigrum, and the heathers, Phyllodoce alanduliflora, Cassiope mertensiana and
Cassiope stelleriana. Jones (1938) lists a total of 98 species of plants from the
Arctic-alpine Life-zone of Mount Rainier. Many of these plants are most
abundant in the next life-zone lower, and are of but incidental occurrence in
the Arctic-alpine Life-zone. No mammalian species is resident but individuals
of several species regularly visit and occasionally breed there.

Below the Arctic-alpine the Hudsonian Life-zone stretches the entire length
of the Cascades. Temperatures are low, especially in winter; then the ther-
mometer does not rise above zero for weeks at a time. The average annual
temperature at Paradise, 5500 feet, Mt. Rainier, is 38.6° (all temperatures
given here are in degrees Fahrenheit). Snowfall is heavy. The average yearly
snowfall, for four years, at Mt. Baker Lodge, at 4200 feet elevation, Whatcom
County, was 478 inches; at Goat Lake, 2900 feet, Snohomish County, 261
inches; Tye, Stevens Pass, 3010 feet. King County, 398 inches; Paradise, 5500
feet, Mt. Rainier, 587 inches. The deepest snow recorded at Paradise was 27
feet, 2 inches on April 2, 1917. Following the spring thaws the mountain
passes are opened to travel, usually in April or May, although nightly tem-
peratures in April and May are still below zero. Spring precipitation is heavy,
the monthly average for a twelve-year period at Paradise being 6.78 inches in
April and 5.5 inches in May. Summer temperatures are high in the daytime,
when the sun beats down through the rarefied atmosphere, but cool at night
when accumulated heat is lost through the thin atmospheric blanket. In
summer precipitation is light, averaging, at Paradise, 3.46 inches in June, .9
inches in July, and 3.44 inches in August. In the autumn the temperature,
both daily and nightly, drops somewhat, and rain and cloudiness are the rule.
At Paradise the average precipitation in September is 8.29 inches and in
October 10.02 inches. The winter snows usually arrive by the middle of
November.

Trees that are characteristic of parts of the Hudsonian Life-zone include the
alpine fir {Abies lasiocarpa), mountain hemlock (Tsuga mertensiana) , Alaska
cedar (Chamaecyparis nootkatensis) and white-barked pine (Pinus albica/ulis).
The following shrubs are listed by Jones (1938) as common in the Hudsonian
Life-zone on Mt. Rainier: Salix barclayi, Salix eommutata, Juniperus sibirica,

26

University of Kansas Publs., Mus. Nat. Hist.

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Climate and Vegetation 27

Alnus sinvata, Ribes howeUii, Lutkea pectinata, Potentilla fruticosa, Sorbus
occidental is, Spiraea densiflora, Pachistima myrsinites, Arctostaphylos neva-
densis, Arctostaphylos uva-ursi, Cassiope mertensiana, Cassiope stelleriana,
Phyllodocc empetriformis, Rhododendron albiflorum and Gaultheria ovali-
Jolia.

There are extensive coniferous forests in the Canadian Life-zone, still lower
on the mountain slopes. This is an area of lesser temperature extremes than
is the Hudsonian Life-zone. The average annual temperature at Longmire,
2761 feet, Mt. Rainier, is 43.8°. The average temperature for the winter
months, however, is below freezing. In July and August the temperatures
are high, especially in the daytime. The eighteen year average for Longmire
during these months is 60.6°. Snow is regular but the fall is lighter than in
the Hudsonian Life-zone. The annual average, over a period of nineteen
years, is 184.4 inches at Longmire. Precipitation is similar to that in the
Hudsonian Life-zone, averaging perhaps slightly less.

In the Cascades the typical feature of the Canadian Life-zone is the ex-
tensive coniferous forest that extends, almost without a break, the entire
length of the Cascades on both sides of the main crest. In addition to Douglas
fir, the following trees occur in this forest: western hemlock (Tsuga helero-
phylla), amabalis fir (Abies amabalis), white pine (Pinus monticola) and
noble fir (Abies nobilis) . Other plants include Vaccinatum ovalifolium, Vac-
cinnium membranaceum, Menziesia jerruginea, Alnus sinuata, Acer circinatum,
Sorbus cascadensis. Cornus canadensis, Clintonia uniflora, Stenauthium oc-
cidentale, Galium oreganum, and Prenanthes lessingii. Saprophytes abundant
in, if not confined to, this zone are listed by Jones as: Monotropa uniflora,
M onotropa hypopitys, Allotropa virguta, N ewberrya congesta, Pterospora an-
dromedea, Corallorrhiza macvlata, Corallorrhiza mertensiana and Corallorrhiza
striata.

West of the Canadian Life-zone in the western Cascades, the coniferous
forests merge with the lowland forests of western Washington. To the east
of the Canadian Life-zone in the eastern Cascades, there is a distinct change
to a more arid climate and flora. At Leavenworth, 1167 feet, Chelan County,
the annual average precipitation is but 19.5 inches and at Cle Elum, 1930 feet.
but 23.23 inches. Temperatures are higher, the annual average of the above
two localities being 47.2° nd 45.4°. respectively. The winter months are cold,
with the average temperature in January and February below freezing. In
summer the averages in July and August at Leavenworth are 68.8° and 68.0°,
with the average maximum being 87.5° and 86.1°. Snowfall is heavy, the
yearly average at Leavenworth being 98.5 inches and at Cle Elum 86.3 inches.
The effect of this more arid climate is seen in the vegetation. The dense
Douglas fir forest is replaced by more open forests of yellow pine (Pinus
ponderosus) . Groves of oak (Quercus garryana) are found near streams. The
open forests give way to the extensive grasslands bordering the desert. The
transition of vegetation is similar to that occurring in the Yakima Valley Area.

In the Yakima Valley Area, arid conditions prevail. The rverage yearly
precipitation at Yakima is 6.67 inches. Only in November, December and
January may more than one inch of precipitation be expected monthly.
Snow may be expected in the winter months and the yearly average snowfall
is 22.1 inches. Winter temperatures are low, the average for December and

28

University of Kansas Publs., Mrs. Nat. Hist.

January being but slightly above freezing. Summer temperatures are ex-
treme; the July average is 73.1° and the average maximum for the same
month is 89°. The highest temperature recorded is 111°.

The open pine forests of the eastern Cascades give way to grasslands.
Grasses of several species are common but the bunchgrass (Agropyron spicatum)
is most important. Other plants include the primrose (Oenothera pallida),
lupines (Lupinus), and Mertensia. In ravines and near wartercourses such
shrubs as hawthorn {Crataegus douglasii), service-berry (Amelanehier cusickii,
Amelanchier utahensis), aspen (Populus tremuloides) , syringa (.Philadelphia
lewisii), snowberry (Symphoricurpus albus), choke-berry (Prunus nu lanocarpa)

Fig. 9. Mount Rainier. Washington: Cowlitz Chimnevs from base ol Dege
Peak, July 19. 1933. (Fish and Wildlife Service photo by Victor B. Scheffer,
No. 859.)

and elderberry (Sambuius cncrulae) form thickets. Lower in the valley the
vegetation is xerophytic, similar to that of the Columbian Plateau. Sage-
brush (Artcmisiae tridentata) is dominant. Other shrubs include rabbit
brush (Chrysol haunt u.s nauseosus, Chrysothamnus mscidiftorus) , hop sage
(Grayia spiyiosa), black sage (Purshia tridentata) and greas< wood (Sarcobatus
vermiculatm) .

Climatic conditions in the Olympic Mountains are, in general, similar to
tlmse of the higher Cascade Mountains. There is a limited area of Arctic-
alpine Life-zone on Mount Olympus. The principal life-zone is the Budsonian.
The Canadian merges with the lowland Humid-Transition and is difficult to
ascertain as a separate zone. In the Hudsonian Life-zone the average tem-
peratures are low. Winter climate is bitter and the snow lies deep. In the

Climate and Vegetation 29

absence of government weather stations in the Olympics, detailed descriptions
of climatic conditions can not be given. Vegetation of the Hudsonian Life-zone
of the Olympic Mountains is, in general, similar to that of the same life-zone
of the Cascades (see Jones, 1936, Botanical Survey of the Olympic Peninsula).

The lowlands of western Washington have a cool, humid climate. The
average annual temperature of the area varies little from 50°. In winter the
temperature, especially in January and February, commonly drops below the
freezing point at night. Summer temperatures are moderate, rarely reaching
90°. Snowfall is light, averaging about 10 inches. The prevailing winds are
from the west and are moisture-laden. They rise over the Olympic Mountains
and loose heavy rains along the coastal area of the lowlands. The average
annual precipitation at La Push. Clallam County, is 97.9 inches; at Clearwater,
Jefferson County, 124.9S inches; at Aberdeen, Grays Harbor County, 81.58
inches; at South Bend. Pacific County, 83.35 inches. In contrast, the more
inland areas receive less than half as much precipitation. The yearly average
at Bellingham is 31.09 inches; at Seattle. 30.07 inches; at Tacoma, 39.53
inches; at Vancouver, Clark County. 37.24 inches.

The outstanding feature of the vegetation of western Washington is the
coniferous forest. Previous to the logging activities a dense cover of Douglas
fir, western hemlock and red cedar spread almost unbroken over the area. The
openings in the forest and the marshy ravines and river valleys supported
growths of underbrush and deciduous trees so dense and luxuriant as to com-
pare with a tropical jungle. In the dense rain forests along the coast, mosses
and lichens develop an understory vegetation many inches deep and clothe
the branches of the forest trees. The mild temperature and excessive rain-
fall cause some species that usually are of bush or shrub size to reach the
proportions of small trees. In some places one can climb twenty feet from
the ground in a huckleberry tree, the trunk of which is five inches in diameter.
The coniferous forest is made up of several species of trees. Most important
among these are the western hemlock (Tsuga heterophylla) , Douglas fir
(Pseudotsuga taxifolia), and red cedar (Thuja plicata). Locally the western
yew (Taxus brevi folia), lodgepole pine (Pinus contorta) and spruce (Picea
sitchensis) may be common. Deciduous trees are numerous and include sev-
eral willows (Salix sp.), aspen (Populus tremuloides) , hazel (Corylus califor-
nica), alder (Alnus oregona), oak (Quercus garryana), broadleaf maple (Acer
macrophyllum) , vine maple (Acer circinatum) , and flowering dogwood (Conius
nuttallii). Mosses and ferns are abundant. The sword fern (Polystichum
munitum) and bracken (Pteridium aquilinum) are especially common. Space
prevents listing all but a fraction of the typical shrubs but these include huckle-
berry (Vaccinium parvifolium, Y actinium ovatnm), Oregon grape (Berberis
nervosa), salal (Gaultheria shallon), rose (Rosa zymnocarpa), thimbleberry
(Rubus jxtrviflorus) , salmonberry (Rubus spectabilis), blackcap raspberry
(Rubus leucodermis) and wild blackberry (Rubus macr ope talus) .

The higher parts of some of the ranges of northeastern Washington are in
the Hudsonian Life-zone, but most of the mountains are in the Canadian Life-
zone. The valleys are in the Transition Life-zone. Climatic conditions are
similar to those of the eastern slopes of the Cascades. Winter temperatures
are low, the average for December, January and February being below freez-
ing. Summer temperatures are high, the July average for Colville being 67.2°

30

University of Kansas Publs., Mus. Nat. Hist.

and the July average maximum being 87.4°. Vegetation consists principally
of coniferous forests in the mountains and deciduous woods in the valleys.
Among the interesting features of the vegetation are the extensive stands of
almost pure larch (Larix occidentalis). In most respects the flora closely
resembles that of the Blue Mountains.

The Blue Mountains of southeastern Washington differ from other ranges
in Washington in their relative aridity. There are few streams and a single
river drains the area. There are no government weather stations in the Blue
Mountains. The winter temperatures are low and the snow deep and lasting.
Summer temperatures are high and humidity and precipitation low. Conif-
erous forests of the type of arid regions form the principal tree cover. Typical
plant species include the white fir (Abies grandis), alpine fir (Abies lasiocarpa),
larch (Lfirix occidentalis), spruce (Picea columbiana) , and such shrubs as fool
huckleberry (Menziesia ferruginea) , Pachystima myrsinites, dogwood (Cornus

Fig. 10. A "pothole" being crowded by drifting sand, ten miles south of
Moses Lake. Washington. March 23. 1940. (Fish and Wildlife Service photo
by Victor B. Scheffer, No. 925.)

canadensis), wild current (Ribes petiolarc). mountain mahogany (Cerco-
carpus ledifolius) . spirea (Spiraea sp.), lupines (Lupinus) of several species,
maple (Acer douglasii), buckbrush (Ceanothus sanguineus), sticky brush
(Ceanothus velutinus), and huckleberry (Vaccinium tin nibranacium) .

The Columbian Plateau and southeastern Washington present desert con-
ditions. At Odessa. 1590 feet, Lincoln County, the average annual precipita-
tion is only 9.38 inches, and only in tin winter may more than one inch of
precipitation per month be expected. The average temperature is 48.5°. In
the winter the average is below freezing but in July it is 71.3°. The average
maximum for July is 90° and an extreme of 111 is recorded. Walla Walla,
991 feet, has a higher annual precipitation (16.66 inches) but higher tempera-
ture (yearly average 53.5 , July average 75.0°. average July maximum 88.6°,
extreme 113°). Winter temperatures on the Columbian Plateau are low. The
January average al Odessa is 25.3° and at Walla Walla 32.4° The average

Climate and Vegetation 31

yearly snowfall at Odessa is 19.4 inches and at Walla Walla 23.5 inches. Vege-
tation of the Columbian Plateau and southeastern Wshington is of the desert
type. A few pines and junipers grow in favored places. Along streams the
cottonwood (Populus hastata) and willow (Salix) of several species are com-
mon. Most typical are grasses and shrubs such as the bunch grass (Agropyron
iaerme, Agropyron spictalum), foxtail (Alopecurus aequalis), cheat grass
(Bromus trctorum), saltbrush (Atriplex tnmcata), greasewood (Sarcobatus
v< rmiculatiis) mustard (Arnbis sp., Brassiea sp.), sagebrush (Artemisia rigida,
Artemisia tridentata) , rabbit brush (Chrysothamnus nauseosus, C hrysothamnus
viscidiflorus) and cactus (Opuntia polyacontha). The arid climate of the
Columbian Plateau affects, to some extent, surrounding areas. Thus the
Yakima Valley Area, the Columbia Valley, where it borders the Plateau, and
the Okanogan Valley possess vegetation typical of the Columbia Plateau.

32

University of Kansas Publs., Mis. Nat. Hist.

LIFE-ZONES AND ECOLOGY

The Transition Life-zone is the principal life-zone in Washington.
It is divisible into three subdivisions: Humid, Arid-timbered
and Arid-grasslands (Fig. 11) subdivisions. The Humid and Arid-
timbered subdivisions of the Transition life-zone are closely related
in some respects but different in others. They are separated by the
Cascade Mountains. All of the Transition Life-zone west of the
Cascades belongs to the Humid subdivision and the timbered Transi-
tion Life-zone east of the Cascades belongs to the Arid-timbered
subdivision.

_ -4»

DISTRIBUTION MAP
STATE OF WASHINGTON

h ->'» p-.jt.t «n-jnf fc,cwj rtir

Fig. 11. Lifp-zones of Washington. Arctic-alpine not shaded. A. Hud-
sonian and Canadian (mapped together). B. Forested Transition (Humid
and Arid subdivisions). C. Arid-grasslands of the Transition. D. Upper
Sonoran.

The Arid-grasslands are of minor geographic extent. Although
this subdivision is relatively distinct as concerns the distribution
of plants, insects and birds, it is of little importance as concerns
the distribution of mammals. For the most part, the mammals

Life-Zones and Ecology

33

occupying it are more representative of surrounding areas. Large
parts of the Arid-grasslands have been taken over for agriculture,
especially wheat raising. Perhaps the greatest extent of the Arid-
grasslands existing in a natural state is along the eastern Cascade
Mountains and along the eastern side of the Columbian Plateau.
These are truly transition areas, situated where the arid pine forests
are replaced by open, sagebrush desert.

Fig. 12. Arctic-alpine Life-zone, Mount Rainier, Washington: Cowlitz
Glacier from elevation of 9,500 feet. (Fish and Wildlife Service photo by Vic-
tor B. Scheffer. No. 900.)

The Upper Sonoran Life-zone includes the desert areas of Wash-
ington (figure 11 D). Its principal extent is the central Columbian
Plateau. From the central Columbian Plateau, fingerlike projec-
tions of desert extend along the principal valleys.

The Arctic-alpine Life-zone occurs in the high Olympic Mountains
and on the higher peaks of the Cascades. This is shown on the life-
zone map, Fig. 11, as white, unshaded areas.

The mammalian faunas of the Hudsonian and Canadian life-
zones resemble each other closely. The boundary between them is
too complex to permit separating them on a small-scale map.
Consequently they are mapped together on Fig. 11 as A.

3—4993

84

University of Kansas Pibls., Mis. Nat. Hist.

The Humid Subdivision of the Transition Life-zone

This subdivision is remarkably uniform in composition over
western Washington. The greatest difference is in precipitation.
Rainfall along the coast is heavier than that in the interior. Con-
sequently vegetation is more dense and luxuriant along the coast.

Four habitats may be distinguished in the Humid subdivision
and further subdivision is possible. The dominant and most ex-
tensive habitat is the forest. Dominant mammalian species include:
Peromyscus maniculatus, Sorex trowbridgii, Sorex obscurus, Tamias-
ciurus douglasii, Clethrionomys californicus, Aplodontia rufa,
( iloiiconi i/s snbrinus and Odocoili us h< mionus. Mammals are scarce
and nocturnal forms prevail. As a rule, a line of mouse traps set in

Fia. 13. Humid subdivision of the Transition Life-zone, Headley's Marsh,
five miles east of Granite Falls. Washington, June 4. 1938. (Fish and Wildlife
Service photo by Victor B. Scheffer, No. 60.)

a foresl habitat will take principally Peromyscus maniculatus with
a few Sorex trowbridgii and Sorex obscurus and rarely a Clc-
thrioHomi/s californicus. In some places, especially where the moss
is deep, a line of mouse traps will catch only shrews.

A second important habitat is the deciduous jungle. This differs
from the forest habitat in that the dominant trees are of the de-
ciduous type and in that understory vegetation, such as shrubs and
annuals, is dense. The jungle habitat occurs in ravines and in val-

Life-Zones and Ecology

35

leys of streams and rivers and, in general, covers the lower, poorly
drained portions of the Humid subdivision of the Transition Life-
zone. Mammals are abundant and varied m the jungle habitat.
The deer mouse [Peromyscus maniculatus) is the most common
mammal but a line of mouse traps might also catch: Neurotrichus
gibbsii, Scapanus orarius, Sorex vagrans, Microtus oregoni, or Zapus
p. trinotatus. The mountain beaver, snowshoe rabbit, and Town-
send chipmunk also occur there.

The prairies form a third habitat. These areas of native grass-
lands are of minor extent but are the principal home of several races
of gophers and the Townsend mole (Scapanus townsendii). Deer
and elk also browse on the prairies.

S- '. <« j

Fig. 14. Timbered arid subdivision of the Transition Life-zone, Kettle
Falls on the Columbia River (now beneath Coulee Dam backwater), Stevens
Countv, Washington, June 15. 1938. (Fish and Wildlife Service photo bv Vic-
tor B.'Scheffer, No. 72.)

A fourth habitat is the marsh. In it there is here included the
sphagnum bogs and marshy shores of lakes and streams. This
habitat is characterized by damp ground, standing water, and dense
vegetation. Typical mammalian species include the water shrew
(Sorex bendirii) , Townsend meadow mouse, muskrat and mink.

Other habitats, such as aerial for the bats and aquatic for the
beaver and otter, might be listed.

The Arid Timbered Subdivision of the Transition Life-zone

In Washington the Arid timbered subdivision of the Transition
Life-zone is the open, pine forest. Because of the aridity of this

36 University of Kansas Pubis., Mrs. Nat. Hist.

habitat, marshes and streamside thickets are uncommon, but where
habitats of this kind do occur they have a fauna distinct from that
of other habitats.

The pine forest habitat includes many diurnal species, such as the
red squirrel, yellow-pine chipmunk, and Columbian ground squirrel.
The white-tailed deer occurs here and, for most of the year, the
mule deer. Snowshoe rabbits are usually present. Near rocks the
bushy-tailed wood rat is common. Mice are scarce, probably be-
cause of the open nature of the surface of the ground. A night's
trapping usually yields only a few Peromyscus manicvlatus.

The mammalian fauna of the marshes and streamside thickets is
similar. Shrews including Sorex vaapans and Sorex obscurus are
uncommon. Meadow mice, including Microtus pennsylvanicus, Mi-
crotia longicaudus, and more rarely Microtus montanus, are taken.

The Arid Grasslands Subdivision of the Transition Life-zone

This subdivision is so much utilized by man where it occupies
any considerable areas, and is of such a transitional nature elsewhere,
that it is important for only a few native wild mammals. The
sagebrush vole (Lagurus curtatus) seems to be confined to the arid
grasslands. The white-tailed jack rabbit is now found principally
in the arid grasslands, but its confinement there has resulted prob-
ably from competition with the black-tailed jack rabbit. The mon-
tane meadow mouse (Microtus montanus) is the only common,
representative species. Many species from the Upper Sonoran Life-
zone extend into the arid grasslands where conditions are suitable.
These include Reithrodontomys megalotis, Perognathus parvus, Ci-
tellus Washington! and Marmota flaviventris. A few species more
typical of the Arid timbered subdivision of the Transition Life-zone
stray onto the arid grasslands. Citellus columbianus and Microtus
longicaudus may be included here.

The Upper Sonoran Life-zone

The sagebrush desert in Washington is relatively uniform in na-
ture. Several different habitats may be distinguished, such as sandy
areas, open sage, dense sage, stony ground, and talus. Qualitatively,
however, the mammalian fauna of these areas is surprisingly similar.
Quantitatively, there are great differences. For example, the grass-
hopper mouse is rare in the open sage areas with hard, claylike soil
but common on drifted sand. The harvest mouse is common in
dense sage but rare in open sage or in open, sandy areas. Mammals
are abundant on the sagebrush desert and typical species include:

Life-Zonks and Ecology 'M

the black-tailed jack rabbit, Nuttall cottontail, Orel kangaroo rat,
Great Basin pocket mouse, Townsend ground squirrel, Washington
ground squirrel.

Marshes are not uncommon on the Columbian Plateau and else-
where in the Upper Sonoran Life-zone in Washington. They do not
possess a fauna that is strictly Upper Sonoran but instead contain
species more typical of the Arid-timbered subdivision of the Transi-
tion Life-zone. Meadow mice found in desert marshes include
Microtus montanus and Microtus pennsylvanicits. The only shrew
we have found is Sorex vagrans. The harvest mouse (Reithro-
dontomi/s megalotis) is often abundant in marshes.

Fig. 15. Upper Sonoran Life-zone, sand and basalt cliffs along the east bank
of the Columbia River, at Vantage, Washington, 1930. (Fish and Wildlife
Service photo by Victor B. Scheffer, No. 848.)

The Canadian Life-zone

The Canadian Life-zone is a relatively uniform area in which
the forest habitat is most important. Talus and swift, cold streams
bring some typical mammalian species into the Canadian Life-zone
from the Huclsonian. Mammals are usually common; they are
abundant only in the upper portion of the Life-zone. Arboreal
species and forms adapted to life beneath the forest cover are domi-
nant. The Douglas squirrel, red squirrel, northern flying squirrel
and Townsend chipmunk are typical arboreal species. Traps set
beneath the trees might catch Peromyscus maniculatus, Clethri-
onomys gapperi, Neotoma cinerea, Sorex obscurus, or Sorcx trow-
bridgii.

'is

University of Kansas Pvbls., Mtjs. Nat. Hist

Fig 16. Canadian Life-zone foresl on Mount Rainier, Washington, eleva-
tion 2,800 feet, September 11, 1934. Western hemlock, Douglas fir, western red
cedar, and grand fir. (Fish and Wildlife Service photo by Victor B. Scheffer,

Life-Zones and Ecology 39

The Hudsonian Life-zone

In Washington the Hudsonian Life-zone is of greater diversity
than any other. Frost and steep slopes have formed great masses
of talus and, in this talus, certain species of mammals, such as
Ochotona princeps, Marmota oaligata and Marmota olympus are
found. Other species, such as the golden-mantled ground squirrels,
mountain chipmunk, bushy-tailed wood rat, red-backed mouse and
long-tailed meadow mouse find the talus an ideal home. Glacial ac-
tion has produced, in the Hudsonian Life-zone, numerous level or
concave areas that contain small lakes and slow-moving streams.
Dense, herbaceous vegetation is abundant nearby. Small mammals
abound and a line of mouse traps will almost certainly catch a few
such typical species as: Sorex palustris, Sorex obscurus, Microtus
oregoni, Microtus richardsoni, Microtus longicaudus and Zapus
princeps as well as the ever present Peromyscus maniculatus. The
shrew-mole or heather vole might also be taken, though the latter
is more apt to be found in nearby heather meadows.

40

University of Kansas Publs., Mis. Nat. Hist.

Table 1. Distribution of mammals in Washington by Life-Zones.
.4. Abundant. C. Common. R. Rare.

Species vnd Subspecies.

'en

X

<

X

1

j. ■-

x ,"*3
~ y.

u -

— r~

<

5

u

c
c
o
■f.

u

<t>

3
■3
c

o

c

DO

"2

Neurotriclms gibbsii gibbsii

?

c

A

A

orarius orarius

A

orarius sr-hefferi

C

A

R

R

R

R

trowbridgii trowbridgii

A

C

trowbridgii dsstructioni

A
A

R

C

c

A

C

R

C

C

A

C

A

A
C

C

R

Myotis lucifugus carissima

c

C

lucifugus alascensis

A

C

vumanensis sociabilis

c

C

A

R

R

R

R

R

C

volans iaterior

C

R

R

ualifomicus caurinus

A

C

subulatus melanorhinus

R

A

A

A

C

' lorynorhinufi rafinesquii townsendii

R

R

Life-Zones and Ecology

41

Table 1. — Continued

Species and Subspecies.

c
o
+-»
"w

•- 03

- —

X

a>.°

<

J5 c
<

oj
M
O
C

o

a
a
P

•5

OS

c

a!

u

c

'3

■a

w

R

A

A

R

R

R

R

R

R

Ursus americanus altifrontalis

A

C

A

A

A

C

R

Procyon lotor psora

A

C

R

R

C

A

C

A

C

R

R

C

C

C

C

R

R

R

C

R

C

C

C

C

f renata effera

A

C

C

C

c

C

A

c

C

C

C

C

C

c

R

R

R

C

R

R

R

C

R

R

gracilis latifrons

A

A

R

R

C

R

mephitis spissigrada

A

C

C

C

R

R

42 University of Kansas Publs., Mus. Nat. Hist.

Table 1. — ( 'ontinm >l

Species a.nd Subspecies

7

2 ~

- -

3 Eh

—
c -
— ~

-= -
1 |

X

j. .~

- -

Is
<

::

3

s

71

:.

ft
ft

u

/.

c

A

A

A

C

R

A

c

C

A

lupus fuscus

R

R

R?

R

C

C

concolor oregonensis

c

C

C

R

A

C

ruf us pallescens

A

c

C

C

C

R

A

A

A

A

A

A

A

A

C

A

C

A

A

C

A

R

R

R

c

A

R

A

C

A

A

c

A

A

c

A

A

lateralis tescorum

C

C

C

A

A

A

A

A

R

A

A

A

c

C

C

Life-Zones and Ecology

4:5

Table 1. — Continued

Species and Subspecies.

c

.2
'■*->

"en

3H
w

-a

.a 5

E 2

GO

-a

J2 3

S3

<

5

c

t-.

c.
a

3
c

03
O

.2
'2

QQ

B

c

A

A

A

A

R

C

c

A

c

A

A

A

A

A

A

A

A

C

R
A

C

C

C

C

A

A

A

C

A

A

A

talpoides tacomensis

A

A

A

talpoides yelrnensis

A

talpoides couchi

A

C

A

Castor canadensis leucodonta

A

A

C

C

canadensis idoneus

A

A

C

A

A

A

A

A

A

A

C

R

A

A

R

R

C

C

44 University of Kansas Publs., Mrs. Nat. Hist.

Table 1. — Continued

Species and Subspecies.

Humid
Transition

.§1

<

BO
J g

U

in

<

c
s

c

DQ

t*

o

a
a

c

n

■6

e
U

5

i
■a

-

c

B

A

c

\

c

A

C

Synaptomys borealis wrangeli

A

Phenacomys intermedius intermedius. .

C

A

intermedius oramontis

C

A

Clethrionomys gapperi saturatus

R

A

A

R

A

\

A

A

A

Mierotus pennsylvanicus funebris

A

R

C

pennsylvanieus kincaidi

A

A

C

C

A

A

A

A

longieaudus halli

A

C

C

longieaudus macrurus

R

C

A

rinhardsoni arvicoloides

C

A

richardsoni maoropus

C

A

A

A

Ondatra zibsthicus osoyoosensis

A

A

C

A

A

princeps kootenayensia

A

A

princeps trinotatus

A

C

A

A

A

Erethizon dorsatum epixanthum

A

R

c

A

R

dorsatum nigrescens

A

R

c

A

R

A

princeps fenisex

A

C

A

Life-Zones and Ecology

45

Table 1. — Concluded

Species and Subspecies.

C

_o

'53

T3

S3 .2

P'itj

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ealifornicus deserticola

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americanus cascadensis

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americanus pineus

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canadensis nelsoni

C

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virginianus oehrourus

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Ovis canadensis canadensis

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canadensis calif orniana

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46 University of Kansas Publs., Mrs. Nat. Hist.

GEOLOGIC HISTORY OF WASHINGTON

The composition of the mammalian fauna of any area is depend-
ent on several factors. These include the composition of the original
fauna, species which have since invaded the area, and quantitative
and qualitative changes that have occurred in the area. The latter
two factors refer to changes in relative numbers or extermination
of species through environmental changes or competition with other
forms and evolutionary changes that have occurred in the species
making up the mammalian fauna.

Our knowledge and understanding of the distribution and history
of the species of mammals occurring in Washington decreases rapidly
as we go back in time. The distribution of the modern fauna at the
present time is fairly well known. The distribution of species 100
years ago is less well understood. This is especially true of certain
game species and carnivores whose distribution has been altered by
man. Our knowledge of the distribution of mammals in the Pleis-
tocene and earlier times is based on fossil skeletons. Such knowl-
edge must necessarily be meager, for conditions favorable to fossil-
ization and the preservation of fossils until their subsequent
discovery by man. were not of common occurrence.

In the Cascades and in eastern Washington, the Miocene was
a time of orogeny and great volcanism. Great Hows of lava, 4.000
feet thick in the Snake River area (Russell, 1893), emerging from
fissure> in the Snake River area, formed the Columbian Plateau.
The Columbian basalt slopes inward eentripetally from the eastern,
northern, and western margins of the Columbian Plateau with an
average descent of 25 feet to the mile (Flint, 1938). The dip of the
lava flows results in the basalt-marginal course of the Spokane
and Columbia rivers today, along the northern edge of the Colum-
bian Plateau. The earlier part of the Pliocene was a period of
erosion and deformation. In the early Pleistocene the five great
volcanic cones of the Cascades, Mount Baker, Glacier Peak, Mount
Rainier, Mount Adams and Mount St. Helens were formed. In
eastern Washington a gentle folding of the Miocene lava flows
occurred. The folding took place slowly and the Columbia River
in it- course along the eastern edge of the Cascades cut through
the folds as they formed, making a series of water gaps. Farther
south, the Simcoe-Frenchman Hills anticline seems to have arisen

Geologic History 47

more rapidly and the Columbia River was forced eastward before
it became impounded and rose over the barrier and plunged down,
tearing out the great Wallula Water Gap (Flint, 1938). This gap is
a mile wide, eight miles long and, in places, a thousand feet
dee]). The impounding of the Columbia by the Simcoe-Frenchman
Hills anticline resulted in a lake several hundred miles in area.
Sediments deposited in this lake form the Ringold formation. The
Ringold formation possesses a very early Pleistocene mammalian
fauna.

The Pleistocene was a time of great change in the mammalian
fauna of the world. Unfortunately the beautiful glacial sequence
revealed in Europe and the Mississippi Valley cannot be detected in
Washington. In western Washington the deposits of the last con-
tinental glaciation and fluvial deposits of the last interglacial period
almost everywhere obscure evidence of earlier glaciations. Deposits
of an earlier glaciation, named Admiralty by Bretz (1913), have
been detected in places. Deposits of greater age, that may rep-
resent a still earlier glaciation, have been noted. In eastern Wash-
ington the only definite proof of multiple glaciation is of one glaci-
ation preceding the last. This is the Spokane glaciation of Bretz
(1923). That multiple glaciation in the sequence reported from the
Mississippi Valley affected Washington seems probable. The lack
of evidence of a complete sequence is negative evidence. In
western Washington the earliest glacial deposits might be beneath
the later deposits or they may have been removed or reworked
by subsequent glaciations, whereas in eastern Washington they
may have been removed by subsequent glaciation and erosion.

The time interval between the two known glaciations appears to
have been of greater duration than the Recent. The drift of the
earlier period is sometimes found covered by the till of the later
glaciation, and preserved by it. The early material is deeply weath-
ered and all save the hardest pebbles and quartzites, for example,
are rotten and disintegrate at the touch. In contrast, the later de-
posits are almost unweathered. Pebbles are hard, and ring when
struck. A zone of leaching and oxidation of the finer materials
reaches a depth of some 30 inches, below which the till is fresh.

Two names are currently applied to the last continental glaciation
of the state of Washington. That west of the Cascade Mountains,
studied and described by Bretz (1913), was termed "Vashon." The
interglacial cycle preceding it was called "Puyallup." The glaciation
of eastern Washington has been called "Wisconsin," after the Mis-

48

University of Kansas Publs., Mus. Nat. Hist.

sissippi Valley terminology, by several writers. Papers by Flint
(1935, 1937) describe and map it.

The Vashon and Wisconsin glaciations probably occupied the
same time interval, although this has not certainly been established.
In the present report I have used the term "Vashon-Wisconsin" in
speaking of the entire period, or the glaciers both east and west of
the Cascades together. Vashon, alone, is restricted to western Wash-
ington and Wisconsin to eastern Washington.

DISTRIBUTION MAP
STATE OF WASHINGTON

MILES

I'n;. 17. Extent of \ a-hon-Wisconsin ice over Washington. The Wisconsin
and Vashon glaciers have been connected through the Mount Rainier section
of the Cascades because it is thought that few or no mammalian species lived
in the Cascades north of Mount Rainier while the ice was in place. Data
generalized from Flint (1937). Bretz (1913), Culver (1936) and other sources.

The Vashon glaciation seems to have consisted of an \w dome
centering in Pnget Sound (the Puget Glacier of Bretz, 1913) and
flooding the lowlands from the Olympic Mountains to the Cascade
Mountains. The southern edge of the Puget Glacier was slightly
south of the present terminus of Pnget Sound. Fingerlike projec-
tions of ice were forced up valleys of the western Cascades and the
northern and eastern Olympics. Some of these upward moving
fingers <>f ice met and coalesced with valley glaciers descending from

the mountains. At the southern edge of the glacier, the Black Hill-

Geologic History 49

and Porcupine Hills remained above the ice although partially sur-
rounded by it.

The Wisconsin glacier, according to Flint (1935), was a great
piedmont glacier, fed by valley glaciers from the Cascades and
Coast Ranges to the west and the Rockies to the east. It extended
from the Idaho boundary to the Cascade Mountains. From the
Canadian Boundary it sloped down to an approximate elevation of
some 6500 feet at Republic and to 2500 feet on the northern edge
of the Columbian Plateau which was the southern edge of the glacier.
The Kettle River Mountains, in almost the center of the glacier,
remained a peninsula or driftless area that divided the glacier into
two lobes. The Pend Oreille, Huckleberry and other mountain
ranges, formed nunataks, or islands above the ice, at the southern
part of the glacier.

The behavior of valley glaciers in the northern Cascade Moun-
tains during Vashon-Wisconsin time, seems to have been variable.
Some depression of the snow line, at least in the north, seems
probable.

The Vashon Glacier impinged on the eastern, northern and to
some extent the western, slopes of the Olympic Mountains. Late
Pleistocene valley glaciers in the Olympics, however, seem to have
been inconsequential.

The time of the retreat of the Vashon-Wisconsin glaciers is a
subject of special interest to the mammalogist in that it represents
time for invasion and dispersal of species and in that it represents
generations of individuals upon which natural selection might act.
It is generally agreed that a period of approximately ten thousand
years has elapsed since the retreat of the Vashon-Wisconsin glaciers
from Washington.

Information on the climate of the state of Washington previous to
the period of the last continental glaciation is understandably
meager. Bretz (1913) considers the Puyallup period a time of ex-
cessive precipitation and erosion. Bits of lignite from Puyallup
sediments seem to be of Douglas fir. Presumably the climate was
slightly warmer and more humid than it is today. Vegetation
possibly consisted of coniferous forests.

With the advance of the Vashon ice, mammals north of the ice
border were all or mostly eliminated. Climatic conditions south
of the border of the ice probably were strongly affected by it. Re-
mains of mammoths have been found in Vashon till. The presence
of many non-boreal species of mammals in southwestern Washing-

4—4993

;,o

University of Kansas Plbls., Mtjs. Nat. Hist.

ton indicates their persistence there and that conditions therefore
were not intolerable for them. Probably the climate of south-
western Washington was cool and dry. Fir, spruce, and Douglas
fir may have been the dominant trees. Hansen (1941 A: 209) found
evidence from studies of pollen that coniferous forests were grow-
ing in west-central Oregon in late glacial time. These pollen studies
of post-glacial peat bogs by Henry P. Hansen give evidence of

i

4.

- y.-. ■

t

Fig. 18. McDowell Lake, Little Pend Oreille Wildlife Refuge, Stevens
County, Washington, September 29, 1939. (Fish and Wildlife Service photo
by Victor H. Scheffer, No. 730.)

post-glacial climatic changes. Hansen points out d!)41 B, 1M41 C)
that climatic changes west of the Cascades were probably slight
because of the influence of the Pacific Ocean. Pollen profiles in-
dicate an early, cool, dry climate followed by a wanner one and
increasing humidity. The present climate' may be considered cool
and humid. Mosf of western Washington lies in the Humid sub-
division of the Transition Life-zone.

The pre-Wisconsin climate of eastern Washington is unknown.

Geologic History 5]

From the loessial nature of the Palouse Soil (Bryan, 1927), a pre-
glacial deposit, the area would seem to have been arid, probably
a grassland or a sagebrush desert. If the fossil fauna discovered at
Washtuckna Lake, Adams County, is of this period, the forest
conditions of the Blue Mountains were slightly more extensive than
at present. The glacial climate of the Columbian Plateau in Wis-
consin time was probably cool and arid. Pollen studies by Hansen
(1939, 1940) indicate increasing dryness and warmth since the
retreat of the Wisconsin ice. Present-day climate on the Columbian
Plateau is warm and dry. The plateau is principally a sagebrush
desert. The glaciated area to the north is cooler and more humid,
supporting extensive forests of yellow pine and other conifers.

52 University of Kansas Publs., Mrs. Nat. Hist.

THE FAUNAS

We have mentioned elsewhere that three different mammalian
faunas occur in Washington. These may be described as follows.

1. The Great Basin Fauna. This fauna is best typified by
such genera as Perognathus and Dipodomys. Species that, at least
in Washington, are confined to this fauna arc:

Sorex merriami Tamms minimus

Myotic thysanodes Perognathus parvus

Myotis subulatus Dipodomys ordii

Pipistrellus hesperus Onychomys leucogash r

Antrozous pallidus R eit hrodontomys megalotis

Taxidea taxus Lagurus curtains

Marmota flaviventris Lepus californicus

( 'it< llus townsendii Sylvilagus nuttallii

Citellus Washington? Sylvilagus idahoensis

The break between the Great Basin Fauna and the other two
faunas is extremely sharp, probably as a consequence of a sharp
break in the flora.

2. The Pacific Coastal Fauna. The mammals of the humid
coastal district range from the Frascr River, British Columbia,
southward to the vicinity of Monterey Bay, California. In Wash-
ington typical genera are Aplodontia, Nevrotrichus and Soa/panus
The following species are typical of the Pacific Coastal Fauna in
Washington:

Nt a ml rich us gibbsii Tamiasciurus douglasii

Scapanus townst ndii Sciurus griseus

Sorex trowbridgii Clethrionomys californicus

Sorex benditii Microtus townsendii

Marmota olympus Microtus oregoni

Tamias townsendii Aplodontia rufa

Some species which range outside this fauna! area have strongly
marked races confined to it. Glaucomys sabrinus oregonensis and
Li pus ami riciituis ivashingtonii are examples. The Pacific Coastal
Fauna is a forest fauna. Like the Great Basin Fauna, it reaches
its northern limit of distribution in Washington and is better repre-
sented farther south. Unlike the Great Basin Fauna, the break
between the Pacific Coastal and the surrounding fauna is not sharp,
because forests continue into the more boreal fauna! areas to the
north ami east. There, some mingling of coastal and Rocky Moun-
tain faunas occurs.

Faunas 53

3. Rocky Mountain Fauna. If this fauna be thought of as in-
cluding mammals of the Rocky Mountains of the United States, and
also those of the subarctic faunal area to the east of these mountains
in Canada, the species in Washington are as follows:

Sorex palustris Synaptomys boreaUs

Microsorex hoyi Phenacomys intermedin*

Lynx canadensis Clethrionomys gapperi

Marmota caligata Microtus richardsoni

Citellus lateralis Ocholo)ia privceps

Citellus columbianus Lepus americanns

Tamias amoenus Rangijer montanus

Tamiasciurus hudsonicus Oreamnos americanns
Glaucomys sabriuus

Of the three, the Great Basin Fauna is the most distinct. Only
about twenty species which occur within the Great Basin Faunal
Area, occur also outside of it in one or both of the two other faunal
areas. Most of these twenty are subspecifically different in the
Great. Basin Faunal Area as contrasted with one or both of the other
areas. Each of the other two areas has no less than 32 species that
are not restricted to it.

54 University of Kansas Publs., Mrs. Nat. Hist.

SPECULATION AS TO EMIGRATIONAL
HISTORY OF THE MAMMALS

The present fauna of the state of Washington was derived in part
from Asia and in part from native forms. Great changes occurred
in early Pleistocene through emigration. By the late Pleistocene
most of the mammals now occurring in the state of Washington were
as they are today. The greatest changes that seem to have oc-
curred in the late Pleistocene are the extinctions of numerous groups,
locally or totally. Among the carnivores, Matthew (1902: 321)
reports remains of the great lion. Fells atrox, associated with such
familiar species as the badger, cougar, lynx and mountain goat. The
great lion was very similar to the modern African lion but was fully
a fourth larger. Associated with the great lion in the California
tar pits are the carnivorous short-faced bears {Tremarctotherium) ,
as large as the Alaskan brown bears; dire wolves [Aenocyon) ,
larger than timber wolves; and saber-tooth tigers (Smilodon) . These
forms were probably also present in Washington in the late Pleis-
tocene. Matthew Hoc. cit.) reports remains of the giant beaver.
Castoroides, from the Silver Lake deposit of Oregon. This greal
beaver, as large as a black bear, was doubtless a residenl of Wash-
ington also. Peccaries, camels, bison, horses and giant ground sloths
have been recorded from Pleistocene deposits of Washington and
nearby areas. Of the elephant tribe, the mastodon and several
species of mammoths were present.

These extinct forms have doubtless exerted some influence on the
past distribution of mammals in Washington and possibly have had
an effect on the distribution of members of the living fauna. One
species of the mammoth, at least, existed in Washington in post-
glacial time. Remains of this form, Elephas columbi, have been
found in Yashon till.

The three faunas of Washington can lie placed in two categories.
One is Sonoran. essentially a desert type, and occupies the Colum-
bian Plateau. The other two are forest, faunas, predominantly
boreal in complexion, and are closely related. Tin 1 Rocky Moun-
tain Fauna is found in the Blue Mountains ami in northeastern
Washington. The Pacific Coastal Fauna is found in western Wash-
ington.

The ice sheet- of Yashon-Wisconsin time descended southward
to southern Pugel Sound and to the northern edge of the Columbian

Emigrational History 55

Plateau. If the area of the ice sheet be superimposed on a map of
distributional areas of Washington, it is seen that the area occupied
by the Rocky Mountain Fauna in northeastern Washington is elim-
inated. Thus, at the maximum descent of Wisconsin ice, the Rocky
Mountain type of mammalian fauna was found only in extreme
southeastern Washington. No point of contact between the forest
fauna of the Rocky Mountains and the fauna of the Pacific coast
exists, anywhere, because desert areas, or at least barren plains, lie
between them from the border of the glaciers south to Mexico. For
the entire period, perhaps thousands of years long, while the glaciers
were in place, the two forest faunas were separated. Repeated sep-
aration of the faunas by successive glaciations is thought to be re-
sponsible for many of the differences now existing between them.

Following the retreat of the ice, the Pacific Coastal Fauna ex-
tended its range northward to the Fraser River and, in part, into
the Cascade Mountains. The Rocky Mountain Fauna invaded
northeastern Washington and boreal Canada, including the Pacific
Coast north of the Fraser River. Certain parts of the Rocky Moun-
tain Fauna also invaded the Cascade Mountains.

Inasmuch as the Cascades were invaded by species from both
faunas, a detailed analysis of the mammals existing there now seems
justified. Several significant features of the composition of the
mammal fauna of the Cascades are apparent. First, several species
typical of the Pacific Coastal Fauna are present, such as Neurotrichus
ffibbsii, Sorex troicbridgii, Sore.v bendirii, Tamias townsendii, Mi-
crotis oregoni and Aplodontia rufa. Each of these species has no
close relatives in the Rocky Mountain Fauna and, save perhaps
Sore.v troicbridgii, occupies a unique ecological niche and has no
counterpart in the Rocky Mountain Fauna.

A second group includes species with close relatives in both the
Rocky Mountain and Pacific Coastal faunas. This group is re-
markable in that it is composed of either very closely related species
or very strongly differentiated subspecies in each fauna. For ex-
ample, the golden-mantled ground squirrel (Citellus saturatus) of
the Cascade Mountains is specifically distinct from Citellus lateralis.
Supposedly the Cascade form was isolated in the southern Cascades
during Vashon-Wisconsin time. The Douglas squirrel (Tamias-
ciurus douglasii) of the Cascades, which has a red belly, is the same
as the squirrel of the lowlands of western Washington but is specifi-
cally distinct from the red squirrel (Tamiasciurus hudsonicus) of
the Rockv Mountain Fauna, which has a white bellv. In the extreme

56 University of Kansas Publs., Mtjs. Nat. Hist.

northeastern Cascades the two species come together. They do not
interbreed but seem to compete, for they do not occur together. The
flying squirrel (Glaucomys sabrinus fvliginosus) of the Cascades is
only slightly differentiated from other races of the Rocky Mountain
Fauna but is much different, as are all Rocky Mountain races, from
the western Washington subspecies {Glaucomys s. oregonensis) . The
red-backed mouse of the Cascades is Clethrionomys gapperi, a
species distinct from Clethrionomys calif ornicus of western Wash-
ington. The jumping mouse of the Cascades is Zapus princeps
trinotatus, the same race that occurs in western Washington. It
is quite distinct from, and has previously been considered a species
separate from, the races of the Rocky Mountain Fauna. The snow-
shoe rabbit of the Cascades is closely related to other races of the
Rocky Mountain Fauna but is distinct from L. a. washingtonii of
western Washington. The pika (Ochotona prina ps I of the Cascades
was apparently isolated in the southern part of the range during
the glaciation. After the retreat of the glaciers it extended its range
northward. Competition between two subspecies has resulted in
parallel distributions due to relative body size. The two races freely
intergrade and the differences between them are not so great as in
the other forms mentioned.

The third group of mammalian species of the Cascades is com-
posed of species typical of the Rocky Mountain Fauna such as:
Marmota caligata, Synaptomys borealis, and Oreamnos americanus.
Each has no ecological counterpart in the Pacific Coastal Fauna.
Each is absent from the Cascades of Oregon.

We interpret the mixture of faunas in the Cascades as follows:
The Vashon-Wisconsin ice sheet was in place for a long period of
time, longer, probably, than the Recent. During this time, forest
mammals of the Pacific Coast were isolated from forest mammals
farther east by glaciers to the north and desert to the east. Changes
took place in both of the separated forest faunas. Certain species.
perhaps, such as the mammoth, became extinct. Other forms were
exterminated then or at an earlier time in one fauna or the other.
If Aplodonlhi, X '<■ urotrichus or Scapanus occurred in the Rocky
.Mountain fauna! area, it lived in an inland area of rigorous climate.
and disappeared there because it was unable to adapt itself to the
cold. In the mild climate caused by proximity of the ocean, mild
even in Vashon-Wisconsin time to judge from evidence yielded by
study of fossil pollens, primitive forms such as moles, the Bendire
sinew, and mountain beaver persisted along the coast, where there

Emigrational History 57

were no boreal conditions. Sonic alpine forms, such as Marmota
olympus, Ochotona princeps brunnescens and Citellus saturatus per-
sisted in the Olympic or Cascade mountains as relic species. On the
whole, however, the glacial divergence resulted in a boreal forest
fauna and a temperate forest fauna.

In addition to change in component species, there were evolu-
tionary changes in the species themselves. In some these were con-
siderable, as shown by the differences between related forms of the
two faunas. In most species, however, evolutionary changes have
resulted in only subspecific differences.

Following the retreat of the glaciers and the establishment of
vegetation on the deglaciated areas, movements of the faunas oc-
curred. The Rocky Mountain Fauna spread northward and west-
ward, to northeastern Washington and, in Canada to the Pacific,
occupying most of the land exposed by the glaciers. The Pacific
Coastal Fauna spread northward only as far as the relatively slight
barrier of the Fraser River. The Cascade Mountains became a
"no-man's land." The pika and golden-mantled ground squirrel
of the southern Cascades spread northward. Boreal Rocky Moun-
tain forms with no ecologic competitors from the Pacific Coastal
Fauna occupied the Cascades. Also, coastal species with no Rocky
Mountain competitors occupied the Cascades. Nevertheless, some
competition between members of the two faunas ultimately oc-
curred, and in instances where closely related forms occurred in
the two faunas, one or the other prevailed in the Cascade Range.
For example, the Douglas squirrel and big jumping mouse are now
established in that range, but the relative of each occurring in
the Rocky Mountains is present in the extreme northeastern Cas-
cades. It is possible that in these two cases, the related form oc-
curring in the Rocky Mountains has just entered the area and that
competition has just begun. With regard to the flying squirrel,
red-backed mouse and snowshoe rabbit, the more boreal Rocky
Mountain representatives have definitely displaced the coastal
forms.

Certain mass movements of mammals are popularly believed to
have occurred with the advance of the ice sheets of the Pleistocene.
The boreal birds and plants on higher peaks of the Cascades and
the Sierra Nevada of California are thought to represent relics of
faunas that moved northward. Such mass movements probably
did occur and there is some evidence of their occurrence in Wash-
ington. Probably the pre-W 7 isconsin flora of coastal British Colum-

58 University of Kansas Publs., Mis. Nat. Hist.

bia consisted of coniferous forest similar to that of western Wash-
ington today. If this were the case, the mammalian species in
British Columbia corresponded closely to those of western Wash-
ington. An influx of such a fauna into coastal Washington would
scarcely be evident today if, indeed, it was noticeable even then. In
eastern Washington, forest species forced southward would conic
upon the barren, inhospitable plains and deserts of the Columbian
Plateau.

The greater part of the southward moving forms found refuge
in the Cascade Mountains where, for mosf of Wisconsin time, they
were isolated in the southern Cascades. Examples are Sorc.v palus-
tris, Martes caurina, Martes pennanti, (lulu luscus, Vulpes fvlva,
Lynx cana r ensis, Tamias amoenus, Thomomys talpoides {douglasii
group). Phenacomys intermedius, Microtus richardsoni and Ocho-
tona princeps. In each of these species little or no subspecific varia-
tion has occurred between the populations in the Cascades of Wash-
ington and the Cascades of Oregon.

While the ice sheet existed in Washington there may have been
relatively little movement of the mammalian fauna. There is defi-
nite evidence of a brief contact between the Rocky Mountain Fauna
of the Blue Mountains and the fauna of the southern Cascades. For
example, the pocket gopher of southeastern Washington {Thomomys
talpoides aequalidens) is most closely related to the gopher of the
Simcoe Anticline, and the long-tailed meadow mouse {Microtus
longicaudus halli) of the Blue Mountains closely resembles the
meadow mouse of the Yakima Valley.

Mammals of the Blue Mountains and those of the southern Cas-
cades may have come into contact on the Simcoe-Horseheaven Hills
Anticline, which now stretches 150 miles from the Cascades to the
Wallula Water Gap. Excepting the easternmost 40 miles, it is
timbered. East of the Columbia, a continuation of the anticline
and other hills reaches to the Blue Mountains. Supposedly, in Wis-
consin Time, this anticline possessed a more humid climate and
the habitat was essentially the same as that of an alpine meadow
today. The forms on the two ends of the anticline that are closely
related inhabit humid, meadow habitat. In an earlier paper, Dal-
quesl and Scheffer tl!»44: 316) named this connection the Simcoe
Bridge. Its existence was so strongly indicated by the distribution
of pocket gophers in Washington that we supposed that the study
df many other species would show that they crossed this bridge.
However, study of additional species show- that for them the Sim-

Emigrational History

59

coe Bridge was of only slight importance; there appears to have
been but little mingling of the fauna of the Blue Mountains and
the Cascades by way of the bridge. The Columbia River probably
acted as an effective harrier to many forms that might otherwise
have utilized it. The forms that did cross on this bridge are specie-
known to he active in winter and to emigrate over considerable
areas through tunnels under the snow (Davis, 1939: 257). The
pocket gopher and long-tailed meadow mouse may have crossed
the Columbia, under a cover of snow, when the river was frozen
over. The Columbia has frozen over at the Wallula Water Gap in
historic times.

Fig. 19. Pend Oreille River (or Clark Fork of the Columbia) from a point
near Newport, Washington, looking south. June 13. 193S. (Fish and Wildlife
Service photo by Victor B. Scheffer, No. 67.)

Great movements of mammal species came after the retreat of
the Vashon-Wisconsin ice. The greatest of these was the spread
of the Rocky Mountain Fauna northward and eastward to the
Pacific. In this process, northeastern Washington was inhabited
by animals that probably came from Idaho and Montana. Some
of the species from farther north, as for example the caribou, may
first have been forced into Idaho and Montana by the glaciers. The
invasion of northeastern Washington probably was not a mass
movement of an entire fauna, because invasion seems to be still
going on. As yet the woodchuck and striped ground squirrel of the
Pend Oreille Mountains have not crossed the Columbia River, a

60 University of Kansas Pibls., Mus. Nat. Hist.

relatively minor barrier in northeastern Washington. The moun-
tains west of the Columbia are occupied instead by the yellow-
bellied marmot, a member of the Great Basin Fauna, and there is
no ecologic counterpart in these mountains of the golden-mantled
ground squirrel.

Also the invasion of the Cascade Mountains by a number of
Rocky Mountain species may have been an intermittant or gradual
movement. The red squirrel and Rocky Mountain subspecies of
jumping mouse now are present in the extreme northeastern Cas-
cades, where they possibly arrived relatively recently. The squirrel
is competing with the coastal species already present and may
eventually supplant it. The same may be true of the two forms
of jumping mouse.

The invasion by the Rocky Mountain Fauna Mas rapid as com-
pared with that of the Pacific Coastal Fauna. Evidence of this
was presented in an earlier paper (Dalquest and Scheffer, 1944:
310), where it was shown that the gophers of the douglasii group,
isolated during the Vashon time in the southern Cascades, made
only a few feeble post-glacial movements and then only when con-
ditions were ideal. In this same time the fuscus group of gophers
moved from Idaho and virtually surrounded the range of the
douglasii group. This tendency to immobility seems to have been
characteristic of every member of the Pacific Coastal Fauna. If
the retreat of the Vashon and Wisconsin glaciers occurred at the
same time, both faunas had an equal opportunity to invade the de-
glaciated area. Yet, the Pacific Coastal Fauna actually moved
northward along the coast only to the Fraser River area, and slightly
farther in the Cascades.

Possibly the two glaciers retreated at different times. The \Yis-
consin Glacier was a piedmont glacier. \rd by valley glaciers to
the west and east, and may have disappeared when the feeder
glaciers dried up. The Vashon Glacier was instead an ice cap.
supposedly self-supporting much in the same manner as is the
Greenland Ice Cap. and may have persisted longer than the Wis-
consin glacier. If it did persist longer it formed a barrier to the
northward emigration of coastal species of mammals.

Possibly, also, the whole of the Pacific Coastal Fauna possessed
an inherent sluggishness resulting from their long residence in the
uniform climate and habitat of the Pacific Coast, Certainly the
species -how today great habitat specialization as compared with
species of the Rocky Mountain Fauna. Also, there are fewer
individual mammals per unit, of area in western Washington than

Emigrational History 61

in northeastern Washington. The persistence of the Vashon Glacier,
an inherent lack of incentive to emigrate, or retention of a favorable
environment, may account for the relatively small area invaded by
the Pacific Coastal Fauna.

The Olympic Mountains, on the Olympic Peninsula, rise above
the timberline and are surrounded by forested lowlands which in a
sense isolates this mountain range. Early workers, notably Elliot,
obtained specimens of mammals from the Olympics and described
numerous races, principally, it appears, on the supposition that be-
cause the range was somewhat isolated it should possess a unique
fauna. Subsequent revisions of groups of mammals have indicated
that most of the names proposed, on the basis of specimens from the
Olympics, were either invalid or pertained to mammals found also
in the Cascades.

The mammals of the Olympic Peninsula appear to be divisible
into three groups. A majority of them fall within the first group,
namely coastal races possessing wide ranges in the lowlands of
western Washington. The second group consists of species of the
Rocky Mountain Fauna but with close relatives in the Cascades.
The third group includes but two forms, both unique and found only
on the Olympic Peninsula.

The first group includes nonalpine forms of the lowlands sur-
rounding the Olympic Mountains. For the most part these are
identical with races of the Puget Sound area. A few are slightly
differentiated from the mammals of the Puget Sound area but are
the same as mammals from southwestern Washington. As will be
shown later, some differentiation in the Pacific Coastal Fauna has
occurred. This is thought to be evolution in situ, rather than the
result of mass movements. Many nonalpine Coastal mammals
occur in alpine habitat in the Olympics.

The second group consists of species of the Rocky Mountain
Fauna. Their relationship to the mammals of the Cascades is indi-
cated in the two parallel columns below.

Olympics Cascades

Sorex palustris navigator Sorex palustris navigator

Martes caurina caurina Martes caurina caurina

Martes pennanti Martes pennanti

Tamias amoenics caurinus Tamias amoenus ludibundus

Phenacomys intermedins oratnontis Phenacomys intermedium oramontis

Clethrionomys gapperi nivarius CIcthrionomys gapped saturatus

Only two of these are racially distinct from their relatives in the
Cascades. Of these the chipmunk is a plastic species and breaks
down into many races in Washington. The chipmunks of the

62 University of Kansas Publs., Mrs. Nat. Hist.

Olympics and of Mt. Rainier arc so similar that Howell (1929: 77)
considered them as identical and mapped Mount Rainier as an
isolated part of the range of the Olympic form (see account of T. a.
caurinus). The relationship of the red-backed mice, also, is close,
but has been obscured by the usual assumption of relationship be-
tween californicus (occidentalis) and gapperi. The principal dif-
ference between the alpine forms is the pallor of nivarius. This
pallor of mammals in general from the Olympic Mountains is note-
worthy, but in the red-backed mouse is exceptionally noticeable.
This pallor is discussed beyond in the paragraphs dealing with dif-
ferentiation. Mention should be made here of Myotis fce< nil. This
is a species which seems to have extended its range to Washington
from the north. The power of flight, of course, removes it from
consideration in attempting to reconstruct routes followed by ter-
restrial mammals.

The route of the pocket gopher (Thomomys) in emigrating from
the Cascades to the Olympics (Dalquest and Scheffer, 1944: 310),
was over the out wash train of the Mount Rainier Glaciers, especially
the Nisqualli Glacier, to the extensive outwash aprons of the Vashon
Glacier around southern Puget Sound, and thence into the Olympic
Mountains. Under the conditions in early post-glacial time this
invasion route, hereinafter termed the Puget Bridge, around the
Pleistocene Lake Russell (present Puget Sound), is thought to have
been mainly an alpine meadow. Indeed, the isolated prairies re-
maining today are the unforested remnants of the outwash aprons
(see Dalquest and Scheffer, P>42: (if) I and possess several species
of alpine plants, notably the shooting star, camas. and bear grass.

[f the Vashon Glacier remained in place considerably longer than
the Wisconsin Glacier, these Rocky Mountain species may have
invaded the Cascades from northeastern Washington and travelled
around the southern edge of the Puget Glacier or of Lake Russell.
The close relationship of the races involved, however, suggests that
the emigration took place much more recently. The barriers to
such movement even today are slight, consisting principally of nar-
row areas of forest. For the water shrew, an almost continuous
water habitat still exists, by way of the Nisqualli River, streams
in the Puget Sound area, and the Satsop River in the Olympics.
Tree-living forms such as the fisher and marten might easily travel
the intervening distance today, and. by going along the forests north
of the Chehalis River, reach the Olympics without crossing more
than small streams and virtually without descending to the ground.

Emigrational History 63

Chipmunks and mice probably utilized the prairie or meadow area
of the Puget Bridge, as did the gophers.

Considering the long existence of the Puget Bridge, it is sur-
prising that such forms as the pika, water rat and golden-mantled
ground squirrel did not cross to the Olympics. These forms are.
however, species of the higher or eastern slopes of the Cascades.

The third group of Olympic mammals includes the white-bellied
water shrew and the Olympic marmot, both indigenous forms.

The Bendire water shrew, Sorex bendirii albiventer, is not re-
stricted to alpine habitat but occurs throughout the Olympic Pen-
insula. Its nearest relative is S. b. bendirii of the rest of western
Washington. »S. b. albiventer differs from bendirii only in possess-
ing a partially white ventral surface. We can only conclude that
the white belly of albiventer is a mutation that the local environ-
ment has favored and that the characters have, therefore, spread
through the population on the Olympic Peninsula. Occasional speci-
mens are taken with dark bellies characteristic of bendirii (Jack-
son, 1928: 199).

The Olympic marmot, Marmota oh/mpus, specifically distinct,
and apparently the only preglacial relic species of alpine mammal
in the Olympics, is most nearly related to Marmota vancouverensis
of the unglaciated mountains of Vancouver Island, British Columbia.
Both olympus and vancouverensis are close relatives of Marmota
caligata which ranges southward into the Cascades of Washington.

The Columbia River in its course westward through the Cascade
Mountains, might be expected to act as a highway for the movement
of mammals, but the extent to which it has done so seems to be
slight, at least in post-glacial time. The pocket gopher of south-
western Washington reached the area about Vancouver from the
southern Cascades by way of meadows on the gravel terraces of
Wisconsin glacial drift. No other mammal seems to have extended
this far. Several Great Basin species, such as the cottontail, extend
westward in the valley of the Columbia to the vicinity of Bingen.

The mammals of western Oregon and southwestern Washington
are closely similar as are the plants and climate, despite the fact
that the broad Columbia River courses through the area and did
so all through Recent and Pleistocene times. Many species would
be expected to have crossed this barrier by swimming and rafting,
and that they did so is indicated by the large number of mammals
which are identical or very closely related on the two sides of the
river.

64 University of Kansas Pibls., Mrs. Nat. Hist.

Mammals which seem not to differ on the two sides of the river
include:

Scapanu-s toumsemlii Citellus l>< > cht yi

Scapanus orarius Tamias townsendii

Sorex trowbridgii Sciurus griseus

Sorex vagrans Glaucomys sabrinus

Canis lupus ( 'astor canacU nsis

Felis concolor Microtus lownst ndii

Lynx rufus Microtus oregoni

Mephitis mephitis Ondatra zibethicus

S/iilogale gracilis Zapus prina ps

Procyon lot or OdocoiU us In mionus
Ursus amcricanus

The following mammals are subspecifically distinct in western
Washington and western Oregon:

Washington Oregon

Sorex bendirii bendirii Sorex bendirii pal men

Sorex obscurus setosus Sorex obscurus bairdi

Neotoma cinerea oceidentalis Neotoma citu rea fusca

Peromyscus maniculatus austerus Peromyscus maniculatus rubidus

Clethrionomys californicus < '/< thrionomys californicus

occidi n talis calijornicus

Microtus longicaudus ma'crurus Microtus longicaudus abditus

Aplodontia raja rufa Aplodontia ruja jxicifica

The following species are found in western Oregon but do not
occur in western Washington:

Vulpes fulva Pht nacomys albipes

Urocyon cinereoargenteus Microtus canicaudus

Neotoma fuscipes Thomomys bulbivorus

Phenacomys silvicola Lepus californicus

Phenacomys longicaudus Sylvilagus bachmani

Several of these mammals which occur south of the river but not
north of it are common on the south bank, a few miles from favor-
able but uninhabited territory on the north.

Seemingly the pre-Yashon faunas of western Oregon and Wash-
ington were similar. Some species became extinct in Washington in
the course of Vashon isolation. Others persisted. The very close
relationship of the mammals of the first group indicates some cross-
ing of the river. The best known of such crossings was that of the
Beechey ground squirrel which, previous to 1915, was unknown in

Washington. In 1915, when there was no man-made bridge at
White Salmon, it crossed the river and since has spread over an
area of at least- 50 square miles. The distribution of the mountain
beavers is unusual in that the form in the lowlands of Washington
is indistinguishable from the subspecies in the Cascades of Oregon.

Emigration al History

65

The mammals that are racially distinct on the two sides of the
Columbia River merit careful scrutiny. The Peromyscus of the
two sides more closely resemble one another than those of southern
Oregon resemble those of northern Oregon or than those of southern
Washington resemble those of northern Washington. For Pero-
myscus maniculatus, the Columbia River is simply a convenient
boundary for the separation of two slightly different races. The
Oregon race of the bushy-tailed wood rat is a coastal type but the
Washington form is the same as that of eastern Washington. Seem-

Fig. 20. Rocky bluff along north bank of the Columbia River near Lyle,
Wadiington. March 20, 1939. Habitat of Beechey ground squirrel and yellow-
bellied marmot. (Fish and Wildlife Service photo bv Victor B. Scheffer.
No. 640.)

ingly the more eastern race spread to an unoccupied habitat in
western Washington. Other races that differ on the two sides of
the Columbia probably developed while separated by the river.

The San Juan Islands now possess a limited mammalian fauna.
Unfortunately the activities of man have somewhat changed the
native populations, especially by the introduction of the domestic
rabbit which is now a serious pest in the islands. The Douglas
squirrel, present on Blakeley Island, is said to have been introduced

5—4993

66 University of Kansas Plbls., Mrs. Nat. Hist.

and one resident claims to have first brought it to the island. Two
different persons claim credit for introducing the Townsend chip-
munk on Ocas Island but do not account for its presence on Lopez
Island. The three mammals most abundant and widely distributed
in the islands are Sorex vagrans, Peromyscus maniculatus and Mi-
crotias townsendii. These species, at least, probably reached the
islands at an early time. The two last named are now subspecific-
ally distinct from their mainland relatives. Other mammals which
probably were established before the arrival of the white man in-
clude the mink, otter, beaver, muskrat, raccoon and black-tailed
deer.

The Great Basin Fauna of eastern Washington exists as three
units, one on the Columbian Plateau, another in southeastern Wash-
ington and the third in the Yakima Valley area. The desert species
of the Yakima Valley are more closely related to the species of
eastern Oregon than they are to those of the Columbian Plateau.
In a number of respects the Columbian Plateau gives indications of
age. The ground squirrel, Citellus irashingtoni, is related to, but
specifically distinct from Citellus to\rvs< ndii of the Yakima Valley
and eastern Oregon. Perognathus parvus lordi is a well-marked race,
as is Microtus pennsylvanicus kincaidi and Thomomys talpoides
devexus. We suppose that these species were present on the Colum-
bian Plateau at least through the Recent and probably through all of
Wisconsin Time. The loess deposits of eastern Washington seem
to have been laid down in Wisconsin and Recent times. These
indicate an arid climate which, although probably cool, was prob-
ably not so cold as to exterminate these species. On the other hand,
some species that are now abundant on the Columbian Plateau seem
io have arrived there relatively recently. The black-tailed jack
rabbit, for example, was unknown in eastern Washington before
1870 when it appeared in Walla Walla County. In 1005 it crossed
the Snake River on ice and invaded the Columbian Plateau where
it rapidly spread over the whole area. In January, 1920, it crossed
the Columbia in two places and spread over the Yakima Valley.

The known facts of this movement were sufficiently impressive
to cause the author to study rather closely the distribution of mam-
mals in this area. The collection of bones from a cave along the
Columbia River near Vantage, Grant County, on the Columbian
Plateau, is especially helpful in this respect. This cave was first
visited in 1!>.'5S. It had been the habitat of owls. bats, and primitive
man. The floor of the cave was buried under from one to three

Emigrational History 67

feet of bat guano, much of which had been hauled away for ferti-
lizer. Here and there we found traces of fire and occasional piles
of mussel shells. Some arrowheads and one beautiful obsidian
spear head were found, all buried in guano and about midway be-
tween the floor and the top of the deposit. Remains of mammals
were abundant through the bat guano, and apparently had been
brought to the cave both by man and owls. The jaw of a mountain
sheep was found. This species was known to be present when
the first settlers reached the area (Cowan, 1940: 558). The remains
of smaller mammals included gopher, pocket mouse, muskrat,
meadow mouse, deer mouse, coyote and white-tailed jack rabbit.
No remains of cottontail, black-tailed jack rabbit or harvest mouse
were found. The absence of the cottontail was especially surpris-
ing, in that fully thirty skulls of white-tailed jack rabbits were
noted. The grasshopper mouse (Onychomys) was also absent,
but this species is not common. The two rabbits and the harvest
mouse, however, are abundant in the area today. The cottontail
and harvest mouse have only recently been recorded from the
Okanogan Valley of British Columbia (Cowan and Hatter, 1940: 9).
The black-tailed jack rabbit has never been taken there.

Apparently then, some species have only recently entered the
Upper Sonoran Life-zone of eastern Washington. They have, of
course, reached the state from Oregon. The first step in the inva-
sion probably was the occupation of southeastern Washington. No
barrier prevents mammals from reaching southeastern Washington
from eastern Oregon but the Columbia to the north and west pre-
vents them from occupying the Yakima Valley, and the Snake River
prevents them from reaching the Columbian Plateau. The kangaroo
rat. Great Basin striped skunk and Great Basin spotted skunk now
are at this stage of invasion. The second stage was the crossing
of the Columbia River to the Yakima Valley. This has been ac-
complished by the black-tailed jack rabbit and, earlier, by the
pocket mouse, Perognathus parvus parvus, and ground squirrel,
Citellus townsendii. The third stage was the crossing of the Snake
River and occupation of the Columbian Plateau. The final stage
is the crossing of the northern Columbia River and occupation of
the Okanogan Valley.

68 University of Kansas Publs., Mus. Nat. Hist.

SPECULATION AS TO THE LATER DISTRIBU-
TIONAL HISTORY OF THE MAMMALS

Whereas it is probable that a few of the species now occurring in

Washington evolved there, most are immigrants from other areas.
The success of a given species in any area is dependent on numerous
factors which may be classified under food, shelter from the ele-
ments, protection from enemies and safe breeding places. The fac-
tors may be of an inorganic^ nature, such as climate, soil and ex-
posure or they may be organic, such as vegetation, competition for
food and response to enemies. Abundance results in population
pressure and a tendency for the range of a species to expand.

Mammalian populations are dynamic and change in accordance
with alterations in environment. Because the later geologic history
of the state of Washington was violent, with resultant changes in
climate and geography, the mammalian populations and the distri-
bution of the species have changed much. With changes in en-
vironment, rare species may become common; common and wide-
spread species may become rare or extinct; species foreign to the
area may enter, become established and affect the distribution of
other forms.

Subspecies are groups of individuals with similar genetic compo-
nents or are groups of microgeographic races. In instances where
the phenotypic expression of these similar genetic factors, or the
"characters," are, as a unit, uniformly different from those of ani-
mals of the same species in another geographic area, it is convenient
to give recognition to the two kinds by separate subspecific name.
Intergradation between two geographically adjacent subspecies oc-
curs, directly or where impassable barriers separate them, indirectly
by way of one or more other subspecies. Subspecies of mammals
are geographic races, which means that to warrant recognition by
subspecific name, there must be a logical geographic range in addi-
tion to morphological characters.

Timofeeff-Ressovsky (1JK32, 1940) advances the theory of har-
moniously stabilized gene-complexes to account for the persistence
of subspecies. The persistence of subspecies as genetic units has
been besl explained, I feel, by Sumner (1932: 84-86) who theorizes
as follows:

1. The number of young produced by ;i subspecies is greater than th(
carrying capacity of the land they occupy, al least at certain times or in sonn-
years.

Distributional History (i!»

2. Population pressure results, with a tendency of individuals to emigrate
<iut wards, to the border of the range of the subspecies, where the population
pressure is less.

3. The outward moving tendency keeps the center of the range of the sub-
species genetically "pure."

4. The peripheral wave continues, as long as favorable habitat is en-
countered, until an oppositely directed wave of another race is encountered.

5. Areas of intergradation represent local mingling of genetic factors and
do not affect the ''pure" individuals of the central part of the range of the

subspecies.

Certain aspects of this hypothesis are strongly supported by the
distribution of mammals in Washington. Witness the rapid in-
vasion of Citellus beecheyi and Lepus calijornicus in Washington,
and the eastern cottontail in western Washington and the domestic
rabbit in the San Juan Islands. The volume of the "wave of popula-
tion pressure" where no opposing force is met, is scarcely believable.
In seven years the eastern cottontails released in southwestern
Washington multiplied from a maximum of 12 individuals to a
minimum of 40,000.

Competition between subspecies where their ranges come into
contact seems to be exceptional. Peromyscus maniculatus oreas
and P. m. austerus seem to afford an example of this. However,
in a few cases subspecies seem to be determined in part by adapta-
tion to restricted environments ; each race lives only w r here local con-
ditions favor its respective adaptations.

In the pocket gophers, where restricted habitat and fossorial habits
cause numerous microgeographic races, these microgeographic races
may be potential subspecies. This is especially true in the Puget
Sound area, where six races occur in a small area. These races
meet all the requirements of subspecies and are recognized as such.
It should be pointed out, however, that these races and probably
many other races produced by isolation, may represent degenerative
mutations of the type mentioned by Wright (in Huxley, 1940).
The principal differences of such races seem to have resulted from
the loss of factors of original multiple factor series, with resultant
homogeneity of the race. Inherent variability is another thing
that has to be taken into account when considering the differentia-
tion of the mammals of Washington into subspecies. The pocket
gopher is an extremely plastic species, especially in Washington,
whereas the Douglas squirrel is less so. The flying squirrels, the
yellow-pine chipmunk and the snowshoe rabbit are the other plastic
species. These species are not so likely to break up into numerous
subspecies over all of their ranges as they are in Washington where

70 University of Kansas Publs., Mis. Nat. Hist.

in a small area the topography is highly varied. The range of the
one subspecies, Tamias amoenus a?noenus, to the southeast of Wash-
ington is larger than the combined ranges of all six races occurring
in Washington but, so far as I can see. the topography and environ-
ment are no more varied in Washington than in the mentioned
area to the southeast of it. The range of one subspecies, Lepus
americanus americanus, in Canada is several times larger than the
entire state of Washington, in which four races are found.

The shrews are poor subjects for a study of differentiation, princi-
pally because their small size makes it difficult to see morphological
variations that may be present. The difficulty is increased because
cranial sutures become ossified at an early age. Although it is
difficult to evaluate the differentiation in them, there is some. The
bats, especially the Myotis, are less restricted by geographic barrier-
than are terrestial mammals. Nevertheless, obvious differentiation
exists. The larger predatory mammals and the artiodactyls are
able to move over large areas, at least in the breeding season, but
in these animals also, some differentiation has occurred.

The greatest changes, other than the extinctions, to occur in the
mammalian fauna of Washington since the late Pleistocene, are
changes in distribution. The interglacial cycle preceding the Vashon-
AYisconsin glaciation was of far greater duration than the Recent.
Presumably the mammalian fauna had, from a distributional stand-
point, reached a relatively stable condition. The descent of the
Yashon-Wisconsin ice destroyed the stability and set parts of the
fauna in motion. Probably no stability was reached before the
ice began to recede, and when it did so the previous movements of
the various species were, at least in part, reversed. Stability has
not yet been reached by the mammalian fauna of Washington.
Great changes have occurred in historic times and other changes
probably are under way at present.

In the following pages an attempt has been made to interpret the
probable late Pleistocene and Recent distributional history of the
species of mammals occurring in Washington. The interpretations
are made in the light of what is known of the physical history of the
state and are to be accepted as such rather than as evidence for
the conclusions made concerning the physical history of the state
of Washington and adjacent areas.

Scapanus townsendii. — Probably this animal was confined to
the Humid Transition Life-zone of the Pacific Coast since the
Pliocene.

Distributional History 71

Scapanus orarius. — This species probably had a history similar
lo that of townsendii up to the late Pleistocene. It seems slightly
more adaptable than townsendii, and to be able to extend higher into
the mountains. The distribution of the subspecies orarius is almost
exactly that of S. townsendii. In Oregon, orarius extended eastward
over the Cascades where the subspecies schefferi developed. Perhaps
this subspecies developed since the Pleistocene and since that time
extended along the Columbia River Valley to southeastern Wash-
ington. The race yakimensis, in the Yakima Valley area, is closely
related to schefferi, and seemingly could have been developed from
a stock of schefferi that migrated westward across the Simcoe Bridge.

Neurotrichus gibbsii. — The history of Neurotrichus in North
America was probably similar to that of the two species of Sca-
panus. It tolerates environmental differences to about the same
degree that Scapanus orarius does but occurs much farther south
(Monterey County, California) than S. orarius. This may be be-
cause Neurotrichus has no counterpart to compete with it in the
south, whereas Scapanus orarius must compete in northern Cali-
fornia with the morphologically similar Scapanus latimanus. S.
orarius stops short at this place and S. latimanus occupies all the
territory to the south.

The shrew-mole of the lowland of Washington (N. g. minor)
probably became distinct from the mountain subspecies (gibbsii) in
Vashon-Wisconsin time.

Sorex cinereus. — It is reasonable to suppose that the cinereous
shrew had a continuous range across the forested area of British
Columbia in pre-Wisconsin time. Without having been isolated,
the dark coastal race (streatori) may have developed from the
wider-ranging inland cinereus, as a response to the denser, humid,
coastal forest-habitat, after having been forced southward to Wash-
ington by the Vashon Glaciation. Since that time it is presumed to
have reoccupied the coast of British Columbia and southern Alaska.
This coastal race might have developed in Vashon time, while iso-
lated in southwestern Washington. The Cascades are populated by
a race of the Rocky Mountain Fauna, S. c. cinereus, which probably
entered the Cascades from northeastern Washington or British Co-
lumbia in Recent Time. The absence of the species in western
Oregon, its rarity in western Washington, and its abundance farther
north suggest a northern origin and northward rather than south-
ward postglacial movement. Had the full species cinereus been a

72 University of Kansas Publs., Mus. Nat. Hist.

preglacial resident of western Washington we would expect strea-
tori or a race related to it to occur in the Cascades.

Sorex merriami. — The periphery of the range of this member
of the Great Basin Fauna may have been in southeastern Washing-
ton since pre -Wisconsin time.

Sorex trowbridgii. — This shrew is a typical Pacific coastal spe-
cies with an extensive range along the Pacific Coast south of Wash-
ington. The Washington population may have been isolated in
southwestern Washington during Vashon time or may have crossed
the Columbia into Washington from western Oregon early in the
Recent. Since the retreat of the ice it has extended northward to
southern British Columbia and eastward to the eastern side of the
Cascades. Save for crossing the Cascades its postglacial move-
ments have been slight, as is typical of Pacific Coastal species. The
race destruction! probably has been isolated on Destruction Island
for several thousand years.

Sorex vagrans. — This species probably has had a continuous
range over the western United States since the late Pleistocene.
The dark coastal race (vagrans) probably was differentiated from
the paler races of the Great Basin in response to the more humid
climate along the coast.

Sorex obsctjrus. — The history of this shrew of alpine predilection
probably corresponded closely to that of Sorex cinereus. The de-
rivation of the dark, long-tailed, coastal race {S. o. setosus) from the
smaller, paler, inland race (obscurus\ probably occurred before
A'ashon-W T isconsin Time. Sorex o. setosus is one of a complex of
laces distributed along the Pacific Coast from Alaska to California.

Sorex palustris. — This species has a wide range in North Amer-
ica and extends southward in the Cascade-Sierra Nevada Chain to
southern California. Its extensive range at present in this mountain
chain suggests that it was resident in the Cascades previous to
Wisconsin time. Mountain water shrews probably reached the
Olympic Mountains from the Cascades by way of the Pugel
Bridge in early Recent Time.

Sorex bendirii. — This Pacific Coastal species probably had a
history very similar to that of Ni wotrichus and Scapanus orarius.

The difference between the Bendire water shrews of western
Washington and western Oregon indicates that the Washington pop-
ulation was separated from the sinews of western Oregon during
Vashon Time. The white-bellied race of the Olympic Peninsula
is probably of local origin.

Distributional History 73

Microsorex hoyi. — The Washington record of this shrew at
Loon Lake, Stevens County, is in an area where mammals typical
of the Rocky Mountain Fauna occur.

Myotis lucifugus. — The dark race of this species (alascensis)
may have persisted through the glacial period in southwestern
Washington. The race carvssima, of the Great Basin Fauna, may
have entered the state since the glacial period, from the south, of
course. Habitat selection determines their range at present.

Myotis yumanensis. — The dark, coastal race (saturatus) seems
to be an established member of the Pacific Coastal Fauna. Unlike
lucifugus, the coastal race is not found east of the Cascades. The
race sociabilis, of the Great Basin, has doubtless entered the desert
of eastern Washington from eastern Oregon.

Myotis keenii. — The southernmost record station for this north
coastal species is on the Olympic Peninsula of Washington. It
probably developed in the humid, northern part of the Pacific
Coastal area previous to the last Pleistocene glaciation and ex-
tended its range to the south in Vashon-Wisconsin Time. The
range of tolerance in M. k. keenii seems to be more restricted than
that of M. lucifugus alascensis.

Myotis evotis. — In Washington, the distribution of this bat is
similar to that of Myotis lucifugus. The dark, forest race probably
originated in the north-coastal region. The paler race, that de-
veloped in the southwest, entered eastern Washington from Oregon.

Myotis thysanodes. — In Washington this species has been re-
corded only in the southeastern part where the Great Basin Fauna
occurs. It probably originated in the southwestern United States,
and a point in British Columbia a little way north of Washington
marks the northern edge of its natural range.

Myotis volans and Myotis californicus. — Remarks made
about Myotis lucifugus apply also to these two species.

Myotis subulatus. — The northwestern periphery of the range of
this species seems to be in eastern Washington.

Lasionycteris xoctivagans. — Undifferentiated subspecifically
from coast to coast, no basis is provided for judging the route by
which this species entered the state.

Pipistrellus hesperus. — The northwestern periphery of the
range of this bat, also, lies in eastern Washington.

Eptesicus fuscus. — Big brown bats from both eastern and
western Washington seem to have been derived from the Pacific

74 University of Kansas Publs., Mrs. Nat. Hist.

Coastal race of the species. Presumably it extended its range west-
ward across the Cascades in early post-Pleistocene Time.

Lasiurus cinereus. — No speculation as to the distributional his-
tory of the hoary bat seems justified at present.

Corynorhinus rafinesqiti. — The dark, coastal race of this bat
probably persisted in southwestern Washington and western Oregon
through Vashon Time and moved northward in the Recent. The
paler intermedins probably invaded eastern Washington from
eastern Oregon in the Recent.

Antrozous pallidus. — This species strays into eastern Washing-
ton from Oregon as part of the Great Basin Fauna.

Ursus americanus. — The dark, western race of the black bear
{altifrontalis) and the paler, inland race (rinnamomum) were prob-
ably separated by a glacial divergence. The inland race has
entered northeastern Washington in the Recent with other members
of the Rocky Mountain Fauna.

Ursus chelan, etc. — The apparent past distribution of chelan in-
dicates it to have invaded Washington from British Columbia since
the Pleistocene.

The apparent absence of grizzly bears from the southern Cascades
and western Washington may indicate their absence from these
a teas immediately before pre- Vashon time, or their extermination
in or shortly after that period.

Procyon lotor. — The raccoon of western Washington seems to
be the Pacific Coastal race which occurs also in western Oregon and
northwestern California. This indicates that the coastal race
(psora) was confined to the coastal area south of Washington during
Vashon Time and has only recently reinvaded western Washington.
It is possible, though less likely, that raccoons existed in south-
western Washington during Vashon Time but did not develop racial
characters, or that the Columbia was crossed so frequently that
gi netic differences were dispersed throughout the entire population.

Reasons why the second hypothesis is inadequate are: ill Rac-
coons range bin little north of the state of Washington, both easl
and west of the Cascades. (2) Raccoons of western Washington
and the area about San Francisco Bay, California, are as much
alike as are raccoon.- from southwestern Washington and north-
western Oregon. It is thoughl that raccoons, if resident in western
Washington since interglacial time, would have developed strong
racial character-, and the fact thai they have not indicates that
iliev have entered the state at a relatively recent date.

Distributional History 75

The raccoon of eastern Washington (excelsus) is a member of the
Great Basin Fauna and has probably included southeastern Wash-
ington in its natural range for a long period of time. The raccoon
has not extended its normal range into northeastern Washington,
although it is seemingly ideal raccoon habitat; only an occasional
vagrant occurs there. A stock of raccoons from which emigrants
might come has existed in southeastern Washington and the Yakima
Valley for some time. The Columbia River might serve as a high-
way by which emigrants could reach northeastern Washington.

Martes caurina. — The earlier distributional history of the west-
ern marten has been postulated by Davis (1939: 181-132), who
stated: "When the ancestral stock split into the two groups, the
one that gave rise to americana may have pushed eastward across
Canada to the Atlantic Coast; the other, giving rise to caurina, may
have migrated southward along the Sierra Nevada-Cascade and
Rocky mountains. Perhaps the great, ice sheet was instrumental in
pushing americana eastward and separating it geographically from
caurina/' The present occurrence of americana in Alaska and
British Columbia is thought to have been by invasion from the east
in postglacial time.

Davis' theory seems basically correct but subject to correction
in detail. The presence of caurina in the southern Rocky Moun-
tains suggests that it is not a Pacific Coastal species in the common
sense. Had americana occupied northern British Columbia in pre-
Wisconsin Time, it and not caurina would be expected to occur in
the southern Rocky Mountains today, for the form found in British
Columbia almost certainly would have been forced into the Rockie-.
The range now occupied by caurina in the Rocky Mountains is so
extensive as to suggest that martens could not have migrated into
all of it from the Pacific Coast since Vashon Time, even had the
region been unoccupied by any species of marten. The presence of
americana in Alaska and British Columbia suggests that it arrived
in those areas before caurina and that had the Rocky Mountains
been unoccupied by martens in pre-Wisconsin time, americana and
not caurina would have reached the Rockies first. It appears that
caurina occupied much of western North America in pre-Wisconsin
Time and was forced southward into the southern Rocky Mountains
and along the Pacific Coast by Vashon-Wisconsin ice.

The separation of americana and caurina may be supposed to
have occurred before the pre-Vashon-Wisconsin interglacial interval,
perhaps by a glacier similar to but antedating the Vashon-Wis-
consin glaciation.

7(i University of Kansas Publs., Mis. Nat. Hist.

The martens of western Washington {Martes caurina caurina) are
a coastal race. Those of northeastern Washington belong to a race
of the Rocky Mountain Fauna, and are referred to M. c. origenes.
Davis (1939: 132) refers the martens of Idaho to Martes caurina
caurina. I have compared specimens from Idaho with animals
trapped for fur from the Pacific Coast proper and feel that the ani-
mals from northeastern Washington and those from Idaho are more
like origenes than caurina, although perhaps not typical. The ani-
mals from the Pacific Coast proper are caurina and have darker
heads and brown instead of yellow patches on the throat.

Martes pennanti. — Fishers are found throughout the Cascade
Mountains and probably were widely distributed over western North
America in pre-Wisconsin Time.

Mustela erminea. — The distribution of ermines along the coast
of northern California and in the Cascade-Sierra Nevada of Oregon
and California indicates, as does their differentiation there, that
they ranged southward to these areas before and during Vashon-
Wisconsin Time.

In immediate pre-Yashon-Wisronsin Time, the dark race strcatori
probably occurred in western Washington. The race murica prob-
ably occurred in the Blue Mountains then, as it does today, but
probably occurred also in the Cascades of Washington. The descent
of the Vashon glaciers probably displaced strcatori from the northern
part of its range, at least temporarily. In the Cascades, murica was
likewise forced southward. Ermines related to the northern richaril-
sonii were forced into northern Washington and Idaho by the Wis-
consin ice. They probably were unable to live on the barren, ungla-
ciated plains of eastern Washington but persisted in Idaho.

The ranges of the three forms at the maximum extent of the
Vashon- Wisconsin may be reconstructed as follows: streatori in
southwestern Washington; murica in the southern Cascades and the
Blue Mountains; inricta stock in northern Idaho. While so isolated,
the ermines of the southern Cascades probably mingled, to a
certain extent, with streatori and developed the characters that
now separate gviosa from both murica and streatori. The inter-
mediate nature of gidosa lias been mentioned by Hall 1 1945: 85).

The retreat of the ice allowed streatori to move north and invicta
to move north and east into Washington and the northeastern
Cascades. To a lesser extent, gulosa may have moved north. The
poorly-marked race oh/mpicu probably evolved from streatori in
the Recent. It is difficult to account for the dark race fallenda.

Distributional History 77

It must have evolved from streatori in the Recent but the origin
of such a strongly marked race in such a short time is surprising.
It might be mentioned that a similarly differentiated race of chip-
munk, Eutamias amoenus felix, occupies much the same range.

Mustela frenata. — The long-tailed weasels of the Pacific Coast
behave as a plastic group and clearly show the effect of the Vashon-
Wisconsin Divergence. The range of the coastal race, altifrontalis,
indicates that it was isolated in southwestern Washington during
Vashon Time. In that period, or shortly after, it extended its range
southward but only along the extreme, coastal area of Oregon (see
Hall, 1936: 101). Following the retreat of the ice it extended its
range northward to the deglaciated area of western Washington.

Also following the retreat of the ice, a Great Basin subspecies
{nevadensis) extended its range northward. This race seems to
have been more adaptable and successful than other kinds of Great
Basin mammals, for it extended its range farther northward, east-
ward and westward than most.

A third race, washingtoni, was isolated in the southern Cascade
Mountains during Vashon Time and became differentiated from
both altifrontalis and nevadensis. It is now found in the Cascades
from central Oregon north to Mount Rainier. It is difficult to see
why it did not extend its range to include the northern Cascades
when the glacial ice left, but it did not. Instead altifrontalis en-
tered the northern Cascades from the west and nevadensis did the
same from the east. Weasels obtained in habitats north of Mt.
Rainier are intergrades between altijrontalis and nevadensis.

One is reminded here of the douglasii group of Thomomys talpoides
in which subspecies did not move north of Mt. Rainier in post-
glacial time. The area north of Mt. Rainier was populated instead
by gophers of the fuscus group, subspecies of which invaded the
area from the east. Perhaps Mt. Rainier itself served as a barrier
to alpine mammals in the immediate post-Pleistocene. Perhaps
Mustela f. washingtoni will eventually extend its range northward,
displacing the altifrontalis-nevadensis intergrades from the habitats
to which washingtoni may be better adapted.

The Blue Mountains of southeastern Washington are occupied
by a weasel (effera) that has a more extensive range in eastern
Oregon. The range of this race has probably not changed materially
for a long period of time.

One would expect the weasels from extreme northeastern Wash-
ington to be referable to the race oribasa, of the Rocky Mountain

78 University of Kansas Publs., Mrs. Nat. Hist.

Fauna. Instead they are intermediate between that race and the
Great. Basin race, nevadensis. Apparently nevadi nsis was so dy-
namic and adaptable that it actually entered the geographic ranges
of surrounding races for some distance. In view of Sumner's theory
for the retention of subspecies, one might say that the population
pressure of nevadensis on the periphery of its range is stronger than
the opposing pressure of some surrounding races.

Mistela vison. — Pending a review of the minks of North
America, little can be said concerning their historical distribution
in the state of Washington. From the general range of the species
in western North America, one would expect some effect of the
Vashon-Wisconsin Divergence to be apparent. There is some evi-
dence for this. Minks from Idaho and adjacent parts of British Co-
lumbia are distinctly less reddish than minks from the area about
Puget Sound, as noted by Davis (1939: 138).

Gilo luscus. — The range of the subspecies luteus, peculiar to
the Cascades and Sierra Nevada suggest that the wolverine may
have been forced southward in the Cascades and there isolated dur-
ing Vashon Time. The differences separating the southern race from
the northern may have been developed while the two populations
were isolated. The range of the wolverine was probably more ex-
tensive in glacial and immediate postglacial time than at present.

Lutra canadensis. — The otter of western Washington seems to
be a member of the Pacific Coastal Fauna. Little can be said re-
garding the distributional history of the species in the state, for
specimens from eastern Washington are not numerous enough to
permit of a person certainly establishing their systematic position.

Spilogale gracilis. — The western race of the civet cat ilatifrons)
seems to be a coastal race, isolated in southwestern Washington
and western Oregon during Vashon time. The eastern race, saxa-
tilis, is a race of the Great Basin Fauna, that has entered the state
from Oregon and that will probably extend its range to the north.

Mephitis mephitis. — Of the four subspecies of skunks occur*
ring in Washington, two seem to have been resident in the state
during Vashon Time. The western race, spissigrada, was probably
isolated in southwestern Washington and extended its range north-
ward, in the deglaciated area of western Washington, after the
retreat of (lie ice. Another race (notata) was probably isolated
in the southeastern Cascades and adjacent Oregon. M. m. hud-
sonica of the Rocky Mountain Fauna entered the northeastern
pari of Washington after the ice retreated from there. A race of

Distributional History 7!>

the Great Basin Fauna, major, entered southeastern Washington
from Oregon and may eventually extend its range farther north.

It is interesting to note that both of the western races, spissi-
grada and notuta, both of which probably developed in Washing-
ton during Vashon Time, occupy limited ranges in adjacent Oregon
(Bailey, 1936: 308).

Taxidea taxis. — This species has probably long been resident
on the Columbian Plateau and in southeastern Washington. For
the early distributional history of the species see Hall !l!>44: 17).
Pleistocene remains, referable to this race, have been found in
Franklin County.

Yulpes ftjlva. — The red fox of the Cascades was probably
isolated there during Vashon Time by glacial ice. Its range ex-
tends southward in the Cascades to Oregon. The fox of eastern
Washington is probably a member of the Rocky Mountain Fauna
that lived in the Blue Mountains of southeastern Washington in
Wisconsin Time and that emigrated to northeastern Washington
in Recent Time.

Canis latrans. — The distributional history of the coyote in
Washington is not clear.

Canis lupus. — The dark wolf (fuscus) of western Washington
is probably a coastal race. The race that may have occurred in
northeastern Washington probably was an invader from the Rocky
Mountain Fauna, and the race that possibly occurred in south-
eastern Washington would be assumed to have long been a resi-
dent of the area.

Fells concolor. — The cougar of western Washington is a
coastal race, probably developed while isolated in southwestern
Washington and western Oregon. The cougar of northeastern
Washington probably entered the state with other Rocky Mountain
species, early in the Recent. The cougar of the Blue Mountains
of southeastern Washington has probably long been resident there.

Lynx rufus. — The bobcat of western Washington seems to be
a coastal race that was isolated in either southwestern Washington
or western Oregon by Vashon ice. It has since extended its range
into southern British Columbia. The bobcat of eastern Washing-
ton seems to be a member of the Great Basin Fauna that has
spread to some forested areas on the periphery of the more arid
life-zones.

Lynx canadensis. — The lynx is an element of a northern fauna
that was probably forced southward into the Cascades and Rocky

80 University of Kansas Publs., Mus. Nat. Hist.

Mountains. Its range was probably more extensive, as is indi-
cated by the scattered records of its occurrence in Oregon (Bailey.
1936:271).

Marmot a monax. — The woodchuck invaded northeastern Wash-
ington in the early Recent with the Rocky Mountain Fauna.

Marmota flaviventris. — The yellow-bellied marmot is a typical
member of the faunas of the Great Basin and the southern Rocky
Mountains. It has doubtless entered southeastern Washington
from eastern Oregon at an early time. In northeastern Washington,
west of the Columbia River, it occupies alpine habitat, but it does
not occur farther east, where Marmota monax is found, or in the
Cascades where Marmota caligata lives.

The yellow-bellied marmots are great wanderers, and commonly
are found in scattered outcrops far out on the Columbian Plateau.
There is even one record for western Washington, near Bellingham,
Whatcom County. This individual must have crossed some low
pass in the Cascades from the area about Lake Chelan. There are
records of eastern Washington birds occurring in this same area, so
it seems likely that the marmot was a natural stray and not an
animal that escaped from captivity.

Marmota caligata. — The absence of the hoary marmot from the
Cascades of Oregon, and the presence there of Marmota flaviventris,
indicates that the species did not occur in the southern Cascades
of Washington during Vashon Time. Presumably the hoary marmot
is a member of the fauna of the northern Rocky Mountains and
entered the Cascades of Washington in the Recent, after which it
spread widely and rapidly owing to lack of competition with any
established species of marmot,

Marmota olympus. — This species has probably lived in the
Olympic Mountains since pre-Vashon Time.

Citellus washingtoni. — This ground squirrel has probably
lived on the Columbian Plateau since before Wisconsin Time.

Citellus townsendii. — The Townsend ground squirrel probably
entered the Yakima Valley area from Oregon. The differences be-
tween it and its relatives in Oregon indicate a considerable period
of isolation but one far shorter than the period during which wash-
ingtoni is presumed to have been isolated from toivusctulii.

Citellus columbianus. — The Columbian ground squirrel might
have been forced southward in the Rocky Mountain area by the Wis-
consin glaciation, might have lived in southeastern Washington since

Distributional History 81

then, and might have invaded northeastern Washington in the
Recent with other species of the Rocky Mountain Fauna.

Citelltjs beecheyi. — This ground squirrel is known to have en-
tered Washington about 1915 from Oregon.

Citellus sattjratus. — The mantled ground squirrel of the Cas-
cades probably evolved, from the lateralis stock, as a separate
species while isolated in the southern Cascades during Vashon Time.
It is a poorly differentiated species and may actually be instead a
strongly marked subspecies.

Citellus lateralis. — The golden-mantled ground squirrels of
northeastern and southeastern Washington are closely similar. It
is deduced that connectens of southeastern Washington developed
the differences that characterize it while isolated, from the main
stock, in the Blue Mountains area of Washington and Oregon.

The race found in extreme northeastern Washington (tescorum)
probably reached that area in relatively recent times. Its range in
Washington is more restricted than that of several other members of
the Rocky Mountain Fauna; areas of suitable habitat west of the
Columbia River are not inhabited by these ground squirrels. Its
range in Washington is almost exactly that of {Marmota monax).

Tamias minimus. — The least chipmunk of the Yakima Valley
is the same race {scrutator) as that occupying the Great Basin area
of Oregon and Nevada. It must have crossed the Columbia in rela-
tively recent times. Had it been resident in the isolated Yakima
Valley area for any considerable period of time, the development
of distinctive racial characters there would be expected. Perhaps,
then, it has not been resident there as long as has the Townsend
ground squirrel which, though closely related to the ground squirrel
of eastern Oregon, is racially distinct.

The least chipmunk of the Columbian Plateau is thought to be
racially distinct from its relatives in the Yakima Valley and east-
ern Oregon. Probably it reached the Plateau very early in the
Recent. It has probably not been separated from the parent stock
as long as has the ground squirrel (Citellus washingtoni) of the
plateau. The ground squirrel is specifically rather than racially
distinct.

Tamias amoenus. — The distributional picture of the yellow pine
chipmunks in Washington is complex. (Fig. 81.) Certain habits
of these mammals doubtless have modified what was probably the
original postglacial distribution of the species. Chipmunks are

6—4993

82 University of Kansas Publs., Mrs. Nat. Hist.

diurnal and natural selective factors for color possibly operate more
strongly on animals active by day than on nocturnal animals. Yel-
low pine chipmunks are neither forest nor deserl inhabitants. In-
deed, dense forests or open deserts serve as barriers to their distri-
bution. They prefer brush lands, open woods, and other habitats
where there is food and cover but abundant sunlight. In such hab-
itats they are almost independent of altitude, temperature and hu-
midity. They live in the Olympic Mountains where rainfall is
heavy and humidity high. They live and breed at considerable
altitudes in the Cascades, even in the crater of Mount Rainier, where
snow, ice and freezing conditions exist the year around. On the
other extreme, they occupy the low. open pine forests and brush
lands at the lower edge of the Arid Transition Life-zone where tem-
peratures, in summer, are high and rainfall scarce.

We find in the present distribution of the species in the Cascade-
Sierra Nevada chain and the Rocky Mountains, indication that the
species had a wide geographic range over western North America
previous to the Vashon-Wisconsin glacial interval. Probably the
range of the species extended in an arc, from the Rocky Mountains
across northern Washington to the Cascades, around the basaltic
plateau desert in eastern Washington and Oregon. Presumably the
descent of the Vashon-Wiseonsin glaciers broke this arc into two
parallel geographic ranges, the Rocky Mountains and the Cascade-
Sierra Nevada chain, with a desert area between.

Almost every species of forest-dwelling animal had its range sep-
arated into two parts by the southward movement of the glaciers.
Most of these forest-dwelling species were composed of relatively
homogeneous stocks, although the yellow pine chipmunk probably
was not. The extensive range of tolerance of the yellow pine chip-
munk to altitude and climate and its unique habital requirements
cause it to meet radically different natural selective factor-. The
predators of the chipmunks near Wenatchee. Chelan County, would
include: rattlesnake, gopher snake, badger, striped skunk, prairie
falcon, red-tailed hawk and other predominantly desert -dwelling
species. The chipmunks at Stevens Pas:-, in the mountain- to the
west, would have to contend with: marten, black bear, goshawk,
bald eagle and other alpine predators. At the pre-eiit lime, the
chipmunk of the eastern Cascades is racially distinct from that of
the higher Cascades. Geologic and botanical evidence indicates
that the Columbian Plateau was a desert in pre-Wisconsin Time.
We suppose that a transition from alpine conditions in the Cascades

Distributional History 83

to desert conditions on the Columbian Plateau existed even in pre-
Wisconsin Times. We suppose also that the chipmunk existed in
this transition area and in the Cascades before Wisconsin Time and
in the southern and southeastern Cascades during Wisconsin Time.
We further suppose that the differences separating the transition
area race (Tamias amoenus affinis) from the mountain race (T. a.
ludibundus) came about through natural selection and not as a re-
sult of geographic isolation. The principal difference between the
two is the paler color of the race in the transition area.

The descent of the Vashon-Wisconsin glaciers, then, found two
races of the yellow pine chipmunk in the Cascades. Chipmunks
living north of the Columbian Plateau, in northern Washington and
British Columbia, were probably forced southward onto the inhospit-
able plains of the plateau and exterminated. Farther east, north of
northeastern Washington, chipmunks from the north were probably
forced southward to compete with resident chipmunks.

The range of Tamias amoenus luteiventris in Washington, Idaho
and Montana is most unusual (See Howell, 1920; Davis, 1939). From
a compact range in Montana, two long fingers reach northward and
westward. The western finger crosses Idaho to end in the Blue
Mountains of Oregon and Washington. The northern finger crosses
northern Idaho, northeastern Washington and extends on into south-
ern British Columbia. Between these two fingers of the range of
lutein ntns another race {canicaudus) is found. This race occupies
a more lowland area than does luteiventris. The range of luteiven-
tris in the northern Rocky Mountains is extensive. Presumably
this race occupied an area farther north in pre-Wisconsin Time and
was forced southward to its present range by the Wisconsin glaciers.
The original population occupying extreme eastern Washington and
adjacent Idaho was Tamias amoenus canicaudus. The pre-Wis-
consin range of this race might have been more extensive. At any
rate, luteiventris which was driven southward displaced canicaudus,
or some other race of chipmunk, from much of the Rocky Mountain-
south of the glacier. The northern chipmunks were adapted to more
boreal conditions and perhaps otherwise better suited to environ-
mental conditions of the northern Rocky Mountains. A small pop-
ulation of the older established race (canicaudus) persisted in low-
land areas of eastern Washington and adjacent Idaho.

Regarding the range of canicaudus, surrounded by the range of
luteiventris on three sides and faced by desert on the west, Davi-
1 1930: 220) writes, "It may be that, of these two races, luteiventris

84 University of Kansas Publs., Mis. Nat. Hist.

has a greater range of tolerance to environmental conditions and.
thus, is able to succeed in areas to which canicaudus is not adapted.
This inference is supported by the fact that luteiventris occupies a
large range which is diversified geographically and climatically,
whereas canicaudus seems to be limited to a much smaller, more
nearly uniform area." Seemingly canicaudus now exists only in an
area ideally suited to it, and one where it can successfully compete
with the generally more adaptable and successsful luteiventris. The
maximum extent of the glacial ice, then, found luteiventris the domi-
nant chipmunk in the northern Rocky Mountains, with an isolated
population of canicaudus in eastern Washington and adjacent Idaho.
The topography of the ground moraine exposed by the retreat of
the Vashon-Wisconsin glaciers was a barrier to many species of
mammals. The rough, rocky surface with thin soil probably first
supported mosses and grasses, then brush, and later trees. The
earlier stages of plant succession on the deglaciated ground probably
presented ideal habitat for yellow pine chipmunks. Certainly the
races immediately adjacent to the glaciers extended their ranges
farther north than many species. In eastern Washington, T. a.
luteiventris spread to the northeastern corner of the state and on
into British Columbia. In the northern Cascades, T . a. affinis spread
northward and eastward, across the Okanogan River, into north-
eastern Washington as far as the range of luteiventris. The chip-
munk of the higher Cascades (ludibundus) likewise extended its
range northward into British Columbia. In the northwestern
Cascades of northern Washington and southern British Columbia,
a richly-colored race, T. a. felix, now occupies a limited geographic
range. This race doubtless originated from ludibundus stock but
the method of its development is unknown. Perhaps in early post-
glacial time, selective factors developed in chipmunks of the western
slopes of the Cascade Mountains the rich, dark color of felix. The
ancestral ludibundus may have given rise to a pale race, affinis,
in the arid eastern Cascades and a dark race, felix, on the
humid western slope of the Cascades. This seems improbable for
there is no trend to darker color on the western border of the range
of ludibundus south of the range of felix, and instead, affinis may
have given rise to ludibundus. A more appealing hypothesis is that
a local mutation in some ludibundus stock so changed the range of
tolerance of a portion of the population that it was allowed to enter
the more dense habitat along the coast north of the Frascr River and,
there, isolated by habitat selection, it developed the characters of

Distributional History 85

felix. Population pressure later forced it eastward until the eastern
border of its range again met the range of the ancestral race,
ludibundus..

The chipmunks of the Olympic Mountains probably reached their
present range from the Cascades. Their probable path of emigration
was westward from Mt. Rainier, along the glacial outwash train of
Nisqualli Glacier, to the moraine and outwash apron of the Vashon
Glacier and thence to the Olympics. So similar are the chipmunks
of Mt. Rainier and the Olympic Mountains that Howell (1929) in-
cluded Mt. Rainier in the range of caurinus.

Briefly summarized, the probable pre-Vashon-Wisconsin distribu-
tion of chipmunks of the species Tamias amoenus in Washington was:
ludibundus in the higher Cascades; affinis in the eastern Cascades;
canicaudus in eastern Washington and adjacent Idaho; and lutei-
vt ntris in the area north of the range of canicaudus. The descent of
the Vashon- Wisconsin ice restricted but did not materially alter the
ranges of ludibundus or affinis. On the east, luteiventris w r as forced
southward to compete with canicaudus and displaced it over a large
region, especially in mountainous areas. Following the retreat of
the ice, luteiventris, affinis, and ludibundus extended their ranges
northward over the deglaciated territory. A stock of ludibundus
that moved westward from Mt. Rainier became isolated and gave
rise to caurinus. In some less obvious development, ludibundus
stock gave rise to felix north of the Fraser River in the Cascades.

Tamias ruficaudus. — Until a better understanding of the range
of this chipmunk and its relation to other Tamias is gained, uncer-
tainty will remain concerning its distribution in the past.

Tamias townsendii. — This is a typical coastal species that ranges
southward, along the coast, to California. The lowland race of
western Oregon and Washington {townsendii) probably occurred no
farther north than southwestern Washington when the Vashon
Glacier was in place. Chipmunks of this species in the Cascades
and in the southern Olympic Mountains probably developed inde-
pendently the slightly paler color that separates coo-peri from town-
sendii. The tendency for species of the Pacific Coastal Fauna of
the Cascades and the Olympic Mountains to be paler than their
lowland relatives is widespread.

After the retreat of the ice, both races probably moved northward.
Perhaps because of its alpine adaptations, cooperi has moved farther
than townsendii. Also, townsendii, in the lowlands, ranges to the
Fraser River, a barrier not encountered by cooperi.

86 University of Kansas Publs., Mus. Nat. Hist.

Sciurus griseus.- — This species of the Pacific Coastal Fauna
probably entered Washington from Oregon since the retreat of the
Vashon Glacier. It has probably entered the state in relatively re-
cent times.

Tamiasciurus hudsonicus. — The two species of red squirrels,
T. hudsonicus and T. douglasii, arc specifically distinct and probably
became differentiated in the Pleistocene when southward moving
glaciers eut in two the range of the ancestral stock. The morpho-
logical differences are too great, comparatively, to have occurred
during the Vashon-Wisconsin Divergence. T. hudsonicus probably
occupied a range in pre-Wisconsin Time that included the Rocky
Mountains and areas to the north. Glacial ice probably restricted
the range of hudsonicus in Wisconsin Time but after the retreat of
the ice hudsonicus moved northward to reoccupy its former range.
It also moved westward across northern Washington to the Cascades,
where it met the range of douglasii. Farther north, it moved west-
ward to the Pacific, thus occupying an area that, in pre-Vashon
time, probably was occupied by douglasii.

Tamiasciurus douglasii. — The Douglas squirrel probably oc-
cupied the coastal region of Oregon, Washington and British Colum-
bia in pre-Vashon Time. The descent of the ice restricted its range
to southwestern Washington and areas to the south. After the re-
treat of the ice it moved northward somewhat but. like other coastal
species, the movement was slow. Meanwhile hudsonicus from the
Pocky Mountain Fauna, had spread to the coast of British Colum-
bia.

Glaucomys sabrinus. — This Hying squirrel is a plastic species.
It inhabits all of the forested parts of Washington. The distribu-
tional picture presented by the 5 races (Fig. 92) which occur in
Washington is complicated. The ranges of 3 of these lie principally
outside the state of Washington.

The race arcgonensis occupies Washington and Oregon west of
the Cascades; juliginosus occupies the Cascades of Washington,
Oregon and southern British Columbia; columbu nsis occupies the
interior valleys of British Columbia and adjoining Washington;
latipes occupies the northern Pocky Mountains of British Columbia.
northern Idaho and extreme northeastern Washington; and bangsi
occupies the Blue Mountains of Washington and Oregon and a wide
range in Idaho and eastward.

The differences separating the race oregont nsis from other sub-
species found in Washington are relatively gnat. This lowland

Distributional History 87

race is smaller and richer in color. The other races exhibit slight
but relatively constant differences. The relatively great difference
between oregonensis and the other races indicates that oregonensis
was isolated from the remainder of the species for a considerable
time. Presumably oregonensis was a strongly differentiated coastal
race in pre-Yashon Time and occupied most of western Washington
and Oregon. The descent of the Vashon ice restricted the range of
oregonensis to southwestern Washington and western Oregon. The
descent of the ice forced a northern race, fuliginosus, southward
into the range of oregonensis. The northern race, adapted
to boreal conditions, was able to compete successfully with the es-
tablished oregonensis only in mountainous areas. In the Cascade
Mountains, fuliginosus extended its range southward to southern
Oregon.

The descent of the Wisconsin ice in eastern Washington forced
the flying squirrels of adjacent British Columbia southward into
the Rocky Mountains. These squirrels were probably closely re-
lated to fuliginosus, or to bangsi, which latter race already may have
been established farther south in the Rocky Mountains. The Blue
Mountains of southeastern Washington were probably inhabited by
hungsi in Wisconsin times, or even earlier. The retreat of the
Vashon-Wisconsin glaciers allowed the flying squirrels to extend
their ranges northward. In western Washington oregonensis moved
to southern British Columbia. In the Cascade Mountains the more
boreal fuliginosus moved much farther northward and. north of the
Okanogan Valley, spread eastward to the arid, interior valleys of
British Columbia. Subsequent differentiation in the population of
the arid, interior valleys developed the slightly differentiated race
columbiensis. Farther east, flying squirrels from the northern Rocky
Mountains moved northward. Northeastern Washington and ad-
jacent British Columbia were occupied by latipes, derived from
bangsi.

Thomomys talpoides. — Views as to the probable historical dis-
tribution ol this plastic group have been presented in an earlier
report (Dalquest and Scheffer, 1944: 308-333). This may be briefly
summarized as follows.

Previous to Vashon-Wisconsin Times, pocket gophers occupied at
least the Cascade Mountains and the Columbian Plateau of Wash-
ington. The race occupying the Columbian Plateau, devexus, was
probably racially distinct in pre-Wisconsin time. The descent of
the Vashon-Wisconsin glaciers isolated gophers in the southern

88 University of Kansas Publs., Mrs. Nat. Hist.

Cascades. Three racial stocks developed there: shawi in the
Mount Rainier area; limosus in the Columbia River Valley south
of the Cascades; and immunis in the mountainous area between the
other two races. At the maximum extent of the Wisconsin ice,
gophers from the Columbia River Valley (limosus) were able to
cross the Simcoe Bridge and reach the Blue Mountains. With the
retreat of the Wisconsin ice, the Simcoe Bridge was closed. Gophers
isolated in the Blue Mountains developed the racial characters of
aequalidens and those between the Blue Mountains and the Colum-
bia River intergraded with the desert race, de vents. These inter-
grades, which have, also, some characters of their own, bear the
name columbianus. Gophers in the southern Cascades (limosus)
moved westward on prairie-like river terraces to Clark County where,
isolated, they became racially distinct (douglasii).

Gophers from the Mount Rainier area {shawi) moved westward
on glacial outwash trains to the extensive outwash aprons of the
Vashon glaciers in the area about Puget Sound. Here they multi-
plied and spread to the Olympic Mountains. Growth of forest on
the original outwash apron broke the area into numerous isolated
prairies. Gophers in the Olympic Mountains (mclanops) were iso-
lated from those in the area about Puget Sound. Six distinct races
originated on the isolated prairie (glacialis, tacomensis, puget( nsis,
yelmensis, tumuli, couchi).

Following the retreat of the glacial ice from eastern Washington,
pocket gophers from the Blue Mountains of Oregon (wallowa)
moved northward into Washington and gophers from the Rocky
Mountain Fauna of Idaho moved onto the deglaciated part of north-
eastern Washington. From northeastern Washington they spread
westward to the Cascades and thence southward to meet the native
gophers of the Cascades in the Yakima Valley Area. No racial
differentiation in these gophers occurred; all are referable to fuscus.
Where fuscus and the native gophers came together in the Yakima
Valley, a new race, yakimensis, developed.

Perognathl's parvus. — Three races of the pocket mouse occur
in Washington. Two of these (lordi and columbianus) occur on the
Columbian Plateau. Pike many desert species that occur on the
Columbian Plateau, the pocket mice are rather different than their
relatives in eastern Oregon. Presumably they have been isolated
on the plateau since before Vashon-Wisconsin Times.

The range of the pocket mouse of southeastern Washington,
Perognathus parvus parvus, is continuous with the range of the race

Distributional History S9

in Oregon. This same race occurs in the Yakima Valley, whence
it probably arrived from Oregon in relatively recent time.

The distribution of pocket mice on the Columbian Plateau, in
eastern Oregon and in the Yakima Valley resembles that of the
least chipmunk in those areas. It is also similar to, but of more
recent origin than, that of the ground squirrels, Citellus Washington',
and townsendii.

Dipodomys ordii. — This kangaroo rat enters the desert area of
southeastern Washington from Oregon. It may be expected event-
ually to cross the Columbia River to the Yakima Valley and the
Snake River to the Columbian Plateau.

Castor canadensis. — Two races of beavers occur in Washington.
One, found in southwestern AVashington and northwestern Oregon,
is dark with a short, wide skull. The other, ranging over most of
the state, is paler with a longer, narrower skull.

The form now found in southwestern Washington and adjacent
Oregon (idoneus) was probably isolated there by the Vashon gla-
ciation and developed its characters while isolated. The other race,
leucodonta, was probably widely spread in Wisconsin Time. Beav-
ers are present in Moses Lake, in almost the center of the Colum-
bian Plateau. Beavers might well have lived in the streams of melt
water that emerged from the Wisconsin Glacier. The beavers of
western Washington, save those in the extreme southwest, are like
the beavers of eastern Washington. It seems likely that the
race leucodonta originated north of the state of Washington and
was forced southward by the Vashon-Wisconsin glaciers. This
northern race, adapted to boreal conditions, competed with the
resident coastal race, idoneus, and occupied much of its range.
The distribution of the races of muskrat in Washington closely re-
sembles that of the beavers.

Onychomys leucogaster. — The desert-dwelling grasshopper
mouse has doubtless entered eastern Washington and the Yakima
Valley from eastern Oregon at a relatively recent time.

Reithrodontomys megalotis. — The harvest mouse, like the
grasshopper mouse, seems to have entered Wahington from Oregon
at a relatively recent date. Within the last ten years it has ex-
tended its range into the Okanogan Valley in British Columbia.

Peromyscus manictjlattjs. — Six subspecies of Peromyscus mani-
culatus occur in the state of Washington. The geographic range
of one of these (rubidus) lies mainly in the states of California and
Oregon and includes, so far as is known, a single small island in

90 University of Kansas Ptjbls., Mrs. Nat. Hist.

the Columbia River that is politically within the state of Wash-
ington. Another {hollisteri) is restricted to certain islands in north-
ern Puget Sound and obviously has become subspecifically differ-
entiated in postglacial time. The remaining four subspecies,
namely oreas, austerus, curtemisiae and gambelii, have extensive
geographic ranges. These subspecies are not confined to their geo-
graphic ranges by geographic barriers. Deer mice occur in the deep
forests and the open desert, on high mountains and in low valleys,
and are almost everywhere the commonest species of mammal
present.

The study of several populations of deer mice from any general
area usually shows small but constant differences between them.
Dice (1939: 21) studied stocks of deer mice from nine localities in
southeastern Washington and found significant differences between
several of them. A statistical study of mice from the San Juan
islands shows that the population of almost every island is different
in one or more respects from the mice of any other island. Geo-
graphically separated populations of "wild caught" mice of the sub-
species austerus, of the mainland, were statistically compared and
significant differences were found between these populations, too.
Small, differentiated populations are to be found in many parts
of the state, and each subspecies appears to be an assemblage of
such tiny genetic units.

These genetic units probably are the microgeographic races of
Dcbzhansky (1937). They have been intensively studied by Sum-
ner (1917 A, Bi and Dice. An especially important paper by the
latter author (1940) summarizes much of the available informa-
tion on speciation in Peromyscus and clearly discusses the micro-
geographic races of Peromyscus.

The numerous microgeographic race- of Peromyscus maniculatus
in Washington present diverse combinations of characters which
could result from the random fixation and elimination of genes
(Wright. 1932:360-362). Such a hypothesis, however, requires at
least partial isolation of the populations involved. The mechanism
of such isolation, in such populations of deer mice as we have
studied, is not readily apparent. Some microgeographic races are
nut separated by noticeable geographic or ecologic barriers and
the distance between their ranges is not too great to be traversed
by a deer mouse. The tendency to remain on a home range may
have the same effect as isolation would be supposed to have.

The work of Mnrie and Murie (1931: 200-209, L932: 79) is en-
lightening in this respect. These authors found that mice residing

Distributional History !>1

in a locality tended to remain in that locality; individuals trapped
and marked were retaken in the same locality a year later. Indi-
viduals released some distance from the point of capture remained
where released or returned to the point of capture. Transported
individuals did not spread at random. The home instinct was de-
veloped in young as well as in old mice. Two mice in the gray
pelage, four to eight weeks old, returned to their home ranges
from distances one and two miles away. The authors fix the home
range of an individual Pero?nyscns m. artemisiae in Teton County.
Wyoming, at approximately one hundred yards in diameter.

This home-range instinct is essentially a lack of incentive for
individual mice to emigrate to new localities wdiere mice of the
same species are already established. This may partly account,
for the microgeographic races of deer mice in Washington.

Dice (1939: 21 1 pointed out that, except in color, the differences
in nine stocks of mice from southeastern Washington could not be
correlated with environmental factors. We have found this to
be true of microgeographic races throughout the state of Washing-
ton.

Of the four subspecies of deer mice that occupy extensive geo-
graphic ranges in Washington, one, oreas, is a long-tailed form
that seems not to intergrade with austerus, a neighbor in western
Washington that has a tail of moderate length. These two and
gambclii, a short-tailed form with which oreas intergrades, are
easily distinguished. In eastern Washington two short-tailed sub-
species, gambclii and artemisiae, are currently recognized. The
taxonomic relationships of these two subspecies are complex. The
subspecies gambelii has an extensive geographic range in Oregon
and California. These mice, with short tails, occur in the Wallula
Water (lap of southeastern Washington and on the Columbian
Plateau. To the west the desert conditions of the Columbian
Plateau fade into the Transition Life-zone forests of the eastern
Cascade Mountains. The pale, short-tailed desert mice {gambelii)
gradually change to the dark, long-tailed subspecies, oreas, that
occupies the Cascade Mountains.

North of the Columbian Plateau, in northeastern Washington, the
deer mice are darker and relatively longer-tailed than on the Colum-
bian Plateau. Some populations are distinctly reddish, almost as
reddish as oreas. Although assigned to artemisiae, they are almost
identical with populations of deer mice from the eastern Cascade
Mountains, known to be intergrades between oreas and gambelii.
This fact, and the presence of surprisingly oreas-\\ke characters in

92 University of Kansas Publs., Mus. Nat. Hist.

some microgeographic races in extreme northeastern Washington,
may indicate that the race called artemisiae is a group of intergrades
between gambelii and an oreas-like mouse that has become extinct.

Intergradation between gambelii and artemisiae is normal and
takes place gradually where the ranges of the two subspecies meet.
The same is true of intergradation between oreas and gambelii in
the eastern Cascade Mountains. West of the Cascade Mountains
the range of oreas meets the range of austerus. These two sub-
species appear not to interbreed in nature, since no intergrades were
taken at any of the numerous localities in western Washington where
the mice were trapped. Pure populations of the two subspecies
exist within a few miles of each other. In the valley of the Sky-
komish River, in the western Cascades Mountains, from the town
of Skykomish, King County, to the lowlands to the west, only
austerus was found. In the coniferous forests of the mountains
above Skykomish, only oreas was taken. Several pairs, each an
oreas and an austerus of the other sex, were kept from four to six
months, and one pair was kept for a year, but they did not repro-
duce. The oreas were from the upper Skykomish Valley and the
austerus were from Seattle, King County. Along the border of the
ranges of the two subspecies in the Skykomish River Valley, a
definite habitat preference was noted. The coniferous forests were
occupied by oreas and brush or deciduous forests by austerus.
Within the range of austerus and within the range of oreas only one
subspecies is found whether the habitat be coniferous forest or
thickets of alder and willow, but where the ranges of the two sub-
species meet austerus occurs only in the thickets of aspen and willow
and oreas occurs only in the coniferous forest.

The morphological characters that separate oreas from austerus,
namely, larger size, richer color, and longer tail, are all features that
might be associated wfith a more arboreal existence in dark, coni-
ferous forests. Our observations show that oreas is, to a large extent,
arboreal. Traps nailed to tree trunks six to eight feet from the
ground ami set for flying squirrels often caught oreas in the Sky-
komish River Valley. On one occasion I walked up a leaning
tree trunk to set a, trap, fifty feet from the ground on the trunk
of another tree that was upright. An orats was found in this (rap
the next morning. Svihla (1933: 13) relates how a specimen of
oreas that escaped from a live trap took refuge in a tree when pur-
sued. We have set numerous traps for Hying squirrels in the area
about Puget Sound. As far as memory and field notes serve, we

Distributional History 93

have never taken austerus in these traps. Our observations on the
habits of austerus are much more abundant than those on oreas,
but for austerus no evidence of arboreal activities has been obtained.

There are, then, two subspecies that do not interbreed, each cap-
able of existing in any ecologic niche that will support deer mice.
"Where the ranges of the two subspecies come together, they compete.
The large size, richer color, longer tail, and perhaps arboreal habits
of oreas give it an advantage in coniferous forests. The small size
and dark color of austerus give it an advantage in other habitats,
especially, perhaps, in winter, when the deciduous trees have shed
their leaves.

The differences separating austerus from oreas would be expected
to have developed under complete isolation. That oreas developed
from austerus or austerus from oreas seems impossible. A glance
at the distribution map (Fig. 107) shows that the range of austerus
clearly separates the range of oreas into two segments. The range
of austerus tapers out to the north, east and west. On the south
austerus intergrades with rubidus from Oregon, a subspecies from
which it is but slightly differentiated.

The geographic behavior of the four subspecies of deer mice that
occupy extensive areas in Washington may be summarized as fol-
lows: P. m. gambelii occupies southeastern Washington and inter-
grades normally with oreas in the eastern Cascade Mountains and
with artemisiae at the northern edge of the Columbian Pleateau;
artemisiae, occupying northeastern Washington, closely resembles
populations of mice that are known to be intergrades between oreas
and gambelii and itself intergrades with both oreas and ga??ibelii;
oreas occupies most of western Washington, intergrades with its
neighbors artemisiae and gambelii to the east, but does not inter-
grade with austerus, its morphologically more similar neighbor in
the west; austerus occupies a range in western Washington that is
almost surrounded by the range of oreas, a subspecies with which it
apparently does not interbreed.

The relations of these four subspecies appear to be the result of
certain great changes in the range of deer mice in the Pacific North-
west that occurred during and since the last Pleistocene glaciation.
We reconstruct these changes as follows. In the Puyallup inter-
glacial cycle, just previous to the last (Vashon-Wisconsin) continen-
tal glaciation, dines, or gradual transitions existed in the ranges of
the deer mice along the Pacific Coast. The small, dark, short-
tailed mice of the coastal redwood forests of California and Oregon

!)4 University of Kansas Publs., Mus. Nat. Hist.

became gradually larger, redder and longer-tailed to the north,
climaxing in a large, red, long-tailed form in the spruce forests of
southern Alaska and northern British Columbia. A similar cline
(\isted inland. Small, pale, short-tailed mice of the (treat Basin
became increasingly larger, darker, and longer-tailed to the north,
reaching a maximum in the spruce forests of northern British Colum-
bia.

The development and advance of the Vashon-Wisconsin ice sheet
exterminated mice over much of British Columbia and the northern
United States. Long-tailed northern mice were forced southward
and contacted the southern, short-tailed forms. The preglacial clines
were thus destroyed.

What might be the southern part of the western cline may be noted
in the deer mice of western Oregon today. From the southern coast
of Oregon the mice \P. in. rubidus) do become larger, redder and
longer-tailed to the north. Tin' climax of this cline is now reached
iii austerus, of the Puget Sound area of Washington. The (dine is not
continued farther northward because the range of on<is is encoun-
tered.

Tin- advance of the Vashon-Wisconsin ice from the north prob-
ably forced species of mammals southward, ahead of it, including
the long-tailed northern deer mice which generation by generation
encountered progressively shorter-tailed forms of resident mice.
Perhaps the unfamiliar, and certainly the extremely frigid, con-
ditions in the fore of the glacier exterminated the short-tailed in-
dividuals but favored the long-tailed mice, since the latter originally
were adapted to a boreal environment. The climax of the ice ad-
vance found the glaciers just within the political limits of the United
State.- and found the long-tailed mice spread before the ice front.

In western Washington the Vashon glacier advanced as far south
as the southern edge of the Puget Sound area. Pong-tailed northern
mice advancing before it reached the Columbia River. This gla-
cially swollen stream served a- a natural barrier and prevented their
southward extension. At this lime the northern mice had traversed
more than half the length of the original western cline. The north-
ern mice, originating in a boreal habitat a thousand miles away,
were unable to interbreed with the southern mice and such indi-
viduals as might have crossed the Columbia River never be-
came established in Oregon. During the existence of the glacier in
western Washington, the long-tailed northern emigrants competed
with the resident deer mice of western Washington to the total elimi-

Distributional History 95

nation of the resident mice. The retreat of the Vashon Glacier
from Washington found the northern mice in complete control of
that part of the state from the Pacific Ocean to the Cascade Moun-
tains.

In eastern Washington the Wisconsin Glacier advanced south to
the northern edge of the Columbian Plateau. Northern mice ad-
vancing before it probably did not survive on the treeless plateau
but existed in forested areas of northern Idaho and driftless areas
of northern Washington. Xo natural barriers separated the north-
ern mice from the pale, short-tailed forms. The nonuniform topog-
raphy perhaps allowed more mingling of the two types where cli-
matic conditions permitted. Intergradation in some places as well
as competition and elimination of one form or another in other places
occurred. Following the withdrawal of the Vashon ice and the es-
tablishment of soil and forests on the deglaciated land, the long-
tailed mice of western Washington (P. m. oreas) apparently spread
northward, unhindered by competition, until they reached southern
Alaska. The deep coniferous forests of western Washington pre-
sented conditions acceptable to oreas and it persisted there despite
postglacial changes in climate.

Postglacial changes in climate did, however, permit the dark,
short-tailed mice (rubidus) to cross the Columbia River and to be-
come established in suitable habitats, namely the deciduous jungles
of the river valleys. From these points the mice spread northward
through the lowlands of western Washington, infiltrating the range
of oreas, competing with it, and driving it from the lowlands. Thi>
movement is in progress today. By logging and clearing of lands
for agriculture man has considerably speeded the invasion of the
southern mice. Slight differentiation of the short-tailed mice north
of the Columbia River {austerus) separates them from their parent
race, rubidus.

In the dense forests along the Pacific Coast of Washington, aus-
terus did not become established. This area includes most of the
land west of Puget Sound. There is a narrow band of austerus
that extends along the eastern and part of the northern edge of the
Olympic Peninsula, where they have probably invaded in relatively
recent time.

On the Oregon side of the Columbia River the range of rubidus
is continuous from the Pacific to the Cascades. On the Washington
side of the river, oreas extends from the Pacific to eastern Wahkia-
kum County, where the range of austerus starts. At the border of

96 University of Kansas Publs., Mus. Nat. Hist.

the ranges of the two subspecies, ecological conditions determine
the ranges; oreas occurs in the Douglas fir and hemlock forests and
austerus occurs in the jungles of alder and salmonberry in the
stream valleys. The range of austerus extends from eastern Wah-
kiakum County east along the Columbia, to western Klickitat
County. In a north-south strip across the Columbia, through the
ranges of rubidus in Oregon and austerus in Washington, normal
intergradation is apparent. By gradual degrees rubidus changes to
austerus. In a north-south strip farther west, through the ranges of
rubidus and oreas, the same transition of the Oregon rubidus is seen,
namely an approach toward austerus. The cline is, of course,
abruptly terminated by the range of the dissimilar oreas.

On Puget Island, a small island lying in the Columbia River in
western Wahkiakum County and politically within the state of
Washington (see Scheffer, 1942) a population of deer mice occurs
that resembles rubidus more closely than austerus. This island lies
in the river between the ranges of rubidus and oreas. The lack of
intergradation between these two subspecies has probably kept the
Puget Island population pure rubidus. Puget Island is a junglelike
lowland locally known as "tideland."

The San Juan Islands of northern Puget Sound were thoroughly
glaciated in Wisconsin Time and probably were under thousands of
feet of ice when the Vashon Glacier was at its maximum extent. The
subspecies of deer mouse occurring there now is hollistcri, a race
similar to oreas in color, body size and cranial characters and prob-
ably derived from oreas. P. m. hollisteri differs from both oreas and
austerus in its much shorter tail. Shortness of tail is apparently
a product of insularity in northwest coastal mice. The transition
from long-tailed mainland mice to short-tailed island mice is shown
by Hall (1938A: 4(H). When areas first occupied the area about
Puget Sound (this area is occupied by austerus today) some indi-
viduals probably reached the San Juan Islands soon thereafter.
Contact between oreas and hollistcri is now prevented by the pres-
ence of austerus between their ranges. Of course, a water barrier
separates austerus from hollistcri but austerus does occur in pure
form on some islands in Puget Sound I not on any of the San Juan
Islands), for instance, on Whidby Island. One wonders why aus-
terus has not established itself on some of the San Juan Islands but
considering the degree of difference between hollisteri and austerus,
I doubt that the two would interbreed if they did occur together.
On Vancouver Island, British Columbia, a short-tailed, austerus-

Distributional History 97

type of mouse occurs in the lowlands and a long-tailed, areas-type
of mouse in the mountains. Vancouver Island is a large island
(16,400 square miles). Apparently a stock of austerus from the
mainland reached Vancouver Island and were able to find suitable
habitat and compete with and drive out the established oreas in the
manner that they probably did in the western Washington lowlands.
The large Vancouver Island offers far more variety of habitat and
opportunity for establishment of emigrating mice than do the
smaller San Juan Islands, the largest of which is Orcas (57 square
miles) .

Following the withdrawal of the Wisconsin ice from eastern
Washington, numerous minor movements and adjustments of deer
mice seem to have taken place. Long-tailed, oreas-type of mice
were exterminated on the Columbian Plateau if, indeed, they eVex
became established there. Long-tailed mice did apparently become
established in driftless areas and forested areas to the south of the
drift border. With the establishment of soil and forests on the de-
glaciated land, the short-tailed gambelii and the long-tailed mice
became thoroughly intermixed. In some areas, especially near the
type locality of artemisiae , the gambelii characters of the mixture
predominate at the present time. Deer mice from the Okanogan
Valley differ from gambelii only in slightly larger size and darker
color. In other areas, notably near Metaline, Pend Oreille County,
•the oreas characters of the mixture dominate at the present time.
Mice from here are large and red and differ from typical oreas in
having relatively short tails. Other less marked segregations of
characters are numerous, in Idaho and British Columbia as well as
in Washington. Cowan (1937) has described Peromyscus m. al-
pinus, isolated in a mountainous area of British Columbia. This
seems to be a form derived from oreas-type stock.

With the reestablishment of soil and forests on the deglaciated
land of British Columbia north of Washington, a northward ex-
tension of the gambelii-oreas mixture occurred. Another invasion
was taking place at this time, however. Peromyscus m. arcticus
spread onto the deglaciated land from the north or east, ahead of
artemisiae (the gambelii-oreas mixture). This new invader inter-
graded with artemisiae as well as with oreas. Further collecting and
studies are required in this area before the relationships of the three
subspecies can be completely understood.

If geographic ranges are assigned to the named subspecies of deer
mice that occur in Washington, it must, in part, be done arbitrarily.
7—4993

98 University of Kansas Publs., Mis. Nat. Hist.

The deer mice of the San Juan Islands are all referred to the sub-
species hollisteri. Only Puget Island, Wahkiakum County, is con-
sidered to be inhabitated by rubidus. The boundary between the
ranges of austerus and oreas is sharply defined and further col-
lecting by resident naturalists should result in detailed mapping
of the boundary.

The ranges and distribution of the deer mice of eastern "Washing-
ton are less clearly defined than those of western Washington.
Peromyscus m. gambelii exists in its purest state on the Columbian
Plateau. Here the Columbia River makes a convenient boundary
to the west. Pure Peromyscus m. oreas exist in the Cascade Moun-
tains. In the area between the Columbian Plateau and the Cascades
the deer mice are variously intermediate between gambelii and oreas
and, over most of the area, are nearest to gambelii. This might
be considered a natural area of intergradation such as commonly
occurs between the ranges of subspecies. These intergrades, how-
ever, are similar to Peromyscus m. artemisiae and the area occupied
by these intermediates is connected on the north with the range
of artemisiae. Thus Osgood (1909: 61) regards specimens from
Easton and Wawawai, in the area of intergradation, as artemisiae
or intergrades between gambelii and artemisiae. Taylor and Shaw
(1929: 22) include the entire area of intergradation in the range of
artemisiae. This treatment, although convenient to the student
of geographic variation, is apt to conceal the evolutionary and
historical influences. These influences may be appreciated better if
the intergrades from each locality arc referred to the subspecies they
most closely resemble.

The specimens upon which the name artemisiae was based came
from a narrow tongue of zonally lower country that extends north-
ward from the range of the lighter-colored gambelii. As might be
expected, topotypes are lighter in color than specimens from the
north, west and east. Nevertheless, the type locality is within a
geographic area that is occupied principally by a darker race,
artemisiae, to the north of gambelii. The topotypes of arti misiae
may be considered to be intermediate between gambelii and the
darker, northern race. Thus the name artemisiae becomes avail-
able for the mice of tin' general area in question. The mice of the
area immediately to the east of the type locality, in Washington and
presumably in British Columbia and Idaho also, are essentially a
mixture of the subspecies gambelii and a now mostly extinct and
unnamed race that probably resembled oreas. Local variations

Distributional History 99

of populations from within this area are extreme but some segregation
of color and length of tail has taken place. Mice from mountainous
areas resemble oreas while mice from the lowlands resemble gambelii,
or, more exactly, mice from coniferous forests resemble oreas while
mice from other areas resemble gambelii.

South of the area of racial mixture in northeastern Washing-
ton, in the Blue Mountains of southeastern Washington, the deer
mice are like gambelii except in slightly darker color. Mice from
the Blue Mountains are darker and browner than gambelii, not more
reddish. There is no indication of adulteration with orcas stock.
Since the Blue Mountains are a forested area and are far south of the
drift border, we suppose that deer mice existed there through the
last glacial period and that their dark color is an adaptation to forest
habitat.

Mice similar to those of the Blue Mountains have an extensive
range in Idaho (Davis 1939: 290). These mice have relatively
uniform racial characters and constitute a "good" subspecies.

At present the deer mice of northern Washington, southeastern
British Columbia, northern Idaho, northeastern Oregon, western
Montana and northwestern Wyoming are called artemisiae. The
mice of this extensive area are, however, of two genetic types: that
type with mixed racial characters that lives in northeastern Washing-
ton and probably also farther east along the Wisconsin drift border,
and that type that occurs farther south in Idaho and seems to con-
stitute a stable subspecies.

The separation of these two types may be desirable. Detailed
study of the deer mice from the area now assigned to the range of
artemisiae may show that the name subarcticus (Peromyscus tex-
anus subarcticus Allen, 1899) is applicable to the southern form. The
type with mixed racial characters must be called artemisiae. If the
two types are eventually separated, the mice from the Blue Moun-
tains of Washington will be referable to the southern form.

Xeotoma ctnerea. — The wood rat found over most of Washing-
ton, Xeotoma c. occidentalis, probably entered the state from eastern
Oregon early in the Recent and spread over most of the state. The
wood rat of the Blue Mountains {alticola) probably developed from
occidentalis.

A coastal race of the wood rat (fusca) occurs in western Oregon.
This race, if it occurred in western Washington in pre-Vashon Time,
was eliminated in Vashon Time or subsequently. In Oregon it lives in
deep forests (Bailey, 1936: 171). In Washington occidentalis occupies

100 University of Kansas Publs., Mus. Nat. Hist.

but a small part of the ecologic niche occupied by fusca in Oregon.
Elimination of fusca from Washington through competition with
occidentalis seems highly improbable. Should fusca ever cross the
Columbia River and become established it would probably spread
to a considerable part of western Washington.

Synaptomys borealis. — The lemming mouse seems now to be
retreating northward and was an arctic species forced southward
by the Vashon-Wisconsin glaciers. Unlike other alpine species, it
seems to be unable to exist for long in isolated mountain areas.

Phenacomys intermedius. — The heather vole, like the lemming
mouse, probably was forced southward by the Vashon-Wisconsin
glaciers. In the Cascade and Rocky mountains it found suitable
habitat and spread southward to almost the lower end of the Sierra
Nevada in California. The Cascade race moved eastward on the
Puget Bridge to the Olympic Mountains after the retreat of the
Vashon ice, and northward in the Cascades. Northeastern Wash-
ington was reinvaded by the subspecies of the Rocky Mountain
Fauna.

Clethrionomys gapperi and californicus. — The two species of
red-backed mouse found in Washington were probably distinct in
pre-Vashon Times. At the maximum extent of the Vashon ice,
californicus was probably found in western Oregon and gapperi in
the Blue Mountains, where idahoensis was developed, and in the
southern Cascades (cascadensis). During or shortly after Vashon
Time, gapperi crossed the Puget Bridge to become established in
the Olympic Mountains. After the retreat of the ice, gapperi moved
northward and eastward from the Cascades and californicus crossed
to western Washington from Oregon and displaced gapperi from the
lowlands.

Hinton (1926: 215) separated the American Clethrionomys into
three groups, suggesting that Old World counterparts of each group
exist. Davis (lD.'W: 308) gives an excellent analysis of the emigra-
tional history of the American species, and points out the close re-
lationship of the Siberian and Arctic-American forms. He suggests
that the rutilus group invaded Arctic- America from Siberia at the
close of the last glaciation.

Two other groups are recognized by Davis, who assumes that
both arose from a common ancestral stock and divided into two
stems, one of which (gapperi group! followed the main Rocky Moun-
tain course southward and worked eastward across Canada while
the other {californicus group) followed down the Pacific Coast.

Distributional History 101

Davis states, "This southward extension of range in America prob-
ably took place in the Pleistocene, but almost certainly the present
southern range of the genus in the West was not reached until late
in, or after, that period."

Our studies of the Clethrionomys of Washington indicate the
essential correctness of Davis' conclusions. The separation of the
two groups, however, probably was caused by southward moving
glaciers and the separation of the mice into two stocks closely
paralleled the divergence of other groups, such as that of the snow-
shoe rabbits. The two groups of red-backed mice are more sharply
separated than was thought by Davis. The intermediate color of
the back of C. g. idahoensis, mentioned by Davis, seems to be merely
coincidental. The essential difference in the two groups is the
sharply marked red band of the gapperi group as opposed to the
general red area on the dorsal surface of the californicus group.
Clethrionomys gapperi nivarius has been considered a derivative of
occidentalis because the range of the latter surrounds the Olympic
Mountains, where nivarius is found. Considering the immediate
post-Pleistocene movements of mammals from the Cascades to the
Olympic Mountains, so clearly illustrated in Thomomys, Tamias
amoenus, and Phenacomys, a means is indicated by which Cle-
thrionomys of the gapperi group might have reached the Olympics.
The apparent lack of intergradation between occidentalis and
nivarius gives proof of their relatively distant relationship. C.
nivarius seems not to have been derived from occidentalis, and
apparently does not belong to the californicus group. It belongs
instead to the gapperi group, and I consider it to be a subspecies of
gapperi. In no sense is it intermediate between the two groups. The
other form considered by Davis to be intermediate between the
two groups is caurinus. This mouse has not yet been taken in Wash-
ington, although it may eventually be found on Point Roberts, on
the Fraser River delta. Its distribution is paralleled by that of
many other mammals that are definitely not of the Pacific Coastal
Fauna.

The californicus group, I feel, contains only the races of Cle-
thrionomys californicus, while the gapperi group contains C. gapperi
and its races, including caurinus, and possible other species.

Possibly intergradation occurs between C. californicus occidentalis
and C. gapperi. Nevertheless, I fail to find evidence of such inter-
gradation. I have taken C. gapperi nivarius and C. californicus
occidentalis within ten miles of each other, but each retained its

102 University of Kansas Publs., Mis. Nat. Hist.

distinctive characters with no evidence of intergradation. In the
case of C. gapperi saturates and C. calif ornicus occidentalis the
proof is less conclusive. In spite of numerous attempts to trap
Clethrionomys in the area geographically intermediate between their
ranges, I have taken none. Though common along the coast, occi-
dentalis becomes progressively scarcer to the east, being rare in the
vicinity of Seattle and apparently absent from the western base of
the Cascades. So far as is known, the ranges of caurinus and occi-
denta 7 is are separated by the Fraser River.

Microtus pennsylvanicus and montanus. — The Pennsylvania
meadow mouse is closely related to Microtus montanus. Certain
races of montanus, notably those from southern Oregon, California,
and northern Nevada, closely resemble pennsylvanicus externally
and cranially. From the central part of its range northward, mon-
tanus becomes progressively less like pennsylvanicus. The races
nanus and caniCaudus are quite different from pennsylvanicus both
externally and cranially, and in addition the anterior loop of the
second molar is less constricted; often it is not constricted at all.
Microtus pennsylvanicus and Microtus montanus occur together over
parts of the northern Rocky Mountains. Where the two species
c;ime together, pennsylvanicus occurs with races of montanus thai
are most unlike it.

Microtus pennsylvanicus kincaidi closely resembles races of Mi-
crotus montanus that occur in southern Oregon, California, and Ne-
vada. It is larger, darker, and longer-furred than Microtus pennsyl-
van'cus funebris from Washington. Seemingly pennsylvanicus and
montanus diverged previous to Vashon-Wisconsin Time. The stock
that gave rise to montanus spread over the Great Basin while penn-
sylvanicus ranged farther east. Some of the montanus stock worked
northward in the Rocky Mountains. Microtus pennsylvanicus had
meanwhile moved westward to the Rocky Mountains. The two
stocks met and behaved as full species.

Microtus pennsylvanicus probably occupied northeastern Wash-
ington in the interglacial cycle preceding the Recent. The advance
of the Wisconsin ice eliminated most of these mice. The glacier
dammed the Columbia River and caused it to turn southward from
its basalt-marginal course and take a path over the Columbian
Plateau. Along this glacial river a population of Microtus persisted
to become kincaidi.

While the Wisconsin ice was at its maximum extent, Microtus
montanv* from the Blue Mountains crossed the Simcoe Bridge to the

Distributional History 103

Yakima Valley and the eastern Cascade Mountains. The closing
of the Simcoe Bridge isolated these mice, wjhich subsequently became
slightly differentiated, as canescens. Another stock moved westward
along the Columbia River to western Oregon. This stock is now
called Microtus canicaudus but is probably racially rather than
specifically distinct from Microtus montanus nanus.

With the retreat of the Wisconsin ice, montanus extended its
range northward from the Yakima Valley along the eastern Cas-
cade Mountains to extreme southern British Columbia. Microtus
pennsylvanicus funebris entered northeastern Washington with other
elements of the Rocky Mountain Fauna.

At present, pennsylvanicus occurs in northeastern Washington
while 7nontanus is found in southeastern Washington and the eastern
Cascade Mountains. The Okanogan River Valley separates their
ranges. Eventually montanus may extend its range to northeastern
Washington and pennsylvanicus to the Cascades, the two forms oc-
curring together as they do in Montana, Idaho and Colorado. The
isolation of kincaidi on the Columbian Plateau seems complete and
the probability of its range reaching that of its related species seems
slight.

Microtus loxgicaudus. — The later distributional history of the
long-tailed meadow mouse is not yet clear.

Microtis townsendii. — This Pacific Coastal species probably
lived west of the Sierra Nevada-Cascade Mountains since the early
Pleistocene. It has extended its range northward since the retreat
of the Vashon ice and has reached some of the islands in Puget
Sound and the Strait of Georgia. This species seems to be related
to Microtus richardsoni but the two species probably separated at
an early time.

Microtus richardsoni. — The water rat of the Cascade Moun-
tains seems to be have been forced southward into Washington and
Oregon by the descent of the Vashon-W T isconsin glaciers where it
became isolated from the water rat of the northern Rocky Moun-
tains. After the retreat of the ice, both forms have moved north-
ward. Bailey (1900) records macropus from Bonner County, Idaho,
not far from northeastern Washington but in Washington up to this
time, macropus has been recorded only from the Blue Mountains of
the southeastern part of the state.

Microtus oregoni. — The later historical distribution of the
creeping mouse was probably the same as that of townsendii.

104 University of Kansas Publs., Mus. Nat. Hist.

Lagurus curtatus. — The distribution of this species of the Great
Basin Fauna has probably changed little if any since the late Pleis-
tocene.

Ondatra zibethicus. — The distribution of the two races of musk-
rat that occur in Washington is almost exactly that of the beavers.
Probably one form was isolated in southwestern Washington and
northwestern Oregon during Vashon Time and another, more adapt-
able, race occurred in eastern Washington. After the retreat of
the ice the adaptable race spread widely but the muskrats of the
lower Columbia River changed their range little or not at all.

Aplodontia rufa. — The race of Aplodontia found in the Cascades
of Washington was probably confined to the southern Cascades in
Vashon Time and has since spread northward to reoccupy the range
as far north as southern British Columbia.

The presence of a mountain beaver in western Washington that
is indistinguishable from the race rufa, found in the Cascades of
Oregon, is most surprising. In Oregon, pacifica occurs in the western
lowlands and rufa in the mountains to the east. In Washington
rufa occurs in the western lowlands and rainieri in the mountains to
the east.

The offset in range of rufa seems best explained by assuming
that all of western Washington was once occupied by mountain
beavers (rufa) that spread from the Cascades of Oregon to the
Cascades of Washington and thence to the western Washington
lowlands. This must have occurred early in the interglacial cycle
preceding the Vashon Glaciation. While confined to the southern
Cascades, rainieri developed the larger size that now separates it
from the ancestral rufa that occurs to the west and south of it.

Zapus princeps. — The coastal race of the jumping mouse was
probably distinct from the more inland population before Vashon-
Wlseonsin Times. The differences between them were probably
accentuated while the Vashon-Wisconsin glaciers separated their
ranges. The western race (trinotatus) was isolated in the southern
Cascades and southwestern Washington. After the retreat of the
glaciers, trinotatus moved northward through western Washington
and the Cascades into British Columbia. The race kootenayemis
of the adjacent Rocky Mountains moved eastward through north-
eastern Washington to the Cascades. The race oregonus, found in
the Blue Mountains, has probably been resident there since it
evolved from the populations to the east.

f

Distributional History 105

Erethizon dorsatum. — I have inadequate basis for speculation
concerning the historical distribution of the porcupine.

Ochotona princeps. — The descent of the Wisconsin ice separated
the western pikas into two populations. One was confined to the
Cascade-Sierra Nevada system and another to the Rocky Moun-
tains. Two races are now found in the Cascades of Washington.
One, brunnescens, inhabits the higher Cascades and another, fenisex,
the eastern edge of the Cascades. The range of brunnescens extends
southward into Oregon while the range of fenisex extends farther
north in British Columbia than does that of brunnescens. The
principal difference between the two races is the smaller size and
paler color of fenisex. In Washington, fenisex occupies a slightly
more arid habitat than brunnescens. Further, fenisex lives in talus
principally of basaltic rock while brunnescens lives in talus of
granitic rock. The basaltic talus is more finely fractured, offering
smaller crevices in which pikas can conceal themselves. The granitic
rock, on the other hand, forms talus composed of fragments of large
size. Freshly fractured granite is pale, whitish gray. After weather-
ing, however, it becomes blackish as the more soluble, pale feldspars
are removed, leaving the black hornblend and biotite exposed.
Freshly fractured Columbian basalt is blackish but, after weather-
ing, becomes rusty, reddish brown.

In Washington the two races maintain their distinctiveness be-
cause selective factors in the basalt talus of the eastern Cascades
favor the smaller size and paler color of fenisex while in the higher
Cascades, selective factors in the granitic batholith favor larger
size and darker color.

The present range of brunnescens in Oregon indicates that this
race was the pre-Wisconsin resident of the Cascades of at least
southern Washington. Probably fenisex evolved in the arid interior
of British Columbia. In color fenisex is intermediate between the
dark brunnescens and the pale cuppes of the western spurs of the
Rocky Mountains. Probably fenisex was forced southward into
the eastern Cascades by the Wisconsin ice and inhabited the area
east of the range of brunnescens. This eastern area was not occu-
pied by brunnescens because the basaltic talus was of small size.
After the retreat of the ice, fenisex (adapted to more arid habitat than
brunnescens) , spread northward to the relatively dry valleys of the
interior of British Columbia.

The distinctness of brunnescens and fenisex is maintained by se-
lective factors of the habitats they occupy but the zone of inter-

10(5 University of Kansas Publs., Mrs. Nat. Hist.

gradation between the two is broad. A tendency towards paleness
is noticeable in specimens from as far west as the eastern side of
Mount Rainier.

The pikas in the Rocky Mountain area have given rise to a num-
ber of races. The pale, small race of northern Idaho, cuppes, en-
tered northeastern Washington with other members of the Great
Basin Fauna after the retreat of the ice.

Lepus townsendii.— The white-tailed jack rabbit has probably
lived on the Columbian Plateau ever since or even before the time
when the Wisconsin ice bordered the plateau on the north.

Lepus americanus. — Snowshoe rabbits were probably spread over
forested parts of Washington in pre-Wisconsin time. A coastal race.
washingtonii, was probably distinct from the remainder of the
species at an early time. It, and its kindred races klamathensis and
tahoensis, are very different from races derived later from the Rocky
Mountain Fauna. From this it is deduced that washingtonii was a
member of the Pacific Coastal Fauna before the advance of the last
continental glaciers.

The snowshoe rabbits now found in the Cascade Mountains of
Washington are racially distinct from populations occurring to the
north and east, but are more closely related to the northern and
eastern rabbits than they are to washingtonii. This race, cascadensis,
probably lived in the area north of the state of Washington in pre-
Wisconsin interglacial time. The snowshoe rabbit of northeastern
Washington was probably a race having an extensive range in the
Rocky Mountains from which pineus in the Blue Mountains de-
veloped. The descent of the Vashon-Wisconsin glaciers found wash-
ingtonii restricted to western Oregon. It might have persisted also
in southwestern Washington but, had this been the case, we would
expect a broader zone of intergradation between washingtonii anil
cascadensis than actually exists.

The glaciers forced cascadensis southward into the Cascades of
southern Washington. The difference between washingtonii and
cascadensis and the narrow zone of intergradation between them,
indicate that the two races were not in contact in pre-Vashon Time.
The presence in the Cascades of Oregon of a race related to wash-
ingtonii rather than cascadensis also supports this view. The snow-
shoe rabbits of northeastern Washington were probably eliminated
from areas where the ice was in place. The effect of the glaciers
in nearby areas was probably less on snowshoe rabbits than upon
ot her species.

Distributional History 107

Following the retreat of the glaciers, washingtonii reinvaded west-
ern Washington and spread northward through the lowlands to the
Fraser River. In the Cascades, cascadensis moved even farther
north. L. a. pineus of the Rocky Mountain Fauna invaded north-
eastern Washington from adjacent areas to the south. Another
race, columbiensis, spread from the interior valleys of British Co-
lumbia to the western part of northeastern Washington in probably
relatively recent times. The post-W 7 isconsin history of the distribu-
tion of this race is uncertain.

Lepus californicus. — The black-tailed jack rabbit occurs
throughout the sagebrush areas of eastern Washington, except in the
Okanogan Valley. It invaded Washington from the south recently
(Couch, 1927: 313). The first black-tails were thought to have
entered Walla Walla County about 1870. Here they were checked
by the Snake River. About 1905 they crossed the Snake River, on
ice. and by 1920 had spread over much of the Columbian Plateau.
In January, 1920, they crossed the Columbia River in two places,
on ice jams, and by 1927 had spread north and west over the sage-
brush areas between the Columbia River and the Cascade Moun-
tains. When Couch, in 1927, published his account, he predicted
that the black-tail would eventually spread to the Okanogan Valley.
This has not yet occurred although it is still to be expected.

It should be noted that the migrational history of the black-
tailed jack rabbit is known only because the animal is large, spec-
tacular, and diurnal, and therefore noticed by resident ranchers,
and because the facts came to the notice of a competent biologist.
Migration of a less conspicuous mammal, for example, a mouse,
would rarely be detected.

Sylvilagus nuttallii. — The cottontail might have been resident
on the Columbian Plateau during Wisconsin Time but probably it
invaded, or perhaps reinvaded, the Plateau from eastern Oregon
in the Recent.

Sylvilagus idahoensis. — Too little is known about this rabbit
in Washington to hazard a guess as to its recent distributional his-
tory.

Cervus canadensis. — The elk emigrated from Siberia to North
America in the late Pleistocene but previous, certainly, to Vashon-
Wisconsin time. Probably the coastal race was distinct from the
interior population in the preceding interglacial cycle. The descent
of the last continental glaciers, in Washington, separated the elk

108 University of Kansas Publs., Mus. Nat. Hist.

of the Pacific Coastal Fauna from the Rocky Mountain Fauna.
After the retreat of the ice the coastal race moved northward to
southern British Columbia and thje Rocky Mountain race moved
northward and westward through British Columbia. Northeastern
Washington was reoccupied by the Rocky Mountain elk.

Remains of elk have been found associated with human artifacts
on the northern part of the Columbian Plateau. The plateau is
poorly suited to elk but a few may have persisted there until late
historic time. The remains may have been brought by Indians
from northeastern Washington. The remains included teeth and
portions of the skull and it seems unlikely that these would have
been carried any great distance.

Odocoileus hemionus. — The American deer probably evolved
in North America. In this respect they are unlike the elk, moose
and caribou, all of which emigrated from Asia to America in the
Pleistocene. Odocoileus hemionus and virginiamis probably di-
verged from a common stock in the Pliocene.

The black-tailed deer was probably a member of the Pacific
Coastal Fauna at an early time and distinct from the mule deer
before the last interglacial cycle. Probably the mule deer lived in
the eastern Cascades and on part of the Columbian Plateau in Wis-
consin Time. They occur on the plateau in limited numbers at
present and their remains have been found associated with human
artifacts in the Grand Coulee area.

Odocoileus virginianus. — The white-tailed deer ranges from the
Atlantic to the Pacific and from Canada to Mexico. The species
was probably abundant in the Pacific Northwest in the Pleistocene,
perhaps in Vashon-W T isconsin time, but has since largely given way
to the black-tailed and mule deer. The race 0. v. leucurus now occu-
pies an extremely small range. The accounts of early naturalists
indicate that it was more abundant and had a wider range 100 years
ago. Probably leucurus was once an important member of the Pa-
cific Coastal Fauna.

The inland race, ochrourm, is a typical member of the Rocky
Mountain Fauna. It and leucurus were probably derived from a
common ancestor in the late Pleistocene.

Alce americanus. — The moose occurs in Washington as a casual
wanderer from the Rocky Mountain Fauna to the east.

Rangifer montanus. — The caribou occurs in northeastern Wash-
ington as a winter migrant from the north. It was probably of more
regular and extensive occurrence in the past.

Distributional History 109

Antilocapra americaxa. — The antelope probably was a casual
wanderer to southeastern Washington and perhaps to the Columbian
Plateau from eastern Oregon before white man reached Washington.

Bison bison. — The buffalo, like the antelope, probably occurred
in southeastern Washington and on the Columbian Plateau only as
a casual wanderer from Oregon in postglacial time.

Ovis canadensis. — Mountain sheep reached North America from
Asia in the Pleistocene. By Vashon-Wisconsin Time they had
spread southward, perhaps to Mexico. The descent of the Vashon-
Wisconsin ice in Washington presumably separated the canadensis
and californiana type of sheep, the former being confined to the
Rocky Mountain Fauna and the latter to the Cascade-Sierra Nevada
chain and adjacent parts of the Great Basin.

Mountain sheep probably persisted in the southern Cascades of
Washington and on the Columbian Plateau during Vashon-Wiscon-
sin times. After the retreat of the ice, these sheep (californiana)
moved northward slightly, both in the Cascades and on the Colum-
bian Plateau. Remains of sheep are to be found with human arti-
facts in the Grand Coulee area today. When white man first reached
the state, sheep existed throughout the Cascades and on part of the
Columbian Plateau. Only a pitiful remnant remains in the extreme
northern Cascades.

Mountain sheep of the Rocky Mountain race probably existed in
small numbers in the Blue Mountains of southeastern Washington
until historic times. Sheep from the Blue Mountains or adjacent
Idaho invaded northeastern Washington shortly after that region
was freed from Wisconsin ice. These sheep were exterminated
shortly after the coming of the white man.

Oreamnos americanus. — Fossil remains of the mountain goat
have been discovered at W T ashtuckna Lake, associated with those
of the lion, horse, and camel. Probably the mountain goat had an
extensive range in Washington at the time of a glacial advance pre-
vious to the Wisconsin glaciation. In Wisconsin Time the mountain
goat was confined to the southern Cascades. Strangely enough, it
seems not to have crossed the Columbia River to become established
in the Cascades of Oregon. After the retreat of the ice it moved
northward through the Cascades.

A mountain goat recently taken in northeastern Washington
seemingly wandered to the area from Idaho.

110 University of Kansas Publs., Mrs. Nat. Hist.

EXPLANATION OF TREATMENT

The order of arrangement of the following accounts is that of Miller (1924)
with some modifications. For example, the Chiroptera are arranged according
to Tate (1942). the Sciuridae according to Bryant (1945) and the Cetacea ac-
cording to Scheffer (1942). A few other minor changes are included.

Although the principal purpose of this report is to describe and interpret
the distribution of the various species and races of mammals that occur in
Washington, a brief description and account of the habits of the animals is
included. Each species account begins with a description, based principally on
external characters. This is followed by pertinent information regarding the
range of the species, its relationships within the genus and a brief account of
its habits. The habits are dealt with in most detail for the species that arc
of greatest economic importance.

The accounts of subspecies are largely technical. The account of the type
specimen is rather complete. Subspecific diagnoses are brief, stating often only
the principal racial characters. Standard external measurements arc usually
given for each subspecies. All measurements, unless otherwise stated, arc in
millimeters and weights are in grams. Specimens are adult, unless otherswise
stated.

No formal list of specimens examined is included. Localities from which
specimens have been examined are usually shown on the distribution maps
by solid circles. Open circles indicate localities from which specimens, not
examined by me. have been recorded by other persons. Most of these records
arc published but a few are based on authentic trappers' reports, photographs,
or other evidence. Unusual occurrences of animals outside their natural ranges
are not. shown on the maps. Not every record from well within the range
of a subspecies has been plotted, but care has been taken to plot all records
ol occurrences from peripheral areas.

When more than one race of a species occurs in Washington, specimens
from localities where the geographic range of one subspecies meets or ap-
proaches that of another are listed as "marginal occurrences" in the paragraph
on the distribution of the race to which they are best referred. Marginal oc-
currences are listed from north to south and from west to east. The authority
for tin- record, if published, is given. When the record i- based on specimens
examined by the author, the collection containing the specimen is indicated
as follows:

(E.S.B.) Kriiest S. Booth collection. College Place,, Washington.

(J.M.E.) .1. M. Edson collection. Bellingham, Washington.

(K.r.) University of Kansas. Museum of Natural History, Lawrence, Kansas.

(M.V.Z.) Museum of Vertebrate Zoology, University of California, Berkeley,
( lalifornia.

(CSX M.) United Slates National Museum, Washington, I). C. Specimens
from the Biological Surveys Collection are included here.

(V.B.S.) Victor B. Scheffer collection. United States National Museum, Wash-
ington, D. C.

Explanation of Treatment 111

(W.S.C.) Washington State College, Charles R. Conner Museum. Pullman,

Washington.
(W.S.M.) Washington State Museum, University of Washington, Seattle,

Washington.
(W.W.D.) Walter W. Dalquest collection, Seattle, Washington.

In all, 230 kinds (species and subspecies) of mammals are listed
for the state of Washington. Of these, f) kinds (opossum, eastern
gray squirrel, fox squirrel, 3 kinds of Old World rats, house mouse,
nutria, and eastern cottontail) have been introduced into Wash-
ington from elsewhere. The dog of the Indians might be listed as
a tenth introduced species. The grizzly bear, the sea otter, the wolf,
the moose, one race of mountain sheep, one race of mountain goat,
and. if they ever occurred in the state, the pronghorn antelope and
bison, are nearly or completely extinct in Washington. Mammals
other than cetaceans possibly occurring in Washington, but of which
satisfactory record is lacking, are mentioned in a hypothetical list
at the end of the accounts of species. Of the 220 kinds of native
mammals, known to occur in the state, 23 are marine (4 pinnipeds,
19 cetaceans). The 197 living, native, land mammals include 6 or-
ders, 20 families, 58 genera and 101 full species.

112

University of Kansas Publs., Mus. Nat. Hist.

CHECK LIST OF MAMMALS

CLASS MAMMALIA— mammals
Order MARSUPALIA— marsupials

Family Didelphiidae — opossums
Genus Didelphis Linnaeus — opossums
Didelphis virginiana virginiana Kerr

opossum

Order INSECTIVORA— insectivores

Family Talpidae — moles

Genus Neurotrichus Gunther — shrew-mole

Neurotrichus gibbsii gibbsii (Baird)
Neurotrichus gibbsii minor Dalquest
and Burgner

Genus Scapanus Pomel — western American
Scapanus townsendii (Bachman)
Scajmnus orarius orarius True
Scapanus orarius yakimcnsis Dalquest

and Scheffer
Scapanus orarius schefferi Jackson

Family Soricidae — shrews

Genus Sorex Linnaeus — long-tailed shrews
Sorex cinereus cinereus Kerr
Sorex cinereus streatori Merriam
Sorex merriami merriami Dobson

Sorex trowbridgii trowbridgii Baird
Sorex trowbridgii destructioni Scheffer

and Dalquest
Sorex vagrans vagrans Baird
Sorex vagrans monticola Merriam

Sorex obscurus obscurus Merriam
Sorex obscurus setosus Elliot

Sorex palustris navigator (Baird)
Sorex bendirii bendirii (Merriam)
Sorex bendirii albiventcr Merriam

Genus Microsorex Coues — pigmy shrew
.\ficrosorex hoyi washing toni Jackson

[Gibbs shrew-mole

moles
Townsend mole

coast mole

rcinereous shrew
Merriam shrew

^Trowbridge shrew

wandering shrew

[dusky shrew
mountain water-shrew
J-Bendire water-shrew

pigmy shrew

Order CHIROPTERA— bats
Family Vespertilionidae — vespertilionid bats

Genus Myotis Kaup — mouse-eared bats
My otis lucijugus carissima Thomas
Myotis lucijugus alascensis Miller

tig myotis

M yotis yumanensis sociabilis
H. W. Grinnell

Myotis i/ununii iisis sat unit us Miller

^Yuma myotis

Check List

118

long-eared myotis
fringe-tailed myotis
[hairy-winged myotis

[■California myotis

Myotis keenii keenii (Merriam) Keen myotis

Myotis ewtis evolis (H. Allen) ]

Myotis evotis pacificus Dalquest /

M yotis thysanodes thysanodes Miller
Myotis volans longicrus (True)
Myotis volans interior Miller
Myotis calijornicus caurinus Miller
Myotis calijornicus calijornicus

(Audubon and Bachman)
Myotic subulatus melanorhinus (Merriam) small-footed myotis

Genus Lasionycteris Peters — silver-haired bat

Lasionycteris noctivagans (Le Conte) silver-haired bat

Genus Corynorhinus H. Allen — long-eared bats

C orynorhinus rafinesquii townsendii
(Cooper)

Corynorhinus rafinesquii intermedins
H. W. Grinnell
Genus Pipistrellus Kaup — pipistrelles

Pipistrellus hesperus hesperus (H. Allen)
Genus Eptesicus Rafinesque — serotine bats

Eptesicus juscus bernardinus Rhoads
Genus Lasiurus Gray — hairy-tailed bats

Lasiurus cinereus cinereus (Beauvois) hoary bat

Genus Antrozous H. Allen — nyctophiline bats

long-eared bat

western pipistrelle

big brown bat

Antrozous pallidus cantwelli Bailey

pallid bat

Order CARNIVORA

Family Ursidae — bears
Genus Ursus Linnaeus — bears

Ursus americanus altifrontalis Elliot
Ursus americanus cinnamomum

(Audubon and Bachman)
Ursus chelan Merriam

[bk

ack bear

J

grizzly bear

Family Procyonidae — raccoons and allies
Genus Procyon Storr — raccoons
Procyon lotor psora Gray

Procyon lotor excelsus Nelson and J-raccoon

Goldman J

Family Mustelidae — weasels and allies

Genus Martes Pinel — martens and fisher

Martes caurina caurina (Merriam) 1 , ,

,j . • • /t>u j \ r w estern marten

Martes caurina ongenes (Rhoads) J

Martes pennanti (Erxleben) fisher

Genus Mustela Linnaeus — weasels, ferrets and minks

M us tela erminea invicta Hall
Mustela erminea jallenda Hall
Mustela erminea olympica Hall
Mustela erminea streatori (Merriam)
Mustela erminea gulosa Hall
Mustela erminea murica (Bangs)

8—4993

ermine

114 University of Kansas Plbls., Mus. Nat. Hist.

■Ion«i-tailed weasel

Mustela jrenata washingtoni (Merriam)

Mustela jrenata altijrontalis Hall

Mustela jrenata ejjcra Hall

Mustela jrenata nevadensis Hall

Mush In vison energumenos (Bangs) mink

Genus Gulo Pallas — wolverines

Gulo luscus luteus Elliot wolverine

Genus Lutra Brisson— river otters

Lutra canadensis pacifica Rhoads "Liver otter

Lutra canadensis vancouvercnsis Goldman /

Genus Enhydra Fleming — sea otter

Enhydra lutris nereis (Merriam) sea otter

Genus Spilogale Gray — civet cats or spotted skunks

Spilogale gracilis saxatilis Merriam 1 iyet Cflt Qr spotted skunk

Spilogale gracilis latijrons Merriam J

Genus Mephitis Geoffroy and Cuvier — striped skunks

Mephitis mephitis hudsonica Richardson

Mephitis mephitis major (Howell) Lped skunk

Mephitis mephitis notata (Howell)

Mephitis mephitis spissigrada Bangs
Genus Taxidea Waterhouse — American badger

Taxidea taxus taxus (Schreber) badger

Family Canidae — foxes, coyote, wolves and dogs
Genus Vulpes Oken — foxes

V id pes julva cascadensis Merriam red fox

Genus Canis Linnaeus — coyote, wolves and dogs

Canis latrans lestes Merriam Icoyote

Canis latrans incolatus Hall J
Canis lupus fuscus Richardson wolf

Canis jamiliaris Linnaeus dog

Family Felidae — cats

Genus Felis Linnaeus — true cats

Felis concolor oregonensis Rafinesque l cougar or mountain lion

Felis concolor missoulensts Goldman J

Genus Lynx Kerr — lynxes and bobcats

Lynx canadensis canadensis Kerr Canadian lynx

Lynx rujus Jasciatus Rafinesque Ibobcat

Lynx rujus pallescens Merriam /

Order PINNIPEDIA— seals and walruses

Family Otariidae — eared seals

Genus Zalophus Gill— Californian sea lions

Zalophus cdifornixmus (Lesson) California sea lion

Genus Eumetopias Gill— Steller sea lion

Eurrn topias jubata (Schreber) Steller sea lion

Genus Callorhinus Gray— Alaska fur seal
Callorhinus ursinus cynocephalus \ y as k a f ur ^ oa i

(Walbaum) 1

Family Phocidae — hair seals

Genus Phoca Linnaeus— hair seals
Phoca vitulina richardii (Cray) hair seal

Check List

115

}-least chipmunk

.yellow-pine chipmunk

Order RODENTIA— rodents

Family Sciuridae — squirrels and allies
Genus Tamias Illiger — chipmunks

Tamias minimus scrutator (Hall and

Hatfield)
Turn ins minimus griscscens (Howell)
Tamias amoenus caurinus (Merriam)
Tamias amoenus felix Rhoads
Tamias amoenus ludibundus (Hollister) [ .
Ta7?iias amoenus ajfinis Allen
Tamias amoenus canicaudus (Merriam)
Tamias amoenus lutt ircntris Allen

Tamias ruficaudus simulans (Howell)
Tamias tovmsendii townsendii Bachman
Tamias townst ndii cooperi Baird

Genus Marmota Blumenbach — marmots

Marmota monax petrensis Howell

Marmota flaviventris avara (Bangs)

Marmota caligata cascadensis Howell

Marmota olympus (Merriam)
Genus Citellus Oken — ground squirrels

Citellus townsendii tovmsendii (Bachman) Townsend ground squirrel

Citellus washingtoni Howell

Citellus columbianus columbianus (Ord)

Citellus columbianus ruficaudus Howell

Citellus beecheyi douglasii (Richardson)

Citellus lateralis tescorum (Hollister)
Citellus lateralis connectens (Howell)
Citellus saturatus (Rhoads)

Genus Sciurus Linnaeus — tree squirrels

Sciurus griseus griseus Ord

Sciurus carolinensis hypophaeus Merriam

Sciurus nigcr Linnaeus, subsp.?
Genus Tamiasciurus Trouessart — red squirrels

Tamiasciurus hudsonicus streatori

(Allen)
Tamiasciurus hudsonicus richardsoni
(Bachman)

Tamiasciurus douglasii douglasii

(Bachman) ]

Genus Glaucomys Thomas — American flying squirrels

Glaucomys sabrinus oregonensis

(Bachman)

Glaucomys sabrinus banqsi (Rhoads) j.i n ■ • 1

n] * , • , i- • xj 11 [northern living squirrel

Glaucomys sabrinus columbiensis Howell • & ^

Glaucomys sabrinus latipes Howell

Glaucomys sabrinus juliginosus (Rhcads) I

Family Heteromyidae — pocket mice, kangaroo mice and kangaroo
rats
Genus Perognathus Weid — pocket mice
Perognathus parvus parims (Peale)

Perognathus parvus lordi (Gray) [great basin pocket, mouse

Perognathus parvus columbianus Merriam I

red-tailed chipmunk
Townsend chipmunk

woodchuck

yellow-bellied marmot
hoary marmot
Olympic marmot

Washington ground squirrel
^Columbian ground squirrel
Beechey ground squirrel

[•golden-mantled ground squirrel

western gray squirrel
eastern gray squirrel
fox squirrel

red squirrel

Douglas squirrel

116 University of Kansas Publs., Mrs. Nat. Hist.

Genus Dipodomys Gray — kangaroo rats
Dipodomys ordii columbiamis (Merriam)

Ord kangaroo rat

northern pocket gopher

Family Geomyidae — pocket gophers

Genus Thomomys Weid — smooth-toothed pocket gophers
Thomomys talpoides devexus Hall and

Dalquest
Thomomys talpoides columbianus Bailey
Thomomys talpoides aequalidens

Dalquest
Thomomys talpoides wallowa Hall

and Orr
Thomomys talpoides juscus Merriam
Thomomys talpoides yakimensis Hall

and Dalquest
Thomomys talpoides shawi Taylor
Thomomys talpoides immanis Hall and

Dalquest
Thomomys talpoides limosus Merriam
Thomomys talpoides doitglasii

(Richardson)
Thomomys talpoides gla<$alis Dalquest

and Scheffer
Thomomys talpoides tacomensis Taylor
Thomomys talpoides pugetensis Dalquest

and Scheffer
Thomomys talpoides tumuli Dalquest

and Scheffer
Thomomys talpoides yelmensis Merriam
Thomomys talpoides couchi Goldman
Thomomys talpoides melanops Merriam

Family Castoridae — beavers
Genus Castor Linnaeus — beavers
Castor canadensis leucodonta Gray
Castor canadensis idoneus Jewet-t and [-beaver

Hall J

Family Muridae — rats and mice
Genus Onychomys Baird — grasshopper mice

Onychomys leucogaster fuscogriseus ) northem ^shopper mouse

Anthony J

Genus Reithrodontomys Giglioli— American harvest mice

Reithrodontomys megalotis megalotis } westem harves f m0U se

(Baird) J

Genus Peromyscus Gloger — white-footed mice
Peromyscus maniculatus oreas Bangs
Peromyscus maniculatus hollisteri Osgood
Peromyscus maniculatus austerus (Baird)
Peromyscus maniculatus rubidus Osgood l-deer mouse
Peromyscus maniculatus gambelii (Baird)
Peromyscus maniculatus artemisiae

(Rhoads)

( icnus Neotoma Say and Ord— wood rats

Neotoma cinerea occidentalis Baird Ibushy-tailed wood rat

Neotoma cinerea alticola Hooper J

Check List

117

heather vole

mice

Genus Synaptomys Baird — lemming mice

Synoptomys borealis wrangeli Meniam northern lemming mouse
Genus Phenacomys Merriam — heather voles and tree mice

Phenacomys intermedins intermedins )

Merriam
Phenacomys intermedins oramontis

Rhoads

Genus Clethrionomys Tilesius — red-backed
Clethrionomys gappcri saturatus

(Rhoads)
( 'lethrionomys gapped idahoensis

(Merriam)
Clethrionomys gapperi cascadensis Booth
Clethrionomys gapperi nivarius (Bailey)

Clethrionomys calijornicus occiden talis
(Merriam)

Genus Microtus Schrank — meadow mice

Microtus pennsylvanicus funebris Dale
Microtus pennsylvanicus kincaidi
Dalquest

Microtus montanus nanus (Merriam)
Microtus montanus canescens Bailey

Microtus townsendii townsendii

(Bachman)
Microtus townsendii pugeti Dalquest
Microtus longicaudus halli Ellerman
Microtus longicaudus macrurus Merriam

Microtus richardsoni arvicoloides
(Rhoads)

Microtus richardsoni macropus (Merriam)

Microtus oregoni oregoni (Bachman)
Genus Lagurus Gloger — short-tailed voles

Lagurus curtatus pauperrimus (Cooper)
Genus Ondatra Link — muskrat

Ondatra zibethicus osoyoosensis (Lord)

Ondatra zibethicus occipitalis (Elliot)
Genus Rattus Fischer — Old World rats

Rattus rattus rattus (Linnaeus)

Rattus rattus alexandrinus (Geoffroy)

Rattus norvegicus norvegicus (Erxleben)
Genus Mus Linnaeus — house mice
Mus musculus Linnaeus, subsp.? house mouse

Family Aplontidae — mountain beaver
Genus Aplodontia Richardson — mountain beaver
Aplodontia ruja rufa (Rafinesque)
Aplodontia rufa rainieri Merriam

Family Dipodidae — jumping mice and allies
Genus Zapus Coues — jumping mice
Zapus princeps trinotatus Rhoads
Zapus princeps kootenayensis Anderson
Zapus princeps idahoensis Davis
Zapus princeps oregonus Preble

Gapper red-backed mouse
California red-backed mouse

Pennsylvania meadow mouse

montane meadow mouse

Townsend meadow mouse

long-tailed meadow mouse

water rat
creeping mouse

sagebrush vole
J muskrat

roof rat
Norway rat

^mountain beaver

big jumping mouse

snowshoe rabbit

118 University of Kansas Publs., Mus. Nat. Hist.

Family Erethizoxtidai: — American porcupines
Genus Erethizon Cuvier — North American procupine
Erethizon dors-alum epixanthum Brandt 1
Erethizon dorsatum nigrescens Allen jporcupine

Family Chinchillidae Bennett — chinchillas and allies
Genus Myocastor Kerr — nutria
Myocastor coypus (Molina), subsp.? nutria

Order LAGOMORPHA— pikas, hares and rabbits

Family Ochotonidae — pikas

Genus Ochotona Link — pikas

Ochotona princeps cuppes Bangs

Ochotona princeps jenisex Osgood r pika

Ochotona princeps brunnescens Howell

Family Leporidae — hares and rabbits
Genus Lepus Linnaeus — hares

Lepus towvsendii toumsendii Bachman white-tailed jack rabbit

L( pus americanus ivashingtonii Baird
Lepus americanus cascadensis Nelson
Lepus americanus pineus Dalquest
Lepus americanus columbiensis Rhoads
Lepus caHJomicus deserticola Mearns black-tailed jack rabbit

Genus Sylvilagus (bay — cottontails and allies
Sylvilagus nuttattii nuttallii (Bachman) Nuttall cottontail
Sylvilagus floridanus (Allen), subsp.? Florida cottontail

Sylvilagus idahoi nsis (Merriam) pigmy rabbitl

Order ARTIODACTYLA— even-toed ungulates

Family Cervidae — deer and allies
Genus Cervus Linnaeus — American elk or wapiti

Cervus canadensis roosevelti Merriam 1 •. , ., ■■■
Cervus canadensis nelsoni Bailey J e 01 NN ''''' '

Genus Odocoileus Rafinesque — northern deer

OdocoiL us virginianus ochrourus Bailev ■ ■. - , , .

, , , •, , ... ; x ^white-tailed deer

Odocoileus virginianus leucurus (Douglas;

Odocoileus hemionus hemionus
(Rafinesque)

Odocoileus hemionus eol u inhianus
(Richardson)

Genus Alee (J ray — Moose and Old-World elk

Ala iiiik ricuna shirasi Nelson moose

( Minis Rangifer Hamilton-Smith — reindeer and i aribou

Rangift r arcticus montanus Seton- ■.

mu ^caribou

I hompson J

Family Bovidae — cattle, sheep, goats and allies
Genus Bison Hamilton-Smith — bison

Bison bison oregonus Bailey bison

mule deer or black-failed deer

Check List 11!)

Genus Ovis Linnaeus — sheep

Ovis canadensis canadensis Shaw . . ,

Ovis canacL nsis califomiana Douglas , mountain sheep

Genus Oreamnos Rafinesque — mountain goat
Orcamnos americanus americanus

(Blainville) [-mountain goat

Oreamnos americanus missoulae Allen

Order CETACEA — whales and porpoises

Family Ziphiidae — beaked whales
Genus Berardius Duvernoy — beaked whales

Berardius bairdii Stejneger Baird beaked whale

Genus Mesoplodon Gervais — beaked whales
Mesoplodon stejnegeri True Stejneger beaked whale

Family Delphinidae — porpoises
Genus Delphinus Linnaeus— dolphins

Delphinus bairdii Dall Baird porpoise

Genus Lissodelphis Gloger — right-whale porpoises

Lissodelphis borealis (Peale) northern right-whale porpoise

Genus Lagenorhynchus Gray — striped porpoises

Lagenorhynchus obliquidens Gill striped porpoise

Genus Grampus Gray — killer whales

Grampus rectipinna (Cope) Pacific killer

Genus Grampidelphis Iredale and Troughton

Grampidelphis griseus (Cuvier) grampus

Genus Globicelphalus Hamilton — blackfishes

Globicephalus scammonii (Cope) blackfish

Genus Phocoena Cuvier — harbor porpoises

Phocoena vomerina (Gill) harbor porpoise

Genus Phocoenoides Andrews — Dall porpoises

Phococnoides dalli (True) Dall porpoise

Family Physeteridae — sperm whales
Genus Physeter Linnaeus — sperm whale
Physeter catodon Linnaeus sperm whale

Family Kogiidae — pigmy sperm whale
Genus Kogia Gray — pigmy sperm whale
Kogia breviceps (Blainville) pigmy sperm whale

Family Rhachianectidae — gray whale
Genus Rhachianectes Cope — gray whale
Rhachianectes glaucus (Cope) gray whale

Family Balaenopteridae— finback whales
Genus Balaenoptera Lacepede — finback whales
Balaenoptera physalus (Linnaeus) finback whale

Balaenoptera borealis Lesson Pollack whale

Balaenoptera acutorostrata Lacepede pike whale

120 University of Kansas Publs., Mus. Nat. Hist.

Genu.* Sibbaldus Gray — blue whale

Sibbaldus musculus (Linnaeus) blue whale

Genus Megaptera Gray — humpbacked whales

Megaptera novaeangliae (Borowski) humpback whale

Family Balaenidae — baleen whales
Genus Eubalaena Gray — baleen whales
Eubalaena sieboldii (Gray) Pacific right whale

Opossum 121

ACCOUNTS OF SPECIES AND SUBSPECIES

Didelphis virgin iana virginiana Ken-
Opossum

Didelphis virginiana Kerr, Anim. Kingd., p. 1P3, 17!)"2.
Type locality. — Virginia

Description. — Slightly smaller than a house cat; body in older
animals heavy and fat ; tail long, naked, scaled and prehensile. Ears
large, naked and black with white tips; muzzle elongate and pointed;
color of fur variable; overhair usually white and underfur white
tipped with black; guard hairs long and coarse but underfur soft and
dense; forefoot with opposable thumb; females with abdominal
pouch.

Remarks. — The opossum has been introduced from the eastern
United States into California and Oregon and has become well
established in those states. Recent records from Clear Lake, Skagit
County, and South Bend, Pacific County, indicate that the opossum
is now resident in Washington and it may be expected to increase
pnd spread (Scheffer, 1943). The animals may also enter the area
about Walla Walla from Oregon. The source of the opossums which
have appeared in Washington is not yet known.

Of this animal, Dr. Carl Hartman (1923: 347) has written:

"In the popular mind, the generation of no animal is so shrouded
in mystery as that of the opossum. Throughout the country, among
both whites and negroes, deeply rooted tradition has it that the
opossum copulates through the nose and that the female blows the
fruit of conception into the pouch. Other myths relating to details
of the reproductive process in this species are current among the
people.

"The growth of such legends need not surprise one, however, for
the early birth of the embryos and the use of the pouch as an
incubator certainly challenge the imagination. These phenomena
attract the attention because they are unique, differing from the
familiar method of rearing the young obtaining among the higher
mammals, including man. Familiarity breeds contempt; the ordinary
ceases to be marvelous. Thus on account of its rareness and its
'different' character the opossum, our only marsupial, figures in the
folklore to a prominent degree."

122

University of Kansas Publs., Mis. Nat. Hist.

Neurotrichus gibbsii
Gibbs shrew-mole

Description. — The shrew-mole is tiny, possessing a head and body
2V2 to 3 inches long and a tail about 1% inches in length. The body
is relatively stout but is less cylindrical than that of Scapanus. The
eyes are nearly buried in the fur. The nose is long and pointed.
The legs are short and the forefeet wide and powerful. The tail is
thick, constricted at the base and clothed with short, stiff bristles.
The fur is short and posteriorly directed. In color the shrew-mole is
dark slate, almost black.

Fio. 21. Gibbs shrew-mole (Neurotrichus gibbsii minor), female in captivity;

Seattle, Washington, September 12, 1939. (Fish and Wildlife Service photo bv
Victor B. Scheffer, No. 719.)

Remarks. — Shrew-moles inhabit moist habitats from sea level to
8,000 feet. They are burrowing mammals and prefer to live in soft
earth, free of sod. In the lowlands of western Washington, shrew-moles
arc most abundant in damp, shady ravines where the vegetation in-
cludes deciduous trees and dense underbrush with but little grass.
In the mountains, shrew-moles are usually found near streams or
rock slides, where the larger annuals grow densely on soil that is
deep, soft, and free of turf. Vertically they range from the Humid

Moles

123

Transition Life-zone through the Canadian, and well into the Hud-
sonian Life-zone.

Shrew-moles are both diurnal and nocturnal. They rest or sleep
periodically, the length of their rest or sleep being longer when much
food is eaten, and the intervals between their periods of rest or sleep
is longer when less food is eaten. Owls and snakes appear to be
their principal enemies. Predatory mammals eat some shrew-moles,
and probably kill many that they do not eat. Shrew-moles are
completely blind, and their long, prehensile nose guides all their
activity. Their ordinary movements on the surface of the ground
are slow and cautious. When frightened they break into a scuttling

DISTRIBUTION

MAP

STATE OF WASHINGTON

■

25 50

75 10

MILES

'■«

d Under A US pc»i of WPA P.ej
1 1

ci 60'9-5T37 t> 3 CM

1

Fig. 22. Distribution of the Gibbs shrew-mole in Washington. A. Neuro-
trickus gibbsii gibbsii. B. Neurotrichus gibbsii minor.

rush which ends beneath a leaf or bit of bark where the animal be-
comes motionless. Shrew-moles, on the surface of the ground, make
considerable noise. They construct molelike burrows, but these are
not as extensive or complicated as those of moles. The shrew-mole
hunts for food in shallow trenches that it makes just under the layer
of dead leaves and vegetable debris that covers the ground in their
habitat. The food of the shrew-moles includes earthworms, isopods,

124 University of Kansas Publs., Mrs. Nat. Hist.

insect larvae, soft-bodied insects, and other animal matter. Some
vegetable matter is eaten.

Breeding takes place at all seasons of the year, save perhaps in
December and January. Embryos vary from one to four. The nest
of a shrew-mole at Seattle, King County, consisted of a handful of
damp leaves in a cavity of a rotten, punky. alder stump. The nest
contained four half-grown young.

Neurotrichus gibbsii gibbsii (Baird)

Urotrichus gibbsii Baird, Mamm. N. Amer., p. 70, 185".

Neurotrichus [sic] gibbsi Giinther, Proc. Zool. Soc. London, pi. 42, 1880.

Neurotrichus Gibbsii True, Proc. U. S. Nat. Mus., 7 (1884): 607, 1885.

Neurotrichus gibbsii Bryant, Zoe, 1:359, February. 1891.

Neurotrichus gibbsii gibbsii Miller, U. S. Nat. Mus. Bull., 79:11, December 31, 1912.

Type. — Obtained at Naches Pass, 4,500 ft.. Pierce County, Washington,
by G. Gibbs on July 15, 1854 (see Dalquest and Burgner. 1941); type in
United States National Museum.

Racial characters. — Size relatively large; tail relatively long; foreclaws
straight on ventral surface.

Measurements. — Two females from Tye, 4,000 ft., Stevens Pass, King
County, average: total length, 121.5; length of tail, 45; length of hind foot. 18.

Distribution. — The Cascade Mountains, from British Columbia south, and
Destruction Island, Jefferson County. Records of occurrence are Baker Lake
(J. M. E.), Tye (M. V. Z.), and Mount Rainier (Mount Rainier Ntl. Park
Mus.)

Remarks. — Shrew-moles are present on Destruction Island, a
small island in the Pacific off the coast of Jefferson County. These
moles are large, and are like gibbsii. It is thought, however, that
this resemblance is due to convergent evolution rather than a once-
continuous range with gibbsii. It is significant that a shrew {Sorex
trowbridgii destructioni) , the only other native land mammal on the
island, differs from its mainland counterpart in much the same way
as does Neurotrichus g. gibbsii from Neurotrichus g. minor.

Neurotrichus gibbsii minor Dalquest and Burgner

Neurotrichus gibbsii minor Dalquest and Burgner, Murrelet, 22:12, April 30, 1941.

Type. — Obtained on the University of Washington Campus, Seattle, King
County, Washington, by W. W. Dalquest on May 19. 1940; type in the
Museum of Vertebrate Zoology.

Racial characters. — Size small; tail short; foreclaws light and weak as com-
pared to those of gibbsii, with ventral surfaces curved.

Afcasurements. — Eighty-five specimens (males and females) from Seattle,
King County, average: total length. 107.0; length of tail. 35.3; length of
hind foot, 15.0.

Distribution. — The lowlands of western Washington. Marginal records are:
Mt. Vernon (Jackson, 1915:97), Cottage Lake (W. W.D.) and Yacolt (M.
V.Z.).

Moles

125

Genus Scapanus Pomel

Moles

Moles are of stocky build and have cylindrical, rounded bodies.
The eyes are tiny, nearly concealed in the fur. They have no ex-
ternal ears. The legs are short; the forefeet wide, spadelike, and
armed with powerful claws. The fur is erect, not posteriorly di-

.biG. 23. Coast mole (Scapanus orarius orarius) left
and Townsend mole (Scapanus townsendii) right; Puyal-
lup. Washington. Mav, 1914. (Fish and Wildlife Service
photo by T. H. Scheffer, No. B-1S637.)

rected. The color of the fur is deep bluish or brownish slate. The
short, nearly naked tail is pale pink or whitish. Moles of this genus
are found only along the Pacific Coast of North America from
southern Canada south into Baja California.

126 University of Kansas Publs., Mrs. Nat. Hist.

Scapanus townsendii (Bachman)

Townsend mole

Scalops townsendii Bachman. Jour. Acad. Nat. Sci. Philadelphia, 8 (pt. 1):58, 1830.
Scapanvs tow[n]sendii Pomel, Arch. Sci. Phys. Nat., Geneva, 9 (ser. 4):247, 1848.
Scapanus Townsendii True, Proc. U. S. Nat. Mus., 7 (1884):607, 1885.

Type. — Probably obtained at Fort Vancouver. Clark County. Washington,
by J. K. Townsend. A cotype was obtained on May 9. 1835; type in Academy

of Natural Sciences of Philadelphia.

Measurements. — Three males and 6 females from southwestern Washington
average, respectively: total length 221. 214; length of tail 51. 46; hind foot
28.3. 26; weight 147. 117 grams.

Distribution. — The lowlands of western Washington. Marginal occurrences
are: Sauk (Jackson. 1915: 61); Skykomish (Jackson. 1915: 61) and YacoU
(M.V.Z.).

Remarks. — Scapanus townsendii occurs only in a narrow belt ex-
tending from southwestern British Columbia to northwestern Cali-
fornia. It seems to prefer a generally damper habitat than the
smaller-sized coast mole, although both species are sometimes found
in the same locality. The larger mole is abundant in the meadows
on the flood plains of rivers at low elevations, and on the glacial
outwash prairies. It is often numerous in the fir forests, although
its workings and mounds are less conspicuous there. The bodies of
nine drowned individuals were found in a well by an old cabin in
dense fir forest near Duvall, King County. Townsend moles occa-
sionally occur in the Canadian Life-zone, as at Staircase on the
north side of Lake Cushman in Mason County where workings were
observed, but most records are from the Humid division of the
Transition Life-zone.

The Townsend mole is mainly nocturnal. If ridges of its runways
are crushed down, they usually remain so throughout the day and
are rebuilt the following night. Only about ten per cent of the ridges
that were crushed were rebuilt in the daytime, and most of these
were repaired in the early morning. The species is almost com-
pletely subterranean. Some individuals are crushed on highways
by cars, showing that these moles occasionally travel on the surface
of the ground.

Townsend moles throw up numerous mounds, each usually con-
taining about a cubic foot of earth. The mounds commonly are
built just about as far apart as a man can step. The general direc-
tion as well as the twists and turns of a mole's burrow can usually
be determined from the mounds. Townsend moles also construct
ridges on the surface of the ground by pushing up sod in building a

Moles

127

tunnel just below the grass roots. Smaller ridges are less commonly
made by coast moles. More extensive tunnels, constructed deeper
in the earth, serve as living quarters.

According to Wight (1928:24), Scheffer (1922:11) and Moore
(1933: 39), the food of this large mole includes earthworms and
ground-inhabiting insects, insect larvae, spiders, centipedes, flesh,
and small amounts of soft vegetation. Scheffer (1922: 10) found
that the large mole breeds in February and produces from two to
four young at a litter, with an average of three.

DISTRIBUTION MAP
STATE OF WASHINGTON

I I I

Fig. 24. Distribution of the Townsend mole, Scapanus townsendii, in

Washington.

Scapanus orarius

Coast mole

Description. — The coast mole is almost identical with the larger
mole in form of body but is smaller. Head and body are about
5V4 and tail about IV2 inches in length.

The coast mole occupies all of the territoiy inhabited by the
Townsend mole and ranges slightly farther northward, southward
and eastward. However it does not range east of the boundaries
of the three Pacific Coast states or British Columbia.

128

University of Kansas Publs., Mus. Nat. Hist.

The mounds and workings of the coast mole are smaller than
those of the Townsend mole and consequently are less noticed. It
seems less prone to make numerous mounds, a pace apart, than the
larger mole, and burrows tend to extend deeper in the ground. Up-
thrust ridges are less commonly built by orarius than by townsendii .

Jackson (1915: 62) mentions the possibility of ecologic differ-
ences between orarius and townsendii. There are some differences
in habitat and habits. The coast mole seems to live deeper in the

Fig. 25. Coast, mole (Scapanus orarius orarius). freshly killed; Seattle,
Washington, June 9, 1938. (Fish and Wildlife Service photo by Victor B.

Scheffer, No. 64.)

ground, prefers better drained soil, and is less colonial than the
Townsend mole. These are average differences, however, and the
two species commonly occur together. Another difference is that
the larger mole rarely enters the dense deciduous woods, Mich as
the brush-grown alder and dogwood jungles along the stream val-
leys. I have taken the coast mole in such localities, and often have
found their workings there. The surface of a nearby meadow may
be dotted with mounds of both species, but the larger mole seems
not to enter ground that i> thickly grown with brush. The coast
mole ascends to greater altitudes in the Cascade Mountains than
does the Townsend mole.

Like the larger mole the coast mole feeds principally on earth-
worms and insects I Moore, 1933: 38). On September 2!), 1939, in

Moles

129

a ravine at Seattle, in two baited traps set six inches apart, a creep-
ing mouse (Microtus oregoni) and a coast mole were found. The
mouse had come to the bait and been caught. The mole had seem-
ingly emerged from the ground through a tunnel a foot away and
had been attracted to the body of the mouse. It had eaten an area a
half inch in diameter and three-quarters of an inch deep into the
body of the mouse just behind the shoulder, when in shifting its
position it had become caught in the unsprung trap behind it. When
found the mole lav dead with its nose inside the body of the mouse

rYT t

DISTRIBUTION MAP
STATE OF WASH I NGTON

WILES

Fig. 26. Distribution of the coast mole in Washington. A. Scapanus orarius
orarius. B. Scapanus orarius yakimensis. C. Scapanus orarius schefferi.

and its back broken. This is the only case known to me of a coast
mole appearing voluntarily on the surface of the ground. Never
have I found coast moles crushed on the highway; several Town-
send moles so killed have been found.

The coast mole seems to breed very early in the spring. Males
with swollen testes are found late in January. The young usually
number four and are born in late March or early April. The breed-
ing season seems to be the same as that of townsendii but the num-
ber of young to a litter may average slightly greater.

9-4993

l.*!() University of Kansas Pibls., Mis. Nat. Hist.

Scapanus orarius orarius True

Scap i is orarius True, Proc. L*. S. Nat. Mus., 19:52, Decembei 21, 1896.
Scapim a orarius orarius Jackson, X. Amer. Fauna, S8:(il. September 30, 1915.

Type. — Obtained at Shoalwater (=Willapa) Bay. Pacific County. Washing-
ton, by J. G. Cooper on August 30. 1855; type in United States National
Museum.

Racial Characters. — Color dark bluish; frontal region of skull not inflated.

Measurements. — Eight males and two females from Seattle. King County.
average, respectively: total length 159. 155; length of tail 33. 31; hind foot
20.7, 20.5; weight 58.5, 55.8.

Distribution. — Western Washington. Marginal occurrences are : Skykomish
(B.S.C.), Merritt (B.S.C.), Wenatchee (B.S.C.), Lester (Jackson, 1915: 64)
and Yacolt (M.V.Z.).

Scapanus orarius yakimensis Dalquesl and Scheffer
Scapanus orarius yakimensis Dalquesl and Scheffer, Murrelet, •2.">:27. September 19, 1944.

Ti/ju . — Obtained % mile north of Union Gap, Yakima County. Washington,
by J. A. Gray, Jr.. on July 3, 1941; type in Museum of Vertebrate Zoology.

Racial Characters. — Color pale, grayish; skull narrow with inflated frontal
region.

Measurements. — Six specimens (males and females) from Selah, Yakima
County, average: total length 164; length of tail 37; length of hind foot 21.5.
The type specimen weighed 58 grams.

Distribution. — The Yakima Valley area; recorded from the type locality
northwestward to Easton (B. S.C.).

Scapanus orarius schefferi Jackson
Scapanus orarius schefferi Jackson, X. Amer. Fauna, 38:63, September 30, 1915.

Type.— Obtained at Walla Walla. Walla Walla County. Washington, by T.
H. Scheffer on August 8. 1914; type in United States National Museum.

Racal Characters. — Large size; pale color; large, wide skull with inflated
frontal area.

Measurements. — Six topotypes average: total length 159; length of tail 35;
hind foot 21.5.

Distribution. — Southeastern Washington; recorded from Fort Walla Walla
(Jackson. 1915: 64). Walla Walla (Jackson. 1915: 64) and Dayton (M.V.Z.).

Shrews 131

Genus Sorex Linnaeus
Long-tailed shrews

Shrews have tiny eyes, almost concealed in the fur. The body is
slim; the nose elongate and pointed. The legs are short and the feet
small and weak. The fur is short but soft and posteriorly directed.
The cinereous shrew, for example, is about 4 inches in length, of
which the tail comprises 1% inches. The upper parts are dark
grayish brown and the underparts dull gray.

Sinews of the genus Sorer are cosmopolitan in distribution. In
North America they range from the arctic south to Central America.
Three subgenera are recognized by Jackson (1928: 27), all three of
which are represented in the state of Washington. The twelve sub-
species present occupy numerous habitats, and their ranges include
almost all of the state.

Jackson pointed out (1928: 1) that "No other group of American
mammals having a wide distribution, and in many localities
an abundance of individuals, is so little known to the nonpro-
fessional mammalogist as the long-tailed shrews." In Wash-
ington, especially in the coastal area of western Washington,
shrews are widespread and abundant in many habitats. In some
places they are the most common mammal present. In spite of this
their presence is often unsuspected by persons that are otherwise
alert to the animal life around them. This is even more surprising
when one takes into account the facts that long-tailed shrews are
diurnal as well as nocturnal and are less apt to detect the presence
of man than are most other species of mammals.

An important factor making long-tailed shrews inconspicuous is
their small size, and contributing factors include their rapid move-
ments and dull colors. On several occasions the writer, after seeing
a long-tailed shrew vanish soundlessly under a log or into a patch of
dead leaves, was left wondering if one actually had been seen or if
instead his imagination had conjured up an animal from a dust mote
or wind-blown leaf.

The environment of the long-tailed shrews, except for the water
shrews, is the zone at the very surface of the earth, just beneath the
layer of moss, grass, dead leaves, and decaying vegetation. The
removal of the covering vegetative layer reveals a maze of tiny
tunnels, the branchings and complexities of which are infinite. Tiny

132 University of Kansas Ptjbls., Mus. Nat. Hist.

traps baited with oats or meat and set in these runways catch the
long-tailed shrews that inhabit them.

The food of long-tailed shrews is varied. It is principally soft-
bodied insects, insect pupae, and earthworms. At times a consider-
able quantity of soft vegetation and some seeds are eaten. Recent
studies i Moore, 1940: 1942) have shown that by destroying seeds
some shrews may adversely affect the reforestation of some coni-
ferous trees. Shrews readily eat meat, and often destroy the small
mammals, including other shrews, taken in the mammal collector's
traps.

Hamilton (1940:485) found that in one species of long-tailed
shrew in the United States individuals rarely lived more than one
year. This seems not to be true of at least some of the species found
in Washington.

Parasites, internal or external, are not commonly found on long-
tailed shrews. They are regularly eaten by owls and snakes, but
most carnivorous mammals, though they readily kill them, rarely
eat them.

Sorex cinereus Ken-
Cinereous shrew

The cinereous shrew ranges over most of Alaska, Canada and the
northern half of the United States. A number of subspecies have
been described, of which two have been reported from Wash-
ington. The cinereous shrew is of medium size and difficult to dis-
tinguish from the dusky and wandering shrews, especially in eastern
Washington, without studying the skulls. In cinereus the fourth
unicuspid tooth is smaller than the third; in uagrans and obscurus
it is larger. The relatively narrow rostrum of cinereus also serve-
to separate it from the other two species.

The cinereous shrew seems to be less restricted to the vicinity of
streams and marshes than Sorex vagrans, resembling Sorex obscurus
and Sorex trowbridgii in this respect. In Washington it seems to
be rare and has not been taken by the writer. Published records
for Sorex C. cinereus indicate that this race is, in Washington, con-
fined to mountainous areas. The coastal race, streatori, seems to be
confined to the humid area.

Sorex cinereus cinereus Kerr

Sorei arcticut cinereus Kerr, Anim. Kingd., p. '_''»i, 1702.

Sorei cinereu cii u Jackson, Jour. Mamm., 6-:66, February 9, L925.

Tiii» . — None. Name based on the account of a shrew -ecu at Fort Severn.
Canada, by J. K Forster in 1772.

Shrews

133

Racial characti rs. — Small size and pale color.

Measurements. — Seven males and 8 females from Indian Point Lake, British
Columbia, average, respectively: total length 99, 95; length of tail 43.5, 42.2;
hind foot 12.7. 12.3.

Distribution. — Northeastern Washington (Metaline and Loon Lake, Jackson,
1928: 49) and the Cascades from Whatcom Pass (Jackson. 1928: 55) south-
ward to Conrad Meadows (Jackson, 1928: 49). There is a specimen in the
collection of the California Academy of Sciences, from Mt. Rainier.

R< marks. — This is the most wide-ranging subspecies of shrew, being re-
corded from Alaska. 10 Canadian provinces and 26 states (Jackson, 1928:
46-50). In the more northern and eastern parts of its range it is sometimes
the commonest shrew.

DISTRIBUTION MAP

STATE OF WASH I NGTON

25

n:

50

75

100

UILES

0'ofted Under Autp.ct, of WPA P.oj.ti 6079-5737 by TMT 1,37
I I I

124

123

122

Fig. 27. Distribution of the cinereous shrew in Washington. A. Sorex cin-
ereus cinereus. B. Sorex cinereus streatori.

Sorex cinereus streatori Merriam

Sorex persotmfii* streatori Merriam, X. Am r. Fauna, 10:62, December 31, 1895.
Sorex cinereus streatori Jackson, Jour. Mamni., 6:56, February 9, 1925.

Type.— Obtained at Yakutat, Alaska, by C. P. Streator on July 9. 1895;
type in the United States National Museum.

Racial characters. — Larger and darker than Sorex cinereus cinereus.
Measurements. — Five females from Alta Lake. British Columbia, average:
total length 103; length of tail 45; hind foot 12.5.

184 University of Kansas Publs., Mrs. Nat. Hist.

Distribution. — The western part of the northern Cascades (Glacier, Jackson.
1928: 55) and the Olympic Peninsula (Xeah Bay south to Cedarville, Jack-
son. 1928: 55).

I!* mark*. — This shrew seems to occupy the same range as Sorex obscurus
S( tosus but is rare where obscurus is common.

Sorex merriami merriami Dobson
Merriam shrew

Sorex merriami Dobson, Monogr. [nsectivora, pt. S, fasc. 1, pi. -3. fig. >'. May, 1S90.
Sorex merriami merriami Benson and Bond, .lour. Mamm., 20:348, Augusl II. 1939

Type. — Obtained on Little Bighorn River, about a mile and a half above
Fort Custer. Crow Indian Reservation. Montana, by Charles E. Bendire on
December 26, 1884; type in United State- National Museum.

Measurements. — Of type: total length 90; length of tail 35; hind foot 11.5
(after Jackson, 1928: 80).

Distribution. — A single specimen of this rare sinew is known from Washing-
ton. Jackson (1928: 81) states that it -was collected by George (i. Cantwell,
November 18, 1919. at the entrance to an old badger digging on top of a
'high hunch grass hill' at Starbuck (altitude 645 feet). Columbia County,
Wash."

Description. — Similar to Sorex cinen us but upper parts pale gray-
ish and underparts white.

The Merriam shrew has been recorded from but a low localities in
the western United States. It is one of the rarest of the small mam-
mals known to occur in North America. All the known specimens
have been found in desert areas.

Sorex trowbridgii Baird

Trowbridge shrew

I )< script ion. — The Trowbridge shrew closely resembles the ciner-
eous shrew in body form but possesses a longer tail. Tin' head ami
body of adults measure about l" i inches and the tail about 2 inches.
The Trowbridge shrew may be separated from all other small shrews
that occur in Washington by its dark bluish upper part- and bluish
or slaty underparts. The tail i> distinctly bieolor.

Trowbridge shrews occur from southern British Columbia south
to central California. Their distribution, to the south, is more
extensive than that of many members of the Pacific Coastal Fauna.
They are forest, animals, ranging widely over the dry ground be-
neath the fir forest, where they are usually the only -brews present.
They are abundant in ravines and in some swampy woods when
other shrews are absent, but they avoid open meadows or marshes.

Shrews

135

Vertically, they occur from the humid division of the Transition
Life-zone to the Hudsonian Life-zone.

These little shrews do not live well in captivity and I have learned
relatively little concerning their habits from live specimens. They
seem to be slower-moving and less aggressive than the wandering
shrew. Population studies showed that the Trowbridge shrew is
unable to compete with the wandering shrew in ravine habitats
(Dalquest, 1941 A: 173). The principal food of the Trowbridge
shrew includes soft-bodied insects and insect pupae.

DISTRIBUTION MAP
STATE OF WASHINGTON

Under A US p.c*. cf * PA P..j.d 6079-5797 b, CM

Fig. 28. Distribution of the Merriam and Trowbridge shrews in Washing-
ton. A. Sorex merriami merriami. B. Sorex trowbridgii trowbridgii. C.

Sorcx trowbridgii destruction/.

Moore (1942) has shown that shrews eat the seeds of the Douglas
fir and may be a serious check on the reproduction of this important
tree. The Trowbridge shrew is the most abundant shrew in the fir
forests and probably constitutes the principal shrew that might be
classed as a pest.

A specimen containing 4 embryos was taken near Shelton. Mason
County, on April 23, 1937. Males with enlarged, greenish testes
were taken in April of 1938, 1939 and 1940. Specimens obtained
in other months showed no indications of breeding.

136 University of Kansas Pixels., Mis. Nat. Hist.

Sorex trowbridgii trowbridgii Baird

Sorex trowbridgii Baird, Rept. Pacific R. R. Survey, 8 (pt. 1):13. 1857.

Type. — Two cotypes were obtained at Astoria, Clatsop County, Oregon. The
skin of one (the lectotype) was entered in the U. S. National Museum cata-
logue in July, 1855. and the skull in January, 1857. The other was obtained
by .1. Wayne on July 10. 1855.

Racial characters. — Small size and narrow skull.

Measurements. — The average measurements of 38 adults from King County.
Washington, are: total length 115.3; length of tail 54.4; hind foot 13.4.

Distribution. — Forested anas from the Pacific Coast eastward through the
Cascades to Stehekin (Jackson, 1928: 96), 2 mi. S Blewitt Pass (Jackson,
1928: 96) and Satus Pass fM.Y.Z.).

Sorex trowbridgii destructioni Scheffer and Dalquest
Sorex trowbridgii destructioni Scheffer and Dalquest, Jour. Mmnm.. 23:334, August 13,

Sj/pe— Obtained on Destruction Island, Jefferson County. Washington, by
V. H. Scheffer on April 22. 1941; type in United States National Museum.
Racial characters. — Large size and wide skull.

Measurements. — Thirty topotypes average: total length 122.5; length of
tail 56.7; hind foot 14.3; weight 7.5 grams.

Distribution. — Known only from Destruction Island, 35 acres in area, lying
4 miles off the Washington Coast.

Sorex vagrans Baird
Wandering shrew

Description. — This species closely resembles the cinereous shrew
in body form. Its head and body measure about 2 1 4 inches; the
tail slightly less than 2 inches. In summer the upper parts are red-
dish brown and the underparts gray tinged with brownish. In win-
ter the upper parts are more dusky.

Several races of the wandering shrew range over western North
America from southwestern British Columbia south to southern
Mexico. Two races occur in the state of Washington.

Marshy areas and damp places are the habitat of the wandering
shrew. Cattail and tule marshes, sphagnum bogs, and meadows are
favored. They frequent streams through forests but rarely are taken
in places away from water. On some of the San Juan Islands, wan-
dering shrews were found along the beaches where they were feeding
on the amphipods that live in the dead seaweed and litter at the
high tide line. In a favorable habitat, wandering shrews may be
the most abundant mammal present. Specimens are occasionally
taken in 90 per cent, or more of ;i mammal collector's traps.

The preference of the wandering shrew for damp areas makes it
more or less independent of life-zones, for marshy areas, whether

Shrews

137

in Transition. Canadian, or Upper Sonoran life-zones, present com-
parable ecological conditions.

Broadbrooks (1939: 65) found that captives taken at Seattle ate
lolled oats, apple, fresh or cooked meat, sow bugs, centipedes, earth-
worms, frogs {Hyla regilla), a salamander iPlethodon vehiculum) ,
and small, black slugs (Arean are an) . Wandering shrews proved in-
capable of destroying snails (Helisoma occidentalis) and large slugs.
The captive shrews kept by Broadbooks ate an average of 1.3 times
their own weight in food each day.

DISTRIBUTION MAP
STATE OF WASH I NGTON

-5T97 b.CWT iJiT

Fig. 29. Distribution of the wandering shrew in Washington. A. Sorer
vagrans vagrans. B. Sorex vagrans monticola.

The wandering shrew changes from winter to summer pelage in
a few days, seemingly in the second week of October (Dalquest,
1944: 147). The spring molt occurs rapidly but perhaps at a less
regular date. Rarely a midsummer molt occurs.

The earliest evidence of breeding in a wandering shrew was dis-
covered on January 27, 1937. Most adult females taken in Febru-
ary, March, April and May were pregnant. Embryos were less often
found in summer and fall, but one pregnant female was taken in
November. Embryos varied in number from 3 to 8 with an average
of six.

138 University of Kansas Pi bls., Mis. Nat. Hist.

Sorex vagrans vagrans Baird

S ... vagrans Bund, Rept. Pacific R. R. Survey, s (pt. l):l">. 1857.

Sores suckleyi Band. Kept. Pacific R. R. Survey, - (pt. 1):18, 1857 (type from Steilac mi,
Pierce County, Washington).

Type. — Obtained at Willapa Bay [Shoalwater Bay I. Pacific County. Wash-
ington, by .1. G. Cooper; entered in V. S. Nat. Mus. catalogue on October
23. 1856.

Racial characti rs. — Dark color.

Measurements. — Twenty-tive males and 25 females from Seattle. King
County, average, respectively: total length 110.5. 107; length of tail 44.6. 45;
hind foot 12.4. 12.4.

Distribution. — This is the common marsh shrew of western Washington and
occurs from the Pacific Ocean east to the Cascades at Lake Keeehelus (W.
S.M.) and 15 mi. NW White Salmon (Jackson. 1928: 106).

Sorex vagrans monticola Merriam

Sorex moulicolvs Merriam, X. Amer. Fauna, 3:43, September 11. 1890.

Sores vagrans monticola Merriam, X. Amer. Fauna, 10:09; December 31, 1895.

Type, — Obtained on San Francisco Mountain. 1150 ft. altitude. Coconino
County. Arizona, by C. H. Merriam and V. Bailey on August 28, 1889; type
in United States National Museum.

Racial characters. — Pah 1 color.

Measurements. — Five males and 5 females from Selah, Yakima C<unty.
average, respectively: total length 100. 98.4; length of tail 40.6. 39.2; hind
foot 12.2. 12.

Distribution. — Eastern Washington. The range of this shrew extend- west
to Bauerman Ridge (Jackson, 1928: 113). Merritt (W.W.D.) and Maryhill
(M.V.Z.).

Remarks. — A series of shrews from Moses Lake. ( Irani County, differs from
monticola in larger size and darker color. They agree rather closely with
Sorex V. amoenus from California and Nevada. Because they are isolated
from that subspecies it seems best to consider them a microgeographic ran
[■< ferable to monticola.

Sorex obscurus Merriam

I )usky shrew

Description. — The dusky shrew is similar in form of body to the
cinereous shrew. The length of head and body is about 2^4 inches.
The tail is about 2% inches. The upper parts are rusty or reddish
brown. The underpays are brownish gray. The dusky shrew (lif-
ters from the wandering shrew in possessing a slightly longer body
and longer tail, but in eastern Washington the two species are almost
indistinguishable.

Dusky shrews range from northern Alaska to southern New Mex-
ico, and from the Rocky Mountains to the Pacific Ocean. Thirteen
subspecies are recognized by Jackson (1928: 115), of which two
occur in Washington.

Shrews 139

Records of the dusky shrew arc not available from the arid sub-
division of the Transition or the Upper Sonoran life-zones. It oc-
curs sparingly in the humid subdivision of the Transition and is
common in the Canadian and Hudsonian life-zones.

The habitat of the dusky shrew is varied. Near Seattle, King
County, several specimens were trapped in marshes where Sorer
vagrans was abundant. Near Stevens Pass, King County, two werG
taken in a marsh; two others were trapped in a dry, coniferous
foicst ; one was taken in a small bed of heather on a barren moun-
tain top; and another was found dead in a pan of pancake batter in
camp. Two specimens were trapped along a small stream at Dewey
Lake. Mt. Rainier, Yakima County. Four specimens were caught
in traps set in a talus slope on a dry hillside at the North Fork of
the Quinault River, Jefferson County. Three others were trapped
in dense, rain-forest thickets along the ocean at La Push, Jefferson
County. These records indicate that the dusky shrew has a wider
environmental range than other Washington shrews. Despite this
wide range of tolerance the dusky shrew is common only locally,
except in the Hudsonian Life-zone.

Little is known of the habits of dusky shrews but they seem to
be as diurnal as they are nocturnal. At Wolf Bar, North Fork of
the Quinault River, Jefferson County, a dusky shrew was seen on
the packed-earth floor of an old trapper's cabin. A hat was care-
fully dropped over the live animal but, when the hat was lifted,
the shrew was found dead. In an old cabin at Stevens Pass, King
County, a dusky shrew was found dead one morning in a pan of
flapjack batter prepared the evening before. As this pan was on a
table about three feet from the floor, the shrew must have climbed
to the table by way of the rough cabin wall, but how the animal
managed to scale the side of the pan is a mystery.

Slipp (1942: 211) discovered the nest of a dusky shrew between
Round Pass and Lake George, 4200 feet elevation, in Mt. Rainier
National Park, on July 25, 1937. The nest was in a rotten fir log
20 inches in diameter. The nest, a ball of dry grass the size of a
man's fist, had no central cavity or passages, the occupants "merely
pushed through wherever they wished." Seven young shrews were
found in and near the nest. Though the eyes of the young were
still (dosed, they were able to creep about and squeal.

A specimen obtained 5 miles west-southwest of Guler, Skamania
Countv, contained 4 embrvos on Julv 10, 1939.

140

University of Kansas Ptjbls., Mrs. Nat. Hist.

T"

^\ op' / ! -r\ !

y !

DISTRIBUTION MAP
STATE OF WASHINGTON

C-=lTd U"d«. Au

WILES

of WPA B.ejtl

tC'^O^r b,CMT |>J7

Fig. 30. Distribution of the dusky sinew in Washington. A. Sorer obscurus
obscurus. B. Sorex obscurus s< in.-us.

Sorex obscurus obscurus Merriam

Sorex vagrant small* Merriam, X. Amer. Fauna, .">:34, July 30, 1891 (not of Hensel, 1855).

Sorex obscurus Merriam, N. Amer. Fauna. 1o:7l>, DimiihIui 81. 1 SK.'i (substitute foi similis

M'i i iam ).

Type. — Obtained on Timber Creek, 8.200 ft.. Lemhi Mountains, Lemhi
County. Idaho, by V. Bailey and B. H. Dutcher on August 26. 1890; type in
United States National Museum.

Racial characters. — Tail relatively short, <• lor pale.

Measurements. — Thirteen male- and 10 females from Indian-point Lake,
British Columbia, average, respectively: total length 105, 107; length of tail
45. 46; hind fool 13.3, 13.3.

Distribution. — Northeastern Washington west, according to Jackson (1928
122). to I'a-avicn River, Stehekin and Wenatchee. Hi cause specimens from
Tye, King County, and Mt. Stuart, Easton and Lake Keechelus are clearly
referable to S. o. setosus, the specimens recorded by Jackson (1928:122),
from Easton and Signal Peak, are mapped in the range of setosus.

Sorex obscurus setosus Elliol

Sorex setosus Elliot, I ield Columb, Mus. Publ. 82, zool. ser., 1:274, March, 1899.

Son • nil in,, is setosus Jackson, I 'i • ■•■. Biol, Soc. Washington, 31 : 127, November 29, 1918.

Sorex obscurus bairdi Jacks N. ^mer, Fauna, 51:140, Julj 24, 1928.

Type. Obtained a1 Bappy Lake. Clallam County, Washington, by I). G.
Elliol on Augusl 18, 1898; type in Field Museum of Natural History.

Shrews

141

Racial characters. — Tail long, color dark.

Measurements. — Twelve males anil 17 females from southwestern Wash-
ington average, respectively: total length 118, 119; length of tail 53, 53; hind
foot 13.8. 13.7; weight 6.6. 5.2 grams.

Distribution. — Western Washington, east through the Cascades to Barron
(Jackson. 1928: 137), Cascade River (Jackson 1928: 137) and Satus Pass
(W.W.D.).

Sorex palustris navigator (Baird)

Mountain water shrew

Neosorex navigator Baird, Rept. Pacific R. R. Survey, 8 (pt. 1 ) : 1 1 . 1857.

Sorex (Xeosorex) palustris navigator Merriam, N. Anier. Fauna, 10:92, December 31, 1895.

Sorex palustris navigator Stephens, California Mammals, p. 254, June, 1906.

Type. — Obtained at head of Yakima River, Kittitas County, Washington,
by J. G. Cooper on August 31, 1853; type in United States National Museum.

Measurements. — Two males and 4 females from Washington average, re-
spectively: total length 150. 150; length of tail 70. 74; hind foot 18.5, 20;
weight ?, 11.0 grams.

DISTRIBUTION MAP
STATE OF WASHINGTON

I

I MM-ifJT », zm ilJT

Fig. 31. Distribution of the mountain water shrew. Sorex palustris navigator,

in Washington.

Distribution.— Mountainous areas of entire state, including the Olympic
Mountains, from Elwah (Jackson, 1928: 188) south to Quinault River (Jack-
son, 1928: 189); the Cascades from Tomyhoi Lake (W.W.D.) south to 15

142 University of Kansas Publs., Mrs. Nat. Hist.

mi. X. Carson (Jackson. 1928: 189); northeastern Washington from Shovel
Creek (W.W.D.) south to Gifford (Jackson. 1928: 189): the Blue Mountains
at Godman Springs (M.V.Z.) and Hompeg Falls (M.V.Z.).

Description. — In general form of body the mountain water shrew
resembles the cinereous shrew but is perhaps more stocky. It is a
large shrew, nearly as large as a house mouse. The head and body
measure about 3 inches; the tail also is about 3 inches long. The
fur is exceedingly soft. The upper parts are blackish in color, lightly
frosted with paler hairs. The underparts, from throat to vent, are
whitish tinged with gray or brown. The stiff, curved fringe of
bristles on the outer part of the hind foot serves as an aid in swim-
ming.

Mountain water shrews range over much of Canada and in moun-
tainous areas of the western United States extend south to Arizona.
The subspecies found in Washington ranges over all of the western
United States.

The mountain water shrew is primarily a mammal of the Hud-
sonian and Canadian life-zones. It sometimes descends to the
Transition Life-zone along clear, cold streams where conditions are
similar to those in the Canadian Life-zone.

The favored habitats of the mountain water shrew are the clear.
cold streams of the alpine cirques and gushing streams on the moun-
tain sides. In the pools and waterfalls, and among the rocks and
mosses that border them, the larvae of aquatic insects, upon which
the mountain water shrew feeds, are abundant. Svihla (1934: 4."))

observed that the fur of a swimming water shrew gathered air bub-
bles and "it had difficulty in forcing its way down to the bottom
[of an aquarium]. On reaching the bottom it literally stood on its
long flexible nose which was thrust into the -and and debris, search-
ing for food, its feet kicking rapidly in order to maintain this posi-
tion. A change in direction was brought about by a twist of the
body. To come to the surface again it merely stopped kicking and
immediately rose like a cork." The buoyancy of the water shrew-
allows it to float in the water, like a duck. Jackson (1928: 9)
observed a water shrew run across the surface of a small pool.
Near Stevens Lass a water shrew dashed from under a stone and
ran, did not swim, across the surface of a small, deep pool to escape
in a burrow on the other side. According to Jackson, an air bubble
held in each foot supports the shrew on the surface of the water.
A mountain water sinew observed at Shovel Creek. Ferry County,
was as agile on land as any other species of shrew. In summary,
water shrews are able to swim, dive, lloat like a duck, and walk on
the surface of the water as well as walk on land.

Shrews

1-J:j

The food of the mountain water shrew includes snails, leeches,
and the larvae of aquatic insects. Mice caught in traps are some-
times eaten by water shrews. Svihla iloc. cit.i found a captive
water shrew unable to capture pollywogs and minnows kept in the

same aquarium.

Sorex bendirii (Merriam)

Bendire water shrew

Description. — The Bendire water shrew is similar in form of body
to the cinereous shrew but possesses a more stocky body which, with
the head, measures about 3V2 inches long; the tail is about 2% inches
long. The Bendire water shrew closely resembles also the mountain
water sinew but has a longer body and shorter tail. The upper
parts are blackish in color, not lightly frosted with gray hairs. The
hind feet lack the fringe of stiff, curved bristles characteristic of
the mountain water shrew. The underparts are black in the race
S. b. bendirii. In the race albiventer the throat is blackish but the
abdomen is pale gray tinged with brownish.

DISTRIBUTION MAP
STATE OF WASHINGTON

Fig. 32. Distribution of the Bendire water shrew and the pigmy shrew in
Washington. A. Sorex bendirii bendirii. B. Sorex bendirii albiventer. C.
Microsorex hoyi washingtoni.

Bendire water shrews are restricted to the Pacific Coast of North
America from southern British Columbia to northern California.

144 University of Kansas Publs., Mtjs. Nat. Hist.

Generally they are found at elevation? lower than are mountain
water shrews. They are typically mammals of the humid division
of the Transition Life-zone but often occur in the Canadian Life-
zone. They occupy marshes, swamps, damp ravines, and the banks
of slow-moving streams. Little is known of their habits, except
what has been deduced from the circumstances of their capture.
They seem less aquatic than the mountain water shrew. Near Jack-
son Guard Station on the Hoh River, Jefferson County, one was
taken by setting traps on dense beds of water cress that floated in
a slow-moving stream. The animal must have swum or walked on
the surface of the mat of vegetation. Near Paradise Lake, King
County, several were caught in a dee]), dark, red cedar swamp. One
was caught in a marsh nearby. Nothing is known of the food habits
of the Bendire water shrew.

Sorex bendirii bendirii (Merriam)

Atophyrax bendirii Merriam, Trans. Linn. Soc. New York, 2:217. August 28, 1S84.
Atophyrax Bendirei True, Proc. U. S. Nat. Mus., 7 (1884):606, 1885.
Sorex bendirii Dobson, Monog. Insectivora, part 3, fasc. 1. pi. 23, 1890.
Neosorex bendirii bendirii Miller, U. S. Nat. Mus. Bull., 70:22. December 31, 1912.

Type. — Obtained approximately 1 mile from Williamson River. 18 miles
southeast of Fort Klamath, Klamath County. Oregon, by C. C. Bendire on
August 1, 1882; type in United States National Museum.

Racial dun deter*. — Underparts everywhere sooty black.

Measurenn nts. — Ten males and 10 females from southwestern Washington
average, respectively: total length 163.9, 131.0; length of tail 71.0. 72.6; hind
foot 20.5, 20.3; weight 16.8. 14.5 grams.

Distribution. — The southern Cascades and the lowlands of western Wash-
ington, exclusive of the Olympic Peninsula. Marginal localities on the west
include Mt. Vernon (Jackson. 1928: 196), Bothell (W.S.M.), Renton (M.
V.Z.), Puyallup (W.W.D.), Steilacoom (Jackson. 1928: 196) and Oakville
(Jackson, 1928: 196).

Sorex bendirii albiventer Merriam

Sorex (Atophyrax) bendirii albiventer Merriam, X. Amer. Fauna, 10:97, December 31, 1895
Neosorei bendirii albiventer Miller, V. S. Nat. Mus. Hull., 79:22, December 31, 1912.

Sorts luiulirii albiventer .Jackson, N. Amer. Fauna, 51 : 198, July 21. 1928

Tape. — Obtained at Lake Oushman. Mason County. Washington, by C. P.
Streator on July 7. 1894; type in United States National Museum.

Racial characters. — Area on abdomen whitish.

Measurements.— A male from tear the type locality measures: total length
167; length of tail 69; hind foot 22. One from Potlatch, Mason County,
mi asurea 167 ; 69; 22.

Distribution. The Olympic Peninsula of Washington. Marginal localities
on the SOUth are: Potlatch (M.V.Z.) and Lake Quinaull (Jackson. 1928: 199).

Shrews 145

Microsorex hoyi washingtoni Jackson
Pigmy shrew

Microsorex hoyi wasliingtord Jackson, Proc. Biol. Soc. Washington, 3S:i2. r >, November 13,
1925.

Type. — Obtained at Loon Lake, Stevens County, Washington, by V. Bailey
(,n September 26. 1897; type in United States National Museum.

Measurements. — Of type: total length 89; length of tail 27; hind foot 9.

Distribution^ — In Washington, known only from the type specimen which
was "found dead in a trail in dry pine woods" (Jackson. 1928: 4).

Description. — The pigmy shrew is similar in form of body to the
cinereous shrew but smaller. The head and body are about 1%
inches in length; the tail is about 1 inch long. The upper parts are
reddish brown and the underparts are gray.

These tiny mammals range widely across central Canada and
northern United States from the Atlantic nearly to the Pacific, and
north to central Alaska. A single species is known, one race of
which occurs in Washington. The subspecies is known from but
two specimens: the type and an individual from Montana (Koford,
1938: 372.)

10—4993

146 University of Kansas Publs., Mis. Xat. Hist.

Genus Myotis Kaup

Mouse-eared bats

Description. — The genus Myotis may he separated from all other
hats that occur in Washington by the presence of 38 teeth (dental

2-2 1-1 3-3 3-3

formula «■ — . c — .p. — ,m. -- = 38). Their small size separates them

3-3 1-1 3-3 3-3 '

from all other genera save Pipistrellus, from which Myotis may be
distinguished by the straight, rather than hooked, anterior border of
the tragus. Species of Myotis found in Washington vary consider-
ably in size, but all are less than 100 mm. in total length. The up-
per parts are various shades of brown in color. The ears, when laid
forward, always extend to the nostrils or beyond.

This genus is one of the most widely ranging groups of Recent
mammals. It occurs on all continents, including Australia and
many of the larger islands. Of the 19 American species recognized
by Miller and Allen (1928), eight occur in the state of Washington.
They are low-flying forms and as a rule appear relatively late in
the evening. Their flight is rapid and erratic. They often hunt over
the surfaces of streams, pools, and lakes. Some kinds hunt in the
shade of forest trees and these are especially difficult to collect.
Others hunt the brushy canyons and coulees of the desert areas of
eastern Washington.

Grinned (1918: 241-242) points out that, although bats are not
subject to isolation by topographic barriers, as most wingless mam-
mals are. they may be restricted by ecologic barriers in the same
way as are other small mammals. This is particularly true of
Myotis in Washington. Of the eight species in the state, five are
represented by one subspecies west of the Cascade Mountains and
another, paler subspecies in the more arid country east of the
Cascades.

The mobility of bats makes it difficuH to determine their origin
and migrational history. Five of the Myotis found in Washington
seem to belong to the Pacific Coastal Fauna, and to have been iso-
lated south of the last continental glacier. Complete isolation is
unlikely as these species occur in the Cascade Mountains as well
as in the Pacific Coastal Fauna] Area, and three occur also in the
Blue Mountains of southeastern Washington. The differentiation
of the Coastal type of Myotis may have come about through habitat
selection, of the type discussed by Miller (1942: 25). One western

Bats

147

Washington Myotis (M. keenii) seems to belong to a northern fauna,
and to have extended its range south to Washington. All seven
species of the desert-living Myotis found in eastern Washington
have subspecies which seem to have been derived from the Great
Basin Fauna! Area.

Myotis lucifugus (Le Conte)

Big myotis

Myotis lucifugus is represented by two geographic races in Wash-
ington. The species ranges across Canada and the United States,
from the Atlantic to the Pacific and from the northern limit of tree
growth to southern Mexico.

Fig. 33. Distribution of the big myotis in Washington. A. Myotis lucifugus
alascensis. B. Myotis lucifugus carissima.

It usually proves rather difficult to separate Myotis lucifugus, on
the basis of external features, from other species with which it may
occur. Its large foot (9-10 mm.) , short car (when laid forward not
extending past nose) and the absence of a keel on the calcar
separate it from all species except Myotis yumanensis. From the
latter species, lucifugus may be distinguished by the gradually

148 University of Kansas Publs., Mis. Nat. Hist.

lather than abruptly rising forehead, as seen in cleaned skulls, and
by more shiny, metallic color of fur.

Little is known of the habits of this bat in Washington. It usually
appears after dusk, and most specimens are shot over ponds or lakes,
where the reflection of light from the sky on the water allows the
hunter enough light to sight a gun. A few specimens were collected
in deep forests. Its flight and feeding habits are not known to dif-
fer from those of other species with which it was associated, except
at the south end of Lake Chelan, Chelan County, where two in-
dividuals were shot as they hovered near the tops of pine trees and
seemed to be picking insects from the branches. I have never found
this bat in its daytime retreat.

A specimen taken at Sportman's Lake, San Juan County, held one
embryo on June 26, 1938.

Myotis lucifugus carissima Thomas

Myotis (Leuconoe) cari-ssitna Thomas, Ann. and Mag. Nat. Hist., 13 (sit. 7): 383, May,
1904.

Myotis lucifugus carissima Cary, N. Amer. Fauna, 42:43, October 3, 1917.

Type. — Obtained at Yellowstone Lake. Yellowstone National Park. Wyo-
ming, by J. Darling in September, 1903; type in British Museum.

Racial characters. — Color pale, rather "brassy" in tone; distal border of
interfemoral membrane paler than proximal part.

Measurements. — Four specimens from eastern Washington average: total
length 77; length of tail 33; hind foot 11; ear 13; height of tragus 7.3.

Distribution. — Easl of the eastern base of the Cascade Mountains, save
for the Blue Mountains of southeastern Washington. Western records are
Stehekin (Miller and Allen. 1928: 52) and Vantage (W.W.D.)

Myotis lucifugus alascensis Miller

Myotis lucifugus alascensis Miller, X. Amer. Fauna, 13:63, Octobei 16, 1897.
Vespertilio gryphus lucifugus Allen, I'. S. Nat. Mus. Bull., i:-i:7s. March 14, ls'.'t (pan
specimens from Washington).

Type. — Obtained at Sitka. Alaska, by C. P. Streator on August 5, 1895;
type in United States National Museum.

Racial characters. — Color dark, almost bronze; wing and tail membranes
uniformly dark in color.

Measurements. — Five specimens from San Juan County, Washington,
average: total length 80.9; length of tail 32.1; hind ....it 12; ear 12; height
of tragus 7; weight 5.4 grams.

Distribution. From tin eastern base of the Cascade Mountains west of the
Pacific, and the Blue Mountains of southeastern Washington. Marginal occur-
rences listed by Miller and Allen (1928:49) are Chilliwack River, bake
Wenatchee, and l.vle.

Bats

149

Myotis yumanensis (H. Allen)

Yuma myotis

Description. — The present species closely resembles Myotis luci-
fugus and specimens in worn pelage can not be distinguished from
that species unless the cleaned skulls are examined. In fresh
pelage, yumanensis is duller than lucifugus.

Myotis yumanensis ranges from southern British Columbia to
central Mexico west of the Mississippi River. Four races are
recognized by Miller and Allen (1928: 62).

DISTRIBUTION MAP
STATE OF WASHINGTON

Draf'ed U~d«

6079-J797 b«Ci1.T. IJ37

Fig. 34. Distribution of the Yuma myotis in Washington. A. Myotis yuma-
nensis saturatus. B. Myotis yumanensis sociabilis.

The habits of Myotis yumanensis and Myotis lucifugus appear to
be the same. In Washington the two species are commonly found
together. In western Washington, Myotis yumanensis seems to be
more common than Myotis lucifugus.

In the San Juan Islands a yumanensis was found hiding in the
attic of an old cabin on Blakeley Island. A specimen of long-eared
bat was taken at the same place. Another Yuma myotis was caught
behind a door of a mill on Blakeley Island (Dalquest, 1940: 4).

150

University of Kansas Ptjbls., Mrs. Nat. Hist.

This species shares with Myotis californicus the habit of apparently
drinking salt water.

A specimen obtained at Sportsmans Lake, San Juan County, held
one embryo on June 27, 1938. One from Peavine Pass, Blakeley
Island. San Juan County, held one embryo on June 22. 1939.

Myotis yumanensis sociabilis H. W. Grinnell

[lyotit yumanensis sociabilis II. W. QSrinnell, Univ. California Publ. Zool., 12:318, De-
cember 4. 1914.

Ti//h . — Obtained at old Fort Tejon. Kern County, California, by J. Grin-
nell on July 23. 1904; type in Museum of Vertebrate Zoology.
Uncial character. — Color pale.

DISTRIBUTION MAP
STATE OF WASH I NGTON

D.o*'»d Under »..p I

MILES

c i WBA P..

Fig. 35. Distribution of the fringe-tailed myotis and the Keen myotis in
Washington. A. Myotis thysanodes thysanodes. B. Myotis keenii keenii.

Measurements. — Two mules and 2 females from Selah, Yakima County, av-
erage: total length 78; length of tail 36; hind fool 10; ear II; height of

t latins 7.

Distribution.— Eastern Washington generally. Marginal records on the west
ar< : Stehekin (Miller and Allen, 1928: (19), and Selah (W.W.D.).

Myotis yumanensis saturatus Miller
Myotk yumanensis saturatus Miller, V Amer, Fauna, 13:68, Octobei 16, 1897.

TyjH obtained al Hamilton, Skagit County, Washington, by T. S. Palmer
on September 13, 1889; type in United States National Museum.
Racial character. Color dark.

Bats 151

Measun merits. — Eighteen adults of both sexes from San Juan County,
Washington, average: Total length 78.2; length of tail 34.4; hind foot 10.1;
ear 15; height of tragus 7.4; weight 5.9 grams.

Distribution. — From the eastern base of the Cascade Mountains to the
Pacific. This is the commonest Myotis found in western Washington. Mar-
ginal localities are: Hamilton (Miller and Allen. 1928: 71). and Goldendale
(Miller and Allen. 1928: 71).

Myotis keenii keenii (Merriam)

Keen myotis

Vespertilio subulatus keenii Merriam, Amer. Nat., 29:860, September, 1895.

Myotis subulatus keetui Miller, N. Amer. Fauna. 13:77, October 16, 1S97.

Myotis keenii keenii Miller and Allen, U. S. Nat. Mus. Bull., 144:104, May 25, 1928.

Type —Obtained at Masset, Graham Island, Queen Charlotte Islands, British
Columbia, by J. H. Keen in 1894; type in United States National Museum.

Measurements.— Miller and Allen (1928: 109) list the measurements of a
male from Sol Due Hot Springs, Clallam County, and a specimen of unknown
sex from Lake Cushman, Jefferson County, as. respectively: total length 89,
87; length of tail 34. 36; hind foot 8.4. 7.4; ear ?. 14.6.

Distribution. — Only the Olympic Peninsula, where it has been recorded by
Miller and Allen (1928: 104) from Sol Due Hot Springs and Lake Cushman.

Description. — Myotis keenii is similar, in general, to Myotis luci-
fugus and Myotis yumanensis, but the ears are longer and when laid
forward reach about 4 mm. past the nose rather than ending at the
nostrils. The foot is of medium size (about 8 mm.) and no keel is
present on the calcar.

The distribution of this species is given by Miller and Allen (1928:
101) as "northern North America from the limits of tree growth
south in the east to South Carolina and Arkansas, and in the west
to northwestern Washington."

I have not observed this bat in Washington and know nothing of
its habits. Its distribution is most unusual. Its range seems to lie
only in the glaciated area of western British Columbia and north-
ern Washington.

Myotis evotis (H. Allen)

Long-eared myotis

Description. — The distinguishing feature of Myotis evotis is its
long ears, which, when laid forward, reach 5 mm. in front of the
nose. Myotis thysanodes and Myotis keenii, other species in which
the ears are rather long, have the ears ending less than 5 mm. an-
terior to the nose when laid forward. The foot of Myotis evotis is
of moderate size (8 to 9 mm.).

This species ranges over the western United States, from British
Columbia to central Mexico. Two subspecies of this interesting bat
are recognized, both of which occur in Washington.

152

University of Kansas Publs., Mrs. Nat. Hist.

Though I have hunted for this species of bat in Washington on
numerous occasions, I have taken no specimens. In the summer of
1939, Myotis identified as this species because of their large ears,
were seen flying at midnight in the light of searchlights over Lake
Washington Canal at Seattle. Mary Greer gave us a specimen
which was struck by her auto near Baker Lake, Whatcom County.
According to Miss Greer, the specimen was seen "hovering in the
road, like a large moth." The time was about midnight. This
evidence indicates that the species does its hunting late at night,
when ordinary methods of hunting bats are useless, and may account
for the scarcity of specimens from the state. Nevertheless, collectors

DISTRIBUTION MAP

STATE OF WASHINGTON

rtP* Prtjaci »cvir>7 b 3 cnT nn

I'M.. 36. Distribution of the long-eared myotis in Washington. A. Myotis
evotis < votis. B. M i/<>ti.< < votis pacificus.

from the California .Museum of Vertebrate Zoology took specimens
in the Blue Mountains where the bats flew slowly, in rather straight
courses, 20 to 25 feel from the ground.

This species li;is not, so far as is known, been taken in Washington
iii its daytime retreat. Daniel Bonell saved two specimens from under
slabs of loose bark on old. dead snags near Tillamook, ( Iregon. Davis
M!K{!): 214! reported them as hiding in the daytime in a cave in

Bats 153

Craters of the Moon National Monument, Idaho. Whitlow and Hall
(1933: 241) report specimens found in an old cabin near Pocatello,
Idaho, two of them containing one embryo each.

Myotis evotis evotis (H. Allen)

VespertUio evotis Allen, Smithsonian Misc. Coll., 7 (no. 165):48, June, 1864.

Myotis evotis Miller, N. Amer. Fauna, 13:77, October 1G, 1897.

Myotis evotis evotis Miller and Allen, U. S. Nat. Mus. Bull., 144:114. April 14. 1928.

Type. — Description based on a series of specimens, one of which came from
Monterey, California. This locality was designated the type locality by
Miller (1897: 78).

Racial character. — Color pale.

Measurements. — Two males and 2 specimens of unknown sex from the
Blue Mountains. Columbia County, average: total length 87; length of tail
40; hind foot 7.5; ear 20; height of tragus 11; weight 5.4 grams.

Distribution. — The Blue Mountains area, of southeastern Washington; re-
corded from South Touchet (Miller and Allen. 1928: 116) and Godman Springs
(W.S.M.).

Remarks.— Miller and Allen (1928: 116) record the dark race of Myotis
evotis from the Blue Mountains. Specimens examined by me are much paler
than pacificus and most of them are indistinguishable from specimens of evotis
from California.

Myotis evotis pacificus Dalquest

Vespertilio evotis Allen, Smithsonian Misc. Coll., 7 (no. 165): 48, June, 1864 (part specimens
from Puget Sound).

Myotis evotis evotis Miller and Allen, U. S. Nat. Mus. Bull., 144:114, May 25, 1928.
.\[yotis evotis pacificus Dalquest, Proc. Biol. Soc. Washington, 56:2, February 25, 1943.

Type .—Obtained from 3^ miles east and 5 miles north of Yacolt, Clark
County, Washington, by John Chattin on August 3, 1940; type in Museum of
Vertebrate Zoology.

Racial character. — Color dark.

Measurements.— Five specimens from the type locality average: Total
length 85; length of tail 41; hind foot 7.4; ear 19.4; height of tragus 10;
weight 5.5 grams.

Distribution. — Western Washington from the Cascade Mountains westward.
Marginal occurrences are: Baker Lake (W. W.D.) and Easton (Miller and
Allen. 1928: 116).

Myotis thysanodes thysanodes Miller
Fringe-tailed myotis

Myotis thysanodes Miller, N. Amer. Fauna, 13:80, October 16, 1897.

Myotis thysanodes thysanodes Miller and Allen, TJ. S. Nat. Mus. Bull., 144:126, May
25, 1928.

Type.— Obtained at Old Fort Tejon, Kern County, California, by T. S.
Palmer on July 5, 1891; type in United States National Museum.

Measurements.— Two males and 3 females from Vernon, British Columbia,
average, respectively: total length 90.5, 82; length of tail 41.5, 37; hind foot
10.5. 10; ear 18.5, 16; height of tragus 14, 13.

154 University of Kansas Publs., Mrs. Nat. Hist.

Distribution. — In Washington known only from the southeastern border of
the state, namely from Dayton (W.S.M.) and Anatone (Miller and Allen.
1928: 127).

Description. — Myotis thysanodt s resembles Myotis evotis, but dif-
fers in larger size, smaller ear (reaching less than 5 mm. past nose
when laid forward), and in possessing a well developed fringe of
hairs along the border of the caudal membrane.

This species of bat ranges over western North America from south-
ern British Columbia to southern Mexico. Two geographic races are
currently recognized, only one of which occurs in the United States.
This bat is not recorded by Davis (1939) as occurring in Idaho and
has been found only once in Oregon. The only published account
of the habits of thymnodes seems to be that of Palmer (in Miller.
1897: 84, also Grinnell, 1918) who found adults and young of various
sizes in company with Myotis yumanensis in the attic of an old
adobe building near Old Fort Tejon, California, in July, 1891. The
specimens obtained in Washington and British Columbia came from
dry areas of pine forest.

Myotis volans ( H. Allen I

Hairy-winged myotis

Description. — This species, in Washington, may be easily recog-
nized by its relatively large size and the presence of a distinct keel
on the side of the calcar, posterior to the loot.

Four subspecies of Myotis volans are recognized by Miller and
Allen (1928: 136). These range over western North America from
southern Alaska to southern Mexico. Two subspecies occur in
Washington.

The record stations in Washington for the pale, southern race are
till in arid places and the dark, coastal race is a forest animal. Most
of the specimens taken by me (all of the dark race) were in clear-
ings or along roads through timber near the crests of hills. They
appeared relatively late in the evening, after the big-brown and
the silver-haired bats had been in the air for some time. Often
they were taken in company with Myotis ludfugus ami Mi/<>tis
ijumanensis. They were appreciably larger than those specie- and
their flight was slower and less erratic. They usually Hew in rela-
tively straight lines or huge circles at from ten to forty feet from
the ground. At hake Kapowsin, Pierce County, they were attracted
by swishing a long pole in the air. At Kenton, King County, one
was shot as it hunted insects at a city street light several hour-
after dark.

Bats

155

Myotis volans Iongicrus (True)

Yc&pertilio Iongicrus True, Science, S : 588, 1886.

Vespertilio nitidus Iongicrus H. Allen, U. S. Nat. Mus. Bull., 43:103, Marcli 14, 1S94.

Myotis lucifugus Iongicrus Miller, X. Amer. Fauna, 13:64, October 16, 1897.

Myotis Iongicrus Lyon and Osgood, U. S. Nat. Mus. Bull., 62:271, January 28, 1909.

Myotis volans Iongicrus Miller and Allen, U. S. Nat. Mus. Bull.. 144:140, May 25. 1928.

Type — Obtained in t ho "vicinity of Puget Sound, Washington" by D. S.
Jordan, and catalogued in the U. S. National Museum on December 16, 1S86.

Racial character. — Color dark.

Measun rru nts. — A female from 6 miles northeast of Kelso, Cowlitz County,
measures: total length 95; length of tail 39; hind foot 8; ear 13; height of
tragus 8.

Distribution. — From the eastern base of the Cascade Mountains to the
Pacific. Marginal records (from Miller and Allen. 1928: 142) are Oroville,
Entiat, and Carson.

DISTRIBUTION MAP

STATE OF WASHINGTON

miles

of WPA P-.

60'»-S7)7 by CUT 1,37

Fig. 37. Distribution of the hairy-winged myotis in Washington. A. Myotis
volans Iongicrus. B. Myotis volans interior.

Myotis volans interior Miller

Myotis Iongicrus interior Miller, Proc. Biol. Soc. Washington. 27:211, October 31, 1914.
Myotis volans interior Miller and Allen, U. S. Nat. Mus. Bull., 144:142, May 25, 1928.

Type.— Obtained 5 miles south of Twining. Taos County, New Mexico, by
Vernon Bailey on July 23, 1904; type in United States National Museum.
Racial characters. — Smaller and paler than Myotis v. Iongicrus.

156 University of Kansas Publs., Mis. Nat. Hist.

Measun merits. — Four males and a female from the Blue Mountains, Colum-
bia County, average: total length 93; length of tail 41.5; hind foot ?; ear
12; height of tragus 6.3.

Distribution. — Known only from the Blue Mountains area of the south-
eastern part of the state, from Walla Walla (E. S. B.) east to Anatone (Miller
and Allen, 1928: 144).

R< murks. — Of 5 specimens available from the Blue Mountains. 4 are like
inU rior and 1 is like longicnis.

DISTRIBUTION MAP
STATE OF WASHINGTON

MILES

Fig. 38. Distribution of the California myotis in Washington. A. Myotis
californicus calij ornicus. B. Myotis calijornicus caurinus.

Myotis californicus (Audubon ami Bachman)

California myotis

Description. — Myotis calijornicus may be separated from all My-
otis that occur in Washington, except Mi/otis subulatus, by its small
foot (about 6 mm.). It is the only small-footed bat found in west-
ern Washington. In eastern Washington, where Myotis subulatus
occurs, the cleaned skulls of the two species must be compared be-
fore certain identification of some specimens i- possible. The skull
of 1\F. californicus possesses a higher cranium and more abruptly
rising forehead than that of M. subulatus.

Four geographic races of this bat recognized by Miller and Allen
(1928: 149) range from southern Alaska southward over western

Bats

157

North America to southern Mexico. Two subspecies occur in Wash-
ington.

Myotis californicus, in western Washington, often occurs in com-
pany with Myotis yumanensis, lucifugus, and volans. In flight it
cannot be distinguished from M. yumanensis or M. lucifugus. Most
of our specimens were collected over water, for these bats usually
fly rather late and can be shot most easily where their reflection
on the water assists the hunter in aiming. They are usually not
common, one or two being taken at a single locality. This species,
like Myotis yumanensis, seems to drink salt water. On May 9,
1936, a living specimen was caught under a loose piece of bark on

a dead tree.

Myotis californicus caurinus Miller

Yespertilio nitidus H. Allen, Proc. Acad. Nat. Sci. Philadelphia, p. 247, 1862 (part of the
specimens were from Fort Stiilacoom, Pierce Co., Washington).

Myotis californicus caurinus Miller, N. Amer. Fauna, 13:72, October 16, 1897.

Type. — Obtained at Masset, Graham Island, Queen Charlotte Islands, British
Columbia, by J. H. Keen in 1895; type in United States National Museum.
Racial character. — Color reddish-brown.

DISTRIBUTION MAP
STATE OF WASHINGTON

cd U«d«- Awip.'dl o' WM Prcjtcl 60*9 -5797 bjC*** Hi*

Fig. 39. Distribution of the small-footed myotis, Myotis subulatus melanorhi-

nus, in Washington.

158 University of Kansas Publs., Mus. Nat. Hist.

Measurements. — Eight specimens, including both sexes, from the San Juan
Islands. San Juan and Skagii counties, average: total length 77.8; length of
tail 36.7; hind foot 6.7; ear ?; height of tragus 7.6; weight 5.2 grains.

Distribution. — Western Washington east through the northern Cascades to
Chelan. Blue Creek and Colville (Miller and Allen. 1928: 1.56) in northeastern
Washington, and. farther south, oast to Mount Rainier (Miller and Allen.
1928: 156) and Carson (Miller and Allen. 192S: 156).

Myotis californicus califomicus (Audubon and Bachman)

Vesperttiio californicus Audubon and Bachman, Jour. Acad. Nat. Sci., Philadelphia, - (ser. 1,

pt. i):lsr,. 1S4-2.

Myotis californicus Miller, X. Amer. Fauna, 13:69, October 10, 1SK7.

Type. — None designated. Type locality fixed at Monterey, Monterey
County. California, by Miller and Allen (1928: 153).

line, til character. — Color pale.

Measurements. — A female from Crooked River. Crook County, Oregon,
measures: Total length 80; length of tail 40; hind foot 7; ear 13; height of
tragus 5; weight 3.2 grains.

Distribution. — Recorded only from the eastern part of the state. Western-
most records, according to Miller and Allen (1928: 155) are: Orondo. Gold-
endale and Lyle.

Myotis subulatus melanorhinus (Merriam)

Small-footed myotis

Vetpertilio melanorhinus Merriam, N. Amer. Fauna, 3:40, September II, 1890.
Myotis subulatus melanorhinus Miller and Allen, U.S. Xat. Mus. Bull., 144:169, May 2">.
1928.

Type. — Obtained on San Francisco Mountain. 8250 It. elevation. Coconino
County, Arizona, by C. H. Merriam and V. Bailey on August 4. 1889; type
in United Stales National Museum.

Measurements. — Three males and 2 females from eastern Washington aver-
age: total length 77; length of tail 37; hind foot 7; ear 15; height of tragus
S. One weighed 5.4 grams.

Distribution. — Desert areas of eastern Washington, including the Columbian
Plateau and the lower Columbia River Valley. Marginal records are: 5 mi. S
Grand Coulee Dam (W. WD.) in the north. Wenatche< (W.W.D.) in the
northwest, Lyle (W.W.D.) in the southwest, ami Bly (Miller and Allen. 1928:
171) in the southeast.

Description. — Myotis subulatus is closely similar to Myotis cali-
fornicus, but is more orange in color and has the skin on the face
more nearly black. Specimens cannot be identified with certainty
until the cleaned skulls are examined. The more flattened cranium
and less abruptly rising forehead separate Myotis subulatus from .1/.
californicus.

This species is confined in the United States and northern Mexico.
Two races are recognized by Miller and Allen (1928), of which one
occurs in Washington. It is the commonest bat in the desert of

Bats 159

eastern Washington. It lives far from trees on plains and in sandy
or rocky areas, emerging rather early in the evening. It is not diffi-
cult to shoot. Its flight is erratic. The animal usually hunts in large,
irregular circles at 10 to 25 feet from the ground. It has not been
found in its daytime retreat but may hide in crevices in rocky out-
crops. Near Vantage, Grant County, individuals were shot as they
hung up in a concrete underpass todigest food. The stomachs of speci-
mens taken were so crammed with the remains of insects that their
abdomens were greatly distended. The underpass seemed to be only
a resting place, not inhabited by day. Others were taken when they
came to rest in the loft of a barn at Selah, Yakima County. They
usually did not arrive at the barn until an hour after sunset and were
-till present there at midnight,

Lasionycteris noctivagans (Le Conte)
Silver-haired bat

V [e&pertilio] . noctivagans LeConte, McMurtrie's Cuvier, Anim. Kingd., 1:431, 1831.

Ycsperugo noctivagans True, Proc. II. S. Nat. Mus., 7:602, 1885.

Lasionycteris noctivagans Allen, U. S. Nat. Mus. Bull., 43:105, March 14, 1894.

Type- — None designated; described from a specimen obtained in the "east-
ern United States."

M< asurcments. — Five males from San Juan County, Washington, average:
total length 96.2; length of tail 46.4; hind foot 8.5; ear 16; height of tragus
7.2; weight 8.2 grams.

Distribution. — Forested areas of the entire state. This species is migratory
and first appears about the middle of May. September 15th is the latest re-
corded occurrence, when one was seen at Seattle, King County. Marginal
records are: Sportsmans Lake (W.W.D.), in the northwest; Carson (Taylor
and Shaw. 1929: 9) in the southwest; Pass Creek Pass (W.W.D.) in the
northeast; and Bly (Taylor and Shaw. 1929: 9) in the southeast.

Remarks. — The chocolate-brown color phase of the silver-haired
bat is most common east of the Cascade Mountains, while almost all
specimens from western Washington represent the black phase.

This medium-sized bat is the darkest-colored species living in
the state, ranging from chocolate-brown to nearly black. Nu-
merous white-tipped hairs give the upper parts a frosted appear-
ance. The upper surface of the interfemoral membrane is well
furred, a character shared only with Lasiurus. The dental formula

o_9 1—1 2—2 3—3

d. — -, c — , p. — ,m. -- = £6) is the same as that of Corynorhinus. The

3-3 1-1 3-3 3-3

small ears and short tragus immediately ' distinguish the silver-
haired bat from the long-eared bat.

The genus Lasionycteris contains but a single species, of which

160

University of Kansas Publs., Mrs. Nat. Hist.

no geographic races have been described. It ranges across North
America from coast to coast and from central Canada southward.
in forested areas, nearly to Mexico. This species is known to be mi-
gratory. -and southern records probably do not represent breeding
individuals.

In Washington these bats have been taken in the Transition.
Canadian, and Hudsonian life-zones. They were found near clear-
ings in forests of open pine woods in the arid section of eastern
Washington, near mountain hemlock thickets in the high mountains,
and in the dense rain-forests of the Pacific Coastal area.

DISTRIBUTION MAP
STATE OF WASHINGTON

J I

Fi<;. 40. Distribution of the silver-haired bat, Lasionycteris noctivagans,

in Washington.

The size and flight of the silver-haired bats are distinctive, and
after some experience it is possible to identity them in the air. It
is an early flier, usually appearing just after the swallows roost.
They fly a1 a considerable height, rarely coining within forty feet
of the ground. The wings are moved with a "flnttery" motion, and
their flight is interrupted by frequent short glides. They fly more
rapidly than the big brown-bats, and twist and dart sideways more
frequently. Compared with big brown-bats, silver-haired bats are
relatively gregarious, and six to a dozen individuals were seen in the

Bats

161

same area. They generally hunt in sweeping circles, from fifty to
one hundred yards in diameter. In the daytime the silver-haired
bats hide beneath slabs of loose bark on dead trees. Near Cottage
Lake, King County, two individuals were found beneath the bark
on an old, lightning-blasted stub. The dark color of the bats
blended with the charred surface of the stub.

This species feeds mainly on forest insects and for this reason is
probably of considerable value to man. The stomachs of specimens
were usually crammed with the remains of small, soft-bodied insects.

Corynorhinus rafinesquii (Lesson)

Long-eared bat

Description. — Diagnostic characters of the long-eared bat are:
medium size (total length about 4 inches) ; dull, grayish-brown
color; exceptionally long ears (over 1 inch from notch) ; thin, tissue-
like membranes; and paired "lumps" on the rostrum. The dental

, , . 2-2 1-1 2-2 3-3

formula is : i- — , c ■ — , p. — , m. — - 36.

3-3 1-1 3-3 3-3

S':"<A : ''$%

Fig. 41. Long-eared bat {Corynorhinus rafinesquii intermedins), female
with young; Boulder Cave, Kittitas County, Washington, July 20, 1928. (Fish
and Wildlife Service photo by T. H. Scheffer, No. B-33332.)

Long-eared bats range from southern British Columbia to southern
Mexico. Three species are listed by Miller (1924: 82), one of which
(rafinesquii) is divisible into five geographic races.

This bat is colonial and is not uncommonly found in caves, mine
shafts, and darkened attics of old buildings. It is of scattered dis-
tribution throughout the state. In Washington its distribution in
winter is unknown. A specimen from Friday Harbor, San Juan
County, taken in March, 1936, indicates that it hibernates in the

11—4993

162

University of Kansas Publs.. Mrs. Nat. Hist.

state. Whitlow and Hall (1933: 24.1i give a detailed account of
individuals found hibernating in winter near Pocatello, Idaho.
John K. Townsend (1839: 325) mentions that the "great-eared

hat - ' at the huts of the Columbia River district i Fort Vancouver)
were protected by the "gentlemen of the Hudson'- Bay Company
for their services in destroying the dermestes which abound in their
fin- establishments." Townsend mention- also that the long-cared
hat- seldom left the "storehouses attached to the fort-." even at
night. My own observations are similar, in that at Boulder Cave,
Kittitas County, on July 7. 1 !»:{(;. when not less than TOO long-eared

"S**v «ft~-3»

Fig. 42. Entrance to Boulder Cave, inhabited by long-eared bats (Cory-
norhinus); Kittitas County. Washington, May 2G. 1938. (Fish and Wildlife
Service photo by Victor 15. Scheffer, No. 56.)

hats were present in the cave, 1 watched the entrances to the cave

until an hour after dark hut no hat- were seen to emerge. All of the

specimens available from Washington were caught in their daytime
hiding place. The number of long-eared hat- at Boulder Cave
has decreased in recent years, (hi July 12, 1930, hats were so
abundanl that 90 were captured with a single sweep of a butterfly
i,et (T. 11. Scheffer. 1930: Hi. <>n .Inly 7. 1936, it was estimated
that there were slightly more than 100 in the cave. On June 11,

1937, the number was less, probably abou! 75.

Little i- known of the food habits of the long-eared hat. The

stomach of a specimen from Blakeley I.-land. San Juan County, was

Bats

lr.3

crammed with the remains of insects, including the wing scales of
Lepidoptera and the wings of small Diptera.

Scheffer noted that the long-eared bats at Boulder Cave were
nearly ready to give birth to young on July 12, 1930, and were carry-
ing naked young a week later. On July 7, 1936, at the same locality,
females contained nearly full-term embryos.

DISTRIBUTION MAP
STATE OF WASHINGTON

ed Under A-

MILES

cf WPA P.c

6C9-5797 b a CHT mi
I

Fig. 43. Distribution of the long-eared bat in Washington. A. Corynorhinus
rafinesquii townsendii, B. Corynorhinus rafinesquii intermedins.

Corynorhinus rafinesquii townsendii (Cooper)

Plecotus townsendii Cooper, Ann. Lye. Nat. Hist. New York, 4:73, November, 1837.
Corynorhinus macrotis townsendii Miller, N. Arner. Fauna, 13:53, October 16, 1897.
Corynorhinus megalotis townsendii G. M. Allen, Bull. Mus. Comp. Zool., 60:344, April,
1916.

Corynorliinus rafinesquii townsendii Miller, U. S. Nat. Mus. Bull., 128:82, April 29, 1924.

Type.— Probably obtained at Fort Vancouver. Clark County, Washington;
type not now in existence.

Racial character. — Dark color.

M < asurements.—A male from Blakeley Island, San Juan County, measured:
total length 83; length of tail 43; hind foot 8.5; car 37.4; tragus 15.1; weight
10 grams.

Distribution.— Western Washington, from Blakeley Island (W. W.D.) in
the north, south to Seattle (W.W.D.) and Fort Vancouver.

164

University of Kansas Publs., Mis. Nat. Hist.

Corynorhinus rafinesquii intermedius H. W. Grinnell

Corynorhinus rafinesquii intermedius H. W. Grinnell, Univ. California Publ. Zool., 12:320,
December I. L914.

Corynorhinus rafinesquii townsendii Dalquest, Jour. Mamm., 19:213, May 14, i:»38.

Type. — Obtained at Auburn, Placer County, California, by J. C. Hawver
en July 31. 1909; type in Museum of Vertebrate Zoology.

Racial characters. — Paler and duller than townsendii.

Measurements. — One male and 6 females from 15 miles east of Tonasket,
Okanogan County, average: total length 96, length of tail 47; hind foot 11;
i ar 35.5; tragus 14.

Distribution. — Scattered localities in the arid subdivision of the Transition
Life-zone of eastern Washington, from 15 mi. E Tonasket (W.W.D.) on the
north to Boulder Cave (W.W.D.) on the west and Spokane (W.S. C.) on
the east.

Remarks, — Of specimens in the University of Kansas, Museum of Natural
History, those from Selah (not plotted on distribution map). Yakima County,
are paler than those from Boulder Cave, Yakima County, but both series are
paler than specimens from the coast of Oregon.

DISTRIBUTION MAP
STATE OF WASHINGTON

0'«ft«4 U' d,, *.

I I

Fig. 44. Distribution of the western pipistrelle, Pipistrellus hesperus hesperus,

in Washington.

Bats 165

Pipistrellus hesperus hesperus (H. Allen)
Western pipistrelle

Scotophilia* hesperus H. Allen, Smithsonian Misc. Coll., 7 (no. 1G5):43, June, 1864.
Vesperugo hesperus True, Proc. U. S. Nat. Mus., 7:602, 1885.
Pipistrellus hesperus Miller, N. Amer. Fauna, 13:S«, October lii, ls!>7.

Type. — Obtained at Old Fort Yuma, Imperial County, California, by G. H.
Thomas; catalogued in U. S. National Museum on October 31. 1861.

Measun incuts. — A specimen from Maryhill, Klickitat County, measures
total length 68; length of tail 27; hind foot 7; ear 10; height of tragus 3
weight 4.2 grams. Two males from Vantage. Grant County, average: 68.5
27.5; 6; 11; 4.

Distribution. — Known only from along the Snake and Columbia rivers
of south-central Washington; recorded from Vantage (W.W.D.), south to
Maryhill (M.V.Z.), and east to Almota (Taylor and Shaw, 1929: 9).

Description. — This is the smallest bat found in Washington, its
body being approximately 1% inches long and the tail l 1 /^. It may
be separated from Myotis by the bent tragus and by the
possession of 34 rather than 38 teeth. The dental formula is:

2-2 1-1 2-2 3-3

i. , c. . p. , m. = 34.

3-3 1-1 2-2 3-3

The genus Pipestrellus is cosmopolitan in distribution. The few
records for the single subspecies found in Washington indicate that
it is a casual, though probably regular, summer visitant from the
south.

Two western pipistrelles were shot at Vantage, Grant County,
on July 23, 1937. They flew in slow circles about 50 feet from the
ground. Xo breeding records are known from the state.

Eptesicus f uscus bernardinus Rhoads
Big brown-bat

Eptesicus fuscus bernardinus Rhoads, Proc. Acad. Nat. Sci. Philadelphia, 1901:619, Feb-
ruary 6, 1902.

Eptesicus fuscus pallidas Engels, Amer. Midland Nat., 17:656, May, 1936 (part specimens
from Washington).

Type. — Obtained near San Bernardino, San Bernardino County, California,
by R. B. Herron on May 26, 1893; type in Academy of Natural Sciences of
Philadelphia.

Measurements. — Three males and 5 females from Washington average :
total length 117; length of tail 48; hind foot 11.5; ear 18.3; height of tragus 9.

Distribution. — Forested areas of the entire state of Washington. Marginal
localities are San Juan Island (W.W.D.) in the northwest, Carson (Taylor
and Shaw, 1929: 9), in the southwest, Newport (W.W.D.) in the northeast,
and Grand Ronde River (Taylor and Shaw, 1929: 9) in the southeast.

Remarks. — Specimens from both eastern and western Washington vary
greatly in color, and series of specimens from eastern Washington average

Kib

University of Kansas Publs., Mis. Nat. Hist.

little, if any. paler than series from western Washington. Washington speci-
men.- iini-1 closely resemble specimens from California of the race bt rnardinus
and average darker than pallidus.

The big brown-bat may be distinguished from other bats by its

large size (about 4% indies), rich, brown color, and small ears

(reaching only to nostril when laid forward). It possesses 32 teeth,

i i i • i , • 2_2 1-1 1_1 3_3

the dental tormina being: »• — .<?. — , p. — ,m.-- = 32.

3-3 1-1 2-2 3-3

Bats of the genus Eptesicus are cosmopolitan in distribution. A
single species occurs in North America, of which Engels (1936)
recognizes four races in western United States.

DISTRIBUTION MAP
STATE OF WASHINGTON

i to>*-\n* i

Fig. r>. Distribution of the big brown-bat, Eptesicus fuscus bernardinus,

in Washington.

The l)iu brown-bat appears early in the evening, often before the
-wallow- have retired. This, and its large size, make- it relatively
easy to study. It is principally a forest bat but also is found in
towns and cities. It is common in Seattle and hunts around street
lights and about the tree- in the city parks. In it- more natural
habitat it (lie- over trees .and clearings. Big brown-bats were re-
peatedly timid, with a car's speedometer, at 17 miles an hour a-
they flew down a road lined with tall tree-. This is a greater -peed

Bats 167

than that at which they usually hunt. When the speed of the car
was increased the bats dodged sideways, around the car. At lesser
speeds they pulled ahead and escaped.

Big brown-hats arc less gregarious, when hunting, than some bats.
One or two may be found in a small area, the limits of which are
definitely fixed. We noted this repeatedly near Cottage Lake, King-
County, where the big brown-bats hunted along roads through sec-
ond-growth conifers. The bats patrolled back and forth along a
section of a road about a quarter of a mile in length. When a bat
reached the end of its personal territory, it would wheel and return.
( )n six successive trips a bat turned, to retrace its course, at points
less than 50 feet distant from the point of the first turn. When
the bat approached the turning point on the seventh trip, a bat from
the adjoining strip of road approached the area. The two animals
fluttered about each other with shrill squeaks, audible 50 feet away.
The fluttering and squeaking continued for nearly a minute, after
which both bats resumed their hunting. Darkness concluded the
observations.

On several occasions we stood on a road patrolled by a big brown-
bat. The bat immediately detected the watcher and fluttered about
his head and face. Big brown-bats ignore other species of bats
hunting on their territory. Indeed, such comparatively slow flyers
could scarcely drive the more speedy bats away.

Two Eptesicus, kept in captivity for several weeks, exhibited very
different personalities. One was shy and sullen. It cowered in
the back of its cage and attempted to bite when handled. It would
eat only small quantities of finely scraped liver tendered to it on a
spoon. If a bit of liver adhered to its face it would shake violently
and refuse to eat more. When it had eaten its fill, it would seize
the spoon in its teeth and spill the remaining food. The second
specimen was tame and greedy. It ate insects, liver, earthworms,
and even pieces of flesh. After two days in captivity it learned to
come to the door of its cage at a person's approach and open its
mouth, in anticipation of food. If it dropped bits of liver on which
it was chewing it would descend from the top of its cage to retrieve
them, squeaking indignantly while doing so. It also descended to
the floor of the cage to lap water from a dish. It habitually hung
head downward from the roof of its cage, but reversed its position
when evacuating urine or feces. Engler (1943: 96) discovered that
big brown-bats will kill and eat smaller bats, at least in captivity.

Hi*

University of Kansas Publs., Mus. Nat. Hist.

Lasiurus cinereus cinereus (Beauvois)
Hoary bat

Vespertilio cinereus (misspelled linereus) Beauvois, Catal. Raisonne Mus. Peale, Philadelphia,
1796:18 (p. 15 of English edition by Peale and Beauvois).

Lasiurus cinereus H. Allen, Smithsonian Misc. Coll., 7 (no. 1C5): 21, 1864.
Atalapha cinerea True, Proc. U. S. Nat. Mus., 7:602, 1885.

Type. — None designated. Type locality Pennsylvania, probably near Phila-
delphia.

Measurements. — Ten specimens from California average: Total length 130.5;
length of tail 54.8; hind foot 10.7; ear 16; height of tragus 9.5; weight (of 3)
20.4 grams.

Distribution. — Recorded from Seattle (W.S. M.) to the north, Westport
(W.S.M.) to the west and Pullman (W.S.M.) to the east.

DISTRIBUTION MAP
STATE OF WASHINGTON

25 25 50 75 100

■ ■ ■ I I I I

MILES

C-o'-n) Undo * -

Fig. 46. Record stations for the hoary bat, Lasiurus <-nt<r,<i. in Washington.

Remarks. — The hoary bat is the largest and most distinctively
marked kind of bat in the state. Adults are usually more than 5
inches in total length. The fur is exceptionally long and soft. The

Bats 169

wing-membranes are thick and leathery. The posterior half of the
wing-membrane is black; the anterior half is pale. The interfemoral
membrane is furred. Dorsally the color of the fur is mottled
white and seal-brown, giving a silvery-gray effect. The ears are
short and thick; the feet short and wide. The dental formula is:

1-1 1-1 2-2 3-3

i. , c. ■ , p. ■ , m. = 32.

:; :: 1-1 2-2 3 3

The genus Lasiurus is found in North America, South America,
and the Hawaiian Islands. Lasiurus cinereus ranges from British
Columbia to southern South America. Osgood (1943: 53) records
two subspecies from Chile.

Information on the natural history of the hoary bat is meager.
It was observed in the Kettle River Mountains, but none was ob-
tained. It flew erratically and rapidly and did not appear until
darkness had set in. At least two were seen above a road through
a forest of western larch and ponderosa pine. Probably it breeds in
the mountains of northeastern Washington and in the northern
Cascades.

The hoary bat is migratory, leaving the state in August and Sep-
tember, rarely lingering until early October. It winters along the
coast of central and southern California (Dalquest, 1943: 23).

Antrozous pa Nidus cantwelli Bailey
Pallid bat

Antrozous pallidas cantwelli Bailey, N. Amer. Fauna, 55:391, August 29, 1930.

Type. — Obtained at Rogersburg, Asotin County, Washington, by G. G. Cant-
well on May 28, 1918; type in United States National Museum.

Measurements. — A male from the south bank of the Columbia River, op-
posite Fallbridge, Klickitat County, measures: total length 113; length of tail
40; hind foot 14; ear 36; height of tragus 13; weight 19.8 grams.

Distribution. — Recorded only from localities near the Columbia and Snake
rivers in eastern Washington; from Wenatchee (W.W.D.) south and east to
Bly and Rogersburg (Taylor and Shaw, 1929: 9).

Description. — Antrozous pallidus is more likely to be confused
with Corynorhinus than with any other kind of bat and can
be distinguished from Corynorhinus by its larger size (4% to 5
inches), lighter color, thick and leathery membranes, and shorter,
wider ears. Antrozous lacks the prominent "lumps" on the ros-
trum, characteristic of the long-eared bat. The dental formula is:

1-1 1-1 1-1 3-3

i. , c. , p. , m. = 28.

2-2 1-1 2-2 3-3

170

University of Kansas Publs.. Mrs. Nat. Hist.

The genus Antrozous is confined to western United States and
northwestern Mexico. Two species arc recognized by Miller il!»24:
841. One of these (pallidus) includes three geographic races. A
single race is known to occur in Washington.

The pallid bat chooses a crevice in a cliff, cave, or building a- its
daytime hiding place. Like Corynorhinus, it is colonial. The basalt
cliffs of eastern Washington offer such ideal hiding places that pallid

DISTRIBUTION MAP
STATE OF WASH 1 NGTCN

Fig. 47. Distribution of the pallid hat. Antrozous i><illi<hi* cantwelli,

in Washington.

bats are seldom seen and consequently little is known of their habit-,
hi the southwestern Tinted States, pallid bats are more abundant
and better known. II. W. Grinnell (1918: :}.">."> i notes that the floor
of a roost of pallid bats was strewn each morning with the head-.
wings, and legs of insects. Most numerous were the remains of the
Jerusalem crickel {Stenopelmatus) , a flightless insect that the bats
must have secured on the ground. Engler (1943: 96) found that in
captivity, pallid bats would kill and eat smaller bats and lizards
confined with them.

Bailey (1936: 392 I believes that the young of this bat are bcin
in late June and early July. One or two young constitute a litter.

Bears

171

Ursus americanus Pallas

Black bear

Description. — The black bear is the largest carnivore found in
Washington if the grizzly is extinct there. The exact size of the
adult male black bear is somewhat in question. Few actual weights
are on record of Washington bears. Grinnell, Dixon and Linsdale
i1!»37: 101) paid special attention to the maximum weight of black
bears in California and concluded that few ever exceeded 500 pounds.
The total length of the largest black bear among specimens from
the entire Pacific Coastal area, especially from California and
Alaska, they give as 5 feet. 10 inches (op. tit., p. 102). It may
safely be assumed that few individuals ever reach a length of six
feet. Females are smaller; recorded weights of adults rarely ex-
ceed 350 pounds. The largest males have hind feet from 10 to lO 1 /^
inches in length.

Fig. 48. Female black bear (Ursus americanus altijrontalis) , and two cubs.
near Big Four Inn. Mt. Baker National Forest, Washington, July. 1939. (For-
i si Service photo by R. L. Fromme.)

The black bear, including its closely related species, the Mexican
bear. Ursus machetes, ranges over Alaska, Canada, the United States
and northern Mexico. The distinction between the black and grizzly
bears has been listed under the account of Ursus chelan.

The fur of black bears in the Cascade Mountains in the fall is

172 University of Kansas Publs., Mrs. Nat. Hist.

long, sleek and glossy but rather stiff. In the spring and early
slimmer the fur of animals at Mount Rainier often appeared coarse,
wooly and patchy or rubbed on older animals. Some smaller be;ir>
possessed sleek, well-groomed pelts. Bears from the lowland areas
are said to possess poor pelts because of constant abrasion in the
forests. In the fall, when the salmon are spawning, the bears are
said by trappers to roll in decayed fish until they ''smell so bad
you can't come near them" and their fur is matted and "crawling
with lice."

It is now known that the cinnamon bear, sometimes called brown
bear, is merely a color phase of the black bear. The brown phase
varies in color from a rich, dark reddish brown to a pale cinnamon
brown. Cowan (1938: 204) has recorded the geographic variation
in the brown and black coats of the bears of British Columbia and,
in part, of Washington. On the Olympic Peninsula the brown phase
is rare. Old residents have told me of seeing but a few brown bear-
in their life. I have seen only black bears in the lowlands of west-
ern Washington. Cow T an (loc. ait.) lists 1,197 black and 79 brown
bears from Fort Nisqually, Pierce County, between 18.34 and 1852.
In the Cascade Mountains the brown phase is not uncommon, per-
haps one out of five bears seen being brown. In northeastern
Washington the brown and black phases are about equal in number,
and some trappers state that the browns arc more numerous. Cowan
lists 3,813 black and 2,871 brown bears from Fort Colville between
1826 and 1856.

The black bear occurs in a variety of habitats in Washington.
It seems to be absent only from the treeless areas of eastern Wash-
ington, and is most abundant in the Cascade Mountains and Olym-
pic Mountains where food is abundant and men are few. It is not
uncommon throughout the timbered lowlands of northeastern Wash-
ington and western Washington, however, and shows a surprising
ability to exist unnoticed near the larger cities. In the dense,
junglelike forests of southwestern Washington il is numerous.
Jackson (1944: 1) estimates that 13,679 black bears live in Wash-
ington, more than in any other slate.

The habits of the black bear have changed greatly where it has
come into contact with man. At Mount Rainier National Park
black bears now ignore people, save to beg for food. In the Cas-
cades the bears are wild hut so seldom see men that in many places
they live much as they did hundreds of years ago. In the lowlands
of western Washington they are in daily contact with evidence of

Beaks

173

man or his guns. As a result they are silent and shy, rarely being
seen.

The black bear in the mountains is active sometimes by day and
sometimes by night. Probably it is principally nocturnal but ac-
tive by day only when the food obtained at night is insufficient to
meet the needs of the animal. In the lowlands it is almost com-

Fig. 49. Black bear (Ursus americanus altifron talis), in "hibernation," Mt.
Baker region, Washington, about 1936. (John E. Candle photo, courtesy
"Field and Stream.")

pletely nocturnal. In the mountains it enters upon its winter sleep
with the first snows or, if the snows are late, when the huckle-
berries are gone. Ordinarily black bears are in "hibernation" by the
middle of November. In the lowlands of western Washington they
are active until after the salmon spawning season, and probably do
not ''hibernate" before the middle of December and may not do so
at all. In the mountains they emerge from the winter sleep in May ;
in the lowlands they emerge in February or March.

U-t I'xiversity of Kansas Publs., Mrs. Nat. Hist.

The black hear is omnivorous in the truest sense of the word.
Because the animal is of large size it requires much nourishment.
This is obtained by eating large quantities of material with low
food value. Material eaten passes rapidly through the digestive
tract and often only easily digested parts seem to be utilized. Feces
commonly contain complete and undigested berries and seeds or
almost undigested piece- of apples or other fruit.

The principal food of the black bear in Washington is berries.
Many species are eaten, but the huckleberry {Vaccinivm sp.) is
favored. Other food items are: Gaultheria shallon (pulpy fruit);
Oregon grape, Bcrberis nervosa i flowers and fruit); salmonberry,
Rubvs parviflorus (leaves and fruit) ; leaves of several plants, in-
cluding Rubus macropt talus and other thorny kinds; grass, succulent
plants, and roots. Insects are eaten and most feces -how remains of
a few. In June, 1938, near Cle Elum, Kittitas County, a species
of sword-tailed cricket fully two inches in length was abundant,
and droppings of a bear there were composed entirely of the remains
of these crickets. Fish, especially spawning salmon, are important
food. When salmon arc spawning the bear- in the vicinity feed on
nothing else. Warm-blooded vertebrate- probably are eaten but
none of the many bear droppings examined by me contained remains
of birds or mammals. Bennett, English and Watts (1943: 30) found
ivw mammals to be eaten by bears studied by them in Pennsylvania.
Locally bears kill pigs and sheep, smash bee-hives and raid fruit
trees.

The huge size and sometimes awkward appearance of black bears
combine to give a comic note to some of their natural actions. A
bear in full flight, lumbering along a trail with head swinging from
side to side and hind feet stretching past the forelegs at each stride,
is more apt than not to amuse the observer. Near Stevens Pass, a
large black bear was observed lying on its back in a mud wallow.
This was ;ni oval opening in the ground in a meadow of heather and
huckleberry near a small stream. The wallow was approximately
f) feet long, •'! feel deep and 4 feet wide. The bear was immersed in
muddy water with only its head and feet visible. It seemed well
content and comfortable, shifting its bulk occasionally and waving
its paw-. When a stray breeze brought to the bear the scent of my
companion and myself the animal'- contentment vanished and it
hastily made a frantic effort to depart. It had wedged itself deeply
in the hole, and a- it twisted and tinned in attempting to sit tip,
waves of water gushed from the wallow. Eventually escaping

Bears

175

from it- wallow, the bear half ran and half rolled to the cover of
alders 100 feet away. This particular wallow had been in use by-
bears for some time, as was attested by old droppings at its edge
and by its well-worn condition.

The fact that Washington has the largest population of black
bears of the states in the union is due, largely, I feel, to abundant
natural cover, food, such as salmon and huckleberries, and an in-
telligent game code. In California and some other states the black
bear has been considered a fur bearer, to be trapped with steel traps.
This is permitted in spite of the fact that their pelts bring but a few

DISTRIBUTION MAP
STATE OF WASHINGTON

MILES

of WP* P.o,, c . 60'9-5797

Fig. 50. Distribution of the black bear in Washington. A. Ursus americanus
altijrontalis. B. Ursus americanus cinnamomum.

dollars, usually less than ten. Their sale scarcely repays the trapper
for his labor in preparing the hide. Townsend (1887: 182) remarked
mi the ease with which bears could be trapped in California although
in the same area they were so shy that they could seldom be shot.
As a result of trapping, the number of bears in some states has been
dangerously reduced. In Washington the bear is a game animal,
to be hunted with a rifle for a few weeks in the fall when the pelt
and flesh are at their best. As a result bears are numerous and can
be hunted with fair chances of success.

]7() University of Kansas Publs., Mis. Nat. Hist.

Ursus americanus altifrontalis Elliot

Ursus altifrontalis Elliot, Field Column. Mus. Publ. 80, zool. ser. 3:234, June, 1903.
Euarctos altifrontalis Miller, U. S. Nat. Mus. Bull., 128:90, April 29, 1924.
Ursus americanus americanus Taylor and Shaw, Mammals and Birds of Mount Rainier Na-
tional Park, U. S. Nat. Park Service, Washington, p. 37, 1927.

Euarctos americanus altifrontalis Bailey, N. Amer. Fauna, 55:321, August 29, 1936.
Ursus americanus altifrontalis Hall, Univ. California Publ. Zool., 30:232, March 2. 1928.

Type. — Obtained at Lake Crescent, Olympic Peninsula, Clallam County,
Washington, by D. G. Elliot in 1898; type in Field Museum of Natural
History.

Racial characters. — Color dark, almost always black instead of brown; skull
wide, high and heavy; molar teeth wide and heavy.

Distribution. — From the eastern base of the Cascade Mountains westward
to the Pacific. Marginal occurrences are Chelan (W.W.D.) and Signal Peak
(Taylor and Shaw, 1929: 10).

Ursus americanus cinnamomum Audubon and Bachman

Ursus americanus car. cinnami'inuin Audubon and Bachman, Quadrupeds of North America,

3:125, 1S54.

Euarctos cinnamomum Miller, U. S. Nat, Mus. Bull., 128:91, April 29, 1924.
Euarctos americanus cinnamomum Bailey, N. Amer. Fauna, 55:319, August 29, 1936.
Ursus americanus cinnamomum Hall, Univ. California Publ. Zool., 30:232, March 2, 1928

Type. — Obtained near the mouth of Jim Ford Creek, Lower Clearwater
River, western Idaho (Bailey. 1936: 319) by Lewis and Clark on May 31, 1806.

Racial characters. — Resembling altifrontalis but skull and molar teeth nar-
rower; color even in black phase paler and browner; brown and black phases of
approximately equal incidence.

Distribution. — Northeastern Washington and the Blue Mountains of south-
eastern Washington, occurring westward as far as Republic (Taylor and Shaw.
1929: 10).

Ursus chelan Merriam

( irizzly bear

Ursus chelan Merriam, Proc. Biol. Soc. Washington, 29:136, September 6, 1916.

Type. — Obtained in Township 30 N, Range 16 E. Willamette Meridian.
Wenalchee National Forest. Chelan County. Washington, by D. S. Rice on
September 1, 1913; type in United States National Museum.

Measurements. The skull ol the type, an old male, measures basal length
314; occipito-nasal length 323; palatal length 170; zygomatic breadth 225;
interorbital breadth 86.

Distribution. — Probably once from the Stevens Pass ana north to British
Columbia. Possibly still remaining in the mountains between Mount Baker
and Lake Chelan.

Remarks. — Grizzly bears occurred in the lowlands of Oregon (Bailey.
1936: 324) and California (C.rinnell, Dixon and Linsdale. 1937: 67) but there
i- no record of their ever having occurred in the lowlands of western Wash-
ington.

Bears of the genus Ursus range over Europe, Asia and North
America. No less than 75 kinds of grizzly and big brown bears are

Bears

177

recognized for North America by Merriam (1918). They are closely
related to the brown bears of Asia. The 13 kinds of black bears
of North America are now thought to belong to but one species,
namely americanus, and have no close relatives in Eurasia.

The grizzly is the largest carnivorous mammal in North America.
From the black bear the grizzly may be distinguished in life by the
prominent muscular hump on the shoulders. Another feature is the
long. slim, gently curved claws of the forefoot as compared with
the shorter, stouter and sharply curved or hooked foreclaws of the
black bear. The grizzly is much larger than the black bear. Cra-

DISTRIBUTION MAP
STATE OF WASH I NGTON

MtLES

c f * PA P.r

6C-!-S737 b. CMT I9J7

Fig. 51. Probable past distribution of grizzly bears in Washington. A. Ursus
chelan. B. Ursus canadensis. C. Ursus idahoensis.

nially, the grizzly differs in possessing a larger skull with lower
frontal area, higher, more extensive sagittal crest, more posteriorly
jutting occipital condyles and much longer tooth row. The color
of both species is variable. Some grizzly bears from Alaska and
British Columbia are dark brown, almost as black as a black bear.
Others are pale cinnamon with the longer guard hairs pale gray.
Individuals of this frosted appearance popularly are known as "sil-
ver tips." In Washington, the black bear varies in color from glossy
black, through various shades of brown to a pale cinnamon.

12—4993

178 University of Kansas Publs., Mrs. Nat. Hist.

The grizzly bear is extinct over most of Washington. A few may
remain in remote parts of the northern Cascades, and are regularly
listed on the game census reports of the Mount Baker National
Forest. Nevertheless, I have found no one who has seen an un-
doubted grizzly bear in Washington. The type specimen was ob-
tained well up in the Cascade Mountains where the animals, like
those in British Columbia, teed on roots, berries, marmots, pikas
and other vegetable and animal life.

Raccoon 17!>

Procyon lotor i Linnaeus)

Raccoon

Description. — The raccoon is a relatively large carnivore with a
stout, heavy body. Large adults measure three feet in length and
weigh up to 20 pounds. The legs are of moderate length. The
broad head tapers sharply to a pointed muzzle. The ears are erect,
broad, low and rounded; the tail is long, about one-half the length
of the head and body, bushy and round; the hind feet are large,
flat and naked-soled but the forefeet are smaller with long, slim,
handlike toes. The color of the body is a grizzled gray heavily
washed with black on the back and sides. A jet-black "mask"
across the eyes, sharply outlined by white muzzle and forehead, is
the most distinctive feature of the head. The tail is alternately
marked with six or seven black and five or six yellowish gray bands.
The long, loose guard hairs give the body a shaggy appearance. The
fur of the wrists is short, smooth, coarse, and directed downward.

The raccoon is not an aquatic mammal, yet it is closely asso-
ciated with water, whether it be streams, rivers, lakes or the ocean.
The favored habitat of the raccoon is the shore line. It is a wader
and, if it can be considered as specialized for any occupation, it is
wading in mud and shallow water. Its long toes, naked feet, and
short-haired wrists are admirable adapted for walking on muddy
bottoms or in shallow water. The coon is also at home on land. It
moves swiftly and silently and when chased by dogs can cover miles
in a few hours. It is an agile climber and lives in dens well up in
trees.

The raccoon is almost completely nocturnal. Individuals are oc-
casionally seen in the morning or evening, especially, when the tide
is low along the ocean beach or Puget Sound. Near Fall City, King-
County, a small raccoon was seen eating a crayfish at 3:30 p. m. on
a warm, sunny June day. In the eastern United States raccoons
sleep during much of the winter, and probably they do the same
in eastern Washington. In western "Washington they are active most
of the winter. Some trappers stated that the animals "hole up" in
spells of unusually cold weather. Along the Tolt River, 10 miles
southeast of Duvall, King County, their tracks were seen daily in
January, 1936. although the temperature fell well below freezing
each night.

Washington is near the northern limit of the range of the raccoon.
The animal is reasonably common in western Washington, and

180 University of Kansas Pvbls., Mis. Nat. Hist.

ranges well up into the Cascade Mountains. The highest altitudinal

record available is Longmire, Mount Rainier National Park (Tay-
lor and Shaw, 1927). Raccoons are not uncommon in southeastern
Washington and in the Yakima Valley. They follow the Columbia
River northward, as shown by tracks at Wenatchee on two occa-
sions. The river valleys that flow into the Columbia in northeastern
Washington seem admirably adapted for raccoons but the animal
is rare there. Trappers who have lived and trapped in northeastern
Washington for many years tell of seeing tracks at intervals of
years along the Okanogan, San Poil, Colville and Kettle rivers.
Several stated that tra< ks had been seen more often in recent years.
Northeastern Washington seems to be the peripheral range of the
species, occupied at rare intervals by animals wandering north from
the Columbia River.

The food habits of raccoons are almost as varied as those of black
bears. Animal matter forms their diet over most of the year; along
stream courses crayfish, fish, thin-shelled fresh-water mussels, frogs
{Hyla and Rama), and aquatic insects are eaten. Along beaches
fish brought in by the tide, crustaceans, and mussels {Mytilus
edalis) are taken; small sharks appear not to be eaten. A recently
dead dogfish that lay on the beach at W T hidby Island, Island County.
was ringed by the tracks of a raccoon but had not been eaten. The
shore crabs (Hemigrapsus nudus and Hemigrapsus orcgonensis) arc
a favored food and regular items of diet. The edible crab (Cancer
productus) is also eaten and in the San Juan Islands the porcelain
crab (Petrolisthes criomerus) was commonly eaten. Small mammals
and birds are eaten regularly by this race of raccoon in California
i< Irinnell, Dixon and Linsdale, 1937: 157), and the eggs and young
of wild birds are sought in the spring lop. cit.: 158). The ability
of the raccoon as a climber makes it a particular menace to nesting
birds. Berries, including domestic blackberries and salmonberries,
are eaten in quantity when available. Apples are dearly loved and
the trees and orchards of abandoned ranches are regularly visited.
The fondness of the raccoon for green corn is well known. Insect-
are present in small quantities in most droppings and in the late
summer some feces were composed entirely of the remains of grass-
hoppers. Chickens, ducks, young turkeys and eggs are stolen and
individuals become extremely adept at raiding hen-houses.

At one time, about 1920, the raccoon became scarce in western
Washington as a result of heavy trapping and high price of the pelts.
A closed season was strictly enforced until it again became common.

Raccoon

181

The price received by the trapper for raw furs of raccoon has since
been rather low, from two to ten dollars. With fur prices in this
range, the number of raccoons probably will not be reduced to a
dangerously low level by trapping, but instead may be expected
to furnish a regular winter income to the trappers who do trap for it.
Raccoons are known to breed in their first vear of life (Pope, 1944:
91).

Procyon lotor psora Gray

Procyon psora Gray. Ann. and Mag. Nat. Hist., 10:261, December, 1842.

Procyon psora pacifica Merriam, N. Amer. Fauna, 16:107, October 28, 1899 (type from
Lake Keechelus, Kittitas County, Washington).

Procyon proteus Brass, Aus dem Reiche der Pelze, p. 564, 1911.

Procyon lotor pacifica Taylor and Shaw, Occ. Pap. Chas. R. Conner Mus., no. 2:10. De-
cember, 1929.

Porcyon lotor psora Grinnell, Dixon, and Linsdale, Fur-bearing Marani. California, Univ.
California Press, Berkeley, p. 137, July 22, 1937.

Type locality. — Sacramento, Sacramento County, California.

Racial characters. — Medium size; dark color; smoothly rounded skull.

Measurements.— A female from Forks, Clallam County, measures: total
length 905; length of tail 355; hind foot 125; ear 50; weight 14V 2 lbs. A
female and 6 males, young animals of the year, taken between November
15 and December 15, average: 772; 284; 114; weight (of 3) 8 % lbs.

DISTRIBUTION MAP
STATE OF WASH I NGTON

MILES
of *PA P.<j#E> <079-S797 b, CflT 1,37

Fig. 52. Distribution of the raccoon in Washington. A. Procyon lotor psora.

B. Procyon lotor excelsus.

182 University of Kansas Pubis., Mis. Nat. Hist.

Distribution. — From the western dope of the Cascade Mountains westward.
Marginal occurrences are Lake Keechelus (type locality) and Mount Rainier
(Taylor and Shaw. 1927: 45).

h'i minks. — The range of variation in color and cranial characters of coastal
raccoons is large. I am unable to find any character or average difference to
separate tin raccoons of western Washington from those of northern California.

Procyon lotor excelsus Nelson and Goldman
Procyon lotor excelsus Nelson and Goldman, Jour. Mamm., 11 : 158, November 11, 1930.

Typt . — Obtained on upper Owyhee River, near the mouth of the
North Fork in southeastern Oregon by J. W. Fisk on April 15, 1920; type
in United State.- National Museum.

Racial characters. — Similar to psora but larger; body paler and grayer;
skull larger, heavier and more angular.

Distribution. — Southeastern Washington, the valleys of the Snake and
Yakima livers and the valley of the Columbia River north to northeastern
Washington. Marginal records, from reports of trappers, include the Okanogan
River. Okanogan County, Wenatchee. Chelan County, and Selah. Yakima
County.

Rf marks. — The assignment of the raccoons of northeastern Wadiington to
excelsus is tentative for I have seen no specimens.

Marten 183

Martes caurina Merriam

Western marten

Description. — The marten is slightly smaller and slimmer than a
house cat, and at first glance resembles a large squirrel. The legs
are longer, the body stouter and the fur more fluffy than those of
a mink or weasel. Adult males weigh from two to two and a half
pounds, and females from one and a half to two pounds. Males
are slightly more than two feet in total length and females about
18 inches, the tail comprising one-third or more of the total length.
The head is broad and narrows rapidly to a sharp muzzle. The
ears are large, erect and prominent. The feet are large with stout
toes and long, sharply curved pinkish-white claws. The body and
head are rich golden brown, the tail, wrists, feet and muzzle being
darker.

The western marten ranges from British Columbia southward
through Idaho and Washington to California. A closely related
species, americana, is found in Alaska, the eastern United States
and Canada. Martens and fishers may be distinguished from weasels
and minks by the presence of 18 rather than 16 teeth in the upper
jaw and 20 instead of 18 teeth in the lower jaw.

The western marten is arboreal. Its principal habitat in Wash-
ington is the Canadian Life-zone forests of the Olympic, Cascade
and Blue mountains and the various ranges in the northeastern part
of the state. At one time it ranged near sea level along the densely
forested coastal belt and may still do so in the more rugged parts
of the Willapa Hills.

The marten is both diurnal and nocturnal. In Mount Rainier
National Park the species has become quite tame and may be seen
in the daytime. Many of the small mammals upon which it feeds
are diurnal, but others are nocturnal. The marten is active through-
out the year. Trappers report that during a storm the marten
"holes up in rock slides" where it lives on conies and chipmunks
until the storm passes. It spends a large part of its time in trees,
and travels through them for long distances. It climbs more skill-
fully than the tree squirrels upon which it feeds. On the ground
or on snow the marten travels in bounds, a yard at a leap, and
its characteristic bounding gait forms tracks that are distinctive
and easily followed.

Mammals of the weasel family mostly are not gregarious but the
marten is exceptional in that in the winter it travels in bands of 6

184 University of Kansas Publs., Mis. Nat. Hist.

to 10 animal:-. Individuals composing these bands are inclined to
wander but nevertheless the whole band travels in a definite general
direction at a good rate of speed. Travel-ways or "runs" may be
used by more than one band, and a run may extend for many miles,
perhaps for as many as 50. A band of martens may take two weeks
to complete the circuit, but usually returns to the starting point
in less than a week. Most runs are about "half-way up the moun-
tain," or mid-way between the crest of the hill or timber-line and
the floor of the stream valley below. In summer the marten ranges
higher; it lives in the trees just below timber-line and in the talus
slides near timber-line. When the snows are unusually late the
martens may keep to these higher areas until November.

The food of the marten consists principally of small mammals
and probably birds; the staple food in winter is the Douglas squirrel.
In summer they feed on pikas, mantled ground squirrels and chip-
munks. Mice, also, are eaten. The deer mouse, Peromyscus mani-
culatus, is usually abundant about old cabins and is successfully
used as bait by trappers. Wood rats and flying squirrels are also
eaten, the latter being especially important in certain areas. The
tracks of martens that had been following snowshoe rabbits were
seen on several occasions but the martens had turned off before a
kill was made. In every instance the rabbit tracks indicated that
the animals were hopping leisurely and browsing; apparently the
tracks were made some time before the marten began to follow them.

Martens are inquisitive, and to judge from their tracks in snow
they investigate almost every object that they pass; a fallen mound
of snow, branch, bit of moss, log or isolated tree is apt to be visited.
Failing to find a meal in or around one of these objects, the marten
visits the next object that catches its eye. Seemingly the animal
always is giving concentrated attention to some definite object al-
though the attention can shift in an instant when a more interesting
object, comes in to view. As a result the trail of a marten in the
snow is an intricate affair composed of numerous straight lines and
sharp turns.

The trapping of martens is specialized work, engaged in by pro-
fessional dappers that follow trap lines many miles in length.
Trappers commonly have a base cabin and one or two shelters
situated a day's march apart, The trap-line is set in a circle, re-
quiring one to three days' travel by the trapper to complete the
circuit.

The standard "set" for marten in the Cascades consists of an

Marten

l s;>

opening 8 to 10 inches wide, 4 inches high and 6 inches deep chopped
into the side of a dead stub. The hole is cut as high as the trapper
can reach. A trap is set in the opening and bait is placed in the
back of the hole. A sapling 1 to 3 inches in diameter inclined at
an angle of 45° from the ground to the hole completes the set. As
the snow deepens, new holes are chopped higher up on the stub.
Such sets seen in the summer may consist of six or seven holes spaced
a foot apart. The chips from the holes are left lying on the snow
and are said to attract the animals. Bait commonly consists of
flying squirrel, red squirrel or deer mouse.

DISTRIBUTION MAP
STATE OF WASHINGTON

MILES
der Ampicai of WM P--

Fig. 53. Distribution of the western marten in Washington. A. Martes
caurina caurina. B. Martes caurina origenes.

The value of the marten's pelt fluctuates from year to year. In
recent years average pelts have sold at from ten to twenty dollars
each. The number of individuals taken by a trapper varies with
the trapper's skill and energy and the location of the trap line. The
largest catch made bj r one trapper in a single winter, of which I
know, was 300 animals, taken near Mt. Adams.

186 University of Kansas Publs., Mtjs. Nat. Hist.

Martes caurina caurina (Merriam)

Mustela caurina Merriam, N. Amer. Fauna, 4:27. October 8, 1890.

Martes caurina caurina Miller, U. S. Nat. Mus. Bull., 79:93, December 31, 1912.

Type— Obtained near Grays Harbor, Grays Harbor County. Washington,
by L. C. Toey on February 4. 1886; type in United States National Museum.

Racial characters. — Color dark; throat patch bright orange or brownish.

Distribution. — From tin Cascade Mountains westward. Marginal localities
are (from Taylor and Shaw. 1929: 10) head of Cascade River, Riverside. Chelan.
Easton and Trout Lake. The record from Chelan seems doubtful, and the
specimen may have come from somewhere around Lake Chelan, rather than
from the town of that name.

Ix'i marks. — Skulls of martens from many localities in western North
America were studied in an effort to determine the relationship of the eastern
pine marten (Maries americana) and the western marten. East of the Rocky
Mountains the ranges of the two species approach closely, but each retains
distinctive characters, notably the shape of the auditory bullae. The two
species have not been taken together and the possibility of intergradation
exists. On the basis of the evidence at hand, the two should be regarded
as full species until positive proof of intergradation is established.

In the absence of sufficient material, the marten of the Cascades i> referred
to Martes caurina caurina. Fur graders distinguish between a dark "coast
marten" and a paler "Cascade marten."

Martes caurina origenes (Rhoads)

Mustela caurina origenes Rhoads, Proc. Acad. at. Sci. Philadelphia, 1902:458, September
30, 1902.

Martes caurina origenes Mill.-,. U. S. Nat. Mus. Bull., 79:93, December 31, 1912.

Type. — Obtained at Marvine Lodge, Garfield County. Colorado, by E. T.
Seton on September 16, 1901; type in Academy of Natural Sciences of Phila-
delphia.

Racial characters. — Paler than caurina with grayer head and yellow or white
rather than deep orange or brown throat patch.

Distribution . — Mountainous areas of northeastern Washington and the Blue
Mountains of southeastern Washington. Trappers have reported this marten
from the mountains near Republic, Ferry County.

Fisher

187

Martes pennanti (Erxleben)

Fisher

[Mustela] pennanti Erxleben, Syst. Regni. Anim., 1:470, 1777.

Martes pennanti pennanti Miller, U. S. Nat. Mus. Bull., 79:94, December 31, 1912.

Mustela canadensis pacifica Rhoads, Trans. Amer. Philos. Soc, n. s., 19:435, September,
1898 (type from Lake Keechelus, Kittitas County, Washington).

Martes pennanti pacifica Miller. 1*. S. Nat. Mus. Bull., 79:94, December 31, 1912.
Martes pennanti Grinnell, Dixon and Linsdale, Fur-bearing Mamm. California, Univ. Cali-
fornia Press, Berkeley, p. 211, July 22, 1937.

Type locality. — Eastern Canada.

Distribution. — Originally forested areas from the eastern base of the Cas-
cade- west \v;in I and possibly the Blue Mountains of southeastern Washington
and the mountains of northeastern Washington; now probably confined to the
Cascade and Olympic mountains.

Remark*. — The fisher is the size of a large cat. In general pro-
portions it resembles the marten. Adult males measure about 31/2
feet in length; the tail comprises 16 inches of this. Adult females

*. *"■

Fig. 54. Fisher (Martes pennanti). 2-year-old male in captivity; New West-
minster, B. C, March 7. 1939. (Fish and Wildlife Service photo by Victor B.
Scheffer. No. 598.)

are slightly less than 3 feet in length of which the tail makes up
approximately 15 inches. Males weigh up to 10 lbs. and females
about 5y 2 pounds (Grinnell, Dixon and Linsdale, 1937: 213). The
fisher has a slim body, bushy tail, short legs, large feet, and wide,
low and triangular head. The ears are low, wide, rounded and erect.
The fur is an ashy, brownish gray in color with an overwash of

188

University of Kansas Publs., Mus. Nat. Hist.

blackish caused by long, dark hairs. The head is slightly paler than
the body. The feet, rump and tail are darkest. The claws are
strong and sharply curved.

The fisher is found in wooded parts of North America, extending
southward in the Cascade-Sierra Nevada Chain to central Cali-
fornia. Its near relatives are the martens. It is active all year.
Like the marten, it is active by day and probably also by night, In
spite of absolute protection for several years, the fisher is rare in

DISTRIBUTION MAP
STATE OF WASHINGTON

i MM-179T b, CJ1T l»S7

Fig. 55. Distribution of the fisher, Maries pennanti, in Washington.

Washington, and seems never to have been common. In consequence
n latively little is known of its habits, and the little that is known
lias been reported by fur-trappers.

The fisher seems not to live in bands as does the marten. Most of
the actual records of fishers trapped arc at higher altitudes but are
misleading because most trappers agree that the fisher occupied a
lower zone, altitudinally, than does the marten. There arc old
records of its occurrence near sea level (Seheffer, 1!>38: 9). The
animals arc usually taken in marten sets or in trap- sei especially for
fishers by trappers who find their tracks on their marten trap lines.

Fisher 189

Since marten trappers are almost the only persons who travel in
the mountains in winter, and since they operate mostly above the
areas where fishers live, relatively few fishers are reported.

Fishers are said to feed on chipmunks, squirrels, mice, birds and
other small, warm-blooded animals, and to climb trees and catch
squirrels in their natural habitat. Also, fishers are said to catch and
kill martens. Their tracks in the snow resemble the marten's in that
the hind feet land in the same places as the forefeet ; both animals
bound rather than walk.

The pelt of the fisher commands a high price. The smaller sized,
females, are the more valuable. The price paid for pelts fluctuates
widely and has ranged from as low as twelve dollars to as high as
one hundred dollars in recent years.

In Washington more fishers live in the Olympic Peninsula and the
northern Cascade Mountains than elsewhere. A few may occur in
northeastern Washington, the Blue Mountains and the Willapa Hills.

The name pacifica was regarded by Grinnell, Dixon, and Linsdale
(1937: 217) as a synonym of pennanti.

190 University of Kansas Publs., Mis. Nat. Hist.

Mustela erminea Linnaeus

Ermine

Description. — Though it is similar in general characters and pro-
portions to the long-tailed weasel, the ermine is much smaller and
has a relatively shorter tail. It is darker and less reddish or yel-
lowish in summer. Adult males measure about 10 inches in length,
of which 3V2 inches is the length of the tail. Females measure about
8 inches and have tails 2 inches long. The upper parts are choco-
late brown; the underparts arc white or pale yellow. Along the
coast of Washington, the pale color of the underparts is more re-
stricted in the ermine than in the long-tailed weasel. The dark
brown tail has a black tip. Ermines east of the summit of the Cas-
cade- become pure white in winter, save for the black tail tip. Wesl
of the summit of the Cascades the winter pelt is similar to the sum-
mer pelt but is slightly paler with denser underfur.

Ermines in America range from the Arctic southward, in moun-
tainous areas, to the southern end of the Sierra Nevada in Cali-
fornia, and in the Rocky Mountains to northern New Mexico. In
Washington they are found over the entire state except the arid
parts of eastern Washington, where only the long-tailed weasel oc-
curs. So far as my observations go, the ermine is principally noc-
turnal in Washington; I have seen only one abroad in the daytime.
Tt dashed from a roadside thicket near Glacier. Whatcom County,
and was crushed beneath the wheels of a car. Taylor and Shaw
(1927: 53) note several instances of diurnal activity of the ermines
at Mount Rainier National Park.

The ermine seems to feed principally upon mice. Its small size
adapts it to entering burrows that larger weasels cannot enter. It
probably eats chipmunks, birds, and other small, warm-blooded
animals. Ermines climb readily and are often taken in imp- set in
trees for martens. Near Skykomish, King County. William Hoff-
man took two ermines in traps set in burrows of mountain beavers.
The mountain beavers were needed for use as bait on his trap line.
The traps were resel and later caught mountain beavers. Seemingly
the ermines were traveling through the burrows, perhaps to catch
the mice which utilize the burrows as highways, rather than to prey
upon mountain beavers. It is difficull to see how the tiny ermine
could kill an adult mountain beaver, which outweighs it many
times. Should the incisors of the mountain beaver close even once
upon an ermine i! would most certainly he seriously wounded or
killed. Judging by the foresl habitai occupied by the ermine in

Ermine 191

Washington, one might expect its principal food to consist of deer
mice (Peromyscus mardculat-as) , red-backed mice {Clethrionomys)
and meadow mice (Microtus).

In observing the activities and habits of mammals in their natural
habitat, I have often relied on tracking in fresh snow. Strangely
enough, tracks of ermines were seldom found, and the few that were
seen came from beneath a log, bush or wind-fall and disappeared
beneath similar cover, rarely extending 20 feet on the surface of the
snow. The larger, long-tailed weasels often traveled for miles on
the surface of the snow. Possibly the ermines w r ere following the
burrows of mice through the snow, or perhaps they kept beneath
the surface from fear of owls. This suggests a reason why ermines
are so seldom seen abroad. They may be following burrows and
runways of mice and seldom come to the surface of the ground.

On November 18, 1936, we saw three ermines and two long-tailed
weasels taken by a trapper on Deception Creek near Stevens Pass,
King County. All were in the white winter coat. In December,
19.38. we obtained two ermines at Skykomish, King County, 18
miles west of Stevens Pass. These were in the brown winter coat.
In that area the break between the brown and white winter coat
seems to come just west of the main Cascade Summit, or at the
same point that the break occurs between the brown and white
winter coats of the long-tailed weasel.

Ermine skins have little value and usually bring from 10 to 35
cents. They are saved incidentally by trappers, for it takes but a
few moments to skin and prepare them. They possess the strong,
musk odor so typical of the long-tailed weasel.

Mustela erminea invicta Hall
Mustela erminea invicta Hall. .Tour. Mamrn., 26:75, February 27, 1945.

Type. — Obtained at Benewah, Benewah County, Idaho, by W. T. Shaw on
October 24. 1926; type in Museum of Vertebrate Zoology.

Racial characters. — Largest of the Washington ermines; winter coat entirely
white; upper lips white; black of tail more than half length of tail vertebrae.

Measurements. — Ten males and 5 females from central Idaho average,
respectively (Hall, 1945): total length 291, 255; length of tail 86. 71; hind foot
40. 32.3.

Distribution.— Northeastern Washington and the northeastern Cascades,
west to Chilliwack River and Hannegan Pass (Hall, 1945: 78).

Mustela erminea fallenda Hall

Mustela erminea fallenda Hall. Jour. Mamm., 26:79, February 27, 1945.

Type.— Obtained at Huntingdon. British Columbia, by C. H. Young on
May 21. 1927; type in National Museum of Canada.

192

University of Kansas Plbls., Mis. Nat. Hist.

Racial characters. — Size large; winter coat usually brown; pale color of
underparts much restricted; color of upper parts dark; color of lips variable;
tail with more than distal half black.

Measurements. — Seven males and 2 females average, respectively (Hall,
1945: 79) : total length 278, 232; length of tail 77. 60; hind foot 36.5. 27.

Distribution. — The extreme north coast of Washington, from the Canadian
boundary south through Whatcom County (Hall, 1945: 80-81).

R( marks. — The range of this ermine is similar to that of the yellow-pine
chipmunk, Eutamias amoenus felix.

DISTRIBUTION MAP
STATE OF WASHINGTON

D-af'ad U~d«. A.ip [

I M'V)7?7 b B CMT mr

Fia. 56. Distribution of the ermine in Washington. A. Mustela erminea
invicta. B. M us tela erminea murica. C. Mustela erminea gulosa. I). M us-
tela erminea fallenda. E. Mustela erminea streatori. V. Mustela erminea
a! y in pica.

Mustela erminea olympica Hall
Mustela erminea olympica Hall, Jour. Mamm., 20:81, February 27, 1945.

Type. — Obtained near head of Sol Due River, Clallam County. Washing-
ton, by V. Bailey, on April 28, 1897; type in United Stairs National Museum.
Racial characters. — Similar to streatori but smaller, especially females.

Measurements. — Twelve males and 6 females average, respectively: total

length 213, 196; length of tail 65. 52; hind loot 31, 23.4 (Hall. 1945: 81).
Distribution. — The Olympic Peninsula, extending southeastward to Olympia.

Ermine 193

Mustela erminea streatori (Merriam)

I'utorni> streatori Merriam, N. Amer. Fauna, 11:13, June 30, 189(5.

Mustela streatori streatori Miller, U. S. Nat. Mus. Bull., 79:96, December 31, 1912.

Mustela cicognanii streatori Taylor and Shaw, Occ. Pap. Chas. R. Conner Mus., no 2 11
December, 1929.

Mustela erminea streatori Hall, Jour. Mamm., 26:76, February 23, 1945.

Type .—Obtained at Mount Vernon, Skagit County, Washington, by D. R.
Lucky on February 29, 1896; type in United States National Museum.

Racial characters. — Similar to fallenda but pale color of underparts less
restricted; tail with less than distal half black.

Measurements. — Two males and 2 females from extreme southwestern Wash-
ington average, respectively: total length 245, 210; length of tail 72, 54; hind
foot 31.5, 25.5; ear 17, 14; weight 72.3, 46.

Distribution.— The lowlands of western Washington; north to Skagit County
and Whidby Island (Taylor and Shaw. 1929: 11), southward and westward to
Ilwaco (M.V.Z.) and eastward to Carson (U. S.N. M.).

Mustela erminea gulosa Hall

Mustela erminea gulosa Hall, Journ. Mamm., 26:84, February 27, 1945.

Type.— Obtained at Trout Lake, Klickatat County, Washington, by P.
Schmid on February 3, 1897; type in United States National Museum.

Racial characters. — Similar to invicta but smaller; winter coat usually white ;
tail less than one-half black; upper lips white.

Measurements. — Five males from Mount Rainier, and 4 females from the
Cascade Mountains, average, respectively (Hall, 1945: 84): total length 253.
20S; length of tail 76. 54; hind foot 30.2, 24.3.

Distribution. — The Cascade Mountains. Known from Skykomish (W.W.
D.) and southward to Mount Adams (Trout Lake, Hall, 1945: 85).

Mustela erminea murica (Bangs)

Putorius (Arctogale) muricus Bangs, Proc. New England Zo61. Club, 1:71. July 31. 1899.
Mustela muricus Miller, U. S. Nat. Mus. Bull., 79:96, December 31, 1912.
Mustela. cicognanii lepta Taylor and Shaw, Occ. Pap. Chas. R. Conner Mus.. no. 2:11. De-
cember, 1929.

Mustela cicognanii muricus Bailey, N. Amer. Fauna, 55:292, August 29, 1936.
Mustela erminea murica Hall, Jour. Mamm., 26:77, February 27, 1945.

Type. — Obtained at Echo, El Dorado County, California, by W. W. Price
and E. M. Nutting on July 15, 1897; type in Museum of Comparative Zoology.

Racial characters. — Similar to invicta but smaller and paler.

Measurements. — A young female from Stay-a-while Spring. Columbia
County, measured: total length 201; length of tail 46; hind foot 24; ear 14
weight 45.8. A female from Butte Creek, Columbia County, measured: 185
50; 26. A male from Baker Creek, White Pine County, Nevada, measured
220; 56; 26; 14.5; weight 57.7.

Distribution. — The Blue Mountains of southeastern Washington.

Remarks. — Two specimens available from the Blue Mountains are darker
than typical murica.

13—4993

194 University of Kansas Publs., Mrs. Nat. Hist.

Mustela frenata Lichtenstein

Long-tailed Weasel

Description.. — Male long-tailed weasels measure about 16 inches
in total length of which 6 inches is the length of the tail. Females are
smaller, measuring about 14 inches in length with a tail 5 inches
long. The body is long and exceedingly thin. The legs are short and
stout with rather large feet and strong, curved claws. The tail is
well-furred, not tapered, but lacks the bushy appearance of the tail
of the fisher. The head is low. wide at the base and abruptly taper-
ing to the muzzle. The ears are erect. low and rounded. The fin-
is short but dense and rather soft, In summer the color of the head.
back, sides and outside of the legs is brown. The throat, chest,
underside of body and insides of legs are yellow or orange. In winter
they may be entirely white, save for the black tip of the tail. The
tail is slightly darker than the back in summer and possesses a long
black tip.

Long-tailed weasels do not hibernate even in the coldest parts of
"Washington. They are active both by day and by night, apparently
doing their hunting at the time of day or night when they can must
easily obtain food. In the Cascade Mountain- where diurnal mam-
mals such as squirrels, chipmunks and pikas are common, weasels
are not uncommonly seen by day. In the lowlands of western Wash-
ington, where they \wi\ on more nocturnal mammals, they are
seldom seen.

The long-tailed weasel is relatively unspecialized in habits. They
climb readily and skillfully. They are active on the surface of the
ground and follow the burrow systems of fossorial animals such
as gophers and mountain beavers.

Weasels seen in the wild rarely exhibit fear of man but rather
are (anions and apt to watch hi- actions. Weasels are also often
hit by cars and the number so killed seems to me to be out of pro-
portion to their actual numbers.

In moving on the surface of the ground the weasel arches it- hack
and contracts the body until the four feel are rather close together.
When the long neck and small head are held upright the animal
presents a surprisingly giraffelike appearance. When climbing, the
long, slim body has a snakelike appearance. A weasel travels swiftly
and erratically in a series of bounds and seems always to know

where the next hole i- situated.

The weasel has been accused of killing birds and doubtless docs

Weasel 195

so when opportunity presents itself. However, in Washington I
have no actual evidence of its killing birds other than domestic fowls.
At Republic, Ferry County, a companion and I saw a weasel enter
the burrow of a ground squirrel (Citelhis columbianus) . The
following day we returned to the area. The weasel was not seen
but a ground squirrel dashed into the hole at our approach. Seem-
ingly the ground squirrel had eluded the weasel. At Conconully,
Okanogan County, we set a number of gopher traps in an alfalfa
field. The following morning an adult male long-tailed weasel was
found in a trap but not a gopher was taken. Near Moses Lake,
Grant County, an adult male weasel was caught in a gopher trap,
but no gophers were taken. At Shelton, Mason County, 50 gopher
traps were set. At daylight the following morning the first trap
visited was found to be pulled into the burrow. When a gentle
tug was given the wire fastening the trap, a decisive jerk at the other
end showed that the catch was alive. If trapped gophers that pull
the traps back into their burrows are pulled out by main strength,
their skins are often torn and damaged. Therefore an attempt was
made to reach into the burrow and pull back the sod. An adult
female long-tailed weasel promptly fastened its teeth into my fore-
finger and clung on, bulldog fashion, to be lifted into the air with
the attached trap swinging. When the left hand was used to force
the animal to release its grip, it fastened onto the left thumb. With
right thumb and forefinger I forced it to release its grip, but was
unable to elude its teeth which again fastened to my right fore-
finger. Only by laying it on the ground and crushing its chest with
my foot could I free myself from the vicious little beast. No gophers
were taken in traps set less than 150 feet from where the weasel
was trapped. In the three instances mentioned above, weasels
had seemingly killed all the gophers in their immediate vicinity.
As regards the gophers near Moses Lake, none was found
a year later in the area where the weasel was taken and only old,
abandoned burrows were seen. T. H. Scheffer (1932: 54) records
other instances of the capture of weasels in gopher burrows.

At the northern limits of the city of Seattle, steel traps were set
for mountain beavers in a rather dense colony of these mammals.
Well-used burrows indicated that approximately 10 individuals were
present. The following morning an exceptionally large male weasel
was found in a trap, but all others were empty. The traps were
left out for two additional nights but no mountain beavers were
taken. A month later the colony seemed abandoned and no evi-

1 •

University of Kansas Ptjbls., Mrs. Nat. Hist.

dence of recent digging was noted. It could only be concluded that
the weasel had killed the animals comprising the colony. A large
mountain beaver weighs three or four pounds, which is 6 or 8 times
as much as a weasel. Edson (1933: 76) recounts trapping 7 weasels
in burrows of mountain beavers near Bellingham. Whatcom County.

Near Forks, Clallam County, a weasel was seen pursuing a young
snowshoe rabbit (Lepus americanus washingtonii) along the edge
of a concrete highway. As our car approached and passed the ani-
mals, they separated, the weasel retreating to the cover of horsetail
[Equisetum) beside the road. The car was stopped 50 feet ahead.
A- we emerged the weasel dashed from cover to intercept the rabbit
in the center of the road. The weasel knocked the rabbit to its side
and. placing its feet on the rabbit's shoulders, bit fiercely at its
neck. It then dashed back to the cover of the horsetails. The
rabbit stood up, made two hops and died. It was approximately
two weeks old.

The pelts of weasels bring the trapper from twenty-five cents to
;i dollar and a half. Only skins in the white winter coat command

DISTRIBUTION MAP
STATE OF WASHINGTON

Via. -VT. Distribution of the Long-tailed weasc-l m Washington. A. Mustela
nata washingtoni. B. Mustela frenata altijrontalis. C. Mustela frenata
adensis. I). Mustela frenata effera.

Weasel 197

the higher price. They are usually taken in traps set for other
animals.

In the western part of the state, long-tailed weasels do not turn
white in the winter; the back is slightly less reddish than in sum-
mer and the underparts are pale yellow or white or may be both
yellow and white. From the summit of the Cascades eastward
weasels become white. The break in winter color seems to occur
slightly west of the main summit of the Cascades. Specimens taken
by marten trappers at Tye and Scenic, near Stevens Pass, w-ere
white on November 15, 1936, but specimens from Skykomish and
Baring, 18 miles to the west, were brown.

The long-tailed w r easel possesses a distinctive odor. It results
from a glandular secretion and, although it has no great carrying
power or lasting quality, it does affect some persons strongly. It
is a heavy, rather sickening scent.

The gestation period of Mustela f. nevadensis has been recorded
as more than 131 days (Hall, 1938B: 250). The gestation period
of the long-tailed weasel of eastern United States has been recorded
as more than 70 days. Three to five young are born in a litter,

usually four.

Mustela frenata nevadensis Hall

Mustela arizonensis Taylor and Shaw, Occ. Pap. Chas. R. Conner Mus., no. 2:11, De-
cember, 1929.

Mustela washingtoni Taylor and Shaw, in part, Occ. Pap. Chas. R. Conner Mus., no. 2:11,
December, 1929.

Mustela frenata nevadensis Hall, Carnegie Inst. Washington, Puhl. 473:91. Novembei 20,
1936.

Type. — Obtained 3 miles east of Baker, White Pine County. Nevada, by
E. R. Hall and W. C. Russell on May 30, 1929; type in Museum of Vertebrate
Zoology.

Racial characters. — Size medium; colors pale; back near Brussels Brown
in summer, chin white and underparts yellow, rarely orange; color in winter
white with black tip on tail.

Measun mints. — A male from Neppel, Grant County, measures: total length
412; length of tail 151; hind foot 43; ear 19. Two males from Yakima,
Yakima County, average: 379; 135; 42.5; 21; weight 176.5. A female from
Ellensburg and one from 4 miles east of Ellensburg. Kittitas County, average:
284; 98; 33; 17.

Distribution. — From the high Cascades eastward, save for the Blue Moun-
tains area. Marginal occurrences are Barron (Hall. 1936: 93) and Easton
(W.W.D.).

It'i murks. — Specimens from the northern Cascades are intergrades between
ru rmii nsis and the form to the west, altijrontalis. Specimens from extreme
northeastern Washington might be referred to oribasa Bangs as readilv as to
nevadensis.

198 University of Kansas Publs., Mus. Nat. Hist.

Mustela f renata effera Hall

Mustela frenata effera Hall, Carnegie Inst. Wa>liinptr>n, Publ. 478:93, November 20, 1986.

Type. — Obtained at Ironside. Malheur County, Oregon, by H. E. Anthony on

September 8. 1912; type in American Museum of Natural History.

Racial characters. — Similar to m vadensix in color but smaller, with smaller,
lighter skull. Color in winter white with black tip on tail.

Measurements. — A female from Prescott, Walla Walla County, measures:
total length 310; length of tail 105; hind foot 34.

Distribution. — Southeastern Washington, south of the Snake River. Speci-
mens from Walla Walla and Prescotl have been examined.

Mustela frenata washingtoni (Merriam)

Putorius washingtoni Merriam, X. Amer. Fauna, 11:18, June 30, 1896.
Mustela washingtoni Miller, U. S. Nat. Mus. Bull.. 79:98, December 31, 1912.
Mustela frenata washingtoni Hall, Carnegie Inst. Washington, Publ. 473:106, November
20, 1936.

Type. — Obtained at Trout Lake, Skamania County. Washington, by D. N.
Kaegi on December 15. 1895; type in United States National Museum.

Racial characters. — Color rich, dark; hind feet free of color of underparts.

Measurements. — A male from Spray Park, Pierce County, measures: total
length 423; length of tail 164; hind foot 52.

Distribution. — Higher Cascades from Alt. Rainer (M. V. Z.) south to Mt.
Adams (Taylor and Shaw, 1929: 11).

Mustela frenata altifrontalis Hall

Mustela saturata Taylor and Shaw, Occ. Papers Chas. R. Conner Mus., no. 2:11, De-
cember, 1 929.

Musi, In frenata altifrontalis Hall, Carnegie Inst. Washington, Publ. 473:94, November 20,
1936.

'/'///« . — ( Obtained at Tillamook, Tillamook County. < Iregon, by A. Walker on
July 10. 1928; type in Museum of Vertebrate Zoology.

Racial characters.- Similar to nevadensis but darker above in summer and
richer orange beneath; winter color of upper parts dark brown, underparts
paid yellow or white.

Measurements. — Four males and 2 females from western Washington ave-
rage, respectively: total length 411.5, 267; length of tail 11."). 1375; hind foot
50, 43; ear 26. 21.5; weight 227. 136.7 grams.

Distribution. — From the Cascade- Mountains to the Pacific. Marginal
occurrences are Etockporl (Hall. 1936: 95) and Tye (W.W.I).).

Mink

1!)!)

Mustela vison energumenos (Bangs)
Mink

Putorius vison energumenos Bangs, Proc. Boston Soc. Nat. Hist., 27 :5, March, 1896.
Mustela vison energumenos Miller, U. S. Nat. Mus. Bull., 79:101, December 31, 1912.

Type .—Obtained at Sumas. British Columbia, by A. C. Brooks on September
23, 1895; type in Museum of Comparative Zoology.

Measurements. — A male from Seattle, King County, measures: total length
540; length of tail 180; hind foot 66; weight 1 pound, 15 ounces.

Distribution. — Throughout the state save for the Columbian Plateau; re-
corded from Neah Bay (Taylor and Shaw, 1929: 11) in the northwest, Ilwaeo
(M.V. Z.) in the southwest, Metaline (W.W.D.) in the northeast, and the
Blue Mountains (Dice, 1919: 12) in the southeast.

DISTRIBUTION MAP
STATE OF WASHINGTON

Fig. 58. Distribution of the mink, Mustela vison energumenos, in Washington.

Description. — Because of the value and lasting popularity of its
fur, the mink is known to all. Few persons, however, recognize the
animal in the wild. The mink possesses the long, slim body and
short, rather stout legs of a weasel but has a more bushy tail. The
mink is larger than the weasel. Large males weigh up to 3 pounds ;
females 1% pounds. Males measure about 2 feet in length, of

200 University of Kansas Publs., Mrs. Nat. Hist.

which the tail comprises eight inches. Females measure about 20
inches and have tails 7 inches long. The color is rich, dark reddish
or chocolate brown. The underparts are slightly paler than the
hack. There are usually small white markings on the chin, chest
or other part of the ventral surface. These markings sometimes take
the form of narrow white lines.

The mink ranges from the Atlantic to the Pacific and from Alaska
south to Florida, New Mexico and central California. It is active
throughout the year and is principally, though not exclusively, noc-
turnal. McMurry (1940: 47) records three mink seen at 3:30 p. m.
on August 8, 1939, at Packwood Lake, Lewis County, one of which
was carrying a garter snake two feet in length. This mustelid is
semiaquatic, living along rivers, streams, lakes and salt water and
spends most of its time along the shore, on muddy or sandy beaches.
In tins respect it resembles the raccoon. Whereas the raccoon
spe&tls a part of its time on the land, the mink spends a corres-
ponding amount of its time in the water. It is an excellent swimmer,
able to overtake and capture fish in the water.

In the San Juan Islands minks have forsaken the shoreline and
roam over the uplands, feeding on the abundant, feral domestic
r ibbits. I found evidence of their presence far inland, miles from
water, in grassy and bushy wastes; along the beaches their tracks
were rarely seen. Along Puget Sound, minks spend part of their
time on the beaches, feeding on dead fish and other marine animal
life. These animals, however, seem to live along the rivers and
streams flowing into the sound. Along the ocean coast, some minks
seem to live exclusively in the marine shoreline habitat.

The mink ascends some distance into the Cascade and Olympic
mountains along the larger watercourses. There are reliable re-
ports of minks from Heart Lake and bake Dorothy, King County,
high in the Cascades. These animals were seen in the summer but
they may live in higher parts of the Cascades, at least, about some
of the larger lakes throughout the year. As far as is known there
are no minks on the Columbian Plateau.

The food probably varies with locality. . Monti the ocean beaches
they e.-it dead sea birds, stranded fish, crustaceans, and mollusks.
Along Puget Sound their Hacks are commonly -ecu following the
caked, decaying seaweed and debris that collects at the high tide
mirk. Svihla and Svihla (1931: 22) captured a mink that was
feeding on the beach of the Olympic Peninsula. This individual, in
captivity, was able to open and \v{^\ on cliims. Fur trappers report
that minks U-{'i\ on mountain beavers, and that the Hcsh of the

Mink 201

mountain beaver is the best bait for attracting minks. Along
streams and lakes, minks are thought to feed on mice, birds, fish,
crayfish and thin-shelled, fresh-water mussels. The muskrat forms
an important item of diet near the larger lakes and streams. Musk-
rats trapped near Seattle were often attacked by minks and either
eaten or so slashed and torn that their pelts were worth but a frac-
tion of what they would bring in an undamaged condition.

The tracks of a mink noted in freshly fallen snow about a gar-
bage dump on the shore of Lake Washington, Seattle, indicated that
the animal had been hunting house rats, which were numerous there.
Minks are not uncommon in the marshes along Lake Washington
near the University of Washington campus, Seattle, where I dis-
covered evidence of their preying on ducks and coots. The ducks
included mallards and green- winged teal, species most apt to alight
in small pools in the rushes where a mink might find cover. I found
the fresh tracks of one mink about a half-eaten golden tench 8
inches long. The tench is a fish of the sucker tribe, introduced
into the lake. Tracks of minks are not uncommonly seen along
rivers and streams under conditions that indicate they were hunting
crayfish.

The raw pelt of the mink sells usually at from 10 to 20 dollars.
The fur is in constant demand and fluctuates in price less than most
furs. The average trapper takes a few minks each year along with
his catch of muskrats, raccoons, and skunks but some trappers have
taken as many as 100 minks in a winter in areas where the animal
is especially numerous.

•Jin'

University of Kansas Ptjbls., .Mrs. Nat. Hist.

Gulo luscus luteus Elliot
Wolverine

Gulo luteus Elliot, Field Columb. Mus., Publ. ST. zobl. ser. 8:260, December, 1903.
Gulo luscus luteus Grinnell, Dixon, and Linsdale, Fur-bearing Mamm, California, Univ.
California Press, Berkeley, p. 251, July 22, 1937.

Type.— Obtained on Mt. Whitney. Tulare County. California, by E. Heller;

type in Field Museum of Natural History.

Measuri merits. — The dried skin of a wolverine taken 3 miles south of Kiwi-
side. Okanogan County, was 4 feet in length. The animal weighed 40 pounds
(Scheffer, 1941: 37).

Distribution. — Timber-line region of the Cascades Mountains. From
Robinson Creek (Scheffer, 1938: 8) south to Mount Rainier (Taylor and Shaw.
1929: 12).

Remarks. — Specimens from Washington are intermediate in character
between luteus and the race inhabiting British Columbia, although available
skulls are closer to luti us.

Description. — Largo males measure more than three feet in length.
of which the tail make- up 10 inches; they weigh as much as 50
pounds (Grinnell. Dixon and Linsdale. 1937: 253). The body is
wide and stocky; the tail is short; the legs, especially the forelegs,
are short, thick and powerful; the feet are large and are armed with
long, curved claws; the head is wide; the jaws are powerful with
heavy teeth. The pelage is long and rather shaggy. Face, legs,

Fia 59. Wolverine (Gulo luscus), mounted specimen, male, trapped by
billy Robinson aboul 1902 on Billy Robinson Creek, Okanogan County, Wash-
ington. (Fish an, I Wildlife Service photo by Victor B. Scheffer. No. 76.)

Wolverine

203

back and terminal portion of the tail are dark, blackish brown but
the forehead is grayish, contrasting with the dark color of face and
top of the head. Two broad, yellowish-brown bands begin on the
side of the shoulders and pass back, low on the sides, to the hips
where they join across the back and base of the tail.

The wolverines, nominally of several species, form a compact
group that ranges over the boreal regions of the Old and New world.
In North America they range southward from the Arctic to California
and Colorado.

Fig. 60. Wolverine (Gulo hiscus), male, trapped by Reuben R. Lamb,
March 22, 1941. three miles south of Riverside, Washington; reported to
weigh 40 pounds; dried pelt 3x4 feet. (Reuben R. Lamb photo.)

204

University of Kansas Publs., Mrs. Nat. Hist.

DISTRIBUTION MAP
STATE OF WASHINGTON

fcc" in? k ~ i

Fig. 61. Distribution of the wolverine. Gulo luscus luU us, in Washington.

The wolverine is adapted to boreal conditions and is most abun-
dant in the arctic. In the Cascades it occurs only at or near timber-
line. Except the wolf and the possibly extinct grizzly bear, the
wolverine is the rarest carnivore in Washington. Probably it has
been rare since the retreat of the continental glaciers, for suit-
able habitat for wolverines is not abundant and each animal requires
a large area over which to ranise. An unusual record, possibly of
a wandering animal, is given by Scheffer (194] : 37). This is of an
adult male taken in the Okanogan Valley 3 miles south of Riverside.
Okanogan County, on March 22, 1941.

River Otter 205

Lutra canadensis (Schreber)

River otter

Description. — The otter has the long body of many mustelids,
but is specialized for aquatic life. Males are slightly larger than
females. Large adults measure more than 3^ feet in length, of
which the tail comprises 18 inches. The largest adults may weigh
as much as 25 pounds. The body is elongate but rather plump. The
forelimbs are small and laterally placed. The hind legs are the
heavier, and are situated posteriorly. The feet are large and webbed
for swimming. The tail is long, thick at the base and gradually
tapering to a narrow tip. The head is small and rounded. The
ears are tiny and the eyes are of moderate size. The upper lips are
large, somewhat spaniellike, and support a mustache of stiff vi-
brissae. The fur is short, dense and soft. The tail is sleek with
short, posteriorly-directed fur, and is not bushy. The fur is rich,
dark chocolate brown in color, slightly paler on the animal's under-
pays than on its upper parts.

Otters range over Eurasia, North America and South America.
Those occurring in North America north of Mexico are thought to
belong to a single species, canadensis. Several species from tropical
America may eventually be shown to also be races of canadensis.

The aquatic habits of the otter allow it to exist in several life-
zones, in both marine and fresh-water habitats. The principal en-
vironment is the streams, rivers and lakes of the Transition Life-
zone, but along the larger rivers, otters range up into the Canadian
Life-zone and along the Snake and Columbia rivers they enter the
Upper Sonoran Life-zone. They are active throughout the year.
As we have observed them, they are principally nocturnal.

Otters are extremely powerful swimmers. Tracks along the North
Fork of the Tolt River, King County, showed where otters had en-
tered water that flowed over a stony bottom at the velocity of
rapids. Their occurrence in the Tolt, Skykomish and Snoqualmie
rivers where these are swift mountain streams indicates exceptional
swimming ability. After emerging from the water, an otter often
follows along the shore, sometimes for miles.

In the San Juan Islands the otters have taken to a marine ex-
istence. In the summer of 1938, abundant opportunity offered to
observe the otters at Thatcher Bay, Blakely Island. Here, in the
evening, bats were shot as they flew over the calm water of the bay.
Specimens killed were retrieved by rowing out to them with a dory.
For the first two nights bats were collected and retrieved without

206 University of Kansas Publs., Mrs. Nat. Hist.

incident. On the third night, several of the hats vanished between
the time they were killed and the time that the boat was launched
from the rocky beach. The presence of a shark or other large fish
was at first suspected, but observation showed a group of three or
possibly four otters to be the thieves. On succeeding nights the
animals became bolder. A dead hat became the object of a race
between otters and collector. They completely ignored shouts,
dodged stones hurled at them, and stole almost all the bats shot,
t Inly a sense of humor and desire to study the animals saved them
from a load of fine shot. Bat collecting was given up in disgust.

These otters were remarkably seallike in many actions. In swim-
ming their heads, shoulders and part of their back were exposed.
In resting in the water, only the round head remained above the
surface. They were never seen to float prone on the surface or rest
on their backs, as does the sea otter. They were noisy swimmers,
splashing with paws or heads as they dived. Their eyesight was
remarkable. Stones the size of a walnut were hurled at them from
distances as short as 25 feet. All were dodged with little effort. In
this connection the experience of two trappers who caught an otter
in the Samamish River near Woodinville, King County, is inter-
esting. The trap holding the otter had been attached by a wire 6
feet long to a pole on the bank. The wire was detached from the
pole; while one trapper held the wire, the other tried to strike the
otter with an oar as the boat drifted over deep water. The otter
was free to swim and dive for the length of the wire and trap chain.
It detected, and evaded by dodging or diving, every blow directed
at it and not until the animal tired, a half hour later, could it be
killed. The trappers then noted that the splashing of oars and otter
had nearly filled the boat with water. The trappers were exhausted.

The otters of Blakely Island were not unique in occupying a ma-
rine habitat. At Strawberry Bay, Cypress Island, on .Inly .">, 1938,
fishermen brought in a "strange animal" caught by them at Black
Rock, a tiny bare and isolated islet f) miles to the west. The heavy
box holding the animal was opened, disclosing a very frightened
young otter. Questioned, the fishermen stated that lour young and

otic adult had been >een in the surf. They had thought the animals

a specie- o\ seal, and were somewhat surprised at being successful

in their attempts to catch one. The otter was too young to tare
for itself and was kept in camp in the hope that it might, he reared
and released. It ale a few fresh herring and candlefish and drank
-oiuc condensed milk. It refused whole salmon and sea bass hut

River Otter 207

ate some skinned and honed strips of these fish. It died a week
after ii was captured. This young otter made a ticking sound, al-
most a chirp. A variety of crying and whimpering sounds were also
made, and when petted it grunted in satisfaction. If surprised or
when first picked up it gave a deep harsh growl, unexpectedly
vicious in sound, for such a small animal.

The otters at Blakely Island fed on a great variety of food. The
only food they were actually seen to eat was candlefish, a slim
silvery fish (i inches in length. On several occasions an otter was
seen swimming with the head of a candlefish held fast in its mouth
and the fish's body extending out in a silvery bow. Many feces of
these otters were composed entirely of the feathers of grebes and
scoters. These birds were probably not killed by the otters but died
a natural death or were shot by men. At that camp scarcely a
day passed in which at least one person was not seen firing a .22
caliber rifle from a cabin cruiser into the flocks of scoters at the
mouth of the bay. Dead sea birds were common on the shore and
doubtless furnished food for otters as they did for crows and eagles.
The majority of the otter feces examined at Blakely Island were
composed of the remains of invertebrates. The small mussel (M y-
tilus edulis) was most abundant. Crabs fonned an important part
of their diet. Strangely enough the shore crab (Hemigrapsns nudus)
and the porcelain crab {Petrolisthes eriomerus) were seldom eaten
though they were abundant beneath the rocks upon which the feces
were found, and formed the principal food item of raccoons. These
species of invertebrates live beneath rocks and the raccoon probably
reaches beneath the rocks to catch the crabs with its handlike fore-
feet, as the otter is unable to do. The edible crab (Cancer produc-
tus) and kelp crab {Telmessus cheiragonus) were most often eaten
by otters. Also eaten were snails (Margarites, Littorina), bitter
oysters (Pododesmns macroschisma) , unidentified pelecypods, barn-
acles (Balanus), one chiton iMopalia muscosa) . and once a starfish.
The invertebrate remains, save for the snails, were crushed and
broken.

Fish of many species were abundant in these waters. Salmon
were often seen leaping near the playing otters. Yet no scales or
other remains of fish were detected in the feces. Candlefish possess
tiny scales that could probably be seen by microscopic examina-
tion only. As for the bird remains, the quantity of feathers
consumed is notable. The otters apparently do not pluck birds, as
do minks. Indeed, of the material eaten, the food value by volume

208 University of Kansas Publs., Mrs. Nat. Hist.

t:

Fig. G2. River olter (Lulra canadensis pacifica), (rapped in Columbia Na-
tional Forest, Washington, February, 1937. (Adolph Roth photo.)

River Otter

I'd!)

seems extraordinarily low. A great quantity must be eaten to
nourish an animal as large and as active as an otter. The number of
fecal droppings seen each morning indicates that this is the case.

The favored food of the otter in fresh water streams and lakes
is the crayfish. The greater part, perhaps 80 per cent, of otter feces
seen in the lowlands of western Washington, were composed of cray-
fish remains. Some fish are also eaten and locally otters may do
damage among the steelhead trout. Even worse, from the fisher-
man's standpoint, they frighten and scatter the schools of steel-
head trout. Frogs are eaten, as well as the thin-shelled mussels.
The feathers of coots were twice seen in otter feces near Cottage
Lake, King County.

The fur of the otter is dense and beautiful but the skin is rather
heavy. . Changes in styles cause great fluctuation in the value of
the otter's pelt.

L- —47

DISTRIBUTION MAP
STATE OF WASHINGTON

50

75

100

MILES
of WPA P. e) tc> 6079-5797 by CUT 1937

I

123

122

Fig. 63. Distribution of the river otter in Washington. Unshaded Lulra
canadensis pacifica. Shaded Lutra canadensis vancouverensis.

14—4993

210 University of Kansas Pibls., Mus. Nat. Hist.

Lutra canadensis pacifica Rhoads

Lutra hudsonica pac\ ca Rhoads, Trans. Amer. Philos. Soc, n. s., 19:420. September, 1898.
Lutra canadensis pacifica Allen, Bull. Amer. Mus. Nat. Hist.. L0: 160, November 10, 1S98.

Type. — Obtained at Lake Keeehelus. Kittitas County. Washington (altitude
3.000, rather than 8000 feel as stated in original citation); typo in Academy
of Natural Sciences of Philadelphia.

Measun nn nts. — An adult male from the Lower Calawah River, near Forks.
Clallam County, measured (skinned carcass): total length 1205; length of tail
490; weight (entire) 23 pounds. A male from Puget Island. Wahkiakum
County, measured (skinned carcass): total length 1250: length of tail 490.
A female from Satsop. Grays Harbor County, measured (skinned carcass):
total length 1205; length of tail 440. A female from Calawah River near
Forks. Clallam County, measured: total length 1062; length of tail 430;
hind foot 120; ear 23; weight 14% pounds.

Distribution. — Watercourses throughout the state, scarcer east of the Cas-
cades; recorded at Forks (V. B. S.) in the northwest, Puget Island (Y.B.S.)
in the southwest, Colville (Taylor and Shaw, 1929: 12) in the northeast, and
Touchet (Taylor and Shaw. 1929: 12) in the southeast.

Lutra canadensis vancouverensis Goldman

Lutra vancouverensis Goldman, Proc Biol. Sue. Washington, 18:186, November If), 1935.

Type. — Obtained at Quatsino, Vancouver Island. British Columbia, by H.
O. Berg in 1905; type in United States National Museum-.

Racial characters. — No adults, referred to this race from Washington, have
been examined. Said to have broader skull than pacifica.

Measurements. — The only available specimen, a young female from Black
Rock. San Juan County, measured: total length 792; length of tail 247; hind
foot 103; ear 22.

Distribution.- — The San Juan Islands of northern Puget Sound and adjacent
marine waters

/,% marks. — The otters of the San Juan [slands are referred to this race on
geographical grounds. Numerous islam Is connect Vancouver Island with the
San Juan Islands, and intervening channels are not too wide for otters to
swim. Adult material may show that the otters of the San Juan Islands are
referable to pacifica or to an undescribed race rather than to vancouverensis.

Sea Otter 211

Enhydra lutris nereis (Merriam)

Sea otter

Latin- lutris nereis Merriam, Proc. Biol. Soc. Washington, 17:159, October 6, 1904.
Enhydra lutris mnis Grinnell, Univ. California Publ. Zool., 21:316, January 27, 1923.

Type. — Obtained on San Miguel Island. Santa Barbara Islands, by G. M.
McGuire in 1904; type in United States National Museum.

Ditiiibution. — Formerly occurred along the ocean coast of Washington.
Now extinct there.

Description. — Adult sea otters reach a length of six feet. The body-
is stout, long and round; the tail is short, flattened and about a foot
in length. The legs are short and thick. The hind feet are webbed
for swimming. The head is rounded with small eyes and ears, and
a seallike mustache of stiff bristles on the muzzle. The upper lip
is thick and heavy, drooping over the sides of the mouth in a spaniel-
like fashion. The fur is long, soft and extremely dense. The color
of the body is various shades of rich, dark brown. Scattered white
hairs often give the coat a frosted appearance. The heads of the
adults are pale grey, occasionally almost white.

The sea otter once ranged from the Gulf of California north
along the west coast of North America to Alaska, the Aleutian
Islands, Siberia and Japan. It is now found only in a small area
along the coast of California and in remote islands of the north
Pacific. Once abundant along the coast of Washington, it is now
completely extinct there.

The habits of sea otters living on the California Coast in the
vicinity of Carmel have been described by Fisher (1939: 21-36).
Available information on the sea otter in Washington has been given
by Scheffer (1940B: 369-388). The search for sea-otter skins influ-
enced the exploration and settlement of the northwest. Captain
James Cook obtained skins of sea otter at Nootka Sound, Vancouver
Island, in 1778. The mouth of the Columbia River was discovered
by Captain Robert Gray on May 11, 1792, while trading for sea
otter skins.

According to Scheffer, the bulk of the sea-otter population on the
Washington coast was concentrated between the mouth of the Co-
lumbia River and Point Grenville at the mouth of the Quinault
River. The otters were found several miles from land in extensive
kelp beds. The principal food of the sea otter in Washington seems
to have been the short-spined purple sea urchin (Strongylocentrotus
purpuratus). The animals were social, living in herds of up to 400
individuals. They never came out on land, living, sleeping, and
bringing forth their young on the kelp beds of the open ocean. The
single young was born at any month of the year.

212 University of Kansas Publs., Mus. Nat. Hist

Spilogale gracilis Merriam

Civet cat or spotted skunk

Description. — The civet eat is slightly less than 18 inches in total
length, of which the tail constitutes 6 inches. Large, fat animals
may weigh more than a pound. The body is long and moderately
stout, heavier than that of the weasel but slimmer than that of the
striped skunk. The tail is long with long, plumelike fur. The legs
are slender and of moderate length. The feet are small with long
claws. The head is small and triangular with low, erect ears and
large, bright eyes. The fur is short but soft and silky. The ground
color of the civet cat is black. Wavy lines and spots of white or
salmon intercept the black in a complex pattern. Variations in color
patterns include width of stripes, waviness of stripes, and breaking
up of the stripes into spots.

Spvlixjiih is restricted to America; it ranges from southern
British Columbia south into Central America. Several species are
now recognized, but additional material probably will show inter-
gradation between some of them and reduce the number.

The name "spotted skunk" is seldom used by trappers or other
persons familiar with the animal. Nor is it appropriate lor the
distinctive color pattern is composed of short stripes. Neither is
the term "civet cat" appropriate, for the true civet is found in Asia.
The name civet cat is well established, however, and will doubtless
remain in general use.

The civet cat is principally a lowland animal, but has been re-
corded from the Nisqually entrance of Mt, Rainier National Park
(Brockman, 1939: 70). Its rather generalized habits allow it to
exist in areas that will not support larger carnivores or species with
specialized food habits. It dwells in areas of thickets, brushy
tangles or deep woods. In southeastern Washington it lives in
rocky places as well as in river-side thickets of willow and cotton-
wood. Where conditions are suitable it ranges up into the Cana-
dian Life-zone but is most abundant in the Transition Life-zone.

The civet cats, in contrast to the striped skunks, are exclusively
nocturnal. They are active the year around in western Washington
but move about less in -pells of unusually cold weather. We have
taken them, however, in freezing weather.

In spite of their abundance and wide range, civet cats are known
to U-w persons other than trappers, perhaps because of their ex-
clusively nocturnal habits. They do occasionally make people

Skunks 213

aware of themselves by taking up residence about old buildings.
They are rather noisy at times. At the Tolt River, 10 miles south-
east of Duvall, King County, we were asleep in an old building
when a stamping noise from the next room awakened us. Investi-
gation revealed a civet cat indulging in a series of short, stiff-legged
hops. The forefeet, were held slightly ahead and six or eight hops
made. The animal would then relax, turn, and prance off in a new
direction. The amount of noise made by its feet striking the board
floor was surprising. The civet cat seemed to enjoy the noise it
made, and the prancing may have been a method of playing.
Trapped civet cats have been seen to make a similar but shorter
series of hops at the trapper's approach, which might have been a
warning. In the case described, however, the noise of the animal's
actions was heard before the animal was seen and presumably
when it was unaware of human presence. The civet cat also makes
a drumming noise similar to the drumming of a wood rat. This
was heard twice from wild animals that did not suspect an ob-
server's presence, and once from a captive animal. While an
animal was drumming I never had the opportunity to watch the
actions closely enough to describe them. Apparently the drumming
is done with the forefeet.

The civet cat is said to be a good climber and to do some of its
hunting in trees. One trapped civet cat climbed to the limit of a
trap chain and wire in a low bush near Cottage Lake, King County,
but, in Washington, I have no other evidence of this animal climb-
ing. Near Cottage Lake, we took a civet cat in a trap set beneath
2 inches of water at the side of a stream. The animal had either
been swimming or wading.

Trapped civet cats rarely dig holes near the traps, as striped
skunks often do. One caught near Ocean Park, Pacific County, did
dig a hole beneath a log and forced earth and leaves over the en-
trance. I walked within a few feet of the animal in searching for
the trap. Not until the trap chain was seen did I find the civet cat.
None was taken in traps set in mountain beaver burrows, although
I have caught both striped skunks and weasels in such burrows.

Some trappers state that the musk of the civet cat smells different
from that of the striped skunk. I think the odor is slightly more
acrid and that it does not carry so far as the skunk musk. The civet
cat is far more active and nervous than the striped skunk. Whereas
the striped skunk almost never throws its scent when trapped, the
civet cat almost invariably does so, apparently when the trap closes

21-4

University of Kansas Ptjbls., Mtjs. Nat. Hist.

about its legs. Striped skunks in traps move slowly and steadily
but civet cats jump, roll and squirm erratically.

In the winter of 1934 a large male civet cat was taken near
Cottage Lake, King County. It had been killed and partially eaten
in the trap. The ground about the set was torn up, indicating that
the civet cat had put up a fight. The area, for many feet about
smelled of skunk musk. The body of the civet cat was used for bait
and the trap reset. The following morning a large male striped
skunk was in the trap. Its skin bore superficial cuts in several places
about its neck and rump, and it was doubtless the animal that had

DISTRIBUTION MAP

STATE OF WASHINGTON

MILES

Fig. fit. Distribution of the civel rat in Washington. A. Spilogale gracilis
latifrons. B. SpilogaU gracilis saxatilis.

killed the civet cat. ruder ordinary circumstances a striped skunk
could scarcely catch the tar swifter and more agile civet cat. One
civet cat in a trap was killed and partly eaten by a horned owl. Save
lor dislodging a few feathers, the civet cat seemed to have done little
damage to the owl. The thick cover inhabited by the civet cat
ordinarily protects it from owls. Dead civet cats run over by cars on
the highway are rarely seen.

The stomach of a specimen from Ocean Park, Pacific County,
contained the remains of three red-backed mice (ClethrionomyS

Skunks 215

calif ornicus) . The stomachs of most specimens trapped were empty.
Mice, birds and insects probably constitute the bulk of the food.

Notes on the early life of Spilogale interrupta, a species related
to the one occurring in Washington, have been published by Crabb
(1944: 213-2211.

The fur of the civet cat is of little value; in recent years trappers
have received from fifteen cents to a dollar for large pelts. Because
of its habit of throwing scent when in the traps, most trappers discard
the animals without skinning them.

Spilogale gracilis saxatilis Merriam

Spilogale saxatilis Merriam, N. Amer. Fauna, 4:13, October 8, 1890.
Spilogale gracilis saxatilis Howell, N. Amer. Fauna, 26:23, November 24. 1906.

Type . — Obtained at Provo, Utah County, Utah, by V. Bailey on November
13, 1890; type in United States National Museum.

Racial characters. — Size small; white areas extensive, especially on tail but
white triangle on head small ; white areas less often tinged with salmon or
orange than latijrons.

Measurements. — Howell (1906: 32) gives the measurements of a male from
Harney, Oregon, and the average of 3 females from Oregon as, respectively:
total length 455. 360; length of tail 155. 129; hind foot 50, 40.

Distribution. — Southeastern Washington, north to Kamiak Butte (Taylor
and Shaw, 1929: 12).

Spilogale gracilis latifrons Merriam

Spilogale phenax latifrons Merriam, N. Amer. Fauna, 4 : 15, October 8, 1890.

Spilogale olympica Elliot, Field Columb. Mus., Publ. 32, zool. ser. 1:270, March, 1899
(type from Lake Sutherland, Clallam County, Washington).

Spilogale phenax olympica Howell. N. Amer. Fauna, 26:33, November 24, 1906.

Spilogale gracilis latifrons Grinnell, Dixon and Linsdale, Fur-bearing Mamm. California,
Univ. California Press, Berkeley, p. 301, July 22, 1937.

Type. — Obtained at Roseburg, Douglas County, Oregon, by T. S. Palmer
on July 13, 1889; type in United States National Museum.

Racial characters. — Larger than saxatilis with less extensive white markings,
especially on tail, but white markings on head larger; white usually tinged
with salmon or pale orange.

Measurements. — Means for four males, from Pacific County, are total
length 411; length of tail 136; hind foot 47.5; ear 27. One weighed 784 grams.

Distribution. — Western Washington from the western edge of the Cascades
westward. Marginal records, given by Taylor and Shaw (1929: 12) are Hamil-
ton, on the north; Lake Keechelus. on the east; and Carson, on the south.

Remarks. — Comparison of adult specimens of civet cats from western
Washington and western Oregon shows no reliable character for recognition
of two races. No difference in tail length exists. Narrowness of rostrum was
the only diagnostic character found by Howell (1908: 34) to separate latifrons
and olympica. We have carefully compared civet cats from western Oregon,
southwestern Washington and the area about Seattle. Some local variation
exists but overlapping is great and specimens from Washington do not have
rostra that average narrower than specimens from Oregon.

216 University of Kansas Pibls., Mus. Nat. Hist.

Mephitis mephitis i Schreber)

Striped skunk

Description. — The striped skunk is a heavy-bodied animal about
the size of a house cat. The legs are of moderate length and stout
and the hind feet are large. The claws of the forefeet are long,
strong and curved. The head is small and pointed, with small eyes
and ears. The tail is long, nearly equal in length to the head and
body. The fur is long, soft and shiny, and is jet black with sharply
contrasting white markings. These consist of a narrow stripe on
the forehead, a broad band on the neck that diverges into two
stripes on the back. The two lateral stripes fuse on the rump. The
tail has long black hairs some of which are white at the base.

Striped skunks range over North America from central Canada
southward to southern Mexico. Two species are recognized by Hall
(1936: 64), namely mephitis and macroura. The latter species' is
found in Mexico and parts of the southwestern United States.

Skunks are principally nocturnal but are sometimes active in the
morning and evening, especially on cloudy days. They prefer rela-
tively open country such as logged-over land, old fields and riverside
and streamside thickets. Their dens usually consist of old Aplo-
dontia burrows or burrows which they, themselves, dig under stumps
or log jams, the floors of old buildings or among rocks. They feed
on a variety of animals and wild fruit. Along Puget Sound they
wander over the beaches when the tide is out, eating stranded fish,
crustaceans and other marine animals. The purple shore crab
(Hemigrapusus nudus) forms a staple article of diet. Along streams
and rivers they wander along low, muddy banks and sand bars
searching for fish, crayfish, insects and insect larvae. The larger
water beetles (Dytiscidae) are often eaten; feces are at times com-
posed entirely of their shells. For the most part, however, skunks
have no regular food habits but eat such insects, small mammals,
birds or refuse as are available.

The skunk is famous for the musk which it uses as a defensive
weapon. This highly volatile liquid is ejected from two small,
nipplelike ducts situated in the edge of the anus. The consistency,
color, and distance to which the musk can be discharged varies with
the amount thrown. The first discharge or two is usually a fine,
pale yellow spray, which can be accurately directed to a distance
of 25 feet. The third discharge consists of small drops of heavy,
bright-yellow liquid thai travels an inching curve, 5 feet high, reach-

Skunks 217

ing a maximum distance of about 10 feet. Later discharges consist
of heavy yellow mucus and can be thrown only a few feet. As many
as seven or eight discharges are possible. Skunk musk is acrid and
pungent in order. In quantity or at close range it is choking. Tn
small quantities it is not unpleasant. It is extremely lasting, some-
times being noticeable for months on clothes or buildings. It is
soluble in gasoline and clothes may be de-scented by several wash-
ings in that fluid.

The striped skunk is an even-tempered animal. Its defensive
fluid is discharged only when it is cornered or attacked. A trapped
skunk rarely releases musk. A man, by speaking softly and mov-
ing slowly, can come within a distance of six feet of a trapped
skunk. Experienced trappers utilize this fact to approach and shoot
trapped skunks through the head or neck and so produce odorless
furs.

The skunk is potentially a source of considerable income to trap-
pers in Washington. The value of their furs varies with the de-
mand but large prime skins usually bring from $1 to $4. In eastern
Washington, where trapping for coyotes and other terrestial mam-
mals is carried on, the skunks taken incidentally are an important
source of revenue. In western Washington they are often abundant
but are seldom trapped. The most sought pelts in western Washing-
ton are the mink, muskrat and raccoon; all semiaquatic species.
Skunks are rarely taken in traps set for these mammals and few
trappers bother to set overland trap lines for skunks.

The young of the striped skunk usually number four to six. They
are born in late May or early June in western Washington ; possibly
later in eastern Washington.

Mephitis mephitis hudsonica Richardson

Mephitis americana var. hudsonica Richardson, Fauna Boreali-Americana, 1 : 55, 1829.

Mephitis hudsonica Bangs, Proc. Boston Soc. Nat. Hist., 26:536, July 31, 1895.

Chincha hudsonica Howell, N. Amer. Fauna, 20:24, August 31, 1901.

Mephitis mephitis hudsonica Hall, Carnegie Inst. Washington, Publ. 473:65, November 20,
1936.

Type. — Obtained on the "plains of the Saskatchewan, Canada."

Racial characters. — Size moderate; stripes diverging anteriorly on back of
neck; tail long with white stripe reaching well out, nearly to tip; zygomatic
arches nearly parallel.

Measurements. — Howell (1901:24) gives the average of 3 males from Sas-
katchewan, Montana and Wyoming, and of 3 females from Montana and
Idaho, as, respectively: total length 726, 602; length of tail 268, 250; hind
foot 82, 71.

218 University of Kansas Ptjbls., Mus. Nat. Hist.

Distribution. — Northeastern Washington and the eastern edge of the north-
em Cascade-, south probably to the Wenatchee Mountains. Recorded west
to Oroville (W.W.D.) and Timentwa (WW. D.) and south to Spokane (Tay-
lor and Shaw. 1929: 12).

Mephitis mephitis major (Howell)

Chincha occidentals major How..]], N. Amer. Fauna, 20:37, Augu-t 31, 1901.
Mephitis mephitis major Hall, Univ. California Publ. Zool., 37:2. April 10, 1931.

Type. — Obtained at Fort Klamath, Klamath County, Oregon, by B. L. Cun-
ningham on January 5, 1898; type in United States National Museum.

Racial characters. — Slightly larger than hudsonica with white stripes diverg-
ing on shoulders rather than neck.

DISTRIBUTION MAP
STATE OF WASHINGTON

SS 26 50 75 100

MM — ■ ' ■ ■

MILES

Fig. 65. Distribution of the striped skunk in Washington. A. Mephitis
mephitis hudsonica. B. Mephitis mephitis major. ('. Mephitis mephitis rio-
tata. 1). Mephitis mephitis spissigrada.

Measurements. A young male from 1 mile aorth of Burbank, Walla Walla
County, measures: total length 171; length of tail 205; hind fool 68; ear 30;

weight 815 grams.

Distribution- Southeastern Washington, south of the Snake River and east
of the Columbia River, occurring wesl to Burbank (M. V.Z.).

Skunks 219

Mephitis mephitis notata (Howell)

Chincha occidentalis notata Howell, N. Amer. Fauna, 20:36, August 31, 1901.
Mephitis mephitis notata Hall, Carnegie Inst. Washington, Publ. 473:67, November 20,
1936.

Type. — Obtained at Trout Lake, south base of Mt. Adams, Klickitat County.
Washington, by P. Schmid on March 22, 1897; type in United States National
Museum.

Racial characters. — Similar to hudsonica but larger; stripes narrower, di-
verging anteriorly on neck or back of head; tail shorter, sometimes without
long, white hairs.

Measurements. — Howell (1901:37) gives the average of 3 adult males from
Trout Lake, Klickitat County, as: total length 633; length of tail 249; hind
foot 76.

Distribution. — The Columbia River Valley of the southern Cascades from
the Wind River east to the Snake River and the Yakima Valley area (trappers'
reports).

Mephitis mephitis spissigrada Bangs

Mephitis spissigrada Bangs. Proc. Biol. Soc. Washington, 12:31, March 24, 1898.
Mephitis foetulenta Elliot, Field Columb. Mus., Publ. 32, zool. ser., 1:269, March. 1899
(type from Lagune, near Port Angeles, Clallam County, Washington).

Chincha occidentalis spissigrada Howell, N. Amer. Fauna, 20:35, August 31, 1901.
Mephitis mephitis spissigrada Hall, Carnegie Inst. Publ. 473 :67, November 20, 1936.

Type. — Obtained at Sumas [prairie], British Columbia, by A. Brooks on
September 30, 1895; type in Museum of Comparative Zoology.

Racial characters. — Similar to hudsonica but larger; stripes broader, usually
diverging on shoulders; hairs of stripes often cream color or yellowish near
base; end of tail usually white.

Measurt mints. — An adult male from 2% miles southeast of Chinook, Pacific
County, measures: total length 578; length of tail 260; hind foot 30. A female
from the same place and one from Renton, King County, average: 575;
233; 72; 27.

Distribution. — Lowlands of western Washington. This form rarely goes far
into the mountains except along the valleys of the larger rivers. Marginal
records are Skykomish (W.W.D.) and Washougal River (W. W.D.).

220

University of Kansas Publs.. Mrs. Nat. Hist.

Taxidea taxus taxus (Schreber)
Badger

Ursus lnj-iis Schreber, Saugethiere, 3:520, 1778.
Taxidea taxus Rhoads. Amer. Nat., 28:524, June, l>!'l.

Taxidea taxus neglecta Taylor and Shaw, Occ. Pap. Clias. R. Conner Mus., no. 2:12, De-
cember, 1929.

Taxidea taxus taxus Hall, Carnegie Inst. Washington, Publ. 473:78, November 20, 193G.

Type. — None. Type locality probably southwest of Hudson Bay (Hall.
1936: 78).

Measurements. — Two males from northern Xevada measure, respectively:
total length 780, 762; length of tail 113, 150; hind foot 136, 120; ear 55, 54;
weight 15, 11 pounds.

Fig. 66. Badger (Taxidea taxus hi.ru*). tame animal in Seattle, Washington,
January 29, 1938. Captured at Lakeside, Chelan County, and photographed
al approximate age of tin months; subadull male. (Eloise Kuntz photo.)

Distribution. — Of general distribution over open country easl of the Cas-
cades. Marginal records are Wauconda (Taylor and Shaw, 1929: 12), Chelan
(Taylor and Shaw, 1929: 13). Goldendale - W N D.) and "Divide above
Trout Lake" ( t ra pper's photograph ) .

Description. — The badger is the size of a small dog, measuring up
to 32 inches in total length and weighing up to 20 pounds. The bodj

Badger

221

is heavy, powerful and remarkably flat and compressed. The tail
and legs are short. The forelegs are thick and strong, armed with
long heavy claws for digging. The ears are wide and low. The color
of the upper parts is a grizzled yellowish brown, not unlike the
color of the yellow-bellied marmot. The underparts are buffy, often
with a white area on the abdomen. The legs, feet, top of head, ears,
and small areas on the cheeks are blackish. Triangular areas about
the eyes are buffy. A white stripe extends from the nose pad back-
wards, between the eyes, to the shoulders and serves as the best
recognition mark.

DISTRIBUTION MAP
STATE OF WASHINGTON

P'iitt: SC79-5M7 =.;-" ir

Fig. 67. Distribution of the badger, Taxidea taxus taxus, in Washington.

Badgers are found over central and western United States,
Canada and northern Mexico. They are commonest on the plains
and desert, principally because the burrowing mammals upon which
they feed are most abundant there. The badger is a powerful and
rapid digger, being able to overtake and capture mice, ground
squirrels, and even pocket gophers. Perry (1939: 49-53) in her
interesting accounts of the habits of a pet badger obtained at Lake-
side, Chelan County, found the animal powerful enough to dig
through a concrete floor! Evidence of badger's activities are usually

222 University of Kansas Ptjbls., Mtjs. Nat. Hist.

seen at any ground squirrel colony in eastern Washington. This
evidence consists of large holes in the ground. Rarely a horizontal
tunnel begins at the depth of two to four feet and extends for an
unknown distance. Earth removed in excavating is heaped beside
one or both of the narrower sides of the surface opening.

In examining badger workings in ground squirrel colonies I have
been impressed by the fact that most of the holes ended not more
than four or five feet from the entrances — perhaps at the places
where the ground squirrel nests were located, although it may be
that the digging of the badger so terrified the squirrels that they
dashed out in an attempt to escape past the badger, before he
reached the nests. Kangaroo rats and pocket mice often attempt
to escape by dashing past a person when he is excavating their bur-
rows.

The power of the short, thick and slightly bowed foreleg of the
badger is tremendous. The claws are stout and fully an inch long.
The animal is able to break up and remove at a scoop, the baked,
clay like loess of the Columbian Plateau. Clods of this same material
are so firm that only by twisting and exerting considerable power
was I able to break them. The soil a foot beneath the sunbaked
crust is softer and more easily worked.

It is difficult to estimate the size of badger populations. In the
arid land of eastern Washington their digging- may exist almosl
unchanged for many years. Interviews with professional trappers
serve to indicate their range and numbers as follows: southeastern
edge of the Cascades and Yakima Valley, not common, rarely stray-
ing up into mountains — average trapper's catch, only one to three a
year; Okanogan Valley and northeastern edge of Cascades, not
common — average trapper's catch is six to ten a year, occa-
sional trappers catching a- many as 35; Columbia. Kettle River,
ami other valleys in northeastern Washington, uncommon, a few
records only; Columbian Plateau, fairly common — average profes-
sional trapper's yearly catch includes 10 to 20 badgers; southeastern
Washington, now rare because of overtrapping, formerly common.

Trappers state thai the badgers taken in northeastern Washington
are usually classified as "hair badger" by fur buyers and bring
only a dollar or two. The badgers of the eastern Cascades are "fur"
badgers but do not bring top prices. The badgers of the Columbian
Plateau bring the best prices. Seemingly some geographic variation
exists among badgers in Washington. Those from the more humid

Badger 223

areas of northeastern Washington and the eastern Cascades are
darker and bring poorer prices than the paler "silver" badgers of
the more open desert areas.

The principal food of the badger in Washington seems to be
ground squirrels, Citellus Washington/, townsencii, columbianus, or
saturatus, depending on locality. Pocket mice, gophers, and other
mammals are also eaten, as are grasshoppers, sword-tailed crickets,
other insects, and birds.

Young of badgers number 3 to 5 and are born in late April, May,
or early June.

224 University of Kansas Pibls.. Mrs. Nat. Hist.

Vulpes fulva cascadensis Merriam

Red fox

Vulpes cascadensis Merriam, Proc. Washington Acid. Sci., 2:665, December 28, 1900.
Vulpes fulvus cascadensis Bailey, N. Amer. Fauna, 55:281, August 29, 1936.

Type. — Obtained at Trout Lake, Klickitat County, Washington, by P.
Schmid on March 3. 1898; type in United States National Museum.
Racial characters. — Skull heavy, narrow; color yellowish.

Measurements. — A male from Crater Lake. Klamath County. Oregon, meas-
ures: total length 1113; length of tail 441; hind foot 180; ear 112; weight 9
pounds.

Distribution. — From Trout Lake northward, through the higher Cascades,
to Loomis (Taylor and Shaw, 1929:13).

Description. — The red fox of the Cascades is large and measures
about 4 feet in total length, of which the tail comprises 15 inches.
The body is slender and doglike; the legs long and slim; the tail
thick and bushy, and the ears are large and erect. In the red phase
the red fox of the Cascades is distinctly more yellowish than the
red fox of the eastern United States; the head is especially yellow.
The body has more red on the shoulder area than posteriorly, and
is darkest on the rump. The tail is rather pale with a dusky, not
black, area distally and a white tip. The ears are dusky. The lower
legs and feet are black. The throat, chest and underparts are white.
The "cross" phase, according to Cowan (1938: 202-206), is rather
common in the Cascades. In cross foxes the color is darker, brown
rather than yellowish, and the area from the nape of the neck back
between the shoulders, including a bar across the shoulders, is deep
blackish or grayish brown. In a pelt that is stretched out a cross is
formed by the dark areas. The black and silver phases of the red
fox are also said by Cowan to be relatively common in the Cascades,
constituting 48 per cent of the population. Of 3,163 foxes traded
at Fort Colville, in northeastern Washington, only 1!» per cent were
silver or cross. One fox, trapped in Okanogan County, is said by
it- captor to have been black above and straw colored beneath, with
no white on the body.

Red foxes range from Alaska and unit hern Canada well south-
ward into (he United State-. Related forms occur in Eurasia.

The red fox of Washington is an alpine animal, ranging at or
slightly below timber-line. Here food is abundant in summer and
fall but must be scarce in winter. In winter it- habitat is difficult
for man to reach and few persons penetrate the dangerous terrain
where the fox lives. A few professional trappers regularly catch

foxes in the Cascades but know little of their habits.

Fox

225

The feces of red foxes examined by Taylor and Shaw on Mt.
Rainier contained remains of insects and berries (Taylor and Shaw,
1927: 43).

The red fox is rare in Washington; it lives in inaccessible terri-
tory and its fur is not especially valuable. The animal is of rela-
tively little economic importance.

The brood den of a red fox found by Livezey and Evendan (1943:
500) near Corvallis, Oregon, was two-thirds of the way up a 300-foot
hill in a strip of oaks [Quercus garryana). Well-packed trails led

DISTRIBUTION MAP

STATE OF WASHINGTON

FtCd Under Aubi

i 6079-5737 buCMT r?37

Fig. 68. Distribution of the red fox in Washington. A. Vulpes julva cas-
cadensis. B. Vulpes julva macroura. (See p. 450.)

to an entrance concealed in poison oak (Rhus diversiloba) . Remains
of a turkey, 5 ground squirrels, and a jack rabbit were found near
the den. The entrance was 8 inches wide and 15 inches high. The
tunnel tapered to 5 inches in diameter and was 47 feet long. Seven
pups. 4 males and 3 females, were found in the den.

15—4993

226

University of Kansas Plbls., Mus. Nat. Hist.

Canis latrans Say

Coyote

Description. — The coyote is a large carnivore, about the size and
general proportions of a small collie dog but with bushier tail, and
more slender body. Adults measure about four feet in length, of
which the tail comprises a fourth. Adult males weigh about 30
pounds. The color is somewhat variable, yellowish, buffy. or
grayish. The muzzle and backs of the ears are more reddish, and
the legs brown. The tail is yellowish gray with a dusky tip. The
throat, chest and underparts are white.

Fie. 69. Coyote (Canis latrans Icslcs), in trap. 5 miles southeast of Mc-
Kenna, Washington, April 10, 1924. (Fish and Wildlife Service photo by G.
R. Bach, No. 26901.)

The coyote ranges from Alaska southward, over western North
America, to Central America.

At the present time the coyote ranges over almost all of Washing-
ton, except for the highest parts of the mountains and the dense
forest areas of western Washington. According to available in-

Coyote 227

formation it was not found about Puget Sound or the Olympic
Peninsula until relatively recent times. Some old trappers are of
the opinion that coyotes did not come into western Washington
until the timber wolves became rare there. The scarcity of the
timber wolves seems to coincide with settlement, clearing and
lumbering. Probably coyotes did not become common in western
Washington until lumbering provided extensive clearings and open
areas more suitable to them than forest. Certainly coyotes and
wolves existed together in eastern Washington.

It is difficult to determine whether coyotes were completely absent
from western Washington in the early days or simply scarce. The
glacial prairies of the Puget Sound area provide suitable habitat
for coyotes and coyotes are abundant there at the present time.
If coyotes were present at all in western Washington in the early
days it is reasonable to suppose that they would have been common
on the prairies. Yet I was told by an old trapper who had lived
near Scotts Prairie, Mason County, for many years, that he had
never seen or heard coyotes there until about 1910. Although a
coyote might be mistaken for a small wolf, the call, as this trapper
pointed out, is distinctive.

The principal habitat requirement of the coyote seems to be
extensive areas of open country. This it finds in the desert area
of the Columbian Plateau, the open forests of northeastern Wash-
ington and the eastern Cascades, and in the extensive timbered and
burned-over lands in western Washington. In summer coyotes
range well up into the Hudsonian Life-zone of the Cascade and
Olympic mountains. Their principal range is lower, in the Transi-
tion and Upper Sonoran life-zones.

The coyotes are both nocturnal and diurnal. In the vicinity of
human habitations they are most active at night. In the heat of
the day they take refuge in brushy areas or small gullies. Many
actions of the coyote are doglike, and were it not for the large,
round, bushy tail, a coyote might easily be mistaken for a dog.
The tail is carried in a drooped position with the tip bowed slightly
backwards. When badly frightened and running at full speed the
tail is stretched out straight behind. The ordinary gait is a pur-
poseful trot with the head held erect, the ears pricked up, and the
legs moving smoothly and effortlessly. Near Moses Lake, Grant
County, I watched a coyote trot along the side of one of a series
of small sand dunes. At the report of the small shotgun fired at it.
the coyote's dignified trot changed to terrified bounds, its feet dug
into the dune, throwing showers of sand into the air, as it crossed

228 University of Kansas Publs., Mus. Nat. Hist.

several successive dunes in full flight before turning to take ad-
vantage of the shelter of a draw between two dunes. A coyote
chased by an automobile attained a speed of 43 miles an hour for
a short distance (Zimmerman, 1943: 400).

I have not watched a coyote stalk game, but as described to me
it creeps up to within a few yards of its prey and catches it with
a sudden dash. In hunting a jack rabbit, the coyote is said to pur-
sue it, taking advantage of short cuts, until close enough to seize it.

Better known than the coyote itself is its howl — several doglike
barks, each successive one of a series shorter and higher pitched, the
last one ending in a long, drawn-out howl. In the winter of 1936
the coyotes near Cottage Lake, King County, were especially vo-
ciferous. They ordinarily began to howl about 9 p. m., but could
be induced to howl earlier by imitating their call. Their howling
was a signal for all nearby ranchers' dogs to howl in reply. In the
desert areas of eastern Washington I heard coyotes most often just
at dusk or at dawn.

The coyote is principally carnivorous, feeding on any mammals
and birds easy to kill. It willingly eats carrion, even when much
decayed. Large insects such as grasshoppers and crickets are eaten
when they are abundant and easily caught. Fruit and berries are
eaten when available.

Sperry (1941) reported on the analysis of the contents of 8,339
coyote stomachs from various places in the United States, 1,186 of
the stomachs being from Washington. The following information
is from his report on all of the 8,339 stomachs: rabbits formed
one-third of the food; Microtus was found in 7 per cent of the
stomachs; Peromyscus in 6 per cent; Neotoma in 4 per cent. Rei-
throdontomys were found in 53 of the 8,339 stomachs and in insig-
nificant numbers. Onychomys occurred in 11 of the 8,339 stomachs;
Clethrionomys in 8. Ondatra occurred in 8 of the 8,339. Two
muskrats were in stomachs obtained from Washington. Citellus
(exclusive of beecheyi, lateralis, and saturatus) were found in 4
per cent and were locally important ; Citellus beecheyi occurred in
84 stomachs, including 1 from Washington; Citellus lateralis and
saturatus occurred in 50 stomachs, including 5 from Washington.
Marmota were found in 1 per cent of the stomachs. They were
included in stomachs from Washington but the number was not
reported. Tarnias (=Euta?nias) were found in 43 stomachs from
western United States and were present in 1 per cent of the 1.186
stomachs from Washington. Sciurus and Tamiasciunts occurred in

Coyote 229

33 stomachs. Included were 3 Tandasciurus douglasii, a Tamias-
ciurus hudsonicus, and a Sciurus griseus from Washington. Glau-
comys occurred in 6 Washington-taken coyotes, of a total of 11
from the entire United States. Perognathus occurred in 3 per cent of
the stomachs, and 274 individuals were represented. They were
found in 10 per cent of the 1,186 Washington stomachs. Thomomys
occurred in 4 per cent of the stomachs examined and in 7 per cent
of the stomachs from Washington. Erethizon appeared in 2 per cent
of the stomachs (135 records), including some from Washington.
Aplodonta occurred in only 11 stomachs, all taken in Washington.
Locally, it is concluded, mountain beavers are important coyote
food. House mice occurred in but five stomachs, including 2 from
Washington. Zapus did not occur in coyote stomachs from Wash-
ington. Domestic sheep and goats formed 7 per cent of the food of
Washington coyotes. Calves occurred in 3 stomachs of coyotes
from Washington. Pigs occurred in 8. Deer formed 3 per cent, by
volume, of coyote food in Washington. A part of the stock and
deer reported was doubtless carrion.

One shrew was found in the stomach of a coyote from Washing-
ton, and two stomachs contained moles. A house cat was eaten by
one Washington coyote, and another coyote had eaten a raccoon.
Birds occurred in 13 per cent of the stomachs examined but formed
only a small part by volume. Poultry formed one-fourth of this
volume. Reptiles formed 0.08 per cent of the food eaten by coyotes
and occurred in 3 per cent of the stomachs. A coyote from Wash-
ington had eaten a garter snake. No frogs were found in coyote
stomachs. A coyote from Washington had eaten a salamander. An-
other had eaten a fish. Insects formed 1.08 per cent of the total
food eaten by coyotes. Fruit formed 3.63 per cent. Carrion con-
stituted 25.2 per cent of the total food eaten.

It is extremely difficult to draw conclusions regarding the economic
value of any species. Much depends on the outlook of the indi-
vidual, his occupation, the locality where the animal occurs and
local conditions there. Furthermore it is impossible to understand
and fairly weigh all the factors involved.

Coyotes destroy game and stock. They also eat carrion and de-
stroy sick animals, thus preventing the spread of disease. They
eat jack rabbits which are pests, and snowshoes and cottontails
which are game. They eat mice, which are a pest in agricultural
areas, a benefit on rangelands where they destroy weed seeds, and
of neutral importance elsewhere. To analyze the economic value

230

University of Kansas Publs., Mus. Nat. Hist.

of the coyote, the economic importance of all animals on which it
feeds must be considered and the "good" and "bad" balanced — a
well-nigh impossible task.

In any event, the coyote has been judged and found guilty.
Coyotes are controlled by poison, trapping, and bounty. In spite of
control measures the coyote is holding its own in numbers or in-
creasing.

Stimulated in part by the bounty, professional trappers take many
coyotes each year in Washington. The pelts fluctuate in value from
year to year but a large, prime skin usually brings from 5 to 10
dollars.

DISTRIBUTION MAP
STATE OF WASHINGTON

25 50 75 100

D.o'-fd Und>> A*

I

Fig. 70. Distribution of the coyote in Washington. A. Canis latrans testes.

B. Canis latrans incolatus.

Canis latrans lestes Merriam

Canis testes Merrian i. Proc. Biol. Soc. Washington, Ll:25, March 16, 1807.

Canis latrans lestes Tayloi and Shaw, Mamm. and Birds Mi. Rainier Nat. Park, p. 41. 1927.

Type. — Obtained in the Toyabe Mountains near Oloverdale, Nye County,
Nevada, by V. Bailey on November 21, 1890; type in United States National
Museum.

Racial characters. — Frontal region of skull only slightly concave.

Coyote 231

Measun nu nts. — A male from 20 miles south of Ephrata, Grant County, on
the Columbian Plateau, measures: total length 1185; length of tail 365;
hind foot 198; ear 125; weight 26% lbs. A female from 10 miles northeast
of Goldendale, Klickitat County, in the southern Cascades, measured: 1105;
280; 197; weight 19 lbs. A female from Cashmere, Chelan County, in the
northern Cascades, measured: 1209; 410; 210; 131. A female from the middle
fork of the Nooksack River, Whatcom County, in western Washington,
measured: total length 1185; length of tail 358.

Distribution. — Suitable areas of the entire state, except for northeastern
Wa^-hington.

Remarks. — Pending a revision of the coyotes, those of western Washington
are referred to lestes.

Canis latrans incolatus Hall

Cauis latrans incolatus Hall, Univ. California Publ. Zool., 40:369, November 5, 1934.

Type. — Obtained at Isaacs Lake, 3,000 ft. elevation, Bowron Lakes Region,
British Columbia, by T. T. and E. B. McCabe on October 23, 1928; type in
Museum of Vertebrate Zoology.

Racial characters. — Similar to lestes but frontal region more concave.

Measurements. — A male from 20 miles east of Tonasket, Okanogan
County, measures: total length 1033; length of tail 291; hind foot 163;
ear 110.

Distribution. — Northeastern Washington, west to Okanogan and Con-
conully (W.W.D.).

Rt marks. — Several series of coyote skulls from northeastern Washington
are more variable than series of lestes from California and incolatus from
British Columbia. The "dish-faced" character of incolatus is more strongly
represented in some coyotes from northeastern Washington than in topotypes
of incolatus but others are more like lestes. In average measurements they
are nearer incolatus.

232

University of Kansas Publs., Mus. Nat. Hist.

Canis lupus fuscus Richardson

Timber Wolf

Cants lupus var. fusca Richardson, Mammalia, Zoology, Captain Beechey's voyage of the
Blossom, p. 5, 1839.

Lupus gigas Townsend, Jour. Acad. Sci. Philadelphia, 2:75, November, 1850 (type from
near Vancouver, Clark County, Washington).

Canis gigas Miller, Smithsonian Misc. Coll., 59 (no. 15): 4, June 8, 1912.

Canis occidcntalis gigas Taylor and Shaw, Birds and Mamm. Mt. Rainier Nat. Park, p. 39,
1927.

Canis lycaon gigas Bailey, N. Amer. Fauna, 55:272, August 29, 1936.

Type locality. — Banks of the Columbia below The Dalles in Oregon or
Washington.

Racial characters. — A large, relatively dark colored wolf with wide skull
and heavy dentition.

Distribution. — Probably occurred from the eastern base of the Cascades
westward to the Pacific. Now extinct over most of its range.

DISTRIBUTION MAP
STATE OF WASHINGTON

MILES

ol »!>« O.,

M'l UV i, IU Ml

Fio. 71. Probable past distribution of the wolf in Washington. A. Canis
lupus fuscus. B. Canis lupus Columbian us. C. Canis lupus irremotus.

Description. — The wolf is a large carnivore of the general pro-
portions of a large collie dog. Wolves closely resemble coyotes but
are larger, with stouter body, larger feet, thicker muzzle and mere
massive, powerful skull and teeth. The fur of wolves is long and

Wolf 233

rather stiff. In general color of the body they resemble coyotes,
but the underparts are less white and the legs and feet are more
contrastingly reddish.

Wolves of the species Cards lupus range over the northern parts
of both the Old and New World. In America they are found from
the Arctic south into Mexico.

Wolves occurred in western, northeastern and southeastern Wash-
ington. They seem not to have occurred on the Columbian Plateau.
Wolves are generally associated with the larger, hooved mammals
upon which they feed. In Washington these probably once included
the elk, deer, mountain sheep and mountain goat. Hooved animals,
except mountain sheep and deer, may not have occurred on the Co-
lumbian Plateau in historic times, and even the deer and sheep prob-
ably were scarce. There was probably little food for wolves on the
Plateau.

The early settlers found wolves to be common and a serious pest.
By 1900 they had nearly disappeared. In the settled parts of west-
ern Washington they were doubtless exterminated at an early date
but it is difficult to account for their disappearance on the Olympic
Peninsula. To the best of my knowledge, two wolves killed on
the north fork of the Quinault River in 1920, or a few years earlier,
were the last ones from that peninsula. Previously they were com-
mon and I doubt that man killed them all ; perhaps some introduced
disease, such as rabies, brought about their extinction.

In the Cascade area they probably still exist in small numbers
and in remote places. There are said to be some near Mount Adams.
They have been reported from Mount Rainier as recently as 1920.
Trappers state that there are a few in the northern Cascades, be-
tween Lake Chelan and Mount Baker. There are no recent records
for southeastern Washington. The last wolves killed on the Colum-
bian Plateau were two taken at Wahluke, Grant County, on Sep-
tember 17, 1917. Only rumors — no authentic reports — of wolves
are available from northeastern Washington in recent years. The
reappearance of the caribou there may attract wolves from British
Columbia.

Any report of wolves, even from experienced trappers, is open to
some question. An unusually large coyote often is mistaken for a
wolf. A large, wild and wolflike dog is even more likely to be mis-
taken for a wolf.

The wolf no longer is an important element in the mammalian
fauna of Washington and will probably never be so again. The spe-
cies, like the grizzly bear, is nearly extinct in the state.

234

University of Kansas Publs., Mrs. Nat. Hist.

Canis familiaris Linnaeus
Dog

Canis familiaris Linnaeus, Syst. Nat., ed. 12, 1 : 56, 1766.

Type locality. — Sweden.

Dogs were present with aboriginal man in Washington, previous
to the arrival of the white man. For greater detail see Bailey
(1936), Suckley and Gibbs (1860) and Allen (1920).

Felis concolor Linnaeus

Cougar or mountain lion

Description. — The cougar is a large cat and has the general pro-
portions of the house cat. Large cougars measure more than seven
feet in length of which the tail comprises two feet. Large males

v , •";

'■■*■

■i.\

fc*

Fig. 72. Cougar or mountain lion {Felis concolor missoulensis) , skin
mounted as n rut:; shol on Mill Creek. Pend Oreille County, Washington,
February 13. 1935, by Ralph Johnson. Skin now measures: snoul to tip of
tail 252 cm., span between tips of forepaws l.">7 em., tail Si) cm.; male. (Fish
ami Wildlife Service photo by Victor B. Scliel'ler. No. 66.)

Cougar

•j:::»

weigh more than 150 pounds. Females are smaller and lighter than
males. In color the head, back, tail and outside of the legs are
reddish brown. The throat, underparts and inside of the legs are
white. The tip of the tail, or area near the tip, is black or blackish.

Cougars, often called mountain lions, pumas and panthers, range
from Canada to southern South America.

The cougar feeds extensively upon deer and its habits and habitat
are accordingly specialized. It is active in both winter and summer
and zonally ranges from the Transition through the Canadian to the
Hudsonian life-zones.

Fig. 73. Cougar or mountain lion (Felis concolor oregonensis) , adults and
young taken by Dewey Schmid in White Salmon Valley, Washington, about
December 1, 1937, and January 1, 1938. Two bobcats show at extreme right.
(Dewey Schmid photo.)

The number of deer killed by the average cougar is unknown.
Some trappers believe that a cougar kills a deer at least every other
day while others feel that only one a week is taken. Other food
is eaten when available. Because cougars kill deer, they are in-
cessantly warred upon by hunters and sportsmen. Bounties, often
generous, have long been paid in Washington. Although many
cougars are killed each year they are still numerous in many areas.
So many remote areas in the mountains are not easily accessible
to man and dogs that the cougar, as a species, probably will persist
for many years.

A cougar is one of the most secretive animals in the wilds.
Rarely are individuals seen by man. They are difficult to trap,
principally because they are such wide-ranging animals and partly
because they prefer to feed on fresh deer meat and hence are not

236

University of Kansas Publs., Mtjs. Nat. Hist.

apt to be attracted to trap bait. For these same reasons they are
difficult to poison. By using poisoned hamburger, a government
agent did poison one near Leavenworth, Chelan County. The
principal method of killing cougar is to hunt them with packs of
especially trained hounds which pursue the cougar until it takes
refuge in a tree or other supposed place of safety, where it is shot.
A common gait is a swift, smooth trot in which the body is kept
low and the tail droops with the terminal part bent backwards.
A wild cougar seen near Leavenworth, Chelan County, traveled
with effortless speed until fired upon. At the sound of the shot
it made two great bounds and disappeared into the brush fringing
a canvon.

DISTRIBUTION MAP
STATE OF WASHINGTON

f.of'.d Ur.d.» A U ,pt

_J

aCa P..

1_

*c-«-s^» b-CMT mr

Fig. 74. Distribution of the cougar in Washington. A. Felis concolor ore-
gonensis. B. Felis concolor tnissoulcusis.

Although the cougar is a large and powerful carnivore it almosl
never makes unprovoked attacks upon man. An exception was
the cougar that partially devoured a thirteen-year old boy near
Lake Chelan, Chelan Counly (Finley, 1925: 197-199).

The hunting range of an individual cougar is many miles in
extent. These ranges are traveled periodically and any particular
area may be visited regularly every few days. The cougars may

Cougar 237

travel many miles each night in search of deer. Their ranges
must overlap to a certain extent for as many as 12 have been taken
from a single drainage area. Over most of the year they are solitary
but breeding females may attract several males and hunters occa-
sionally capture a female and several males at one locality. Breed-
ing occurs in almost any month of the year. The young are cared
for by the female and follow her for perhaps a year. Young indi-
viduals have been taken at the same time and in the same locality
as an adult female and several adult males that supposedly were
breeding. There are from one to three young in a litter. For the
first months of their life cougar kittens are spotted. In this they
differ from the young of the bobcat which are plain or slightly
mottled and do not become spotted until later in life. Cougar
kittens differ further from bobcat kittens in possessing a long tail.
The pelage of the cougar is short and of no value as fur although
the skins often are sold at a good price as trophies when prepared

as rugs.

Felis concolor oregonensis Rafinesque

Felix [sic] oregonensis Rafinesque, Atlantic Journal, 1:62, 1832.

Felis hippolestes olympus Merriam, Proc. Biol. Soc. Washington, 11:220, July 15, 1897
(type from Lake Cushman, Mason County, Washington).

Felis oregojiensis Stone, Science, n. s., 9:35, January 6, 1899.

Felis oregonensis oregonensis Miller, U. S. Nat. Mus. Bull., 128:158, April 29, 1924.

Felis concolor oregonensis Nelson and Goldman, Jour. Mamm., 10:347, November 11, 1929.

Type locality. — "Oregon Mountains, or east or west of them."

Racial characters. — Light, rounded skull, dark color, extensive black on tip
of tail and short, rather coarse fur.

Distribution. — From the eastern base of the Cascade Mountains westward
to the Pacific. Marginal records are (trappers' specimens) : Ruby, Leaven-
worth, and Goldendale.

Felis concolor missoulensis Goldman
Felis concolor missoulensis Goldman, Jour. Mamm., 24:229, June 8, 1943.

Type. — Obtained 10 miles southwest of Missoula, Missoula County, Mon-
tana, by R. and C. Thompson, on December 30, 1936; type in United States
National Museum.

Racial characters. — A large cougar with heavy, wide skull, pale color and
rather long, soft fur.

Distribution. — Northeastern Washington west at least to Republic (W.W.D.)
and the Blue Mountains of southeastern Washington.

Remarks. — Goldman (1943: 229) states "Between the Rocky Mountains and
the Cascade Range missoulensis intergrades with oregonensis." My own com-
parison of specimens reveals differences of considerable magnitude between the
cougars of western and northeastern Washington. I suppose that intergra-
dation takes place in a limited area in, and west of, the Okanogan River Valley
in Washington.

238 University of Kansas Publs., Mus. Nat. Hist.

Fig. 75. Canadian lynx, Lynx canadensis, male, taken February 16, 1939,
on Ha My Mountain, northwest Idaho, by Lloyd Robinson of Sandpoint.
(Ross A. Hall photo )

Lynx

239

Lynx canadensis canadensis Kerr
Canadian lynx

Lynx canadensis Kerr, Anim. Kingd., 1: systematic catalogue inserted between pages 82 and
33 (description, p. 157), 1792.

Lynx borealis canadensis True, Proc. U. S. Nat. Mus., 7:611, 1885.

Lynx canadensis canadensis Miller, U. S. Nat. Mus. Bull., 128:160, April 29, 1924.

Type locality. — Eastern Canada.

Measurements. — A female from Buttermills Creek, Twisp River, Okanogan
County, measured: total length 900; length of tail 95; hind foot 205.

Distribution. — Higher parts of the Cascade Mountains, Blue Mountains and
mountains of northeastern Washington, recorded from Oroville on the north
(W.W.D.) to Mount Adams (Taylor and Shaw, 1929: 13) on the south.

Fig. 76. Canadian lynx (Lynx canadensis) , catch of nine, with two coyotes,
taken by Lester Fairbrother in hills west of Oroville. Washington, March.
1938. (Lester Fairbrother photo.)

Description. — The Canadian lynx weighs about 20 pounds and is
catlike in general proportions but differs in possessing longer, stouter
legs, much larger feet, a short tail, tufted ears and long, very soft
fur. The pelage of the upper parts is soft gray with a slightly yel-
lowish tone ; the ears and tail are black ; and the underparts are gray
with indistinct black spots. The pelage is shorter and more reddish
in summer. Bailey (1936: 271) records a maximum weight of 28
pounds.

240

University of Kansas Publs., Mrs. Nat. Hist.

The lynx ranges over the forested parts of North America from
the Arctic south into the northern United States. It has a restricted
range in Washington, occurring in the same areas as does the red
fox. Although the lynx is an important fur bearer in Canada and
Alaska, it is unimportant in Washington because only a few are
trapped each winter. Most of the natural range is in the remote
and wilder parts of the mountains. Here, each of several trappers

DISTRIBUTION MAP

STATE OF WASH I NGTON

C-d'ied U-de. A u

MILES
B < .*=* p..

6C'V1*}7 •-^" T »7

Fig. 77. Distribution of the Canadian lynx, Lynx canadi nsis, in Washington.

regularly takes a dozen or more each year. Mr. Lester Fairbrother
of Oroville, Okanogan County, regularly traps lynxes in the north-
ern Cascades. They are taken in wooded areas where snowshoe rab-
bits, their principal food in winter, arc abundant. In the more ac-
cessible parts of the animal's range, such as the Blue Mountains
and the mountains of northeastern Washington, lynxes are rare. As
much as sixty dollars each is offered for large skin-.

Bobcat

241

Lynx ruf us (Schreber)

Bobcat

Description. — The average male bobcat weighs approximately 20
pounds. The female is about one-fourth lighter. A bobcat has
longer, stouter legs and larger feet than a house cat and a short tail.
The ears are short, with pointed tufts of hair.

Fig. 78. Bobcat (Lynx rufus jasciatus) , two-year-old male captured as a
kit near Lyman, Washington, in the spring of 1937, by Earl Scott ; photo-
graphed March 9. 1939. (Fish and Wildlife Service photo bv Victor B. Schef-
fer, No. 588.)

Bobcats range frcm southern Canada south to central Mexico.
Whereas the Canadian lynx occupies the boreal region, the bobcat
occupies the temperate region. It is thought to be principally noc-
turnal but is occasionally active by day. On? that I watched near
Lake Samamish, King County, when it was unaware of my presence,
16—4993

242 University of Kansas Publs., Mus. Nat. Hist.

was decidedly uncatlike as it strolled with a smooth but stiff-legged
gait on a forest trail, with head held up, short tail erect and wagging
back and forth with each step. The general impression was of a
large, extremely long-legged animal. There was nothing stealthy
in its movements. Another individual seen in the same locality on
a later date saw me. As it bounded away the body was kept low
and the legs were bent with the forelegs appearing almost bow-
legged.

Like many other carnivores, each bobcat has a home range
which varies with the available food supply. The range may include
deep forest, dense thickets and open grasslands, but country with
considerable edge-environment seems to be preferred to dense cover,
and rocky areas to smooth soil. Perhaps the abundance of mice
and wood rats attracts bobcats to the rocks, but the cover afforded is
also a factor.

The bobcat is almost universally regarded as a predator and the
state of Washington now offers a bounty on it. The animals doubt-
less do kill a certain amount of game, but kill also other animals
which man regards as pests because they interfere with reforestation
and growing of food crops. One of the best natural checks on these
pests is the bobcat which, in certain areas, does more good than
harm. Where doing damage to game or livestock bobcats may be
eliminated by trapping or hunting with dogs.

Trappers report that bobcats have a poor sense of smell but
very keen sight. They are easily trapped. The fur is soft and
handsome but does not wear well. Fur buyers designate the large,
pale bobcats of eastern Washington as lynx cats and reserve the
name bobcat for the more reddish race of western Washington. Im-
mature and unprime pelts from eastern Washington also are classed
as "bobcat."

The difference between the pale bobcat of eastern Washington and
the dark race of western Washington is greater than that which
separates many subspecies. Dewey Smith of Guler, Klickitat
County, showed me skins of bobcats taken on his trap-line along
the White Salmon River, which drains southward to the Columbia,
and over into the watershed of the Lewis River. Bobcats from the
White Salmon River were pale and gray, and those from along the
Lewis River were more reddish. The difference was striking. A
very few intermediate skins indicated that interbreeding occurs.
The geographic variation between the two races is reminiscent of
that in the snowshoe rabbits of the western Cascades.

Bobcat

243

Lynx rufus fasciatus Rafinesque

Lynx fasciatus Rafinesque, Amer. Monthly Mag., 2:40, November, 1817.
Lynx fasciatus fasciatus Miller, U. S. Nat. Mus. Bull., 128:160, April 29, 1924.
Lynx rufus fasciatus Bailey, N. Amer. Fauna, 55:269, August 29, 1936.

Type locality. — Near mouth of the Columbia River on "Netul" River (Lewis
and Clark River) near Astoria (Bailey, 1936:269).

Racial characters. — Size moderate; fur short; color distinctly reddish.

Measurements.— A male from Forks, Clallam County, measured : total length
890; length of tail 190; hind foot 167; ear 80; weight 24 l / 2 pounds.

Distribution. — From the Cascade Mountains westward, including the valleys
of rivers draining westward in the Cascades. Marginal occurrences are: Sky-
komish (W.W.D.). Mt. Rainier (Taylor and Shaw, 1927: 60) and head-
waters of Lewis River (W.W.D.).

DISTRIBUTION MAP
STATE OF WASHINGTON

Fig. 79. Distribution of the bobcat in Washington. A. Lynx rufus fasciatus.

B. Lynx rufus pallescens.

Lynx rufus pallescens Merriam

Lynx fasciatus pallescens Merriam, N. Amer. Fauna, 16:104, October 28, 1899.
Lynx rufus uinta Bailey, N. Amer. Fauna, 55 : 267, August 29, 1936.
Lynx rufus pallescens Bailey, N. Amer. Fauna, 55 : 268, August 29, 1936.

Type. — Obtained at Trout Lake. Klickitat County, Washington, by D. Kaegi
on January 10, 1895; type in United States National Museum.

Racial characters. — Size large; skull heavy with extensive crests and ridges;
fur long and soft; color pale, more grayish and less reddish than in fasciatus.

Distribution. — From the Cascade Mountains eastward, including the valleys
of rivers in the Cascades which drain southward and eastward. Marginal
records of occurrences are Oroville (W.W.D.), Lake Keechelus (Taylor and
Shaw, 1929:15) and Trout Lake (W.W.D.).

244 University of Kansas Plbls., Mus. Nat. Hist.

Zalophus californianus (Lesson)
California sea lion

Otaria californiana Lesson, Dirt. Class Xat. Hist.. 13:420, 1828.

Zalophus californianus Allen. Monogr. X. Amer. Pinnipeds, U. S. Geol. and Geogr. Surv.
Terr., Misc. Publ., 12:270, 1880.

Type locality. — California.

Distribution. — Rare or casual along the coast of Washington.

Description. — Adult males are 7 to 8 feet in length and females
about 6 feet. Bulls weigh as much as 1,000 pounds, and females up
to 600 pounds. Both fore and hind limbs are modified for swim-
ming; they are flippers directed posteriorly. The body is cylindrical
and streamlined, the neck thick and the head small. Adult males
possess a high sagittal crest resulting in a high forehead. Eyes and
ears are small. The reddish brown pelage is short and coarse. Fe-
males are darker than the males.

The California sea lion occurs along the Pacific Coast from Mex-
ico to northern California and has been recorded from Oregon
(Bailey. 1936: 332), Washington and British Columbia (Green-
wood, Newcombe, and Fraser, 1918: 1-39). It is of rare or casual
occurrence along the coast of Washington.

Eumetopias jubata (Schreber)

Steller sea lion

Phoca jubata Schreber, Siiugthiere, 3 r 300, 1771;.

Eumetopias Stelleri True, Proc. U. S. Nat. Mus., 7 (1884) : 007, 1885.

Eumetopias jubata Allen, Hull. Amer. Mus. Xat. Hist., 10:118, March 15, l!">-_>.

Type locality. — North Pacific Ocean

Distribution. — Ocean coast, breeding on small, rocky islet> such as the Quil-
layute Needles.

Description. — The Steller sea lion resembles the California sea
lion but is larger. Old hulls are from 10 to 12 feet in length and
weigh from 1,200 to 1. 500 pounds, perhaps even a ton. Females are
8 to 9 feet in length and may weigh up to 1000 pounds. The female
Steller sea lion is nearly as large as the male of the California sea
lion. The bull Steller sea lion lacks the high forehead characteristic
of the California sea lion, and the body is heavier and stouter, es-
pecially in the neck and chest. Bulls are distinctly bicolored, the
head, neck and chest being cinnamon and the rest of the body
darker, females are a uniform dark brown color. The Steller sea
lion makes a loud, dee]) roaring sound.

Steller sea lions occur alone, the coast of Asia south to Japan
and the coast of North America from the Bering Strait south to
central California. The range thus meets and overlaps that of the
Cab t'oi nia sea lion.

Steller sea lions are splendid swimmers and spend much of their
time in the surf. They often lie in the water a few yards from

Sea Lions and Seals 245

where the waves pound some jagged cliff, retaining their dangerous
position without discernible effort. They are curious and will raise
their head and neck vertically out of the water to observe a passing
boat. Much time is spent beneath the surface, presumably hunting
fish. They sometimes leap up out of the water in an arching dive
or may simply lower their heads beneath the surface and appear
several minutes later at another place. They may come to the sur-
face with a porpoiselike roll, breathe, and again dive. Many of their
actions seem inspired by playfulness — a means of working off ex-
cess energy.

Sea lions haul out on a number of rocky islands along the coast.
They are slow and clumsy on land. Their rear flippers can be
directed forward and so give them some assistance in moving about.
When startled they are able, with much struggling, to move with
fair speed to the water. They can dive from considerable heights.

The studies of Greenwood, Newcombe and Fraser (1918: 1-39)
show that the Steller sea lions eat a great variety of marine fish
and other sea life. Included in their diet are squid, starfish, crabs,
clams, mussels, salmon, herring, flounder, rock cod, sea bass and
dogfish. They seem not to be selective in their food habits but eat
the food most easily available at the time. Unfortunately this is
occasionally salmon in nets or traps and for this reason fishermen
usually kill sea lions on sight. The number of sea lions along the
coast of Washington has been greatly reduced by shooting and dy-
namiting the animals on their hauling-out grounds. There is no
doubt that sea lions do occasionally eat salmon, especially in traps
or nets. They also become entangled in the nets, and damage them.
On the other hand, investigations of their food habits have
shown that they do not eat any great number of salmon or other
important food fishes. They are a relatively harmless and exceed-
ingly interesting animal and might well be preserved in numbers
exceeding their present population.

Until relatively recent times the sea lion was an important source
of food to the Indians living along the ocean coast. The bones of
sea lions are often the most numerous vertebrate remains in shell
mounds.

The Steller sea lion now breeds only in a few places along the
coast of Washington. They are polygamous and each of the stronger
bulls has eight or ten cows in his harem. Fierce battles between
bulls are said to take place in the breeding season, late in May. A
single pup is born after a gestation period of about one year. Pups
are darker in color than adults.

246 University of Kansas Publs., Mus. Xat. Hist.

Callorhinus ursinus cynocephalus (Walbaum)

Alaska fur seal

Siren cynocephalus Walbaum, Petri Artedi Sueci Genera Pise. p. 560, 1702.

Callorhinus vrsinus True, Proc. U. S. Nat. Mus., 7 (1884) :607, 1885.

Callorhinus alascanus Jordan and Clark, Fur Seals and Fur Seal Islands of the North Pacific,
pt. 3, p. 2. November, 1899.

Calliiinriii ursina cynocephala Stejneger, George Wilhelm Steller, Harvard Univ. Press, p. 285,
1936.

Type locality. — Pribilof Islands. Alaska.

Distribution. — Breeds on the Pribilof Islands, Alaska. Abundant off the
coast of Washington on migration.

Description. — The difference in the size of male and female fur
seals is great. Males are about 8 feet in length when fully grown but
the females are only about 4 feet long. Males weigh up to 700
pounds but females only about 100 pounds. In general appearance
the fur seal resembles the sea lion but the fur is longer, denser and
softer. Males are very dark brown in color. Females and young
are grayish brown. The fur seal, like the sea lion, can reverse its hind
nippers and use them to a certain extent in walking.

The Alaskan fur seal breeds on the Pribilof Islands, Alaska. The
seals emerge from the water to certain favored liauling-out places in
May and June. They leave the Islands in November to begin their
long migration. The bulls spend the winter off the coast of Alaska
south of the Aleutians but the females and young travel south to
the ocean off California. In the spring they start north, arriving off
the coast of Washington in February. March and April. Young,
rarely adults, are washed up on the ocean beaches of Washington
(Scheffer, 1939: 43).

The United States Government, by international treaty, manages
the fur seal herds on a sustained yield basis and pelagic scaling by
independent hunters is prohibited. Indians are allowed to take the
fur seals on migration but are subject to certain restrictions. Modern
boats, outboard motors and guns are prohibited. Indians living on
the coast of Washington hunt the fur seals from dugoul canoes.
The weapons arc double-headed harpoons with long cedar shaft-.
One harpoon head is at the tip of the shaft and the other is on a short
fork that projects downward and forward at a 30° angle from the
main shaft. The harpoon is thrown with the aid of hand grips at
the end of the shaft. The seal is recovered by rawhide lines con-
nected to the harpoon heads. The hunters leave shore at dawn and
travel ten to twenty miles from land, at which distance the seal
herds are usually encountered. The hunting is dangerous work and
is carried on only by skilled and brave men. Relatively calm

Sea Lions and Seals 247

weather is required. The flesh of the seals is eaten and the skins
sold.

Schultz and Rafn (1936: 13-15) examined the stomachs of 41
fur seals taken within 30 miles of La Push, Washington, in March,
April and May, 1930. Food found included squids, shrimps, herring
and lampreys.

Phoca vitulina richardii (Gray)

Hair seal or harbor seal

Halicyon richardii Gray, Proc. Zool. Soc. London, p. 28, 1 S ' i 4 .
Phoca richardii Allen, Bull. Amer. Mus. Nat. Hist., 16:491, December 12, 1902.
Phoca richardii richardii Miller, U. S. Nat. Mus. Bull., 128 : 164, April 29, 1924.
Phoca vitulina richardii Doutt, Ann. Carnegie Mus., 29:117, May 12, 1942.

Type. — Described from a specimen obtained at the Fraser River, British
Columbia, probably on March 23, 1861, by C. B. Wood and another obtained
by Wood at Queen Charlotte Sound, British Columbia, in 1862 (see Scheffer
and Slipp, 1944: 374); type in British Museum of Natural History.

Distribution. — Puget Sound, the Strait of Juan De Fuca and the coast
of Washington, extending at times up the Columbia River to The Dalles.
Has been recorded in Lake Washington, Seattle (Bonham. 1942: 76).

Description. — Adults are about 5 feet in length and weigh up to
250 pounds; males are approximately a quarter larger than females.
The hair seals differ from the sea lions and fur seals in a number of
respects. The body is widest in the midsection rather than in the
chest. The neck is short and slim. The eyes are large and there
are no external ears. The hind flippers are not reversible but are
permanently directed posteriorly. The pelage is short, stiff, and
directed posteriorly. The ground color is silver gray or yellow;
usually it is yellow, blotched and marbled with black, dusky or
gray.

Hair seals range over the northern Atlantic and Pacific oceans,
occurring southward along the Pacific Coast of North America to
central Mexico. Six races are recognized by Doutt (1942: 115).

The hair seal is the common seal in Puget Sound, the Strait of
Juan De Fuca and on the ocean coast. Scheffer and Slipp (1944:
373) estimate that 5,000 live along the coast of Washington. Hair
seals are social to some degree but are often seen singly. In
Puget Sound, where there are relatively few hauling-out areas,
they rarely leave the water but on the ocean coast they emerge to
bask on the reefs. On Destruction Island, Jefferson County, V. B.
Scheffer and I watched a herd of about 20 animals that lay on a
reef exposed by low tide. According to the lighthouse keeper the
reef was occupied daily by these seals. Scheffer and Slipp (loc. cit.,
p. 388) report herds of up to 200 hair seals. In Puget Sound 10 to

248 University of Kansas Publs., Mus. Nat. Hist.

20 seals seem to be the usual number in a herd, but occasionally
there are as many as 50.

Hair seals often exhibit curiosity concerning small boats; the
seals lie in the water with only their round heads above the surface,
staring at the boat for many minutes. Constant shooting has made
them shy and any quick movement will cause them to dive; several
dived instantly when I pointed either a stick or gun at them.

Hair seals are less spectacular than sea lions. When basking
on rocks the seals are silent. When an observer is yet a long dis-
tance away the seals raise their small heads and remain on the
alert, When they take to the water, they travel with a jerking
motion. In the water one rarely sees more than their heads. When
they dive they usually sink below the surface, never making the
spectacular arching dive so typical of the sea lions.

At Useless Bay, Whidby Island, Island County, a herd of five
or six hair seals was studied in July, 1936. These animals played
and slept in the shallow water beneath a high, forested bluff and
could be watched from the bluff above. The seals seemed to
gather here in the early afternoon. Much of their time was spent
sleeping on the surface with the body in a bowed position and drift-
ing freely. Occasional waving of the rear flippers kept them from
drifting away. They played a great deal, splashing, diving and
swimming in circles or spirals. Occasionally one would dive down to
swim slowly along just above the bottom. At times one would put
on a sudden burst of speed, apparently in pursuit of some fish,
rarely continued for more than 30 or 40 feet after which the seal
usually rose to the surface to resume play. Never did I see one with
a fish in its mouth and these pursuits, if pursuits they were, seemed
to have been prompted more by a spirit of play than by hunger.

A young seal was kept as a pet for several weeks at Friday
Harbor. San Juan County. It was tame and affectionate but de-
cidedly temperamental. It demanded constant attention and whim-
pered, cried or moaned it' left alone. It was more active by night
than by day and made a characteristic mooing cry, which continued
at intervals throughout the night. This habit led to its eventual
disposal. The temper of this young seal was shown by it- actions
when it encountered strands of kelp while swimming. It' the kelp
Strands held back its progress it would turn back, seize the kelp
in its teeth and bite viciously. Never did it attempt to bite persons.

The haii- seal was generally thought to feed almost exclusively
on salmon, but the work of Schel'fer (1928: 10-16) -bowed this view
to be incorrect. Of 14 hair seal stomachs examined, all of which

Sea Lions and Seals 249

were full, only two contained salmon. In a later study (Scheffer
and Sperry, 1931: 214-226), only two of 100 stomachs examined
contained salmon. Other food items were tomcod, flounder, sculpin,
herring, shiner, hake, skate, blenny, unidentified fish, squid, octopus,
shrimp, crab and starfish. In spite of Scheffer's work, hair seals
are killed at every opportunity by fisherman and boatman. Boun-
ties were paid on their scalps for many years, and more than 1,000
bounties were paid for each of several years previous to 1930.
Their small, round heads bobbing on the waves offer a poor target
and many seals, after having been fired at, become extremely shy.
They seem to be holding their own in numbers at the present time.

Scheffer and Slipp (1944: 401) found that the young were born
in late May along the ocean coast and in June and July in Puget
Sound. The young seal mentioned as having been kept captive
at Friday Harbor was obtained from an Indian on July 26, 1938,
and was said to be two weeks old at the time. The Indian said that
he had watched the birth of the young and then killed the mother
for bounty. On July 28 the young seal weighed approximately 20
pounds and was in good health. The seal could swim well. It
was said to have been born "on the rocks" at Long Island, San
Juan County. The seal drank milk from a baby's bottle but
refused fresh scallops, clams and fish of several species. When
put into a large, screened box sunk in the water it at once investi-
gated the other animals in the box. It showed no fear of a large
bull cod weighing 50 pounds, or of a 20-pound skate and several
sharks 5 feet long but seemed to be frightened by a large octopus
weighing about 30 pounds. In swimming, the front flippers were
held flat against the body and the actual swimming was accom-
plished by the vertically-held rear flippers and the rapid swinging
of the hips. Its eyes were very dark brown, almost black, but
soft and appealing. The bases of the vibrissae were thick and soft.
The belly was silvery white and unspotted. The sides and back
were iron gray spotted with dark, bluish gray, the whole overlaid
with a silvery tint. The claws were long, round, and sharply
pointed.

When sleeping, the seal usually lay on its side, occasionally upon
its back or belly. The front flippers were held tight to its sides
but the back flippers were held straight back with the digits bent
inward at right angles and laid so that the right digits were against
the left. In moving on land the front flippers were folded into
fists and used to push the animal forward while the body was moved
by snakelike motions of the hips. It breathed in short gasps.

250 University of Kansas Publs., Mrs. Nat. Hist.

Genus Tamias Illiger
Chipmunks

The chipmunks of Asia and western North America have usually
been separated under the generic name Eutamias from those of the
genus Tamias of eastern America. Ellerman (1940: 428) placed
both in the same genus and Bryant (1945: 257-390) reached the
same conclusions after intensive study of American sciurids. Bry-
ant's treatment is followed here. The sciurid genera as they occur in
Washington, are listed by Bryant as follows: Tamias, Marmota,
Citellus, Sciurus, Tamiasciurus, Glaucomys. This order, rather than
that of Miller (1924) is used here. Four species of Tamias are listed
for Washington: minimus represented by two subspecies; amoenus,
by six: ruficaudus by one; and townsendii, by two.

Chipmunks from Washington vary in size from less than 8 inches
in total length to more than 10 inches in total length. Some race
of chipmunk occurs in almost every part of Washington. Their
striped color pattern serves as a universal recognition mark. The
somewhat similarly striped mantled ground squirrel is often mis-
takenly called chipmunk. The mantled ground squirrel is larger
than any chipmunk, has but two dark stripes as compared with
five dark stripes of chipmunks, and has a plain, reddish head unlike
the distinctly striped head of Tamias.

Like most members of the squirrel family, chipmunks are active
by day and are therefore better known to man than are most of
the other kinds of small mammals, most of which arc nocturnal.
The attractive color and sprightly actions of chipmunks make them
a delightful feature of the outdoors. They feed on fruit, seeds, and
fungus and eagerly eat food that can be begged or stolen from
man. They have been known to kill mice and they have been ac-
cused of destroying nests and eggs of birds. They often eat insects
and occasionally eat the flesh of mice or other chipmunks held in
collector's traps.

Chipmunks climb trees and bushes readily but only townsoxlii
can be called arboreal, and even it prefers to climb on stumps and
dead trunks rather than in living trees. All species are fond of
climbing about rocky outcrops and talus slides.

The call of the chipmunks is a birdlike cheep. In minimus it is
shrill and uttered rapidly, but it is low-pitched and is ottered by
toirnsctxlii with longer intervals between the note-. The call of
amoenus is of an intermediate nature.

Chipmunks 251

Young of chipmunks vary in number from four to six. Nests are
constructed of dry grass and are placed under rocks, logs, and in
burrows in the ground. There are four pairs of mammae, one pec-
toral, two abdominal, and one inguinal.

Tamias minimus Bachman
Least chipmunk

Description. — The least chipmunk is the smallest chipmunk found
in Washington. The head and body of adults measure about 3%
inches; the tail about 3Vo inches. Its fur is short and sleek. The
dorsal stripe is black; the upper pale stripe is buffy gray; the lower
dark stripe is rich brown; the lower stripe is white. The sides are
pale buff and the head, rump and thighs are buffy gray. The tail
is brownish above, yellowish beneath.

Tamias minimus has a wide range, being found from the Cas-
cade-Sierra Nevada Chain to the Great Lakes and from northern
Canada to central Arizona and New Mexico. Two races occur in
Washington, both in the sagebrush desert area.

Least chipmunks are only locally common in Washington. I
have found them in areas where the soil was firmly packed and
sagebrush the dominant vegetation. All were far from water. Two
miles west of Vantage, Kittitas County, several were found near
an old sheep corral, where one took shelter in a pile of boards. In
my experience, least chipmunks are wary and difficult to collect.
Many times as I crept silently through the sagebrush chipmunk
after chipmunk scampered with tail aloft into a hole at the base of
same sage bush, each far out of gunshot, voicing alarmed chirps.
The extreme caution of least chipmunks, as compared with other
species, is doubtless a necessary adaptation to living in an exposed
situation. The open sagebrush desert is a favored hunting place
of hawks and eagles; also coyotes, wildcats, and badgers sometimes
abound there. All these probably find the least chipmunk a suit-
able food item and only the most cautious chipmunk survives to re-
produce. The least chipmunk has been timed at a speed of 10 miles
per hour (Cottam and Williams, 1943: 262).

The food of the least chipmunk in eastern Washington is almost
entirely seeds of the annuals that flourish briefly in the spring. In-
sects probably are eaten and one specimen had the remains of two
scorpions in its stomach.

252

University of Kansas Publs., Mrs. Nat. Hist.

Tamias minimus scrutator (Hall and Hatfield)

Eutamias minimus pictus Howell, X. Ahum. Fauna. 52:39, November 30, 1!»29.

Eutamias minimus scrutator Hall and Hatfield, Univ. California Publ. Zool., 40:321, Feb-
ruary 12, 1934.

Tamias minimus scrutator Ellerman, Fam. and Genera of Living Rodents, British Mus. Xat.
Hist., p. 432. June S, 1940.

Type.— Obtained near Blanco Mountain. 10500 ft. elevation. Mono County,
California, by J. Grinnell on July 28. 1917; type in Museum of Vertebrate Zo-
ology.

Racial characters. — Buffy color; wide dark stripes and narrow pale stripes.

Measun rm nts.—A male and a female from Sunnyside. Yakima County,
measure respectively; total length 186, 186; length of tail 81, 82; hind foot 31,
31 ; ear 9, 10.

DISTRIBUTION MAP
STATE OF WASHINGTON

MILES

Fig. 80. Distribution of the least chipmunk in Washington. A. Tamias mini-
mus scrutator. H. Tamias minimus grisescens.

Distribution. — The sagebrush areas west of the Columbia River, Present in
scattered and widely separated areas, ranging, according to Howell (1929: 41),
north to Ellensburg and south to Wiley City.

This species is a member <>! the Greal Basin fauna thai entered the state
from Oregon. The population in Washington is now isolated north and west
of the Columbia River but seems not to differ from leasl chipmunks from
Oregon and Nevada.

Chipmunks 253

Tamias minimus grisescens (Howell)

Eutamias minimus grisescens Howell, Jour. Mamrn., 6:52, February 9, 1925.
Tamias minimus grisescens Ellerman, Fam. and Genera of Living Rodents, British Mus. Nat.
Hist., p. 431, June 8, 1940.

7'///" .—Obtained at Farmer. Douglas County, Washington, by J. A. Loring,
on July 31, 1897; type in United States National Museum.

Racial characters. — Similar to scrutator but smaller; more grayish, less buffy
in color, dark stripes narrower and pale stripes wider.

Measurements. — Seven topotypes and near topotypes average: Total length
177; length of tail 78.7; hind foot 26.8; ear 10.6.

Distribution. — Known only from the Columbian Plateau. This race is rare
and though we hunted for it in localities where specimens have been collected,
including the type locality, it was found but twice. Ranchers living in the area
know the chipmunk but see individuals only occasionally. Marginal occur-
rences are Douglas (Howell, 1929: 41), Vantage (V. B.S.) and Pasco (Howell,
1929: 41).

h'i marks. — The geographic range of this race is separated from that of
scrutator by the Columbia River and many miles of country uninhabited by
chipmunks of this species.

Tamias amoenus Allen

Yellow-pine chipmunk

Description. — The yellow-pine chipmunk resembles the least chip-
munk but is larger. The underside of the tail is more ochraceous,
less yellowish. The color of the sides varies considerably in the
various races. The stripes are narrow and sharply delineated, the
dorsal one being black. Next lower is a grayish stripe, followed by
one of brownish black. The lower stripe is white. The underparts,
in most races, are white but in luteiventris are buffy.

The geographic range of the species is west of the Great Plains
from central British Columbia to central California. There are
twelve races, six of them occurring in Washington.

The yellow-pine chipmunk is a small animal, being but little
larger than the least chipmunk, and much smaller than townsendii.
Externally it may be separated from the Townsend chipmunk by
its small size, sleek, appressed pelage and brighter color. Separa-
tion from minimus is more difficult but, in the Washington races of
minimus, the colored fur of the underside of the tail is pale yellow-
ish while in amoenus it is more ochraceous or buffy. So far as is
known, the two species do not occur together in Washington.

The yellow-pine chipmunks live in open woods, brushy areas,
clearings, and rocky outcrops. Suitable conditions are abundant
in mountainous areas and the distribution of mountain ranges affects
the distribution of these chipmunks. Where yellow pine forests
descend to relatively low altitudes, the chipmunks enter the low-

254 University of Kansas Publs., Mrs. Nat. Hist.

lands. Such conditions are present in some places along the eastern
base of the Cascades, the inter-mountain river valleys of north-
eastern Washington, and along the central-eastern border of the
state.

Yellow-pine chipmunks arc sprightly and active. They seem
always to be moving restlessly about, running, investigating for food,
and watching for enemies. They are far tamer than the least chip-
munks, usually allowing the observer to approach within twenty
feet or closer. Some, after coaxing, will take food from a person's
hand.

Near Stevens Pass, King County, numbers of Tamias townsendii
cooperi and Tamias amoenus ludibundus, apparently on good terms,
were feeding together on blue huckleberries which grew in abun-
dance on an extensive snowslide area. The yellow-pine chipmunks
had been drawn from surrounding open areas by the berries, while
the Townsend chipmunks had been attracted from the forest by
the same food. The nervous movements of amoenus contrasted
strongly with the more sedate behavior of townsendii. At the ob-
server's close approach the yellow-pine chipmunks went scampering
off through the brush and tangles of logs and branches, to emerge
again and watch from a hundred feet away. The same individual-,
when repeatedly followed, always remained in sight. Most of the
Townsend chipmunks, when frightened, ran into the nearest dense
cover and vanished, not to appear again. A goodly number, perhaps
ten per cent, climbed high up in fir trees. None of the yellow-pine
chipmunks took refuge in trees.

The yellow-pine chipmunk is usually found at considerable alti-
tude and consequently there is deep snow and bitter cold in winter
where it lives. Hibernation is probably complete. Seemingly these
chipmunks depend on stored food rather than on accumulated fat
to tide them over the winter, for animals collected in autumn an'
no fatter than those taken in the spring. Svihla (1IKWB: 290) found
that Tamias a. canicaudus, hibernating in captivity at Pullman,
Whitman County, awoke at intervals to eat stored food. The time
of retirement of yellow-pine chipmunks for the winter seems to co-
incide with the coming of winter weather. In mid-November of
one year, when no snow had vet fallen in the Cascades, yellow-pine
chipmunks were common near Stevens Pass, although their actions
were noticeably slow. In another autumn, when the winter snows
came early, I looked in vain for chipmunks in October where they
had been common earlier. George C. Cantwell noted a yellow-pine
chipmunk at Republic, Ferry County, on November !), 190.'}, after
the ground was "well frozen." but apparently free of snow (Howell,

Chipmunks 255

1929: 7). Like other species that hibernate, they, at times, seem to
become active in winter; J. B. Flett reported seeing a yellow-pine
chipmunk at Longmire, Mt. Rainier, on February 14, 1920, and
again on March 31 (Howell, loc. cit., p. 7). At Deer Park, Clallam
County, at timber-line, several Tamias tOwnsendii cooperi were
active in early April, 1938, but only one Tamias amoenus was seen.
In the previous June they were abundant there and townsendii was
scarce.

I have watched these chipmunks eat the berries of the red huckle-
berry (Vaccinium parvifolium) , salmonberry (Rubus spectabilis),
thimbleberry (Rubus parviflorus) , devil's club (Fatsia horrida) , and
mountain ash (Sorbus casoadensis and S. occidentalis) . Of these,
the blue huckleberry (Vaccinum occidentale and V. membranaceum)
are probably the most important to the chipmunks. At Sherman
Creek Pass, in the Kettle River Mountains, Ferry County, I watched
a pika (Ochotona) busily harvesting wild raspberry plants {Rubus
leucodermis) , and laying them in a pile under a rock. A yellow-
pine chipmunk waited under the rock and ate the ripe berries from
each branch as it was laid away.

A brood nest of the yellow-pine chipmunk was discovered by
Shaw (1944: 274) at Hurricane Ridge, Clallam County. The en-
trance was a hole l 1 /^ inches in diameter constructed among the
grass and alpine flowers of a meadow at 6,450 feet elevation. The
burrow itself was 2 inches in diameter, and had a turning-around
pocket 9 inches from the entrance. The nest was situated just be-
neath the sod, 4 feet from the entrance. The nest chamber was 7
inches high by 7 x /2 inches in diameter and in the shape of a "round-
bottomed flask." It was filled snugly with nest material composed
of a grasslike sedge {Carex spectabilis) mixed with feathers of the
blue grouse. Earth excavated from the nest cavity had been forced
upwards through the sod in the manner of a mole in forming mounds.
The single burrow was unbranched. Seven young of about 16 to
18 days of age were found in the nest.

Tamias amoenus caurinus (Merriam)

Eutamias caurinus Merriam, Proc. Acad. Nat. Sci. Philadelphia, p. 352, October 4, 1898.
Eutamias amoenus caurinus Howell, Jour. Mamm., 3:184, August 4, 1922.
Tamias amoenus caurinus Ellerman, Fam. and Genera of Living Rodents, British Mus. Xat.
Hist., p. 432, June 8, 1940.

■Type. — Obtained at timberline, head of Sol Due River, Olympic Moun-
tains, Clallam County, Washington, by C. H. Merriam and Vernon Bailey on
August 27, 1897; type in United States National Museum.
Racial characters. — Small size and pale color.

256

University of Kansas Publs., Mrs. Nat. Hist.

Measurements * — Seven males and 3 females from Deer Park, Clallam
County, average: total length 207.4; length of tail 93.2; hind foot 32.5; ear 16.

Distribution. — Higher parts of the Olympic Mountains, from Deer Park
(W.W.D.) south to head of Dosewallips River (Howell, 1929:77).

Tamias amocnus felix Rhoads

Tamias quadrivittatus felix Rhoads, Amer. Nat., 29:941, October, 1895.

Eutamias quadrivittatus felix Miller and Rehn, Proc. Boston Soe. Nat. Hist., 30:44, De-
cember 27, 1901.

Eutamias amoenus felix Howell, Jour. Mamm., 3:1S4. August 4. 1922.

Tamias amoenus felix Ellerman, Fatn. and Genera of Living Rodents, British Mus. Nat.
Hist., p. 432, June 8, 1940.

Type. — Obtained at Church Mountain, British Columbia, near the United
States boundary by Allan Brooks on August 13. 1895; type in Philadelphia
Academy of Natural Sciences.

DISTRIBUTION MAP
STATE OF WASHINGTON

50 75 100

D.oft*d Und». Aviptol of WPA P-.j.c. iM-mr b,CJ1T. rw

Fig. SI. Distribution of tin- yellow-pine chipmunk in Washington. A.
Tamias amoenus luteiventris. IV Tamias amoenus canicaudus. C. Tamias
mum mis afjinis. 1). Tamias amoenus ludibvndus. E. Tamias. amoenus felix.

V. Tamias unm< nus ca u nit us.

Racial characters^. — Large size and rich, dark color.

Measurements.— Ten topotypes average: total length 224.7; length of tail
C8.8; hind foot. 34.1; ear 1 1.1.

Distribution. The extreme aorthwestern Cascades, north and wesi of Mt
Baker.

Chipmunks 257

Remarks. — This richly-colored coastal race barely enters Washington. It
is abundant near Goldrun Pass and Tomyhoi Lake, Whatcom County, just
south of the international boundary.

Tamias amoenus ludibundus (Hollister)

Eutamias ludibundus Hollister, Smithsonian Misc. Coll.. 50 (no. 26) :1, December 5, 1911.
Eutamias amoenus ludihundus Howell, Jour. Mamm., 3:184, August 4, 1932.
Tamias amoenus ludibundus Ellerman, Fam. and Genera of Living Rodents, British Mus.
Nat. Hist., p. 432, June 8, 1940.

Type.— Obtained at Yellowhead Lake, 3,700 ft., British Columbia, by N.
Hollister on August 29. 1911; type in United States National Museum.

Racial charactt rs. — Moderate size, brownish rump, ochraceous sides.

Measun rm nts. — Five males and 4 females from the higher parts of the
Cascades average, respectively: total length 210, 217; length of tail 89, 90;
hind foot 33, 33; ear 17.4, 16.7; weight 50, 59.7 grams.

Distribution. — The higher Cascade Mountains. Marginal records are:
Barron (Howell. 1929:75) Lyman Lake (Howell, 1929:75); Cascade Tunnel
(W.W.D.), Mt, Stuart (W.W.D.), Lake Kachees (W.W.D.), Boulder Cave
(W.W.D.), and Mt. St. Helens (Howell, 1929:75).

Ri marks. — At the higher altitudes this race seems to be the equivalent of
affinis. The latter race lives in relatively arid yellow pine forests and ludi-
bundus occupies more moist and varied habitats higher in the mountains.

Tamias amoenus affinis Allen

Tamias quadrivittatus affinis Allen, Amer. Mus. Nat. Hist. Bull., 3:103, June, 1890.

Eutamias quadrivittatus affinis Miller and Rehn, Proc. Boston Soc. Nat. Hist., 30:44, De-
cember 27, 1901.

Eutamias amoenus affinis Howell, Jour. Mamm., 3:184, August 4, 1922.

Tamias amoenus affinis Ellerman, Fam. and Genera of Living Rodents, British Mus. Nat.
Hist., p. 432. June 8, 1940.

Type. — Obtained at Ashcroft, British Columbia, by C. P. Streator on July
3. 1889; type in United States National Museum.

Racial characters. — Small size, grayish color including rump, and white un-
derparts.

Mcasifn nil <)tts. — Fourteen males and 7 females from Washington average:
total length 201.5; length of tail 86.5; hind foot 31.7; ear 17.2.

Distribution. — The eastern slope of the Cascade Mountains. Marginal rec-
ords on the west are: Bald Mountain (Howell, 1929:73), Mazama (Howell.
1929:73), Hart Lake (Howell. 1929:73), Lake Wenatchee (W.W.D.), 10 mi.
S. Dryden (W.W.D.). Blewett Pass (W. W.D.), 10 mi. N. W. Ellensburg
(W.W.D.). Wenas Creek (W.W.D.), Mt. Adams (Howell. 1929:73), and
Lyle (Howell. 1929: 73). Marginal occurrences on the east are: Mt. Chopaka
(Howell, 1929:73). 20 mi. E. Tonasket (W.W.D.) and Omak Lake (Howell,
1929:73).

Tamias amoenus canicaudus (Merriam)

Eutamias canicaudus Merriam, Proc. Biol. Soc. Washington, 16:77, May 29, 1903.
Eutamias amoenus canicaudus Howell, Jour. Mamm., 3:184, August 4, 1922.
Tamias amoenus canicaudus Ellerman, Fam. and Genera of Living Rodents, British Mus.
Nat, Hist., p. 432, June 8, 1940.

17—4993

258 University of Kansas Publs., Mrs. Nat. Hist.

Type. — Obtained at Spokane. Spokane County, Washington, by C. P.
Streator, on April 11. 1891; type in United States National Museum.

Racial characters. — Large size, pale color, grayish tail, white or huffy under-
pays.

Measurements. — Thirteen topotypes average: total length 227.2; length
of tail 104.4; hind foot 33.7; ear 14.

Distribution. — The pine-covered lowlands along the central-eastern border
of the state, ranging, according to Howell (1929: 71). from Spokane County
south to Pullman.

Tamias amoenus luteiventris Allen

Tamias quadrivittatus luteiventris Allen. Amer. Mus. Nat. Hist. Bull., 3:101, June, 1890.

Eutamias quadrivittatus luteiventris Miller and Rehn, Proc. Boston Sue. Nat. Hist., 30:44,
December 27. 1901.

Eutamias amoi nus luteiventris, Howell, .lour. Mamm., 3:170, Augusl 4. 1922

Tamias amoenus luteiventris Ellerman, Fain, and Genera of Living Rodents, British Mus.
Nat. Hist., p. 432. June 8, 1940.

Type.— Obtained at Chief Mountain Lake (Waterton Lake). Alberta (3%
mi. N. United States boundary) by Elliott Cones on August 24. 1874; type in
United States National Museum.

Racial characters. — Small size, rich color, huffy underparts.

Measuremt nts. — Twelve males and 12 females from the Blue Mountains,
Columbia County, average respectively: total length 212, 219; length of tail
96.7. 101; hind foot 31.7. 32.5; ear 17.3. 18; weight 46.5. 52.8 grams.

Distribution. — The Blue Mountains of southeastern Washington, and the
Pend Oreille Mountains of northeastern Washington, wesl to Eureka, in
the Kettle River Mountains. Ferry County (Howell. 1929: 69), and south
to Newport (W.W.D.).

Tamias ruficaudus simulans (Howell)

Red-tailed chipmunk

Eutamias ruficaudus simidans Howell, Jour. Mamm., 3:179, Augusl 4, 1922.
Tamias rati rum hi* simulans Ellerman, Fam. and Genera of Living Rodents, British Mus.
Nat. Hist., p. 434, June 8, L940.

Type. — Obtained at Coeur d'Alene, Kootenai County, Idaho, by C. P.
Streator on June 1. 1891.

Measurements. — Six males and 3 females from northeastern Washington
average: total length 234; length of tail 109; hind foot 31.6; ear 18.

Distribution. — Northeastern Washington, reported from Pend Oreille,
Stevens and Ferry counties by Howell (1929: 98).

Description. — The red-tailed chipmunk closely resembles Tamias
amoenus. From T. <i. luteiventris and T. a. canicaudus it differs

principally in larger size, wider brain ease and especially in its pure
white underparts. From T. a. affinis it differs in richer coloration,
especially the brownish lather than gray rump. 'The difference-
separating it from amoenus are -light, and only adult specimens
can he identified in the field.

Chipmunks 259

According to Howell (1929: 81) this specie? occurs in northern
Idaho, western Montana, northeastern Washington, southeastern
British Columbia and extreme southwestern Alberta. Two sub-
species are recognized, only one of which occurs in Washington.

A number of large, white-bellied chipmunks have been taken in
northeastern Washington that answer well to the description of rufi-
caudus. Also, there are a number of specimens that I cannot defi-
nitely identify as either amoenus or ruficaudus. Some buff-bellied
chipmunks from Idaho are as large, and possess brain cases as wide,
as specimens from Washington unhesitatingly called ruficaudus,
while some white-bellied individuals match amoenus luteiventris in
all other characters. When all the specimens available from Pend
Oreille, Stevens and Ferry counties are separated into amoenus and
ruficaudus and the skulls are examined, it is noticeable that all the
ruficaudus are old, fully adult animals and that most of the amoenus
are younger, showing less wear on the teeth. Perhaps the buffy
underparts are lost with increasing age.

This leads to the suspicion that ruficaudus, as applied to chip-
munks in Washington, is a synonym of amoenus. Until considerable
material is collected in northeastern Washington, showing individual
and age variation, it seems best to retain the name ruficaudus.

Most of the specimens referable to ruficaudus were taken in talus
slides high in the Pend Oreille Mountains. A smaller series was
collected at a lower altitude in open pine forests near Pend Oreille
Lakes, Stevens County. I could detect no difference between
amoenus and ruficaudus in habitat or habits.

Tamias townsendii Bachman
Townsend chipmunk

Description. — The Townsend chipmunk is the largest of the chip-
munks that occur in Washington. Head and body measure about
5V 2 inches; the tail about 4% inches. The fur of the Townsend
chipmunk is more lax and less sleek than that of other species. The
upper parts are duller and darker ochraceous. Stripes are wide and
not sharply delineated. The dark stripes are deep chestnut or black-
ish. The upper pale stripe is pale buffy gray; the lower is whitish.
The tail is blackish frosted with white above and rich ochraceous
below. Under parts are dull white.

The Townsend chipmunk ranges from the Fraser River in south-
ern British Columbia through western Washington and Oregon, to
central California. Like other members of the Pacific Coastal
Fauna which extend southward to California, its geographic range

260 University of Kansas Publs., Mrs. Nat. Hist.

extends farther inland to the south and geographic variation is
greater; 3 races are listed by Johnson (1943: 114 ) in California.

The Townsend chipmunk is the largest and darkest chipmunk in
Washington. Over much of its range it is the only chipmunk found
although in some mountainous areas both townsendii and amoenus
occur together. The larger size and richer coloration, especially the
rich tawny color of the underside of the tail, separate Townsend
chipmunks from amoenus.

Townsend chipmunks are closely associated with the coniferous
forest where they live in clearings and tangles of underbrush such

Fig. 82. Townsend chipmunk (Tamias townsendii coopcri), captured on
Goat Creek, 3,000 feet, western Cascade Mountains near Chinook Pass, Wash-
ington, September 16, 1940. bv Earl J. Larrison; photographed February 1,
1941. (Fish and Wildlife Service photo by Victor B. Scheffer. No. 1139.)

as on steep hillsides, fire slashings, snow-slide areas or mountain
glades. Though capable of swift movement, they are less nervous
and active than other chipmunks. Rarely are they as abundant,
even locally, as other species, and the home range of an individual
seems to be larger than that, of an amoenus or a minimus. Rarely
are more than two seen in a locality. Their more sedentary habits
make them less conspicuous than other species. In the fall, when
beiries are dried out and seeds are ripe, they are most often seen.
The Townsend chipmunk seems to be the most arboreal species found
in Washington and individuals are not uncommonly seen in trees.

The brushy cover inhabited by Townsend chipmunks protects
them from most hawks while their diurnal habits prevent owls from
feeding on them. Predatory mammals probably constitute their

Chipmunks

261

greatest menace. A weasel (Mustela frenata) was seen to follow
a Townsend chipmunk into a pile of timbers near Cottage Lake,
King County, and another was seen carrying a dead Townsend chip-
munk at Stevens Pass, King County. Tracks around an area of
blood and fur showed where a mink {Mustela vison) had killed a
Townsend chipmunk near Cottage Lake, King County.

DISTRIBUTION MAP
STATE OF WASHINGTON

MH.C5

cfred Under A.;p-i> c' «■. PA I

Fig. 83. Distribution of the Townsend chipmunk in Washington. A. Tamias
tovmsendii townsendii. B. Tamias toumsendii cooperi.

Shaw (1944: 278) discovered the brood nest of a Townsend chip-
munk on Hurricane Ridge, Clallam County, 4,500 feet elevation.
The burrow was in a cool, damp area among surface runs of moles
(Scapanus), and led to an underground nest among the roots of a
tree. The nest was formed of the gray, moss-like lichen (Usnea) ,
lined within with sedge leaves (Carex spectabilis) and covered out-
side with leaves of the same sedge. A turning-about chamber was
constructed near the entrance. The burrow was single and not
branched. Three young, only two or three days old, were in the nest.

2(i2 University gf Kansas Publs., Mis. Nat. Hist.

Tamias townsendii townsendii Bachman

Tamias townsendii Bachman, Jour. Acad. Nat. Sci. Philadelphia, 8 (pt. 1):G8, 1839.
Tamias quadrivittatus townsendii Allen, Proc. Bostcn Sue. Nat. tL-t., 16:290, 1^74.
Tamias asiaticus var. townsendi Allen, Monog. N. Amer. Rodentia, Kept. U. S. Geol.
Surv. Terr.. 11 : 794, ls77.

Eutamias townsendi Merriam, Proc. Biol. Soc. Washington, 11:195, July 1. 1897.

'I'hjii . — Eectotype obtained near The lower mouth of the Willamette River.
Multnomah County. Oregon, by .1. K. Townsend in 1834; in Philadelphia
Academy of Natural Sciences.

Racial Characters. — Color of sides rich tawny; dark stripes black or dark
brown, and pale stripes cinnamon; underside of tail tawny.

Measurements. — Fifteen males and 10 females from western Washington
average, respectively: total length 254.7, 258.6; length of tail 116. 122; hind
foot 36.7, 37.1; ear 20.5, 20.4; weight 72. 81.2 grams.

Distribution. — The humid coastal belt of western Washington, from the
western base of the Cascade Mountains to the Pacific, exclusive of the Olympic
Mountains. When A. H. Howell revised the chipmunks in 1929, he employed
a concept of a subspecies different from the writer's own. The locality records
listed by Howell (1929: 109-112) for Tamias townsendii tovmsendii and T. t.
cooperi are not in agreement with Howell's own distribution map (op. cit.:
107). When the localities listed by Howell are plotted on a map of Wash-
ington, the ranges of the two races overlap in some critical areas. Not all of
the material examined by Howell was seen by the writer, and, consequently,
the ranges shown in Fig. 83 are plotted, in part, on geographic grounds.

Marginal localities on the east, so plotted, for T. t. towiixcialii, are: Hamil-
ton (U.S.N.M.), 5 mi. E. Monroe (W.W.D.), Redmond (W.W.I).). Roy
(U.S.N.M.), and Vancouver (U.S.N.M.).

Tamias townsendii cooperi Baird

Tamias cooperi Baird, Proc. Acad. Nat. Sci. Philadelphia, 7:MK4. 1855.
Tamia* townsendii var. cooperi Baird, Mamm. X. Amer., p. 7:-!7, is.">7.
Eutamias cooperi Lyon, Smithsonian Misc. ('"II.. 50:89, June 27. 1907.
Eutamias townsendii cooperi Unwell. Jour. Mamm., :f:ls4, August 4, 1922.

Type.— Cotypes obtained at Klickitat Pass, 4,500 ft., Skamania County.
Washington, by J. S. Cooper in July. 1853; in United States National Museum.

Racial Characters. — Similar to T. t. tovmsendii but paler with pale stripes
whitish rather than cinnamon.

Measurements. — Ten males and 10 female- from the Cascades average, re-
spectively: total length 246 4. 246.8; length of tail 111.7. 107.6; hind foot 35,
35.4; ear 20. 20; weight 77. 89.9 gram.-.

Distribution. — The higher and eastern Cascade Mountains and the Olympic
Mountains. Marginal localities along the Cascades probably include: Swamp
Creek (U.S.N.M.), Index (W.S.C.). North Bend (U.S.N.M.), Mt. St.
Helen- (U.S. N.M.). and Yacolt (M.V.Z.).

Marmots 2(>3

Marmota monax petrensis Howell
Woodchuck

Marmota monax petrensis Howell, N. Amer. Fauna, 37:33, April 7, 3915.

Type. — Obtained at Revelstoke, British Columbia, by W. Spreadborough on
May 12. 1890; type in United States National Museum.

Measurements. — Howell (1915: 33) gives the measurements of an adult
male (the type) as: total length 540; length of tail 127; hind foot 76. An
adult female from Barkerville, British Columbia, measured: total length 505;
length of tail 125; and hind foot 68.

Distribution. — The Pend Oreille Mountains. Pend Oreille County. The
woodchuck was seen and positively identified in northeastern Washington but
no specimens were collected.

Description. — The woodchuck is the smallest member of the genus
Marmota that occurs in "Washington. Adults are about 22 inches
in length, of which the tail comprises 5 inches. The body is stout
and plump. The legs are short. The ears are low and rounded.
The eyes are large but not prominent. The fur is rather stiff but
dense. The upper parts are cinnamon, frosted with white-tipped
guard hairs. The underparts are tawny. There is no white bar
across the nose.

Woodchucks occur from Alaska to Idaho and eastward to the
Atlantic, extending southward in the eastern United States. Miller
(1924: 173-1751 lists seven subspecies, one of which enters the ex-
treme northeastern corner of Washington.

The habits of the eastern woodchuck {Marmota monax rufescens)
have been studied by Hamilton (1934: 85-178), but the northern
races are less well known. Cowan (1939: 77-79) gives observa-
tions on the habits, nests and burrows of Marmota monax cana-
densis.

Marmota flaviventris avara (Bangs)

Yellow-bellied marmot

Arctomys ftaviventer munis Bangs, Pioc. New England Zool. Club, 1:68, July 31, 1899.
[Marmota ftaviventer] avarus Trouessart, Catal. Mamm., viv. foss., suppl., p. 344, 1904.
Marmota flaviventris avara Miller, U. S. Nat. Mus. Bull., 128:175, April 29, 1924.

Type— Obtained at Okanogan, British Columbia, by A. C. Brooks on July
17. 1897; type in Museum of Comparative Zoology.

M( asurements.—A male from 5 miles north of Entiat. Chelan County, meas-
ured: total length 610; length of tail 182; hind foot 70; ear 28.

Dist ribution.— From the eastern edge of the Cascade Mountains eastward,
except, apparently, extreme northeastern Washington. Marginal records are:

264 University of Kansas PrBLS., Mus. Nat. Hist.

Okanogan (W.W. D.) in the north. Wenas (W. W.D.) in the west, Pasco
(M.V.Z.) in the south, and 14 mi. S. W. Pullman (Howell, 1915:42) in the
east.

Description. — The yellow-bellied marmot is similar to the wood-
chuck but is slightly larger. It is decidedly paler in color, less
reddish, but possesses white-tipped hairs on the dorsal surface, as
does the woodchuck. The yellow-bellied marmot also differs from
the woodchuck in