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Title: Mammals of Washington, Volume 2 - University of Kansas Publications Museum of Natural History
Author: Dalquest, Walter Woelber
Language: English
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                             MAMMALS OF WASHINGTON


                               WALTER W. DALQUEST

                      University of Kansas Publications
                          Museum of Natural History

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

                                  April 9, 1948

                              UNIVERSITY OF KANSAS


                             MAMMALS OF WASHINGTON

  [Illustration: Mount Rainier from Indian Henry's Hunting Ground, July,
                 1934. (Fish and Wildlife Service photo by Victor B.
                 Scheffer. No. 864.)]

                             MAMMALS OF WASHINGTON


                               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

                             UNIVERSITY OF KANSAS

             Editors: E. Raymond Hall and Donald F. Hoffmeister

                  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

  [Illustration: decoration]




  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


    MAMMALS                                                       68

  EXPLANATION OF TREATMENT                                       110

  CHECK LIST OF MAMMALS                                          112

  ACCOUNTS OF SPECIES AND SUBSPECIES                             121

  ADDENDA                                                        416

  BIBLIOGRAPHY                                                   417

  INDEX                                                          430



  FIGURE                                                        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    35

    15. Upper Sonoran Life-zone                                   37

    16. Canadian Life-zone                                        38

    18. McDowell Lake. Little Pend Oreille Wildlife Refuge        50

    19. Pend Oreille River near Newport                           59

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


  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-eared bats                 162

    48. Female black bear and two cubs                           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 or mountain lion: skin                            234

    73. Cougar or mountain lion: pelts                           235

    75. Canadian lynx                                            238

    76. Trapper's catch of nine Canadian lynx                    239

    78. Bobcat                                                   241

    82. Townsend chipmunk                                        260

    87. Golden-mantled ground squirrel                           282

    89. Feeding station of Douglas squirrel                      287

    91. Northern flying squirrel                                 292

    95. Northern pocket gopher                                   302

    96. Giant mounds formed by pocket gophers                    304

    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

   115. Runways of Townsend meadow mice                          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                                          133

    28. Merriam shrew and Trowbridge shrew                       134

    29. Wandering shrew                                          137

    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                                            166

    46. Hoary bat                                                168

    47. Pallid bat                                               170

    50. Black bear                                               175

    51. Grizzly bears                                            177

    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. Great Basin pocket mouse                                 298

    94. Ord kangaroo rat                                         300

    98. Northern pocket gopher                                   308

   103. Beaver                                                   321

   104. Northern grasshopper mouse                               323

   105. Western harvest mouse                                    325

   107. Deer mouse                                               329

   108. Bushy-tailed wood rat                                    335

   109. Northern lemming mouse                                   337

   110. Heather vole                                             339

   111. Gapper and California red-backed mice                    342

   113. Pennsylvania meadow mouse                                346

   114. Montane and Townsend meadow mice                         348

   116. Long-tailed meadow mouse                                 352

   117. Water rat                                                356

   118. Creeping mouse                                           357

   120. Muskrat                                                  362

   122. Mountain beaver                                          368

   124. Big jumping mouse                                        372

   125. Porcupine                                                375

   126. Pika                                                     378

   127. Snowshoe rabbit                                          383

   128. Black-tailed jack rabbit                                 386

   129. Nuttall cottontail                                       388


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 subspecies, 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 interest to zoölogists. The
evolutionary changes in several species are relatively great. In color,
for example, the bobcat in the humid coastal area of western Washington
is notable for its dark coloration, 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 diversity. 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 "Mammals 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
Washington 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 classification 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 Washington would be given by reducing
the taxonomic accounts to the minimum and dealing principally with the
problems of distribution.

As originally planned, the present report was to be of joint authorship
by Dr. Victor B. Scheffer of the United States Fish and Wildlife
Service, Seattle, and the writer. The press of other work prevented
Dr. Scheffer 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 assisted 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 report was
not reached until 1938. Intensive field work was done between 1938
and 1940. In August, 1941, the author took up residence at Berkeley,
California. Drafting of the manuscript was begun 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. Scheffer have given assistance in the
course of this work. Dr. E. Raymond Hall, in particular, encouraged
the project and gave assistance in various ways including 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 Wildlife Service, Dr. George E.
Hudson of the Charles R. Conner Museum, Mr. Burton Lauckhart of the
State of Washington Department of Game, and Mr. Ernest Booth of Walla
Walla College. Mrs. Peggy B. Dalquest typed and edited the several
preliminary 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 Zoölogy. The writer's 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 Museum of Vertebrate Zoölogy at the
University of California and 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.


The state of Washington was divided into seven physiographic provinces
by Culver (1936). Culver points out that the physiography, 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 volcanic 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
barrier 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.

  [Illustration: FIG. 1. Cascade Mountains at Canadian boundary, looking
  west along boundary trail. Monument 100 in foreground, headwaters of
  Ashnola River in broad valley at right. (Forest Service photo, No.

The Puget Sound Trough, or Puget Sound Province, is immediately 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 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.

The San Juan Islands represent the glaciated remnants of mountains
that, in preglacial 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 all 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, constitutes 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 4,000 feet in some
places (Russell, 1893) and slope inward from the 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 valleys of the 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.

  [Illustration: FIG. 2. Columbia River one mile west of Kellers Ferry,
  Washington, elevation 1,060 feet, April 16, 1940. (Fish and Wildlife
  Service photo by Victor B. Scheffer, No. 933.)]

The Blue Mountains Province is an area of relatively small extent 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 Mountains Province.


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 combine to
produce the various life-zones and influence the distribution 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 distribution of mammals is
influenced also by certain other factors. Among these are historical
factors and isolation by geographic barriers.

  [Illustration: FIG. 3. Mammalian distributional areas of Washington.
  A. Western Washington. B. Cascade Mountains. C. Northeastern
  Washington. D. Blue Mountains. E. Southeastern Washington.
  F. Yakima Valley. G. Columbian Plateau.]

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

  [Illustration: 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 Northeastern Washington
Distributional Area.

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 Distributional Area
are the same.

  [Illustration: FIG. 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.)]

  [Illustration: FIG. 6. Columbia River at Hunters Ferry, Stevens County,
  Washington, April 15, 1940. (Fish and Wildlife Service photo by Victor
  B. Scheffer. No. 932.)]

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


  Cascade Mountains            { Cascade Mountains
                               { Yakima Valley

  Puget Sound       }
  Willapa Hills     }            Western Washington
  Olympic Mountains }

  Okanogan Highlands             Northeastern Washington

  Columbia Lava                { Columbian Plateau
                               { Southeastern Washington

  Blue Mountains                 Blue Mountains


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 zoölogists. 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 cascadensis_, _Salix nivalis_, _Gaultheria humifusa_,
_Empetrum nigrum_, and the heathers, _Phyllodoce glanduliflora_,
_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 thermometer 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 temperatures 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

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 albicaulis_). The following shrubs are listed
by Jones (1938) as common in the Hudsonian Life-zone on Mt. Rainier:
_Salix barclayi_, _Salix commutata_, _Juniperus sibirica_, _Alnus
sinuata_, _Ribes howellii_, _Lutkea pectinata_, _Potentilla fruticosa_,
_Sorbus occidentalis_, _Spiraea densiflora_, _Pachistima myrsinites_,
_Arctostaphylos nevadensis_, _Arctostaphylos uva-ursi_, _Cassiope
mertensiana_, _Cassiope stelleriana_, _Phyllodoce empetriformis_,
_Rhododendron albiflorum_ and _Gaultheria ovalifolia_.

  [Illustration: FIG. 8. North side of Mount Rainier, 14,408 feet, with
  Mount Adams at left and Mount St. Helens at right. June 19, 1932.
  (Photo by 116th Photo Section, Washington National Guard, No.

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 extensive 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 heterophylla_), amabalis fir
(_Abies amabalis_), white pine (_Pinus monticola_) and noble fir
(_Abies nobilis_). Other plants include _Vaccinnium ovalifolium_,
_Vaccinnium membranaceum_, _Menziesia ferruginea_, _Alnus sinuata_,
_Acer circinatum_, _Sorbus cascadensis_, _Cornus canadensis_,
_Clintonia uniflora_, _Stenauthium occidentale_, _Galium oreganum_,
and _Prenanthes lessingii_. Saprophytes abundant in, if not confined
to, this zone are listed by Jones as: _Monotropa uniflora_, _Monotropa
hypopitys_, _Allotropa virguta_, _Newberrya congesta_, _Pterospora
andromedea_, _Corallorrhiza maculata_, _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°
and 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 average 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 January being but slightly above freezing. Summer
temperatures are extreme; the July average is 73.1° and the average
maximum for the same month is 89°. The highest temperature recorded is

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 watercourses such shrubs as hawthorn (_Crataegus
douglasii_), service-berry (_Amelanchier cusickii_, _Amelanchier
utahensis_), aspen (_Populus tremuloides_), syringa (_Philadelphus
lewisii_), snowberry (_Symphoricarpos albus_), choke-berry (_Prunus
melanocarpa_) and elderberry (_Sambucus caerulae_) form thickets.
Lower in the valley the vegetation is xerophytic, similar to that
of the Columbian Plateau. Sagebrush (_Artemisiae tridentata_) is
dominant. Other shrubs include rabbit brush (_Chrysothamnus nauseosus_,
_Chrysothamnus viscidiflorus_), hop sage (_Grayia spinosa_), black sage
(_Purshia tridentata_) and greasewood (_Sarcobatus vermiculatus_).

  [Illustration: FIG. 9. Mount Rainier. Washington: Cowlitz Chimneys
  from base of Dege Peak, July 19, 1933. (Fish and Wildlife Service
  photo by Victor B. Scheffer, No. 859.)]

Climatic conditions in the Olympic Mountains are, in general, similar
to those of the higher Cascade Mountains. There is a limited area of
Arctic-alpine Life-zone on Mount Olympus. The principal life-zone is
the Hudsonian. The Canadian merges with the lowland Humid-Transition
and is difficult to ascertain as a separate zone. In the Hudsonian
Life-zone the average temperatures are low. Winter climate is bitter
and the snow lies deep. In the 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,
124.98 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 compare 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 rainfall 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 brevifolia_), lodgepole pine (_Pinus contorta_) and
spruce (_Picea sitchensis_) may be common. Deciduous trees are numerous
and include several willows (_Salix_ sp.), aspen (_Populus
tremuloides_), hazel (_Corylus californica_), alder (_Alnus oregona_),
oak (_Quercus garryana_), broadleaf maple (_Acer macrophyllum_), vine
maple (_Acer circinatum_), and flowering dogwood (_Cornus 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 huckleberry (_Vaccinium parvifolium_, _Vaccinium ovatum_),
Oregon grape (_Berberis nervosa_), salal (_Gaultheria shallon_), rose
(_Rosa gymnocarpa_), thimbleberry (_Rubus parviflorus_), salmonberry
(_Rubus spectabilis_), blackcap raspberry (_Rubus leucodermis_) and wild
blackberry (_Rubus macropetalus_).

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 freezing. Summer temperatures are high, the July
average for Colville being 67.2° 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. Coniferous 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 (_Larix
occidentalis_), spruce (_Picea columbiana_), and such shrubs as fool
huckleberry (_Menziesia ferruginea_), _Pachystima myrsinites_, dogwood
(_Cornus canadensis_), wild current (_Ribes petiolare_), mountain
mahogany (_Cercocarpus ledifolius_), spirea (_Spiraea_ sp.), lupines
(_Lupinus_) of several species, maple (_Acer douglasii_), buckbrush
(_Ceanothus sanguineus_), sticky brush (_Ceanothus velutinus_), and
huckleberry (_Vaccinium membranaceum_).

  [Illustration: 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.)]

The Columbian Plateau and southeastern Washington present desert
conditions. At Odessa, 1590 feet, Lincoln County, the average annual
precipitation is only 9.38 inches, and only in the 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 temperature (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 at
Odessa is 25.3° and at Walla Walla 32.4° The average yearly snowfall at
Odessa is 19.4 inches and at Walla Walla 23.5 inches. Vegetation of the
Columbian Plateau and southeastern Washington 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 common. Most typical are grasses and shrubs such as the bunch grass
(_Agropyron inerme_, _Agropyron spictatum_), foxtail (_Alopecurus
aequalis_), cheat grass (_Bromus tectorum_), saltbrush (_Atriplex
truncata_), greasewood (_Sarcobatus vermiculatus_) mustard (_Arabis_
sp., _Brassica_ sp.), sagebrush (_Artemisia rigida_, _Artemisia
tridentata_), rabbit brush (_Chrysothamnus nauseosus_, _Chrysothamnus
viscidiflorus_) and cactus (_Opuntia polyacantha_). 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.


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 Transition Life-zone east of
the Cascades belongs to the Arid-timbered subdivision.

  [Illustration: FIG. 11. Life-zones of Washington. Arctic-alpine not
  shaded. A. Hudsonian 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 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.

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

The Upper Sonoran Life-zone includes the desert areas of Washington
(figure 11 D). Its principal extent is the central Columbian Plateau.
From the central Columbian Plateau, fingerlike projections 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.


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. Consequently 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 extensive habitat is the
forest. Dominant mammalian species include: _Peromyscus maniculatus_,
_Sorex trowbridgii_, _Sorex obscurus_, _Tamiasciurus douglasii_,
_Clethrionomys californicus_, _Aplodontia rufa_, _Glaucomys sabrinus_
and _Odocoileus hemionus_. Mammals are scarce and nocturnal forms
prevail. As a rule, a line of mouse traps set in a forest habitat will
take principally _Peromyscus maniculatus_ with a few _Sorex trowbridgii_
and _Sorex obscurus_ and rarely a _Clethrionomys californicus_. In some
places, especially where the moss is deep, a line of mouse traps will
catch only shrews.

  [Illustration: FIG. 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 second important habitat is the deciduous jungle. This differs from
the forest habitat in that the dominant trees are of the deciduous type
and in that understory vegetation, such as shrubs and annuals, is dense.
The jungle habitat occurs in ravines and in valleys 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 in 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 Townsend chipmunk also occur there.

The prairies form a third habitat. These areas of native grasslands 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.

  [Illustration: FIG. 14. Timbered arid subdivision of the Transition
  Life-zone, Kettle Falls on the Columbia River (now beneath Coulee Dam
  backwater), Stevens County, Washington, June 15, 1938. (Fish and
  Wildlife Service photo by Victor 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.


In Washington the Arid timbered subdivision of the Transition Life-zone
is the open, pine forest. Because of the aridity of this 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 because of the open nature of
the surface of the ground. A night's trapping usually yields only a few
_Peromyscus maniculatus_.

The mammalian fauna of the marshes and streamside thickets is similar.
Shrews including _Sorex vagrans_ and _Sorex obscurus_ are uncommon.
Meadow mice, including _Microtus pennsylvanicus_, _Microtus
longicaudus_, and more rarely _Microtus montanus_, are taken.


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 probably from
competition with the black-tailed jack rabbit. The montane 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_, _Citellus washingtoni_ 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 sagebrush desert in Washington is relatively uniform in nature.
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
grasshopper 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: the black-tailed
jack rabbit, Nuttall cottontail, Ord kangaroo rat, Great Basin pocket
mouse, Townsend ground squirrel, Washington ground squirrel.

Marshes are not uncommon on the Columbian Plateau and elsewhere 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 Transition Life-zone. Meadow mice
found in desert marshes include _Microtus montanus_ and _Microtus
pennsylvanicus_. The only shrew we have found is _Sorex vagrans_. The
harvest mouse (_Reithrodontomys megalotis_) is often abundant in

  [Illustration: 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 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
Hudsonian. 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 dominant. The Douglas squirrel, red
squirrel, northern flying squirrel and Townsend chipmunk are typical
arboreal species. Traps set beneath the trees might catch _Peromyscus
maniculatus_, _Clethrionomys gapperi_, _Neotoma cinerea_, _Sorex
obscurus_, or _Sorex trowbridgii_.

  [Illustration: FIG. 16. Canadian Life-zone forest on Mount Rainier,
  Washington, elevation 2,800 feet, September 14, 1934. Western hemlock,
  Douglas fir, western red cedar, and grand fir. (Fish and Wildlife
  Service photo by Victor B. Scheffer, No. 325.)]


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 caligata_ 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 action 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.

TABLE 1. Distribution of mammals in Washington by Life-Zones. _A._
Abundant. _C._ Common. _R._ Rare.

    Column headers:

    A: Humid Transition
    B: Arid-timbered Transition
    C: Arid-grasslands Transition
    D: Upper Sonoran
    E: Canadian
    F: Hudsonian

            SPECIES AND SUBSPECIES.          | A | B | C | D | E | F
  Neurotrichus gibbsii gibbsii               |   |   |   |   | ? | C
      gibbsii minor                          | A |   |   |   |   |
  Scapanus townsendii                        | A |   |   |   |   |
      orarius orarius                        | A |   |   |   |   |
      orarius schefferi                      |   |   |   | C |   |
      orarius yakimensis                     |   |   |   | A |   |
  Sorex cinereus cinereus                    |   | R |   |   | R |
      cinereus streatori                     | R |   |   |   |   |
      merriami merriami                      |   |   |   | R |   |
      trowbridgii trowbridgii                | A |   |   |   | C |
      trowbridgii destructioni               | A |   |   |   |   |
      vagrans vagrans                        | A |   |   |   | R |
      vagrans monticola                      |   | C | C | A | C |
      obscurus obscurus                      |   |   |   |   | R |
      obscurus setosus                       | C |   |   |   | C | A
      palustris navigator                    |   |   |   |   | C | A
      bendirii bendirii                      | A |   |   |   | C |
      bendirii albiventer                    | C |   |   |   |   |
  Microsorex hoyi washingtoni                |   | R |   |   |   |
  Myotis lucifugus carissima                 |   |   | C | C |   |
      lucifugus alascensis                   | A |   |   |   | C |
      yumanensis sociabilis                  |   |   | C | C |   |
      yumanensis saturatus                   | A |   |   |   |   |
      keenii keenii                          | R |   |   |   | R |
      evotis evotis                          |   | R |   |   |   |
      evotis pacificus                       | R |   |   |   |   |
      thysanodes thysanodes                  |   |   |   | R |   |
      volans longicrus                       | C |   |   |   |   |
      volans interior                        |   | C |   |   |   |
      californicus californicus              |   |   | R | R |   |
      californicus caurinus                  | A | C |   |   |   |
      subulatus melanorhinus                 |   |   | R | A |   |
  Lasionycteris noctivagans                  | A | A |   |   | C |
  Corynorhinus rafinesquii townsendii        | R |   |   |   |   |
      rafinesquii intermedius                |   | R |   |   |   |
  Pipistrellus hesperus hesperus             |   |   |   | R |   |
  Eptesicus fuscus bernardinus               | A | A |   |   |   |
  Lasiurus cinereus cinereus                 | R | R | R | R |   |
  Antrozous pallidus cantwelli               |   |   | R | R |   |
  Ursus americanus altifrontalis             | A | C |   |   | A | A
      americanus cinnamomum                  |   | A |   |   | C |
      chelan                                 |   |   |   |   | R |
  Procyon lotor psora                        | A |   |   |   | C |
      lotor excelsus                         |   | R | R | C |   |
  Martes caurina caurina                     |   |   |   |   | A | C
      caurina origenes                       |   |   |   |   | A | C
      pennanti                               | R |   |   |   | R |
  Mustela erminea invicta                    |   | C |   |   | C |
      erminea gulosa                         |   |   |   |   | C | C
      erminea murica                         |   | R |   |   |   |
      erminea fallenda                       | R |   |   |   |   |
      erminea streatori                      | R |   |   |   | C |
      erminea olympica                       | R |   |   |   |   |
      frenata nevadensis                     |   | C | C | C | C |
      frenata effera                         |   | A | C | C | C |
      frenata washingtoni                    |   |   |   |   | C | C
      frenata altifrontalis                  | A |   |   |   | C | C
      vison energumenos                      | C | C | C | C | C |
  Gulo luscus luteus                         |   |   |   | R | R | R
  Lutra canadensis pacifica                  | C | R | R | R | C |
  Spilogale gracilis saxatilis               |   |   | R | R |   |
      gracilis latifrons                     | A |   |   |   |   |
  Mephitis mephitis hudsonica                |   | A |   |   |   |
      mephitis major                         |   |   | R | R |   |
      mephitis notata                        |   | C | R |   |   |
      mephitis spissigrada                   | A |   |   |   |   |
  Taxidea taxus taxus                        |   | C | C | C | R |
  Vulpes fulva cascadensis                   |   |   |   |   |   | R
  Canis latrans lestes                       | C | A | A | A | C | R
      latrans incolatus                      |   | A | C | C | A |
      lupus fuscus                           | R | R | R?|   | R |
  Felis concolor missoulensis                |   | C |   |   | C |
      concolor oregonensis                   | C | C |   |   | C |
  Lynx canadensis                            |   |   |   |   | R |
      rufus fasciatus                        | A |   |   |   | C |
      rufus pallescens                       |   | A | C | C | C |
  Tamias minimus scrutator                   |   |   |   | C |   |
      minimus grisescens                     |   |   |   | R |   |
      amoenus caurinus                       |   |   |   |   | A | A
      amoenus felix                          |   |   |   |   | A | A
      amoenus ludibundus                     |   |   |   |   | A | A
      amoenus affinis                        |   | A |   |   |   |
      amoenus canicaudus                     |   | A |   |   |   |
      amoenus luteiventris                   |   | C |   |   | A |
      ruficaudus simulans                    |   | C |   |   | A |
      townsendii townsendii                  | A |   |   |   |   |
      townsendii cooperi                     |   | C |   |   | A | R
  Marmota monax petrensis                    |   |   |   |   | R |
      flaviventris avara                     |   | R | C | A |   |
      caligata cascadensis                   |   |   |   |   | R | A
      olympus                                |   |   |   |   | C | A
  Citellus townsendii townsendii             |   |   |   | A |   |
      washingtoni                            |   |   | C | A |   |
      columbianus columbianus                |   | A | C |   |   |
      columbianus ruficaudus                 |   | A |   |   |   |
      beecheyi douglasii                     |   | A |   |   |   |
      lateralis tescorum                     |   |   |   |   | C |
      lateralis connectens                   |   |   |   |   | C |
      saturatus                              |   | C |   |   | A |
  Tamiasciurus hudsonicus richardsoni        |   | A |   |   | A |
      hudsonicus streatori                   |   | A |   |   | A | R
      douglasii douglasii                    | A | A |   |   | A | C
  Sciurus griseus griseus                    | C | C |   |   |   |
  Glaucomys sabrinus oregonensis             | C |   |   |   |   |
      sabrinus fuliginosus                   |   |   |   |   | A |
      sabrinus columbiensis                  |   | A |   |   |   |
      sabrinus latipes                       |   | A |   |   | A |
      sabrinus bangsi                        |   | R |   |   | C |
  Perognathus parvus parvus                  |   |   | C | A |   |
      parvus lordi                           |   |   | C | A |   |
      parvus columbianus                     |   |   |   | A |   |
  Dipodomys ordii columbianus                |   |   |   | A |   |
  Thomomys talpoides devexus                 |   |   |   | A |   |
      talpoides columbianus                  |   |   |   | A |   |
      talpoides aequalidens                  |   |   | A |   |   |
      talpoides wallowa                      |   |   |   |   | A |
      talpoides fuscus                       |   | A | C | R | C | C
      talpoides yakimensis                   |   |   | C | A |   |
      talpoides shawi                        |   |   |   |   | C | A
      talpoides immunis                      |   |   |   |   | A | A
      talpoides limosus                      |   | C | A |   |   |
      talpoides douglasii                    | A |   |   |   |   |
      talpoides glacialis                    | A |   |   |   |   |
      talpoides tacomensis                   | A |   |   |   |   |
      talpoides pugetensis                   | A |   |   |   |   |
      talpoides tumuli                       | A |   |   |   |   |
      talpoides yelmensis                    | A |   |   |   |   |
      talpoides couchi                       | A |   |   |   |   |
      talpoides melanops                     |   |   |   |   | C | A
  Castor canadensis leucodonta               | A | A |   | C | C |
      canadensis idoneus                     | A |   |   |   |   |
  Onychomys leucogaster fuscogriseus         |   |   |   | A |   |
  Reithrodontomys megalotis megalotis        |   |   | C | A |   |
  Peromyscus maniculatus oreas               | A |   |   |   | A | A
      maniculatus hollisteri                 | A |   |   |   |   |
      maniculatus austerus                   | A |   |   |   |   |
      maniculatus rubidus                    | A |   |   |   |   |
      maniculatus gambelii                   |   | C | R | A |   |
      maniculatus artemisiae                 |   | A | R | R | C | C
  Neotoma cinerea occidentalis               |   | C | R | A | C | A
      cinerea alticola                       |   | C |   |   | A | C
  Synaptomys borealis wrangeli               |   |   |   |   |   | A
  Phenacomys intermedius intermedius         |   |   |   |   | C | A
      intermedius oramontis                  |   |   |   |   | C | A
  Clethrionomys gapperi saturatus            |   | R |   |   | A | A
      gapperi idahoensis                     |   | R |   |   | A | A
      gapperi nivarius                       |   |   |   |   | A | A
      californicus occidentalis              | A |   |   |   |   |
  Microtus pennsylvanicus funebris           |   | A | R |   | C |
      pennsylvanicus kincaidi                |   |   |   | A |   |
      montanus nanus                         |   |   | A | C |   |
      montanus canescens                     |   | C | A | A |   |
      townsendii townsendii                  | A |   |   |   |   |
      townsendii pugeti                      | A |   |   |   |   |
      longicaudus halli                      |   | A | C | C | C |
      longicaudus macrurus                   | R |   |   |   | C | A
      richardsoni arvicoloides               |   |   |   |   | C | A
      richardsoni macropus                   |   |   |   |   | C | A
      oregoni oregoni                        | A |   |   |   | A |
  Ondatra zibethicus osoyoosensis            | A | A |   | C |   |
      zibethicus occipitalis                 | A |   |   |   |   |
  Zapus princeps oregonus                    |   |   |   |   | A |
      princeps kootenayensis                 |   |   |   |   | A |
      princeps idahoensis                    |   |   |   |   | A |
      princeps trinotatus                    | A |   |   |   | C | A
  Aplodontia rufa rufa                       | A |   |   |   |   |
      rufa rainieri                          |   |   |   |   | A |
  Erethizon dorsatum epixanthum              |   | A | R | C | A | R
      dorsatum nigrescens                    |   | A | R | C | A | R
  Ochotona princeps cuppes                   |   |   |   |   |   | A
      princeps fenisex                       |   |   |   |   |   | A
      orinceps brunnescens                   |   |   |   |   | C | A
  Lepus townsendii townsendii                |   |   | C | A |   |
      californicus deserticola               |   |   | C | A |   |
      americanus washingtonii                | A |   |   |   | A |
      americanus cascadensis                 |   | C |   |   | A | C
      americanus pineus                      |   | C |   |   | A | C
      americanus columbiensis                |   | A |   |   |   |
  Sylvilagus nuttallii nuttallii             |   |   |   | A |   |
      idahoensis                             |   |   |   | A |   |
  Cervus canadensis roosevelti               | A |   |   |   | A | R
      canadensis nelsoni                     |   | C |   |   | A |
  Odocoileus virginianus leucurus            | A |   |   |   |   |
      virginianus ochrourus                  |   | A |   |   |   |
      hemionus hemionus                      |   | A |   |   | A | C
      hemionus columbianus                   | A |   |   |   | C |
  Ovis canadensis canadensis                 |   | A | A | A | A | A
      canadensis californiana                |   | A | A | A | A |
  Oreamnos americanus americanus             |   |   |   |   |   | A


The composition of the mammalian fauna of any area is dependent 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 Pleistocene and earlier times is
based on fossil skeletons. Such knowledge must necessarily be meager,
for conditions favorable to fossilization and the preservation of
fossils until their subsequent discovery by man, were not of common

In the Cascades and in eastern Washington, the Miocene was a time of
orogeny and great volcanism. Great flows of lava, 4,000 feet thick in
the Snake River area (Russell, 1893), emerging from fissures in the
Snake River area, formed the Columbian Plateau. The Columbian basalt
slopes inward centripetally 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 Columbian 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 its
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 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 deep. 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 continental 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 represent a still earlier glaciation, have been
noted. In eastern Washington the only definite proof of multiple
glaciation is of one glaciation 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 weathered and all save the
hardest pebbles and quartzites, for example, are rotten and disintegrate
at the touch. In contrast, the later deposits 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 Mississippi 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 Washington and
Wisconsin to eastern Washington.

  [Illustration: FIG. 17. Extent of Vashon-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 ice dome centering
in Puget 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 Puget Sound. Fingerlike projections of ice were forced up
valleys of the western Cascades and the northern and eastern Olympics.
Some of these upward moving fingers of ice met and coalesced with valley
glaciers descending from the mountains. At the southern edge of the
glacier, the Black Hills and Porcupine Hills remained above the ice
although partially surrounded 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 Mountains
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 excessive
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. Remains of mammoths
have been found in Vashon till. The presence of many non-boreal species
of mammals in southwestern Washington indicates their persistence
there and that conditions therefore were not intolerable for them.
Probably the climate of southwestern 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 growing in west-central Oregon in late glacial time. These pollen
studies of postglacial peat bogs by Henry P. Hansen give evidence of
postglacial climatic changes. Hansen points out (1941 B, 1941 C) that
climatic changes west of the Cascades were probably slight because of
the influence of the Pacific Ocean. Pollen profiles indicate an early,
cool, dry climate followed by a warmer one and increasing humidity.
The present climate may be considered cool and humid. Most of western
Washington lies in the Humid subdivision of the Transition Life-zone.

  [Illustration: FIG. 18. McDowell Lake, Little Pend Oreille Wildlife
  Refuge, Stevens County, Washington, September 29, 1939. (Fish and
  Wildlife Service photo by Victor B. Scheffer, No. 730.)]

The pre-Wisconsin climate of eastern Washington is unknown. From
the loessial nature of the Palouse Soil (Bryan, 1927), a preglacial
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 Wisconsin 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.


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 are:

    _Sorex merriami_                   _Tamias minimus_
    _Myotis thysanodes_                _Perognathus parvus_
    _Myotis subulatus_                 _Dipodomys ordii_
    _Pipistrellus hesperus_            _Onychomys leucogaster_
    _Antrozous pallidus_               _Reithrodontomys megalotis_
    _Taxidea taxus_                    _Lagurus curtatus_
    _Marmota flaviventris_             _Lepus californicus_
    _Citellus townsendii_              _Sylvilagus nuttallii_
    _Citellus washingtoni_             _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

2. THE PACIFIC COASTAL FAUNA. The mammals of the humid coastal district
range from the Fraser River, British Columbia, southward to the
vicinity of Monterey Bay, California. In Washington typical genera are
_Aplodontia_, _Neurotrichus_ and _Scapanus_. The following species are
typical of the Pacific Coastal Fauna in Washington:

    _Neurotrichus gibbsii_             _Tamiasciurus douglasii_
    _Scapanus townsendii_              _Sciurus griseus_
    _Sorex trowbridgii_                _Clethrionomys californicus_
    _Sorex bendirii_                   _Microtus townsendii_
    _Marmota olympus_                  _Microtus oregoni_
    _Tamias townsendii_                _Aplodontia rufa_

Some species which range outside this faunal area have strongly marked
races confined to it. _Glaucomys sabrinus oregonensis_ and _Lepus
americanus washingtonii_ 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 represented 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 faunal areas to the north and east. There, some mingling of
coastal and Rocky Mountain faunas occurs.

3. ROCKY MOUNTAIN FAUNA. If this fauna be thought of as including
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 borealis_
    _Microsorex hoyi_                  _Phenacomys intermedius_
    _Lynx canadensis_                  _Clethrionomys gapperi_
    _Marmota caligata_                 _Microtus richardsoni_
    _Citellus lateralis_               _Ochotona princeps_
    _Citellus columbianus_             _Lepus americanus_
    _Tamias amoenus_                   _Rangifer montanus_
    _Tamiasciurus hudsonicus_          _Oreamnos americanus_
    _Glaucomys sabrinus_

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


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 occurred 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, _Felis 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 Pleistocene. Matthew (_loc. cit._)
reports remains of the giant beaver, _Castoroides_, from the Silver
Lake deposit of Oregon. This great beaver, as large as a black bear,
was doubtless a resident of Washington 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 postglacial time. Remains
of this form, _Elephas columbi_, have been found in Vashon till.

The three faunas of Washington can be placed in two categories. One is
Sonoran, essentially a desert type, and occupies the Columbian Plateau.
The other two are forest faunas, predominantly boreal in complexion,
and are closely related. The Rocky Mountain Fauna is found in the Blue
Mountains and in northeastern Washington. The Pacific Coastal Fauna is
found in western Washington.

The ice sheets of Vashon-Wisconsin time descended southward to southern
Puget Sound and to the northern edge of the Columbian 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 eliminated. 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 separation of the faunas by successive glaciations is thought
to be responsible for many of the differences now existing between them.

Following the retreat of the ice, the Pacific Coastal Fauna extended
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 Mountain Fauna also invaded the Cascade

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 gibbsii_,
_Sorex trowbridgii_, _Sorex bendirii_, _Tamias townsendii_, _Microtus
oregoni_ and _Aplodontia rufa_. Each of these species has no close
relatives in the Rocky Mountain Fauna and, save perhaps _Sorex
trowbridgii_, 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 remarkable in that it
is composed of either very closely related species or very strongly
differentiated subspecies in each fauna. For example, 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 (_Tamiasciurus douglasii_) of the Cascades, which
has a red belly, is the same as the squirrel of the lowlands of western
Washington but is specifically distinct from the red squirrel
(_Tamiasciurus hudsonicus_) of the Rocky Mountain Fauna, which has a
white belly. In the extreme 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 fuliginosus_)
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 californicus_ of
western Washington. 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 snowshoe
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 princeps_) 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

The third group of mammalian species of the Cascades is composed of
species typical of the Rocky Mountain Fauna such as: _Marmota caligata_,
_Synaptomys borealis_, and _Orcamnos 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 _Aplodontia_, _Neurotrichus_
or _Scapanus_ occurred in the Rocky Mountain faunal 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 shrew, and mountain beaver persisted along the coast,
where there were no boreal conditions. Some alpine forms, such as
_Marmota olympus_, _Ochotona princeps brunnescens_ and _Citellus
saturatus_ persisted 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 evolutionary
changes in the species themselves. In some these were considerable, 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 occurred.
The Rocky Mountain Fauna spread northward and westward, 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
Mountain 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 occurred, 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 Cascades. It is possible that in these two cases,
the related form occurring 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 Washington. Probably the
pre-Wisconsin flora of coastal British Columbia consisted of coniferous
forest similar to that of western Washington today. If this were the
case, the mammalian species in British Columbia corresponded closely to
those of western Washington. 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
come upon the barren, inhospitable plains and deserts of the Columbian

The greater part of the southward moving forms found refuge in the
Cascade Mountains where, for most of Wisconsin time, they were isolated
in the southern Cascades. Examples are _Sorex palustris_, _Martes
caurina_, _Martes pennanti_, _Gulo luscus_, _Vulpes fulva_, _Lynx
canadensis_, _Tamias amoenus_, _Thomomys talpoides_ (_douglasii_ group),
_Phenacomys intermedius_, _Microtus richardsoni_ and _Ochotona
princeps_. In each of these species little or no subspecific variation
has occurred between the populations in the Cascades of Washington 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 definite 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 Cascades 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 Wisconsin 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,
Dalquest and Scheffer (1944: 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 of many
other species would show that they crossed this bridge. However, study
of additional species shows that for them the Simcoe 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 barrier to many forms that
might otherwise have utilized it. The forms that did cross on this
bridge are species known to be 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.

  [Illustration: FIG. 19. Pend Oreille River (or Clark Fork of the
  Columbia) from a point near Newport, Washington, looking south,
  June 13, 1938. (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
relatively minor barrier in northeastern Washington. The mountains 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 Cascades, 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 was rapid as compared 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 postglacial
movements and then only when conditions 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 deglaciated
area. Yet, the Pacific Coastal Fauna actually moved northward along the
coast only to the Fraser River area, and slightly farther in the

Possibly the two glaciers retreated at different times. The Wisconsin
Glacier was a piedmont glacier, fed 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 Wisconsin 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 show
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 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
timber-line 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 because 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 surrounding 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 indicated 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 amoenus caurinus_            _Tamias amoenus ludibundus_
  _Phenacomys intermedius oramontis_   _Phenacomys intermedius oramontis_
  _Clethrionomys gapperi nivarius_     _Clethrionomys gapperi 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 Olympics and of Mt.
Rainier are 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 between _californicus_ (_occidentalis_) and
_gapperi_. The principal difference between the alpine forms is the
pallor of _nivarius_. This pallor of mammals in general from the Olympic
Mountains is noteworthy, but in the red-backed mouse is exceptionally
noticeable. This pallor is discussed beyond in the paragraphs dealing
with differentiation. Mention should be made here of _Myotis keenii_.
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
terrestrial mammals.

The route of the pocket gopher (_Thomomys_) in emigrating from the
Cascades to the Olympics (Dalquest and Scheffer, 1944: 310), was over
the outwash 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 postglacial 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 remaining today are the unforested
remnants of the outwash aprons (see Dalquest and Scheffer, 1942: 69) and
possess several species of alpine plants, notably the shooting star,
camas, and bear grass.

If 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 narrow 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.
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 surprising
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 restricted
to alpine habitat but occurs throughout the Olympic Peninsula. Its
nearest relative is _S. b. bendirii_ of the rest of western Washington.
_S. b. albiventer_ differs from _bendirii_ only in possessing a
partially white ventral surface. We can only conclude that the white
belly of _albiventer_ is a mutation that the local environment has
favored and that the characters have, therefore, spread through the
population on the Olympic Peninsula. Occasional specimens are taken with
dark bellies characteristic of _bendirii_ (Jackson, 1928: 199).

The Olympic marmot, _Marmota olympus_, 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
postglacial time. The pocket gopher of southwestern 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.

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

    _Scapanus townsendii_              _Citellus beecheyi_
    _Scapanus orarius_                 _Tamias townsendii_
    _Sorex trowbridgii_                _Sciurus griseus_
    _Sorex vagrans_                    _Glaucomys sabrinus_
    _Canis lupus_                      _Castor canadensis_
    _Felis concolor_                   _Microtus townsendii_
    _Lynx rufus_                       _Microtus oregoni_
    _Mephitis mephitis_                _Ondatra zibethicus_
    _Spilogale gracilis_               _Zapus princeps_
    _Procyon lotor_                    _Odocoileus hemionus_
    _Ursus americanus_

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

               WASHINGTON                         OREGON

    _Sorex bendirii bendirii_          _Sorex bendirii palmeri_
    _Sorex obscurus setosus_           _Sorex obscurus bairdi_
    _Neotoma cinerea occidentalis_     _Neotoma cinerea fusca_
    _Peromyscus maniculatus austerus_  _Peromyscus maniculatus rubidus_
    _Clethrionomys californicus        _Clethrionomys californicus
        occidentalis_                      californicus_
    _Microtus longicaudus macrurus_    _Microtus longicaudus abditus_
    _Aplodontia rufa rufa_             _Aplodontia rufa pacifica_

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

    _Vulpes fulva_                     _Phenacomys 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 favorable but
uninhabited territory on the north.

Seemingly the pre-Vashon faunas of western Oregon and Washington 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 crossing 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

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 _Peromyscus 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. Seemingly 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

  [Illustration: FIG. 20. Rocky bluff along north bank of the Columbia
  River near Lyle. Washington. March 20. 1939. Habitat of Beechey ground
  squirrel and yellow-bellied marmot. (Fish and Wildlife Service photo
  by Victor B. Scheffer. No. 640.)]

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 and one
resident claims to have first brought it to the island. Two different
persons claim credit for introducing the Townsend chipmunk on Orcas
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 _Microtus townsendii_. These
species, at least, probably reached the islands at an early time. The
two last named are now subspecifically distinct from their mainland
relatives. Other mammals which probably were established before the
arrival of the white man include 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 Washington 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
washingtoni_, is related to, but specifically distinct from _Citellus
townsendii_ 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 Columbian 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
probably not so cold as to exterminate these species. On the other hand,
some species that are now abundant on the Columbian Plateau seem to 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 1905 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 mammals 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 1938. It had been the
habitat of owls, bats, and primitive man. The floor of the cave was
buried under from one to three feet of bat guano, much of which had
been hauled away for fertilizer. 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
between 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
surprising, 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 invasion 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 prevents 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 accomplished 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.


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 elements,
protection from enemies and safe breeding places. The factors may be of
an inorganic nature, such as climate, soil and exposure 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 distribution of the species
have changed much. With changes in environment, rare species may become
common; common and widespread 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 components 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 animals 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 occurs, 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 addition to morphological characters.

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

      1. The number of young produced by a subspecies is greater than the
      carrying capacity of the land they occupy, at least at certain
      times or in some years.

      2. Population pressure results, with a tendency of individuals to
      emigrate outwards, 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 subspecies genetically "pure."

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

      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 invasion of
_Citellus beecheyi_ and _Lepus californicus_ in Washington, and the
eastern cottontail in western Washington and the domestic rabbit in the
San Juan Islands. The volume of the "wave of population 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 adaptation to restricted
environments; each race lives only where local conditions 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 differentiation 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 in a small area the topography is highly varied. The
range of the one subspecies, _Tamias amoenus amoenus_, to the southeast
of Washington is larger than the combined ranges of all six races
occurring in Washington but, so far as I can see, the topography and
environment 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, principally
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
barriers 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-Wisconsin
glaciation was of far greater duration than the Recent. Presumably the
mammalian fauna had, from a distributional standpoint, reached a
relatively stable condition. The descent of the Vashon-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.

SCAPANUS ORARIUS.--This species probably had a history similar to 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 Washington. 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 _Scapanus_. 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 because _Neurotrichus_ has no
counterpart to compete with it in the south, whereas _Scapanus orarius_
must compete in northern California 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 Washington 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 isolated 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 Columbia 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 southward postglacial movement. Had the full species _cinereus_
been a preglacial resident of western Washington we would expect
_streatori_ 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 Washington since pre-Wisconsin

SOREX TROWBRIDGII.--This shrew is a typical Pacific coastal species with
an extensive range along the Pacific Coast south of Washington. 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 movements have been slight, as is typical of Pacific Coastal
species. The race _destructioni_ 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 OBSCURUS.--The history of this shrew of alpine predilection
probably corresponded closely to that of _Sorex cinereus_. The
derivation of the dark, long-tailed, coastal race (_S. o. setosus_) from
the smaller, paler, inland race (_obscurus_) probably occurred before
Vashon-Wisconsin Time. _Sorex o. setosus_ is one of a complex of races
distributed along the Pacific Coast from Alaska to California.

SOREX PALUSTRIS.--This species has a wide range in North America 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 Puget Bridge in early Recent Time.

SOREX BENDIRII.--This Pacific Coastal species probably had a history
very similar to that of _Neurotrichus_ and _Scapanus orarius_.

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

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 _carissima_, 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 extended 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 developed in the
southwest, entered eastern Washington from Oregon.

MYOTIS THYSANODES.--In Washington this species has been recorded 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 NOCTIVAGANS.--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 Coastal race of
the species. Presumably it extended its range westward across the
Cascades in early post-Pleistocene Time.

LASIURUS CINEREUS.--No speculation as to the distributional history of
the hoary bat seems justified at present.

CORYNORHINUS RAFINESQUII.--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 _intermedius_ probably
invaded eastern Washington from eastern Oregon in the Recent.

ANTROZOUS PALLIDUS.--This species strays into eastern Washington 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 (_cinnamomum_) were
probably 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_ indicates
it to have invaded Washington from British Columbia since the

The apparent absence of grizzly bears from the southern Cascades and
western Washington may indicate their absence from these areas
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 southwestern Washington
during Vashon Time but did not develop racial characters, or that the
Columbia was crossed so frequently that genetic differences were
dispersed throughout the entire population.

Reasons why the second hypothesis is inadequate are: (1) Raccoons range
but little north of the state of Washington, both east and west of the
Cascades. (2) Raccoons of western Washington and the area about San
Francisco Bay, California, are as much alike as are raccoons from
southwestern Washington and northwestern Oregon. It is thought that
raccoons, if resident in western Washington since interglacial time,
would have developed strong racial characters, and the fact that they
have not indicates that they have entered the state at a relatively
recent date.

The raccoon of eastern Washington (_excelsus_) is a member of the Great
Basin Fauna and has probably included southeastern Washington 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 highway by which emigrants could reach northeastern

MARTES CAURINA.--The earlier distributional history of the western
marten has been postulated by Davis (1939: 131-132), 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 Mountains
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 Rockies. 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-Wisconsin glaciation.

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 animals from northeastern
Washington and those from Idaho are more like _origenes_ than _caurina_,
although perhaps not typical. The animals 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-Vashon-Wisconsin Time, the dark race _streatori_
probably occurred in western Washington. The race _murica_ probably
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 _streatori_ from the northern part of its
range, at least temporarily. In the Cascades, _murica_ was likewise
forced southward. Ermines related to the northern _richardsonii_ were
forced into northern Washington and Idaho by the Wisconsin ice. They
probably were unable to live on the barren, unglaciated 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; _invicta_ 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 _gulosa_
from both _murica_ and _streatori_. The intermediate nature of _gulosa_
has been mentioned by Hall (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
_olympica_ probably evolved from _streatori_ in the Recent. It is
difficult to account for the dark race _fallenda_.

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 chipmunk, _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, eastward 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_ entered 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 _altifrontalis_ and _nevadensis_.

One is reminded here of the _douglasii_ group of _Thomomys talpoides_ in
which subspecies did not move north of Mt. Rainier in postglacial 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

One would expect the weasels from extreme northeastern Washington to be
referable to the race _oribasa_, of the Rocky Mountain Fauna. Instead
they are intermediate between that race and the Great Basin race,
_nevadensis_. Apparently _nevadensis_ was so dynamic 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.

MUSTELA 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 evidence for this. Minks from Idaho and
adjacent parts of British Columbia are distinctly less reddish than
minks from the area about Puget Sound, as noted by Davis (1939: 138).

GULO 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 during 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 extensive 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 regarding 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 (_latifrons_)
seems to be a coastal race, isolated in southwestern Washington and
western Oregon during Vashon time. The eastern race, _saxatilis_, 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 occurring 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 northward, in the
deglaciated area of western Washington, after the retreat of the ice.
Another race (_notata_) was probably isolated in the southeastern
Cascades and adjacent Oregon. _M. m. hudsonica_ of the Rocky Mountain
Fauna entered the northeastern part of Washington after the ice
retreated from there. A race of 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, _spissigrada_
and _notata_, both of which probably developed in Washington during
Vashon Time, occupy limited ranges in adjacent Oregon (Bailey, 1936:

TAXIDEA TAXUS.--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 (1944: 17). Pleistocene
remains, referable to this race, have been found in Franklin County.

VULPES FULVA.--The red fox of the Cascades was probably isolated there
during Vashon Time by glacial ice. Its range extends 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 southeastern Washington would be
assumed to have long been a resident of the area.

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

MARMOTA MONAX.--The woodchuck invaded northeastern Washington 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_

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 between it and its
relatives in Oregon indicate a considerable period of isolation but one
far shorter than the period during which _washingtoni_ is presumed to
have been isolated from _townsendii_.

CITELLUS COLUMBIANUS.--The Columbian ground squirrel might have been
forced southward in the Rocky Mountain area by the Wisconsin glaciation,
might have lived in southeastern Washington since then, and might have
invaded northeastern Washington in the Recent with other species of the
Rocky Mountain Fauna.

CITELLUS BEECHEYI.--This ground squirrel is known to have entered
Washington about 1915 from Oregon.

CITELLUS SATURATUS.--The mantled ground squirrel of the Cascades
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 relatively 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

The least chipmunk of the Columbian Plateau is thought to be racially
distinct from its relatives in the Yakima Valley and eastern 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 diurnal and natural
selective factors for color possibly operate more strongly on animals
active by day than on nocturnal animals. Yellow pine chipmunks are
neither forest nor desert inhabitants. Indeed, dense forests or open
deserts serve as barriers to their distribution. They prefer brush
lands, open woods, and other habitats where there is food and cover but
abundant sunlight. In such habitats they are almost independent of
altitude, temperature and humidity. 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
temperatures, 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-Wisconsin
glaciers broke this arc into two parallel geographic ranges, the Rocky
Mountains and the Cascade-Sierra Nevada chain, with a desert area

Almost every species of forest-dwelling animal had its range separated
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 chipmunk to altitude and climate and its
unique habitat requirements cause it to meet radically different natural
selective factors. 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 Pass, in the mountains to the west,
would have to contend with: marten, black bear, goshawk, bald eagle and
other alpine predators. At the present time, 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 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
result 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 inhospitable 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, 1929; 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 southern British Columbia. Between these
two fingers of the range of _luteiventris_ another race (_canicaudus_)
is found. This race occupies a more lowland area than does
_luteiventris_. The range of _luteiventris_ 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-Wisconsin 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 Mountains south of the glacier. The northern chipmunks were
adapted to more boreal conditions and perhaps otherwise better suited to
environmental conditions of the northern Rocky Mountains. A small
population of the older established race (_canicaudus_) persisted in
lowland 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, Davis
(1939: 220) writes, "It may be that, of these two races, _luteiventris_
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 successful _luteiventris_. The maximum extent of the
glacial ice, then, found _luteiventris_ the dominant 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 northeastern Washington as far as the range of
_luteiventris_. The chipmunk 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
postglacial 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 Fraser River and,
there, isolated by habitat selection, it developed the characters of
_felix_. Population pressure later forced it eastward until the eastern
border of its range again met the range of the ancestral race,

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) included Mt. Rainier in the range of

Briefly summarized, the probable pre-Vashon-Wisconsin distribution 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
_luteiventris_ 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_ was 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, uncertainty 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 independently the slightly paler color that
separates _cooperi_ from _townsendii_. 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_.

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 recent times.

TAMIASCIURUS HUDSONICUS.--The two species of red squirrels, _T.
hudsonicus_ and _T. douglasii_, are specifically distinct and probably
became differentiated in the Pleistocene when southward moving glaciers
cut in two the range of the ancestral stock. The morphological
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 westward to the Pacific, thus
occupying an area that, in pre-Vashon time, probably was occupied by

TAMIASCIURUS DOUGLASII.--The Douglas squirrel probably occupied the
coastal region of Oregon, Washington and British Columbia in pre-Vashon
Time. The descent of the ice restricted its range to southwestern
Washington and areas to the south. After the retreat of the ice it moved
northward somewhat but, like other coastal species, the movement was
slow. Meanwhile _hudsonicus_ from the Rocky Mountain Fauna, had spread
to the coast of British Columbia.

GLAUCOMYS SABRINUS.--This flying squirrel is a plastic species. It
inhabits all of the forested parts of Washington. The distributional
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 _oregonensis_ occupies Washington and Oregon west of the
Cascades; _fuliginosus_ occupies the Cascades of Washington, Oregon and
southern British Columbia; _columbiensis_ occupies the interior valleys
of British Columbia and adjoining Washington; _latipes_ occupies the
northern Rocky 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 _oregonensis_ from other subspecies
found in Washington are relatively great. This lowland 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-Vashon 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 established _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 related 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 _bangsi_ 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 adjacent British Columbia were occupied by _latipes_, derived from

THOMOMYS TALPOIDES.--Views as to the probable historical distribution of
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 Washington. 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 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
_acqualidens_ and those between the Blue Mountains and the Columbia
River intergraded with the desert race, _devexus_. These intergrades,
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 multiplied 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 (_melanops_) were isolated from those in the area about Puget
Sound. Six distinct races originated on the isolated prairie
(_glacialis_, _tacomensis_, _pugetensis_, _yelmensis_, _tumuli_,

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 northeastern 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.

PEROGNATHUS PARVUS.--Three races of the pocket mouse occur in
Washington. Two of these (_lordi_ and _columbianus_) occur on the
Columbian Plateau. Like 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 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 washingtoni_, and _townsendii_.

DIPODOMYS ORDII.--This kangaroo rat enters the desert area of
southeastern Washington from Oregon. It may be expected eventually 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 Washington 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 glaciation and
developed its characters while isolated. The other race, _leucodonta_,
was probably widely spread in Wisconsin Time. Beavers are present in
Moses Lake, in almost the center of the Columbian 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 resembles
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 Washington from Oregon at a relatively
recent date. Within the last ten years it has extended its range into
the Okanogan Valley in British Columbia.

PEROMYSCUS MANICULATUS.--Six subspecies of _Peromyscus maniculatus_
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 the Columbia
River that is politically within the state of Washington. Another
(_hollisteri_) is restricted to certain islands in northern Puget Sound
and obviously has become subspecifically differentiated in postglacial
time. The remaining four subspecies, namely _oreas_, _austerus_,
_artemisiae_ and _gambelii_, have extensive geographic ranges. These
subspecies are not confined to their geographic 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. Geographically separated populations
of "wild caught" mice of the subspecies _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 Debzhansky
(1937). They have been intensively studied by Sumner (1917 A, B) and
Dice. An especially important paper by the latter author (1940)
summarizes much of the available information on speciation in
_Peromyscus_ and clearly discusses the microgeographic races of

The numerous microgeographic races 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 not 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

The work of Murie and Murie (1931: 200-209, 1932: 79) is enlightening in
this respect. These authors found that mice residing in a locality
tended to remain in that locality; individuals trapped and marked were
retaken in the same locality a year later. Individuals 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 developed 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 _Peromyscus 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 where mice of the same
species are already established. This may partly account for the
microgeographic races of deer mice in Washington.

Dice (1939: 21) 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 Washington.

Of the four subspecies of deer mice that occupy extensive geographic
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 _gambelii_, a short-tailed form
with which _oreas_ intergrades, are easily distinguished. In eastern
Washington two short-tailed subspecies, _gambelii_ 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 Gap 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 Columbian
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_-like characters in 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 subspecies 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 Skykomish 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
reproduce. 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 subspecies 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 with a more arboreal existence in dark,
coniferous forests. Our observations show that _oreas_ is, to a large
extent, arboreal. Traps nailed to tree trunks six to eight feet from the
ground and set for flying squirrels often caught _oreas_ in the
Skykomish 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 _oreas_ was found in this trap the next morning.
Svihla (1933: 13) relates how a specimen of _oreas_ that escaped from a
live trap took refuge in a tree when pursued. We have set numerous traps
for flying squirrels in the area about Puget Sound. As far as memory and
field notes serve, we 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 capable 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 follows: _P. m.
gambelii_ occupies southeastern Washington and intergrades normally with
_oreas_ in the eastern Cascade Mountains and with _artemisiae_ at the
northern edge of the Columbian Plateau; _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 _gambelii_; _oreas_ occupies most of
western Washington, intergrades with its neighbors _artemisiae_ and
_gambelii_ to the east, but does not intergrade 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 Northwest
that occurred during and since the last Pleistocene glaciation. We
reconstruct these changes as follows. In the Puyallup interglacial
cycle, just previous to the last (Vashon-Wisconsin) continental
glaciation, clines, 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 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 existed inland. Small, pale,
short-tailed mice of the Great Basin became increasingly larger, darker,
and longer-tailed to the north, reaching a maximum in the spruce forests
of northern British Columbia.

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

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. m. rubidus_) do become larger, redder and longer-tailed to the
north. The climax of this cline is now reached in _austerus_, of the
Puget Sound area of Washington. The cline is not continued farther
northward because the range of _oreas_ is encountered.

The advance of the Vashon-Wisconsin ice from the north probably 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, conditions in the fore
of the glacier exterminated the short-tailed individuals but favored the
long-tailed mice, since the latter originally were adapted to a boreal
environment. The climax of the ice advance found the glaciers just
within the political limits of the United States 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. Long-tailed northern mice
advancing before it reached the Columbia River. This glacially swollen
stream served as a natural barrier and prevented their southward
extension. At this time the northern mice had traversed more than half
the length of the original western cline. The northern mice, originating
in a boreal habitat a thousand miles away, were unable to interbreed
with the southern mice and such individuals as might have crossed the
Columbia River never became 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
elimination 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 Mountains.

In eastern Washington the Wisconsin Glacier advanced south to the
northern edge of the Columbian Plateau. Northern mice advancing before
it probably did not survive on the treeless plateau but existed in
forested areas of northern Idaho and driftless areas of northern
Washington. No natural barriers separated the northern mice from the
pale, short-tailed forms. The nonuniform topography perhaps allowed more
mingling of the two types where climatic 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 establishment 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 presented 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 become
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. This 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, _austerus_
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 Wahkiakum County,
where the range of _austerus_ starts. At the border of 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 Wahkiakum 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

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 _hollisteri_, a race similar to _oreas_ in
color, body size and cranial characters and probably 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: 461).
When _oreas_ first occupied the area about Puget Sound (this area is
occupied by _austerus_ today) some individuals probably reached the San
Juan Islands soon thereafter. Contact between _oreas_ and _hollisteri_
is now prevented by the presence of _austerus_ between their ranges. Of
course, a water barrier separates _austerus_ from _hollisteri_ but
_austerus_ does occur in pure form on some islands in Puget Sound (not
on any of the San Juan Islands), for instance, on Whidby Island. One
wonders why _austerus_ 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_-type
of mouse occurs in the lowlands and a long-tailed, _oreas_-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 ever 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 deglaciated 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. alpinus_, isolated in a mountainous area of
British Columbia. This seems to be a form derived from _oreas_-type

With the reëstablishment of soil and forests on the deglaciated land of
British Columbia north of Washington, a northward extension 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 intergraded 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. The
deer mice of the San Juan Islands are all referred to the subspecies
_hollisteri_. Only Puget Island, Wahkiakum County, is considered to be
inhabitated by _rubidus_. The boundary between the ranges of _austerus_
and _oreas_ is sharply defined and further collecting by resident
naturalists should result in detailed mapping of the boundary.

The ranges and distribution of the deer mice of eastern Washington 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 Mountains. 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, however, 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 are 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 northward 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 _artemisiae_ may be considered to be
intermediate between _gambelii_ and the darker, northern race. Thus the
name _artemisiae_ becomes available for the mice of the 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 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 Washington, 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 _oreas_ 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 Washington and probably also
farther east along the Wisconsin drift border, and that type that occurs
farther south in Idaho and seems to constitute 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 texanus 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 Mountains of Washington
will be referable to the southern form.

NEOTOMA CINEREA.--The wood rat found over most of Washington. _Neotoma
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: 174). In Washington _occidentalis_ occupies 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 Washington 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 (1939: 308) gives an excellent analysis of the emigrational
history of the American species, and points out the close relationship
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 Mountain course southward and
worked eastward across Canada while the other (_californicus_ group)
followed down the Pacific Coast. Davis states, "This southward
extension of range in America probably 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 snowshoe 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
_Clethrionomys_ 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 Washington, 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
_Clethrionomys californicus_, while the _gapperi_ group contains _C.
gapperi_ and its races, including _caurinus_, and possible other

Possibly intergradation occurs between _C. californicus occidentalis_
and _C. gapperi_. Nevertheless, I fail to find evidence of such
intergradation. I have taken _C. gapperi nivarius_ and _C. californicus
occidentalis_ within ten miles of each other, but each retained its
distinctive characters with no evidence of intergradation. In the case
of _C. gapperi saturatus_ and _C. californicus 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, _occidentalis_
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 _occidentalis_ 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, _montanus_ 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 came together, _pennsylvanicus_ occurs
with races of _montanus_ that are most unlike it.

_Microtus pennsylvanicus kincaidi_ closely resembles races of _Microtus
montanus_ that occur in southern Oregon, California, and Nevada. It is
larger, darker, and longer-furred than _Microtus pennsylvanicus
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 _pennsylvanicus_ 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 Washington 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 montanus_
from the Blue Mountains crossed the Simcoe Bridge to the Yakima Valley
and the eastern Cascade Mountains. The closing of the Simcoe Bridge
isolated these mice, which 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 Cascade 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
_montanus_ 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 occurring 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 LONGICAUDUS.--The later distributional history of the
long-tailed meadow mouse is not yet clear.

MICROTUS 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 Mountains seems to
be have been forced southward into Washington and Oregon by the descent
of the Vashon-Wisconsin glaciers where it became isolated from the water
rat of the northern Rocky Mountains. After the retreat of the ice, both
forms have moved northward. 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_.

LAGURUS CURTATUS.--The distribution of this species of the Great Basin
Fauna has probably changed little if any since the late Pleistocene.

ONDATRA ZIBETHICUS.--The distribution of the two races of muskrat 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 adaptable, 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-Wisconsin 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 _kootenayensis_ of the adjacent Rocky Mountains moved
eastward through northeastern 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.

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 Mountains. 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 weathering, 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 weathering, becomes rusty, reddish brown.

In Washington the two races maintain their distinctiveness because
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

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 occupied 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

The distinctness of _brunnescens_ and _fenisex_ is maintained by
selective factors of the habitats they occupy but the zone of
intergradation between the two is broad. A tendency towards paleness is
noticeable in specimens from as far west as the eastern side of Mount

The pikas in the Rocky Mountain area have given rise to a number of
races. The pale, small race of northern Idaho, _cuppes_, entered
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

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 developed. The descent of the Vashon-Wisconsin
glaciers found _washingtonii_ 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_ and _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 _washingtonii_
rather than _cascadensis_ also supports this view. The snowshoe 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 other species.

Following the retreat of the glaciers, _washingtonii_ reinvaded western
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 northeastern Washington from
adjacent areas to the south. Another race, _columbiensis_, spread from
the interior valleys of British Columbia to the western part of
northeastern Washington in probably relatively recent times. The
post-Wisconsin history of the distribution 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
blacktails 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
sagebrush areas between the Columbia River and the Cascade Mountains.
When Couch, in 1927, published his account, he predicted that the
blacktail 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, spectacular, 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 history.

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 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 the 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 _virginianus_ probably diverged from a common stock in the

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 Wisconsin 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-Wisconsin time, but has since largely given way to the
black-tailed and mule deer. The race _O. v. leucurus_ now occupies 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 Pacific Coastal Fauna.

The inland race, _ochrourus_, 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 Washington as a
winter migrant from the north. It was probably of more regular and
extensive occurrence in the past.

ANTILOCAPRA AMERICANA.--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-Wisconsin times. After the
retreat of the ice, these sheep (_californiana_) moved northward
slightly, both in the Cascades and on the Columbian Plateau. Remains of
sheep are to be found with human artifacts 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 Washtuckna 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 previous 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.


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 according to Scheffer (1942). A few other minor changes are

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 are 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 are usually given for each subspecies. All measurements,
unless otherwise stated, are in millimeters and weights are in grams.
Specimens are adult, unless otherwise 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 are 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 of 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
approaches that of another are listed as "marginal occurrences" in the
paragraph on the distribution of the race to which they are best
referred. Marginal occurrences are listed from north to south and from
west to east. The authority for the record, if published, is given. When
the record is based on specimens examined by the author, the collection
containing the specimen is indicated as follows:

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

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

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

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

      (U.S.N.M.) United States National Museum, Washington, D. C.
      Specimens from the Biological Surveys Collection are included

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

      (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, 9 kinds (opossum, eastern gray squirrel,
fox squirrel, 3 kinds of Old World rats, house mouse, nutria, and
eastern cottontail) have been introduced into Washington 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 orders, 20 families, 58 genera and 101 full species.



            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     }  Gibbs shrew-mole
       and Burgner                              }

    Genus =Scapanus= Pomel--western American moles
      _Scapanus townsendii_ (Bachman)              Townsend mole
      _Scapanus orarius orarius_ True           }
      _Scapanus orarius yakimensis_ Dalquest    }  coast mole
        and Scheffer                            }
      _Scapanus orarius schefferi_ Jackson      }

  Family SORICIDAE--shrews
    Genus =Sorex= Linnaeus--long-tailed shrews
      _Sorex cinereus cinereus_ Kerr            }
      _Sorex cinereus streatori_ Merriam        }  cinereous shrew
      _Sorex merriami merriami_ Dobson             Merriam shrew
      _Sorex trowbridgii trowbridgii_ Baird     }
      _Sorex trowbridgii destructioni_ Scheffer }  Trowbridge shrew
        and Dalquest                            }
      _Sorex vagrans vagrans_ Baird             }
      _Sorex vagrans monticola_ Merriam         }  wandering shrew
      _Sorex obscurus obscurus_ Merriam         }
      _Sorex obscurus setosus_ Elliot           }  dusky shrew
      _Sorex palustris navigator_ (Baird)          mountain water-shrew
      _Sorex bendirii bendirii_ (Merriam)       }
      _Sorex bendirii albiventer_ Merriam       }  Bendire water-shrew

    Genus =Microsorex= Coues--pigmy shrew
      _Microsorex hoyi washingtoni_ Jackson        pigmy shrew

            Order CHIROPTERA--bats

  Family VESPERTILIONIDAE--vespertilionid bats
    Genus =Myotis= Kaup--mouse-eared bats
      _Myotis lucifugus carissima_ Thomas       }
      _Myotis lucifugus alascensis_ Miller      }  big myotis
      _Myotis yumanensis sociabilis_            }
        H. W. Grinnell                          }
      _Myotis yumanensis saturatus_ Miller      }  Yuma myotis
      _Myotis keenii keenii_ (Merriam)             Keen myotis
      _Myotis evotis evotis_ (H. Allen)         }  long-eared myotis
      _Myotis evotis pacificus_ Dalquest        }
      _Myotis thysanodes thysanodes_ Miller        fringe-tailed myotis
      _Myotis volans longicrus_ (True)          }  hairy-winged myotis
      _Myotis volans interior_ Miller           }
      _Myotis californicus caurinus_ Miller     }  California myotis
      _Myotis californicus californicus_        }
       (Audubon and Bachman)                    }
      _Myotis 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
      _Corynorhinus rafinesquii townsendii_     }  long-eared bat
       (Cooper)                                 }
      _Corynorhinus rafinesquii intermedius_    }
       H. W. Grinnell                           }

    Genus =Pipistrellus= Kaup--pipistrelles
      _Pipistrellus hesperus hesperus_ (H. Allen)  western pipistrelle

    Genus =Eptesicus= Rafinesque--serotine bats
      _Eptesicus fuscus bernardinus_ Rhoads        big brown bat

    Genus =Lasiurus= Gray--hairy-tailed bats
      _Lasiurus cinereus cinereus_ (Beauvois)      hoary bat

    Genus =Antrozous= H. Allen--nyctophiline bats
      _Antrozous pallidus cantwelli_ Bailey        pallid bat

            Order CARNIVORA

  Family URSIDAE--bears
    Genus =Ursus= Linnaeus--bears
      _Ursus americanus altifrontalis_ Elliot   }
      _Ursus americanus cinnamomum_             }  black bear
       (Audubon and Bachman)                    }
      _Ursus chelan_ Merriam                       grizzly bear

  Family PROCYONIDAE--raccoons and allies
    Genus =Procyon= Storr-raccoons
      _Procyon lotor psora_ Gray                }
      _Procyon lotor excelsus_ Nelson and       }  raccoon
       Goldman                                  }

  Family MUSTELIDAE--weasels and allies
    Genus =Martes= Pinel--martens and fisher
      _Martes caurina caurina_ (Merriam)        }  western marten
      _Martes caurina origenes_ (Rhoads)        }
      _Martes pennanti_ (Erxleben)                 fisher

    Genus =Mustela= Linnaeus--weasels, ferrets and minks
      _Mustela erminea invicta_ Hall            }
      _Mustela erminea fallenda_ Hall           }
      _Mustela erminea olympica_ Hall           }  ermine
      _Mustela erminea streatori_ (Merriam)     }
      _Mustela erminea gulosa_ Hall             }
      _Mustela erminea murica_ (Bangs)          }
      _Mustela frenata washingtoni_ (Merriam)   }
      _Mustela frenata altifrontalis_ Hall      }
      _Mustela frenata effera_ Hall             }  long-tailed weasel
      _Mustela frenata nevadensis_ Hall         }
      _Mustela vison energumenos_ (Bangs)          mink

    Genus =Gulo= Pallas--wolverines
      _Gulo luscus luteus_ Elliot                  wolverine

    Genus =Lutra= Brisson--river otters
      _Lutra canadensis pacifica_ Rhoads        }
      _Lutra canadensis vancouverensis_ Goldman }  river otter

    Genus =Enhydra= Fleming--sea otter
      _Enhydra lutris nereis_ (Merriam)            sea otter

    Genus =Spilogale= Gray--civet cats or spotted skunks
      _Spilogale gracilis saxatilis_ Merriam    } civet cat or
      _Spilogale gracilis latifrons_ Merriam    }  spotted skunks

    Genus =Mephitis= Geoffroy and Cuvier--striped skunks
      _Mephitis mephitis hudsonica_ Richardson  }
      _Mephitis mephitis major_ (Howell)        }
      _Mephitis mephitis notata_ (Howell)       }  striped skunk
      _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
      _Vulpes fulva cascadensis_ Merriam           red fox

    Genus =Canis= Linnaeus--coyote, wolves and dogs
      _Canis latrans testes_ Merriam            }
      _Canis latrans incolatus_ Hall            }  coyote
      _Canis lupus fuscus_ Richardson              wolf
      _Canis familiaris_ Linnaeus                  dog

  Family FELIDAE--cats
    Genus =Felis= Linnaeus--true cats
      _Felis concolor oregonensis_ Rafinesque   }  cougar or
      _Felis concolor missoulensis_ Goldman     }   mountain lion

    Genus =Lynx= Kerr--lynxes and bobcats
      _Lynx canadensis canadensis_ Kerr            Canadian lynx
      _Lynx rufus fasciatus_ Rafinesque         }
      _Lynx rufus pallescens_ Merriam           }  bobcat

            Order PINNIPEDIA--seals and walruses

  Family OTARIIDAE--eared seals
    Genus =Zalophus= Gill--Californian sea lions
      _Zaluphus californianus_ (Lesson)            California sea lion

    Genus =Eumetopias= Gill--Steller sea lion
      _Eumetopias jubata_ (Schreber)               Steller sea lion

    Genus =Callorhinus= Gray--Alaska fur seal
      _Callorhinus ursinus cynocephalus_
       (Walbaum)                                   Alaska fur seal

  Family PHOCIDAE--hair seals
    Genus =Phoca= Linnaeus--hair seals
      _Phoca vitulina richardii_ (Gray)            hair seal

            Order RODENTIA--rodents

  Family SCIURIDAE--squirrels and allies
    Genus =Tamias= Illiger--chipmunks
      _Tamias minimus scrutator_ (Hall and      }
       Hatfield)                                }  least chipmunk
      _Tamias minimus grisescens_ (Howell)      }
      _Tamias amoenus caurinus_ (Merriam)       }
      _Tamias amoenus felix_ Rhoads             }
      _Tamias amoenus ludibundus_ (Hollister)   }
      _Tamias amoenus affinis_ Allen            }  yellow-pine chipmunk
      _Tamias amoenus canicaudus_ (Merriam)     }
      _Tamias amoenus luteiventris_ Allen       }
      _Tamias ruficaudus simulans_ (Howell)        red-tailed chipmunk
      _Tamias townsendii townsendii_ Bachman    }
      _Tamias townsendii cooperi_ Baird         }  Townsend chipmunk

    Genus =Marmota= Blumenbach--marmots
      _Marmota monax petrensis_ Howell             woodchuck
      _Marmota flaviventris avara_ (Bangs)         yellow-bellied marmot
      _Marmota caligata cascadensis_ Howell        hoary marmot
      _Marmota olympus_ (Merriam)                  Olympic marmot

    Genus =Citellus= Oken--ground squirrels
      _Citellus townsendii townsendii_ (Bachman)   Townsend ground
      _Citellus washingtoni_ Howell                Washington ground
      _Citellus columbianus columbianus_ (Ord)  }
      _Citellus columbianus ruficaudus_ Howell  }  Columbian ground
      _Citellus beecheyi douglasii_
       (Richardson)                                Beechey ground squirrel
      _Citellus lateralis tescorum_ (Hollister) }
      _Citellus lateralis connectens_ (Howell)  }  golden-mantled ground
      _Citellus saturatus_ (Rhoads)             }   squirrel

    Genus =Sciurus= Linnaeus--tree squirrels
      _Sciurus griseus griseus_ Ord                western gray squirrel
      _Sciurus carolinensis hypophaeus_ Merriam    eastern gray squirrel
      _Sciurus niger_ Linnaeus, subsp.?            fox squirrel

    Genus =Tamiasciurus= Trouessart--red squirrels
      _Tamiasciurus hudsonicus streatori_       }
       (Allen)                                  }
      _Tamiasciurus hudsonicus richardsoni_     }  red squirrel
       (Bachman)                                }
      _Tamiasciurus douglasii douglasii_        }
       (Bachman)                                }  Douglas squirrel

    Genus =Glaucomys= Thomas--American flying squirrels
      _Glaucomys sabrinus oregonensis_          }
       (Bachman)                                }
      _Glaucomys sabrinus bangsi_ (Rhoads)      }  northern flying
      _Glaucomys sabrinus columbiensis_ Howell  }   squirrel
      _Glaucomys sabrinus latipes_ Howell       }
      _Glaucomys sabrinus fuliginosus_ (Rhoads) }

  Family HETEROMYIDAE--pocket mice, kangaroo mice and kangaroo rats
    Genus =Perognathus= Weid--pocket mice
      _Perognathus parvus parvus_ (Peale)       }  great basin pocket
      _Perognathus parvus lordi_ (Gray)         }   mouse
      _Perognathus parvus columbianus_ Merriam  }

    Genus =Dipodomys= Gray--kangaroo rats
      _Dipodomys ordii columbianus_ (Merriam) Ord kangaroo rat

  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 fuscus_ Merriam       }
      _Thomomys talpoides yakimensis_ Hall      }
       and Dalquest                             }
      _Thomomys talpoides shawi_ Taylor         }
      _Thomomys talpoides immunis_ Hall and     }
       Dalquest                                 }  northern pocket
      _Thomomys talpoides limosus_ Merriam      }   gopher
      _Thomomys talpoides douglasii_            }
       (Richardson)                             }
      _Thomomys talpoides glacialis_ 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_ Jewett and    }  beaver
       Hall                                     }

  Family MURIDAE--rats and mice
    Genus =Onychomys= Baird--grasshopper mice
      _Onychomys leucogaster fuscogriseus_      }  northern grasshopper
       Anthony                                  }   mouse

    Genus =Reithrodontomys= Giglioli--American harvest mice
      _Reithrodontomys megalotis megalotis_     }  western harvest
       (Baird)                                  }   mouse

    Genus =Peromyscus= Gloger--white-footed mice
      _Peromyscus maniculatus oreas_ Bangs      }
      _Peromyscus maniculatus hollisteri_       }
       Osgood                                   }
      _Peromyscus maniculatus austerus_ (Baird) }
      _Peromyscus maniculatus rubidus_ Osgood   }  deer mouse
      _Peromyscus maniculatus gambelii_ (Baird) }
      _Peromyscus maniculatus artemisiae_       }
       (Rhoads)                                 }

    Genus =Neotoma= Say and Ord--wood rats
      _Neotoma cinerea occidentalis_ Baird      }  bushy-tailed wood rat
      _Neotoma cinerea alticola_ Hooper         }

    Genus =Synaptomys= Baird--lemming mice
      _Synaptomys borealis wrangeli_ Merriam       northern lemming mouse

    Genus =Phenacomys= Merriam--heather voles and tree mice
      _Phenacomys intermedius intermedius_      }
       Merriam                                  }  heather vole
      _Phenacomys intermedius oramontis_        }
       Rhoads                                   }

    Genus =Clethrionomys= Tilesius--red-backed mice
      _Clethrionomys gapperi saturatus_         }
       (Rhoads)                                 }
      _Clethrionomys gapperi idahoensis_        }
       (Merriam)                                }  Gapper red-backed
      _Clethrionomys gapperi cascadensis_       }   mouse
       Booth                                    }
      _Clethrionomys gapperi nivarius_          }
       (Bailey)                                 }
      _Clethrionomys californicus               }  California red-backed
       occidentalis_ (Merriam)                  }    mouse

    Genus =Microtus= Schrank--meadow mice
      _Microtus pennsylvanicus funebris_ Dale   }  Pennsylvania meadow
      _Microtus pennsylvanicus kincaidi_        }   mouse
       Dalquest                                 }
      _Microtus montanus nanus_ (Merriam)       }  montane meadow
      _Microtus montanus canescens_ Bailey      }   mouse
      _Microtus townsendii townsendii_          }
       (Bachman)                                }  Townsend meadow
      _Microtus townsendii pugeti_ Dalquest     }   mouse
      _Microtus longicaudus halli_ Ellerman     }  long-tailed meadow
      _Microtus longicaudus macrurus_ Merriam   }   mouse
      _Microtus richardsoni arvicoloides_       }
       (Rhoads)                                 }  water rat
      _Microtus richardsoni macropus_ (Merriam) }
      _Microtus oregoni oregoni_ (Bachman)         creeping mouse

    Genus =Lagurus= Gloger--short-tailed voles
      _Lagurus curtatus pauperrimus_ (Cooper)      sagebrush vole

    Genus =Ondatra= Link--muskrat
      _Ondatra zibethicus osoyoosensis_ (Lord)  }
      _Ondatra zibethicus occipitalis_ (Elliot) }  muskrat

    Genus =Rattus= Fischer--Old World rats
      _Rattus rattus rattus_ (Linnaeus)         }  roof rat
      _Rattus rattus alexandrinus_ (Geoffroy)   }
      _Rattus norvegicus norvegicus_ (Erxleben)    Norway rat

    Genus =Mus= Linnaeus--house mice
      _Mus musculus_ Linnaeus, subsp.?             house mouse

  Family APLONTIDAE--mountain beaver
    Genus =Aplodontia= Richardson--mountain beaver
      _Aplodontia rufa rufa_ (Rafinesque)       }
      _Aplodontia rufa rainieri_ Merriam        }  mountain beaver

  Family DIPODIDAE--jumping mice and allies
    Genus =Zapus= Coues--jumping mice
      _Zapus princeps trinotatus_ Rhoads        }
      _Zapus princeps kootenayensis_ Anderson   }  big jumping mouse
      _Zapus princeps idahoensis_ Davis         }
      _Zapus princeps oregonus_ Preble          }

  Family ERETHIZONTIDAE--American porcupines
    Genus =Erethizon= Cuvier--North American porcupine
      _Erethizon dorsatum epixanthum_ Brandt    }  porcupine
      _Erethizon dorsatum nigrescens_ Allen     }

  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 fenisex_ Osgood        }  pika
      _Ochotona princeps brunnescens_ Howell    }

  Family LEPORIDAE--hares and rabbits
    Genus =Lepus= Linnaeus--hares
      _Lepus townsendii townsendii_ Bachman        white-tailed jack
      _Lepus americanus washingtonii_ Baird     }
      _Lepus americanus cascadensis_ Nelson     }  snowshoe rabbit
      _Lepus americanus pineus_ Dalquest        }
      _Lepus americanus columbiensis_ Rhoads    }
      _Lepus californicus deserticola_ Mearns      black-tailed jack

    Genus =Sylvilagus= Gray--cottontails and allies
      _Sylvilagus nuttallii nuttallii_ (Bachman)   Nuttall cottontail
      _Sylvilagus floridanus_ (Allen), subsp.?     Florida cottontail
      _Sylvilagus idahoensis_ (Merriam)            pigmy rabbit

            Order ARTIODACTYLA--even-toed ungulates

  Family CERVIDAE--deer and allies
    Genus =Cervus= Linnaeus--American elk or wapiti
      _Cervus canadensis roosevelti_ Merriam    }  elk or wapiti
      _Cervus canadensis nelsoni_ Bailey        }

    Genus =Odocoileus= Rafinesque--northern deer
      _Odocoileus virginianus ochrourus_ Bailey }
      _Odocoileus virginianus leucurus_         }  white-tailed deer
       (Douglas)                                }
      _Odocoileus hemionus hemionus_            }
       (Rafinesque)                             }  mule deer or
      _Odocoileus hemionus columbianus_         }   black-tailed deer
       (Richardson)                             }

    Genus =Alce= Gray--Moose and Old-World elk
      _Alce americana shirasi_ Nelson              moose

    Genus =Rangifer= Hamilton-Smith--reindeer and caribou
      _Rangifer arcticus montanus_ Seton-Thompson  caribou

  Family BOVIDAE--cattle, sheep, goats and allies
    Genus =Bison= Hamilton-Smith--bison
      _Bison bison oregonus_ Bailey                bison

    Genus =Ovis= Linnaeus--sheep
      _Ovis canadensis canadensis_ Shaw         }  mountain sheep
      _Ovis canadensis californiana_ Douglas    }

    Genus =Oreamnos= Rafinesque--mountain goat
      _Oreamnos americanus americanus_          }  mountain goat
       (Blainville)                             }
      _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
      _Phocoenoides 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= Lacépède--finback whales
      _Balaenoptera physalus_ (Linnaeus)           finback whale
      _Balaenoptera borealis_ Lesson               Pollack whale
      _Balaenoptera acutorostrata_ Lacépède        pike whale

    Genus =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


=Didelphis virginiana virginiana= Kerr


     _Didelphis virginiana_ Kerr. Anim. Kingd., p. 193, 1792.

     _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 and 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."

=Neurotrichus gibbsii=

Gibbs shrew-mole

_Description._--The shrew-mole is tiny, possessing a head and body
2-1/2 to 3 inches long and a tail about 1-1/2 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.

  [Illustration: FIG. 21. Gibbs shrew-mole (_Neurotrichus gibbsii
  minor_), female in captivity; Seattle, Washington, September 12,
  1939. (Fish and Wildlife Service photo by Victor B. Scheffer, No.

_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 are most
abundant in damp, shady ravines where the vegetation includes 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 Transition Life-zone
through the Canadian, and well into the Hudsonian Life-zone.

  [Illustration: FIG. 22. Distribution of the Gibbs shrew-mole in
  Washington. A. _Neurotrichus gibbsii gibbsii._ B. _Neurotrichus
  gibbsii minor._]

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 rush which ends beneath a leaf or
bit of bark where the animal becomes 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, 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. 76, 1857.

      _Neurotrichus [sic] gibbsii_ Günther, Proc. Zool. Soc. London, pl.
      42, 1880.

      _Neurotrichus Gibbsii_ True, Proc. U. S. Nat. Mus., 7 (1884):607,

      _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 Zoölogy.

      _Racial characters._--Size small; tail short; foreclaws light and
      weak as compared to those of _gibbsii_, with ventral surfaces

      _Measurements._--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.).

Genus =Scapanus= Pomel


  [Illustration: FIG. 23. Coast mole (_Scapanus orarius orarius_) left
  and Townsend mole (_Scapanus townsendii_) right; Puyallup. Washington,
  May, 1914. (Fish and Wildlife Service photo by T. H. Scheffer, No.

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

=Scapanus townsendii= (Bachman)

Townsend mole

      _Scalops townsendii_ Bachman. Jour. Acad. Nat. Sci. Philadelphia, 8
      (pt. 1):58, 1839.

      _Scapanus tow[n]sendii_ Pomel. Arch. Sci. Phys. Nat., Geneva, 9
      (ser. 4):247, 1848.

      _Scapanus Townsendii_ True. Proc. U. S. Nat. Mus., 7 (1881):607,

      _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 Yacolt (M.V.Z.).

_Remarks._--_Scapanus townsendii_ occurs only in a narrow belt extending
from southwestern British Columbia to northwestern California. 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 occasionally 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 completely 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 containing about a
cubic foot of earth. The mounds commonly are built just about as far
apart as a man can step. The general direction 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 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.

  [Illustration: FIG. 24. Distribution of the Townsend mole, _Scapanus
  townsendii_, in Washington.]

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.

=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 5-1/4 and tail
about 1-1/2 inches in length.

The coast mole occupies all of the territory 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.

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. Upthrust ridges are less commonly
built by _orarius_ than by _townsendii_.

  [Illustration: 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.)]

Jackson (1915: 62) mentions the possibility of ecologic differences
between _orarius_ and _townsendii_. There are some differences in
habitat and habits. The coast mole seems to live deeper in the 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, such as the brush-grown alder
and dogwood jungles along the stream valleys. 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 is 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 earthworms and
insects (Moore, 1933: 38). On September 29, 1939, in a ravine at
Seattle, in two baited traps set six inches apart, a creeping mouse
(_Microtus oregoni_) and a coast mole were found. The mouse had come to
the bait and been caught. The mole had seemingly 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 lay dead with its nose inside the body of
the mouse 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 Townsend moles
so killed have been found.

  [Illustration: FIG. 26. Distribution of the coast mole in Washington.
  A. _Scapanus orarius orarius._ B. _Scapanus orarius yakimensis._
  C. _Scapanus orarius schefferi._]

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 breeding season seems to
be the same as that of _townsendii_ but the number of young to a litter
may average slightly greater.

=Scapanus orarius orarius= True

      _Scapanus orarius_ True. Proc. U. S. Nat. Mus., 19:52, December 21,

      _Scapanus orarius orarius_ Jackson, N. Amer. Fauna, 38:61,
      September 30, 1915.

      _Type._--Obtained at Shoalwater (= Willapa) Bay, Pacific County,
      Washington, 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= Dalquest and Scheffer

      _Scapanus orarius yakimensis_ Dalquest and Scheffer, Murrelet,
      25:27, September 19, 1944.

      _Type._--Obtained 3/4 mile north of Union Gap, Yakima County,
      Washington, by J. A. Gray, Jr., on July 3, 1941; type in Museum of
      Vertebrate Zoölogy.

      _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, N. 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

      _Racial 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.).

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 1-3/4 inches. The upper parts are dark grayish brown and the
underparts dull gray.

Shrews of the genus _Sorex_ 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 subspecies present
occupy numerous habitats, and their ranges include almost all of the

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 nonprofessional mammalogist as
the long-tailed shrews." In Washington, 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 movements 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

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

The food of long-tailed shrews is varied. It is principally soft-bodied
insects, insect pupae, and earthworms. At times a considerable quantity
of soft vegetation and some seeds are eaten. Recent studies (Moore,
1940: 1942) have shown that by destroying seeds some shrews may
adversely affect the reforestation of some coniferous 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

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= Kerr

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 Washington. The cinereous shrew is
of medium size and difficult to distinguish 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
_vagrans_ and _obscurus_ it is larger. The relatively narrow rostrum of
_cinereus_ also serves 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, confined to
mountainous areas. The coastal race, _streatori_, seems to be confined
to the humid area.

=Sorex cinereus cinereus= Kerr

      _Sorex arcticus cinereus_ Kerr. Anim. Kingd., p. 206, 1792.

      _Sorex cinereus cinereus_ Jackson. Jour. Mamm., 6: 56, February 9,

      _Type._--None. Name based on the account of a shrew seen at Fort
      Severn, Canada, by J. R. Forster in 1772.

      _Racial characters._--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) southward to Conrad Meadows (Jackson, 1928: 49). There
      is a specimen in the collection of the California Academy of
      Sciences, from Mt. Rainier.

_Remarks._--This is the most wide-ranging subspecies of shrew, being
recorded 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.

  [Illustration: FIG. 27. Distribution of the cinereous shrew in
  Washington. A. _Sorex cinereus cinereus._ B. _Sorex cinereus

=Sorex cinereus streatori= Merriam

      _Sorex personatus streatori_ Merriam. N. Amer. Fauna, 10:62,
      December 31. 1895.

      _Sorex cinereus streatori_ Jackson. Jour. Mamm., 6:56. February 9,

      _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

      _Measurements._--Five females from Alta Lake, British Columbia,
      average: total length 103; length of tail 45; hind foot 12.5.

      _Distribution._--The western part of the northern Cascades
      (Glacier, Jackson, 1928: 55) and the Olympic Peninsula (Neah Bay
      south to Cedarville, Jackson, 1928: 55).

_Remarks._--This shrew seems to occupy the same range as _Sorex obscurus
setosus_ but is rare where _obscurus_ is common.

=Sorex merriami merriami= Dobson

Merriam shrew

      _Sorex merriami_ Dobson. Monogr. Insectivora, pt. 3, fasc. 1, pl.
      23, fig. 6, May, 1890.

      _Sorex merriami merriami_ Benson and Bond. Jour. Mamm., 20: 348,
      August 14, 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 States National

      _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 shrew is known
      from Washington. Jackson (1928: 81) states that it "was collected
      by George G. Cantwell, November 18, 1919, at the entrance to an
      old badger digging on top of a 'high bunch grass hill' at Starbuck
      (altitude 645 feet), Columbia County, Wash."

_Description._--Similar to _Sorex cinereus_ but upper parts pale grayish
and underparts white.

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

=Sorex trowbridgii= Baird

Trowbridge shrew

_Description._--The Trowbridge shrew closely resembles the cinereous
shrew in body form but possesses a longer tail. The head and body of
adults measure about 2-1/4 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 parts and bluish or slaty
underparts. The tail is distinctly bicolor.

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 beneath the fir forest,
where they are usually the only shrews present. They are abundant in
ravines and in some swampy woods when other shrews are absent, but they
avoid open meadows or marshes. 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, 1941A:
173). The principal food of the Trowbridge shrew includes soft-bodied
insects and insect pupae.

  [Illustration: FIG. 28. Distribution of the Merriam and Trowbridge
  shrews in Washington. A. _Sorex merriami merriami._ B. _Sorex
  trowbridgii trowbridgii._ C. _Sorex trowbridgii destructioni._]

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

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.

=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 catalogue in July, 1855, and the skull in January,
      1857. The other was obtained by J. 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 areas from the Pacific Coast eastward
      through the Cascades to Stehekin (Jackson, 1928: 96), 2 mi. S.
      Blewitt Pass (Jackson, 1928: 96) and Satus Pass (M.V.Z.).

=Sorex trowbridgii destructioni= Scheffer and Dalquest

      _Sorex trowbridgii destructioni_ Scheffer and Dalquest, Jour.
      Mamm., 23:334, August 13, 1942.

      _Type._--Obtained on Destruction Island, Jefferson County,
      Washington, by V. B. 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 reddish brown
and the underparts gray tinged with brownish. In winter 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, wandering 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 a
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 in
Transition, Canadian, or Upper Sonoran life-zones, present comparable
ecological conditions.

Broadbrooks (1939: 65) found that captives taken at Seattle ate rolled
oats, apple, fresh or cooked meat, sow bugs, centipedes, earthworms,
frogs (_Hyla regilla_), a salamander (_Plethodon vehiculum_), and
small, black slugs (_Arean arean_). Wandering shrews proved incapable
of destroying snails (_Helisoma occidentalis_) and large slugs. The
captive shrews kept by Broadbrooks ate an average of 1.3 times their
own weight in food each day.

  [Illustration: FIG. 29. Distribution of the wandering shrew in
  Washington. A. _Sorex 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 discovered
on January 27, 1937. Most adult females taken in February, 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.

=Sorex vagrans vagrans= Baird

      _Sorex vagrans_ Baird, Rept. Pacific R. R. Survey, 8 (pt. 1):15,

      _Sorex suckleyi_ Baird, Rept. Pacific R. R. Survey, 8 (pt. 1):18,
      1857 (type from Steilacoom Pierce County, Washington).

      _Type._--Obtained at Willapa Bay [Shoalwater Bay], Pacific County,
      Washington, by J. G. Cooper; entered in U. S. Nat. Mus. catalogue
      on October 23, 1856.

      _Racial characters._--Dark color.

      _Measurements._--Twenty-five 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 Keechelus (W.S.M.) and 15 mi. NW White Salmon (Jackson,
      1928: 106).

=Sorex vagrans monticola= Merriam

      _Sorex monticolus_ Merriam, N. Amer. Fauna, 3:43, September 11,

      _Sorex vagrans monticola_ Merriam, N. Amer. Fauna, 10:69, 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._--Pale color.

      _Measurements._--Five males and 5 females from Selah, Yakima
      County, 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
      extends west to Bauerman Ridge (Jackson, 1928: 113), Merritt
      (W.W.D.) and Maryhill (M.V.Z.).

      _Remarks._--A series of shrews from Moses Lake, Grant County,
      differs from _monticola_ in larger size and darker color. They
      agree rather closely with _Sorex r. amoenus_ from California and
      Nevada. Because they are isolated from that subspecies it seems
      best to consider them a microgeographic race referable to

=Sorex obscurus= Merriam

Dusky shrew

_Description._--The dusky shrew is similar in form of body to the
cinereous shrew. The length of head and body is about 2-1/4 inches. The
tail is about 2-1/2 inches. The upper parts are rusty or reddish brown.
The underparts are brownish gray. The dusky shrew differs 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 Mexico, and from
the Rocky Mountains to the Pacific Ocean. Thirteen subspecies are
recognized by Jackson (1928: 115), of which two occur in Washington.

Records of the dusky shrew are not available from the arid subdivision
of the Transition or the Upper Sonoran life-zones. It occurs 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 _Sorex vagrans_ was
abundant. Near Stevens Pass, King County, two were taken in a marsh; two
others were trapped in a dry, coniferous forest; one was taken in a
small bed of heather on a barren mountain 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 carefully 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 closed, they were able to
creep about and squeal.

A specimen obtained 5 miles west-southwest of Guler, Skamania County,
contained 4 embryos on July 10, 1939.

  [Illustration: FIG. 30. Distribution of the dusky shrew in Washington.
  A. _Sorex obscurus obscurus._ B. _Sorex obscurus setosus._]

=Sorex obscurus obscurus= Merriam

      _Sorex vagrans similis_ Merriam, N. Amer. Fauna, 5: 34, July 30,
      1891 (not of Hensel, 1855).

      _Sorex obscurus_ Merriam, N. Amer. Fauna, 10: 72, December 31,
      1895 (substitute for _similis_ Merriam).

      _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, color pale.

      _Measurements._--Thirteen males and 10 females from Indian-point
      Lake, British Columbia, average, respectively: total length 105,
      107; length of tail 45, 46; hind foot 13.3, 13.3.

      _Distribution._--Northeastern Washington west, according to
      Jackson (1928: 122), to Pasayten River, Stehekin and Wenatchee.
      Because 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= Elliot

      _Sorex setosus_ Elliot, Field Columb. Mus. Publ. 32. zoöl. ser.,
      1:274. March, 1899.

      _Sorex obscurus setosus_ Jackson, Proc. Biol. Soc. Washington.
      31:127. November 29, 1918.

      _Sorex obscurus bairdi_ Jackson, N. Amer. Fauna, 51:140. July 24,

      _Type._--Obtained at Happy Lake, Clallam County, Washington, by D.
      G. Elliot on August 18, 1898; type in Field Museum of Natural

      _Racial characters._--Tail long, color dark.

      _Measurements._--Twelve males and 17 females from southwestern
      Washington 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

  [Illustration: FIG. 31. Distribution of the mountain water shrew.
  _Sorex palustris navigator_, in Washington.]

      _Neosorex navigator_ Baird, Rept. Pacific R. R. Survey, 8 (pt. I):
      11, 1857.

      _Sorex (Neosorex) palustris navigator_ Merriam, N. Amer. 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,
      respectively: total length 150, 150; length of tail 70, 74; hind
      foot 18.5, 20; weight?, 11.0 grams.

      _Distribution._--Mountainous areas of entire state, including the
      Olympic Mountains, from Elwah (Jackson, 1928: 188) south to
      Quinault River (Jackson, 1928: 189); the Cascades from Tomyhoi
      Lake (W. W. D.) south to 15 mi. N. 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 swimming.

Mountain water shrews range over much of Canada and in mountainous 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 Hudsonian 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 mountain 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: 45) observed that the fur of a
swimming water shrew gathered air bubbles 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 sand and debris, searching for food, its feet kicking rapidly
in order to maintain this position. 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 Pass 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 shrew
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,
float like a duck, and walk on the surface of the water as well as walk
on land.

The food of the mountain water shrew includes snails, leeches, and the
larvae of aquatic insects. Mice caught in traps are sometimes eaten by
water shrews. Svihla (_loc. cit._) 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 3-1/2 inches long; the tail is about 2-3/4 inches
long. The Bendire water shrew closely resembles also the mountain water
shrew 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.

  [Illustration: 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.
Generally they are found at elevations 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 Jackson 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 deep, 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, 1884.

      _Atophyrax Bendirei_ True, Proc. U. S. Nat. Mus., 7 (1884):606,

      _Sorex bendirii_ Dobson, Monog. Insectivora, part 3, fasc. 1, pl.
      23, 1890.

      _Neosorex bendirii bendirii_ Miller, U. S. Nat. Mus. Bull., 79:22,
      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 characters._--Underparts everywhere sooty black.

      _Measurements._--Ten males and 10 females from southwestern
      Washington average, respectively: total length 163.9, 161.0;
      length of tail 71.0, 72.6; hind foot 20.5, 20.3; weight 16.8, 14.5

      _Distribution._--The southern Cascades and the lowlands of western
      Washington, 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, N. Amer. Fauna,
      10:97, December 31, 1895.

      _Neosorex bendirii albiventer_ Miller, U. S. Nat. Mus. Bull.,
      79:22, December 31, 1912.

      _Sorex bendirii albiventer_ Jackson, N. Amer. Fauna, 51:198, July
      24, 1928.

      _Type._--Obtained at Lake Cushman, 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 near the type locality measures:
      total length 167; length of tail 69; hind foot 22. One from
      Potlatch, Mason County, measures 167; 69; 22.

      _Distribution._--The Olympic Peninsula of Washington. Marginal
      localities on the south are: Potlatch (M.V.Z.) and Lake Quinault
      (Jackson, 1928: 199).

=Microsorex hoyi washingtoni= Jackson

Pigmy shrew

      _Microsorex hoyi washingtoni_ Jackson, Proc. Biol. Soc. Washington,
      38:125, November 13, 1925.

      _Type._--Obtained at Loon Lake, Stevens County, Washington, by V.
      Bailey on September 26, 1897; type in United States National

      _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 1-3/4 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.)

Genus =Myotis= Kaup

Mouse-eared bats

_Description._--The genus _Myotis_ may be separated from all other bats
that occur in Washington by the presence of 38 teeth (dental formula i.
2-2/3-3, c. 1-1/1-1, p. 3-3/3-3, m. 3-3/3-3 = 38). Their small size
separates them 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 considerably in size, but all are less than 100 mm. in total
length. The upper 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.

Grinnell (1918: 241-242) points out that, although bats are not subject
to isolation by topographic barriers, as most wingless mammals 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 difficult 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 isolated 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
Faunal 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 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 Faunal Area.

=Myotis lucifugus= (Le Conte)

Big myotis

_Myotis lucifugus_ is represented by two geographic races in Washington.
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.

  [Illustration: 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 ear (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 rather 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 differ from
those of other species with which it was associated, except at the south
end of Lake Chelan, Chelan County, where two individuals 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

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

=Myotis lucifugus carissima= Thomas

      _Myotis (Leuconoë) carissima_ Thomas, Ann. and Mag. Nat. Hist., 13
      (ser. 7): 383, May, 1904.

      _Myotis lucifugus carissima_ Cary, N. Amer. Fauna, 42:43, October
      3, 1917.

      _Type._--Obtained at Yellowstone Lake, Yellowstone National Park,
      Wyoming 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._--East 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, N. Amer. Fauna, 13:63,
      October 16, 1897.

      _Vespertilio gryphus lucifugus_ Allen, U. S. Nat. Mus. Bull.,
      43:78, March 14, 1894 (part 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 foot 12; ear
      12; height of tragus 7; weight 5.4 grams.

      _Distribution._--From the eastern base of the Cascade Mountains
      west of the Pacific, and the Blue Mountains of southeastern
      Washington. Marginal occurrences listed by Miller and Allen
      (1928:49) are Chilliwack River, Lake Wenatchee, and Lyle.

=Myotis yumanensis= (H. Allen)

Yuma myotis

_Description._--The present species closely resembles _Myotis lucifugus_
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).

  [Illustration: FIG. 34. Distribution of the Yuma myotis in Washington.
  A. _Myotis yumanensis 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).

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

      _Myotis yumanensis sociabilis_ H. W. Grinnell, Univ. California
      Publ. Zoöl., 12:318, December 4, 1914.

      _Type._--Obtained at old Fort Tejon, Kern County, California, by J.
      Grinnell on July 23, 1904; type in Museum of Vertebrate Zoölogy.

      _Racial character._--Color pale.

      _Measurements._--Two males and 2 females from Selah, Yakima
      County, average: total length 78; length of tail 36; hind foot 10;
      ear 14; height of tragus 7.

      _Distribution._--Eastern Washington generally. Marginal records on
      the west are: Stehekin (Miller and Allen, 1928: 69), and Selah

  [Illustration: FIG. 35. Distribution of the fringe-tailed myotis and
  the Keen myotis in Washington. A. _Myotis thysanodes thysanodes._
  B. _Myotis keenii keenii._]

=Myotis yumanensis saturatus= Miller

      _Myotis yumanensis saturatus_ Miller, N. Amer. Fauna, 13:68,
      October 16, 1897.

      _Type._--Obtained at Hamilton, Skagit County, Washington, by T. S.
      Palmer on September 13, 1889; type in United States National

      _Racial character._--Color dark.

      _Measurements._--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

      _Distribution._--From the eastern base of the Cascade Mountains to
      the Pacific. This is the commonest _Myotis_ found in western
      Washington. Marginal 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 keenii_ Miller, N. Amer. Fauna, 13:77, October
      16, 1897.

      _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 Duc 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 Duc Hot Springs
      and Lake Cushman.

_Description._--_Myotis keenii_ is similar, in general, to _Myotis
lucifugus_ 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

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 northern 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. anterior 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.

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 from the California Museum of Vertebrate
Zoölogy took specimens in the Blue Mountains where the bats flew
slowly, in rather straight courses, 20 to 25 feet from the ground.

  [Illustration: FIG. 36. Distribution of the long-eared myotis in
  Washington. A. _Myotis evotis evotis._ B. _Myotis evotis pacificus._]

This species has not, so far as is known, been taken in Washington in
its daytime retreat. Daniel Bonell saved two specimens from under slabs
of loose bark on old, dead snags near Tillamook, Oregon. Davis (1939:
214) reported them as hiding in the daytime in a cave in 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)

      _Vespertilio evotis_ Allen, Smithsonian Misc. Coll., 7 (no.
      165):48, June, 1864.

      _Myotis evotis_ Miller, N. Amer. Fauna, 13:77, October 16, 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

      _Distribution._--The Blue Mountains area, of southeastern
      Washington; recorded 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.

      _Myotis evotis pacificus_ Dalquest, Proc. Biol. Soc. Washington,
      56:2, February 25, 1943.

      _Type._--Obtained from 3-1/2 miles east and 5 miles north of
      Yacolt, Clark County, Washington, by John Chattin on August 3,
      1940; type in Museum of Vertebrate Zoölogy.

      _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,

      _Myotis thysanodes thysanodes_ Miller and Allen, U. 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.

      _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 thysanodes_ resembles _Myotis evotis_, but
differs 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 southern
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
_thysanodes_ 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)

Hairy-winged myotis

_Description._--This species, in Washington, may be easily recognized by
its relatively large size and the presence of a distinct keel on the
side of the calcar, posterior to the foot.

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 all 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 clearings 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 lucifugus_ and _Myotis yumanensis_. They were appreciably larger
than those species and their flight was slower and less erratic. They
usually flew in relatively straight lines or large circles at from ten
to forty feet from the ground. At Lake Kapowsin, Pierce County, they
were attracted by swishing a long pole in the air. At Renton, King
County, one was shot as it hunted insects at a city street light several
hours after dark.

=Myotis volans longicrus= (True)

      _Vespertilio longicrus_ True, Science, 8:588, 1886.

      _Vespertilio nitidus longicrus_ H. Allen, U. S. Nat. Mus. Bull.,
      43:103, March 14, 1894.

      _Myotis lucifugus longicrus_ Miller, N. Amer. Fauna, 13:64,
      October 16, 1897.

      _Myotis longicrus_ Lyon and Osgood, U. S. Nat. Mus. Bull., 62:271,
      January 28, 1909.

      _Myotis volans longicrus_ Miller and Allen, U. S. Nat. Mus. Bull.,
      144:140, May 25, 1928.

      _Type._--Obtained in the "vicinity of Puget Sound, Washington" by
      D. S. Jordan, and catalogued in the U. S. National Museum on
      December 16, 1886.

      _Racial character._--Color dark.

      _Measurements._--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.

  [Illustration: FIG. 37. Distribution of the hairy-winged myotis in
  Washington. A. _Myotis volans longicrus._ B. _Myotis volans interior._]

=Myotis volans interior= Miller

      _Myotis longicrus_ 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.

      _Measurements._--Four males and a female from the Blue Mountains,
      Columbia 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
      southeastern part of the state, from Walla Walla (E. S. B.) east
      to Anatone (Miller and Allen, 1928: 144).

      _Remarks._--Of 5 specimens available from the Blue Mountains, 4
      are like _interior_ and 1 is like _longicrus_.

  [Illustration: FIG. 38. Distribution of the California myotis in
  Washington. A. _Myotis californicus californicus._ B. _Myotis
  californicus caurinus._]

=Myotis californicus= (Audubon and Bachman)

California myotis

_Description._--_Myotis californicus_ may be separated from all _Myotis_
that occur in Washington, except _Myotis subulatus_, by its small foot
(about 6 mm.). It is the only small-footed bat found in western
Washington. In eastern Washington, where _Myotis subulatus_ occurs, the
cleaned skulls of the two species must be compared before certain
identification of some specimens is possible. The skull of _M.
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 North America to
southern Mexico. Two subspecies occur in Washington.

_Myotis californicus_, in western Washington, often occurs in company
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

      _Vespertilio nitidus_ H. Allen, Proc. Acad. Nat. Sci. Philadelphia,
      p. 247, 1862 (part of the specimens were from Fort Steilacoom,
      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.

  [Illustration: FIG. 39. Distribution of the small-footed myotis,
  _Myotis subulatus melanorhinus_, in Washington.]

      _Measurements._--Eight specimens, including both sexes, from the
      San Juan Islands, San Juan and Skagit counties, average: total
      length 77.8; length of tail 36.7; hind foot 6.7; ear?; height of
      tragus 7.6; weight 5.2 grams.

      _Distribution._--Western Washington east through the northern
      Cascades to Chelan, Blue Creek and Colville (Miller and Allen,
      1928: 156) in northeastern Washington, and, farther south, east to
      Mount Rainier (Miller and Allen, 1928: 156) and Carson (Miller and
      Allen, 1928: 156).

=Myotis californicus californicus= (Audubon and Bachman)

      _Vespertilio californicus_ Audubon and Bachman, Jour. Acad. Nat.
      Sci., Philadelphia, 8 (ser. 1, ser. 1, pt. 2):285, 1842.

      _Myotis californicus_ Miller, N. Amer. Fauna, 13:69, October 16,

      _Type._--None designated. Type locality fixed at Monterey, Monterey
      County, California, by Miller and Allen (1928: 153).

      _Racial 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 grams.

      _Distribution._--Recorded only from the eastern part of the state.
      Westernmost records, according to Miller and Allen (1928: 155)
      are: Orondo, Goldendale and Lyle.

=Myotis subulatus melanorhinus= (Merriam)

Small-footed myotis

      _Vespertilio melanorhinus_ Merriam, N. Amer. Fauna, 3:46, September
      11, 1890.

      _Myotis subulatus melanorhinus_ Miller and Allen, U. S. Nat. Mus.
      Bull., 144:169, May 25, 1928.

      _Type._--Obtained on San Francisco Mountain, 8250 ft. elevation,
      Coconino County, Arizona, by C. H. Merriam and V. Bailey on August
      4, 1889; type in United States National Museum.

      _Measurements._--Three males and 2 females from eastern Washington
      average: total length 77; length of tail 37; hind foot 7; ear 15;
      height of tragus 8. 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. W. D.) in the north,
      Wenatchee (W. W. D.) in the northwest, Lyle (W. W. D.) in the
      southwest, and Bly (Miller and Allen, 1928: 171) in the southeast.

_Description._--_Myotis subulatus_ is closely similar to _Myotis
californicus_, 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 _M.

This species is confined to 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 eastern
Washington. It lives far from trees on plains and in sandy or rocky
areas, emerging rather early in the evening. It is not difficult 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 outcrops.
Near Vantage, Grant County, individuals were shot as they hung up in a
concrete underpass to digest food. The stomachs of specimens 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 still present there at

=Lasionycteris noctivagans= (Le Conte)

Silver-haired bat

      _V[espertilio]. noctivagans_ Le Conte, McMurtrie's Cuvier, Anim.
      Kingd., 1:431, 1831.

      _Vesperugo noctivagans_ True, Proc. U. 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
      "eastern United States."

      _Measurements._--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 recorded 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. Numerous
white-tipped hairs give the upper parts a frosted appearance. The upper
surface of the interfemoral membrane is well furred, a character shared
only with _Lasiurus_. The dental formula (i. 2-2/3-3, c. 1-1/1-1, p.
2-2/3-3, m. 3-3/3-3 = 36) is the same as that of _Corynorhinus_. The
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 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 migratory, 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 clearings 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.

  [Illustration: FIG. 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 identify them in the air. It is an
early flier, usually appearing just after the swallows roost. They fly
at a considerable height, rarely coming within forty feet of the ground.
The wings are moved with a "fluttery" 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 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 formula is: i.
2-2/3-3, c. 1-1/1-1, p. 2-2/3-3, m. 3-3/3-3 = 36.

  [Illustration: FIG. 41. Long-eared bat (_Corynorhinus rafinesquii
  intermedius_), 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 distribution
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 state. Whitlow and Hall
(1933: 245) give a detailed account of individuals found hibernating in
winter near Pocatello, Idaho.

  [Illustration: FIG. 42. Entrance to Boulder Cave, inhabited by
  long-eared bats (_Corynorhinus_); Kittitas County, Washington, May 26,
  1938. (Fish and Wildlife Service photo by Victor B. Scheffer, No. 56.)]

John K. Townsend (1839: 325) mentions that the "great-eared bat" at the
forts of the Columbia River district (Fort Vancouver) were protected by
the "gentlemen of the Hudson's Bay Company for their services in
destroying the _dermestes_ which abound in their fur establishments."
Townsend mentions also that the long-eared bats seldom left the
"storehouses attached to the forts," even at night. My own observations
are similar, in that at Boulder Cave, Kittitas County, on July 7, 1936,
when not less than 100 long-eared bats were present in the cave, I
watched the entrances to the cave until an hour after dark but no bats
were seen to emerge. All of the specimens available from Washington were
caught in their daytime hiding place. The number of long-eared bats at
Boulder Cave has decreased in recent years. On July 12, 1930, bats were
so abundant that 90 were captured with a single sweep of a butterfly net
(T. H. Scheffer, 1930: 11). On July 7, 1936, it was estimated that there
were slightly more than 100 in the cave. On June 11, 1937, the number
was less, probably about 75.

Little is known of the food habits of the long-eared bat. The stomach of
a specimen from Blakeley Island, San Juan County, was 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 carrying naked
young a week later. On July 7, 1936, at the same locality, females
contained nearly full-term embryos.

  [Illustration: FIG. 43. Distribution of the long-eared bat in
  Washington. A. _Corynorhinus rafinesquii townsendii._ B. _Corynorhinus
  rafinesquii intermedius._]

=Corynorhinus rafinesquii townsendii= (Cooper)

      _Plecotus townsendii_ Cooper, Ann. Lyc. Nat. Hist. New York, 4:73,
      November, 1837.

      _Corynorhinus macrotis townsendii_ Miller, N. Amer. Fauna, 13:53,
      October 16, 1897.

      _Corynorhinus megalotis townsendii_ G. M. Allen, Bull. Mus. Comp.
      Zoöl., 60:344, April, 1916.

      _Corynorhinus 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.

      _Measurements._--A male from Blakeley Island, San Juan County,
      measured: total length 83; length of tail 43; hind foot 8.5; ear
      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.

=Corynorhinus rafinesquii intermedius= H. W. Grinnell

      _Corynorhinus rafinesquii intermedius_ H. W. Grinnell, Univ.
      California Publ. Zoöl., 12:320, December 4, 1914.

      _Corynorhinus rafinesquii townsendii_ Dalquest, Jour. Mamm.,
      19:213, May 14, 1938.

      _Type._--Obtained at Auburn, Placer County, California, by J. C.
      Hawver on July 31, 1909; type in Museum of Vertebrate Zoölogy.

      _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; ear 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.

  [Illustration: FIG. 44. Distribution of the western pipistrelle,
  _Pipistrellus hesperus hesperus_, in Washington.]

=Pipistrellus hesperus hesperus= (H. Allen)

Western pipistrelle

      _Scotophilus hesperus_ H. Allen, Smithsonian Misc. Coll., 7 (no.
      165):43, June, 1864.

      _Vesperugo hesperus_ True, Proc. U. S. Nat. Mus., 7:602, 1885.

      _Pipistrellus hesperus_ Miller, N. Amer. Fauna, 13:88, October 16,

      _Type._--Obtained at Old Fort Yuma, Imperial County, California, by
      G. H. Thomas; catalogued in U. S. National Museum on October 31,

      _Measurements._--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 1-3/4 inches long and the tail 1-1/4. It may be
separated from _Myotis_ by the bent tragus and by the possession of 34
rather than 38 teeth. The dental formula is: i. 2-2/3-3, c. 1-1/1-1, p.
2-2/2-2, m. 3-3/3-3 = 34.

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. No
breeding records are known from the state.

=Eptesicus fuscus bernardinus= Rhoads

Big brown-bat

      _Eptesicus fuscus bernardinus_ Rhoads, Proc. Acad. Nat. Sci.
      Philadelphia, 1901:619, February 6, 1902.

      _Eptesicus fuscus pallidus_ 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 little, if any, paler than series from western Washington.
Washington specimens most closely resemble specimens from California of
the race _bernardinus_ and average darker than _pallidus_.

_Description._ The big brown-bat may be distinguished from other bats by
its large size (about 4-1/2 inches), rich, brown color, and small ears
(reaching only to nostril when laid forward). It possesses 32 teeth, the
dental formula being: i. 2-2/3-3, c. 1-1/1-1, p. 1-1/2-2, m. 3-3/3-3 =

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.

  [Illustration: FIG. 45. Distribution of the big brown-bat, _Eptesicus
  fuscus bernardinus_, in Washington.]

The big brown-bat appears early in the evening, often before the
swallows have retired. This, and its large size, makes 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 trees in the city parks. In its more natural habitat it flies
over trees and clearings. Big brown-bats were repeatedly timed, with a
car's speedometer, at 17 miles an hour as they flew down a road lined
with tall trees. This is a greater speed 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

Big brown-bats are 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 second-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. On 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.

=Lasiurus cinereus cinereus= (Beauvois)

Hoary bat

      _Vespertilio cinereus_ (misspelled _linereus_) Beauvois, Catal.
      Raisonné Mus. Peale, Philadelphia, 1796:18 (p. 15 of English
      edition by Peale and Beauvois).

      _Lasiurus cinereus_ H. Allen, Smithsonian Misc. Coll., 7 (no.
      165): 21, 1864.

      _Atalapha cinerea_ True, Proc. U. S. Nat. Mus., 7:602, 1885.

      _Type._--None designated. Type locality Pennsylvania, probably near

      _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.

  [Illustration: FIG. 46. Record stations for the hoary bat, _Lasiurus
  cinerea_, 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
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: i. 1-1/3-3, c. 1-1/1-1,
p. 2-2/2-2, m. 3-3/3-3 = 32.

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

Information on the natural history of the hoary bat is meager. It was
observed in the Kettle River Mountains, but none was obtained. 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 September,
rarely lingering until early October. It winters along the coast of
central and southern California (Dalquest, 1943: 23).

=Antrozous pallidus cantwelli= Bailey

Pallid bat

      _Antrozous pallidus cantwelli_ Bailey, N. Amer. Fauna, 55:391,
      August 29, 1936.

      _Type._--Obtained at Rogersburg, Asotin County, Washington, by G.
      G. Cantwell on May 28, 1918; type in United States National Museum.

      _Measurements._--A male from the south bank of the Columbia River,
      opposite 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 (4-1/2 to 5 inches), lighter
color, thick and leathery membranes, and shorter, wider ears.
_Antrozous_ lacks the prominent "lumps" on the rostrum, characteristic
of the long-eared bat. The dental formula is: i. 1-1/2-2, c. 1-1/1-1, p.
1-1/2-2, m. 3-3/3-3 = 28.

The genus _Antrozous_ is confined to western United States and
northwestern Mexico. Two species are recognized by Miller (1924: 84).
One of these (_pallidus_) includes three geographic races. A single race
is known to occur in Washington.

  [Illustration: FIG. 47. Distribution of the pallid bat, _Antrozous
  pallidus cantwelli_, in Washington.]

The pallid bat chooses a crevice in a cliff, cave, or building as its
daytime hiding place. Like _Corynorhinus_, it is colonial. The basalt
cliffs of eastern Washington offer such ideal hiding places that pallid
bats are seldom seen and consequently little is known of their habits.
In the southwestern United States, pallid bats are more abundant and
better known. H. W. Grinnell (1918: 355) notes that the floor of a roost
of pallid bats was strewn each morning with the heads, wings, and legs
of insects. Most numerous were the remains of the Jerusalem cricket
(_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) believes that the young of this bat are born in late
June and early July. One or two young constitute a litter.

=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 (1937: 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. cit._, 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 exceed 350 pounds. The largest males
have hind feet from 10 to 10-1/2 inches in length.

  [Illustration: FIG. 48. Female black bear (_Ursus americanus
  altifrontalis_), and two cubs, near Big Four Inn, Mt. Baker National
  Forest, Washington, July, 1939. (Forest Service photo by R. L.

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 long,
sleek and glossy but rather stiff. In the spring and early summer the
fur of animals at Mount Rainier often appeared coarse, wooly and patchy
or rubbed on older animals. Some smaller bears 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 bears in their life. I have seen only black bears
in the lowlands of western Washington. Cowan (_loc. cit._) lists 1,197
black and 79 brown bears from Fort Nisqually, Pierce County, between
1834 and 1852. In the Cascade Mountains the brown phase is not uncommon,
perhaps 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 are 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 Washington, and is
most abundant in the Cascade Mountains and Olympic Mountains where food
is abundant and men are few. It is not uncommon throughout the timbered
lowlands of northeastern Washington and western Washington, however, and
shows a surprising ability to exist unnoticed near the larger cities. In
the dense, junglelike forests of southwestern Washington it is numerous.
Jackson (1944: 1) estimates that 13,679 black bears live in Washington,
more than in any other state.

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 Cascades the bears are wild but 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 man or his guns. As a result they are silent
and shy, rarely being seen.

  [Illustration: FIG. 49. Black bear (_Ursus americanus altifrontalis_),
  in "hibernation," Mt. Baker region, Washington, about 1936. (John E.
  Candle photo, courtesy "Field and Stream.")]

The black bear in the mountains is active sometimes by day and sometimes
by night. Probably it is principally nocturnal but active by day only
when the food obtained at night is insufficient to meet the needs of the
animal. In the lowlands it is almost completely nocturnal. In the
mountains it enters upon its winter sleep with the first snows or, if
the snows are late, when the huckleberries 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.

The black bear 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 pieces of
apples or other fruit.

The principal food of the black bear in Washington is berries. Many
species are eaten, but the huckleberry (_Vaccinium_ sp.) is favored.
Other food items are: _Gaultheria shallon_ (pulpy fruit); Oregon grape,
_Berberis nervosa_ (flowers and fruit); salmonberry, _Rubus parviflorus_
(leaves and fruit); leaves of several plants, including _Rubus
macropetalus_ and other thorny kinds; grass, succulent plants, and
roots. Insects are eaten and most feces show 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 are spawning
the bears in the vicinity feed on nothing else. Warm-blooded vertebrates
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 few 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 large 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 an oval opening in
the ground in a meadow of heather and huckleberry near a small stream.
The wallow was approximately 5 feet long, 3 feet 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 paws. When a stray breeze brought to the
bear the scent of my companion and myself the animal's contentment
vanished and it hastily made a frantic effort to depart. It had wedged
itself deeply in the hole, and as it twisted and turned in attempting to
sit up, waves of water gushed from the wallow. Eventually escaping from
its 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

  [Illustration: FIG. 50. Distribution of the black bear in Washington.
  A. _Ursus americanus altifrontalis._ B. _Ursus americanus cinnamomum._]

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 intelligent 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 dollars,
usually less than ten. Their sale scarcely repays the trapper for his
labor in preparing the hide. Townsend (1887: 182) remarked on 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.

=Ursus americanus altifrontalis= Elliot

      _Ursus altifrontalis_ Elliot, Field Columb. Mus. Publ. 80, zoöl.
      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 National 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.
      Zoöl., 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 var. cinnamomum_ Audubon and Bachman, Quadrupeds
      of North America, 3:125, 1854.

      _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. Zoöl.,
      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 narrower; color even in black phase paler and browner;
      brown and black phases of approximately equal incidence.

      _Distribution._--Northeastern Washington and the Blue Mountains of
      southeastern Washington, occurring westward as far as Republic
      (Taylor and Shaw, 1929: 10).

=Ursus chelan= Merriam

Grizzly bear

      _Ursus chelan_ Merriam, Proc. Biol. Soc. Washington, 29:136,
      September 6, 1916.

      _Type._--Obtained in Township 30 N. Range 16 E. Willamette
      Meridian, Wenatchee National Forest, Chelan County, Washington, by
      D. S. Rice on September 1, 1913; type in United States National

      _Measurements._--The skull of 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 area 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 (Grinnell, Dixon and Linsdale,
      1937: 67) but there is no record of their ever having occurred in
      the lowlands of western Washington.

Bears of the genus _Ursus_ range over Europe, Asia and North America. No
less than 75 kinds of grizzly and big brown bears are 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.

  [Illustration: FIG. 51. Probable past distribution of grizzly bears in
  Washington. A. _Ursus chelan._ B. _Ursus canadensis._ C. _Ursus

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. Cranially, 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 "silver tips." In Washington, the black bear
varies in color from glossy black, through various shades of brown to a
pale cinnamon.

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 undoubted grizzly bear in Washington.
The type specimen was obtained well up in the Cascade Mountains where
the animals, like those in British Columbia, feed on roots, berries,
marmots, pikas and other vegetable and animal life.

=Procyon lotor= (Linnaeus)


_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

The raccoon is not an aquatic mammal, yet it is closely associated 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 occasionally
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 ranges well up
into the Cascade Mountains. The highest altitudinal record available is
Longmire, Mount Rainier National Park (Taylor 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 occasions. 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 tracks 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 _Rana_), and aquatic insects are eaten. Along beaches fish
brought in by the tide, crustaceans, and mussels (_Mytilus edulis_) are
taken; small sharks appear not to be eaten. A recently dead dogfish that
lay on the beach at Whidby Island, Island County, was ringed by the
tracks of a raccoon but had not been eaten. The shore crabs
(_Hemigrapsus nudus_ and _Hemigrapsus oregonensis_) are 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
eriomerus_) was commonly eaten. Small mammals and birds are eaten
regularly by this race of raccoon in California (Grinnell, Dixon and
Linsdale, 1937: 157), and the eggs and young of wild birds are sought in
the spring (_op. 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. Insects are present in small quantities in
most droppings and in the late summer some feces were composed entirely
of the remains of grasshoppers. 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. 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 year of life (Pope, 1944:

=Procyon lotor psora= Gray

      _Procyon psora_ Gray, Ann. and Mag. Nat. Hist., 10: 261. December,

      _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. December, 1929.

      _Procyon lotor psora_ Grinnell, Dixon, and Linsdale, Fur-bearing
      Mamm. California, Univ. California Press, Berkeley, p. 137, July
      22, 1937.

      _Type locality._--Sacramento, Sacramento County, California.

      _Racial characters._--Medium size; dark color; smoothly rounded

      _Measurements._--A female from Forks, Clallam County, measures:
      total length 905; length of tail 355; hind foot 125; ear 50;
      weight 14-1/2 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) 8-2/3 lbs.

  [Illustration: FIG. 52. Distribution of the raccoon in Washington. A.
  _Procyon lotor psora._ B. _Procyon lotor excelsus._]

      _Distribution._--From the western slope of the Cascade Mountains
      westward. Marginal occurrences are Lake Keechelus (type locality)
      and Mount Rainier (Taylor and Shaw, 1927: 45).

      _Remarks._--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 the raccoons of western Washington
      from those of northern California.

=Procyon lotor excelsus= Nelson and Goldman

      _Procyon lotor excelsus_ Nelson and Goldman, Jour. Mamm., 11:458,
      November 11, 1930.

      _Type._--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 States 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 rivers 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.

_Remarks._--The assignment of the raccoons of northeastern Washington to
_excelsus_ is tentative for I have seen no specimens.

=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 Washington 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 throughout 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 skillfully 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 to 10
animals. 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 mountain," or midway 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 chipmunks. Mice,
also, are eaten. The deer mouse, _Peromyscus maniculatus_, 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 although 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 professional
trappers 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, requiring one to three days' travel by
the trapper to complete the circuit.

The standard "set" for marten in the Cascades consists of an 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.

  [Illustration: 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 by
one trapper in a single winter, of which I know, was 300 animals, taken
near Mt. Adams.

=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

      _Distribution._--From the 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

_Remarks._--Skulls of martens from many localities in western North
America were studied in an effort to determine the relationship of the
eastern pine marten (_Martes 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 is
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_ Miller, 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 Philadelphia.

      _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

=Martes pennanti= (Erxleben)


      [_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, U. S. Nat. Mus. Bull., 79:94,
      December 31, 1912.

      _Martes pennanti_ Grinnell, Dixon and Linsdale, Fur-bearing Mamm.
      California, Univ. California Press, Berkeley, p. 211, July 22,

      _Type locality._--Eastern Canada.

      _Distribution._--Originally forested areas from the eastern base
      of the Cascades westward and possibly the Blue Mountains of
      southeastern Washington and the mountains of northeastern
      Washington; now probably confined to the Cascade and Olympic

  [Illustration: FIG. 54. Fisher (_Martes pennanti_). 2-year-old male in
  captivity; New Westminster, B. C., March 7, 1939. (Fish and Wildlife
  Service photo by Victor B. Scheffer, No. 598.)]

_Remarks._--The fisher is the size of a large cat. In general
proportions it resembles the marten. Adult males measure about 3-1/2
feet in length; the tail comprises 16 inches of this. Adult females
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
5-1/2 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 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 California. 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 Washington, and
seems never to have been common. In consequence relatively little is
known of its habits, and the little that is known has been reported by

  [Illustration: FIG. 55. Distribution of the fisher, _Martes pennanti_,
  in Washington.]

The fisher seems not to live in bands as does the marten. Most of the
actual records of fishers trapped are at higher altitudes but are
misleading because most trappers agree that the fisher occupied a lower
zone, altitudinally, than does the marten. There are old records of
its occurrence near sea level (Scheffer, 1938: 9). The animals are
usually taken in marten sets or in traps set especially for fishers
by trappers who find their tracks on their marten trap lines. 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_.

=Mustela erminea= Linnaeus


_Description._--Though it is similar in general characters and
proportions to the long-tailed weasel, the ermine is much smaller and
has a relatively shorter tail. It is darker and less reddish or
yellowish in summer. Adult males measure about 10 inches in length, of
which 3-1/2 inches is the length of the tail. Females measure about 8
inches and have tails 2 inches long. The upper parts are chocolate
brown; the underparts are white or pale yellow. Along the coast of
Washington, the pale color of the underparts is more restricted in the
ermine than in the long-tailed weasel. The dark brown tail has a black
tip. Ermines east of the summit of the Cascades become pure white in
winter, save for the black tail tip. West of the summit of the Cascades
the winter pelt is similar to the summer pelt but is slightly paler with
denser underfur.

Ermines in America range from the Arctic southward, in mountainous
areas, to the southern end of the Sierra Nevada in California, 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 occurs. So far as my observations go, the
ermine is principally nocturnal in Washington; I have seen only one
abroad in the daytime. It 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 traps set in trees for martens. Near
Skykomish, King County, William Hoffman 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 reset 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 difficult 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 it would most certainly be seriously wounded or killed.
Judging by the forest habitat occupied by the ermine in Washington, one
might expect its principal food to consist of deer mice (_Peromyscus
maniculatus_), red-backed mice (_Clethrionomys_) and meadow mice

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 were 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, 1938, 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, Jour. Mamm., 26:75, February 27,

      _Type._--Obtained at Benewah, Benewah County, Idaho, by W. T. Shaw
      on October 24, 1926; type in Museum of Vertebrate Zoölogy.

      _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:

=Mustela erminea fallenda= Hall

      _Mustela erminea fallenda_ Hall, Jour. Mamm., 26:79, February 27,

      _Type._--Obtained at Huntingdon, British Columbia, by C. H. Young
      on May 21, 1927; type in National Museum of Canada.

      _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:

_Remarks._--The range of this ermine is similar to that of the
yellow-pine chipmunk, _Eutamias amoenus felix_.

  [Illustration: FIG. 56. Distribution of the ermine in Washington. A.
  _Mustela erminea invicta._ B. _Mustela erminea murica._ C. _Mustela
  erminea gulosa._ D. _Mustela erminea fallenda._ E. _Mustela erminea
  streatori._ F. _Mustela erminea olympica._]

=Mustela erminea olympica= Hall

      _Mustela erminea olympica_ Hall, Jour. Mamm., 26:81, February 27,

      _Type._--Obtained near head of Sol Duc River, Clallam County,
      Washington, by V. Bailey, on April 28, 1897; type in United States
      National Museum.

      _Racial characters._--Similar to _streatori_ but smaller,
      especially females.

      _Measurements._--Twelve males and 6 females average, respectively:
      total length 243, 196; length of tail 65, 52; hind foot 31, 23.4
      (Hall, 1945: 81).

      _Distribution._--The Olympic Peninsula, extending southeastward to

=Mustela erminea streatori= (Merriam)

      _Putorius streatori_ Merriam, N. Amer. Fauna, 11:13, June 30. 1896.

      _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,

      _Type._--Obtained at Mount Vernon, Skagit County, Washington, by D.
      R. Lucky on February 29, 1896; type in United States National

      _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
      Washington 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

=Mustela erminea gulosa= Hall

      _Mustela erminea gulosa_ Hall, Journ. Mamm., 26:84, February 27,

      _Type._--Obtained at Trout Lake, Klickatat County, Washington, by
      P. Schmid on February 3, 1897; type in United States National

      _Racial characters._--Similar to _invicta_ but smaller; winter
      coat usually white; tail less than one-half black; upper lips

      _Measurements._--Five males from Mount Rainier, and 4 females from
      the Cascade Mountains, average, respectively (Hall, 1945: 84):
      total length 253, 208; length of tail 76, 54; hind foot 30.2,

      _Distribution._--The Cascade Mountains. Known from Skykomish
      (W.W.D.) and southward to Mount Adams (Trout Lake, Hall, 1945:

=Mustela erminea murica= (Bangs)

      _Putorius (Arctogale) muricus_ Bangs, Proc. New England Zoöl. 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, December, 1929.

      _Mustela cicognanii muricus_ Bailey, N. Amer. Fauna, 55:292,
      August 29, 1936.

      _Mustela erminea murica_ Hall, Jour. Mamm., 26:77, February 27,

      _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 Zoölogy.

      _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_.

=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 tapering to the muzzle. The
ears are erect, low and rounded. The fur 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 most easily
obtain food. In the Cascade Mountains where diurnal mammals such as
squirrels, chipmunks and pikas are common, weasels are not uncommonly
seen by day. In the lowlands of western Washington, where they feed 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
curious and apt to watch his actions. Weasels are also often hit by cars
and the number so killed seems to me to be out of proportion to their
actual numbers.

In moving on the surface of the ground the weasel arches its back and
contracts the body until the four feet 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 is

The weasel has been accused of killing birds and doubtless does 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 (_Citellus 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. Seemingly 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 forefinger 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 forefinger.
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 evidence 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 animals, they separated,
the weasel retreating to the cover of horsetail (_Equisetum_) beside
the road. The car was stopped 50 feet ahead. As 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.

  [Illustration: FIG. 57. Distribution of the long-tailed weasel in
  Washington. A. _Mustela frenata washingtoni._ B. _Mustela frenata
  altifrontalis._ C. _Mustela frenata nevadensis._ D. _Mustela frenata

The pelts of weasels bring the trapper from twenty-five cents to a
dollar and a half. Only skins in the white winter coat command 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 summer 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, were white on November 15, 1936, but specimens from
Skykomish and Baring, 18 miles to the west, were brown.

The long-tailed weasel 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, December, 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,
      Publ. 473:91, November 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 Zoölogy.

      _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.

      _Measurements._--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 Mountains area. Marginal occurrences are Barron (Hall, 1936:
      93) and Easton (W. W. D.).

      _Remarks._--Specimens from the northern Cascades are intergrades
      between _nevadensis_ and the form to the west, _altifrontalis_.
      Specimens from extreme northeastern Washington might be referred
      to _oribasa_ Bangs as readily as to _nevadensis_.

=Mustela frenata effera= Hall

      _Mustela frenata effera_ Hall, Carnegie Inst. Washington, Publ.
      473:93, November 20, 1936.

      _Type._--Obtained at Ironside, Malheur County, Oregon, by H. E.
      Anthony on September 8, 1912; type in American Museum of Natural

      _Racial characters._--Similar to _nevadensis_ 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. Specimens from Walla Walla and Prescott have been examined.

=Mustela frenata washingtoni= (Merriam)

      _Putorius washingtoni_ Merriam, N. Amer. Fauna, 11:18, June 30,

      _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

      _Racial characters._--Color rich, dark; hind feet free of color of

      _Measurements._--A male from Spray Park, Pierce County, measures:
      total length 423; length of tail 164; hind foot 52.

      _Distribution._--Higher Cascades from Mt. 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, December, 1929.

      _Mustela frenata altifrontalis_ Hall, Carnegie Inst. Washington,
      Publ. 473:94, November 20, 1936.

      _Type._--Obtained at Tillamook, Tillamook County, Oregon, by A.
      Walker on July 10, 1928; type in Museum of Vertebrate Zoölogy.

      _Racial characters._--Similar to _nevadensis_ but darker above in
      summer and richer orange beneath; winter color of upper parts dark
      brown, underparts pale yellow or white.

      _Measurements._--Four males and 2 females from western Washington
      average, respectively: total length 411.5, 267; length of tail
      145, 137.5; hind foot 50, 43; ear 26, 21.5; weight 227, 136.7

      _Distribution._--From the Cascade Mountains to the Pacific.
      Marginal occurrences are Rockport (Hall, 1936: 95) and Tye (W. W.

=Mustela vison energumenos= (Bangs)


      _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 Zoölogy.

      _Measurements._--A male from Seattle, King County, measures: total
      length 540; length of tail 180; hind foot 66; weight 1 pound, 15

      _Distribution._--Throughout the state save for the Columbian
      Plateau; recorded from Neah Bay (Taylor and Shaw, 1929: 11) in the
      northwest, Ilwaco (M.V.Z.) in the southwest, Metaline (W.W.D.) in
      the northeast, and the Blue Mountains (Dice, 1919: 12) in the

  [Illustration: 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 1-1/2 pounds. Males
measure about 2 feet in length, of 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 back. 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, nocturnal. 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 this respect it resembles the
raccoon. Whereas the raccoon spends a part of its time on the land, the
mink spends a corresponding 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 rabbits. 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

The mink ascends some distance into the Cascade and Olympic mountains
along the larger watercourses. There are reliable reports of minks from
Heart Lake and Lake 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. Along the ocean beaches they eat
dead sea birds, stranded fish, crustaceans, and mollusks. Along Puget
Sound their tracks are commonly seen following the caked, decaying
seaweed and debris that collects at the high tide mark. 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
feed on clams. Fur trappers report that minks feed on mountain beavers,
and that the flesh of the 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. Muskrats trapped near Seattle were often attacked by minks and
either eaten or so slashed and torn that their pelts were worth but a
fraction of what they would bring in an undamaged condition.

The tracks of a mink noted in freshly fallen snow about a garbage 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 discovered 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.

=Gulo luscus luteus= Elliot


      _Gulo luteus_ Elliot, Field Columb. Mus., Publ. 87, zoöl. ser.
      3:260, 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.

      _Measurements._--The dried skin of a wolverine taken 3 miles south
      of Riverside, 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 _luteus_.

  [Illustration: FIG. 59. Wolverine (_Gulo luscus_), mounted specimen,
  male, trapped by Billy Robinson about 1902 on Billy Robinson Creek,
  Okanogan County, Washington. (Fish and Wildlife Service photo by
  Victor B. Scheffer, No. 76.)]

_Description._--Large males measure more than three feet in length, of
which the tail makes 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, 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.

  [Illustration: FIG. 60. Wolverine (_Gulo luscus_), male, trapped by
  Reuben R. Lamb, March 22, 1941, three miles south of Riverside,
  Washington; reported to weigh 40 pounds; dried pelt 3 × 4 feet.
  (Reuben R. Lamb photo.)]

  [Illustration: FIG. 61. Distribution of the wolverine, _Gulo luscus
  luteus_, in Washington.]

The wolverine is adapted to boreal conditions and is most abundant 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 suitable habitat for wolverines
is not abundant and each animal requires a large area over which to
range. An unusual record, possibly of a wandering animal, is given by
Scheffer (1941: 37). This is of an adult male taken in the Okanogan
Valley 3 miles south of Riverside, Okanogan County, on March 22, 1941.

=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 3-1/2 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 spaniel-like, and support a
mustache of stiff vibrissae. 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
underparts 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 environment 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 entered 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

In the San Juan Islands the otters have taken to a marine existence. 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 incident. On the third night,
several of the bats 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 bat 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. Only 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 swimming 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 interesting. 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 marine
habitat. At Strawberry Bay, Cypress Island, on July 5, 1938, fishermen
brought in a "strange animal" caught by them at Black Rock, a tiny bare
and isolated islet 5 miles to the west. The heavy box holding the animal
was opened, disclosing a very frightened young otter. Questioned, the
fishermen stated that four young and one adult had been seen in the
surf. They had thought the animals a species of seal, and were somewhat
surprised at being successful in their attempts to catch one. The otter
was too young to fare for itself and was kept in camp in the hope that
it might be reared and released. It ate a few fresh herring and
candlefish and drank some condensed milk. It refused whole salmon and
sea bass but ate some skinned and boned strips of these fish. It died a
week after it was captured. This young otter made a ticking sound,
almost 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
6 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
(_Mytilus edulis_) was most abundant. Crabs formed an important part of
their diet. Strangely enough the shore crab (_Hemigrapsus 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 forefeet, as the
otter is unable to do. The edible crab (_Cancer productus_) and kelp
crab (_Telmessus cheiragonus_) were most often eaten by otters. Also
eaten were snails (_Margarites_, _Littorina_), bitter oysters
(_Pododesmus macroschisma_), unidentified pelecypods, barnacles
(_Balanus_), one chiton (_Mopalia 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 examination 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 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.

  [Illustration: FIG. 62. River otter (_Lutra canadensis pacifica_),
  trapped in Columbia National Forest, Washington, February, 1937.
  (Adolph Roth photo.)] 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 crayfish remains.
Some fish are also eaten and locally otters may do damage among the
steelhead trout. Even worse, from the fisherman's standpoint, they
frighten and scatter the schools of steelhead 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.

  [Illustration: FIG. 63. Distribution of the river otter in Washington.
  Unshaded _Lutra canadensis pacifica_. Shaded _Lutra canadensis

=Lutra canadensis pacifica= Rhoads

      _Lutra hudsonica pacifica_ Rhoads, Trans. Amer. Philos. Soc., n.
      s., 19: 429, September, 1898.

      _Lutra canadensis pacifica_ Allen, Bull. Amer. Mus. Nat. Hist.,
      10: 460, November 10, 1898.

      _Type._--Obtained at Lake Keechelus, Kittitas County, Washington
      (altitude 3,000, rather than 8,000 feet as stated in original
      citation); type in Academy of Natural Sciences of Philadelphia.

      _Measurements._--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 14-3/4 pounds.

      _Distribution._--Watercourses throughout the state, scarcer east
      of the Cascades; recorded at Forks (V. B. S.) in the northwest,
      Puget Island (V. 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. Soc. Washington, 48:
      186, November 15, 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

      _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.

      _Remarks._--The otters of the San Juan Islands are referred to
      this race on geographical grounds. Numerous islands 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

=Enhydra lutris nereis= (Merriam)

Sea otter

      _Latax lutris nereis_ Merriam, Proc. Biol. Soc. Washington, 17:
      159, October 6, 1904.

      _Enhydra lutris nereis_ Grinnell, Univ. California Publ. Zoöl.,
      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.

      _Distribution._--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 influenced 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 Columbia
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.

=Spilogale gracilis= Merriam

Civet cat or spotted skunk

_Description._--The civet cat 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.

_Spilogale_ 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 intergradation 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, for 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

The civet cat is principally a lowland animal, but has been recorded
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 cottonwood. Where conditions are
suitable it ranges up into the Canadian 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 spells 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 few
persons other than trappers, perhaps because of their exclusively
nocturnal habits. They do occasionally make people aware of themselves
by taking up residence about old buildings. They are rather noisy at
times. At the Tolt River, 10 miles southeast of Duvall, King County, we
were asleep in an old building when a stamping noise from the next room
awakened us. Investigation 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 observer'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 climbing. 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 entrance. 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 about its legs.
Striped skunks in traps move slowly and steadily but civet cats jump,
roll and squirm erratically.

  [Illustration: FIG. 64. Distribution of the civet cat in Washington.
  A. _Spilogale gracilis latifrons._ B. _Spilogale gracilis saxatilis._]

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 killed the civet cat. Under ordinary
circumstances a striped skunk could scarcely catch the far swifter and
more agile civet cat. One civet cat in a trap was killed and partly
eaten by a horned owl. Save for 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_ _californicus_). 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:

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,

      _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 _latifrons_.

      _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, zoöl.
      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 Hamilton, 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 (1906: 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.

=Mephitis mephitis= (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 relatively open
country such as logged-over land, old fields and river-side and
streamside thickets. Their dens usually consist of old _Aplodontia_
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 composed 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 that travels an
arching curve, 5 feet high, reaching 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. In small quantities it is not unpleasant. It is extremely
lasting, sometimes being noticeable for months on clothes or buildings.
It is soluble in gasoline and clothes may be de-scented by several
washings 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 moving 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 trappers in
Washington. The value of their furs varies with the demand but large
prime skins usually bring from $1 to $4. In eastern Washington, where
trapping for coyotes and other terrestrial mammals 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 Washington 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,

      _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 Saskatchewan, 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.

      _Distribution_.--Northeastern Washington and the eastern edge of
      the northern Cascades, south probably to the Wenatchee Mountains.
      Recorded west to Oroville (W. W. D.) and Timentwa (W. W. D.) and
      south to Spokane (Taylor and Shaw, 1929: 12).

=Mephitis mephitis major= (Howell)

      _Chincha occidentalis major_ Howell, N. Amer. Fauna, 20:37, August
      31, 1901.

      _Mephitis mephitis major_ Hall, Univ. California Publ. Zoöl.,
      37:2, April 10, 1931.

  [Illustration: FIG. 65. Distribution of the striped skunk in
  Washington. A. _Mephitis mephitis hudsonica._ B. _Mephitis mephitis
  major._ C. _Mephitis mephitis notata._ D. _Mephitis mephitis

      _Type._--Obtained at Fort Klamath, Klamath County, Oregon, by B. L.
      Cunningham on January 5, 1898; type in United States National

      _Racial characters._--Slightly larger than _hudsonica_ with white
      stripes diverging on shoulders rather than neck.

      _Measurements._--A young male from 1 mile north of Burbank, Walla
      Walla County, measures: total length 474; length of tail 205; hind
      foot 68; ear 30; weight 815 grams.

      _Distribution._--Southeastern Washington, south of the Snake River
      and east of the Columbia River, occurring west to Burbank (M. V.

=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, diverging 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, zoöl.
      ser., 1:269, March, 1899 (type from Laguna, 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

      _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.

      _Measurements._--An adult male from 2-1/2 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.).

=Taxidea taxus taxus= (Schreber)


      _Ursus taxus_ Schreber, Saugethiere, 3:520, 1778.

      _Taxidea taxus_ Rhoads, Amer. Nat., 28:524, June, 1894.

      _Taxidea taxus neglecta_ Taylor and Shaw, Occ. Pap. Chas. R.
      Conner Mus., no. 2:12, December, 1929.

      _Taxidea taxus taxus_ Hall, Carnegie Inst. Washington, Publ.
      473:78, November 20, 1936.

      _Type._--None. Type locality probably southwest of Hudson Bay
      (Hall, 1936: 78).

      _Measurements._--Two males from northern Nevada measure,
      respectively: total length 780, 762; length of tail 113, 150; hind
      foot 136, 120; ear 55, 54; weight 15, 11 pounds.

      _Distribution._--Of general distribution over open country east of
      the Cascades. Marginal records are Wauconda (Taylor and Shaw,
      1929: 12), Chelan (Taylor and Shaw, 1929: 13), Goldendale (W. W.
      D.) and "Divide above Trout Lake" (trapper's photograph).

  [Illustration: FIG. 66. Badger (_Taxidea taxus taxus_), tame animal in
  Seattle, Washington. January 29, 1938. Captured at Lakeside, Chelan
  County, and photographed at approximate age of ten months; subadult
  male. (Eloise Kuntz photo.)]

_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 body 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 backwards, between the eyes, to
the shoulders and serves as the best recognition mark.

  [Illustration: 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 Lakeside, Chelan County, found the animal powerful
enough to dig through a concrete floor! Evidence of badger's activities
are usually 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 burrows.

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, claylike 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 diggings may exist almost 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 straying 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, occasional trappers catching as many as 35;
Columbia, Kettle River, and other valleys in northeastern Washington,
uncommon, a few records only; Columbian Plateau, fairly common--average
professional trapper's yearly catch includes 10 to 20 badgers;
southeastern Washington, now rare because of overtrapping, formerly

Trappers state that 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 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 washingtoni_, _townsendii_, _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

=Vulpes fulva cascadensis= Merriam

Red fox

      _Vulpes cascadensis_ Merriam, Proc. Washington Acad. 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,
      measures: 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 19 per cent
were silver or cross. One fox, trapped in Okanogan County, is said by
its captor to have been black above and straw colored beneath, with no
white on the body.

Red foxes range from Alaska and northern Canada well southward into the
United States. 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 its 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.

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 territory
and its fur is not especially valuable. The animal is of relatively
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 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.

  [Illustration: FIG. 68. Distribution of the red fox in Washington. A.
  _Vulpes fulva cascadensis._ B. _Vulpes fulva macroura._ (See p. 450.)]

=Canis latrans= Say


_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

  [Illustration: FIG. 69. Coyote (_Canis latrans lestes_), in trap, 5
  miles southeast of McKenna, 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 Washington,
except for the highest parts of the mountains and the dense forest areas
of western Washington. According to available information 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 Washington 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 Transition 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 purposeful 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
several successive dunes in full flight before turning to take advantage
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 pursue 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 vociferous. 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. _Reithrodontomys_ were found in 53 of
the 8,339 stomachs and in insignificant 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. _Tamias_ (= _Eutamias_) 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 _Tamiasciurus_ occurred in 33 stomachs.
Included were 3 _Tamiasciurus douglasii_, a _Tamiasciurus hudsonicus_,
and a _Sciurus griseus_ from Washington. _Glaucomys_ 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 Washington. 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 Washington, 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 Washington had eaten a garter snake. No frogs were found
in coyote stomachs. A coyote from Washington had eaten a salamander.
Another had eaten a fish. Insects formed 1.08 per cent of the total food
eaten by coyotes. Fruit formed 3.63 per cent. Carrion constituted 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 individual, 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 destroy 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 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 increasing.

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.

  [Illustration: FIG. 70. Distribution of the coyote in Washington. A.
  _Canis latrans lestes._ B. _Canis latrans incolatus._]

=Canis latrans lestes= Merriam

      _Canis lestes_ Merriam, Proc. Biol. Soc. Washington, 11:25, March
      15, 1897.

      _Canis latrans lestes_ Taylor and Shaw, Mamm. and Birds Mt.
      Rainier Nat. Park, p. 41, 1927.

      _Type._--Obtained in the Toyabe Mountains near Cloverdale, Nye
      County, Nevada, by V. Bailey on November 21, 1890; type in United
      States National Museum.

      _Racial characters._--Frontal region of skull only slightly

      _Measurements._--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 26-1/2 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 Washington.

_Remarks._--Pending a revision of the coyotes, those of western
Washington are referred to _lestes_.

=Canis latrans incolatus= Hall

      _Canis latrans incolatus_ Hall, Univ. California Publ. Zoöl.,
      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 Zoölogy.

      _Racial characters._--Similar to _lestes_ but frontal region more

      _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
      Conconully (W. W. D.).

      _Remarks._--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_.

=Canis lupus fuscus= Richardson

Timber Wolf

      _Canis lupus_ var. _fusca_ Richardson, Mammalia, Zoölogy, 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,

      _Canis gigas_ Miller, Smithsonian Misc. Coll., 59 (no. 15):4, June
      8, 1912.

      _Canis occidentalis 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,

      _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

  [Illustration: FIG. 71. Probable past distribution of the wolf in
  Washington. A. _Canis lupus fuscus._ B. _Canis lupus columbianus._ C.
  _Canis lupus irremotus._]

_Description._--The wolf is a large carnivore of the general proportions
of a large collie dog. Wolves closely resemble coyotes but are larger,
with stouter body, larger feet, thicker muzzle and more massive,
powerful skull and teeth. The fur of wolves is long and 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 _Canis 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 Washington.
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 Columbian Plateau in historic times, and
even the deer and sheep probably 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 western 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 common and I doubt that man killed them all;
perhaps some introduced disease, such as rabies, brought about their

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, between Lake Chelan and
Mount Baker. There are no recent records for southeastern Washington.
The last wolves killed on the Columbian Plateau were two taken at
Wahluke, Grant County, on September 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 mistaken for a

The wolf no longer is an important element in the mammalian fauna of
Washington and will probably never be so again. The species, like the
grizzly bear, is nearly extinct in the state.

=Canis familiaris= Linnaeus


      _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

  [Illustration: FIG. 72. Cougar or mountain lion (_Felis concolor
  missoulensis_), skin mounted as a rug; shot on Mill Creek, Pend Oreille
  County, Washington, February 13, 1935, by Ralph Johnson. Skin now
  measures: snout to tip of tail 252 cm., span between tips of forepaws
  157 cm., tail 89 cm.; male. (Fish and Wildlife Service photo by Victor
  B. Scheffer, No. 66.)]

_Description._--The cougar is a large cat and has the general
proportions of the house cat. Large cougars measure more than seven feet
in length of which the tail comprises two feet. Large males 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

  [Illustration: 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 incessantly 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 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

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 canyon.

  [Illustration: FIG. 74. Distribution of the cougar in Washington. A.
  _Felis concolor oregonensis._ B. _Felis concolor missoulensis._]

Although the cougar is a large and powerful carnivore it almost never
makes unprovoked attacks upon man. An exception was the cougar that
partially devoured a thirteen-year old boy near Lake Chelan, Chelan
County (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 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 occasionally capture a female and several males at one
locality. Breeding occurs in almost any month of the year. The young are
cared for by the female and follow her for perhaps a year. Young
individuals 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,

      _Felis oregonensis_ 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, Leavenworth, and Goldendale.

=Felis concolor missoulensis= Goldman

      _Felis concolor missoulensis_ Goldman, Jour. Mamm., 24:229, June 8,

      _Type._--Obtained 10 miles southwest of Missoula, Missoula County,
      Montana, 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 comparison of specimens reveals differences
      of considerable magnitude between the cougars of western and
      northeastern Washington. I suppose that intergradation takes place
      in a limited area in, and west of, the Okanogan River Valley in

  [Illustration: FIG. 75. Canadian lynx, _Lynx canadensis_, male, taken
  February 16, 1939, on Baldy Mountain, northwest Idaho, by Lloyd
  Robinson of Sandpoint. (Ross A. Hall photo.)]

=Lynx canadensis canadensis= Kerr

Canadian lynx

      _Lynx canadensis_ Kerr, Anim. Kingd., 1: systematic catalogue
      inserted between pages 32 and 33 (description, p. 157), 1792.

      _Lynx borealis canadensis_ True, Proc. U. S. Nat. Mus., 7:611,

      _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.

  [Illustration: 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 yellowish 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.

  [Illustration: FIG. 77. Distribution of the Canadian lynx, _Lynx
  canadensis_, in Washington.]

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 regularly takes a dozen or
more each year. Mr. Lester Fairbrother of Oroville, Okanogan County,
regularly traps lynxes in the northern Cascades. They are taken in
wooded areas where snowshoe rabbits, their principal food in winter, are
abundant. In the more accessible 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 skins.

=Lynx rufus= (Schreber)


_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.

  [Illustration: FIG. 78. Bobcat (_Lynx rufus fasciatus_), two-year-old
  male captured as a kit near Lyman, Washington, in the spring of 1937,
  by Earl Scott; photographed March 9, 1939. (Fish and Wildlife Service
  photo by Victor B. Scheffer, No. 588.)]

Bobcats range from 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 nocturnal but is
occasionally active by day. One that I watched near Lake Samamish, King
County, when it was unaware of my presence, 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 bowlegged.

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 doubtless 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

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. Immature 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.

=Lynx rufus fasciatus= Rafinesque

      _Lynx fasciatus_ Rafinesque, Amer. Monthly Mag., 2:46, November,

      _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,

      _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

      _Measurements._--A male from Forks, Clallam County, measured:
      total length 890; length of tail 190; hind foot 167; ear 80;
      weight 24-1/2 pounds.

      _Distribution._--From the Cascade Mountains westward, including
      the valleys of rivers draining westward in the Cascades. Marginal
      occurrences are: Skykomish (W. W. D.), Mt. Rainier (Taylor and
      Shaw, 1927: 60) and headwaters of Lewis River (W. W. D.).

  [Illustration: 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,

      _Lynx rufus pallescens_ Bailey, N. Amer. Fauna, 55:268, August 29,

      _Type._--Obtained at Trout Lake, Klickitat County, Washington, by
      D. Kaegi on January 10, 1895; type in United States National

      _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.

=Zalophus californianus= (Lesson)

California sea lion

      _Otaria californiana_ Lesson, Dict. Class Nat. Hist., 13:420, 1828.

      _Zalophus californianus_ Allen, Monogr. N. Amer. Pinnipeds, U. S.
      Geol. and Geogr. Surv. Terr., Misc. Publ., 12:276, 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 swimming; 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. Females are darker than the males.

The California sea lion occurs along the Pacific Coast from Mexico to
northern California and has been recorded from Oregon (Bailey, 1936:
332), Washington and British Columbia (Greenwood, Newcombe, and Fraser,
1918: 1-39). It is of rare or casual occurrence along the coast of

=Eumetopias jubata= (Schreber)

Steller sea lion

      _Phoca jubata_ Schreber, Säugthiere, 3:300, 1776.

      _Eumetopias Stelleri_ True, Proc. U. S. Nat. Mus., 7 (1884):607,

      _Eumetopias jubata_ Allen, Bull. Amer. Mus. Nat. Hist., 16:113,
      March 15, 1902.

      _Type locality._--North Pacific Ocean.

      _Distribution._--Ocean coast, breeding on small, rocky islets such
      as the Quillayute Needles.

_Description._--The Steller sea lion resembles the California sea lion
but is larger. Old bulls 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, especially 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, deep roaring sound.

Steller sea lions occur along 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 California sea

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 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 surface with a porpoiselike roll, breathe, and
again dive. Many of their actions seem inspired by playfulness--a means
of working off excess 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 dynamiting 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 exceedingly interesting
animal and might well be preserved in numbers exceeding their present

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

=Callorhinus ursinus cynocephalus= (Walbaum)

Alaska fur seal

      _Siren cynocephalus_ Walbaum, Petri Artedi Sueci Genera Pisc., p.
      560, 1792.

      _Callorhinus ursinus_ True, Proc. U. S. Nat. Mus., 7 (1884):607,

      _Callorhinus alascanus_ Jordan and Clark, Fur Seals and Fur Seal
      Islands of the North Pacific, pt. 3, p. 2, November, 1899.

      _Callotaria 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 flippers 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 hauling-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 sealing 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 dugout canoes. The weapons are
double-headed harpoons with long cedar shafts. 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 connected 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 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. Zoöl. Soc. London, p. 28, 1864.

      _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 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 distance 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

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 drifting 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 decidedly temperamental.
It demanded constant attention and whimpered, cried or moaned if 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 its actions when it encountered strands of kelp while swimming.
If 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

The hair seal was generally thought to feed almost exclusively on
salmon, but the work of Scheffer (1928: 10-16) showed this view to be
incorrect. Of 14 hair seal stomachs examined, all of which 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. Bounties 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 investigated 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 accomplished 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.

Genus =Tamias= Illiger


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. Bryant'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 mistakenly 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 are 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 accused 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 _townsendii_ 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 uttered by
_townsendii_ with longer intervals between the notes. The call of
_amoenus_ is of an intermediate nature.

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 pectoral, 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 3-1/2 inches; the
tail about 3-1/2 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 Cascade-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 suitable food item and only
the most cautious chipmunk survives to reproduce. The least chipmunk has
been timed at a speed of 10 miles per hour (Cottam and Williams, 1943:

The food of the least chipmunk in eastern Washington is almost entirely
seeds of the annuals that flourish briefly in the spring. Insects
probably are eaten and one specimen had the remains of two scorpions in
its stomach.

=Tamias minimus scrutator= (Hall and Hatfield)

      _Eutamias minimus pictus_ Howell, N. Amer. Fauna, 52:39, November
      30, 1929.

      _Eutamias minimus scrutator_ Hall and Hatfield, Univ. California
      Publ. Zoöl., 40:321, February 12, 1934.

      _Tamias minimus scrutator_ Ellerman, Fam. and Genera of Living
      Rodents, British Mus. Nat. Hist., p. 432, June 8, 1940.

  [Illustration: FIG. 80. Distribution of the least chipmunk in
  Washington. A. _Tamias minimus scrutator._ B. _Tamias minimus

      _Type._--Obtained near Blanco Mountain, 10500 ft. elevation, Mono
      County, California, by J. Grinnell on July 28, 1917; type in Museum
      of Vertebrate Zoölogy.

      _Racial characters._--Buffy color; wide dark stripes and narrow
      pale stripes.

      _Measurements._--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._--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 of the Great Basin Fauna that 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
      least chipmunks from Oregon and Nevada.

=Tamias minimus grisescens= (Howell)

      _Eutamias minimus grisescens_ Howell, Jour. Mamm., 6:52, February
      9, 1925.

      _Tamias minimus grisescens_ Ellerman, Fam. and Genera of Living
      Rodents, British Mus. Nat. Hist., p. 431, June 8, 1940.

      _Type._--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 occurrences are Douglas
      (Howell, 1929: 41), Vantage (V. B. S.) and Pasco (Howell, 1929:

_Remarks._--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 chipmunk
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. Separation from _minimus_ is more
difficult but, in the Washington races of _minimus_, the colored fur of
the underside of the tail is pale yellowish 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 lowlands. Such conditions are
present in some places along the eastern base of the Cascades, the
inter-mountain river valleys of northeastern Washington, and along the
central-eastern border of the state.

Yellow-pine chipmunks are 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 chipmunks, 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 abundance 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 observer'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
individuals, 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 altitude 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 are no fatter than those
taken in the spring. Svihla (1936B: 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 coincide with the coming of winter
weather. In mid-November of one year, when no snow had yet 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 9, 1903, after the ground was
"well frozen," but apparently free of snow (Howell, 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 huckleberry
(_Vaccinium parvifolium_), salmonberry (_Rubus spectabilis_),
thimbleberry (_Rubus parviflorus_), devil's club (_Fatsia horrida_), and
mountain ash (_Sorbus cascadensis_ 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 entrance was a hole 1-1/2
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 beneath the sod, 4 feet from the entrance.
The nest chamber was 7 inches high by 7-1/2 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,

      _Tamias amoenus caurinus_ Ellerman, Fam. and Genera of Living
      Rodents, British Mus. Nat. Hist., p. 432, June 8, 1940.

      _Type._--Obtained at timber-line, head of Sol Duc River, Olympic
      Mountains, 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.

      _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,

=Tamias amoenus felix= Rhoads

      _Tamias quadrivittatus felix_ Rhoads, Amer. Nat., 29:941, October,

      _Eutamias quadrivittatus felix_ Miller and Rehn, Proc. Boston Soc.
      Nat. Hist., 30:44, December 27, 1901.

      _Eutamias amoenus felix_ Howell, Jour. Mamm., 3:184, August 4,

      _Tamias amoenus felix_ Ellerman, Fam. and Genera of Living
      Rodents, British Mus. Nat. Hist., p. 432, June 8, 1940.

  [Illustration: FIG. 81. Distribution of the yellow-pine chipmunk in
  Washington. A. _Tamias amoenus luteiventris._ B. _Tamias amoenus
  canicaudus._ C. _Tamias amoenus affinis._ D. _Tamias amoenus
  ludibundus._ E. _Tamias amoenus felix._ F. _Tamias amoenus caurinus._]

      _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.

      _Racial characters._--Large size and rich, dark color.

      _Measurements._--Ten topotypes average: total length 224.7; length
      of tail 98.8; hind foot 34.1; ear 14.4.

      _Distribution._--The extreme northwestern Cascades, north and west
      of Mt. Baker.

_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., 56 (no.
      26):1, December 5, 1911.

      _Eutamias amoenus ludibundus_ Howell, Jour. Mamm., 3:184, August
      4, 1922.

      _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

      _Racial characters._--Moderate size, brownish rump, ochraceous

      _Measurements._--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

      _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,

      _Remarks._--At the higher altitudes this race seems to be the
      equivalent of _affinis_. The latter race lives in relatively arid
      yellow pine forests and _ludibundus_ 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, December 27, 1901.

      _Eutamias amoenus affinis_ Howell, Jour. Mamm., 3:184, August 4,

      _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 underparts.

      _Measurements._--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 records 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,

=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.

      _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 buffy underparts.

      _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 Soc. Nat. Hist., 30:44, December 27, 1901.

      _Eutamias amoenus luteiventris_, Howell, Jour. Mamm., 3:179,
      August 4, 1922.

      _Tamias amoenus luteiventris_ Ellerman, Fam. and Genera of Living
      Rodents, British Mus. Nat. Hist., p. 432, June 8, 1940.

      _Type._--Obtained at Chief Mountain Lake (Waterton Lake), Alberta
      (3-1/2 mi. N. United States boundary) by Elliott Coues on August
      24, 1874; type in United States National Museum.

      _Racial characters._--Small size, rich color, buffy underparts.

      _Measurements._--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, west 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 simulans_ Howell, Jour. Mamm., 3:179, August
      4, 1922.

      _Tamias ruficaudus simulans_ Ellerman, Fam. and Genera of Living
      Rodents, British Mus. Nat. Hist., p. 434, June 8, 1940.

      _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. a. luteiventris_ and _T. a. canicaudus_ it differs
principally in larger size, wider brain case and especially in its pure
white underparts. From _T. a. affinis_ it differs in richer coloration,
especially the brownish rather than gray rump. The differences
separating it from _amoenus_ are slight, and only adult specimens can be
identified in the field.

According to Howell (1929: 81) this species occurs in northern Idaho,
western Montana, northeastern Washington, southeastern British Columbia
and extreme southwestern Alberta. Two subspecies 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
_ruficaudus_. Also, there are a number of specimens that I cannot
definitely 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 chipmunks
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 chipmunks
that occur in Washington. Head and body measure about 5-1/2 inches; the
tail about 4-1/2 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 blackish. The upper pale stripe
is pale buffy gray; the lower is whitish. The tail is blackish frosted
with white above and rich ochraceous below. Underparts are dull white.

The Townsend chipmunk ranges from the Fraser River in southern 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 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_.

  [Illustration: FIG. 82. Townsend chipmunk (_Tamias townsendii
  cooperi_), captured on Goat Creek, 3,000 feet, western Cascade
  Mountains near Chinook Pass, Washington, September 16, 1940, by
  Earl J. Larrison; photographed February 1, 1941. (Fish and Wildlife
  Service photo by Victor B. Scheffer, No. 1139.)]

Townsend chipmunks are closely associated with the coniferous forest
where they live in clearings and tangles of underbrush such as on steep
hillsides, fire slashings, snowslide 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 berries 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 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 chipmunk 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.

  [Illustration: FIG. 83. Distribution of the Townsend chipmunk in
  Washington. A. _Tamias townsendii townsendii._ B. _Tamias townsendii

Shaw (1944: 278) discovered the brood nest of a Townsend chipmunk 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 outside 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.

=Tamias townsendii townsendii= Bachman

      _Tamias townsendii_ Bachman, Jour. Acad. Nat. Sci. Philadelphia, 8
      (pt. 1):68, 1839.

      _Tamias quadrivittatus townsendii_ Allen, Proc. Boston Soc. Nat.
      Hist., 16:290, 1874.

      _Tamias asiaticus_ var. _townsendii_ Allen, Monog. N. Amer.
      Rodentia, Rept. U. S. Geol. Surv. Terr., 11:794, 1877.

      _Eutamias townsendii_ Merriam, Proc. Biol. Soc. Washington,
      11:195, July 1, 1897.

      _Type._--Lectotype obtained near the lower mouth of the Willamette
      River, Multnomah County, Oregon, by J. 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

      _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 townsendii_ 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 Washington, 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.
      townsendii_, are: Hamilton (U. S. N. M.), 5 mi. E. Monroe (W. W.
      D.), Redmond (W. W. D.), Roy (U. S. N. M.), and Vancouver (U. S.
      N. M.).

=Tamias townsendii cooperi= Baird

      _Tamias cooperi_ Baird, Proc. Acad. Nat. Sci. Philadelphia, 7:334,

      _Tamias townsendii_ var. _cooperi_ Baird, Mamm. N. Amer., p. 737,

      _Eutamias cooperi_ Lyon, Smithsonian Misc. Coll., 50:89, June 27,

      _Eutamias townsendii cooperi_ Howell, Jour. Mamm., 3:184, 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. townsendii_ but paler with
      pale stripes whitish rather than cinnamon.

      _Measurements._--Ten males and 10 females from the Cascades
      average, respectively: 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

      _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. Helens (U. S. N. M.), and Yacolt (M. V.

=Marmota monax petrensis= Howell


      _Marmota monax petrensis_ Howell, N. Amer. Fauna, 37:33, April 7,

      _Type._--Obtained at Revelstoke, British Columbia, by W.
      Spreadborough on May 12, 1890; type in United States National

      _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: 173-175)
lists seven subspecies, one of which enters the extreme 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 observations on the habits, nests
and burrows of _Marmota monax canadensis_.

=Marmota flaviventris avara= (Bangs)

Yellow-bellied marmot

      _Arctomys flaviventer avarus_ Bangs, Proc. New England Zoöl. Club,
      1:68, July 31, 1899.

      [_Marmota flaviventer_] _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 Zoölogy.

      _Measurements._--A male from 5 miles north of Entiat, Chelan
      County, measured: total length 610; length of tail 182; hind foot
      70; ear 28.

      _Distribution._--From the eastern edge of the Cascade Mountains
      eastward, except, apparently, extreme northeastern Washington.
      Marginal records are:

      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 woodchuck
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
having a distinct white bar on the nose. Its pelage is coarse and rather

The yellow-bellied marmot is typically an animal of the basalt talus of
eastern Washington but occurs in mountainous areas in northeastern
Washington. These animals are usually found near streams, ponds, lakes,
or rivers. They wander considerably, however, and are often found far
from water. Their wandering habits probably account for their presence
near temporary streams and ponds on the Columbian Plateau. When these
temporary sources of water dry up in July or early August, the marmots
go into hibernation. Edson (1935: 68) records a marmot from Bellingham,
Whatcom County, far west of the usual range of the species.

The "ground hog" is often hunted for sport and sometimes for food. Near
centers of human population the yellow-bellied marmots are extremely
shy. Along the highways of the Columbia River on any Sunday in June, it
is not unusual to see a dozen cars in an hour, moving slowly past a
talus slide while eager hunters scan the rocks for marmots. In the late
afternoon, when the marmots leave the protection of the talus slides to
drink at the river, they fall easy prey to rifles with telescope sights.
Near cities in eastern Washington yellow-bellied marmots have become
partially nocturnal.

Couch (1930: 2-6) attempted to excavate several dens of yellow-bellied
marmots, but decided to leave the task "to some future road-building
crew." Embryos found by Couch numbered three to six. Couch thought the
young were born about March 15 in the Snake River area and about April
15 in the upper Okanogan area. The young appear above ground
approximately 30 days after birth.

The yellow-bellied marmots enter aestivation from late June to early
August, depending on the locality and local conditions. They are active
longer in northeastern Washington. Couch records a yellow-bellied marmot
seen in Okanogan County on October 10, but regards this as exceptional.
The marmots near Wenatchee, Chelan County, emerge from their burrows in
early March (March 5 to 10, from reports of residents in 1937, 1938).
Couch (1930: 5) gives February 20 to March 15 as the date of

A principal requirement for marmots is the presence of rocks. At Cle
Elum, Kittitas County, I took a marmot from an alfalfa field where a
farmer had placed all the surface stones in a loose pile. Fifteen miles
east of Tonasket, Okanogan County, marmots were living in the stones
piled by road builders to support the ends of a small bridge. A hundred
feet away another marmot was living under an abandoned building. A high,
convenient rock near their burrow serves the yellow-bellied marmots as a
look-out post. These look-out posts seem, in many cases, to have been
used by many generations of marmots, for their feces sometimes fill
nearby crevices to a depth of several feet and cover the look-out rock
itself. The glacial boulders on the plateau between the Okanogan River
and Omak Lake, Okanogan County, furnish the best examples of look-out
posts. These numerous isolated boulders, ten to fifty feet in diameter
and ten to thirty feet high, each seem to furnish shelter to one or more
marmots. Well-worn trails lead from the boulders to burrows and feeding

The food of the yellow-bellied marmot includes grasses and succulent
plants found in their habitat. Fondness for alfalfa makes them a serious
pest in some areas, for their large size enables them to make
considerable inroads on a field. Natural enemies probably include most
larger predaceous mammals. Hawks and eagles probably kill their young.
Near Tonasket, Okanogan County, Robert Dalquest surprised a coyote as it
ran across a small wash. A shot caused the coyote to drop a half-grown
marmot which it had been carrying.

=Marmota caligata cascadensis= Howell

Hoary marmot

      _Marmota caligata cascadensis_ Howell, Proc. Biol. Soc. Washington,
      27:17, February 2, 1914.

      _Type._--Obtained on Mt. Rainier, 6,000 ft., Pierce County,
      Washington, by A. K. Fisher on August 11, 1897; type in United
      States National Museum.

      _Measurements._--A young adult male from Stevens Pass, King
      County, measures: total length 773; length of tail 221; hind foot
      93; ear 27.

      _Distribution._--The Cascade Mountains, ranging from Mt. Baker (W.
      W. D.) and Mt. Chopaka (Taylor and Shaw, 1929: 15) south to Mt.
      Adams (Taylor and Shaw, 1929: 15).

_Description._--The hoary marmot is the largest of the American marmots;
adults are 28 inches or more in length, of which the tail makes up about
8 inches. The pelage is dense and rather woolly. The upper parts are not
frosted with white-tipped hairs, but are gray washed with blackish. The
head is blackish with white facial markings, and the shoulders, legs
and underparts are gray. The tail is dark reddish brown.

Hoary marmots range from Alaska south to Washington and Idaho. Howell
(1915: 57-67) recognizes seven races of this species.

  [Illustration: FIG. 84A. Distribution of marmots in Washington. A.
  _Marmota monax petrensis._ B. _Marmota flaviventris avara._
  C. _Marmota caligata cascadensis._ D. _Marmota olympus._]

This mammal of the higher altitudes rarely goes below the Hudsonian
Life-zone. It is most common in the talus slides at the lower edge of
the Arctic-Alpine Life-zone. Like other marmots, it prefers to live amid
loose boulders. The steep talus or "scab rock" slides in the glacial
cirques provide an ideal habitat. The crevices and caves beneath the
rocks offer concealment for young and adults. A large boulder with a
flat top is usually selected as a look-out. Well-worn trails lead from
the talus slides to nearby grassy slopes. When surprised in the open,
the hoary marmot exhibits a peculiar bounding run, reminiscent of that
of tree squirrels. The short tail "follows through" in the leaps. In the
rock slides the hoary marmot is surprisingly agile.

The clear, shrill whistle of the hoary marmot is familiar to all who
penetrate its haunts. The whistle is remarkably similar to the whistle
of a person. Locally the hoary marmot is known as "whistler" or "whistle
pig." Individuals emerge from hibernation early in June; most adults
retire again by the middle of September. On September 14, 1937, a young
of the year was shot and few were seen where they had been common in
June. All were shy. Only one adult was seen.

Large hawks and eagles commonly hunt over the rock slides inhabited by
marmots, and probably kill very young individuals. Only the larger
predators such as bear, cougar, wolf, coyote, lynx, and bobcat would be
expected to kill an adult hoary marmot.

=Marmota olympus= (Merriam)

Olympic marmot

      _Arctomys olympus_ Merriam, Proc. Acad. Nat. Sci. Philadelphia, p.
      352, October 4, 1898.

      [_Marmota_] _olympus_ Trouessart, Catal. Mamm., viv. foss. suppl.,
      p. 344, 1904.

      _Type._--Obtained at timber-line, head of Sol Duc River, Clallam
      County, Washington, by C. H. Merriam, on August 27, 1897; type in
      United States National Museum.

      _Measurements._--Two small but adult females from Deer Park,
      Clallam County, measure respectively: total length 758, 691;
      length of tail 163, 161; hind foot 106, 97; ear 31, 29.

      _Distribution._--This species is confined to the Olympic

_Description._--The Olympic marmot closely resembles the hoary marmot,
and differs in being slightly larger, and reddish or rusty brown where
the hoary marmot is gray. It lacks the blackish overwash of the hoary
marmot. The nose is whitish.

Although the Olympic marmot belongs to the hoary marmot group it is a
distinct species, most closely related to the marmot of the mountains of
Vancouver Island. Its habits differ but little from those of the hoary
marmot. It lives in burrows in talus slides and boulder piles near
timber line. A few burrows are dug beneath logs. Well worn trails lead
from burrows to feeding grounds on nearby grassy slopes and heather
meadows. High rocks or logs serve as lookouts. The alarm whistle is
similar to that of the hoary marmot.

=Citellus townsendii townsendii= (Bachman)

Townsend ground squirrel; sage rat

      _Spermophilus townsendii_ Bachman, Jour. Acad. Nat. Sci.
      Philadelphia, 8:61, 1839.

      _Spermophilus mollis yakimensis_ Merriam, Proc. Biol. Soc.
      Washington, 12:70, March 24, 1898.

      [_Citellus mollis_] _yakimensis_ Trouessart, Cat. Mamm., Sup., p.
      339, 1904.

      _Citellus townsendii townsendii_ Howell, N. Amer. Fauna, 56:60,
      May 18, 1938.

  [Illustration: FIG. 84B. Distribution of the Townsend and Washington
  ground squirrels in Washington. A. _Citellus townsendii townsendii._
  B. _Citellus washingtoni._]

      _Type._--Obtained on the western bank of the Columbia River "about
      300 miles above its mouth" (probably near the mouth of the Walla
      Walla or Touchet river; more exactly, opposite Wallula, in Benton
      County, Washington) by J. K. Townsend in July, 1836; type in
      Academy of Natural Sciences of Philadelphia.

      _Measurements._--Ten adults from Mabton and North Yakima, Yakima
      County, average: total length 212.2; length of tail 45.7; hind
      foot 33.9.

      _Distribution._--The Upper Sonoran Life-zone area of the Yakima
      Valley from Ellensburg (Howell, 1938: 63) south to the Columbia at
      Kennewick (W. W. D.).

_Remarks._--This race is probably derived from _C. t. mollis_ of Oregon,
from which it is but slightly differentiated. It was known for many
years as _Citellus mollis yakimensis_. For use of the name _townsendii_
see Howell (1938: 62).

The head and body are about 6-1/2 inches long and the tail is about 2
inches in length. The head is large with low, rounded ears and large
eyes. The neck is distinctly constricted and the body is plump. Adults
are "pot-bellied." The legs are short and the feet small. The tail is
short, round at the base but bushy throughout the rest of its length.
The pelage is short and rather harsh. In color the upper parts are
yellowish gray flecked with tiny, pale dots. The underparts are buffy.
The face, thighs and tail are reddish buff.

Ground squirrels occur in parts of Europe, Asia, and much of western
North America. Howell (1938: 36-37) recognizes eight subgenera and
thirty-one species in North America. Three subgenera occur in
Washington, namely: _Citellus_, the spotted ground squirrels,
represented by _Citellus townsendii_, _Citellus washingtoni_ and
_Citellus columbianus_; _Otospermophilus_, the long-tailed ground
squirrels, represented by _Citellus beecheyi_; and _Callospermophilus_,
the striped ground squirrels, represented by _Citellus lateralis_ and
_Citellus saturatus_.

All ground squirrels are diurnal and consequently are familiar to man.
Most species bear local names and the striped or mantled ground
squirrels are often mistaken for chipmunks. Some species are destructive
to crops, especially grain. In addition they harbor sylvatic plague.
Economically the genus _Citellus_ is of great importance in Washington.

All Washington ground squirrels live in burrows which they construct
themselves. The life histories of the two striped species are less well
known than those of the more economically important species. All species
hibernate. The striped species disappear in October and reappear the
following March. This can be called true hibernation. The long-tailed
ground squirrels probably hibernate at high altitudes but aestivate in
more arid localities. In Washington the Douglas ground squirrel occupies
an area that is relatively temperate and humid. They may hibernate or
aestivate, depending on local conditions, or they may remain active all
year. The spotted ground squirrels disappear in midsummer and sleep
(aestivate) until the following spring, for in their arid habitat the
disappearance of green food in late summer and fall makes living
conditions almost as unfavorable as in winter.

Townsend ground squirrels range from central Washington south to
southern Nevada and from the Cascade Mountains east to eastern Idaho and
central Utah. Five races are recognized, only one of which occurs in
Washington, where it is confined to the Upper Sonoran Life-zone. It
occupies the sagebrush area, being most common where the sage is in
scattered clumps separated by grassy areas. Occasionally these squirrels
occupy extensive grasslands where sage is scarce and in the Yakima area
may enter pastures and fields. They live in colonies, often with more
than ten separate burrows to the acre.

Burrows are dug in dusty ground, either near sage bushes or in openings
among them. Well-worn trails lead from feeding places to openings of
burrows. Mouths of burrows are usually situated on flat ground and are
surrounded by a rim of excavated earth four to six inches high. If
located on a slope, the excavated earth collects as a mound on the lower
side, and serves as a look-out post for the squirrels. Burrows excavated
by Scheffer are described by Howell (1938: 5). One reached a depth of
5-1/4 feet at a point where it branched 11 feet from the entrance. One
branch led to the nest chamber three feet to the right of the main
tunnel. The other reached a depth of six feet at a point 14 feet from
the entrance, then turned upwards at a 70 degree angle and reached the
surface through a partly obstructed entrance, 12 feet from the original
entrance. The nest chamber was 6-1/2 inches in diameter and filled with
a perfectly dry nest of fine grass, partly broken and shredded.

Food consists of soft green vegetation and seeds. Foods listed by Howell
(1938: 5) include: _Sphaeralida munroana_, _Plantago purshii_, _Bromus
tectorum_, _Agropyron pauciflorum_, _Oryzopsis hymenoides_, _Norta
altissima_, _Artemisiae spinescens_, sunflower, alfalfa, wheat, barley,
potato, beets, carrots, lettuce, and insects (grasshoppers, cicadas). In
1917, these squirrels were reported to have practically destroyed a
ten-acre field of beets at White Swan, Yakima County.

Aestivation of older individuals begins in late May and the last young
disappear in early June. The squirrels appear again in late January,
before the snow has disappeared (Scheffer, 1941: 272).

The voice of the Townsend ground squirrel is a faint, high pitched
"pe-eee-ep," of remarkable carrying power for so faint a sound, and
extremely difficult to trace to its source. Scheffer (in Howell, _op.
cit._, p. 6) mentions also a chirping sound and a chatter of alarm which
I have not heard.

Five to seven embryos were found in specimens collected near Yakima.
Scheffer (1941: 270) found the number of fetuses in 52 pregnant females
from the Kennewick area to vary from 4 to 16 with an average of 8.6.
Francis (1922: 5) reported tularemia in this species. They probably
harbor also sylvatic plague.

=Citellus washingtoni= Howell

Washington ground squirrel; sage rat

      _Citellus washingtoni washingtoni_ Howell, N. Amer. Fauna, 56:69,
      May 18, 1938.

      _Citellus washingtoni loringi_ Howell, N. Amer. Fauna, 56:71, May
      18, 1938 (type from Douglas, Douglas County, Washington).

      _Type._--Obtained at Touchet, Walla Walla County, Washington, by C.
      P. Streator on May 18, 1891; type in United States National Museum.

      _Distribution._--The Columbian Plateau and southward into Oregon,
      ranging from Farmer (Howell, 1938: 71) and Moses Coulee (W. W. D.)
      south to Wallula (M. V. Z.).

      _Measurements._--Fifteen specimens from the vicinity of the type
      locality average: total length 229.2; length of tail 50; hind foot
      35.3. A male from 4 miles west of Pasco, Franklin County, weighed
      201 grams.

      _Remarks._--_Citellus w. loringi_ allegedly differs from typical
      _washingtoni_ in smaller size. The specimens collected by us from
      within the range of _loringi_ are as large as those from farther
      south. _C. w. loringi_ is regarded as a synonym of _C. w.

_Description._--The Washington ground squirrel closely resembles the
Townsend ground squirrel in size and appearance. It differs principally
in color. The upper parts are brownish gray marked with distinct white
spots, and the underparts are buffy. The face, thighs and tail are dull

This species is closely related to the Townsend ground squirrel but its
habitat is more varied. It is most common in areas of low sage bushes
surrounded by grasslands and on extensive grasslands, but occurs also in
sandy places, wheat fields, and rocky hillsides. The animals live in
colonies, in some places 50 or more individuals to the acre. They occur
also as scattered individuals or small colonies in smaller areas of
suitable habitat. For example, along the highway from Farmer, Douglas
County, to Waterville, a distance of 15 miles, a band of natural grass,
50 feet or less in width, lies between the road and the extensive wheat
fields beyond. This strip of natural cover possessed about one
_Citellus_ to each 200 feet along most of its length. The squirrels were
often seen sitting on rolls of wire drift fence, used to keep snow
drifts from the road in winter but in summer rolled into bundles 3 feet
in diameter and left standing at intervals of one each 100 feet. The
squirrels had constructed their burrow entrances beneath these rolls and
used the rolls as look-out posts. At my approach they would dodge into
the rolls of wire and, unless scared, remained in the wire roll. By
approaching carefully and overturning the rolls I was able to trap and
capture a good series of living specimens.

Burrows, nests, habits, and food of this species seem identical to those
of _townsendii_. Aestivation dates seem to be the same (Scheffer, 1941:
270-279). The hibernation of this species has been discussed in detail
by Svhila (1939: 6-10). Food species listed by Scheffer at Wallula
(Howell, 1938: 8) are identical to those of _townsendii_ at Kennewick,
across the Columbia River (see account of _townsendii_).

Scheffer (1941: 270-279) examined 26 pregnant females of this species
and found the fetuses to vary from 5 to 11 in number, with an average of

=Citellus columbianus= (Ord)

Columbian ground squirrel

_Description._--The Columbian ground squirrel has the general body
proportions, large head, large eyes, low, rounded ears, plump body and
short tail of the Townsend ground squirrel, but is much larger. The head
and body of an adult measure about 10 inches and the tail about 4
inches. The upper parts are grayish buff mottled with round white dots.
The underparts and forefeet are pale ochraceous. The back of the head
and neck are gray. The face, thighs and tail are reddish.

The Columbian ground squirrel occupies inland mountainous areas from
central British Columbia to central Oregon and Idaho and from eastern
Washington and Oregon to western Montana and southwestern Alberta. It is
closely related to the arctic ground squirrels of the _Citellus parryii_
group, which are in turn related to the Siberian forms, _buxtoni_ and

The habitat of the Columbian ground squirrel is varied but is usually
more humid than that of the Townsend and Washington ground squirrels.
The most arid of the lands occupied by them in Washington are the
grasslands and wheat fields of the Poulouse country and the eastern
border of the state. In northeastern Washington they live in meadows and
grasslands in the valleys, in openings in the coniferous forest at
higher altitudes and in parks and alpine meadows almost to timber-line
on the mountains. Some individuals were in brushy places or even in
forests, far from grassy clearings. Near Republic, Ferry County, a
Columbian ground squirrel was killed as it ran along a log in dense
larch woods. Its burrows were found in a tangle of brush and fallen
logs. No clearings, meadow or grassland existed nearby. Narrow bands of
cleared land beside roads and railroad tracks are extensively used by
these squirrels. Where small meadows or pastures occur, this species
lives in dense colonies. If grasslands are extensive it lives in loose
colonies, often with considerable distances between individual burrow
systems. In the mountains the squirrels are scattered or live in small

Burrows of the Columbian ground squirrel are often constructed in the
open. Mouths of burrows, especially if the animals are living in
colonies, usually are marked by a large pile of excavated earth. If,
however, there is some large object on the surface of the ground, such
as a stone, stump, or log, the entrance to the burrow is located beside
this. Such objects are often undermined and made to settle, eventually
becoming buried. The squirrels live under houses, and the yards of
abandoned ranch buildings are often infested with them.

The burrows of the Columbian ground squirrel have been described by Shaw
(1919, 1924B, 1925, A, B, E, 1926) and Bailey (1918: 47). Two types of
burrows are constructed, summer burrows and hibernation dens. Summer
burrows are used year after year, perhaps by succeeding generations of
squirrels. They vary considerably but usually are 3 or 4 inches in
diameter and possess several entrances. The depth to which the burrow
descends, as well as the number of forks and subsidiary burrows, depends
to some extent on the whims of the individual squirrel and the length of
time the burrow has been occupied. Usually the burrow reaches a depth of
four feet and two or more branches are present. A turning-about chamber
is present, not far from the entrance. Here a startled squirrel can turn
about and watch the entrance and, if the alarm was false, the squirrel
resumes its activities above ground. Nests are above the level of the
main burrow system and thus are protected from flooding by rain or
melting snow. Stored food and excrement are deposited in separate
compartments. Escape entrances, concealed in grass and weeds, for use in
case the burrow is invaded, are dug from beneath the surface of the
ground and thus are unmarked by any telltale mound of earth.

Separate dens are used for aestivation. While the squirrel is in
aestivation the entrances to the summer burrow are solidly plugged with
tamped earth. Aestivation dens are short and may or may not communicate
with the summer burrow. Shaw (1925B: 58) measured 50 such dens and found
the aestivation cell to average 2 feet 6 inches beneath the surface with
extreme depths of 6 inches and 4 feet 11 inches. Ordinarily a drainage
burrow is dug beneath the level of the nest.

  [Illustration: FIG. 85. Distribution of the Columbian and Beechey
  ground squirrels in Washington. A. _Citellus columbianus columbianus._
  B. _Citellus columbianus ruficaudus._ C. _Citellus beecheyi douglasii._]

The time and length of aestivation is variable. Old males enter
aestivation before the females and young, and emerge earlier in the
spring. In the lowlands, as about Pullman, Whitman County, the squirrels
begin to disappear in mid July. In the mountains of northeastern
Washington they are active until late August. They emerge again in late
February or early March. Shaw (1925B) mentions that food is occasionally
stored in aestivation dens of male squirrels. This food is not eaten in
the winter but in the spring, when the squirrels awaken.

The gestation period was determined by Shaw (1925C: 108) as 24 days. Two
to five, rarely seven, young are born. The average litter is about 3.5.
Mating occurs in late March and the young are born about the middle of

Most green vegetation occurring in its habitat is food for the Columbian
ground squirrel. Bulbs, seeds, fruit, berries, grain, clover, alfalfa,
and garden truck are eagerly eaten. The squirrels are especially fond of
wheat and great damage results from their depredations. In northeastern
Washington some wheat fields are almost entirely destroyed. Shaw (1925G)
showed that one squirrel destroyed an average of more than 50 pounds of
wheat in a season. When populations range as high as ten squirrels to
the acre, little wheat is left.

In Montana the Columbian ground squirrel is a known reservoir of Rocky
Mountain spotted fever (Birdseye, 1912: 1-46) while sylvatic plague has
been reported in this species in Oregon (Meyer, 1936: 965). Control of
the Columbian ground squirrel by traps, poison, and shooting is only
locally successful.

=Citellus columbianus columbianus= (Ord)

      _Arctomys columbianus_ Ord, Guthrie's Geography, 2d American
      Edition, 2:292 (description, p. 303), 1815.

      _Spermophilus columbianus_ Merriam, N. Amer. Fauna, 5:39, July 30,

      _Citellus columbianus_ Allen, Bull. Amer. Mus. Nat. Hist., 19:536,
      October 10, 1903.

      _Type._--None designated. Description based on Lewis and Clark's
      account of animals taken by them between the forks of the
      Clearwater and Kooskooskie rivers, Idaho.

      _Racial characters._--Smaller size and paler color than in

      _Measurements._--Five females from northeastern Washington
      average: total length 346; length of tail 77; hind foot 48.6; ear

      _Distribution._--Northeastern Washington, from 15 mi. E. Tonasket
      (W. W. D.) east to Pass Creek Pass (U. S. N. M.) and south to
      Pullman (W. S. C.) along the eastern edge of the state.

=Citellus columbianus ruficaudus= Howell

      _Citellus columbianus ruficaudus_ Howell, Proc. Biol. Soc.
      Washington, 41:212, December 18, 1928.

      _Type._--Obtained at Wallowa Lake, Wallowa County, Oregon, by G. G.
      Cantwell on April 13, 1919; type in United States National Museum.

      _Racial characters._--Similar to _Citellus columbianus
      columbianus_ but tail redder, especially above, and red color
      everywhere deeper.

      _Distribution._--Restricted to the Blue Mountains of extreme
      southeastern Washington.

      _Remarks._--This is at best a slightly differentiated race and
      upon further study, animals assigned to it may prove unworthy of
      recognition as a distinct subspecies.

=Citellus beecheyi douglasii= (Richardson)

Beechey ground squirrel

      _Arctomys? (Spermophilus?) douglasii_ Richardson, Fauna
      Boreali-Americana, 1:172, 1829.

      _Spermophilus douglasii_ F. Cuvier, Sup. a l'hist. natur. Buffon,
      1:333, 1831.

      [_Spermophilus grammurus_] var. _douglasii_ Allen, Proc. Boston
      Soc. Nat. Hist., 16:293, 1874.

      _Citellus v[ariegatus]. douglasii_ Elliot, Field Columb. Mus.
      Publ. 76, zoöl. ser. 3:183, May, 1903.

      _Citellus beecheyi douglasi_ Grinnell, Proc. California Acad.
      Sci., 3 (ser. 4):345, August 28, 1913.

      _Otospermophilus grammurus douglasii_ Miller, U. S. Nat. Mus.
      Bull., 128:18, April 29, 1924.

      _Citellus douglasii_ Taylor and Shaw, _Occ._ Papers Chas. R.
      Conner Mus., no. 2:15, December, 1929.

      _Citellus beecheyi douglasii_ Howell, N. Amer. Fauna, 56:150, May
      18, 1938.

      _Type._--None. Description based on a hunter's skin from "Banks of
      the Columbia," probably near The Dalles, Wasco County, Oregon.

      _Measurements._--A male and 6 females from Klickitat County,
      average, respectively: total length 469, 480; length of tail 181,
      198; hind foot 58, 62; ear 26, 26.

      _Distribution._--The Columbia River Valley from Goldendale (W. W.
      D.) west to the White Salmon River (W. W. D.).

_Remarks._--Ground squirrels have been common on the Oregon side of the
Columbia River for a long time. The specimen upon which the description
was based was presumably obtained there in the early 1800's. Yet the
species did not become established in Washington until 100 years later,
in about 1915.

The head is large with eyes and ears of moderate size. The body is
rather stout; thicker than that of a tree squirrel but slimmer than that
of the Columbian ground squirrel. Head and body are about 11 inches in
length and the tail is about 7 inches long. The upper parts are dark
brownish or blackish gray heavily marked with white spots. A triangular
area at the shoulders is clear black outlined in whitish. The head and
neck are grayish brown. The underparts are buffy. The tail is gray above
and buffy beneath.

This species belongs to the subgenus _Otospermophilus_. Externally the
species _beecheyi_ differs from _Citellus washingtoni_, _townsendii_,
and _columbianus_, all of which belong to the subgenus _Citellus_, in
possessing more slender limbs, a longer, thinner body, a rather bushy
tail that is nearly two-thirds, rather than less than half, the length
of the body, and larger, more prominent ears. The Beechey ground
squirrel lacks also the hazel color of nose and thigh that is
characteristic of the subgenus _Citellus_ in Washington.

_Citellus beecheyi_ ranges from the southern edge of the state of
Washington southward through western Oregon and California into northern
Lower California. It is a western coastal species and reaches eastward
only as far as Nevada. A related species, _Citellus variegatus_, occurs
farther inland, from Utah, Colorado, and Texas, southward to central
Mexico. Eight subspecies of _Citellus beecheyi_ are recognized by Howell
(1938), all but three of which are restricted to California.

At present the Beechey ground squirrel occupies a limited area of
Washington, which it has invaded in recent years (Scheffer and Dalquest,
1939: 44). However, it is extending its range and may be expected
eventually to occupy a considerable part of the state.

The habitat of this ground squirrel is varied. It occupies a more humid
terrain than do most members of the genus. At the present time it is
most common in grassy fields and rocky outcrops along the Columbia
River. It is common also in the open oak groves on the hillsides back
from the river valley and some individuals were seen near Guler, well up
on the slopes of Mount Adams. As observed in Washington, its favored
habitat is about rock outcrops and talus slides near extensive fields or
grasslands. In California and Oregon, however, it occurs among other
places, in scattered fields and meadows of the coniferous forests in the
humid subdivision of the Transition Life-zone. The range of tolerance of
the species is such that it might extend its range into much of western
Washington, including the Puget Sound area. It may be expected to move
northward through the eastern Cascades.

Because of its limited range and the short time of its establishment,
the Beechey ground squirrel has been studied little in Washington.
According to Howell (1938: 28), this race is less prolific and abundant
than other members of the species. Of the Beechey ground squirrel,
certainly the subspecies _C. b. douglasii_, is less common, locally and
over wide areas, in its range in California and Oregon, than are the
subspecies _C. b. beecheyi_ and _C. b. fisheri_ in California. In
Washington, on the other hand, _douglasii_ is locally abundant, perhaps
more so than elsewhere.

The habits of this race in California have been studied by Grinnell and
Dixon (1919: 595-807). Burrows have been excavated near Eugene, Oregon,
and the findings published (Edge, 1934: 189-193). Burrows were dug on
sloping or well-drained ground. Mounds of earth at the entrances were
usually inconspicuous but well-worn trails led to the burrows. The
burrows entered the ground at an angle of 35° for about two feet, then
flattened out horizontally for six feet or more. The burrows were from 1
to 4 feet deep, usually about 2 feet. Ordinary burrows had a single
entrance but some older burrows had two or more. Burrows branched
frequently. Nest chambers were of the shape of a flattened sphere 10
inches in diameter. Frequently more than one nest was found in a burrow,
but only one was in use at a time. Some nests were infested with fleas.
Nests frequently contained food. Most of the burrows observed by me had
their entrances beside rocks or oak roots. In alfalfa fields near Bingen
and Lyle, Klickitat County, the burrows were in open fields and the
entrances were marked by large mounds of earth. In actual diameter the
burrows seemed smaller than those of _Citellus columbianus_.

Aestivation and hibernation of the ground squirrels in the lower
Columbia River Valley seem unnecessary, for the climate is temperate and
humid. It is not established that _douglasii_ hibernates in other parts
of its range where the climate is mild. Specimens shot at White Salmon
in early March were lean and gave no evidence of having hibernated.

The Beechey ground squirrel is an excellent climber; I have seen
individuals in oak trees on a number of occasions. They like to sit on
fence posts and when alarmed descend head foremost, with speed and
agility. In running on the ground their movements are more graceful than
those of _Citellus columbianus_ but are unlike the flowing, bounding run
of tree squirrels.

Near White Salmon and Lyle, these ground squirrels were eating burr
clover (_Medicago_), grasses (_Bromus_), and alfalfa. In some places
they do considerable damage to alfalfa fields. They probably eat acorns
and are said to eat some insects.

In California the young are born in May and number five to seven.

No disease has been recorded for this subspecies but a related form in
California (_Citellus b. beecheyi_) is known to harbor plague (Kellogg,
1935: 857) and tularemia (McCoy, 1911: 53-71).

=Citellus lateralis= (Say)

Golden-mantled ground squirrel

_Description._--The head and body measure about 6 inches and the tail
about 4 inches. The head is large and rounded with eyes and ears of
moderate size. The body is stouter than that of a chipmunk. The tail is
long and well furred. Upper parts are buffy gray with one pale stripe
bordered by two black stripes extending from shoulders to rump, and the
underparts are buffy. The head and shoulders are tawny. The tail is
buffy beneath and darker above.

Externally golden-mantled ground squirrels somewhat resemble chipmunks
but actually are no more closely related to chipmunks than are other
ground squirrels. They are larger than chipmunks and their stripes
differ in that there is but one pale stripe on each side bordered with
black stripes rather than two pale stripes, and in that the stripes end
at the shoulders rather than continuing on to the face.

These ground squirrels are distributed in mountainous areas of western
North America from central British Columbia south to central Arizona and
New Mexico. A closely related species (_Citellus madrensis_) occupies a
limited area in northern Mexico and another lives in the Cascades of
Washington. There are two subspecies of _lateralis_ in Washington, each
with a major part of its range outside of Washington.

This species inhabits mountain clearings, parks and talus slides, where
it is most common about rocks, stumps, and logs. Its fondness for talus
slides and outcrops has earned it the common name of "rock squirrel" in
some places.

Entrances to burrows are usually beside a rock, stump, or root, or are
concealed under talus slides. A burrow excavated by Hatt (1927) in
Colorado was three inches in diameter at the entrance and uniformly two
inches in diameter for the rest of its length. Most of the burrow was 8
inches beneath the surface. The tunnel branched twice and contained one
pocket, probably a turning or passing chamber, in addition to the nest.
"A runway surrounded the nest on three sides, from which there were four
passages leading in. The nest cavity was 4 inches deep, the nest not
filling the space available, but occurring more as a mat in the bottom
of a cup."

The habits of this ground squirrel in Washington are little known. Since
it occupies areas that are cold and under deep snow in winter, it
probably hibernates. According to Howell (1938: 32) these animals become
fat in the fall and must retire for the winter about the middle of

The food habits of these ground squirrels in Washington are little
known. In the Pend Oreille Mountains, Pend Oreille County, I found them
eating the fruit of the western raspberry (_Rubus leucodermis_). Howell
lists, as food of this species, the seeds of yellow pine and douglas
fir, serviceberries, other berries and seeds, grain, mushrooms, and
several kinds of insects.

=Citellus lateralis tescorum= (Hollister)

      _Callospermophilus lateralis tescorum_ Hollister, Smithsonian Misc.
      Coll., 56 (no. 26):2, December 5, 1911.

      _Citellus lateralis tescorum_ Elliot, Check-list Mamm. N. Amer.,
      Supp., p. 29, 1917.

      _Type._--Obtained at the head of Moose Pass Branch, Smoky River,
      Alberta (near Moose Pass, British Columbia), 7,000 ft., by N.
      Hollister on August 2, 1911; type in United States National Museum.

      _Racial characters._--Large size, gray color.

      _Measurements._--A female from Pass Creek Pass, Pend Oreille
      County, measures: total length 257; length of tail 90; hind foot
      40; ear 14.

      _Distribution._--Known only from the Pend Oreille Mountains, Pend
      Oreille County, at Pass Creek Pass (W. W. D.).

  [Illustration: FIG. 86. Distribution of golden-mantled ground
  squirrels in Washington. A. _Citellus lateralis tescorum._
  B. _Citellus lateralis connectens._ C. _Citellus saturatus._]

=Citellus lateralis connectens= (Howell)

      _Callospermophilus chrysodeirus connectens_ Howell, Jour. Mamm.,
      12:161, May 14, 1931.

      _Citellus lateralis connectens_ Howell, N. Amer. Fauna, 56:205,
      May 18, 1938.

      _Type._--Obtained at Homestead, Oregon, by H. H. Sheldon on June 1,
      1916; type in United States National Museum.

      _Racial Characters._--Similar to _C. l. tescorum_ but browner,
      more buffy and less grayish, mantle brighter, size slightly

      _Measurements._--Seven males from northeastern Oregon average
      (Howell, 1938: 206): total length 266; length of tail 92; hind
      foot 41.7; ear 14.8 (dry). An adult female from Godman Springs,
      Columbia County, measures: total length 257; length of tail 87;
      hind foot 37; ear 14.

      _Distribution._--The Blue Mountains of extreme southeastern

      _Remarks._--This race differs markedly from the subspecies that
      occupies northeastern Washington, but differs but little from
      _Citellus l. chrysodeirus_ of the Cascades of Oregon.

=Citellus saturatus= (Rhoads)

Golden-mantled ground squirrel

      _Tamias lateralis saturatus_ Rhoads, Proc. Acad. Nat. Sci.
      Philadelphia, 1895:43, April 9, 1895.

      [_Spermophilus lateralis_] _saturatus_ Elliot, Field Columb. Mus.
      Publ. 45, zoöl. ser., 2:83, 1901.

      _Citellus lateralis saturatus_ Elliot, Field Columb. Mus. Publ.
      105, zoöl. ser., 6:106, 1905.

      _Callospermophilus lateralis saturatus_ Miller, U. S. Nat. Mus.
      Bull., 79:316, December 31, 1912.

      _Citellus saturatus_ Howell, N. Amer. Fauna, 56:212, May 18, 1936.

      _Type._--Obtained at Lake Keechelus, 3000 ft., Kittitas County,
      Washington, by A. Rupert, in September, 1893; type in Academy of
      Natural Sciences of Philadelphia.

      _Measurements._--Ten males from the Cascade Mountains average:
      total length 305; length of tail 110.9; hind foot 46.5; ear 17
      (dry). The weights of 3 males and 5 females average, respectively:
      281 grams; 259.4 grams.

      _Distribution._--The higher and eastern Cascade Mountains from
      Barron (Howell, 1938: 213) and Bauerman Ridge (Howell, 1938: 213)
      south to Cleveland (Howell, 1938: 213) and Goldendale (W. W. D.).

_Remarks._--The golden-mantled ground squirrel of the Cascades is
similar to _Citellus lateralis_ but is larger and duller-colored. Head
and body are about 8 inches in length and the tail is about 4-1/2 inches
long. The upper parts are brownish gray with one pale and two dark
stripes on each side. Head and shoulders are tawny. The underparts are
dull buffy gray.

_Citellus saturatus_ occupies the Cascade Mountains of Washington and
southern British Columbia. It inhabits talus slides and clearings. The
rock embankment of the Great Northern Railroad between the Cascade
Tunnel and Leavenworth, Chelan County, is a favored habitat and the
population along the railroad probably averages one squirrel each
hundred yards. To the east it extends well into the Transition
Life-zone, being abundant in clearings in the open forest of yellow
pine. In places its range nearly reaches the Upper Sonoran Life-zone. A
few miles east of Leavenworth, Chelan County, we found this squirrel in
chapparal and brush, principally composed of _Ceanothus creneatus_. Here
the animals were unusually abundant and formed a colony almost as dense
as colonies of _Citellus washingtoni_.

Entrances of burrows are usually placed beside rocks, stumps or logs.
In the area near Leavenworth, mentioned above, entrances to burrows
were in the open or among roots of bushes. The entrances to many
burrows are doubtless concealed under talus slides. No complete records
of excavations of burrows are available. One burrow, uncovered by road
construction operations three miles east of Scenic, Chelan County, was
dug in the earth-filled cleft of a great granite boulder. The cleft
was 5 feet wide at the surface of the ground but narrowed until at a
depth of six feet the stones were in contact. The cleft was at least
10 yards long. The burrow descended at an angle of 45 degrees, to a
depth of three feet. Here the construction work had passed the cleft,
but digging into the almost vertical wall uncovered the nest at the end
of a horizontal tunnel two feet farther on. It was a matted cup of dry
grass with two fresh, green fern fronds lying loose in the cup. Two
divergent burrows emerged on opposite sides of the nest cavity, but a
fall of the loose, gravelly soil prevented further observation. The
body of the occupant, crushed and mangled by the steam-shovel scoop,
was that of a small male.

  [Illustration: FIG. 87. Golden-mantled ground squirrel (_Citellus
  saturatus_), captured when young at Tye, Washington, by Earl J.
  Larrison, June 20, 1940; photographed February 1, 1941. (Fish and
  Wildlife Service photo by Victor B. Scheffer, No. 1139.)]

In spite of their vivacious appearance, these squirrels are rather
sedentary. When undisturbed they move leisurely over rocks and stumps,
pausing often, and occasionally sitting on their hindquarters to gaze
about for minutes at a time. They are good climbers and often ascend
smooth-barked trees to heights of 20 feet or more. At the approach
of danger they descend and enter the nearest burrow. In August these
ground squirrels become exceedingly fat and by late September only a
few may be seen, basking in the midday sun. These take alarm at the
slightest excuse. All have usually gone into hibernation by October 15.

The food includes the berries of salal, huckleberry, mountain ash, and
seeds of lupine. Near Liberty, Kittitas County, a squirrel killed by a
car had its cheek pouches stuffed with garden peas. The source of its
loot was doubtless the garden of a farmhouse 100 feet away. Numerous
individuals are run over by cars, and on many occasions I have found
other individuals eating the sun-dried flesh of their mates, parents,
or young. These cannibals are often run over while so engaged, and it
is not unusual to find two or three dead on a section of pavement 20
feet in length.

Economically this species is of little importance. Its principal enemies
probably include hawks, weasels, martens, bobcats and coyotes.

=Sciurus griseus griseus= Ord

Western gray squirrel

      _Sciurus griseus_ Ord, Jour. de phys., 87:152, 1818.

      _Sciurus griseus griseus_ Miller, U. S. Nat. Mus. Bull., 128:222,
      April 29, 1924.

      _Type._--None. Described from a squirrel seen by Lewis and Clark.
      Type locality, The Dalles, Wasco County, Oregon.

      _Measurements._--A female from midway between Satus Pass and
      Goldendale, Klickitat County, measured: total length 560; length
      of tail 264; hind foot 76; weight 897 grams.

      _Distribution._--Southwestern Washington north to the glacial
      prairies near Tacoma (W. W. D.); the Columbia River Valley of the
      southern Cascades; the eastern edge of the Cascades north to Lake
      Chelan (Taylor and Shaw, 1929: 18).

  [Illustration: FIG. 88. Distribution of the western gray squirrel,
  _Sciurus griseus_, in Washington.]

_Description._--This is the largest tree squirrel found in Washington.
The head and body measure about 12 inches, and the tail about 11 inches.
The body is long and slender. The fur is long and soft, that on the tail
being exceptionally long. The upper parts are silvery gray and the
underparts white.

The western gray squirrel is restricted to the region of the Pacific
Coast and ranges from central Washington south to northern Lower
California. Three subspecies are recognized, only one of which occurs in
Washington. Like the red squirrels, the western gray squirrel is
arboreal. Its favored habitat is the oak woods rather than coniferous
forest. Its range in Washington is largely regulated by the distribution
of oaks, especially the garry oak (_Quercus garryana_). Altitudinally it
ranges from near sea level at Puget Sound to above 2,500 feet elevation
in the eastern Cascade Mountains. It seems to be restricted to the
Transition Life-zone.

The habits of the western gray squirrel are modified by its arboreal
existence. Homes consist of holes in hollow trees or outside nests of
sticks and twigs. It is extremely active in trees and travels from tree
to tree on branches that seem scarcely strong enough to bear the weight
of so large an animal. The oak woods in Washington are usually rather
open and the trees more scattered than is the case in coniferous
forests. Consequently gray squirrels must more often descend to the
ground than Douglas and red squirrels. On the ground, western gray
squirrels travel in smooth leaps each of two feet or more in length. The
long tail is held out behind and "follows through" the animal's leaps in
a remarkable way. The tip of the tail may be descending from the
previous leap while the animal's fore parts are already at the height of
a new jump. This results in a "flowing" movement that is extremely
graceful. The squirrel may stop momentarily to dig in the ground or
search for an acorn. At such times the tail is immediately switched up,
over the back.

The western gray squirrel is able to drop unharmed for considerable
distances. Near Fort Lewis, Pierce County, a gray squirrel was cornered
in a tall fir tree and an agile friend volunteered to climb the tree.
The squirrel ascended to the very topmost branches. When only a few feet
separated them the squirrel leaped far out into the air. Its legs were
stretched out stiffly, the tail was extended and the body slightly
arched. It struck the ground with an audible thud and bounced fully 18
inches. At the height of its bounce, the squirrel's legs began moving
rapidly, and it struck the ground the second time at a full run.

The principal food is acorns, although the seeds of the Douglas fir and
probably other conifers are eaten. Four embryos were found by Victor B.
Scheffer in a specimen from Klickitat County on March 20, 1939.

=Sciurus carolinensis hypophaeus= Merriam

Eastern gray squirrel

      _Sciurus carolinensis hypophaeus_ Merriam, Science, 7:351, April
      16, 1886.

      _Type._--Obtained at Elk River, Sherburne County, Minnesota.

_Description._--Size large, slightly smaller than the western gray
squirrel; color of upper parts less silvery, more reddish, especially on
the dorsal area and top of the tail.

_Remarks._--The eastern gray squirrel was introduced at Woodland Park,
Seattle, in 1925. The original stock (7 pairs) came from Minneapolis,
Minnesota. It has spread through the nearby woods, around Green Lake,
Cowan Park, and to the woods on the University of Washington Campus.
Occasional individuals are found outside the city limits, but the
species seems not to spread away from the city.

=Sciurus niger= Linnaeus subsp?

Fox squirrel

      [_Sciurus_] _niger_ Linnaeus, Syst. Nat., 1 (10th ed.):64, 1758.

_Description._--Similar in size and appearance to the eastern gray
squirrel but upper parts more reddish and underparts reddish orange
rather than white.

_Remarks._--Occasional fox squirrels are encountered near Seattle where
they have been introduced from the southeastern United States. The
source and date of the introduction are unknown.

=Tamiasciurus hudsonicus= (Erxleben)

Red squirrel

_Description._--The head and body of the red squirrel measure about
7-3/4 inches, the tail about 5 inches. It may be recognized by its trim
body, bushy tail and white underparts. The upper parts are reddish gray,
reddest on the dorsal area. The red color of the center of the back
extends to the tail. A black line separates the dark upper parts from
the white underparts.

Red squirrels range over North America from the northern limit of tree
growth south through the United States to Tennessee and North Carolina.
A related species, _douglasii_, is found along the Pacific coast from
British Columbia to California. The genus _Tamiasciurus_ differs from
_Sciurus_ of Washington in lacking a penis bone or baculum. There are
other fundamental differences in anatomy (see Mossman, Lawlah and
Bradley, 1932: 89-155).

The habitat of the red squirrels is the coniferous forests from which
they rarely stray. Zonally they range through the Transition and
Canadian life-zones into the Hudsonian Life-zone. Red squirrels are
arboreal and most of their habits are modified by arboreal existence.
They are swift and agile climbers, able to travel from tree to tree on
slender twigs or by leaping as much as ten feet to span the distance
from one branch to another. They ascend and descend trees head first.
They hang by their hind feet, high in the air, to clip the cones of
conifers. If cornered in a tree they leap far out and, by extending the
legs and tail stiffly, fall to the ground unharmed by leaps of fifty
feet or more.

  [Illustration: FIG. 89. Douglas squirrel (_Tamiasciurus douglasii
  douglasii_): feeding station with remnants of Douglas fir cone,
  Longmire, Washington, elevation 2,700 feet, June 25, 1937. (Fish
  and Wildlife Service photo by Victor B. Scheffer, No. 268.)]

The homes of red squirrels are woodpecker holes or other holes in hollow
trees. More rarely they build outside nests of twigs and branches, about
two feet in diameter, or add onto old nests of crows or jays. Most nests
and holes are some distance from the ground but some holes are between
roots on the ground itself.

The food consists principally of the seeds of coniferous trees,
especially Douglas fir and various species of pines. Cones are clipped
when green or just before ripening and are either allowed to fall to the
ground to be retrieved later or are at once carried to a favored feeding
place to be consumed. Cones are held between the forepaws while the
squirrel sits on its hind feet, tail curved up over its back, and
rapidly clips the cone apart to get the seeds it contains. The cone is
rotated between the paws and a steady stream of husks drops to the
ground. Soon only the core is left and this too joins the husks on the
ground. Favored feeding stations are used continually, perhaps by
generation after generation of squirrels, and debris from thousands of
cones accumulates in great piles.

Cones are stored. In the vicinity of a favored feeding place, at times
virtually every hiding place is filled with green cones. Cones may be
jammed into cracks or crevices in logs or stumps without effort at
concealment or may be carefully covered with leaves or dry needles. Many
are placed in craterlike pits dug in the ground. Most of these pits
probably are later covered over but many are left open to the weather.
Hollows in trees are probably also used for storage, as are holes dug
into the piles of accumulated cone debris beneath feeding stations.
Other food eaten by squirrels includes hazelnuts, berries, maple seeds,
and mushrooms. A variety of fruits and seeds are doubtless eaten when
opportunity offers.

Red squirrels do not hibernate. In the lowlands they are active all
winter long but are noticeably shy and quiet. In the mountains they
disappear after the snow falls but tracks may be seen in the snow about
their dens and occasional individuals are seen. Specimens collected at
this time are not fat, as would be the case if hibernation had been
interrupted. Seemingly they stay close to their homes and feed on stored

The call of the red squirrel is harsher, more metallic, than that of the
Douglas squirrel. The common call is a rolling "bur-r-r," starting loud
but fading out entirely in a half minute. A lower-pitched "pauf" is
uttered at intervals when the squirrel is going about its business. The
danger cry is a loud "pee-ee," not unlike the call of a red-tailed hawk
but less shrill. A low pitched "chauf-chauf-chauf," repeated at
intervals of about two seconds, is occasionally given in the fall of the

=Tamiasciurus hudsonicus streatori= (Allen)

      _Sciurus hudsonicus streatori_ Allen, Bull. Amer. Mus. Nat. Hist.,
      10:267, July 22, 1898.

      _T[amiasciurus]. h[udsonicus]. streatori_ Howell, Proc. Biol. Soc.
      Washington, 49:135, August 22, 1936.

      _Type._--Obtained at Ducks, British Columbia; type in American
      Museum of Natural History.

      _Racial characters._--Similar to _richardsoni_ but smaller and
      darker, redder above.

      _Measurements._--Seven males from north-central Washington average:
      total length 330; length of tail 129; hind foot 51; ear 23.

      _Distribution._--Northeastern Washington. Marginal occurrences are
      (from Taylor and Shaw, 1929: 17): Beaver Creek, Ruby Creek, and
      head of Lake Chelan.

      _Remarks._--This race reaches Washington from the interior of
      British Columbia. Intergradation between _hudsonicus streatori_
      and _h. richardsoni_ takes place over much of northeastern

  [Illustration: FIG. 90. Distribution of the red squirrel and Douglas
  squirrel in Washington. A. _Tamiasciurus hudsonicus richardsoni._ B.
  _Tamiasciurus hudsonicus streatori._ C. _Tamiasciurus douglasii

=Tamiasciurus hudsonicus richardsoni= (Bachman)

      _Sciurus richardsoni_ Bachman, Proc. Zoöl. Soc. London, p. 100,

      _Sciurus hudsonius Richardsoni_ True, Proc. U. S. Nat. Mus.,
      7:595, 1884.

      _Sciurus hudsonicus richardsoni_ Allen, Bull. Amer. Mus. Nat.
      Hist., 10:265, July 22, 1898.

      _Sciurus hudsonicus richardsoni_ Miller, U. S. Nat. Mus. Bull.,
      128:211, April 29, 1924.

      _Tamiasciurus hudsonicus richardsoni_ Svihla and Svihla, Murrelet,
      21:55, December 20, 1940.

      _Type._--Obtained at the head of the Big Lost River, Custer
      County, Idaho.

      _Racial characters._--Color paler, especially on back.

      _Measurements._--Four males and 4 females average, respectively:
      total length 339, 339; length of tail 133, 131; hind foot 54, 51;
      ear 24, 24; weight 256, 266.

      _Distribution._--The Blue Mountains of southeastern Washington.

=Tamiasciurus douglasii douglasii= (Bachman)

Douglas squirrel

      _Sciurus douglasii_ Bachman, Proc. Zoöl. Soc. London, p. 99,

      _Sciurus hudsonius Douglasii_ True, Proc. U. S. Nat. Mus.,
      7:595, 1884.

      _Sciurus douglasii douglasii_ Miller, U. S. Nat. Mus. Bull.,
      128:212, April 29, 1924.

      _Sciurus douglasii cascadensis_ Taylor and Shaw, Occ. Papers Chas.
      R. Conner Mus., no. 2:18, December, 1929.

      _Tamiasciurus douglasii_ Holdenried, Jour. Mamm., 21:406, November
      14, 1940.

      _Type._--Obtained near the mouth of the Columbia River by David

      _Measurements._--Fifteen males and 10 females from the lowlands of
      western Washington average, respectively: total length 317, 320;
      length of tail 121, 126.7; hind foot 50, 49.4; ear 23.2, 22.6;
      weight 204, 202 grams.

      _Distribution._--From the eastern base of the Cascade Mountains
      westward to the Pacific. Marginal occurrences are: Nooksack River
      (Taylor and Shaw, 1929: 18), Lake Wenatchee (W. W. D.) and
      Wenatchee (Taylor and Shaw, 1929: 18).

_Remarks._--Squirrels from the Puget Sound area are slightly less
intensely colored than specimens from the ocean coast and the Olympic
Mountains. Those from the higher and eastern Cascade Mountains are
paler still. When, however, squirrels from the Cascades of Washington
are compared with individuals of the race inhabiting the Cascades of
Oregon, it is apparent that the Washington squirrels are nearest to
_douglasii_. The pale tips of the hairs in the tail of the Oregon race
are strikingly white while in Washington specimens they are orange or

The Douglas squirrel resembles the red squirrel in size and proportions
but differs in being dusky olive (less reddish) on the upper parts and
orange instead of white on the underparts.

Douglas squirrels range from southern British Columbia southward
to Lower California. They are confined to the Pacific Coastal
region. Although closely related to _Tamiasciurus hudsonicus_, no
intergradation with that species has been found.

The rich coloration of the Douglas squirrel matches the background of
the humid forests it inhabits. It occurs at sea level along the ocean
coast, the shores of Puget Sound and on some islands in Puget Sound.
It ranges through the Transition and Canadian life-zones well into the

It makes its home in holes in trees or in nests constructed of conifer
twigs, needles and bark. Old nests of birds may be modified and taken
over, but most outside nests seem to have been made entirely by the
squirrel. Nests are usually placed near the top of denser fir and
cedar trees. Nests are hemispherical in shape, 12 inches or more in
diameter, and open on top. The central cup, 4 inches in diameter, is
lined with strips of inner bark of red cedar coiled but unshredded.
Nests in holes are usually some distance from the ground, but the
entrances to some are at ground level.

The habits of the Douglas squirrels vary with the season. In spring
they are shy. They rarely call and are skillful in keeping branches
or tree trunks between themselves and persons. In June and July
they become bolder, calling more frequently and moving about more.
In September and October they become exceptionally bold. They call
almost incessantly from late morning until early afternoon, and spend
considerable time on the ground.

The call notes of the Douglas squirrel are similar to those of the
red squirrel but are softer, more slurred. The common call is the
long, trilling "burr," loud at first and becoming gradually softer.
On hot afternoons in late summer a barking "pauf" note is given. This
sound is repeated several times at intervals of a minute or so. It
has remarkable carrying power. The danger signal is a loud, explosive
"pe-ee." Another common call is a short barking or chirping "bauf" with
a musical, questioning sound. This is repeated at intervals of a few
seconds when the presence of danger is suspected. At times it, rather
than the loud danger signal, is given also when a person is sighted. A
low chirping note is often given while the squirrel is busily at work.
This is made while the squirrel is moving about. Other calls described
are made from a standing or sitting position and are accompanied by a
jerk or a flick of the tail.

The Douglas squirrel eats seeds of the Douglas fir, seeds of maple,
hazelnuts, dogwood berries and mushrooms. The berries of forest shrubs,
such as the red huckleberry, salal and Oregon grape, may also be eaten.

Two broods of young may be born in a year. The first is born in early
June. Embryos found from June 11 to 28 varied in number from 6 to 8. A
nursing female was taken as early as June 10, 1938, and one was taken as
late as October 10, 1938.

Douglas squirrels commonly have a few fleas and 2 to 5 ticks at the
bases of the ears. No lice, cases of mange, or serious infestations of
parasites have been seen.

=Glaucomys sabrinus= (Shaw)

Northern flying squirrel

  [Illustration: FIG. 91. Northern flying squirrel (_Glaucomys
  sabrinus_), probably from near Seattle, in Woodland Park Zoo, Seattle,
  Washington, January 28, 1941. (Fish and Wildlife Service photo by
  Victor B. Scheffer, No. 1137.)]

_Description._--The northern flying squirrel is slightly smaller than
the red squirrel (_Tamiasciurus_). The head and body measure about 7
inches and the tail about 5 inches. The most distinctive feature is
the loose fold of skin that stretches from the wrist of the foreleg to
the ankle of the hind leg. The fur is extremely soft and plushlike. It
is not separated into guard hair and underfur, and because all hairs
are of approximately the same length, the fur appears extremely sleek.
The fur of the tail is of the same texture as that on the body but is
dorso-ventrally compressed. The eyes are large and dark. The color of
the upper parts varies from reddish brown in some races to brownish
gray in others. The underparts are buffy gray.

Flying squirrels range over the forested parts of North America from
Guatemala to Alaska and northern Canada and from the Atlantic to
the Pacific. A closely related genus (_Pteromys_) occurs in Europe.
Two species are found in North America. The small _volans_ ranges
in eastern United States, Mexico, and Guatemala, while the larger
_sabrinus_ occurs in parts of the western United States, most of
Canada, and central Alaska. Near the Great Lakes the two species occur
in the same area, probably in different habitats.

Howell (1918: 16) recognized 18 subspecies of _sabrinus_, five of which
were recorded in the state of Washington. One of these (_olympicus_)
proves to be a synonym of an earlier-named subspecies but another form
(_bangsi_), not recorded for Washington by Howell, has been found in
the Blue Mountains in the southeastern corner of the state (Taylor and
Shaw, 1929: 18).

Flying squirrels are strictly arboreal and occur only in woods and
forests. They prefer areas where trees grow close together and to
considerable heights. Older woods with dead stubs and hollow trees
are preferred. They occasionally invade attics of cabins and other
habitations of man. The old Forestry Building on the University of
Washington Campus, later the home of the Washington State Museum,
was inhabited by flying squirrels for many years. Several specimens
preserved in the Museum prove the poor judgment of the animal in
choosing a natural history museum as a home.

Relatively little is known of the detailed habits of _Glaucomys
sabrinus_ although the life history of the eastern _Glaucomys volans_
has been described. Flying squirrels are active all winter, even in
the high mountains. They are often caught in traps set for fur bearers
and, where trapping is an important means of livelihood, they are
serious pests. Flying squirrels do not actually fly but only glide
through the air. The loose fold of fur between their limbs is stretched
by extending the legs. With it the flying squirrel is able to sail 50
yards or more. The flat tail serves as a rudder and allows the squirrel
to change direction while in flight. Glides end with an upward swoop,
allowing the squirrel to alight, head upwards, on a tree trunk slightly
lower than their starting point.

Unlike other members of the squirrel family, flying squirrels are
completely nocturnal. They are seen in the daytime only when frightened
from their retreats. Homes consist principally of old woodpecker holes
or other cavities in trees. They are said to build outside nests,
similar to those of tree squirrels, but I have found none of these in
Washington. The animals can be frightened from their holes by pounding
on the bases of trees in which their nests are situated. Certain holes
seem to be preferred nesting places. Near Cottage Lake, King County,
two flying squirrels were taken from a woodpecker hole in succeeding
years. Cowan (1936B: 58) discovered remains of 14 nests of flying
squirrels in a single hollow tree near Alta Lake, British Columbia.
According to Cowan, hollow trees are used in winter but the young are
born in outside nests of shredded bark and lichens. The young usually
number 3 and are born in May and June.

  [Illustration: FIG. 92. Distribution of flying squirrel in Washington.
  A. _Glaucomys sabrinus oregonensis._ B. _Glaucomys sabrinus
  fuliginosus._ C. _Glaucomys sabrinus columbiensis._ D. _Glaucomys
  sabrinus latipes._ E. _Glaucomys sabrinus bangsi._]

Only once have I heard the voice of a flying squirrel. Near Dewey Lake,
Yakima County, the squirrels were unusually abundant in the black
hemlock and Douglas fir forests of the mountain sides. Here a birdlike
twittering note caught my attention and occasional dark, sailing shapes
were glimpsed against the clear sky. Eight traps set in the afternoon
and visited at 11 p. m. held two flying squirrels. In the morning
another was in a trap. Seemingly the squirrels are active most of the

The food consists principally of conifer seeds and probably other nuts,
seeds, and fruit. They eat the meat bait of traps set for fur bearers
and probably eat the eggs of birds.

=Glaucomys sabrinus oregonensis= (Bachman)

      _Pteromys oregonensis_ Bachman, Jour. Acad. Nat. Sci. Philadelphia,
      8:101, 1839.

      _Sciuropterus alpinus oregonensis_ Rhoads, Proc. Acad. Nat. Sci.
      Philadelphia, p. 324, June, 1897.

      _Sciuropterus alpinus olympicus_ Elliot, Field Columb. Mus. Publ.
      30, zoöl. ser., 1:225, February 1, 1899 (type from Happy Lake,
      Clallam County, Washington).

      _Glaucomys sabrinus oregonensis_ Howell, N. Amer. Fauna, 44:44,
      June 13, 1918.

      _Glaucomys sabrinus olympicus_ Howell, N. Amer. Fauna, 44:49, June
      13, 1918.

      _Type._--Obtained "in pine [= conifer] woods of the Columbia near
      the sea" by J. K. Townsend in 1839. Probably near St. Helen,
      Columbia County, Oregon (Rhoads, 1897:324).

      _Racial characters._--Small size, rich color.

      _Measurements._--A male from Cottage Lake, King County, measured:
      total length 287; length of tail 125; hind foot 38. A female from
      5 miles southeast of Sequim, Clallam County, measured: 303; 133;
      41; ear 27. A male from Quilcene, Jefferson County, measured: 311;
      140; 41.

      _Distribution._--Western Washington west of the Cascade Mountains.
      The locality records for flying squirrels in the northern Cascade
      Mountains, given by Taylor and Shaw (1929: 18), when plotted on a
      distribution map, show overlapping of ranges in this area. The
      ranges of the three races involved have been drawn on the
      accompanying map (fig. 92) on the basis of geographic probability
      and are subject to revision because the specimens from this area
      have not been examined.

      Marginal records that may apply to _G. s. oregonensis_ are, from
      Taylor and Shaw (1929: 18): Nooksack River, Rockport, North Bend
      and Skamania.

      _Remarks._--Of all the races occurring in Washington,
      _oregonensis_ is the most distinct. _Glaucomys s. olympicus_
      Elliot must be regarded as a synonym of _oregonensis_.

=Glaucomys sabrinus bangsi= (Rhoads)

      _Sciuropterus alpinus bangsi_ Rhoads, Proc. Acad. Nat. Sci.
      Philadelphia, p. 321, June, 1897.

      _Glaucomys sabrinus bangsi_ Howell, N. Amer. Fauna, 44:38, June
      13, 1918.

      _Type._--Obtained in Idaho County, Idaho, by Harbison and Bargamin
      on March 8, 1897; type in Philadelphia Academy of Natural Sciences.

      _Racial characters._--Similar to _oregonensis_ but larger and
      paler throughout.

      _Measurements._--A female from Wildcat Spring, Columbia County,
      measured: total length 333; length of tail 147; hind foot 45; ear
      29; weight 151 grams.

      _Distribution._--Found only in the Blue Mountains of extreme
      southeastern Washington.

=Glaucomys sabrinus columbiensis= Howell

      _Glaucomys sabrinus columbiensis_ Howell, Proc. Biol. Soc.
      Washington, 28:111, May 27, 1915.

      _Type._--Obtained at Okanogan, British Columbia, by Allan Brooks on
      May 9, 1898; type in United States National Museum.

      _Racial characters._--Larger than _oregonensis_ and paler. Similar
      to _bangsi_ but paler, especially beneath, and less reddish above.

      _Measurements._--Howell (1918: 46) gives the average of two
      subadult topotypes as: total length 313; length of tail 143; hind
      foot 42.

      _Distribution._--Northeastern Washington, probably extending, from
      records of Taylor and Shaw (1929: 18), west to Mazama and Stehekin
      and east to Molson.

=Glaucomys sabrinus latipes= Howell

      _Glaucomys sabrinus latipes_ Howell, Proc. Biol. Soc. Washington,
      28:112, May 27, 1915.

      _Type._--Obtained at Glacier, British Columbia, by J. A. Loring, on
      August 13, 1894; type in United States National Museum.

      _Racial characters._--Similar to _bangsi_ and _columbiensis_ but
      with larger feet and grayer color.

      _Measurements._--Howell (1918: 49) gives the average of 10
      specimens as: total length 342; length of tail 153; hind foot

      _Distribution._--The Pend Oreille Mountains of extreme
      northeastern Washington, north (from records of Taylor and Shaw,
      1929: 18-19, as revised) to Sullivan Lake and south to Loon Lake.

=Glaucomys sabrinus fuliginosus= (Rhoads)

      _Sciuropterus alpinus fuliginosus_ Rhoads, Proc. Acad. Nat. Sci.
      Philadelphia, p. 321, June, 1897.

      _Glaucomys sabrinus fuliginosus_ Howell, N. Amer. Fauna, 44:47,
      June 13, 1918.

      _Type._--Obtained at Martin Station, Kittitas County, Washington,
      by Allan Rupert in March, 1893; type in Philadelphia Academy of
      Natural Sciences.

      _Racial characters._--Similar to _columbiensis_ but underparts
      darker and tail paler. Larger and paler than _oregonensis_.

      _Measurements._--Three females from the Cascades average: total
      length 327; length of tail 145; hind foot 40.7; ear 25.

      _Distribution._--The Cascade Mountains, according to Taylor and
      Shaw (1929: 18), as revised here, east to Entiat River and south
      to Carson.

=Perognathus parvus= (Peale)

Great Basin pocket mouse

_Description._--Great Basin pocket mice are slightly larger than house
mice. The ears are tiny and round; the tail is relatively long, slightly
longer than the head and body; the forefeet are small but the hind feet
are large and powerful, more than one-fourth as long as the head and
body. They have relatively large, fur-lined, external cheek pouches. The
color of the upper parts varies with the subspecies from blackish
olive-gray to buffy olive-gray; a line of clear buff or buffy-olive
extends along the lower part of side, separating the darker color of the
upper parts from the white underparts; the tail is short-haired,
blackish above, olive beneath.

Pocket mice of the genus _Perognathus_ range from Mexico northward to
British Columbia. Only one species, _parvus_, occurs in Washington where
it is represented by three subspecies. Pocket mice are most common in
the Upper Sonoran Life-zone in sandy areas dotted with desert shrubs.
They are found occasionally in dry, grassy places in the Arid Transition
Life-zone. They may be locally abundant in rocky areas and are often
trapped high on talus slides, many yards from the nearest soil. Gray
(1943: 191-193) estimates their numbers on the sagebrush areas of the
Yakima Valley at 32 per acre. They are completely nocturnal. Their
ordinary movements are rather slow and specimens studied by the aid of a
searchlight usually crept quietly into the protection of the nearest
desert shrub. The trail of a pocket mouse in soft sand may be recognized
by the distinctive mark left by the dragging tail.

Burrows of pocket mice usually are at the bases of shrubs where tough
roots furnish protection. They are kept closed during the day by means
of a plug of fresh earth or sand. Openings may often be recognized by a
fan-shaped pile of fresh sand before the hole. Burrows excavated were
usually less than four feet in length and branched from two to four
times. No nests were found in the burrows but one contained a few fresh
stalks of desert annuals. The air in the burrows seemed warm and humid.

The pouches of pocket mice from Washington often contain the fresh,
green tips of desert plants, grass seeds, seeds of plants other than
grasses, and plant leaves. Because of their occasional great abundance,
pocket mice may be a menace to agriculture. Fortunately much of their
habitat is unsuited to farming.

In Washington pocket mice breed in March and April. From 4 to 8 embryos
were found in pregnant females.

=Perognathus parvus parvus= (Peale)

      _Cricetodipus parvus_ Peale, U. S. Explor. Exped., 8 (mamm. and
      ornith.):53, 1848.

      _Perognathus parvus_ Cassin, U. S. Explor. Exped., 8 (mamm. and
      ornith.):48, 1858.

      _Perognathus parvus parvus_ Miller, Bull. U. S. Nat. Mus.,
      128:278, April 29, 1924.

  [Illustration: FIG. 93. Distribution of the Great Basin pocket mouse
  in Washington. A. _Perognathus parvus parvus._ B. _Perognathus parvus
  columbianus._ C. _Perognathus parvus lordi._]

      _Type._--Probably obtained in the neighborhood of The Dalles, Wasco
      County, Oregon.

      _Racial characters._--Size small; upper parts brownish-buff washed
      with blackish or, in gray phase, ashy gray washed with blackish;
      sides buffy yellow; a buffy spot often present on throat; tail
      blackish above, olive below; facial markings usually brownish-buff
      but in older animals indistinct and washed with blackish.

      _Measurements._--Thirty-one males and 19 females from Washington
      average, respectively: total length 169, 164; length of tail 90,
      86; hind foot 22.6, 21.8; ear 5, 5.

      _Distribution._--Southeastern Washington south of the Snake River,
      the area on the north side of the Columbia River in Klickitat
      County, and the Yakima Valley area as far north as the Vantage,
      Kittitas County (W. W. D.). Other marginal records are: Kennewick
      (W. W. D.), Atilla (W. W. D.) and Walla Walla (E. S. B.).

      _Remarks._--Dichromatism seems to be rather common in _Perognathus
      p. parvus_--so common that Osgood (1900: 35) recognized both a
      "red" and a "gray" phase. Anderson (1932: 102) found no
      dichromatism in _P. p. lordi_ in British Columbia. A single
      specimen from the Grand Coulee at Dry Falls, Grant County, does
      show dichromatism. It is even more red than the reddest topotypes
      of _parvus_ examined.

      Seemingly the recessive gene for red is still present in _lordi_,
      or has mutated anew. When present, the color is deeper and brighter
      than in the parent population, in keeping with the heavier
      pigmentation of the race _lordi_.

=Perognathus parvus lordi= (Gray)

      _Abromys lordi_ Gray, Proc. Zoöl. Soc. London, p. 202, 1868.

      _Perognathus lordi_ Merriam, N. Amer. Fauna, 1:28, October 25,

      _Perognathus lordi lordi_ Miller, U. S. Nat. Mus. Bull., 128:279,
      April 29, 1924.

      _Perognathus parvus lordi_ Davis, Recent Mamm. of Idaho, p. 266,
      Caxton Printers, Caldwell, Idaho, April 5, 1939.

      _Type._--Obtained in southern British Columbia (probably near Lake
      Osoyoos) by J. K. Lord, probably in 1860.

      _Racial characters._--Size large; color of upper parts
      buffy-olive, washed with blackish; sides buff-olive; facial
      markings absent or, if present, indistinct and pale olive; tail
      blackish above, olive below.

      _Measurements._--Twenty-nine males and 10 females average,
      respectively: total length 175, 171; length of tail 93, 89; hind
      foot 23.4, 22.9; ear 5.3, 5.1.

      _Distribution._--Okanogan Valley and the Columbian Plateau, except
      for the southwestern part. Marginal localities are: Vantage, Grant
      County (W. W. D.), 10 mi. S. Moses Lake (W. W. D.), Washtucna (M.
      V. Z.) and Pullman (M. V. Z.).

_Remarks._--There are constant differences between _Perognathus parvus
parvus_ and _Perognathus parvus lordi_. The latter is larger, darker,
and differs slightly in average cranial measurements. The Snake River,
the Columbia River, and the Wenatchee Mountains separate the geographic
ranges of the two subspecies and prevent intergradation between them.
Nevertheless their close similarity and probable common origin indicate
that both belong to one species, namely _parvus_.

=Perognathus parvus columbianus= Merriam

      _Perognathus columbianus_ Merriam, Proc. Acad. Nat. Sci.
      Philadelphia, p. 263, September 27, 1894.

      _Perognathus lordi columbianus_, Osgood, N. Amer. Fauna, 18:40,
      September 20, 1900.

      _Type._--Obtained at Pasco, Franklin County, Washington, by Clark
      P. Streator on May 9, 1891; type in United States National Museum.

      _Racial characters._--Size and color as in _lordi_ but skull
      considerably wider in mastoid region.

      _Measurements._--Twenty-one male and 9 female topotypes average,
      respectively: total length 173, 168; length of tail 91, 89; hind
      foot 23.9, 22.8; ear 5, 5.

      _Distribution._--The vicinity of the type locality and the part of
      the Columbian Plateau north of the Snake and east of the Columbia

_Remarks._--This race is separated from _parvus_ by river barriers and
the two do not intergrade. It differs significantly from _lordi_ only in
the wider mastoid region. No barrier separates the range of _lordi_ from
that of _parvus_, and the two races intergrade over a wide area (north
to Moses Lake, east to Washtucna). The race _columbianus_ must have
originated _in situ_ from _lordi_. The habitat of _columbianus_ seems
not to differ from that of _parvus_ or _lordi_.

=Dipodomys ordii columbianus= (Merriam)

Ord kangaroo rat

      _Perodipus ordi columbianus_ Merriam, Proc. Biol. Soc. Washington,
      9:115, June 21, 1894.

      _Dipodomys ordii columbianus_ Grinnell, Jour. Mamm., 2:96, May 2,

      _Type._--Obtained at Umatilla, Umatilla County, Oregon, by C. P.
      Streator on October 18, 1890; type in United States National

      _Measurements._--Fourteen males and 9 females from Walla Walla
      County average, respectively: total length 261.5, 248.4; length of
      tail 137.2, 139.1; hind foot 40.6, 40.8; ear 13.1, 13.0; weight
      52.1, 49.0 grams.

      _Distribution._--Western Walla Walla County from the Oregon
      boundary north to the Snake River and on Blalock Island in the
      Columbia River, Benton County.

  [Illustration: FIG. 94. Distribution of the Ord kangaroo rat,
  _Dipodomys ordii columbianus_, in Washington.]

_Description._--The kangaroo rat with a body about the size of that of a
chipmunk has an exceptionally large head and large, black eyes. The
forelegs and forefeet are tiny but the hind feet and legs are large and
powerful. The hind foot is almost a third as long as the head and body.
The tail is long, longer than the head and body. Kangaroo rats possess
fur-lined, external cheek pouches, like those of the pocket mice. The
upper parts are soft buff in color. The underparts and a stripe on each
flank are white. The tail is dusky above and below, with white sides; it
is tufted at the tip.

Kangaroo rats are typical of the desert regions of the southwestern
United States, where numerous species and subspecies are found. A single
subspecies of the wide-ranging species _ordii_ occurs into southeastern
Washington, where it is restricted to sandy areas in the Upper Sonoran
Life-zone. In the soft, drifted sand along the Columbia River where
sagebrush and other desert shrubs are low and widely spaced kangaroo
rats are abundant.

These rats are strictly nocturnal. When individuals are dug from their
burrows in the daytime they usually hop about in a dazed manner and
appear to be blinded by sunlight. Near Wallula, Walla Walla County,
these rats were caught at night with a butterfly net as they stood
"paralyzed" in the beam of a powerful searchlight. Such night hunting
was unsuccessful on cloudy or windy nights when kangaroo rats seem not
to move about.

As might be guessed from their powerful hind legs, kangaroo rats travel
in bounds. Near Wallula, where we watched them in their natural habitat,
they traveled, when unfrightened, in slow hops, each hop followed by a
pause. As they struck the surface of the ground an audible thud could be
heard for a distance of several feet. After each jump they paused for a
second or so, perhaps to allow a pursuing enemy to over-run them.

Near Wallula the burrows of kangaroo rats were dug in large mounds of
wind-blown sand. The burrows entered these natural mounds horizontally
and branched two or three times. Their average length was about five
feet. No nests or food stores were discovered although several kangaroo
rats were caught as they burst from entrances at sides of the mounds.
All entrances to burrows were plugged with soft sand. The air in the
burrows seemed warm and humid.

Food found in the cheek pouches of kangaroo rats from Washington
included the seeds of desert annuals, short sections of sprouts of an
unidentified plant, grass seeds, and the leaves of the hop-sage.

A female taken March 22, 1939, contained 3 embryos.

=Thomomys talpoides= (Richardson)

Northern pocket gopher

_Description._--The pocket gopher is a fossorial animal, being but
slightly less adapted to an underground existence than the moles. The
body is stout, the legs short and the head broad. The tail is short,
sparsely haired, cylindrical and blunt-ended. The fur is soft and dense.
The eyes are small and the ears tiny and naked. The incisor teeth are
external, being separated from the mouth cavity by a furry strip of
skin. Like the pocket mouse and kangaroo rat, the pocket gopher
possesses external, fur-lined cheek pouches. The openings of these begin
just below and posterior to the nostrils, sweep out and down in a
semicircle, and end at the chin posterior to the base of the lower
incisors. They extend laterally to the shoulders and easily accommodate
a fifty-cent piece.

  [Illustration: FIG. 95. Northern pocket gopher (_Thomomys talpoides
  yelmensis_), from two miles southwest of Tenino, Washington, January
  28, 1941. (Fish and Wildlife Service photo by Victor B. Scheffer, No.

The family Geomyidae is composed of eight genera, so similar in
appearance that the name "pocket gopher" is applied to all of them. The
family is confined to North and Central America. Three genera occur in
the United States but only one, _Thomomys_, occurs in Washington.
_Thomomys_ is restricted to western North America where it ranges from
central Canada south to the southern edge of the table land of Mexico.

Several hundred kinds of _Thomomys_ have been described and as
systematic work with this genus has been continued, more and more kinds,
originally thought to be species, are found to intergrade and to be only
subspecies. All of the 17 kinds of pocket gophers occurring in
Washington belong to a single species.

The pocket gopher is principally nocturnal or crepuscular but sometimes
it is active at midday, especially if the day be dark and cloudy.
Pocket-gopher activity is indicated by fresh mounds of earth on the
surface of the ground. Rarely, an observer may see movement of plants as
the gopher molests the roots of the plants, or even see the head and
shoulders of an animal that partly emerges from an open burrow. The
ordinary gopher mound consists of less than a cubic foot of earth. The
earth is forced up from a single opening and usually is pushed out in
one direction. In consequence it forms in a fan-shaped pile about the
opening, and the last load forms a circular plug above and to one side
of the burrow opening. When so much earth has been forced out of one
opening that expulsion of additional loads of earth is overly difficult,
the burrow is extended slightly to one side, or even extended into the
newly formed mound, and another fan formed. Usually not more than three
coalesced fans form a mound, but where the soil is exceedingly soft and
fluffy, hundreds of fans may form a composite mound and the one mound
may include a cubic yard of earth. Large composite mounds probably are
formed gradually over a period of weeks or even months.

The earth in a fresh gopher mound is usually "scratched," and gives the
appearance of having been sieved. Pebbles weighing more than 100 grams
are included in material ejected from burrows. The entrances to the
burrows of gophers are usually solidly plugged with earth. The plug may
be from a few inches to more than a foot in length. At times a burrow
entrance may appear to be open, but in such cases investigation will
usually reveal it to be plugged some distance back--sometimes several

In contrast to the gopher mounds described above, the mounds of moles
are not fan-shaped but volcano-shaped. The earth from a mole's burrow is
forced straight upwards, whence it falls to either side. Later loads are
pushed up from beneath, raising the entire mound, with the last material
ejected at the center and bottom. The earth of a fresh mole-mound is not
of fine texture but instead is "clotted" and, if damp, gives the mound
a fractured appearance. When mounds are older, perhaps changed by rain
and sun, their identity as of mole-origin or pocket gopher-origin is
more difficult to establish. In such cases, if no fresh mounds can be
found, the observer must rely on the spacing of the mounds. Mole-mounds
are spaced along a burrow, about as far apart as a man can step. Gopher
mounds are irregularly spaced, and the course of the burrow cannot be
traced merely by observing the arrangement of the mounds as can that of
the mole.

In addition to mounds, gopher burrows have plugged openings where a
gopher has come to the surface, probably to cut plants. Such entrances
are marked by a plug of earth several inches long. Mounds and feeding
entrances of the gopher burrow are usually not constructed in the main
burrow system itself, but at the ends of lateral burrows of varying
length. If one traces the burrow back from the mound, a junction with
the main, better-constructed burrow is found. The junction is usually
T-shaped, with the lateral burrow at right angles to the main burrow.
More rarely the junction has a Y-shape.

  [Illustration: FIG. 96. Giant mounds raised by pocket gophers on Mima
  Prairie, Thurston County, Washington, July 13, 1941. (Fish and
  Wildlife Service photo by Victor B. Scheffer, No. 1209.)]

The burrow system of the pocket gopher may be divided into three main
parts. These are the laterals, just discussed, the main burrow, and the
deep nesting burrow. The main burrow is a sinuous tube or tunnel at a
relatively uniform depth, that marks the extent of the gopher's home
territory. This tunnel may branch, or even intersect. As it is extended
in one direction, the earth excavated by the gopher may in part be
thrust into an unused portion of the burrow.

The deep, nesting burrows may be used only in the breeding season. They
are connected with the main tunnel system but descend to a greater
depth. Usually they descend into the harder, consolidated layers of soil
below the zone where plant roots penetrate. Here chambers are
constructed in which nests and food are stored. Usually a vertical shaft
is dug in the burrow ahead of the nest to lead rain water away from the

In areas where gophers live in a thin layer of soil underlain by a more
or less impenetrable layer of rock, clay or gravel, it has been
suggested that they form unique structures known as Mima Mounds. The
formation of these mounds has been discussed in detail elsewhere
(Dalquest and Scheffer, 1942: 68-84). At least in the breeding season
the gophers work mainly and make their nest where the soil is deepest.
In the vicinity of this nest, considerable mounding and cultivation of
soil ensues. This stimulates plant growth in the area. Much observation
indicates that cultivation of earth by gophers stimulates plant growth
to a greater extent than the depredations of the gopher deplete the
ensuing growth. Thus the gopher, by cultivation of the soil in the area
about its burrow, stimulates the growth of vegetation and so increases
his own food supply. Consequently there is but little incentive for the
gopher to leave the vicinity of the nest. The gopher does, however,
construct lateral tunnels into surrounding areas. Earth from these
lateral tunnels is, in part, thrown to the surface in mounds and in part
transported back to fill the abandoned burrows near the nest. The earth
from the burrows about the nest was earlier ejected on the surface.
Slowly, then, earth is transported from surrounding areas to burrows in
the vicinity of the original nest. Each succeeding generation finds in
the vicinity of the original nest, better food and deeper soil, while
areas surrounding the nest possess thinner soil and less vegetation.
Over a period of thousands of generations of gophers, large mounds,
known as Mima Mounds, are formed. Since the removal of earth from the
surrounding areas and its accumulation in the Mima Mound are chance
affairs, the contours of the mound are smooth and flowing while the
contours of the intermound areas are smoothly convex.

The pocket gophers in Washington are economically important. In truck
and flower gardens they are a pest, especially if the crop be bulb
plants. In grain fields they are a pest because their mounds cover
considerable grain and are apt to clog or dull the knife of the mower.
In fields of young alfalfa they are apt to crop back the plant more
rapidly than it can grow. Once the alfalfa plant is well established,
however, the cultivation resulting from activities of gophers, some
persons believe, stimulates the plant to such an extent that it grows
larger and healthier in spite of the gopher's feeding on it. In the
White Salmon Valley, Klickitat County, I examined numerous alfalfa
fields. The most luxuriant growth was invariably in fields where gophers
were common. In these fields, the largest plants were those in the
immediate vicinity of gopher activity. The commensal relation between
the gopher and alfalfa was understood by many farmers, who forbade us to
take gophers for specimens from their fields. Several told us that they
always trapped the gophers from the fields of young alfalfa and from hay
meadows but encouraged their presence in fields of older alfalfa.

  [Illustration: FIG. 97. Food cache of northern pocket gopher
  (_Thomomys talpoides tacomensis_), from chamber four inches below
  surface of ground, Tacoma, Washington, December 1, 1940. Contents
  575 grams (about 2 liters) of roots, principally quackgrass,
  _Agropyron repens_. (Fish and Wildlife Service photo by Victor
  B. Scheffer, No. 1119.)]

Another economic factor is the gopher's removal of certain weeds from
grazing land. A number of introduced weeds form dense rosettes that
prevent the growth of grass from several square inches of ground and
themselves lie so close to the ground as to be unavailable as food to
grazing stock. These weeds seem to be favored food of gophers which cut
not only the rosettes but the roots of the weeds as well.

In irrigated parts of eastern Washington the gopher is a serious pest.
It burrows in the banks of the main ditches, causing cave-ins and
occasional breaks with resultant water loss. The mounds of gophers fill
in the smaller channels and divert the streams. Constant attention is
necessary to keep the ditches free of gopher mounds.

In uncultivated land the gopher is a distinct asset unless the land is
near enough to cultivated land to serve as a reservoir of pests. In the
mountains and on the desert the gopher cultivates and keeps the soil
soft and stimulates the growth of water-retaining vegetation, thus
preventing rapid run-off and erosion and keeping the flow of springs and
streams constant. Boulders, logs, and other obstructions are undermined
and, as a result of gopher activity, in time sink under the surface of
the ground. Thus a larger surface is available for plant growth. Lastly
the gopher furnishes an important food source for certain fur-bearing
mammals and eases the predator pressure on game species.

The subspecies of pocket gopher occupying the Puget Sound area are
highly restricted in their habitat preference; they occur only on the
grassy prairies of the glacial outwash aprons. They do not occur in
woods, brush or even small openings on the borders of the prairies. The
alpine forms occupy the mountain meadows and are slightly less selective
in their habits. It should be noted that in western Washington the
forests are far more open at higher altitudes than at low elevations.
The races inhabiting the desert are found in open areas, often in sandy
places. They occur more rarely in areas where the soil is baked and
claylike, and then usually in the vicinity of springs or watercourses.
The race _T. t. fuscus_ has a wide range of tolerance as regards
environmental factors; it occurs near Wenatchee in essentially desert
conditions, in alpine meadows of northeastern Washington, and in many
habitats at intermediate elevations. It occurs also in brushy areas and
is often abundant in open pine forests.

Since the gopher usually has a narrow range of tolerance as regards its
environmental adaptations, this has resulted in considerably more
isolation than is the case with other mammals, and has probably
contributed to the formation of the many subspecies. Within the range of
almost every race, microgeographic races, or local populations with
distinctive characters, are found. Many subspecies of _Thomomys_ are
probably the result of chance fixation of genetic characters already
present in a more genetically variable ancestral population, and the
loss of other genetic factors. Such races might be considered
degenerative (see Dalquest and Scheffer, 1944: 24).

  [Illustration: FIG. 98. Distribution of the northern pocket gopher in
  Washington. A. _Thomomys talpoides devexus._ B. _Thomomys talpoides
  columbianus._ C. _Thomomys talpoides aequalidens._ D. _Thomomys
  talpoides wallowa._ E. _Thomomys talpoides fuscus._ F. _Thomomys
  talpoides yakimensis._ G. _Thomomys talpoides shawii._ H. _Thomomys
  talpoides immunis._ I. _Thomomys talpoides limosus._ J. _Thomomys
  talpoides douglasii._ K. _Thomomys talpoides pugetensis._ L. _Thomomys
  talpoides tacomensis._ M. _Thomomys talpoides glacialis._ N. _Thomomys
  talpoides tumuli._ O. _Thomomys talpoides yelmensis._ P. _Thomomys
  talpoides couchii._ Q. _Thomomys talpoides melanops._]

The history of the pocket gophers of Washington has been traced
previously (Dalquest and Scheffer, 1942, 1944). It may be briefly
summarized as follows: At the close of Vashon-Wisconsin times, gophers
were found in the southern Cascades, on the Simcoe Bridge, the Columbian
Plateau, and in southeastern Washington. Following the retreat of the
ice, the gophers in the Mount Rainier area spread westward on the
outwash of the Nisqually and perhaps other glaciers to the Vashon
Outwash about southern Puget Sound and thence to the Olympic Mountains.
In the southern Cascades, gophers spread westward on glacial terraces of
the Columbia River to the vicinity of Vancouver, Clark County. The
establishment and growth of the forests split up the original
populations, and continued spread of forest has exterminated many units.
All the races in the lowlands of western Washington face extermination
as the prairies are reclaimed by forest.

Pocket gophers also invaded northeastern Washington from Idaho and
spread westward to the Cascades, thence southward until the native
gophers were encountered north of Mount Rainier and in the Yakima
Valley. The invading gophers nearly surrounded the Columbian Plateau.

T. H. Scheffer (1938B: 220-224) found the gestation period of the pocket
gopher to be approximately 28 days. No second brood is raised in
Washington. Near Kennewick, Yakima County, the young are born from
February to April. The average number of embryos found in 76 female
gophers was 6.3. Near Olympia, Thurston County, the young are born from
March to June. The average number of embryos from 312 females was 5.0.

=Thomomys talpoides devexus= Hall and Dalquest

      _Thomomys talpoides devexus_ Hall and Dalquest, Murrelet, 20:3,
      April 30, 1939.

      _Thomomys talpoides ericaeus_ Goldman, Jour. Mamm., 20:243, May
      15, 1939 (type from Badger Mountains, Douglas County, Washington).

      _Type._--Obtained 1 mile west-southwest of Neppel (now Moses Lake),
      Grant County, Washington, by W. W. Dalquest on May 30, 1938; type
      in Museum of Vertebrate Zoölogy.

      _Racial characters._--Size medium; ears tiny; color of upper parts
      pale brownish gray; underparts white; postauricular spots dark.

      _Measurements._--Two male and 4 female topotypes average,
      respectively: total length 184, 184; length of tail 54, 55; hind
      foot 25, 25; weight 89, 71 grams.

      _Distribution._--The Columbian Plateau. Marginal occurrences of
      the races of pocket gophers occurring in Washington are not listed
      here because they have recently been placed on record (Dalquest
      and Scheffer, 1944: 308-333, 423-450).

_Remarks._--This is the smallest and palest race of pocket gopher found
in Washington.

=Thomomys talpoides columbianus= Bailey

      _Thomomys fuscus columbianus_ Bailey, Proc. Biol. Soc. Washington,
      27:117, July 10, 1914.

      _Thomomys columbianus_ Bailey, N. Amer. Fauna, 39:106, November
      15, 1915.

      _Thomomys talpoides columbianus_ Goldman, Jour. Mamm., 20:234, May
      15, 1939.

      _Type._--Obtained at Touchet, Walla Walla County, Washington, by C.
      P. Streator on September 10, 1890; type in United States National

      _Racial characters._--Larger and brighter in color than _devexus_.
      Near ochraceous orange in color.

      _Measurements._--Five male and three female topotypes average,
      respectively: total length 213, 209; length of tail 60, 58; hind
      foot 29, 28.

      _Distribution._--Walla Walla County, between the Snake River and
      the Oregon boundary and from the Columbia River east to the
      Columbia County line.

=Thomomys talpoides aequalidens= Dalquest

      _Thomomys talpoides aequalidens_ Dalquest, Murrelet, 23:3, May 14,

      _Type._--Obtained at Abel Place, 2,200 ft., 6 miles south-southeast
      of Dayton, Columbia County, Washington, by S. H. Lyman, on April 6,
      1934; type in Museum of Vertebrate Zoölogy.

      _Racial characters._--Large size, very dark color.

      _Measurements._--The average of four male topotypes and the
      measurements of one female topotype are, respectively: total
      length 202, 201; length of tail 57, 59; hind foot 26, 27.

      _Distribution._--Southeastern Washington east of the range of
      _columbianus_ and north of the higher parts of the Blue Mountains.

=Thomomys talpoides wallowa= Hall and Orr

      _Thomomys quadratus wallowa_ Hall and Orr, Proc. Biol. Soc.
      Washington, 46:41, March 24, 1933.

      _Thomomys talpoides wallowa_ Goldman, Jour. Mamm., 20:234, May 15,

      _Type._--Obtained at Catherine Creek, 7 miles east of Telocaset,
      3,500 ft., Union County, Oregon, by R. T. Orr on June 29, 1932;
      type in Museum of Vertebrate Zoölogy.

      _Racial characters._--Similar to _devexus_ but much darker.
      Similar (in Washington) to _aequalidens_ but much smaller.

      _Measurements._--Four males and 4 females from Mountain Top and
      Stay-a-while Spring, Columbia County, average, respectively: total
      length 191, 180; length of tail 56, 52; hind foot 26, 25.

      _Distribution._--The higher parts of the Blue Mountains.

_Remarks._--Washington specimens referred to this race are intermediate
between _wallowa_ and _aequalidens_ but are colored like _aequalidens_.

=Thomomys talpoides fuscus= Merriam

      _Thomomys clusius fuscus_ Merriam, N. Amer. Fauna, 5:69, July 30,

      _Thomomys myops_ Merriam, Proc. Biol. Soc. Washington, 14:112,
      July 19, 1901 (type from Conconnully, Okanogan County,

      _Thomomys fuscus fuscus_ Bailey, N. Amer. Fauna, 39:126, November
      15, 1915.

      _Thomomys talpoides fuscus_ Hall and Dalquest, Murrelet, 20:4,
      April 30, 1939.

      _Type._--Obtained at the head of Big Lost River, Custer County,
      Idaho, by B. H. Dutcher on September 23, 1890; type in United
      States National Museum.

      _Racial characters._--Similar to _devexus_ but upper parts rich
      tawny in color.

      _Measurements._--Three males and 2 females from Newport, Pend
      Oreille County, average, respectively: total length 189, 186;
      length of tail 54, 57; hind foot 27, 26.

      _Distribution._--Along the eastern border of the state to the
      north of the Snake River, northeastern Washington, and the
      northeastern Cascades.

=Thomomys talpoides yakimensis= Hall and Dalquest

      _Thomomys talpoides yakimensis_ Hall and Dalquest, Murrelet, 20:4,
      April 30, 1939.

      _Thomomys talpoides badius_ Goldman, Jour. Mamm., 20:242, May 15,
      1939 (type from Wenatchee, Chelan County, Washington).

      _Type._--Obtained at Selah, Yakima County, Washington, by P.
      Burgner, on November 27, 1938; type in Museum of Vertebrate

      _Racial characters._--Similar to _fuscus_ but more orange, less

      _Measurements._--Four male and 3 female topotypes average,
      respectively: total length 209, 191; length of tail 60, 56; hind
      foot 27, 26.

      _Distribution._--The eastern edge of the Cascades from the
      Wenatchee Mountains south to the Simcoe Anticline.

=Thomomys talpoides shawi= Taylor

      _Thomomys douglasii shawi_ Taylor, Proc. Biol. Soc. Washington,
      34:121, June 30, 1921.

      _Thomomys talpoides shawi_ Hall and Dalquest, Murrelet, 20:4,
      April 30, 1939.

      _Type._--Obtained at Owyhigh Lakes, Mount Rainier, Pierce County,
      Washington, by G. G. Cantwell, on August 9, 1919; type in United
      States National Museum.

      _Racial characters._--A large, tan-colored pocket gopher, similar
      to _aequalidens_ but paler.

      _Measurements._--Two male and 7 female topotypes average,
      respectively: total length 227, 213; length of tail 72, 64; hind
      foot 32, 30.

      _Distribution._--The higher Cascade Mountains from Mount Rainier
      southward. Southern limits of range unknown.

=Thomomys talpoides immunis= Hall and Dalquest

      _Thomomys talpoides immunis_ Hall and Dalquest, Murrelet, 20:4,
      April 30, 1939.

      _Type._--Obtained 5 miles south of Trout Lake, Klickitat County,
      Washington, by W. W. Dalquest on July 27, 1937; type in Museum of
      Vertebrate Zoölogy.

      _Racial characters._--Similar to _shawi_ but paler and grayer.

      _Measurements._--Two males and 2 females from Morrison Springs
      Ranger Station, Skamania County, average, respectively: total
      length 211, 212; length of tail 64, 58; hind foot 28, 29.

      _Distribution._--The Cascade Mountains from the vicinity of Mount
      Adams north. The zone of intergradation between _shawi_ and
      _immunis_ is in the rugged, inaccessible mountains between Mount
      Rainier and Mount Adams.

=Thomomys talpoides limosus= Merriam

      _Thomomys limosus_ Merriam, Proc. Biol. Soc. Washington, 14:116,
      July 19, 1901.

      _Thomomys talpoides limosus_ Goldman, Jour. Mamm., 20:235, May 15,

      _Type._--Obtained at White Salmon, Klickitat County, Washington, by
      J. A. Loring on June 26, 1897; type in United States National

      _Racial characters._--Similar to _immunis_ but darker with
      smaller, shorter skull.

      _Measurements._--One male and 13 female topotypes average,
      respectively: total length 210, 198; length of tail 55, 56; hind
      foot 30, 28.

      _Distribution._--The lower Columbia River Valley, from the type
      locality east to the Columbian Plateau.

=Thomomys talpoides douglasii= (Richardson)

      _Geomys douglasii_ Richardson, Fauna Boreali-American, 1:200, 1829.

      _Geomys fuliginosus_ Schinz, Syn. Mamm., 2:136, 1846 (type from
      "Habitat ad fluvium Columbia").

      _Thomomys douglasii_ Allen, Bull. Amer. Mus. Nat. Hist., 5:66,
      April 28, 1893.

      _Thomomys douglasi douglasi_ Bailey, N. Amer. Fauna, 39:116,
      November 15, 1915.

      _Thomomys talpoides douglasii_ Goldman, Jour. Mamm., 20:234, May
      15, 1939.

      _Type._--Obtained at Fort Vancouver (now the city of Vancouver),
      Clark County, Washington, by David Douglas, probably in 1825.
      Probably not now in existence.

      _Racial characters._--A medium-sized, yellowish gopher with tiny,
      pointed ears and very small postauricular patches.

      _Measurements._--Two male and 10 female topotypes average,
      respectively: total length 227, 213; length of tail 68, 63; hind
      foot 30, 30; and weight 148, 117 grams.

      _Distribution._--Known only from Clark County.

=Thomomys talpoides glacialis= Dalquest and Scheffer

      _Thomomys talpoides glacialis_ Dalquest and Scheffer, Proc. Biol.
      Soc. Washington, 55:97, August 13, 1942.

      _Type._--Obtained 2 miles south of Roy, Pierce County, Washington,
      by W. W. Dalquest, on December 19, 1941; type in United States
      National Museum.

      _Racial characters._--A dark, yellowish-brown gopher with
      orange-tinged underparts.

      _Measurements._--Twenty male and 17 female topotypes average,
      respectively: total length 225, 220; length of tail 72, 71; hind
      foot 30, 30; and weight 128, 116 grams.

      _Distribution._--Known only from Roy Prairie, Pierce County.

=Thomomys talpoides tacomensis= Taylor

      _Thomomys douglasii tacomensis_ Taylor, Proc. Biol. Soc.
      Washington, 32:169, September 30, 1919.

      _Thomomys talpoides tacomensis_ Goldman, Jour. Mamm., 20:235, May
      15, 1939.

      _Type._--Obtained 6 miles south of Tacoma, Pierce County,
      Washington, by G. G. Cantwell on December 24, 1918; type in United
      States National Museum.

      _Racial characters._--A large, dark form; rich hazel in color with
      large, black postauricular patches and ochraceous underparts.

      _Measurements._--Thirteen male and 15 female topotypes average,
      respectively: total length 224, 196; length of tail 71, 57; hind
      foot 31, 29; and weight 127, 104 grams.

      _Distribution._--Restricted to the area about Steilacoom and
      Tacoma, Pierce County, Washington.

=Thomomys talpoides pugetensis= Dalquest and Scheffer

      _Thomomys talpoides pugetensis_ Dalquest and Scheffer, Proc. Biol.
      Soc. Washington, 55:96, August 13, 1942.

      _Type._--Obtained 4 miles south of Olympia, Thurston County,
      Washington, by W. W. Dalquest, on December 31, 1941; type in United
      States National Museum.

      _Racial characters._--Similar to _glacialis_ but underparts not
      tinged with orange and conspicuous dusky areas present on sides of

      _Measurements._--Fourteen male and 19 female topotypes average,
      respectively: total length 223, 205; length of tail 62, 59; hind
      foot 30, 29; and weight 123, 96 grams.

      _Distribution._--Known only from the type locality.

=Thomomys talpoides tumuli= Dalquest and Scheffer

      _Thomomys talpoides tumuli_ Dalquest and Scheffer, Proc. Biol. Soc.
      Washington, 55:96, August 13, 1942.

      _Type._--Obtained on Rocky Prairie, 7 miles north of Tenino,
      Thurston County, Washington, by W. W. Dalquest on January 2, 1942;
      type in United States National Museum.

      _Racial characters._--Similar to _pugetensis_ but grayer, less

      _Measurements._--Eleven male and 14 female topotypes average,
      respectively: total length 225, 216; length of tail 60, 64; hind
      foot 31, 30; and weight 140, 118 grams.

      _Distribution._--Known only from the type locality.

=Thomomys talpoides yelmensis= Merriam

      _Thomomys douglasi yelmensis_ Merriam, Proc. Biol. Soc. Washington,
      13:21, January 31, 1899.

      _Thomomys douglasii yelmensis_ Taylor, Proc. Biol. Soc.
      Washington, 32:169, September 30, 1919.

      _Thomomys talpoides yelmensis_ Goldman, Jour. Mamm., 20:235, May
      15, 1939.

      _Type._--Obtained at Tenino, Thurston County, Washington, by C. P.
      Streator on October 24, 1891; type in United States National

      _Racial characters._--A rather small, yellowish race with whitish

      _Measurements._--Twenty-one male and 21 female topotypes average,
      respectively; total length 213, 202; length of tail 64, 61; hind
      foot 29, 28; and weight 121, 101 grams.

      _Distribution._--Known only from Grand Mound Prairie (type
      locality), Vail Prairie, near Vail, and Rochester Prairie, near
      Rochester, all in Thurston County.

=Thomomys talpoides couchi= Goldman

      _Thomomys talpoides couchi_ Goldman, Jour. Mamm., 20:243, May 15,

      _Type._--Obtained on Scotts Prairie, 4 miles north of Shelton,
      Mason County, Washington, by L. K. Couch, on June 27, 1922; type in
      United States National Museum.

      _Racial characters._--A small race; rich hazel in color.

      _Measurements._--Thirteen male and 9 female topotypes average,
      respectively: total length 196, 191; length of tail 55, 53; hind
      foot 27, 27; and weight 87, 79 grams.

      _Distribution._--Known only from the type locality and from Lost
      Lake Prairie, near Satsop, Mason County.

=Thomomys talpoides melanops= Merriam

      _Thomomys melanops_ Merriam, Proc. Biol. Soc. Washington, 13:21,
      January 31, 1899.

      _Thomomys douglasi melanops_ Bailey, N. Amer. Fauna, 39:119,
      November 15, 1915.

      _Thomomys talpoides melanops_ Goldman, Jour. Mamm., 20:235, May
      15, 1939.

      _Type._--Obtained at head of Sol Duc River, Clallam County,
      Washington, by V. Bailey on August 28, 1897; type in United States
      National Museum.

      _Racial characters._--Most like _shawi_ but smaller, with larger
      postauricular patches and dusky areas on side of head.

      _Measurements._--A male topotype measures: total length 211;
      length of tail 67; hind foot 28.

      _Distribution._--Higher Olympic Mountains.

=Castor canadensis= Kuhl


_Description._--The beaver is the largest rodent occurring in
Washington. Large individuals weight approximately 50 pounds. They are
heavily-built, robust animals with large heads and short necks. The
large, flat, naked tail immediately separates them from all other
mammals occurring in the state. The forelegs are short and the forefeet
handlike. The hind legs are long, thick and powerful. The hind feet are
large and webbed for swimming. The ears are small and the eyes, although
of moderate size, are not prominent. The incisors are large and
prominent with a distinct yellow or orange color. The close, dense
underfur is overlaid with long, lax, rather stiff overfur or guardhairs.

  [Illustration: FIG. 99. Beaver (_Castor canadensis_). Silvana,
  Washington, August 15, 1921. (Fish and Wildlife Service by Leo K.
  Couch, No. B-21912.)]

Beavers are found in Europe, Asia and North America. In America they
range from Alaska to Mexico and from the Atlantic to the Pacific. Many
American subspecies have been described, all thought to belong to a
single species.

Beavers live in streams and lakes of Washington under diverse climatic
conditions. They are perhaps most abundant in the western Washington
lowlands where there are numerous watercourses. In the Cascade
Mountains they are found well up into the Canadian Life-zone, where the
streams are swift and clear with stony bottoms. The highest altitudinal
record known to us is Reflection Lake, Mt. Rainier, 4,861 feet
(Brockman, 1939: 71). Farther east, in the timbered regions of the
eastern Cascades and in northeastern Washington, beavers live in deeper,
more sluggish streams. Abundant beaver signs were noted at the junction
of the Snake and Columbia rivers where the animals were occupying one of
the hottest, most desertlike portions of the state. They are found in
Moses Lake, in the center of the arid Columbian Plateau. Beavers once
occurred in the San Juan Islands and have recently been reintroduced

  [Illustration: FIG. 100. Beaver (_Castor canadensis_): lodge and
  beaver pond, Elbe, Washington, August 24, 1926. (T. H. Scheffer

The part the beaver played in man's exploration of the state of
Washington is a story in itself. The establishment of Fort Vancouver and
Fort Spokane and other settlements was principally due to the trade in
beaver pelts. These forts served as headquarters for Douglas, Suckley,
Townsend, Nuttall and other early naturalists who contributed so greatly
to the knowledge of the mammalian fauna of the western United States.

  [Illustration: FIG. 101. Cottonwood pole carved by beaver, Lake
  Wenatchee, Washington, May 13, 1938. (Fish and Wildlife Service photo
  by Victor B. Scheffer, No. 33.)]

Trapping for fur drastically reduced the number of beavers in
Washington. After trapping was prohibited they were slow in recovering
and, until about 1930, they were uncommon. After that date they seemed
to increase rapidly, becoming abundant about 1940. The present system of
removing beavers only from areas where they are doing damage, and then
only under strict supervision from the State Department of Game, has
held their numbers at a high level.

The food of the beaver varies greatly with locality. Along the Columbia
River, in eastern Washington, cottonwood (_Populus hastata_) and willow
(_Salix_ sp.) seem to constitute favorite foods. Few other trees grow in
this arid region. On Puget Island, near the mouth of the Columbia,
willow, alder (_Alnus oregonus_) and probably other shrubs are eaten. In
the many streams that run from the western Cascade Mountains to Puget
Sound, various species of willow seem to be the principal food eaten.
Some alder, cascara, Douglas fir, red huckleberry and salmonberry are
eaten. In some streams on the eastern slope of the Cascades, coniferous
trees, including Douglas fir, yellow pine (_Pinus ponderosa_), and red
cedar (_Thuja plicata_) are used.

Beaver dams are abundant in certain sections of Washington, notably the
area between the Cascade Mountains and Puget Sound. Dams several hundred
feet in length were seen in the valley of Cherry Creek, 7 miles
northeast of Duvall, King County. However, in most parts of the state
few dams are built. No dams are necessary in the larger streams and
lakes while the smaller mountain streams possess a gradient too steep
for successful damming. Spring floods and high water following rains
would destroy such dams.

True beaver houses, or lodges, are seldom seen in Washington. In the
larger streams, rivers and lakes the beavers are "bank beavers," living
in burrows with underwater entrances. In the mountain streams, log jams
furnish protection. Great piles of logs and debris accumulate at bends
of streams in periods of high water. These jams commonly contain large,
well-anchored key logs and persist for years. Back-cutting by the stream
often forms deep pools before the jams while the sloping bank opposite
may support willows and alders. Thus beavers may find a home, food, and
a protecting pool of deep water at a log jam. In periods of low water,
and when meandering of the stream causes it to leave the log jam, canals
are constructed by the beavers to connect the log jam with the stream.

Where dams are built a few lodges may occur. For the most part, however,
the beavers dig holes in the bank or in small islands that result from
their dam-building activities. Burrows are occasionally easy to locate
because of the mud and sticks laid on the ground above. At best these
sticks form a layer only a few inches thick and are too loosely packed
to furnish any additional protection. They probably represent a
concession to the beavers' lodge-building instincts.

The beaver probably breeds in January. A specimen from Cle Elum, taken
on March 1, contained 4 embryos and one from the Teanaway River near Cle
Elum held 2 embryos on February 28. One from Swauk Creek, Kittitas
County, on the eastern slope of the Cascades, had 3 embryos on February
12. A young beaver was born to a captive animal on May 18. Its mother
was taken at Soos Creek in southeastern King County. This baby weighed
884 grams.

The majority of the beavers trapped are small animals, measuring from
725 to 900 mm. (total length) in February, and weighing from 10 to 20
pounds. These are kits, probably one and two years old. It is rather
difficult to divide these animals into two age groups but probably the
first year young (7 to 11 months old) weigh from 10 to 15 pounds and
measure between 725 and 800 mm. in total length. Two-year olds (19 to 23
months) measure from 800 to 925 mm. and weigh from 15 to 25 pounds. They
measure approximately 1050 mm. the third year, when they become adult,
and weigh in the neighborhood of 45 pounds. Individuals occasionally
weigh more than 50 pounds; these are probably old adults.

Damage by beavers is limited. Their dams occasionally flood roads and
rarely fields. In some parts of eastern Washington, where fruit raising
is an important industry, an individual beaver may cause considerable
damage by cutting valuable trees. Ornamental trees may be damaged near
the larger cities. Beavers living in Lake Washington in almost the
center of the city of Seattle, caused some damage to ornamental trees on
the University of Washington Golf Course. Canals and dams in small
streams and ditches in areas where stock raising is practiced, flood
stock food and sometimes form mudholes or bogs where stock might be
trapped. Where individual beavers are doing damage they are trapped and
killed, or are live-trapped and transported to more favorable habitats.
In places where any beaver may be potentially dangerous, as in the
fruit-growing areas of eastern Washington or stock-raising areas in
western Washington, their numbers should be kept down by continued

  [Illustration: FIG. 102. Forest road flooded by beavers which built a
  dam on the upstream (left) side of a bridge; Tronson Creek, Blewett
  Pass, Wenatchee National Forest, May 13, 1938. (Fish and Wildlife
  Service by Victor B. Scheffer, No. 30.)]

Probably the greatest value of beavers lies in the impounding of water
by their dams. This is especially true in the arid mountains of eastern
Washington and in the logged-off or burned-over parts of western
Washington. These ponds prevent erosion, raise the water table and so
stimulate the growth of vegetation, insure greater regularity of stream
flow, provide suitable water for trout and muskrats as well as some
breeding grounds for waterfowl. On burned-over or logged-off land, the
greatest, and often the only, growth of new conifers is in the vicinity
of beaver ponds. In the vicinity of beaver dams, rich growths of willows
and other deciduous vegetation provides food and cover for deer and
fur-bearing mammals.

  [Illustration: FIG. 103. Distribution of the beaver in Washington. A.
  _Castor canadensis leucodonta._ B. _Castor canadensis idoneus._]

Trapping of beaver for their fur at present is not a private enterprise
but controlled trapping by the state adds to public funds.

=Castor canadensis leucodonta= Gray

      _Castor canadensis leucodonta_ Gray, Ann. and Mag. Nat. Hist., 4
      (ser. 4):293, October, 1869.

      _Castor canadensis pacificus_ Rhoads, Trans. Amer. Philos. Soc.,
      19 (n. s.):422, September, 1898 (type from Lake Keechelus,
      Kittitas County, Washington).

      _Type._--Obtained by Dr. Robert Brown, probably on Vancouver
      Island, British Columbia.

      _Racial characters._--Large size; reddish color; elongate skull.

      _Distribution._--Found over all but the southwestern corner of the
      state. The most westward locality from which specimens have been
      examined is Lake Crescent (U. S. N. M.).

_Remarks._--The good series of beaver skulls available from several
areas of Washington has led to careful comparison of Washington material
with specimens from Vancouver Island. Skulls from extreme southwestern
Washington are small and decidedly short, being referable to the race
_idoneus_. No satisfactory way of distinguishing between the others
could be found and all are consequently referred to the earliest named
form, _leucodonta_.

=Castor canadensis idoneus= Jewett and Hall

      _Castor canadensis idoneus_ Jewett and Hall, Jour. Mamm., 21:87,
      February 15, 1940.

      _Type._--Obtained at Foley Creek, tributary to Nehalem River,
      Tillamook County, Oregon, by C. Leach on December 15, 1914; type in
      California Museum of Vertebrate Zoölogy.

      _Description._--Similar to _leucodonta_ but smaller, darker, with
      shorter and wider skull.

      _Distribution._--Lowlands near the mouth of the Columbia River.
      Specimens from Puget Island (U. S. N. M.) have been examined.

=Onychomys leucogaster fuscogriseus= Anthony

Northern grasshopper mouse

      _Onychomys leucogaster fuscogriseus_ Anthony, Bull. Amer. Mus. Nat.
      Hist., 32: 11, March 7, 1913.

      _Type._--Obtained at Ironside, Malheur County, Oregon, by H. E.
      Anthony, in 1912; type in American Museum of Natural History.

      _Measurements._--Nine males and 5 females from Franklin, Benton,
      and Walla Walla counties, Washington, average, respectively: total
      length 132.3, 133.0; length of tail 35.8, 35.6; hind foot 19.2,
      19.6; ear 17.6, 17.6; weight 23.3, 24.1 grams.

      _Distribution._--The Columbian Plateau, southeastern Washington,
      and the Yakima Valley, ranging north to Douglas (Taylor and Shaw,
      1929: 21), west to Yakima (M. V. Z.), east to Asotin (Taylor and
      Shaw, 1929: 21) and south to Wallula (Taylor and Shaw, 1929: 21).

  [Illustration: FIG. 104. Distribution of the northern grasshopper
  mouse, _Onychomys leucogaster fuscogriseus_, in Washington.]

_Description._--A mature grasshopper mouse is slightly larger than a
house mouse; the head and body measure about 4 inches and the tail about
1-1/4 inches; it may be recognized by the rather plump body, naked
ears, short, thick and tapering tail and soft, dense fur. The upper
parts of adults are buffy. Younger animals are blue-gray above.
Underparts and tail are white.

Grasshopper mice of two species range over much of northern Mexico and
the western United States. The species occurring in Washington,
_leucogaster_, is found from southern Manitoba to northern Mexico, and
from the Pacific states east to North Dakota. This species is
characteristic of the Upper Sonoran Life-zone, and usually is associated
with sagebrush and greasewood. It prefers open areas of sand and avoids
heavy cover, and seems to be strictly nocturnal. It eats vegetation,
seeds, insects, or the flesh of other mice. According to Bailey (1936:
178), grasshopper mice pursue, catch and kill other mice. They hunt by
scent and follow tracks like a wolf. They make a sound, or "call," like
a miniature wolf howl. They are said to be rather docile and make
friends with humans quickly.

On some occasions grasshopper mice are locally abundant, but for the
most part they are rather uncommon over their range. Usually one is
taken in about 200 traps set in suitable habitats. They do not
hibernate; one was caught at Moses Lake, Grant County, on a freezing
winter night.

The grasshopper mouse is almost always associated with the pocket mouse
(_Perognathus parvus_), deer-mouse (_Peromyscus maniculatus_), and often
the harvest mouse (_Reithrodontomys megalotis_). The same snakes, owls,
and carnivorous mammals that prey on these mice doubtless eat also the
grasshopper mouse.

R. D. Svihla (1936: 172) determined the gestation period of 3 lactating
grasshopper mice captured at Lind, Adams County, as 33, 39 and 47 days.
A non-lactating female gave birth to a litter in 32 days. Young varied
from 2 to 5 in number and were pink and hairless at birth, with eyes and
ears closed.

=Reithrodontomys megalotis megalotis= (Baird)

Western harvest mouse

      _Reithrodon megalotis_ Baird, Mamm. N. Amer., p. 451, 1857.

      _Reithrodontomys megalotis nigrescens_ Howell, N. Amer. Fauna,
      36:32, June 5, 1914 (type from Payette, Idaho).

      _Reithrodontomys megalotis megalotis_ Bailey, N. Amer. Fauna,
      55:189, August 29, 1936.

      _Type._--Obtained between Janos, Chihuahua, and San Luis Springs,
      New Mexico, by C. B. R. Kennerly, in 1855; type in United States
      National Museum.

      _Measurements._--Five males and 6 females, showing moderate wear on
      the third upper molars, from Walla Walla County, average,
      respectively: total length 141, 137.5; length of tail 71.8, 68.1;
      hind foot 17.2, 16.7; weight (males only) 10.5 grams. A female, 136
      mm. long, from Grant County, weighed 9.5 grams. Over the range of
      _megalotis_ in the United States, males average consistently larger
      than females in length of the hind foot and consistently less in
      length of head and body.

      _Distribution._--The Columbian Plateau, the Upper Sonoran
      Life-zone area west of the Columbia River and south of the
      Wenatchee Mountains in Kittitas, Yakima, and Benton counties, the
      north side of the Columbia River in Klickitat County, and Okanogan
      County east of the Okanogan River. Marginal localities are:
      Timentwa (W. W. D.) on the north, Maryhill (M. V. Z.) on the west,
      Wallula (M. V. Z.) on the south, and Colfax (Taylor and Shaw,
      1929: 21) on the east.

  [Illustration: FIG. 105. Distribution of the western harvest mouse,
  _Reithrodontomys megalotis megalotis_, in Washington.]

_Remarks._--Harvest mice from the states of Washington, Oregon, Idaho,
Utah, Nevada, California, Arizona, and New Mexico were examined to
ascertain the validity of the name _Reithrodontomys megalotis
nigrescens_ Howell (type from Payette County, Idaho). No cranial
characters or measurements could be found to separate _nigrescens_ from
_megalotis_. Specimens from Washington, Oregon, and west-central Idaho,
within the range ascribed to _nigrescens_, averaged slightly darker in
color with a more distinct, blackish dorsal line than specimens from New
Mexico. This difference is slight, and specimens from any one locality
vary greatly in color. Some Washington specimens are as pale as the
palest material from New Mexico, and some of the darkest _nigrescens_
can be matched by specimens of _megalotis_ from Arizona and New Mexico.
This slight color variant is not worthy of recognition as a distinct
subspecies, and I agree with Howell (1939: 390) that _nigrescens_ is a
synonym of _megalotis_.

_Description._--The western harvest mouse is about the size and shape of
the common house mouse. The length of the tail is approximately equal to
the length of the head and body. The tail is slim and lightly haired.
The upper parts are brownish buff lightly washed with blackish. A faint
but distinct dark dorsal stripe is usually present. The underparts are
white. From _Mus_ and _Peromyscus_, _Reithrodontomys_ can be
distinguished by the grooves on the anterior face of the upper incisors
and the long brown hairs at the bases of the ears.

This genus reaches the extreme northern limit of its distribution in the
state of Washington, where it ranges to the Canadian border. A single
subspecies occurs in Washington. The harvest mouse is a resident of the
Upper Sonoran Life-zone and it is most abundant in dense growths of
grass, weeds, and other vegetation. Along the Yakima River, near
Ellensburg, it was abundant in thick, river-side jungle. Near Yakima
many specimens were taken in a marsh of cattails and tules. In the Grand
Coulee they live in thick grass. Near Moses Lake several were found in
their nests in a thick growth of tumbleweed. Harvest mice occur also,
although never in great numbers, in relatively open areas. At the
Potholes near Moses Lake, Grant County, a few were caught on sand dunes
among scattered bushes of sage. They were taken on sagebrush flats in
several localities. Gray (1943: 191) estimates their numbers in the
sagebrush area of the Yakima Valley at 5 per acre.

Where harvest mice live in dense vegetation they make small runways and
food piles similar to those of _Microtus_ but in more open areas they
leave no perceptible signs. Near Yakima the nest of a harvest mouse was
found among the roots of an overturned apple tree. This nest was an
irregular ball composed of fine grasses and shredded bark closed above,
with a side entrance and a central cavity three inches in diameter. Near
Moses Lake three nests of harvest mice were discovered on the ground
under cover of a dense growth of tumbleweed, and six live harvest mice
were captured in the same area. These nests were cuplike, open above,
and were composed of dry grasses, bits of weeds, and newspaper. The
smoothly rounded inner cups measured about three inches in diameter.

Embryos were found in harvest mice in Washington from March to April and
numbered 3 to 6.

=Peromyscus maniculatus= (Wagner)

Deer mouse

_Description._--The deer mouse is slightly larger than the house mouse,
the head and body measuring from 3-1/2 to 4 inches. The length of the
tail varies considerably in the several races, ranging from
approximately 2-1/2 inches in _gambelii_ to 4 inches or more in some
individuals of _oreas_. The eyes are large and protrude slightly. The
ears are large, erect and naked. The tail is sparsely furred with short,
posteriorly directed hairs. The color of the upper parts varies from
yellowish buff in _gambelii_ to reddish brown in _oreas_ and dusky in
_austerus_. Young deer mice are gray or bluish gray above. Underparts
and feet are white. The tail is bicolored, being dusky above and white

  [Illustration: FIG. 106. Deer mouse (_Peromyscus maniculatus
  austerus_), in captivity, Seattle, Washington, September 18, 1939.
  (Fish and Wildlife Service photo by Victor B. Scheffer, No. 723.)]

In Washington _Peromyscus_ may be separated from other naked-eared mice
by ungrooved teeth and slightly tapering, bicolor tail. Nearly every
part of North America is inhabited by one or more kinds of _Peromyscus_.
The single species, _maniculatus_, which occurs in Washington, ranges
from the Atlantic to the Pacific and from the Arctic to tropical Mexico.
Osgood (1909: 17) remarks: "... it is probable that a line, or several
lines, could be drawn from Labrador to Alaska and thence to southern
Mexico throughout which not a single square mile is not inhabited by
some form of this species."

Deer mice are the most abundant mice in Washington. They occur at sea
level on the ocean beaches and above timber-line, even breeding in the
Arctic-alpine Life-zone. They occur on the islands in Puget Sound, the
lowland marshes and deciduous thickets of western Washington, the great
conifer forests, the alpine cirques and mountain parks, and the deserts
of eastern Washington. Almost everywhere they are the commonest mammal
encountered. They make their homes in city dwellings, under logs or in
hollow trees in the forest, in the burrows of larger rodents and, if
necessary, in burrows of their own construction.

Nests are usually cup-shaped masses of soft material, 4 to 8 inches in
diameter with central cavities 2 to 3 inches in diameter. Materials
utilized include dry grass, shredded bark, rope, rags, newspaper, moss,
cattail fluff, fur, wool, and feathers. One nest discovered in the attic
of an old building near Coulee City, Grant County, consisted of the
mummified body of a wood rat (_Neotoma cinerea_). The fur of the arched
body of the rat formed one side of the nest, while the fur of the far
side of the body had been plucked to form the remainder. In the
extensive marshes along Lake Washington, King County, deer mice occupy
the ball-shaped nests of tule wrens (_Telamtodytes palustris_). These
are constructed entirely of fragments of cattail leaves and cattail
fluff and are placed in the cattail rushes about four feet above the wet
ground of the marsh. Other deer mouse nests were found in an unused
typewriter, in the pocket of an old coat, in a window frame, under a
piece of tar paper on the ground, in a cavity in the ground under a
board, between a ceiling and a loose bit of roofing paper, under rocks
and logs, in hollows in logs, and in an unused nest of a wood rat. Where
plant cover is dense, nests are placed on the surface of the ground, as
under thick growths of tumbleweed (_Salsola_) in eastern Washington.

Deer mice are definitely nocturnal. Of thousands of specimens trapped
only a few were taken in the daytime. Two of these were taken in a rock
slide at Pass Creek Pass, Pend Oreille County, on the same day, and some
unusual conditions may have incited them to travel in daylight. Deer
mice are active throughout the winter and their lacy footprints are seen
on the snow in below-freezing weather. They are mainly terrestial but
one race (_oreas_) is at least partly arboreal for it is often taken in
traps set in trees for flying squirrels. One was caught in a trap set 50
feet from the ground. All are good climbers, and _oreas_ is perhaps the
best. One was watched as it climbed the rough cedar shake wall of a
cabin, ran upside-down across three feet of rough ceiling, and
descended the opposite wall head first. They are equally agile in
climbing rocks and cliffs.

The ordinary gait of the deer mouse is a steady run. When frightened it
may leap a distance of one foot or more. When surprised in its nest it
usually remains in the vicinity, hopping about in confusion, returning
time after time to the exposed nest.

The food of deer mice consists principally of vegetable material such as
seeds, nuts, berries, and the soft parts of fleshy plants. Mushrooms and
lichens are eaten. Insect remains are present in small quantities in
most stomachs examined. Flesh is also eaten and small mammals caught in
traps are often eaten by deer mice. Cannibalism appears to be common
only in captivity.

  [Illustration: FIG. 107. Distribution of the deer mouse in Washington.
  A. _Peromyscus maniculatus oreas._ B. _Peromyscus maniculatus
  austerus._ C. _Peromyscus maniculatus hollisteri._ D. _Peromyscus
  maniculatus gambelii._ E. _Peromyscus maniculatus artemisiae._]

Deer mice are often heavily parasitized by tapeworms and roundworms.
Virtually all of the deer mice taken on Jones Island in the San Juan
Islands had livers so infected by the eggs of a nematode worm as to be
swollen to several times their natural size, and had a yellow,
crystalline appearance. The maggots of parasitic flies often occur in
the anal and genital region of deer mice, effectively sterilizing some

Predatory snakes, birds, and mammals doubtless kill and eat deer mice.
Strangely enough, although deer mice may be the commonest mammal in an
area, the pellets of owls collected in the same area usually contain the
remains of relatively few deer mice. The impact of owls seems to be
greatest upon mice, such as _Microtus_, which follow definite runways.

The breeding season of the deer mouse extends over most of the spring
and summer. Growth and development of the young of _Peromyscus m. oreas_
have been discussed by A. Svihla (1936A). Embryos found varied in number
from 2 to 8 with a mean of 4.5.

=Peromyscus maniculatus oreas= Bangs

      _Peromyscus oreas_ Bangs, Proc. Biol. Soc. Washington, 12:84, March
      24, 1898.

      _Peromyscus akeleyi_ Elliot, Field Columb. Mus. Publ. 30., zoöl.
      ser., 1:226. February 1, 1899 (type from Elwha River, Olympic
      Mountains, Washington).

      _Peromyscus maniculatus oreas_ Osgood, N. Amer. Fauna, 28:51,
      April 17, 1909.

      _Type._--Obtained on Church Mountain, 6,500 ft., Mt. Baker Range,
      near the U. S. boundary, British Columbia, by Allan Brooks on
      August 29, 1896; type in Museum of Comparative Zoölogy.

      _Racial characters._--Size large; tail more than 50 per cent of
      total length; color of upper parts reddish.

      _Measurements._--Twenty-five males and 15 females average: total
      length 202; length of tail 110; hind foot 22.6; ear 18.7.

      _Distribution._--From the Cascade Mountains west to the Pacific
      Ocean, save for the Puget Sound area and a narrow band extending
      southward nearly to the Columbia River. Marginal localities along
      the west side of Puget Sound are: Deer Park (W.W.D.), Lake Cushman
      (W.W.D.), and Kelso (M.V.Z.). Marginal localities on the west side
      of the Cascade Mountains are: Tomyhoi Lake (W.W.D.), Forks of
      Skykomish River (W.W.D.), North Bend (W.W.D.), Mt. St. Helens
      (V.B.S.) and Yacolt (M.V.Z.). Marginal localities on the east
      slope of the Cascade Mountains are: Grouse Creek (W.W.D.), Lake
      Wenatchee (W.W.D.), Easton (W.W.D.), Morrison Springs Ranger
      Station (W.W.D.) and Wind River C.C.C. Camp (W.W.D.).

=Peromyscus maniculatus hollisteri= Osgood

      Peromyscus maniculatus hollisteri Osgood, N. Amer. Fauna, 28:62,
      April 17, 1909.

      _Type._--Obtained at Friday Harbor, San Juan Island, San Juan
      County, Washington, by N. Hollister on October 23, 1903; type in
      United States National Museum.

      _Racial characters._--Similar to _oreas_ but darker, less reddish,
      and with much shorter tail. Length of tail about 40 per cent of
      total length.

      _Measurements._--Populations of deer mice on the various islands
      vary slightly in average measurements. The average measurements of
      10 males and 10 females from San Juan Island, 20 males and 15
      females from Blakeley Island, and 20 males and 15 females from
      Cypress Island, are respectively: total length 189, 183, 179;
      length of tail 87, 82, 79; hind foot 22.8, 22, 19.5; ear 19.4,
      18.7, 19.5; weight 33, 29, 32 grams.

      _Distribution._--The San Juan Islands, San Juan and Skagit
      counties, in northern Puget Sound.

=Peromyscus maniculatus austerus= (Baird)

      _Hesperomys austerus_ Baird, Proc. Acad. Nat. Sci. Philadelphia,
      7:336, 1855.

      _Sitomys americanus austerus_ Allen, Bull. Amer. Mus. Nat. Hist.,
      5:192, August 18, 1893.

      _P[eromyscus]., austerus_ Bangs, Amer. Nat., 31:75, January 1,

      _Peromyscus maniculatus austerus_ Osgood, N. Amer. Fauna, 28:63,
      April 17, 1909.

      _Type._--Obtained at Old Fort Steilacoom, Pierce County,
      Washington, by J. G. Suckley, probably on January 20, 1854; type in
      United States National Museum.

      _Racial characters._--Similar to _oreas_ but smaller with
      relatively shorter tail; tail narrower at base, less sharply
      bicolor; upper parts duller, browner, less reddish. In _oreas_ the
      tail is usually more than 100 mm. In _austerus_ the tail is
      usually less than 90 mm.

      _Measurements._--Twenty-seven males and 23 females from Seattle,
      King County, average: total length 180.5; length of tail 88.6;
      hind foot 21; ear 18. The average weight of 15 adults is 21 grams.

      _Distribution._--The Puget Sound area and south to the Columbia
      River in a narrow band through the lowlands. Marginal localities
      on the west side of Puget Sound are: Port Angeles (W.W.D.),
      Bremerton (W.W.D.), 4 miles north of Shelton (W.W.D.), Tenino
      (W.W.D.) and Mouth of Klama River (M.V.Z.). Marginal localities on
      the east side of Puget Sound are: Bellingham (W.W.D.), Arlington
      (W.W.D.), Everett (W.W.D.), 4 miles east of Monroe (W.W.D.), Fall
      City (W.W.D.), and Vancouver (W.W.D.).

=Peromyscus maniculatus rubidus= Osgood

      _Peromyscus oreas rubidus_ Osgood, Proc. Biol. Soc. Washington,
      14:193, December 12, 1901.

      _Peromyscus maniculatus rubidus_ Osgood, N. Amer. Fauna, 28:65,
      April 17, 1909.

      _Type._--Obtained at Mendocino City, California, by J. A. Loring on
      November 17, 1897; type in United States National Museum.

      _Racial characters._--Closely similar to _austerus_ but paler with
      longer skull. Specimens from Washington are almost purplish-brown
      in appearance.

      _Measurements._--Six males and 2 females from Puget Island,
      Wahkiakum County, average: total length 177; length of tail 89;
      hind foot 21.9; ear 18.7.

      _Distribution._--This is the deer mouse of the humid coastal belt
      of Oregon and California. It enters Washington only at Puget
      Island, Wahkiakum County, in the Columbia River.

=Peromyscus maniculatus gambelii= (Baird)

      _Peromyscus gambelii_ Baird, Mamm. N. Amer., Pacific R. R. Reports,
      8:464, 1857.

      _Sitomys americanus gambelii_ Allen, Bull. Amer. Mus. Nat. Hist.,
      5:190, August 18, 1893.

      _P[eromyscus]., t[exanus]. gambelii_ Mearns, Proc. U. S. Nat.
      Mus., 18:446, March 25, 1896.

      _Peromyscus maniculatus gambeli_ Osgood, N. Amer. Fauna, 28:67,
      April 17, 1909.

      _Type._--Obtained at Monterey, Monterey County, California, by W.
      P. Trowbridge; skin catalogued on January 4, 1853; type in United
      States National Museum.

      _Racial characters._--Size small; tail short; color pale, varying
      from buffy gray to rich buff; rarely dark reddish brown.

      _Measurements._--Twenty-five males and 15 females average: total
      length 160; length of tail 71; hind foot 19.8; ear 17.7.

      _Distribution._--The Columbian Plateau. Intergradation with
      _oreas_ occurs between the Columbia River and the Cascade
      Mountains. Marginal records on the west are: Chelan (W.W.D.),
      Leavenworth (W.W.D.), Cle Elum (W.W.D.), Satus Pass (W.W.D.), Lyle
      (V.B.S.). Marginal localities on the north are: Chelan (W.W.D.), 5
      miles north of Coulee (W.W.D.), Sprague (V.B.S.) and Steptoe Butte

=Peromyscus maniculatus artemisiae= (Rhoads)

      _Sitomys americanus artemisiae_ Rhoads, Proc. Acad. Nat. Sci.
      Philadelphia, p. 260, October 23, 1894.

      _Peromyscus texanus artemisiae_ Miller and Rehn, Proc. Boston Soc.
      Nat. Hist., 30:84, December, 1901.

      _Peromyscus maniculatus artemisiae_ Osgood, N. Amer. Fauna, 28:58,
      April 17, 1909.

      _Type._--Obtained at Ashcroft, British Columbia, by S. N. Rhoads on
      July 5, 1892; type in Philadelphia Academy of Sciences.

      _Racial characters._--Similar to _gambelii_ but darker, more
      reddish; often with longer tail.

      _Measurements._--Ten males and 10 females from northeastern
      Washington average: total length 167; length of tail 80; hind foot
      20; ear 17.8.

      _Distribution._--Northeastern Washington and the Blue Mountains of
      southeastern Washington, extending west, in northeastern
      Washington, to Conconully (W.W.D.) and south to Duly Lake

=Neotoma cinerea= Ord

Bushy-tailed wood rat

_Description._--The bushy-tailed wood rat is slightly larger than the
common Norway or brown rat. The head and body measure about 12-1/2
inches and the tail about 3-1/2 inches. It resembles the deer mouse in
general proportions. The ears are large and naked and the black eyes are
large and protruding. The tail is bushy, squirrel-like. The feet are
small and have furry soles. The fur is soft and silky. Adults are
brownish gray above with white underparts and a gray tail. Young
individuals have blue-gray upper-parts.

Wood rats range over most of North America. They exhibit great
variation, especially in the southwestern United States. The
bushy-tailed species occurs in the western United States and Canada.
Wood rats are notorious for invading buildings in the mountains and in
the desert. However, their natural habitat is broken rock or talus. This
habitat preference accounts for their distribution in Washington, for
talus is common except in the humid subdivision of the Transition
Life-zone. Wood rats are probably most abundant in the talus slides of
the Columbian Plateau, especially in the canyon of the Columbia River
and in Moses and Grand coulees. They are common in all the mountainous
areas in the state where high altitudes and steep slopes result in the
accumulation of talus. Wood rats range from sea level to 10,000 feet
elevation on Mount Rainier and from the Upper Sonoran to the
Arctic-alpine life-zones. Great horned owls and probably all of the
carnivores that share the range of the wood rat prey on it to some
extent. Sperry (1941: 15) lists _Neotoma_ in four per cent of 8,339
coyote stomachs gathered throughout the United States.

Wood rats in Washington are definitely nocturnal and are rarely seen in
daylight. On January 10, 1939, however, near Colville, Stevens County, a
companion and I were sitting on the porch of a deserted shack eating
lunch. Suddenly a wood rat darted out of the open door into the full
sunlight and tugged at a can, containing a few drops of tomato juice,
which one of us was holding in his hand. When the rat looked up and saw
the human, it hastily retreated. The wood rat has a habit of flattening
its body tightly against the ground when observed and also of "drumming"
with its feet when excited. This habit is shared by the snowshoe rabbit
and the spotted skunk. A captive specimen "drummed" by lifting its back
feet, alternately, about one-half inch from the surface of a piece of
wood and striking downward with surprising power. The agility of the
wood rat was demonstrated near Wallula where we watched a specimen by
the light of a flashlight as it climbed an almost vertical rock wall,
taking advantage of small fractures for toe-holds.

Wood rats are far from noiseless in their nocturnal activities. Near
Moses Coulee, Douglas County, wood rats were heard from a distance of 50
feet as they scampered back and forth through a concrete culvert under
the highway.

Many species of wood rats build complicated stick houses in which they
live. In Washington, elaborate stick houses are sometimes built but are
usually not occupied. In the attic of an old building near Pend Oreille
Lakes, Stevens County, two large and complicated houses were discovered.
These were built of sticks, paper and other debris and measured more
than three feet in diameter. Three small, cup-shaped nests resembling
those of a bird, made of soft grasses, moss, and shredded paper were
found on the attic floor ten feet or more from the stick houses. These
nests gave every indication of being used, while the piles of sticks
contained no nests and seemed never to have been occupied. Near Moses
Lake, Grant County, a wood rat house built in the rafters of an old shed
was constructed entirely of tumbleweeds (_Salsola pestifer_). This house
was spherical in shape and measured more than five feet in diameter. It
appeared to be unoccupied and a wood rat, probably its builder, was
living under the floor of the shed. Most wood rat houses consist of a
half-dozen sticks arranged in a crevice in a rock pile or a cave. Wood
rats were placing fresh material on a house in the Wenatchee National
Forest on January 17, 1939, when there was two feet of snow on the
ground and the temperature was hovering around zero. The house measured
about one cubic yard and consisted of _Ceanothus_ leaves and steins,
with some twigs of fir (_Abies_). It was built under a bunk in a
broken-down cabin left by deer hunters.

The collecting habit of the wood rat has gained it the name "pack-rat."
Taylor (1920A: 91) lists the following materials from a wood rat nest on
Mt. Rainier: _Nesting material_: rags, leaves, paper, thumb of a glove,
string, thongs, oakum; _Food_: apple core, onion peel, bacon rind,
raisins, 10 bars of chocolate, figs, puff balls, bread crust, meat
scraps, cantaloupe rind, potatoes, dried apricots, lemons, mushrooms,
beans, peanuts, banana, 15 lumps of sugar; _Miscellaneous_: dime, coffee
can cover, paraffin from jelly glass, bones, 19 pieces of candles and
several cakes of soap.

As a rule only one or two wood rats are trapped at a given locality,
indicating that the species is not gregarious. The presence of wood rats
in a building, cave, mine tunnel, or talus slide can be detected in
several ways. The white, mineral-like incrustation formed by the urine
of many generations of wood rats is usually conspicuous on rocks near
their homes. A strong musky odor pervades the atmosphere in every
habitat occupied by wood rats. Wood rat feces, consisting of hard, black
cylinders one-half inch long and three-sixteenths of an inch in
diameter, are invariably found scattered on stones or exposed areas of

  [Illustration: FIG. 108. Distribution of the bushy-tailed wood rat in
  Washington. A. _Neotoma cinerea occidentalis._ B. _Neotoma cinerea

The food of the bushy-tailed wood rat is varied and includes insects and
vegetation. A captive specimen that escaped in the University of
Washington Zoölogical Laboratories killed and ate a number of lizards.

Embryos found from April 18 to June 14 varied in number from 3 to 5.

=Neotoma cinerea occidentalis= Baird

      _Neotoma occidentalis_ Baird, Proc. Acad. Nat. Sci. Philadelphia,
      p. 335, 1855.

      _Neotoma cinerea occidentalis_ Merriam, N. Amer. Fauna, 5:58, July
      30, 1891.

      _Type._--Obtained at Shoalwater Bay (Willapa Bay), Pacific County,
      Washington, by J. G. Cooper, in June, 1854; type in United States
      National Museum.

      _Racial characters._--Size large; color of upper parts cinnamon

      _Measurements._--A male and a female from Glade Creek, 1/2 mile
      north of the Columbia River, Benton County, measure, respectively:
      total length 425, 395; length of tail 181, 176; hind foot 50, 44;
      ear 32, 31; weight 532.5, 330.0 grams.

      _Distribution._--Throughout the state except in the Blue

_Remarks._--Specimens from the Olympic Mountains and western Washington
are slightly duller-colored than specimens from eastern Washington and
young from western Washington are more bluish, less gray, than young
individuals from the Columbian Plateau. The specimens from the Columbian
Plateau show some resemblance to _alticola_. Specimens from northeastern
Washington are more like typical _occidentalis_.

=Neotoma cinerea alticola= Hooper

      _Neotoma cinerca alticola_ Hooper, Univ. California Publ. Zoöl.,
      42:409, May 17, 1940.

      _Type._--Obtained at Parker Creek, 5,500 ft., Warner Mountains,
      Modoc County, California, by N. B. Stern on June 22, 1916; type in
      Museum of Vertebrate Zoölogy.

      _Racial characters._--Similar to _Neotoma c. occidentalis_ but
      slightly paler, with paler hind feet and narrower sphenopalatine

      _Measurements._--A female from Stay-a-while Spring, Columbia
      County, measures: total length 387; length of tail 171; hind foot
      43; ear 33; weight 330 grams.

      _Distribution._--Southeastern Washington, south of the Snake

_Remarks._--The only adult specimen available from the Blue Mountains
shows well the characters attributed to _alticola_.

=Synaptomys borealis wrangeli= Merriam

Northern lemming mouse

      _Synaptomys (Mictomys) wrangeli_ Merriam, Proc. Biol. Soc.
      Washington, 10:63, March 19, 1896.

      _Synaptomys (Mictomys) truei_ Merriam, Proc. Biol. Soc.
      Washington, 10:62, March 19, 1896 (type from Skagit Valley, Skagit
      County, Washington).

      _Synaptomys borealis wrangeli_ Howell, N. Amer. Fauna, 50:26, June
      30, 1927.

      _Type._--Obtained at Wrangel, Alexander Archipelago, Alaska, by C.
      P. Streator on September 6, 1895; type in United States National

      _Racial character._--Dark color.

      _Distribution._--The northwestern Cascades, from Mt. Baker

_Remarks._--The relationship of this race to _artemisiae_ remains
to be worked out.

  [Illustration: FIG. 109. Distribution of the northern lemming mouse in
  Washington. A. _Synaptomys borealis wrangeli._ B. Possible range of
  _Synaptomys borealis artemisiae_.]

_Description._--Lemming mice may be recognized by their short, thick
bodies, slightly larger than the bodies of the common house mouse
(_Mus_); short tails, which are less than 20 per cent of their total
length; small, inconspicuous ears; and grooved upper incisors.

Mice of the genus _Synaptomys_ range over much of boreal North America.
Two species and eleven races are recognized by Howell (1927B: 9).
_Synaptomys borealis_ ranges westward across Canada from Labrador to the
Pacific Coast and from Alaska south to Washington. The lemming mice are
terrestrial and inhabit runways similar to those of meadow mice
(_Microtus_). They are alpine in distribution. Shaw (1930: 7-10) found
them among typical annual herbs in a cold, boggy mountain valley. Other
than this, nothing is known of the species in Washington. Indeed little
is known of the life history of any member of the genus.

The lemming mouse possesses four pairs of mammae, two pectoral and two
inguinal. It is the most primitive of the microtines. It seems to be of
boreal origin but is not known from the Old World. It was evidently
forced southward by the Pleistocene glaciers into Washington and is now
retreating northward in the wake of the ice. The only specimens recorded
from Washington are from the northern border of the state.

=Phenacomys intermedius= Merriam

Heather vole

_Description._--Heather voles are heavy-bodied, short-legged mice,
closely similar in general appearance to other microtines that occur in
Washington. Their dull, brownish upper parts, lacking a distinct reddish
dorsal stripe, separate them from the red-backed mice (_Clethrionomys_)
and the lack of grooves on their upper incisors separates them from
_Synaptomys_. Their relatively short tail (30 per cent or less of their
total length) separates them from most meadow mice (_Microtus_). There
are no external characters which serve to separate them from all species
of _Microtus_ and the teeth must be examined certainly to identify
_Phenacomys_. In _Microtus_ the angles between the cusps of the inner
and outer sides of the lower molars are of approximately equal depth,
but in _Phenacomys_ the angles on the inner side of the lower molars are
at least twice as deep as those of the outer side of the jaw. In
addition the molars of _Phenacomys_ are rooted while those of _Microtus_
are not.

The heather voles and their relatives are primitive microtines that
range through boreal North America including the higher parts of the
Rocky Mountains and the cool area along the Pacific Coast. Several
species are included in the genus: an arctic form (_ungava_), a lowland
Pacific Coastal species (_albipes_), two tree-inhabiting species
(_longicaudus_ and _silvicola_) and a mountain species (_intermedius_).
Thus far only _intermedius_ has been found in Washington but further
search may reveal that one or more of the arboreal species and possibly
_albipes_ are also present.

Heather voles are alpine animals, seldom occurring below the Hudsonian
Life-zone. They are rare as compared with most microtines, and are
seldom taken in traps, even by the experienced collector of small
mammals. Evidence of their presence is most often found in the dense
meadows of heather (_Cassiope_ and _Phyllodoce_) high on the mountains.
Here their feces, runways, and old winter nests are not uncommon, and
heather voles may be far more abundant than trapping records indicate.
They are not restricted to the vicinity of heather, however, for one of
my specimens was taken in a marshy willow thicket near Stevens Pass in
King County, one in a patch of skunk-cabbage in a grassy glacial cirque
near Tomyhoi Lake in Whatcom County, and one at the edge of a snowbank
on a steep hillside at Deer Park, Clallam County. In each of these
places, however, there were heather meadows within a thousand yards.
Shaw (1924A: 12-15) also found the heather mouse on "comparatively dry
hill slopes" and reports that it "rather characteristically occurs in
such log-tangled areas as former fire zones."

  [Illustration: FIG. 110. Distribution of the heather vole in
  Washington. A. _Phenacomys intermedius oramontis._ B. _Phenacomys
  intermedius intermedius._]

The winter nests of _Phenacomys intermedius_ are placed on the surface
of the ground and in the season when occupied are deeply buried under
the snow. They are spherical or oval in shape, six to eight inches in
diameter and with a side entrance. One examined by Shaw was "formed of
white heather twigs, the interstices being filled with leaves of
huckleberry." It was "lined with finely shredded grass." All those
examined by me were composed of grass and lichens.

The principal foods of the heather vole seem to be white heather
(_Cassiope mertensiana_) and huckleberry (_Vaccinium_). Red heather
(_Phyllodoce empetriformis_) is not eaten. Shaw also lists beargrass
(_Xerophyllum tenax_) and lousewort (_Pedicularis_) as eaten.

A burrow excavated and figured by Shaw, was slightly more than a yard
long and contained four young in a nest situated but a few inches from
one of the three entrances. The nest was made of grass and moss. A
female taken at Tye, King County, contained 4 embryos on May 28, 1939.
One taken at Pass Creek Pass, Pend Oreille County, contained 3 small
embryos on June 14, 1938.

=Phenacomys intermedius intermedius= Merriam

      _Phenacomys intermedius_ Merriam, N. Amer. Fauna, 2:32, October 30,

      _Phenacomys intermedius intermedius_ Howell, N. Amer. Fauna,
      48:15, October 12, 1926.

      _Type._--Obtained 20 miles north-northwest of Kamloops, 5,500 ft.
      elevation, British Columbia, by G. M. Dawson on October 2, 1888;
      type in United States National Museum.

      _Racial characters._--Size small; color of upper parts brownish
      gray; skull small.

      _Measurements._--A female from Pass Creek Pass, Pend Oreille
      County, measured: total length 127; length of tail 29; hind foot
      17; weight 25 grams.

      _Distribution._--Known only from extreme northeastern Washington
      at Pass Creek Pass (W.W.D.) and the Blue Mountains.

=Phenacomys intermedius oramontis= Rhoads

      _Phenacomys oramontis_ Rhoads, Amer. Nat., 29:941, October, 1895.

      _Phenacomys olympicus_ Elliot, Field Columb. Mus. Publ. 30, zoöl.
      ser., 1:225, February 1, 1899 (type from Happy Lake, 5,000 ft.,
      Clallam County, Washington).

      _Microtus (Lagurus) pumilus_ Elliot, Field Columb. Mus. Publ. 30,
      zoöl. ser., 1:226, February 1, 1899 (type from Happy Lake, 5,000
      ft., Clallam County, Washington).

      _Phenacomys intermedius olympicus_ Howell, N. Amer. Fauna, 48:21,
      October 12, 1926.

      _Phenacomys intermedius oramontis_ Anderson, Canadian Field Nat.,
      56:59, June 8, 1942.

      _Type._--Obtained on Church Mountain, 6,000 ft., Mt. Baker Range,
      British Columbia, just north of international boundary, by A. C.
      Brooks on August 6, 1895; type in Philadelphia Academy of Natural

      _Racial characters._--Similar to _intermedius_ but darker, browner
      above and deeper gray beneath; skull large and heavy.

      _Measurements._--A female from Tye, King County, measures: total
      length 152; length of tail 40; hind foot 19; ear 12. A female from
      Tomyhoi Lake, Whatcom County, measures: 170; 45; 20; 15.

      _Distribution._--The Cascade and Olympic mountains, from Tomyhoi
      Lake (W. W. D.) on the north, to Mt. Adams (Taylor and Shaw,
      1929:23) on the south.

=Clethrionomys gapperi= (Vigors)

Gapper red-backed mouse

_Description._--Red-backed mice are heavy-bodied, short-tailed and
short-legged mice, similar in general appearance to the meadow mice
(_Microtus_). Unlike the meadow mice, they possess rooted molars, a
primitive character. They do not have grooved incisors, like
_Synaptomys_, or the great difference in the depth of the angles of the
lower molars that characterizes both _Synaptomys_ and _Phenacomys_.
Externally _Clethrionomys_ may be recognized by the broad red dorsal
area from which they obtain their common name. Their sides are grayish
or dusky and the undersides buffy white.

The red-backed mice, including more than one species, range over the
boreal parts of the Old and New World. In America they are found in most
of Alaska, Canada and the northern United States, and extend southward
in the Rocky Mountains and along the Pacific Coast. The one species
_Clethrionomys gapperi_ ranges across southern Canada from the Atlantic
to the Pacific, extending southward in forested areas to New Mexico.
They usually live in clearings in the forest. In the Pend Oreille
Mountains _saturatus_ was common in damp, mossy talus slides, along with
_Microtus longicaudus_ and _Phenacomys intermedius_. The _Clethrionomys_
outnumbered the two latter species combined by 25 to 1. In nearby
forest, red-backed mice were scarce but no other microtines were found.
In the Kettle River Mountains a week later, red-backed mice were rare. A
single specimen was taken in a damp place in the forest; none was found
in talus slides. Near Stevens Pass, King County, in the Cascades,
_cascadensis_ was taken in equal numbers in talus slides and under logs
in the forest. Near Dewey Lake, Yakima County, in the Cascades, I took
them only in an extensive grassy meadow. In the Blue Mountains I found
_idahoensis_ in the dense chaparral, far from forests.

Unlike _Microtus_, red-backed mice do not make runways, although they
sometimes follow the runways of other mammals.

Taylor (1920B: 92) found red-backed mice breeding on Mount Rainier from
early July to the middle of September. One female gave birth to four
young in a nest in his duffle bag.

  [Illustration: FIG. 111. Distribution of the Gapper and California
  red-backed mice in Washington. A. _Clethrionomys gapperi saturatus._
  B. _Clethrionomys gapperi idahoensis._ C. _Clethrionomys gapperi
  nivarius._ D. _Clethrionomys gapperi cascadensis._ E. _Clethrionomys
  californicus occidentalis._]

=Clethrionomys gapperi saturatus= (Rhoads)

      _Evotomys gapperi saturatus_ Rhoads, Proc. Acad. Nat. Sci.
      Philadelphia, p. 284, October 23, 1894.

      [_Clethrionomys gapperi_] _saturatus_ Whitlow and Hall, Univ.
      California Publ. Zoöl., 40:265, September 30, 1933.

      _Type._--Obtained at Nelson, British Columbia, by S. N. Rhoads on
      August 27, 1892; type in Philadelphia Academy of Sciences.

      _Racial characters._--Size small, about 140 mm. in total length;
      tail short, about one-third of total length; ears large,
      projecting above fur; color of head and sides gray tinged with
      yellowish; back with distinct red stripe; underparts whitish
      tinged with buff.

      _Measurements._--Five males and 10 females from Pass Creek Pass,
      Pend Oreille County, average: total length 147; length of tail 43;
      hind foot 18.3; ear 14.

      _Distribution._--Northeastern Washington from Sherman Creek Pass
      (W. W. D.) on the west to Pass Creek Pass on the east.

=Clethrionomys gapperi idahoensis= (Merriam)

      _Evotomys idahoensis_ Merriam, N. Amer. Fauna, 5:66, July 30, 1891.

      _Clethrionomys gapperi idahoensis_ Whitlow and Hall, Univ.
      California Publ. Zoöl., 40:265, September 30, 1933.

      _Type._--Obtained at Sawtooth (Alturas) Lake, 7,200 ft., Blaine
      County, Idaho, by C. H. Merriam and V. Bailey, on October 4, 1890;
      type in United States National Museum.

      _Racial characters._--Similar to _saturatus_ but brain case longer
      and narrower and dorsal area more reddish.

      _Measurements._--Seven males and 6 females from the Blue Mountains
      average, respectively: total length 138, 142; length of tail 41,
      41; hind foot 18.7, 19.0; ear 13.6, 13.5; weight 20.5, 22.7 grams.

      _Distribution._--The Blue Mountains of southeastern Washington.

=Clethrionomys gapperi cascadensis= Booth

      _Evotomys gapperi saturatus_ Taylor and Shaw, Occ. Pap. Chas. R.
      Conner Mus., no. 2:23, December, 1929.

      _Clethrionomys gapperi cascadensis_ Booth, Murrelet, 26:27, August
      10, 1945.

      _Type._--Obtained 2 miles south of Blewett Pass, 3,000 ft.
      elevation, Kittitas County, Washington, by G. G. Cantwell, on
      October 30, 1921; type in United States National Museum.

      _Racial characters._--Dark and dull with underparts dull buffy.

      _Measurements._--Ten adults from the Cascade Mountains average
      (Booth, 1945: 27): total length 162; length of tail 50; hind foot
      19; ear 14.

      _Distribution._--The Cascade Mountains, according to Booth (_loc.
      cit._, p. 28), from Hannegan Pass south to Mount Adams. Marginal
      localities listed are: Swamp Creek, Glacier Peak, McKenna, and Mt.
      St. Helens. These, and other localities listed by Booth are not
      plotted on the accompanying distribution map (fig. 111).

=Clethrionomys gapperi nivarius= (Bailey)

      _Evotomys nivarius_ Bailey, Proc. Biol. Soc. Washington, 11:136,
      May 13, 1897.

      _Evotomys gapperi nivarius_ Taylor and Shaw, Occ. Papers Chas. R.
      Conner Mus., no. 2:23, 1929.

      _Clethrionomys nivarius_ Svihla, Murrelet, 12:54, May, 1931.

      _Type._--Obtained on the northwest slope of Mt. Ellinor, 4,000 ft.
      elevation, Mason County, Washington, by C. P. Streator, on July 9,
      1894; type in United States National Museum.

      _Racial characters._--Similar to _saturatus_ but paler throughout.

      _Measurements._--Two males and a female from 1 mile northwest of
      Lake Cushman, Mason County, average: total length 140; length of
      tail 42; hind foot 18; ear 13.

      _Distribution._--The Olympic Mountains, from Sol Duc Hot Springs
      (W.S.M.) south and west at least to Staircase, on Lake Cushman

=Clethrionomys californicus occidentalis= (Merriam)

California red-backed mouse

      _Evotomys occidentalis_ Merriam, N. Amer. Fauna, 4:25, October 8,

      _Evotomys pygmaeus_ Rhoads, Proc. Acad. Nat. Sci. Philadelphia, p.
      284, October 23, 1894 (type from mouth of Nisqually River, Pierce
      County, Washington).

      _Evotomys gapperi occidentalis_ Taylor and Shaw, Occ. Papers Chas.
      R. Conner Mus., no. 2:23, 1929.

      _Clethrionomys occidentalis_ Hall, Murrelet, 13:79, September,

      _Type._--Obtained at Aberdeen, Grays Harbor County, Washington, by
      T. S. Palmer, on August 16, 1889; type in United States National

      _Measurements._--Ten males and 10 females average: total length
      137; length of tail 45; hind foot 18; ear 12.3.

      _Distribution._--The lowlands of western Washington, east at least
      to Cottage Lake (W. W. D.).

_Remarks._--Specimens from the type locality of _occidentalis_ and other
places along the ocean coast are larger and brighter in color than
specimens from farther inland, but the difference appears to me to be
insufficient to warrant subspecific separation of the two lots.

_Clethrionomys californicus_ resembles _Clethrionomys gapperi_ but is
darker and duller in color with the red dorsal area more obscured and
forming less of a stripe. _C. californicus_ is found in the forested
lowlands of western Washington, Oregon, and northern California, and in
the Cascades of Oregon. In Washington, it is confined entirely to the
forest where it is trapped under logs and on the layer of dead needles
at the bases of conifers. Mice of this species were numerous in the
forest along the Pacific Coast on the Long Beach Peninsula and at
Aberdeen, Grays Harbor County. At Lost Lake Prairie, Mason County, at
the southeastern base of the Olympic Mountains, they were rather scarce,
but were the only mammals taken in the deep woods. Near Shelton, Mason
County, at the eastern edge of the Olympic Mountains, a number of
specimens were taken along with twice as many _Peromyscus maniculatus_.
At Cottage Lake, King County, near Seattle, they were rare, comprising
about two per cent of the mammals taken in two weeks' trapping.

Almost nothing was learned of the habits of these mice. They seem to be
rigidly restricted to a habitat where few plants other than trees grow.
The stomachs examined contained pasty masses of finely chewed white
vegetation with occasional gray particles that might have been bits of
lichens. The mice do not make runways like those of _Microtus_.

=Microtus pennsylvanicus= (Ord)

Pennsylvanian meadow mouse

_Description._--The upper parts are dark blackish brown and the
underparts grayish or whitish. The tail comprises about a fourth of the
total length and the foot does not exceed 21 mm. These two features
separate it from most other species of meadow mice. From _Microtus
oregoni_ it may be separated by its larger size, blackish color and
well-developed eyes. From _Microtus montanus_ it differs in being
larger, darker, and in having a closed posterior loop on the innerside
of the second upper molar.

  [Illustration: FIG. 112. Pennsylvania meadow mouse (_Microtus
  pennsylvanicus kincaidi_), from ten miles south of Moses Lake,
  Washington, April 28, 1940. (Fish and Wildlife Service photo by Victor
  B. Scheffer, No. 937.)]

_Microtus pennsylvanicus_ is the most common meadow mouse of the eastern
United States. It ranges westward through Canada and southward in the
Rocky Mountains. A number of races occur in this extensive range. In
northeastern Washington the eastern meadow mouse is locally common,
being confined to marshes and damp meadows. Well-used runways are made
through the dense vegetation and piles of feces and blades of grass are
deposited therein. A number of broods are raised in a season, for young
of many different ages are taken together in midsummer.

On the Columbian Plateau this mouse lives in marshy areas about lakes
and potholes. Narrow trails are constructed through the dense
vegetation. It burrows in damp earth and some occupied burrows are
half-full of seepage water. Cuttings found in runways show that the mice
feed on a variety of plants, including sedge (_Carex_). In the Moses
Lake area the breeding season begins in March. Embryos found March 24,
1940, varied from 4 to 7 in number. This species is cyclically abundant.

  [Illustration: FIG. 113. Distribution of the Pennsylvania meadow mouse
  in Washington. A. _Microtus pennsylvanicus funebris._ B. _Microtus
  pennsylvanicus kincaidi._]

=Microtus pennsylvanicus funebris= Dale

      _Microtus pennsylvanicus modestus_ Bailey, N. Amer. Fauna, 17:20,
      June 6, 1900.

      _Microtus drummondii_ Bailey, N. Amer. Fauna, 17:22, June 6, 1900.

      _Microtus pennsylvanicus funebris_ Dale, Jour. Mamm., 21:338,
      August 13, 1940.

      _Type._--Obtained at Coldstream. 1,450 ft., 3-1/2 miles southeast
      of Vernon. British Columbia, by T. P. Maslin. Jr., on August 2,
      1937; type in Museum of Vertebrate Zoölogy.

      _Racial characters._--Size medium; color reddish brown; fur short,
      harsh; skull small and narrow.

      _Measurements._--Three males from Newport, Pend Oreille County,
      average: total length 165; length of tail 40; hind foot 20; ear

      _Distribution._--Northeastern Washington, west to Conconully
      (Taylor and Shaw, 1929: 24) and east to Newport (W. W. D.).

_Remarks._--Specimens from northeastern Washington are larger and
darker, more reddish and less gray, than _drummondii_. They are larger,
more reddish, and have relatively narrower skulls, than _modestus_ from
Colorado and Idaho. They closely resemble _funebris_ from south-central
British Columbia, and are best referred to that race. Rand (1943: 123)
considers _funebris_ a synonym of _modestus_ but I feel is incorrect in
doing so.

=Microtus pennsylvanicus kincaidi= Dalquest

      _Microtus montanus kincaidi_ Dalquest, Proc. Biol. Soc. Washington,
      54:145, September 30, 1941.

      _Type._--Obtained at The Potholes, 10 miles south of Moses Lake,
      Grant County, Washington, by W. W. Dalquest on March 24, 1940; type
      in Museum of Vertebrate Zoölogy.

      _Racial characters._--Large size; dark blackish color; long fur;
      wide, angular skull.

      _Measurements._--Six male and 12 female topotypes average,
      respectively: total length 176.6, 168.0; length of tail 45.1,
      43.8; hind foot 20.8, 20.3; ear 14.0, 13.4.

      _Distribution._--The Columbian Plateau, in the Grand Coulee area.

=Microtus montanus= (Peale)

Montane meadow mouse

_Description._--Montane meadow mice have short tails, flaring zygomatic
arches, and heavily ridged skulls. They are small, about 6-1/2 inches
long, and buffy-gray in color. Their short tails, less than one-third
the length of head and body, and gray color separate them from all other
Washington meadow mice.

Montane meadow mice are widely distributed in the southwestern United
States, where numerous races are found. They occupy damp meadows and
marshes in the arid subdivision of the Transition Life-zone of eastern
Washington. They seem to require deep, dense cover of grasses, reeds, or
sedges, near water. They are usually associated with harvest mice
(_Reithrodontomys megalotis_), wandering shrews (_Sorex vagrans
monticola_), and muskrats (_Ondatra zibethica_). Hawks and owls are
their principal enemies, but predatory mammals and snakes probably kill

Montane meadow mice evidently are both diurnal and nocturnal; specimens
were taken in the daytime as well as at night, and captives were active
periodically day and night. Captives were extremely quick in their
movements. If disturbed they sat up on their haunches and chattered
indignantly. It was almost impossible to handle them without being

The trails of montane meadow mice are narrower than those of other
meadow mice, and appear to be used the year around. Burrows are
numerous where the mice are plentiful. A heap of damp earth marks the
entrance to each burrow. One nest of _M. m. canescens_ near Lake Chelan,
Chelan County, was under a log on a small knoll in a marsh. The nest was
round, about 8 inches in diameter, and was composed of cattails,
grasses, and moss.

  [Illustration: FIG. 114. Distribution of the montane and Townsend
  meadow mice in Washington. A. _Microtus montanus nanus._ B. _Microtus
  montanus canescens._ C. _Microtus townsendii townsendii._ D. _Microtus
  townsendii pugeti._]

_Microtus montanus nanus_ (Merriam)

      _Arvicola (Mynomes) nanus_ Merriam, N. Amer. Fauna, 5:63, July 30,

      _Microtus nanus_ Miller, Proc. Biol. Soc. Washington, 11:67, April
      21, 1897.

      _Microtus nanus nanus_ Miller, Bull. U. S. Nat. Mus., 128:409,
      April 29, 1924.

      _Microtus montanus nanus_ Hall, Proc. Biol. Soc. Washington,
      51:133, August 23, 1938.

      _Type._--Obtained in the Pahsimeroi Mountains, 9,300 ft., Custer
      County, Idaho, by C. H. Merriam and V. Bailey on September 16,
      1890; type in United States National Museum.

      _Racial characters._--Size medium; color of upper parts brownish
      gray; sides paler; underparts grayish white.

      _Measurements._--A large male from Prescott, Walla Walla County,
      measures: total length 168; length of tail 48; hind foot 20. A
      female from Prescott, Walla Walla County, and a female from
      Pullman, Whitman County, average: 133; 31; 18.5.

      _Distribution._--Southeastern Washington, westward as far as 5
      miles east of Wallula (M.V.Z.).

=Microtus montanus canescens= Bailey

      _Microtus nanus canescens_ Bailey, Proc. Biol. Soc. Washington,
      12:87, April 30, 1898.

      _Microtus montanus canescens_ Hall, Proc. Biol. Soc. Washington,
      51:133, August 23, 1938.

      _Type._--Obtained at Conconully, Okanogan County, Washington, by J.
      A. Loring, on September 12, 1897; type in United States National

      _Racial characters._--Size small; color of upper parts pale
      brownish gray; sides yellowish gray. This race differs from
      _Microtus m. nanus_ in generally paler, less brownish coloration.
      It does not differ from _nanus_ in any distinctive cranial

      _Measurements._--Two male and 2 female topotypes average,
      respectively: total length 151, 143; length of tail 35, 33.5; hind
      foot, 19.5, 19.5. A series of 8 males and 6 females from Selah,
      Yakima County, average, respectively: total length 151.7, 150.5;
      length of tail 41.5, 40.1; hind foot 19.7, 18.8; ear 12.8, 13.0;
      weight 47.2, 36.8 grams.

      _Distribution._--The eastern foothills of the Cascade Mountains
      from the British Columbian boundary south, probably to the
      Columbia River. Recorded east to Benton City (Taylor and Shaw,
      1929: 24).

_Remarks._--A large series from Selah, Yakima County, is somewhat
intermediate between _nanus_ and _canescens_. These specimens from south
of the Wenatchee Mountains are, however, more like _canescens_ in color
than they are like _nanus_.

Bailey (1900: 32) records a specimen of this race from North Yakima as
_Microtus canicaudus_.

=Microtus townsendii= (Bachman)

Townsend meadow mouse

_Description._--The Townsend meadow mouse is a large-bodied, long-furred
mouse with a tail of moderate length. Head and body measure about 8
inches, the tail about 2-1/2 inches. The legs are short and the ears
scarcely project through the fur on the head. In summer the color is
dark reddish-brown. The winter color is dark brownish-black. Underparts
are paler, more grayish brown, than are the upper parts. The tail is
sparsely haired.

Townsend meadow mice occur west of the Cascade Mountains in California,
Washington, Oregon, British Columbia, and on some British Columbian and
Washington islands. The habitats of the two races of the Townsend meadow
mice found in Washington are wholly in the humid subdivision of the
Transition Life-zone but vary somewhat in nature. The race _townsendii_
lives in marshes or damp meadows, under cover of deep, rank vegetation.
These mice avoid forested areas or dry brush, but sometimes occur in dry
grass when it is deep enough to conceal them from enemies. The race
_pugeti_ has been recorded from meadows, salt marshes, driftwood strewn
on sea-beaches, areas of sparse, dry grass, and piles of rocks.

Townsend meadow mice are as diurnal as they are nocturnal; specimens
have been trapped at almost all hours of the day and night. Their
ordinary method of traveling is a slow run. When startled they make a
dash for the nearest cover, into which they dive headlong. They make
considerable noise while moving about and often may be heard from
several feet away. Captives in the laboratory seemed rather dull and
stupid as compared with other meadow mice.

  [Illustration: FIG. 115. Runways of Townsend meadow mice (_Microtus
  townsendii_) worn to grooves in the damp soil at Seattle; May 18,
  1938; dense cover of cattails has been burned off (W. W. Dalquest

In the fall, winter, and early spring the Townsend meadow mice keep to
their runways. These runways are used by successive generations of mice,
and often are worn into ditches several inches deep. In the late spring
and summer, when the grass and other vegetation in their habitat is
tall and rank, offering complete concealment, the runways are abandoned
and cuttings of rejected food are found scattered over the surface of
the ground.

The food of the Townsend meadow mice includes the succulent leaves and
stems of many grasses and annuals. Near Seattle the staple summer food
is the velvet grass (_Holcus lanatus_), although many other plants,
including the horsetail (_Equisetum arvense_), are eaten. In winter the
common cattail is eaten. Couch (1925: 200) found caches of the roots of
mint (_Mentha canadensis_) stored by this species. As much as 14 quarts
was found in a single cache.

The cup-shaped nests of Townsend meadow mice near Seattle were below
ground. Embryos were found from May 4 to May 20 and varied in number
from 5 to 8 with a mean of 7. In the San Juan Islands the nests of _M.
t. pugeti_ were under driftwood.

=Microtus townsendii townsendii= (Bachman)

      _Arvicola townsendii_ Bachman, Jour. Acad. Nat. Sci. Philadelphia,
      8 (pt. 1):60, 1839.

      _Arvicola occidentalis_ Peale, U. S. Expl. Exped., Mammalogy, p.
      45, 1848 (type from Puget Sound).

      _M[icrotus]. townsendi_ Miller, N. Amer. Fauna, 12:66, July 23,

      _Microtus townsendii townsendii_ Svihla and Svihla, Murrelet,
      14:40, May, 1933.

      _Type._--Obtained on the Columbia River (probably on or near Sauvie
      Island, Multnomah County, Oregon); type in Philadelphia Academy of
      Natural Sciences.

      _Racial characters._--Size large; skull narrow in interorbital

      _Measurements._--Eight males and 5 females from Clark and Pacific
      counties, southwestern Washington, average, respectively: total
      length 208.0, 208.4; length of tail 65.7, 66.0; hind foot 26.5,
      25.4; ear 15.0, 14.6; weight 80.8, 76.7 grams. Thirteen males and
      9 females from Seattle average, respectively: 211.8, 209.0; 71.2,
      68.6; 25.7, 26.0; 15.8, 15.4.

      _Distribution._--The lowlands of western Washington from
      Bellingham (J.M.E.) south to Puget Island (V.B.S.).

=Microtus townsendii pugeti= Dalquest

      _Microtus townsendii pugeti_ Dalquest, Murrelet, 21:7, April 1,

      _Type._--Obtained at Neck Point, northwest corner of Shaw Island,
      San Juan County, Washington, by D. H. Johnson, on July 10, 1938;
      type in Museum of Vertebrate Zoölogy.

      _Racial characters._--Size small; skull wide in interorbital
      region, averaging about 4.0 mm. (3.8-4.2); basi-sphenoid truncate
      posteriorly; upper incisors strongly curved.

      _Measurements._--Two males and 6 females average: total length
      182.6; length of tail 50.3; hind foot 23.2; ear 15.0.

      _Distribution._--Found only on the San Juan Islands, San Juan and
      Skagit counties.

_Remarks._--The islands occupied by this race of mouse were heavily
glaciated by the last continental glacier (Vashon). Mice of the species
_townsendii_ apparently migrated to the islands early in the Recent era,
and under isolation developed the differences which now separate them
from the mainland population.

=Microtus longicaudus= (Merriam)

Long-tailed meadow mouse

_Description._--The several races of this species vary from small to
large in size. Their bodies are relatively longer and slimmer than those
of the other meadow mice that occur in Washington. Their most
distinctive feature is the long tail, only slightly shorter than the
head and body. Their fur is rather coarse. The color varies from grayish
brown to dull reddish brown with a brighter brown dorsal stripe. The
tail is bicolor; black or dark brown above, yellowish below. The
underparts are whitish gray.

  [Illustration: FIG. 116. Distribution of the long-tailed meadow mouse
  in Washington. A. _Microtus longicaudus macrurus._ B. _Microtus
  longicaudus halli._]

Long-tailed meadow mice are widely distributed over the western United
States, Canada, and Alaska. In Washington the long-tailed meadow mouse
has been taken in many habitats. One specimen was taken along a small,
temporary stream through sagebrush in the Upper Sonoran Life-zone.
Others were found in marshes and near water in the arid subdivision of
the Transition Life-zone. In the humid subdivision of the Transition
Life-zone they are not uncommon in damp areas along the ocean coast, but
are rather rare in dry, grassy habitats. In the Canadian and Hudsonian
life-zones they are fairly common in forest-free, grassy places, being
most abundant near talus slides. Altitudinally they range from sea level
to 6,000 feet.

Long-tailed meadow mice are rather rare. Several were taken in the
daytime at Round Top Mountain, Pend Oreille County, and two others were
taken in daytime two miles south of Tenino, Thurston County. However
most of the specimens were caught at night.

Long-tailed meadow mice do not, at least ordinarily, make trails as do
other meadow mice. Specimens are usually taken unexpectedly, and
intensive trapping in the area where a specimen or two is taken rarely
yields additional individuals.

Two specimens taken in the Cascade Mountains in September were pregnant;
one contained two embryos and the other four.

=Microtus longicaudus halli= Ellerman

      _Microtus mordax angustus_ Hall, Univ. California Publ. Zoöl.,
      37:13, April 10, 1931 (not of Thomas, 1908).

      _Microtus longicaudus angustus_ Goldman, Jour. Mamm., 19:491,
      November 14, 1938.

      _Microtus mordax halli_ Ellerman, Fam. and Genera of Living
      Rodents, British Mus. Nat. Hist., 2:603, March 21, 1941 (new name
      for _Microtus mordax angustus_ Hall).

      _Type._--Obtained at Godman Springs, 5,700 ft., Blue Mountains,
      Columbia County, Washington, by S. H. Lyman, on September 1, 1927;
      type in Museum of Vertebrate Zoölogy.

      _Racial characters._--Size small; color of sides pale grayish
      brown; brown dorsal stripe conspicuous; tail relatively short.

      _Measurements._--Four males and 5 females from eastern Washington
      average, respectively: total length 164.8, 166.6; length of tail
      55.8, 55.8; hind foot 22.0, 21.4.

      _Distribution._--From the Blue Mountains of southeastern
      Washington north, along the eastern edge of the state, to British
      Columbia, and thence west, north of the Columbia River, to the
      Cascades, and south along the eastern edge of the Cascades to the
      Wenatchee Mountains. Marginal occurrences are: Pasayten River
      (Taylor and Shaw, 1929: 24), Hart Lake (Taylor and Shaw, 1929:
      24), Blewett Pass (W.W.D.), Boulder Cave (W.W.D.), and Satus Pass

_Remarks._--Four specimens from Satus Pass, Klickitat County, are
somewhat intermediate between this race and _macrurus_, and indicate
that _halli_ crossed the Columbia River when an alpine meadow land
extended from the Simcoe Anticline to southeastern Washington.

One specimen from Selah, Yakima County, is colored somewhat like
_halli_, and is smaller than _macrurus_. Possibly the _halli_ type of
meadow mouse spread northward to the Yakima Valley. This specimen was
taken in the Upper Sonoran Life-zone. A specimen from the Arid
Transition Zone at Naches, Yakima County (taken in almost Canadian
Life-zone habitat), is like _macrurus-halli_ intergrades from the higher
Cascade Mountains.

=Microtus longicaudus macrurus= Merriam

      _Microtus macrurus_ Merriam, Proc. Acad. Nat. Sci. Philadelphia, p.
      353, October 4, 1898.

      _Microtus mordax macrurus_ Dice, Murrelet, 13:49, May, 1932.

      _Microtus longicaudus macrurus_ Goldman, Jour. Mamm., 19:491,
      November 14, 1938.

      _Type._--Obtained at Lake Cushman, Mason County, Washington, by C.
      P. Streator on June 26, 1894; type in United States National

      _Racial characters._--Size large; tail relatively long, almost as
      long as head and body; color of upper parts dull brown; dorsal
      stripe obscure; underparts gray.

      _Measurements._--Two males and 4 females from the Olympic
      Mountains and the coast of Washington, average, respectively:
      total length 212.5, 220.5; length of tail 82.5, 86.8; hind foot
      25.5, 24.7; ear 15.0, 13.7.

      _Distribution._--The Olympic Mountains, the coast region of
      western Washington, and the Cascade Mountains, save in the
      southeastern part. Marginal occurrences are: Sauk (Taylor and
      Shaw, 1929: 24), Tye (W.W.D.), Naches River (W.W.D.), and Wind
      River (W.W.D.).

_Remarks._--Specimens from the Cascade Mountains are referred to
_macrurus_ but are intermediate between that race and _halli_.

=Microtus richardsoni= (De Kay)

Water rat

_Description._--This is the largest meadow mouse found in Washington. It
is too large to be called a mouse and the term water rat, applied to it
by Merriam, suits it well. It is the only Washington microtine measuring
more than 8 inches in total length. The upper parts are dark, reddish
brown in color; the underparts are grayish brown. The tail comprises
about one-third of the total length.

This species of meadow mouse ranges from Canada south to Colorado in the
Rocky Mountains and in the Cascades south into Oregon. The water rats
are strictly alpine animals, occurring about streams, marshes and damp
meadows. In the Cascade Mountains they are most common in the glacial
cirques where tiny streams flow through grassy meadows to plunge over
the lip of the cirque on a rocky course of cascades to the valley below.
Here their broad trails occur along the stream banks, commonly entering
the water where it is swift. These trails are well-worn roads, usually
about four inches wide but often wider. The burrows of the water rat are
about three inches in diameter and are constructed with no effort at
concealment; large mounds of earth mark their entrances. Freshly dug
burrows are so abundant that it seems likely more are dug than are
actually inhabited. Burrows are often dug beneath rocks.

The water rat is mainly nocturnal but not uncommonly is seen in the
daytime. Twice I cornered a water rat away from its burrow and each time
it escaped by swimming. The rats swam with great speed but with much
splashing. One dived under the surface of a small pool and disappeared.
Stones along the bank were pulled out until the rat was captured in a
small chamber at the end of a burrow. The burrow entered the bank at the
base of a large stone six inches beneath the surface. This burrow
resembled a miniature muskrat burrow and apparently had been dug when
the rat was under water.

Like _Phenacomys_, the water rat constructs sub-snow nests on the
surface of the ground. These are recognizable by their large size and by
piles of ovoid droppings a quarter of an inch in length. These nests are
loosely built and fall apart soon after the snow melts.

The water rat is sometimes a pest to the mammal collector for they
spring mouse traps set for other mammals without becoming caught. At
times the greater part of an entire trap line was thus rendered
ineffective by these mammals. Fully adult animals are uncommon in
collections for a rat trap or steel trap is needed to take them and
these items, when packed on back up mountains to water-rat habitat, are
usually set for still larger animals. Racey (Racey and Cowan, 1935: H27)
recounting his difficulty in securing specimens when no suitable traps
were at hand, writes: "Killed one with my hands, shot another, and a
third was stunned by a mouse trap."

Taylor and Shaw (1927: 76) list food eaten by the water rat on Mount
Rainier as the avalanche lily, dogtooth violet, _Ligusticum purpureum_,
_Valeriana sitchensis_, _Polygonum bistortoides_, _Petasites frigida_,
_Phyllodoce empetriformis_, _Potentilla flabellifolia_, _Aster_ sp.,
grass, wild clover, conifer seeds, two kinds of blueberry (_Vacinnium_)
and _Xerophyllum tenax_. Racey and Cowan (1935) list foods eaten in the
Cascades of southern British Columbia as _Lupinus polyphyllus_, _Senecio
balsamitae_, _Pedicularis bractiosa_ and _Arnica alpinus_.

A female from Dewey Lake, Yakima County, contained 4 embryos on
September 1, 1940. One from Tye, King County, had 2 embryos on September
8, 1940.

=Microtus richardsoni arvicoloides= (Rhoads)

      _Aulacomys arvicoloides_ Rhoads, Amer. Nat., 28:182, February,

      _Microtus richardsoni arvicoloides_ Bailey, N. Amer. Fauna, 17:62,
      June 6, 1900.

      _Type._--Obtained at Lake Keechelus, Kittitas County, Washington,
      by A. Rupert in September, 1893. Rhoads gives the altitude as 8,000
      ft. This apparently is an error, for the elevation of the lake is
      2,458 ft. and the summit of Snoqualmie Pass, to the west, is 3,100
      ft. Probably 3,000 ft. was intended; type in Philadelphia Academy
      of Natural Sciences.

      _Racial characters._--Size large; color of upper parts dark
      reddish brown; underparts paler.

  [Illustration: FIG. 117. Distribution of the water rat in Washington.
  A. _Microtus richardsoni arvicoloides._ B. _Microtus richardsoni

      _Measurements._--A female from Tomyhoi Lake, Whatcom County,
      measured: total length 242; length of tail 68; hind foot 27; ear
      17. A female from Tye, King County, measured: 257; 83; 26; 17.

      _Distribution._--The Cascade Mountains, from Tomyhoi Lake (W.W.D.)
      south to Potato Hill (Taylor and Shaw, 1929: 25).

=Microtus richardsoni macropus= (Merriam)

      _Arvicola (Mynomes) macropus_ Merriam, N. Amer. Fauna, 5:60, July
      30, 1891.

      _Microtus richardsoni macropus_ Bailey, N. Amer. Fauna, 17:61,
      June 6, 1900.

      _Microtus richardsonii macropus_ Taylor and Shaw, Occ. Pap. Chas.
      R. Conner Mus., no. 2:25, December, 1929.

      _Type._--Obtained in the Pashimeroi Mountains, 9,700 ft., Custer
      County, Idaho, by C. H. Merriam and V. Bailey in 1890; type in
      United States National Museum.

      _Racial characters._--Similar to _arvicoloides_ but slightly
      smaller and redder.

      _Measurements._--A female from Stay-a-while Spring, Columbia
      County, measures: total length 228; length of tail 73; hind foot
      25; ear 14.

      _Distribution._--The Blue Mountains of southeastern Washington.

=Microtus oregoni oregoni= (Bachman)

Creeping mouse

      _Arvicola oregoni_ Bachman, Jour. Acad. Nat. Sci. Philadelphia,
      8:60, 1839.

      _Microtus oregoni_ Miller, N. Amer. Fauna, 12:9, July 23, 1896.

      _Microtus morosus_ Elliot, Field Columb. Mus. Publ. 30, zoöl.
      ser., 1:227, February 1, 1899 (type from Boulder Lake, 5,000 ft.,
      Clallam County, Washington).

      _Microtus oregoni oregoni_ Miller, U. S. Nat. Mus. Bull., 79:227,
      December 31, 1912.

      _Microtus oregoni cantwelli_ Taylor, Jour. Mamm., 1:180, August
      24, 1920 (type from Glacier Basin, 5,935 ft., Mt. Rainier, Pierce
      County, Washington).

  [Illustration: FIG. 118. Distribution of the creeping mouse, _Microtus
  oregoni oregoni_, in Washington.]

      _Type._--Obtained at Astoria, Clatsop County, Oregon, by J. K.
      Townsend in 1836; type in Philadelphia Academy of Natural Sciences.

      _Measurements._--Ten males and 10 females from southwestern
      Washington, average, respectively: total length 133, 126; length
      of tail 35, 32; hind foot 16.9, 16.7; ear 9.8, 9.7; weight 19.3,
      19.1 grams.

      _Distribution._--The Cascade Mountains, the Olympic Mountains, and
      the lowlands of western Washington.

_Remarks._--A large series of topotypes of _Microtus o. oregoni_ in the
California Museum of Vertebrate Zoölogy, and the rather large series
from Cowlitz County, Washington, show a wide range of variation in
color, size, and cranial characters. Specimens from the Cascade and
Olympic Mountains seem to average a bit paler than topotypes of
_oregoni_, but are not worthy of recognition as distinct races. Too few
topotypes of _Microtus o. serpens_ from British Columbia are available
to judge the status of that race with any certainty, but specimens from
northwestern Washington are certainly _oregoni_.

_Description._--The creeping mouse is a small species of general
microtine form. The head and body measure about 4 inches and the tail
about 1-1/2 inches. The ears are small and the eyes tiny, nearly buried
in the fur. The fur is short and rather rough. It does not lie back
smoothly, thereby giving a woolly appearance. The upper parts are
reddish or grayish brown and the underparts are grayish white. The
nondescript appearance suggests a young rather than adult mouse.

The creeping mouse (subgenus _Chilotus_) is restricted to the Pacific
Coast and ranges from British Columbia to California. In Washington it
occupies almost every conceivable "mouse" habitat in its range,
including wet marshes, damp ravines, dry forest, damp, mossy forest,
meadows, alpine meadows and fields of short grass. It is rare in all but
the latter habitat. In fields of short or dry grass it is often
abundant. In the Cascade Mountains it was in relatively dry places along
streams or rock slides. Altitudinally it ranges from sea level to at
least 6,000 feet, and from the Humid Transition well into the Hudsonian

Creeping mice construct tiny tunnels among the grass roots and seldom
venture out of them. In suitable habitat the surface of the ground
beneath the grass is a maze of these tunnels, which cross, intersect,
and divide in a complex network. An observer standing in a field
occupied by creeping mice can scarcely conceive of the extent and
perfection of the tiny tunnel system at his feet.

Creeping mice lived but a day or two in captivity. Save for the bits of
grass blades left in their runways, little is known of their food or
other life habits. Their nests are round balls of dry grasses placed in
cavities under logs. None of the many examined possessed a lining of
softer materials. Embryos found in pregnant females from April 10 to May
18 numbered from 2 to 4.

=Lagurus curtatus pauperrimus= (Cooper)

Sagebrush vole

      _Arvicola pauperrima_ Cooper, Amer. Nat., 2:535, December, 1868.

      _Arvicola pauperrimus_ Merriam, N. Amer. Fauna, 5:64, July 30,

      _L[agurus]. pauperrimus_ Thomas, Ann. and Mag. Nat. Hist., ser. 8,
      9:401, April, 1912.

      _Microtus pauperrimus_ Bailey, N. Amer. Fauna, 55:214, August 29,

      _Lemmiscus pauperrimus_ Davis, Recent Mamm. Idaho, Caxton
      Printers, p. 327, April 5, 1939.

      _Lemmiscus curtatus pauperrimus_ Goldman, Proc. Biol. Soc.
      Washington, 54:70, July 31, 1941.

      _Type._--Obtained on the "Plains of the Columbia" near the Snake
      River, southwestern Washington by J. G. Cooper on October 9, 1860.
      Probably from the Bunchgrass Hills near Wallula (Old Fort Walla
      Walla), Walla Walla County: type in United States National Museum.

      _Measurements._--Bailey (1900: 69) gives the average of 3 adults
      from the vicinity of Antelope, Oregon, as: total length 115;
      length of tail 20; hind foot 16.

      _Distribution._--Known in Washington only from the type and a
      specimen from Badger Mountains, 8 miles southwest of Waterville
      (Taylor and Shaw, 1929: 25).

_Remarks._--This rare vole is the smallest microtine rodent in
Washington. The head and body measure about 4 inches and the tail about
1 inch. The upper parts are grayish or yellowish brown and the
underparts grayish. The upper incisors are not grooved and the inner and
outer angles of the molars are about equal in length.

Mice of the genus _Lagurus_ occur in Siberia and in prairie areas of the
northwestern United States and Canada. The sagebrush vole inhabits the
Upper Sonoran Life-zone. It prefers upland areas of low sagebrush with
sparse grass. Poorly formed runways and small piles of feces indicate
its presence. The type of _Lagurus pauperrimus_ was obtained in
southwestern Washington 80 years ago, and a single additional record has
since been obtained. Nothing is known of its habits in Washington. The
life history of related forms has been reported on by Hall (1928:
201-204) from Nevada and Moore (1943: 188-191) from Oregon.

Davis (1939: 326) raised the subgenus _Lemmiscus_ Thomas (for American
forms) to generic rank and Goldman (1941: 69) accepted this usage.
Comparison of the Siberian and American species reveals but three
impressive differences: the dorsal stripe of the Siberian mice, the
greater development of prisms in the molars of the American species, and
the presence of cement in the angles of the molars of the Siberian
specimens and its lack in the American. These differences seem to be of
no more than subgeneric value. Goldman (1941: 69) showed that all
American _Lagurus_ belong to a single species.

=Ondatra zibethicus= (Linnaeus)


_Description._--The muskrat is a large aquatic rodent. Head and body
measure about 14 inches; the tail about 10 inches. The body is plump and
the head small. Eyes and ears are relatively small. The forefeet are
small and handlike with furred wrists. The hind feet are large with
webbed toes and naked wrists. The tail is narrow, constricted at the
base and flattened vertically. It is scaled and possesses scattered,
stiff hairs. The underfur is dense and soft. The guard hair is stiff and
shiny. The upper parts are rich, dark brown. The underparts are gray
washed with cinnamon.

  [Illustration: FIG. 119. Muskrat (_Ondatra zibethicus osoyoosensis_),
  male, Lake Washington, Seattle, October 13, 1939; weight 906 grams on
  November 4, 1939. (Fish and Wildlife Service photo by Victor B.
  Scheffer, No. 736.)]

Muskrats are found in Canada and the United States. They are lowland
animals, rarely ranging into the Canadian Life-zone. Their aquatic
habitat makes them relatively independent of temperature, and
consequently they occur in both the Transition and Upper Sonoran

The muskrat is well adapted to aquatic life. The toes of the large hind
feet are webbed at the base, the hind feet are turned slightly outward,
a fringe of stiff bristles at the rear edge of each foot furnishes
additional aid in swimming, and the laterally compressed tail makes it
an efficient rudder. In the water the surface of the fur flattens down
to entrap air in the dense underfur, keeping the body dry.

The typical habitat of the muskrat is slow-moving water or still water,
such as in lakes, ponds, marshes and sluggish rivers and streams.
Muskrats occur commonly, though not in large numbers, in more swiftly
moving streams. In Puget Sound and the San Juan Islands they occupy a
marine habitat.

In the extensive marshes along Lake Washington, King County, muskrats
are abundant. They occupy sluggish water, often water polluted by
garbage and sewage. In these marshes, banks suitable for the
construction of burrows are absent and houses are made of cattail stalks
and leaves. The food of these marsh-living muskrats consists principally
of cattail and other marsh vegetation.

In contrast to the marsh-occupying muskrats, muskrats along the open,
marsh-free shores of the lake live in deep clear water where waves lap
the shore. These muskrats live exclusively in burrows dug in the banks
and feed upon fresh-water mussels.

In still greater contrast were muskrats living 20 miles away, near
Cottage Lake, King County. Here we found them in small clear streams, 4
to 10 feet wide. Depth of the water varied from a few inches to three
feet. The streams flowed through meadows, pastures and junglelike,
deciduous woods. Muskrats were slightly more common along wooded
stretches than in open areas. Some were trapped where the streams flowed
at high velocity over shallow, gravel bottoms. The animals lived in
burrows and fed upon fresh-water mussels and a variety of plants.

Near Richmond Beach, Snohomish County, muskrats took up residence in a
small tidal pool along Puget Sound. The nearest fresh water stream large
enough to support a muskrat was two miles away. Two muskrats were
trapped here. Investigation of a tidal pool a mile to the north
disclosed unmistakable muskrat signs. Traps set in the culvert
connecting the pool with Puget Sound at high tide took several
specimens. Study showed that the muskrats were not living in the pool
but among the large boulders forming the breakwater for the Great
Northern Railroad, along the sound itself. They were feeding on marine
mussels (_Mytilus_). These mussels lived in the salt water of the sound,
not in the tidal pool.

At Peavine Pass, Blakely Island, in the San Juan Islands, muskrats were
living in the swift tidal current and deep, marine waters. Several were
seen in late afternoon. All were swimming parallel to the shore about 50
feet out. Here also they fed on _Mytilus_, but their homes were not
discovered. Certainly they were not living in the tidal pool at Flat
Point, a half-mile away.

In the interior of Blakely Island a colony of muskrats was discovered
living in a marsh of about one acre. In the rainy season the ground of
the marsh was covered with less than one inch of water. Residents said
that in the dry season springs kept the ground moist. Muskrats were
living in burrows whose entrances descended at a 45-degree angle and
were filled with water. The ground about some occupied burrows was dry,
the only water visible being in the burrow itself. A variety of marsh
vegetation provided food.

  [Illustration: FIG. 120. Distribution of the muskrat in Washington.
  A. _Ondatra zibethicus osoyoosensis._ B. _Ondatra zibethicus

Burrows of muskrats always have entrances under water. Usually they
enter a vertical bank 6 to 15 inches below water line and occasionally 3
feet below it. About half the burrows excavated near Lake Washington,
King County, had a single entrance. About 40 per cent had double or
triple entrances situated 2 to 3 feet apart and converging within a yard
to a single burrow. About 10 per cent had double burrows more than 3
feet in length. Burrows were from 5 to 8 inches in diameter. Nest
chambers were from 12 to 15 inches in diameter, spherical, and from 6 to
30 feet from the burrow entrance. The nests themselves were bulky, loose
masses of cattail leaves. Embryos found in late February and early
March numbered 4 to 8.

Because it is abundant, widely spread and easy to trap, the muskrat is
one of the most important fur bearers in the state. The fur is
relatively stable in value. In recent years the average skin has brought
the trapper slightly less than a dollar. Muskrat flesh is eaten and sold
on the market in the eastern United States but has never been popular in
Washington. The muskrat does little damage to agriculture, most
complaints arising from its burrows which interfere with irrigation

=Ondatra zibethicus osoyoosensis= (Lord)

      _Fiber osoyoosensis_ Lord, Proc. Zoöl. Soc. London, p. 97, 1863.

      _F[iber]. z[ibethicus]. osoyoosensis_ Hollister, Proc. Biol. Soc.
      Washington, 23:1, February 2, 1910.

      _Ondatra zibethica osoyoosensis_ Miller, U. S. Nat. Mus. Bull.,
      79:231, December 31, 1912.

      _Type._--Obtained at Osoyoose Lake, British Columbia, on British
      Columbia-Washington boundary at head of Okanogan River, by J. K.
      Lord, in 1861 or 1862.

      _Racial characters._--Color of the upper parts rich, dark brown.

      _Measurements._--Two males and a female from Seattle, King County,
      average and measure respectively: total length 565, 555; length of
      tail 262, 257; hind foot 80.5, 79; weight 2 pounds 13 ounces and 2
      pounds 3 ounces.

      _Distribution._--Eastern Washington generally and all but the
      southern part of western Washington. Specimens from as far
      southwest as Tenino (W.W.D.) are typical of _osoyoosensis_.

=Ondatra zibethicus occipitalis= (Elliot)

      _Fiber occipitalis_ Elliot, Field Columb. Mus. Publ. 74, zoöl.
      ser., 3:162, April, 1903.

      _Ondatra zibethica occipitalis_ Miller, U. S. Nat. Mus. Bull.,
      79:231, December 31, 1912.

      _Type._--Obtained at Florence, Lane County, Oregon, by E. Heller,
      in 1901; type in Chicago Natural History Museum.

      _Racial characters._--Similar to _osoyoosensis_ but redder; fur
      shorter and interpterygoid spaces of skull narrower.

      _Measurements._--A male from Chinook, Pacific County, measures:
      total length 580; length of tail 248; hind foot 78; ear 21; weight
      863 grams.

      _Distribution._--The southwestern corner of the state, extending
      north to Aberdeen (V.B.S.) and east to Cathlamet (V.B.S.).

=Rattus rattus= (Linnaeus)

Roof rat

_Description._--Size large, total length approximately 400 mm.; tail
long, naked, comprising 50 per cent or more of total length; color sooty
black or brown; in brown phase, whitish beneath.

The roof rat became established in Central America some 350 years ago,
and entered the United States (English Colonies) late in the seventeenth
century. Subsequent to the introduction of the Norway rat the roof rat
decreased in numbers and is now found only in restricted areas.

=Rattus rattus rattus= (Linnaeus)

      _[Mus] rattus_ Linnaeus, Syst. Nat., 1 (ed. 10):61, 1758.

      _Rattus rattus_ Hollister, Proc. Biol. Soc. Washington, 26:126,
      June 6, 1916.

      _Type._--From Uppsala, Sweden.

_Remarks._--In Washington I have taken specimens of the roof rat in the
San Juan Islands and in the coniferous forests on the west slope of the
Cascades. In the latter area it seems to be widely spread and to live in
the wild.

=Rattus rattus alexandrinus= (Geoffroy)

      _Mus alexandrinus_ Geoffroy, Catal. Mam. du Mus. Nat. d'Hist. Nat.
      Paris, p. 192, 1803.

      _R[attus]. rattus alexandrinus_ Hinton, Jour. Bombay Nat. Hist.
      Soc., 26:63, December 20, 1918.

      _Type._--From Alexandria, Egypt.

_Remarks._--This subspecies seems to be rare in Washington. My specimens
are all from small, isolated islands in the San Juan group. All were
taken in the vicinity of human habitations. _R. r. alexandrinus_
resembles _R. r. rattus_, differing only in brown color of upper parts.

=Rattus norvegicus norvegicus= (Erxleben)

Norway rat

      _[Mus] norvegicus_ Erxleben, Syst. Regni Anim., 1:381, 1777.

      _Rattus norvegicus_ Hollister, Proc. Biol. Soc. Washington,
      29:126, June 6, 1916.

      _Type._--From Norway.

      _Description._--Larger, heavier-bodied and coarser-furred than the
      black rat or roof rat with shorter, heavier tail. The tail is less
      than 50 per cent of the total length. The color of the back is
      dull, reddish brown, the sides are paler and the underparts are
      dirty gray.

_Remarks._--The Norway rat was absent from the Pacific Coast of the
United States before 1851. It probably reached the coast slightly after
that date. It is common about all large cities in Washington. In the
western part of the state it lives along streams and marshes under feral

=Mus musculus= Linnaeus, subsp?

House mouse

      _[Mus] musculus_ Linnaeus, Syst. Nat., 1 (ed. 10):62, 1758.

_Description._--Size small; tail about 50 per cent of total length,
naked; ears small, about 12 mm. in height; upper incisors not grooved;
color of upper parts reddish or grayish brown; underparts brown or gray.

_Remarks._--The house mouse became established in North America soon
after its settlement by Europeans. It is now common throughout the state
of Washington, principally near human habitations, but often lives in
the wild.

Races of the house mouse have been dealt with by Schwartz and Schwartz
(1943: 59-72), and by Nichols (1944: 82-89), but lack of adequate
material prevents subspecific identification of house mice from
Washington at this time.

=Aplodontia rufa= (Rafinesque)

Mountain beaver

_Description._--The mountain beaver is a stout-bodied animal about 14
inches in length, with a tiny tail that is almost invisible externally.
The head is large, wide and low with small eyes, small ears and long
vibrissae. The legs are short and heavy, but the forefeet are small and
handlike; the hind feet are large and powerful. The claws of both
forefeet and hind feet are long and strong. The pelage is short, coarse
and rough. The upper parts are dark reddish brown and the underparts are
grayish brown. The feet are pink.

  [Illustration: FIG. 121. Mountain beaver (_Aplodontia rufa rufa_),
  Seattle, Washington, March 19, 1940. (Fish and Wildlife Service photo
  by Victor B. Scheffer, No. 919.)]

Mountain beavers are confined to the Pacific Coast and range from
southern British Columbia to central California. The genus contains a
single species of which Taylor (1918) recognized nine races. The
principal habitat of the mountain beaver is clearings at the edge of
coniferous forests. The animals are most abundant near springs, streams
and damp places, although they are not aquatic. The tangled jungles of
deciduous trees and shrubs that grow in the ravines and stream valleys
of the Puget Sound area present optimum habitat. They occur also on
hillsides, on logged-off land and along roadside clearings. In the
mountains they occur in thickets and forests, always, in our experience,
near streams.

The most conspicuous evidence of the presence of mountain beavers is
their burrows. These are large tunnels, four to eight inches in
diameter. To each set of tunnels there are numerous entrances, some
partly concealed in brush or beside logs or stumps, and some are in the
open. Those in the open are less used as entrances than as places for
receiving the loose earth which the animals excavate. A pile may contain
nearly a cubic yard of earth and stones. Many of the burrows are
shallow, and cave-ins are common. Breaks in the roof of a burrow are not
repaired, although debris is removed from the burrow itself. The burrows
seem not to be constructed according to a system, but are extended to
take in whatever brush, logs or other cover is available. They are
commonly dug through damp or muddy soil. Small streams flow through some
burrows. Such partly flooded runs seem to be favored by the animals.

The nest of a mountain beaver excavated by Scheffer (1929: 15), under
the roots of a fallen tree, was oval in shape, twenty inches high and 13
inches wide. The nest was protected from flooding by a basal chamber, or
basin, six inches beneath the nest. Two drainage tunnels lead away from
this basal chamber. The nest was composed of the leaves and stems of
bracken laced together with grass and fine twigs. Two other nests
examined by Scheffer measured 17 by 18 inches and 19 by 17 inches. Both
were about two feet beneath the surface.

Around Puget Sound the mountain beavers mate in early March. The young
number two to three, rarely four, per litter and are born in early

Mountain beavers enter water readily but wade rather than swim. They are
rather noisy, splashing in water and breaking twigs or rustling leaves
on the ground. They climb bushes and saplings, clipping off branches for
food as they ascend. According to Scheffer (1929: 15) they leave the
stubs of branches attached to the trunk to facilitate their descent.
Twice a mountain beaver was found several feet up in a sapling. In both
saplings the animal had clipped the branches close to the trunk and was
desperately reaching with its hind feet for missing branches. When the
observer came near, one animal squealed, tumbled to the ground, and
scurried frantically to its burrow.

Although principally nocturnal, mountain beavers are not infrequently
active by day, especially in the fall. At this season they harvest food
and spread it on logs to dry. The cured hay is removed to their burrows
for nesting material and food. In winter mountain beavers are more
restricted in habits and are rarely seen by day. Presumably they feed on
stored food at this time but they forage somewhat. In winter they eat
such evergreen shrubs as salal (_Gaultheria shallon_) and Oregon grape
(_Berberis nervosa_). They eat also the bark of trees, especially that
of the willow (_Salix_). Under cover of snow, in the mountains, they
burrow to some extent and pack excavated earth in snow burrows. The
melting of the snow in the spring reveals the earth core, six to eight
inches in diameter and two to four feet long. Several such earth cores
were forked, showing that part of the earth had been pushed into a
branching burrow.

  [Illustration: FIG. 122. Distribution of the mountain beaver in
  Washington. A. _Aplodontia rufa rufa._ B. _Aplodontia rufa rainieri._]

The mountain beaver holds its food in its forefeet, squirrel-like, when
it eats. Its food consists of the leaves and bark of woody plants and
entire herbs, including roots. The mountain beaver is the only mammal so
far as known that eats the bracken fern. It feeds on the branches of
coniferous trees, including Douglas fir, red cedar, and hemlock. Such
thorny species as the blackberry, blackcap and devil's club are eaten.
The odiferous skunk cabbage and the stinging nettle are on its bill of
fare. A list of its food would include most plants found in its habitat,
and we know of no species that it refuses as food.

The mountain beaver is more of a nuisance than a pest. In most of its
range there is but little farming although where crops are raised the
mountain beaver may do some damage. It undermines roads and trails and
defiles springs and streams. Control is simple for the animals readily
enter steel traps set in their burrows.

=Aplodontia rufa rufa= (Rafinesque)

      _Anisonyx? rufa_ Rafinesque, Amer. Monthly Mag., 2:45, November,

      _Haplodon rufus_ True, Proc. U. S. Nat. Mus., 7 (1884):596, 1885.

      _Aplodontia rufa_ Merriam, Ann. New York Acad. Sci., 3:316, May,

      _Aplodontia olympica_ Merriam, Proc. Biol. Soc. Washington, 13:20,
      January 31, 1899 (type from Lake Quiniault, Grays Harbor County,

      _Aplodontia rufa grisea_ Taylor, Univ. California Publ. Zoöl.,
      12:497, May 6, 1916 (type from Renton, King County, Washington).

      _Aplodontia rufa rufa_ Taylor, Univ. California Publ. Zoöl.,
      12:497, May 6, 1916.

      _Type._--None. Based on a description by Lewis and Clark. Taylor
      (1918: 455) regarded as typical specimens collected at "Marmot,
      Clackamas County, Oregon (western slope of Mount Hood, not far from
      the Columbia River)."

      _Racial characters._--Size small; skull small.

      _Measurements._--Eight males and 7 females, from the area about
      Puget Sound, average, respectively: total length 343, 338; length
      of tail 37.5, 33.5; hind foot 57.8, 56; ear 25, 24.5; weight 1342,
      1300 grams.

      _Distribution._--Western Washington, between Puget Sound and the
      Cascade Mountains and southward. Marginal localities are:
      Bellingham (U.S.N.M.), Sauk (U.S.N.M.), Forks of Skykomish River
      (W.W.D.), North Bend (U.S.N.M.), and mouth of Klama River

_Remarks._--Individual variation in mountain beavers is considerable.
Two weakly defined races are recognized in Washington.

=Aplodontia rufa rainieri= Merriam

      _Aplodontia major rainieri_ Merriam, Proc. Biol. Soc. Washington,
      13:21, January 31, 1899.

      _[Haplodontia rufa] raineri_ Elliot, Field Columb. Mus. Publ. 45,
      zoöl. ser., 2:112, 1901.

      _[Aplodontia rufa] raineri_ Trouessart, Catal. Mamm., viv. foss.,
      suppl. p. 348, 1904.

      _Aplodontia rufa columbiana_ Taylor, Univ. California Publ. Zoöl.,
      12:499, 1916 (type from Hope, British Columbia).

      _Type._--Obtained at Paradise Creek, 5, 200 ft., Mount Rainier,
      Pierce County, Washington, by V. Bailey on August 6, 1897; type in
      United States National Museum.

      _Racial characters._--Similar to _rufa_ but larger with larger

      _Measurements._--Three males and a female from Tye, King County,
      average and measure respectively: total length 352, 340; length of
      tail 35, 40; hind foot 60, 59; ear 24, 19.

      _Distribution._--The higher Cascade Mountains from the Columbia
      River northward to the Canadian boundary. Marginal localities are:
      Canyon Creek (U.S.N.M.), Cascade Pass (U.S.N.M.), Tye (W.W.D.),
      Mt. Rainier (U.S.N.M.), Mt. St. Helens (U.S.N.M.), and Yacolt

_Remarks._--Intergradation between _rufa_ and _raineri_ is seen in
specimens from the area between Stevens Pass and Skykomish, King County.

=Zapus princeps= Allen

Big jumping mouse

  [Illustration: FIG. 123. Big jumping mouse (_Zapus princeps
  trinotatus_) in hibernation. Puyallup, Washington, January 30, 1939.
  (Fish and Wildlife Service photo by Victor B. Scheffer, No. 576.)]

_Description._--The big jumping mouse is a medium-sized mouse, slightly
larger than the house mouse, with an exceptionally long, tapering tail.
Head and body measure about 4 inches and the tail about 5 inches. The
body is slender with a small head, small eyes and small, naked ears. The
forefeet are small and handlike but the hind feet are large with long,
powerful legs. The long, naked tail is smoothly tapering to a narrow
point. The fur is short, posteriorly directed, stiff and bristlelike.
The dorsal area is dusky, the sides are orange or yellow, and the
underparts are creamy white. Each upper incisor tooth has a groove on
its anterior face.

Jumping mice are boreal mammals occurring in wooded regions ranging from
the Arctic region southward to North Carolina, New Mexico and
California. Jumping mice are of rather general distribution in
Washington, being only locally common. They are partial to damp, boggy
areas but avoid true swamps. They occur in clearings in the forests in
the Puget Sound area and in meadows in river bottoms and in jungles in
ravines. They reach their greatest abundance in the boggy meadows and
glacial cirques of the mountains and spread out from such areas to
seemingly less favored habitat on dryer slopes and heather meadows.
Unlike many species, they do not seem to inhabit talus slides. They are
entirely absent from the eastern Washington desert.

Sometimes the jumping mouse walks on all four feet but the ordinary
means of progression is by short hops on the hind feet alone. When
startled they travel in great bounds, covering six feet or more at a
jump. When jumping they make considerable noise, swishing or rustling
through the grass and landing with an audible thud. The long tail serves
as a balancing organ. A specimen whose tail had been lost was reported
by Svihla and Svihla (1933: 133) to turn somersaults in the air and
invariably to land on its back rather than its feet.

Jumping mice become very fat and hibernate in middle summer or early
fall. In the lowlands they disappear by late July but in the mountains
they remain active until the middle of September. They spend the winter
in nests of grass several feet beneath the surface. A hibernating
individual figured by Flahaut (1939: 17) was curled in a ball, head down
with the tail wrapped completely around the greatest circumference of
the ball.

Near Seattle the principal food of the jumping mouse was the velvet
grass (_Holchus lanatus_), and the seeds of grasses and the broad-leaved
dock. The fruit of the blackberry (_Rubus macropetalus_) is eaten and
occasional individuals are seen with the chin stained a deep purple from
the juice.

The new-born young of the jumping mouse were reported by Svihla and
Svihla (1933: 132) to average 0.8 grams in weight. They are pink,
hairless, lacking even the facial vibrissae, with eyes closed and ears

=Zapus princeps trinotatus= Rhoads

      _Zapus trinotatus_ Rhoads, Proc. Acad. Nat. Sci. Philadelphia,
      1894, p. 421, January 15, 1895.

      _Zapus imperator_ Elliot, Field Columb. Mus. Publ. 30, zoöl. ser.,
      1:228, February 1, 1899 (type from Sieg's Ranch, Elwha River,
      Olympic Mountains, Clallam County, Washington).

      _Type._--Obtained on Lulu Island, mouth of Fraser River, British
      Columbia, by S. N. Rhoads on May 31, 1892; type in Philadelphia
      Academy of Natural Sciences.

      _Racial characters._--Similar to _oregonus_ but brighter; sides
      orange; underparts creamy white; buffy area often present on chest.

      _Measurements._--Twenty males and 15 females from western
      Washington average, respectively: total length 230, 233; length of
      tail 140, 140; hind foot 33, 32.8; ear 14.9, 16.3; weight 23.7,
      25.7 grams.

      _Distribution._--Western Washington and the Cascade Mountains,
      east in the northern Cascades at least to Tomyhoi Lake (W.W.D.).

_Remarks._--Actual intergrades between _trinotatus_ and _kootenayensis_
have not been examined but the differences separating the two forms are
of the degree that usually distinguish subspecies. Since _trinotatus_
occurs in the high Cascades as far east as Mount Baker, at least, and
_kootenayensis_ probably occurs in the northeastern Cascades, the two
forms doubtless come together in the rugged, inaccessible area between
these two localities. Further collecting will probably show a narrow
zone of intergradation in extreme western Okanogan County.

  [Illustration: FIG. 124. Distribution of the big jumping mouse in
  Washington. A. _Zapus princeps oregonus._ B. _Zapus princeps
  idahoensis._ C. _Zapus princeps kootenayensis._ D. _Zapus princeps

=Zapus princeps kootenayensis= Anderson

      _Zapus princeps kootenayensis_ Anderson, Ann. Rept. Nat. Mus.
      Canada for 1931, p. 108, November 24. 1932.

      _Type._--Obtained on Green Mountain, 6,000 ft., 10 miles north of
      Rossland, West Kootenay District, British Columbia, by R. M.
      Anderson, on July 18, 1929; type in National Museum of Canada.

      _Racial characters._--Similar to _oregonus_ but paler; yellow more

      _Measurements._--Twenty females, including 15 topotypes, average
      (Anderson, 1932: 109): total length 245; length of tail 140; hind
      foot 30.5.

      _Distribution._--Northeastern Washington. Specimens from Sullivan
      Lake (E.S.B.) have been examined.

=Zapus princeps idahoensis= Davis

      _Zapus princeps idahoensis_ Davis, Jour. Mamm., 15:221, August 10,

      _Type._--Obtained 5 miles east of Warm Lake, 7,000 feet elevation,
      Valley County, Idaho, by W. B. Davis; type in Museum of Vertebrate

      _Racial characters._--Similar to _kootenayensis_ but brighter in
      color, more ochraceous. Similar to _oregonus_ but paler, more

      _Measurements._--Davis (1939:339) gives the measurements of six
      adult topotypes as: total length 240; length of tail 144; hind
      foot 31.

      _Distribution._--A single specimen in the Charles R. Conner Museum
      from Kamiak Butte, Whitman County, is referable to this race.

=Zapus princeps oregonus= Preble

      _Zapus princeps oregonus_ Preble, N. Amer. Fauna. 15:24, August 8,

      _Type._--Obtained at Elgin, Union County, Oregon, by E. A. Preble,
      on May 29, 1896; type in United States National Museum.

      _Racial characters._--Small size, pale color.

      _Measurements._--Three males and 3 females from the Blue Mountains
      average, respectively: total length 233, 234; length of tail 138,
      139; hind foot 31.8, 31.8; ear 16, 16; weight 29, 33 grams.

      _Distribution._--The Blue Mountains of southeastern Washington.

=Erethizon dorsatum= (Linnaeus)


_Description._--The porcupine is one of the largest rodents found in
Washington, being exceeded in size only by the beaver. Its body is heavy
and stocky, its legs short, its tail long and thick and its eyes small.
It is best known for the modified hairs, or quills, of its tail and
dorsal area. These vary in greatest diameter from one-sixteenth to
three-sixteenths of an inch and from three-quarters of an inch to five
inches in length. They are ivory-white with black tips. In addition to
quills, the porcupine possesses wooly, black underfur and long, banded
guard hair. The bands of the guard hairs are black and yellow, varying
in width. Commonly they are of a single color, black, yellow or brown.

Porcupines range over virtually all wooded parts of North America north
of Mexico, in and above the Transition Life-zone. The Canadian
porcupines have been studied by Anderson and Rand (1943A) and
intergradation between the eastern _dorsatum_ and the western
_epixanthum_ has been shown.

The porcupines are commonly considered to be forest animals. However,
they are rather rare in the denser coniferous forests. In the more open
areas on the Cascade Mountains, especially on the eastern slopes, they
are not uncommon. They are rather common in the coniferous forests of
northeastern Washington and the Blue Mountains. They seem to be rather
common also in desert areas at the southern edge of the Columbian

The huge incisors of the porcupine are adapted to feeding on bark. They
do feed on bark to a certain extent in Washington, but it is my
observation that more herbs and bushes are eaten than bark. In areas
where porcupines are common, trees are commonly girdled, usually close
to the top. Trees girdled in this manner in the Kettle River Mountains
included western larch, ponderosa pine, and grand fir. The tops of some
trees were killed.

The ordinary walking gait of the porcupine is a slow deliberate walk in
which he appears to waddle somewhat. They can increase their speed to a
slow trot. They are slow, deliberate climbers, ascending and descending
trees with head upward. They are able also to climb rocks and cliffs,
sometimes being seen on the tops of large boulders.

Despite their large size, porcupines are not commonly seen. They are
mainly nocturnal and, in the daytime, find concealment high in the
branches of some conifer or a cave between the rocks in a talus slide.

While encamped near Sherman Creek Pass in the Kettle River Mountains my
companion and I heard a crunching of gravel from the road fifty feet
away. We listened intently, wondering what person would be abroad in the
mountains at midnight. In the vicinity of our car, concealed from our
view by trees, the noise stopped, to be followed a few minutes later by
a rasping and clattering that could be heard far away. We raced to the
car to discover a large porcupine crouched on the running board by a
pile of "pick-up" antlers of the white-tailed deer left there by us. The
"porkie" had been chewing on these, heedless of the noise made by the
loose antlers clashing against the metal side of the car.

  [Illustration: FIG. 125. Distribution of the porcupine, _Erethizon
  epixanthum_ in Washington. Boundaries between ranges of subspecies are

In Washington the single young is born late in May or early in June.
There are two pairs of mammae, both pectoral, of which only the anterior
are functional.

=Erethizon dorsatum epixanthum= Brandt

      _Erethizon epixanthus_ Brandt, Mem. Acad. Imp. Sci. St.
      Pétersbourg, ser. 6, 3 (Sci. Nat. vol. =1=): 390, 1835.

      _Erethizon dorsatus epixanthus_ True, Proc. U. S. Nat. Mus.,
      7:600, 1885.

      _Erethizon epixanthum epixanthum_ Miller, U. S. Nat. Mus. Bull.,
      128:437, April 29, 1924.

      _Erethizon dorsatum epixanthum_ Anderson and Rand, Canadian Jour.
      Research, 21:293, September 24, 1943.

      _Type._--None. Type locality California.

      _Racial characters._--Size large, total length of adults
      approximately 30 inches; tail long (nearly one-third of total
      length), thick, heavy and spiny; body stout; legs short; claws
      long and curved; ears and eyes small; body spines short, thick and
      most abundant on posterior part of back, longer and more slender
      on sides and shoulders; guard hairs of shoulders and sides long,
      almost concealing spines; fur of underparts shorter; color
      variable, brown, black or yellow. In winter the fur is longer and
      woolly, concealing spines.

      _Distribution._--The Columbian Plateau and the Blue Mountains.

_Remarks._--Anderson and Rand (1943A: 295) ascribe two races to
Washington. With inadequate material myself to verify this ascription. I
think it probable that the northern forest porcupine and the Great Basin
animal are racially different. In consequence the available names,
_nigrescens_ and _epixanthum_, are here applied, pending a revision of
the entire genus.

=Erethizon dorsatum nigrescens= Allen

      _Erethizon epixanthus nigrescens_ Allen, Bull. Amer. Mus. Nat.
      Hist., 19:558, October 10, 1903.

      _Erethizon epixanthum nigrescens_ Miller, U. S. Nat. Mus. Bull.,
      128:437, April 29, 1924.

      _Erethizon dorsatum nigrescens_ Anderson and Rand, Canadian Jour.
      Research, 21:293, September 24, 1943.

      _Type._--Obtained on the Shesley River, British Columbia, by M. P.
      Anderson on August 23, 1902; type in American Museum of Natural

      _Measurements._--A female from Sherman Creek Pass, Ferry County,
      measured: total length 770; length of tail 250; hind foot 95; ear
      37. A female from Tye, King County, measured: 930; 280; 125;
      weight 20 pounds.

      _Distribution._--Forested parts of the state, exclusive of the
      Blue Mountains.

_Remarks._--Porcupines are extremely rare west of the Cascades but are
occasionally reported from as far west as the Olympic Peninsula.

=Myocastor coypus= (Molina) subsp?

Coypu, nutria

      _Mus coypus_ Molina, Sagg. Stor. Nat. Chili, p. 287, 1782.

      _Myocastor coypus_ Kerr, Anim. Kingd., p. 225, 1792.

      _Type locality._--Chile.

_Description._--Size large, slightly smaller than a beaver; color rich,
reddish brown; tail long, round; hind feet webbed for swimming.

_Remarks._--The nutria, a native of South America, has been brought to
the United States and raised commercially on "fur farms." The species
has become established in the wild in several localities in western
Washington and at the Colville Indian Reservation in northeastern
Washington. For further details see Larrison (1943).

=Ochotona princeps= (Richardson)

Pika or cony

_Description._--The pika is of guinea-pig size, with a short, chunky
body about 200 mm. in length. The tail is represented externally merely
by a tuft of white fur. The short, wide head has large, circular ears,
large black eyes, and long whiskers. The legs are short and the soles of
the feet are furred. The color of the Washington races varies from
grayish-yellow to dark reddish brown. Like rabbits, all pikas have two
pairs of upper incisors. The second pair, located just in back of the
first, is small and delicate.

_Ochotona_ is a wide-ranging genus with many more species in Asia and
extreme eastern Europe than in North America. Three races of the species
_princeps_ occur in the state of Washington, where they are confined to
the Cascade, Kettle River, and Pend Oreille Mountains. The ranges of all
three races extend northward into British Columbia; one (_brunnescens_)
occurs also in Oregon, and one (_cuppes_) in Idaho.

In Washington pikas live only in talus slides and rock piles, where they
find refuge from most of the carnivores that prey on small mammals.
Their distribution in the state seems to be regulated by the
distribution of talus slides, and areas free of talus act as effective
barriers to pikas. They are abundant throughout the Cascades but are
absent from the Olympic Mountains although conditions there are well
suited to them. Probably the lowlands of western Washington which, owing
to moderate temperature and low relief, have little talus and exposed
rock, serve as a barrier. The Columbian Plateau is also free of pikas.
This may be due to the relative scarcity of talus as compared with
mountainous areas, and the fact that much of the talus on the Plateau is
composed of fragments of basalt too small to afford the shelter needed
by pikas. The aridity of the Columbian Plateau may contribute to the
absence of pikas, although this seems unlikely in view of the fact that
they occur on arid lands in Nevada and elsewhere.

Altitudinally, pikas range from 300 feet, in Clark County, to 6,000 feet
on Mt. Rainier, Pierce County, and on Round Top Mountain, Pend Oreille
County. They occur from the arid subdivision of the Transition
Life-zone, at Milk Creek, Kittitas County, to the upper edge of the
Hudsonian Life-zone, at Glacier Basin, Mt. Rainier. Generally speaking,
they are mammals of the mountains.

Common enemies of the pikas are the weasel (_Mustela frenata_), marten
(_Martes caurina_), and hawks of several species. Pikas are active by
day, especially in the early morning. Their call note is a short "eek!"
which carries a long distance. This squeaking note is often heard
throughout the night when rain threatens their drying hay.

  [Illustration: FIG. 126. Distribution of the pika in Washington. A.
  _Ochotona princeps brunnescens._ B. _Ochotona princeps fenisex._ C.
  _Ochotona princeps cuppes._]

Vegetation used as food, either for immediate consumption or for winter
use, includes almost all grasses, vines, shrubs, and trees available
near the pika's home. The subalpine lupines are especially favored. Even
such a thorny growth as the devil's club (_Oplopanax horridum_) is
eaten. Heather (_Phyllodoce, Cassiope_) has not been found in any of the
numerous hay piles examined, even when it is the commonest plant in the
vicinity. Large bundles of plants are carried in the pika's mouth. The
forefeet do not assist in transporting the load. If intended for
immediate consumption, the plants are deposited on one of last year's
hay piles and are eaten at leisure. The eating habits of the pika are
rabbitlike. A large leaf is seized at the tip and drawn into the mouth
with rapid chewing motions without assistance from the forefeet. Plants
destined to become hay are carefully spread out and exposed to the sun.
In cloudy or rainy weather the exposed plants are gathered and stored
under large rocks, to be reëxposed for curing when the weather improves.
Large hay piles often include more than fifty pounds of perfectly cured
grasses, annuals, bushes and evergreens.

No record of embryos is available for Washington pikas, but a male with
enlarged testes was taken at Lake Keechelus, Kittitas County, on March
22, 1940. Half-grown young of _fenisex_ were taken at Sawtooth Mountain,
Skamania County, on July 13, 1939, and of _brunnescens_ at Slate Creek,
Whatcom County, on August 16, 1937. Nearly full-grown young of _fenisex_
are reported taken at Bald Mountain, head of Ashnola River, Okanogan
County, on September 16, 1920. Young _cuppes_ of several sizes were
taken at Sherman Creek Pass, Ferry County, on September 11, 1938. The
breeding season possibly extends from March to August with a tendency to
be earlier at lower elevations.

=Ochotona princeps cuppes= Bangs

      _Ochotona cuppes_ Bangs, Proc. New England Zoöl. Club, 1:40, June
      5, 1899.

      _Ochotona princeps cuppes_ A. H. Howell, N. Amer. Fauna, 47:27,
      August 21, 1924.

      _Type._--Obtained by Allan Brooks at the Monashee Divide, 4,000
      feet, Gold Range, British Columbia, on August 2, 1897; type in
      Museum of Comparative Zoölogy.

      _Racial characters._--Smallest and palest of the Washington pikas;
      total length less than 8 inches; color of upper parts
      grayish-yellow, grayest on posterior third of back; underparts
      pale buff; skull small, but with relatively wide zygomatic and
      interorbital regions.

      _Measurements._--Eight males and 3 females from Round Top
      Mountain, Pend Oreille County, average, respectively: total length
      183.5, 181.5; hind foot 30.7, 31.0; ear 22.4, 23.7; one adult male
      from the same locality weighed 141.6 grams.

      _Distribution._--This pika has been found at Round Top Mountain
      (W.W.D.) and Pass Creek Pass (W.W.D.) in northeastern Washington.

=Ochotona princeps fenisex= Osgood

      _Lagomys minimus_ Lord, Proc. Zoöl. Soc. London, p. 98, 1863 (not
      of Schinz, 1821).

      _Ochotona minimus_ Bangs, Proc. New England Zoöl. Club, 1:39, June
      5, 1899.

      _Ochotona fenisex_ Osgood, Proc. Biol. Soc. Washington, 26:80,
      March 22, 1913 (substitute for _minimus_ Lord).

      _Ochotona princeps fenisex_ A. H. Howell, N. Amer. Fauna, 47:28,
      August 21, 1924.

      _Type._--Obtained by J. K. Lord at "Ptarmigan Hill," near head of
      Ashnola River, Cascade Range, British Columbia, in early fall of
      1860 (?); type in British Museum.

      _Racial characters._--Size and color intermediate between
      _brunnescens_ and _cuppes_; length about 8 inches; color of upper
      parts near Pinkish Cinnamon, becoming gray on posterior third of
      back; underparts washed with buff; skull of medium size and

      _Measurements._--Nine males and 5 females from Okanogan and Chelan
      counties average, respectively: total length 190.0, 197.4; hind
      foot 31.1, 32.8; ear 21.6, 22.0.

      _Distribution._--Specimens referable to this race occur from the
      British Columbian boundary south, through the eastern Cascade
      Mountains. Along the western border of its range, _fenisex_
      becomes larger and darker, merging into the race _brunnescens_.
      Marginal records are: Hidden Lakes (U.S.N.M.), Lyman Lake
      (U.S.N.M.), Mt. Stuart (W.W.D.), Easton (U.S.N.M.), Mt. Aix
      (U.S.N.M.), Steamboat Mt. (M.V.Z.).

=Ochotona princeps brunnescens= Howell

      _Ochotona fenisex brunnescens_ A. H. Howell, Proc. Biol. Soc.
      Washington, 32:108, May 20, 1919.

      _Ochotona princeps brunnescens_ A. H. Howell, N. Amer. Fauna.
      47:31, August 21, 1924.

      _Type._--Obtained by George G. Cantwell at Lake Keechelus, Kittitas
      County, Washington, on August 23, 1917; type in United States
      National Museum.

      _Racial characters._--Largest and darkest of the three races of
      Washington pikas; total length 8 inches or more; color of upper
      parts rich cinnamon, heavily washed with blackish; posterior part
      of back slightly paler; underparts buffy cinnamon; skull large and
      heavy with wide zygomatic arches but relatively narrow
      interorbital region and relatively narrow across maxillary tooth

      _Measurements._--Eight males and 9 females from within three miles
      of Stevens Pass, King and Chelan counties, average, respectively:
      total length 201.8, 208.3; hind foot 33.3, 33.0; ear 22.3, 22.6.
      The average weights of 4 males and 3 females from 3 mi. S E Tumtum
      Mountain, Clark County, are 178.0 and 174.3 grams, respectively.

      _Distribution._--From the British Columbia boundary south to the
      Columbia River and from the western Cascades east to the area of
      intergradation with _fenisex_. Marginal occurrences are: Whatcom
      Pass (U.S.N.M.), Stevens Pass (W.W.D.), Keechelus (U.S.N.M.),
      Cowlitz Pass (U.S.N.M.), Tumtum Mountain (M.V.Z.).

=Lepus townsendii townsendii= Bachman

White-tailed jack rabbit

      _Lepus townsendii_ Bachman, Jour. Acad. Nat. Sci. Philadelphia, 8
      (pt. 1):90, pl. 2, 1839.

      _Lepus campestris townsendi_ Merriam, Proc. Biol. Soc. Washington,
      17:132, July 14, 1904.

      _Lepus townsendii townsendii_ Hollister, Proc. Biol. Soc.
      Washington, 28:70, March 12, 1915.

      _Type._--Obtained by J. K. Townsend at old Fort Walla Walla
      (present town of Wallula), Walla Walla County, Washington; type in
      Philadelphia Academy of Natural Sciences.

      _Measurements._--A male from Miller's Island, Klickitat County, in
      the Columbia River, measured: total length 564; length of tail
      117; hind foot 156; ear 110; weight 337.5 grams.

      _Distribution._--Previously found over the grasslands of eastern
      Washington. Now restricted and scarce except in the Okanogan

_Remarks._--The white-tailed jack rabbit is the largest rabbit in the
state, adults measuring 24 or more inches in length. The long legs and
long ears accentuate the impression of large size. Its body is more
bulky than that of its relative, the black-tailed jack rabbit. In summer
the pelage of the upper parts is dark gray and in winter it is white
over nearly the entire body.

The white-tailed jack rabbit occurs from southern Saskatchewan south to
extreme northern New Mexico, and from eastern Washington east to
Wisconsin. A single race occurs in Washington. In eastern Washington
"whitetails" favor the hilly, bunchgrass territory of the arid
subdivision of the Transition and Upper Sonoran life-zones. In winter
they descend to the lower sagebrush valleys.

The principal enemies of the white-tailed jack rabbit are the eagle,
coyote, and bobcat. Of 1,186 stomachs of coyotes from Washington, Sperry
(1941: 11) found that 27 percent contained rabbits, including jack
rabbits, snowshoe hares, and cottontails.

In the daytime, white-tailed jack rabbits hide in forms which consist of
shallow holes dug at the bases of bushes or beside rocks. They feed in
the morning, evening, and in the night along wide, well-defined trails
through the bunchgrass. If startled from their forms they dash off in
bounding, erratic leaps, skimming away until lost to sight. A whitetail
has been timed at a speed of 34 miles per hour (Cottam and Williams,
1943: 262).

The early explorers and settlers found the white-tailed jack rabbits
abundant in eastern Washington. With the invasion and spread of the
black-tailed jack rabbit, and the reduction of native bunchgrass through
overgrazing by livestock, the whitetail has become rare. In several
years of field work on the Columbian Plateau, I saw none. Near Wallula,
the type locality, residents had not seen whitetails for years, but
thought there might be a few left "back in the hills." There are thought
to be a few left near Ellensburg and Yakima.

Only in the Okanogan Valley are the whitetails holding their own; they
are reasonably common there. In winter they come down from the hills on
to the sagebrush flats along the Okanogan River in Okanogan County. In
January it is not unusual to see as many as five in a day's drive. When,
as will most certainly occur, the black-tailed jack rabbit enters the
Okanogan Valley, the splendid whitetail may be expected to disappear
from Washington.

Because this species has been so reduced in numbers, no distributional
map has been included. Taylor and Shaw (1929: 28) give its range as:
"north to Oroville, east to Pullman, south to Asotin, Walla Walla, and
Kennewick, and west to Lake Chelan (Manson), Yakima Valley, and
Klickitat County." This range is similar to that of the Nuttall
Cottontail (Fig. 129).

=Lepus americanus= Erxleben

Snowshoe rabbit

_Description._--The appearance, size, and proportions of the snowshoe
rabbit are similar to those of the Belgian hare. The body is about 16
inches in length, the ears are midway in size between those of the
cottontail and the jack rabbit, and the feet are relatively long and the
tail is short. In summer the color of the upper parts is reddish brown,
varying with the subspecies. The winter pelage of _Lepus a.
washingtonii_ is a slightly paler brown than the summer coat. In the
other three races in Washington the winter coat is entirely white,
except for the dusky borders of the ears.

Snowshoe rabbits occur in Alaska, Canada, and the northern United
States, from the Atlantic to the Pacific. They are absent in desert or
prairie regions but range far southward in the United States in
mountainous areas. They are found throughout Washington, except on the
Columbian Plateau and in the Okanogan River Valley. None of the four
races found in Washington is restricted exclusively to the state.
Snowshoe rabbits live only in wooded areas. Their habitat varies from
dense, impenetrable rain-forests along the ocean to the alpine parks,
dotted with trees, of the Hudsonian Life-zone. They occur in humid and
arid subdivisions of the Transition, Canadian, and Hudsonian life-zones.
Altitudinally they range from sea level to 6,000 feet (Mt. Rainier).

Enemies of the snowshoe rabbit include the coyote, bobcat, lynx,
long-tailed weasel, and great horned owl.

Snowshoe rabbits are largely nocturnal or crepuscular in habit. They are
secretive and slip away quietly at the least threat of danger. Persons
often live for years in localities where snowshoe rabbits are abundant
without seeing a live individual. Those that are seen ordinarily have
been startled from their forms at midday, or surprised while feeding on
clover along a highway in the early morning. More commonly they are seen
crossing a road in the lights of an automobile. Tracks, easily found
after a fresh snowfall, give some indication of their numbers in any

Little information is available on the fluctuations of numbers of
snowshoe rabbits in Washington. Floyd Thornton, a trapper living at
Forks, Clallam County, states that they were numerous in 1924, scarce in
1930-31, and fairly common in 1938-39. More rabbits are seen shortly
after the breeding season than at other times of the year. From April
8-10, 1941, I saw none on a highway extending about 100 miles along the
west coast of the Olympic Peninsula, but on June 4-5 here counted 3 dead
on the road and saw 3 running across it. One was about one-third grown
and another two-thirds grown.

  [Illustration: FIG. 127. Distribution of the snowshoe hare in
  Washington. A. _Lepus americanus washingtonii._ B. _Lepus americanus
  cascadensis._ C. _Lepus americanus columbiensis._ D. _Lepus americanus

The snowshoe rabbits are classed as game animals in Washington but few
people hunt them. Their winter food includes buds and needles of
hemlock, Douglas fir, and probably other evergreens. Annuals, grasses,
and shrubs, as well as Douglas fir needles, are eaten in the summer.
Snowshoe rabbits do some damage by eating the bark of trees and the
boughs of newly planted evergreens. Together with rodents they are
responsible for serious damage to plantations of Douglas fir and hemlock
on the Olympic Peninsula.

At least in summer, they are heavily parasitized by fleas and ticks and
may in addition carry tularemia, or rabbit fever.

Scheffer (1933: 77-78) found that the young were born from May 5 to July
4 in the Puget Sound area and that there were from 2 to 5, usually 5 per

=Lepus americanus washingtonii= Baird

      _Lepus washingtonii_ Baird, Proc. Acad. Nat. Sci. Philadelphia,
      7:333, 1855.

      [_Lepus americanus_] var. _Washingtoni_ J. A. Allen, Proc. Boston
      Soc. Nat. Hist., 17:434, February 17, 1875.

      _Lepus americanus Washingtoni_ True, Proc. U. S. Nat. Mus., 7:601,

      _Type._--Obtained by G. Suckley at Steilacoom, Pierce County,
      Washington, on April 1, 1854; type in United States National

      _Racial characters._--Size small for a snowshoe rabbit; color of
      upper parts dark, in summer between Sayal Brown and Cinnamon, in
      winter slightly paler, near Pale Cinnamon Buff (capitalized color
      terms in the accounts of the lagomorphs are after Ridgway, Color
      Standards and Color Nomenclature, Washington, D. C., 1912);
      underparts white; soles of feet usually stained yellowish,
      brownish, or blackish.

      _Measurements._--Two males and 6 females from the Olympic
      Peninsula average, respectively: total length 407, 402.5; hind
      foot 114, 119. Weight of a 408 mm. male from the same locality
      2-1/4 lbs.

      _Distribution._--This snowshoe rabbit occupies the humid
      subdivision of the Transition Life-zone of western Washington from
      the British Columbian boundary south to the Columbia River.
      Marginal occurrences are (from Dalquest, 1942: 175): Mt. Vernon,
      Paradise Lake, Lake Kapowsin, and White Salmon.

=Lepus americanus cascadensis= Nelson

      _Lepus bairdi cascadensis_ Nelson, Proc. Biol. Soc. Washington,
      20:87, December 11, 1907.

      _Lepus americanus cascadensis_ Racey and Cowan, Ann. Rept.
      Provincial Mus. British Columbia, p. H 18, 1935.

      _Type._--Obtained by W. C. Colt near Hope, British Columbia, on
      June 12, 1894; type in Museum of Comparative Zoölogy.

      _Racial characters._--Size medium for a snowshoe rabbit; color of
      upper parts in summer near Orange Cinnamon; head paler, sharply
      marked off from body; underparts white. Color in winter: entire
      body pure white except for dusky borders of ears and eyelids.

      _Measurements._--Three males and 5 females from Kittitas County,
      Washington, average, respectively: total length 405, 440; hind
      foot 124, 133.

      _Distribution._--The Cascade Mountains from the British Columbia
      boundary south to Mount Adams. Marginal occurrences are: Skykomish
      (Dalquest, 1942: 177), Vance (Dalquest, 1942: 177) and 7 mi. W
      Guler (W.W.D.).

=Lepus americanus pineus= Dalquest

      _Lepus americanus pineus_ Dalquest, Jour. Mamm., 23:178, May 14,

      _Type._--Obtained by P. G. Putnam at Cedar Mountain (now Moscow
      Mountain), Latah County, Idaho, on May 29, 1921; type in University
      of Michigan, Museum of Zoölogy.

      _Racial characters._--Size small for a snowshoe rabbit; in summer,
      upper parts Cinnamon Brown, with sides slightly paler and rump
      patch blackish; color of the head between Sayal Brown and
      Cinnamon; hips Light Olivaceous Buff; chest-band light Cinnamon;
      ears blackish, often edged with white. In winter the color of the
      entire body is white, save for the dusky edges of the ears and the
      blackish eyelids. The color of the underfur in winter is usually
      Pale Pinkish Cinnamon, rarely Light Vinaceous Cinnamon or Orange
      Cinnamon, with the basal portion slaty.

      _Measurements._--Two males and 5 females, from northeastern
      Washington, average, respectively: total length 419, 439; hind foot
      138.5, 138. The average measurements of 3 males and 7 females from
      the Blue Mountains are: 407, 422; 128, 131.

      _Distribution._--The pine forests of the arid subdivision of the
      Transition Life-zone along the eastern border of Washington, in
      the Blue Mountains, and in the forested parts of northeastern
      Washington as far west as the Kettle River Range. Western records
      of occurrence are (Dalquest, 1942: 179): Deep Lake, Colville,
      Calispel Peak.

=Lepus americanus columbiensis= Rhoads

      _Lepus americanus columbiensis_ Rhoads, Proc. Acad. Nat. Sci.
      Philadelphia, p. 242, June, 1895.

      _Type._--Obtained by S. N. Rhoads at Vernon, British Columbia, on
      July 29, 1892; type in Philadelphia Academy of Natural Sciences.

      _Racial characters._--Size large; color in summer: upper parts
      near Cinnamon Brown, sides and head slightly paler; top of tail
      and small rump-patch blackish; chest-band pale Cinnamon Brown;
      hips light Ochraceous Buff; chin and belly white.

      _Measurements._--A male from Molson, Okanogan County, measures:
      total length 435; length of tail 150. A male and a female from
      Danville, Ferry County, measure, respectively: 460, 430; 150, 142.

      _Distribution._--Only a small part of the range of this rabbit
      lies within the state of Washington; the greater part is in
      British Columbia. In Washington it occupies the timbered areas
      north of the Columbia River, east of the Okanogan River, and west
      of the Kettle River Range. Records are (Dalquest, 1942: 182):
      Molson, Danville and Republic. In the 1942 paper, the latter
      locality was erroneously listed under _pineus_.

=Lepus californicus deserticola= Mearns

Black-tailed jack rabbit

      _Lepus texianus deserticola_ Mearns, Proc. U. S. Nat. Mus., 18:564,
      June 24, 1896.

      _Lepus texianus wallawalla_ Merriam, Proc. Biol. Soc. Washington,
      17:137, July 14, 1904 (type from Touchet, Walla Walla County,

      _Lepus californicus deserticola_ Nelson, N. Amer. Fauna, 29:137,
      August 31, 1909.

      _Type._--Obtained at western edge of Colorado desert, Imperial
      County, California; type in American Museum of Natural History.

      _Measurements._--Three males and 4 females from Union Gap, Yakima
      County, average, respectively: total length 538, 539; length of
      tail 72, 71; hind foot 128, 124; ear 120, 124. Two females from
      the same locality weighed 5 and 6-1/2 pounds, respectively.

      _Distribution._--The Columbian Plateau, southeastern Washington,
      and the Yakima Valley area. The northernmost locality record is
      Moses Coulee (W.W.D.).

_Remarks._--The blacktail is the commonest jack rabbit of the West. It
is larger than the domestic rabbit and its enormous ears, long, gangling
legs, and bounding gait make it appear even larger than it really is.
Its iron-gray color, black tail and black ear tips match its sagebrush

  [Illustration: FIG. 128. Distribution of the black-tailed jack rabbit,
  _Lepus californicus deserticola_, in Washington.]

Black-tailed jack rabbits reach the northern limit of their distribution
on the Columbian Plateau of eastern Washington. They extend from
Washington south to the Valley of Mexico and from the Pacific Coast east
to Missouri (Nelson, 1909: 127). About 20 races are recognized, of which
only one is native to Washington.

The blacktail is restricted to the Upper Sonoran Life-zone. Seldom is it
found far from sagebrush (_Artemisia_) and rabbitbrush (_Chrysothamus_).
It is well adapted to desert life, and is able to withstand the bitterly
cold winters and hot, dry summers of eastern Washington. Blacktails are
active in the evening, night, morning, and cooler parts of the day. In
the daytime they crouch in forms consisting of shallow depressions at
the bases of shrubs.

Blacktails eat the twigs and leaves of sagebrush, rabbitbrush, other
desert shrubs, and grasses. They are particularly fond of alfalfa and
cultivated crops, of which they destroy great quantities. They are
serious pests during periodic years of abundance. Some measure of their
abundance may be gained by counting the bodies of jack rabbits killed by
cars on well-traveled highways through sagebrush areas. In years of
abundance these may number 50 to 100 per mile, while in years of
scarcity these may number only one or two.

Black-tailed jack rabbits are susceptible to numerous parasites and
diseases including tularemia. Diseased rabbits are especially noticeable
during years of abundance. Jack rabbits are almost never used as food
although many are shot for sport or to protect crops.

The black-tailed jack rabbit is a swift runner. Cottam and Williams
(1943: 263) timed 6 individuals while running under varying conditions.
Full speeds for 50 to 300 yards varied from 27 to 38 miles per hour. The
maximum speed was attained by two individuals, each for 100 yards.

Embryos found in March numbered 4, 5 and 6.

=Sylvilagus nuttallii nuttallii= (Bachman)

Nuttall cottontail

      _Lepus nuttallii_ Bachman, Jour. Acad. Nat. Sci. Philadelphia,
      7:345, 1837.

      _Lepus artemisia_ Bachman, Jour. Acad. Nat. Sci. Philadelphia,
      8:94, 1839 (type from Wallula, Walla Walla County, Washington).

      [_Lepus sylvaticus_] var. _Nuttallii_ Allen, Proc. Boston Soc.
      Nat. Hist., 17:434, February 17, 1875.

      _Lepus sylvaticus Nuttalli_ True, Proc. U. S. Nat. Mus., 7:601,

      _Sylvilagus (Sylvilagus) nuttallii_ Lyon, Smithsonian Misc. Coll.,
      45 (no. 1456):336, June 15, 1904.

      _Sylvilagus nuttalli_ Nelson, N. A. Fauna, 29:201, August 31,

      _Type._--Obtained near mouth of Malheur River, Malheur County,
      Oregon. by T. Nuttall in August, 1834; type in Philadelphia Academy
      of Natural Sciences.

      _Measurements._--Two males and 4 females from Moses Lake, Grant
      County, average, respectively: total length 355, 348; length of
      tail 33, 35; hind foot 83, 83; ear 63.5, 60.

      _Distribution._--The Columbian Plateau, Okanogan Valley, Yakima
      Valley and Columbia Valley in southeastern Washington; in general,
      the sagebrush area of eastern Washington; north in the Okanogan
      River Valley to Oroville and in the Columbia Valley to Kettle
      Falls (W.W.D., records not all shown on map).

_Remarks._--Nuttall cottontails are small, grayish-brown rabbits with
relatively short, rounded ears and short legs. Their small size and
small ears, which lack black tips, distinguish them from jack rabbits
where the two occur together.

  [Illustration: FIG. 129. Distribution of the Nuttall cottontail.
  _Sylvilagus nuttallii nuttallii_, in Washington.]

The genus _Sylvilagus_ is found in both North and South America. The
species _nuttallii_, with three subspecies, is restricted to the western
part of the United States. It ranges from southern Canada south to
central New Mexico and from western South Dakota west to the Cascades. A
single race occurs in Washington. Nuttall cottontails depend on cover
for concealment from enemies. They frequent thick stands of tall
sagebrush, riparian thickets, or rocky coulees. Seldom are they
encountered in the open. In the sand-dune areas near Moses Lake
cottontails were abundant in the dense, thorny thickets about potholes
and in areas of tall sagebrush. They are especially common near the
talus at the bases of the walls of Grand Coulee, and Moses Coulee where
they do not hesitate to enter crevices in rock slides for protection.
Indeed, cottontails are abundant everywhere within their range in the
state of Washington, where suitable cover and food are present. They
seem to be confined to the Upper Sonoran Life-zone.

Cottontails are most active at night, as is attested by the number
killed then by automobiles on highways. The greatest number are seen by
observers in the morning and evening but it is not unusual to see an
animal feeding at midday. When startled they dash for the nearest
thicket or pile of rocks with their tiny white tails erect. They travel
in relatively straight lines and do not dash from side to side in flight
as do jack rabbits. They sometimes seek concealment by "freezing"
motionlessly in plain sight. When feeding undisturbed they travel by
slow hops.

The trails of cottontails are characteristic of thickets in sagebrush
country. The trails are narrow, less than four inches wide, and often
enter thickets of strong, thorny growths which can scarcely be
penetrated by man. Near Okanogan Lake the trails of cottontails were
found among greasewood bushes on hard-packed gravel. Trails are usually
most abundant in thickets near water. Permanent trails are not made
through low sagebrush or over sandy areas where the animals prefer to
pick their way when traveling from one clump of cover to another.

Nuttall cottontails probably eat many desert grasses, annuals, and
shrubs; observation indicates that sagebrush (_Artemisiae tridentata_)
and rabbit brush (_Chrysothamnus nauseosus_) are particularly important
as food.

The young of cottontails in Washington seem to vary from one to four per
litter and are born between April and June.

=Sylvilagus floridanus= (Allen) subsp.?

Florida cottontail

      _Lepus sylvaticus floridanus_ Allen, Bull. Amer. Mus. Nat. Hist.,
      3:160, October 8, 1890.

      _Sylvilagus floridanus_ Lyon, Smithsonian Misc. Coll., 45 (no.
      1456):322, June 15, 1904.

_Description._--Slightly larger than the sagebrush cottontail; smaller
than the snowshoe rabbit; ears small; color of upper parts pinkish
cinnamon-brown; sides pale grayish-cinnamon; underparts white; nape of
neck cinnamon; chest band paler cinnamon; tail brown above, white
beneath; forefeet and sides of hind feet cinnamon.

_Remarks._--The Florida cottontail is not native to the state of
Washington but has been introduced at several localities. It is
spreading rapidly at the present time. Points of introduction include
Pullman, Whitman County (1926-1927); Battleground, Clark County (1933);
Auburn, King County (1927); and Whidby Island, Island County (1931). At
least two subspecies (_mearnsi_, _alacer_) have been introduced and a
third (_similis_) may have been introduced.

For further information on introduced cottontails see Dalquest (1941B:

=Sylvilagus idahoensis= (Merriam)

Pigmy rabbit

      _Lepus idahoensis_ Merriam. N. Amer. Fauna, 5:75, July 30, 1891.

      _Brachylagus idahoensis_ Lyon, Smithsonian Misc. Coll., 45 (no.
      1456):323, June 15, 1904.

      _Sylvilagus idahoensis_ Grinnell, Dixon and Linsdale, Univ.
      California Publ. Zoöl., 35:553, October 10, 1930.

      _Type._--Obtained in the Pahsimeroi Valley, Custer County, Idaho,
      by V. Bailey and B. H. Dutcher on September 16, 1890; type in
       United States National Museum.

      _Distribution._--Known only from the central part of the Columbian

_Description._--The pigmy rabbit is a tiny species, differing from the
cottontail in smaller size, paler, grayer color, shorter ears and
smaller legs.

The pigmy rabbit is restricted to the Great Basin region. No subspecies
has been described. It is rare and of local occurrence in Washington,
having been recorded only from the central part of the Columbian
Plateau. Orr (1940), who studied the species in California, found them
only in stands of tall, dense sage (_Artemisiae tridentata_). It is a
burrowing form, not straying far from its hole.

=Cervus canadensis= (Erxleben)

Elk or wapiti

_Description._--The elk, next to the moose, is our largest deer. The
legs of the elk are slender. The tail is a short, pointed stub a few
inches long. The neck is thick in proportion to the head. Both males and
females possess the canine teeth familiar as "elk tooth charms." Only
the males possess antlers. These are huge, slender beams that curve up,
out and back with the basal tine or "dog killer" and four to six points
on each antler. The antlers are deciduous and are shed annually. The
body is grayish or tan in color. The head, neck, chest and legs are
rich, dark brown, strongly contrasting with the paler body. The
distinctive rump patch is pale tan or white.

In the past the elk was found over most of the forested areas of
Washington. Lumbering, agriculture and settlement as well as excessive
hunting removed it from parts of eastern Washington and all except the
most inaccessible parts of the lowlands of western Washington. Only in
the Cascade and Olympic Mountains and the coastal strip between the
Columbia River and the Olympic Mountains, did the elk survive in
appreciable numbers. Conservationists and a more enlightened game policy
began to protect the elk at the turn of the century. It was already too
late to save the species in eastern Washington, where it seems never to
have been truly abundant and where relatively open country afforded
little protection from the high-powered rifle. In the dense, rugged
forests of western Washington a sizable number remained on the Olympic
Peninsula and these, under protection, increased to their present
numbers. The dense, tangled forests of the southwestern coastal area and
the western Cascades lack conditions suitable to support truly large elk
herds. These areas probably now have as large an elk population as can
safely be supported and fed.

The elk of eastern Washington had disappeared or nearly disappeared by
1910. The race originally occurring there was the Rocky Mountain form;
it has been reintroduced from Montana and Wyoming into northeastern
Washington and the Blue Mountains area. These plantings have not been
very successful. Introduced in the eastern Cascades, however, the Rocky
Mountain elk thrived and increased on what was probably once the
peripheral range of the coastal elk.

The habits of the elk are best known from the herds in the Olympic
Mountains and on the Rattlesnake Game Refuge in the eastern Cascade
Mountains. Here the animals are numerous and relatively tame. Their
habits seem to differ somewhat in the forests of the lowlands from
those of the animals in the higher Olympics where the topography and
climate are very different.

  [Illustration: FIG. 130. Elk or wapiti (_Cervus canadensis nelsoni_),
  Banff, Alberta, October, 1939. (G. A. Thomas photo.)]

The elk is a social animal, gathering in herds over most of the year.
The old males leave the herds in the spring but seem to stay in small
bands while their antlers are growing. In the Olympic Mountains, herds
of 100 or more animals have been seen. In the lowlands there are
ordinarily from five to ten in a herd.

  [Illustration: FIG. 131. Group of elk or wapiti (_Cervus canadensis
  nelsoni_), Banff, Alberta, October 10, 1939. (G. A. Thomas photo.)]

The elk is a browsing animal, feeding on twigs and leaves of deciduous
trees, shrubs and evergreens. In spring and summer it eats grasses and
succulent annuals, but in winter twigs and needles of evergreens,
perennial ferns, dry grass and even moss is utilized. To a certain
extent the elk are migratory, ascending to the open meadows of the
Hudsonian Life-zone in the early summer and returning to the dense
forests of the Transition and lower Canadian Life-zones with the winter
snows. The lowland elk make no such migrations, merely leaving the
riverbottom jungles when the leaves are off the deciduous plants
growing there, and live in the coniferous timber.

In the forests the elk is capable of swift and almost silent movement.
It is an eerie experience to trail a herd of elk through a dense forest
in a winter rain, knowing that a number of the large animals are within
a few feet, moving swiftly but silently away. When a herd is feeding and
does not suspect the presence of an observer, the animals rustle
branches, break twigs, snort and wheeze as they breathe.

The antlers of the males are shed in March. The bulls retire from the
herd until the new horns are well grown, in late August or September.
Breeding takes place in September or October and the young are born the
following April or May.

=Cervus canadensis roosevelti= Merriam

      _Cervus roosevelti_ Merriam, Proc. Biol. Soc. Washington, 11:272,
      December 17, 1897.

      _Cervus canadensis occidentalis_ Taylor and Shaw, Occ. Papers
      Chas. R. Conner Mus., 2:29, December, 1929.

      _Type._--Obtained on Mount Elaine, Mason County, Washington, by H.
      and C. Emmet on October 4, 1897; type in United States National

      _Racial characters._--A large, dark elk with short, heavy antlers.

      _Measurements._--The type measured, in the flesh (Bailey,
      1936:81): total length 2,490; tail 80; ear (dry) 208.

      _Distribution._--From the Cascade Mountains westward; possibly
      occurred formerly in the eastern Cascades, and perhaps still
      present there, in places, or mixed with introduced _nelsoni_.

      _Remarks._--For use of the name _roosevelti_ rather than
      _occidentalis_ see Bailey (1936:81).

=Cervus canadensis nelsoni= Bailey

      _Cervus canadensis canadensis_ Taylor and Shaw, Occ. Papers Chas.
      R. Conner Mus., 2:29, December, 1929.

      _Cervus canadensis nelsoni_ Bailey, Proc. Biol. Sec. Washington,
      48:188, November 15, 1935.

      _Type._--Obtained at Yellowstone National Park, Wyoming, by J.
      Pitcher; died in captivity on September 21, 1904; type in United
      States National Museum.

      _Description._--Smaller and paler than _roosevelti_ with longer,
      slimmer antlers.

      _Distribution._--Formerly occurred in northeastern Washington and
      the Blue Mountains of southeastern Washington. Since then
      exterminated and reintroduced to both areas from the Rocky
      Mountains. Also introduced and established in the eastern

=Odocoileus virginianus= (Boddaert)

White-tailed deer

_Description._--The white-tailed deer differs from the black-tailed and
mule deer in several anatomical details. The tail is long, wide, and
when the animal is frightened is carried upright with the long, white
hair of the underside spread out. The antlers of the buck are not of a
biramous system of branching but instead consist of a main beam which
curves sharply out and forward, remaining low. All subsidiary tines,
save the basal one, emerge from the dorsal side of the main beam. The
basal tine is usually a spurlike point arising vertically from the main
beam not far from the base of the antler. In exceptional specimens the
basal tine is large and divided into several points. The body of the
"whitetail" is more smoothly rounded, and more graceful, than that of
the mule deer.

  [Illustration: FIG. 132. White-tailed deer (_Odocoileus virginianus
  leucurus_), doe at least nine years old, kept as a pet by Mrs. Jack
  Hovis, Puget Island, Washington, December 16, 1939. (Fish and Wildlife
  Service photo by Victor B. Scheffer, No. 750.)]

White-tailed deer range from Canada south to Panama and from the
Atlantic to the Pacific. Their range is much more extensive to the east
and south than that of the "blacktail" and mule deer. In western
Washington the whitetail occupies a limited habitat near the mouth of
the Columbia River. Its geographic range was probably somewhat greater
in the past. Its habitat includes the low, damp, marshy islands and
floodplain of the Columbia. Blacktail range in the wooded hills
surrounding the whitetail's range but rarely enter it to compete with
the whitetail (Scheffer, 1940A: 282).

In northeastern Washington the whitetail shows the same tendency to
occupy low, marshy areas and valleys. The habitat preference of the
whitetail in northeastern Washington is less exact than in western
Washington, for the species ranges up from the valleys into the forests
of larch and cottonwood of the Transition Life-zone. Its habitat
includes denser forest and brushy areas. Rarely does it occur in the
open type of forest occupied by the mule deer.

  [Illustration: FIG. 133. White-tailed deer (_Odocoileus virginianus
  ochrourus_): fawn kept as a pet by L. E. Borud, Ione, Washington, June
  13, 1938. (Fish and Wildlife Service photo by Victor B. Scheffer, No.

In Washington the whitetail is not an important game animal. The race
occurring in western Washington is characterized by small size and small
antlers. Further, its total population is estimated at but 400 to 900
individuals. The whitetail of northeastern Washington is larger and
possesses larger antlers. Indeed, some antlers from Ferry County are the
largest antlers of whitetail that I have ever seen. Nevertheless, the
whitetail of northeastern Washington is smaller than the mule deer and
far less common. The habitat is dense and difficult to traverse. The
animal is shy and silent, fleeing soundlessly when approached. For these
reasons most hunters in northeastern Washington prefer to hunt the mule

  [Illustration: FIG. 134. White-tailed deer (_Odocoileus virginianus
  ochrourus_): left antler found on ground at Park Rapids, Pend Oreille
  County, Washington, October 1, 1937. (Fish and Wildlife Service photo
  by Victor B. Scheffer, No. 756.)]

=Odocoileus virginianus ochrourus= Bailey

      _Odocoileus virginianus macrourus_ Taylor and Shaw, Occ. Papers
      Chas. R. Conner Mus., 2:30, December, 1929.

      _Odocoileus virginianus ochrourus_ Bailey, Proc. Biol. Soc.
      Washington, 45:43, April 2, 1932.

  [Illustration: FIG. 135. White-tailed deer (_Odocoileus virginianus
  leucurus_), antlers of large buck killed in the fall of 1939 at
  Cathlamet, Washington, by Paul Lewis. (Fish and Wildlife Service photo
  by Victor B. Scheffer. No. 752.)]

      _Type._--Obtained at Coolin, south end of Priest Lake, Bonner
      County, Idaho, by F. Lemmer on December 27, 1908; type in United
      States National Museum.

      _Racial characters._--A large, pale white tail with large, low
      antlers and distinctive cranial features.

      _Measurements._--The type, an adult male, measures (Bailey, 1936.
      p. 43): total length 1,752; tail 265; hind foot 483; ear (dry)

      _Distribution._--Northeastern Washington. Probably once occurred
      in southeastern Washington.

=Odocoileus virginianus leucurus= (Douglas)

      _Cervus leucurus_ Douglas, Zoöl. Jour., 4:330, 1829.

      _Odocoileus leucurus_ Thompson, Forest and Stream, 51:286, October
      8, 1898.

      _Odocoileus virginianus leucurus_ Lydekker, Cat. Ung. Mamm.
      British Mus., 4:162, 1915.

      _Type._--Obtained on the North Umpqua River, Oregon, by David
      Douglas on October 17, 1826; type originally, and perhaps still, in
      British Museum.

      _Racial characters._--A small, handsome whitetail with slender,
      erect antlers and brownish or grayish color.

      _Measurements._--A young adult female from 1 mile south of
      Skamokawa, Wahkiakum County, measured: total length 1,545; length
      of tail 250; hind foot 458; ear 143; weight 88 pounds.

      _Distribution._--Previously occurred over much of southwestern
      Washington; now confined to the banks and islands of the Columbia
      River in Wahkiakum County.

=Odocoileus hemionus= (Rafinesque)

Mule deer and black-tailed deer

_Description._--The mule and black-tailed deer are among deer of medium
to large size. Adult mule deer may weigh up to 400 pounds while some
fully adult blacktail bucks may weigh as little as 100 pounds. The body
is heavily muscled, the legs long and the tail only about 6 inches in
length. The ears are long, from 6 to 8 inches from notch to tip. The
head is long and the male has well-developed antlers. The first antlers
are almost straight spikes. Those of the second year are bent slightly
outward and forward and are equally or subequally forked with the
anterior branch usually the stouter. In the third year the
anteriolateral curvature is more pronounced and one or both of the forks
again fork. In succeeding years forks are larger and more numerous but
the essential biramous arrangement of forking is maintained. The
"blacktail" and mule deer are strongly marked races. The blacktail
differs from the mule deer in: smaller size; darker color, especially on
face and tail; tail dark brown above with dark tip rather than whitish
with black tip; tail haired beneath, not naked for half its length;
antlers smaller and lighter; and skull and teeth smaller.

Mule deer and black-tailed deer range over western North America from
southeastern Alaska southward into northern Mexico. They inhabit
forested parts of the state of Washington. Blacktails occupy the San
Juan Islands, the islands in Puget Sound, the Olympic Mountains, the
lowlands of western Washington, and the Cascade Mountains. Mule deer
occupy the Cascades, including their eastern slope, northeastern and
southeastern Washington and parts of the Columbian Plateau. Over this
large range there is considerable local geographic variation.

Jackson (1944: 1-56) estimated that 109,600 blacktail and 175,725 mule
deer live in Washington. Thus Washington is second only to California
in number of blacktail and ranks fifth in number of mule deer.

Individual variation over the range of the mule deer is considerable
but no trends of variation are distinguishable. Mule deer from the
Blue Mountains, northeastern Washington and the eastern Cascades are
essentially similar.

There is geographic variation as well as great individual variation
in the black-tailed deer of Washington. The deer of the San Juan
Islands and the islands of Puget Sound are smaller and darker than
those of the mainland and possess smaller, lighter antlers. The deer
of Whidby Island are sometimes contemptuously referred to by residents
as jackrabbit deer. Fully grown bucks on the Islands weigh in the
neighborhood of 100 pounds, rarely exceeding 150 pounds, whereas bucks
on the mainland commonly weigh more than 150 pounds dressed.

  [Illustration: FIG. 136. Mule deer (_Odocoileus hemionus hemionus_),
  subadult male from Okanogan County, Washington, raised in captivity;
  photographed June 29, 1938, on Hurricane Ridge, Olympic National Park.
  (Fish and Wildlife Service photo by Victor B. Scheffer, No. 99.)]

The dividing line between the ranges of the black-tailed and mule deer
in general is the summit of the Cascade Mountains. In summer the two
races come together and doubtless mingle in the Cascades but as fall
approaches they migrate. The mule deer moves eastward to the yellow-pine
areas on the lower slopes while the blacktails descend westward to the
denser Douglas fir and hemlock forests on the western flanks of the
Cascades. In the breeding season the two races are separated. However,
as shown by Cowan (1936A: 219), in the Pemberton Valley, British
Columbia, the two forms occur together during the breeding season and
intergradation occurs there. Intergradation occurs also in the Lake
Wenatchee area of Chelan County. Observation of hunters' specimens
showed some undoubted intergrades among the more abundant, typical mule

Both the mule deer and blacktail have a characteristic bounding gait,
unlike the smooth run of the white-tailed deer. The tail is usually held
down, rarely erect. When not frightened they move with dainty steps,
making little noise.

  [Illustration: FIG. 137. Black-tailed deer (_Odocoileus hemionus
  columbianus_), buck resting at midday, Van Trump Park, Mt. Rainier,
  Washington, August, 1931. (Fish and Wildlife Service photo by Victor
  B. Scheffer, No. 260.)]

The black-tailed deer in certain areas is more nocturnal than diurnal.
They feed principally in the early morning and evening. In the Olympic
Mountains, on the North Fork of the Quinault River, in July, 1937,
blacktails were watched feeding and playing until midnight. Also in
December, 1939, deer, near the forks of the Skykomish River, were moving
about, browsing as late as 11 p. m. while a light snow was falling. In
the San Juan Islands, in the summer of 1939, deer were observed feeding
at all hours of the day.

The blacktail lives in some of the most dense jungle areas of western
Washington. On Whidby Island and some of the San Juan Islands the brush
and vines grow so densely that a man can scarcely penetrate them. Were
it not for the deer trails, formed by generations of deer, our small
mammal investigations would have been far more difficult. The blacktail
lives also in the dense forests where fir, hemlock and cedars grow to
heights of more than 150 feet. Here lack of light allows only ferns and
moss to form an understory vegetation. These forests often clothe the
steep glacial hills and the trails of the deer on such hills show them
to be adept climbers. Deer trails generally avoid fallen trees and other
obstacles. When startled, however, a stump, fence or log is easily
leaped by a blacktail. Often they are in small bands of 6 to 10
individuals, but almost as often are solitary, or in pairs.

The habitat of the mule deer is generally more open than that of the
blacktail. In the summer, it may occupy rough and rugged country; in the
higher Cascades individuals were found in the rocky and brushy country
and in open glades and meadows. Farther east they were in the open
yellow-pine forests where extensive grassy slopes, free of trees,
existed. In some parts of northeastern Washington they lived in the
larch and lodgepole pine forests almost dense enough to be "blacktail
country." In the Okanogan Valley and on the northwestern corner of the
Columbia Plateau mule deer lived in open prairie country where a few
cottonwoods and willows were the only trees.

The mule deer seem more social than the blacktail. In the late summer
they gather in bands of 10 to 20 or more. In winter, under pressure of
hunger, they gather in herds and raid haystacks and pastures. Farmers in
the Methow Valley, Okanogan County, report herds of 200 to 400 mule deer
about a single haystack.

The mule deer and blacktail are the principal game mammals of the state
of Washington. Thousands of hunters enter the woods each year in search
of a buck. Eastern Washington is the favored hunting grounds for that is
the home of the mule deer, the larger size of which makes it a more
desirable trophy. Also the open country which it inhabits makes hunting
more productive. Sums spent on equipment, gasoline and hunting licenses
are tremendous but the feeling is that the return in recreational value
and venison are worth the cost. A few casualties result each year among
the army of deer hunters.

=Odocoileus hemionus hemionus= (Rafinesque)

      _Cervus hemionus_ Rafinesque, Amer. Month. Mag., 1:436, October,

      _Cariacus macrotis_ True, Proc. U. S. Nat. Mus., 7:592, 1885.

      _Odocoileus hemionus_ Merriam, Proc. Biol. Soc. Washington,
      12:100, April 30, 1898.

      _Odocoileus hemionus macrotis_ Bailey, National Geographic, 20:64,

      _Type locality._--Vicinity of Big Sioux River, South Dakota.

      _Racial characters._--Large size, pale color.

      _Measurements._--Two adult males from Stay-a-while Spring,
      Columbia County, measure respectively: total length 1,751, 1,559;
      length of tail 172, 205; hind foot 515, 485; ear 210, 211; length
      of metatarsal gland 150, 135.

      _Distribution._--Summit of Cascades eastward, in forested areas,
      exact limits not certainly known.

=Odocoileus hemionus columbianus= (Richardson)

      _Cervus macrotis_ var. _columbiana_ Richardson, Fauna
      Boreali-Americana, 1:257, 1829.

      _Cariacus columbianus_ True, Proc. U. S. Nat. Mus., 7 (1884):592,

      _Cervus columbianus_ Baird, U. S. Pacific R. R. Exp. and Surveys,
      p. 659, 1857.

      _Eucervus columbianus_ Gray, Ann. and Mag. Nat. Hist., 18 (ser.
      3):338, 1866.

      _Odocoileus columbianus_ Merriam, Proc. Biol. Soc. Washington,
      12:100, April 30, 1898.

      _Odocoileus columbianus columbianus_ Swarth, Univ. California
      Publ. Zoöl., 10:85, February 13, 1912.

      _Odocoileus hemionus columbianus_ Cowan, California Fish and Game,
      22:215, July, 1936.

      _Type._--Obtained at Cape Disappointment, Pacific County,
      Washington, by Lewis and Clark on November 19, 1805 (Cowan, 1936A:

      _Racial characters._--Small size, dark color.

      _Distribution._--Summit of Cascades westward, including islands in
      Puget Sound and San Juan Islands; exact limits of range uncertain.

=Alce americana shirasi= Nelson


      _Alces americanus shirasi_ Nelson, Proc. Biol. Soc. Washington,
      27:72, April 25, 1914.

      _Type._--Obtained near the Snake River, Lincoln County, Wyoming, by
      J. Shire on December 11, 1913; type in United States National

      _Distribution._--Casual wanderer into northeastern Washington from

_Remarks._--The moose is the largest North American deer. Its large size
and huge, palmate antlers serve to separate it from any other member of
the deer tribe.

Moose range from northern United States to central Canada and Alaska.
The European elk and the American moose are subspecies of the same
species. The moose is of only casual occurrence in Washington. The
latest authentic record for Washington is a bull that wandered southward
and westward from Canada until it was killed on an Indian reservation in
Ferry County. Other moose are reported to have wandered into
northeastern Washington from Canada in past years.

Dice (1919) was told that moose once occurred in the Blue Mountains of
southeastern Washington but there is no confirmation of this report.

=Rangifer arcticus montanus= Seton-Thompson


      _Rangifer montanus_ Seton-Thompson, Ottawa Naturalist, 13:129-30,
      August, 1899.

      _Rangifer arcticus montanus_ Jacobi, Erganzungsband, Zoöl. Anz.,
      96:92, November, 1931.

      _Type._--Obtained near Revelstoke, Selkirk Range, British Columbia.

      _Distribution._--Rare or casual along the Canadian boundary in
      northeastern Washington.

_Remarks._--The caribou is a rather stout-bodied deer with large hoofs,
short, rounded muzzle and long, erect, flattened antlers.

Caribou and their relatives, the reindeer, range over Arctic Europe,
Asia, Greenland and America. In North America they range from the
Atlantic to the Pacific and from the northern border of the United
States northward into the Arctic. The caribou was, until recent years, a
regular winter resident in small numbers in northeastern Washington near
the Canadian Boundary. Their wintering grounds in Washington were said
to have been destroyed by fire in 1915 and the species has appeared in
the state only casually since then. Two were killed in 1940 by hunters
who thought they were deer. Caribou are protected by law in Washington.

=Bison bison oregonus= Bailey


      _Bison bison bison_ Taylor and Shaw, Occ. Papers Chas. R. Conner
      Mus., 2:31, December, 1929.

      _Bison bison oregonus_ Bailey, Proc. Biol. Soc. Washington, 45:48,
      April 2, 1932.

      _Type._--Skull and skeleton picked up at Malheur Lake, Oregon, by
      G. M. Benson in November, 1931; type in United States National

      _Distribution._--Perhaps casual in eastern Washington before
      coming of the white man.

_Remarks._--Bison, or buffalo, occurred in southeastern Oregon but
disappeared before white men reached the country (Bailey, 1936: 57).
Gibbs (1860: 138) was told by an Indian hunter in 1853 that a lost bull
had been killed in the Grand Coulee (state of Washington) 25 years
before but that "this was an extraordinary occurrence, perhaps before
unknown." In the days before horses reached the Indian tribes of eastern
Washington and Oregon, wandering bison from herds in Oregon probably
strayed into Washington, in somewhat the manner that the moose today
stray in from Canada.

=Ovis canadensis= Shaw

Mountain sheep

_Description._--The mountain sheep is the size of a small deer. The
horns of the males are massive spirals. Those of the females are
smaller, curve upward and backward, rarely forming a semicircle. The
horns are permanent structures, enlarged each year by growth at the
base. Growth is irregular, probably as the result of seasonal
reproductive activities. As a result the horns are "ringed" with
concentric ridges. The color of the upper parts is dark, grayish brown.
The face is paler, yellowish brown. The outer sides of the legs are dark
brown. The rump, abdomen, and insides of legs are white.

Mountain sheep of the genus _Ovis_ are abundantly represented in Asia.
Two species occur in North America, _Ovis dalli_ in Alaska, and _Ovis
canadensis_ in western North America. They range from Alaska south to
northern Mexico.

In the past the mountain sheep inhabited most of the eastern Cascade
Mountains, the Blue Mountains, Pend Oreille Mountains, and the cliffs of
the Columbia River Valley in eastern Washington. They occurred on the
eastern or Columbian Plateau side of the river and therefore probably
occupied the cliffs of Moses Coulee and the Grand Coulee. Their habitat
seems thus to have included rocky areas from the Upper Sonoran to the
Hudsonian life-zones. At the present time they are extinct over most of
their range. A small band still remains in the extreme northeastern
Cascades near Mount Chopaka.

Little has been published concerning the habits of the mountain sheep in
Washington. In caves along the Columbia River in Grant County, bones of
sheep are found in association with stone arrowheads and other human
artifacts. Presumably the sheep were killed and eaten by the Indians.

The history of the mountain sheep in North America is outlined by Cowan
(1940: 506). The genus is thought to have crossed from Asia to America
by the land bridge now under Bering Strait in the early Pleistocene and
spread southward through the Rockies. The advance of the glacial ice
forced them farther southward and the southern (_canadensis_) sheep were
separated from their relatives farther north. The present differences
between the Rocky Mountain and western sheep seem to have resulted from
separation by glaciers during Wisconsin Time.

=Ovis canadensis canadensis= Shaw

      _Ovis canadensis_ Shaw, Nat. Misc., vol. 15, text to pl. 610, about
      December, 1803.

      _Type._--Obtained in the mountains on Bow River; W. B. Davis (1939:
      377) gives Dew River near Exshaw, Alberta.

      _Racial characters._--Large size; heavy, closely coiled horns.

      _Measurements._--Cowan (1940: 533) gives the measurements of a
      fully adult (6-year old) ram from Colorado as: total length 1,953;
      length of tail 127; hind foot 394; ear (dry) 63.

      _Distribution._--Now extinct in Washington. Cowan (_op. cit._:
      535) refers to this race as the sheep that formerly occurred in
      the Blue Mountains. This view seems logical but I feel that the
      sheep formerly occurring in the Pend Oreille Mountains of extreme
      northeastern Washington should, on the basis of geographic
      probability, also be referred to _canadensis_.

=Ovis canadensis californiana= Douglas

      _Ovis californianus_ Douglas, Zoöl. Jour., 4:332, 1829.

      _Ovis californica_ Richardson, Fauna Boreali-Americana, 1:272,

      _Ovis californiana_ Biddulph, Proc. Zoöl. Soc. London, p. 683,

      _Ovis canadensis californiana_ Lydekker, The Sheep and its
      Cousins, p. 288, 1912.

      _Type._--Obtained from near Mount Adams, Yakima County, Washington,
      by D. Douglas on August 27, 1826; type in British Museum.

      _Racial characters._--Similar to _canadensis_ but smaller with
      more slender, spreading horns and horn tips less blunt.

      _Measurements._--Cowan (1940: 545) gives the measurements of a
      ram, five years old, from Owens Valley, California, as: total
      length, 1,582; length of tail, 110; hind foot, 240.

      _Distribution._--Formerly occupied the eastern Cascades, the
      valley of the Columbia and possibly the cliffs bordering Moses
      Coulee and The Grand Coulee. Now it is found only near Mt. Chopaka
      in the extreme northeastern Cascades.

=Oreamnos americanus= (Blainville)

Mountain goat

_Description._--The mountain goat is of deer-size, with a stout body and
a pronounced hump on the shoulders. The legs are short but the hoofs are
large with large dewclaws. The neck is short and thick. The head is
large and goatlike in appearance. The tail is tiny. The horns are slim,
round and curve up and slightly backwards. They are hollow and are
permanent structures, added to each year. The body is snow white,
consisting of long, soft wool, which is longer and coarser on forelegs,
neck and chin than on the body. Males have a distinct beard.

Mountain goats range from Washington and Idaho north to Alaska. Their
nearest relatives are the Old-World antelopes, especially the alpine
species of Europe and Asia.

The mountain goat is an animal of the high mountains. Their habitat is
the bare-rock cliffs and rock-strewn slopes of the Arctic-alpine and
Hudsonian Life-zones. Where extensive, open rocky areas occur they
descend to the Canadian Life-zone. Even in winter they keep to the high
cliffs where steep slopes and strong winds keep the snow from the plants
on which they feed.

Mountain goats are considered a game animal in Canada and Idaho. The
species has been protected in Washington for many years. As a result
they are common, although not present in numbers sufficient to withstand
hunting. The high country which they occupy is unsuited to any of man's
domestic animals and no reason is apparent at present why the mountain
goat should not be protected and conserved for many years.

  [Illustration: FIG. 138. Mountain goat (_Oreamnos americanus
  americanus_), old female, Lake Chelan, Washington, March 6, 1937.
  (Forest Service photo by Oliver T. Edwards, No. 348491.)]

In spite of the protection accorded them, the mountain goat has not
become tame. In driving over Stevens Pass, King County, one can, with
the aid of glasses, usually pick out one or more mountain goats on
nearby mountains. Nevertheless, I have never seen one within 100 yards
of a highway, nor heard of one being killed by cars on a highway.

The mountain goat does not, as a rule, allow close approach by man. At
Goat Basin, Deception Creek, King County, circumstances are particularly
favorable for goats and several are usually to be seen. While studying
them on several occasions, I was never able to get truly close to them.
On the few occasions when I came upon a band unexpectedly, they rapidly
went over the mountain or up the nearest cliff.

  [Illustration: FIG. 139. Group of mountain goats (_Oreamnos americanus
  americanus_), northern Cascade Mountains, Washington. (Forest Service
  photo, No. 348490.)]

When unfrightened, these mountain goats spent much time standing, or
lying with forefeet folded under them, looking into space. For such
large animals they seemed to spend little time feeding. At intervals of
about five minutes they plucked the ferns or other plants that grew in
abundance on ledges or in tiny crevices in the cliffs. Some animals did
browse for several minutes at a time. They seemed rather particular as
to their food, plucking only one or two stems from a clump of

The ordinary movements of the mountain goats are deliberate. They rarely
move more than a few feet at a time. They climb with sure-footed ease
but usually slowly. In climbing, the forelegs are spread and the knees
are bent. The animal moves as if it were climbing steps. The greatest
action is in the knee joints. If surprised in the open they run with a
smoother gait, the legs moving from the shoulder and hip. If near a
cliff they climb rapidly, jumping when necessary, and rarely stop while
in sight. Evidence of the presence of mountain goats is usually seen in
the form of tufts and strands of wool. Their wool becomes caught in
bushes and rough rocks and seemingly pulls out easily.

The range of the mountain goat in Washington includes the entire Cascade
range from Mt. Adams and Mt. Saint Helens north to the Canadian
boundary. They extend west to Mt. Baker, Mt. Higgins and Mt. Index and
east to Lake Chelan and, in the Wenatchee Range, east of Mt. Stuart,
perhaps almost to the Columbia River.

Dice (1919: 21) was told that mountain goats once occurred in the Blue
Mountains of southeastern Washington. We know of no suitable goat
country in the Blue Mountains of Washington and feel that the report was
probably based on an erroneous identification.

There is a single record, supported by specimen, of a mountain goat from
northeastern Washington. This individual probably wandered into
Washington from northern Idaho, Montana, or adjacent British Columbia.

=Oreamnos americanus americanus= (Blainville)

      _Ovis montanus_ Ord, Guthrie's Geol., 2d Amer. Ed., p. 292, 1815

      _R[upicapra]. americana_ Blainville, Bull. Sci. Soc. Philomath,
      Paris, p. 80, 1816.

      _Mazama dorsata_ Rafinesque, Amer. Month. Mag., 2:44, 1817 (new
      name for _Ovis montanus_ Ord).

      _Mazama montana_ True, Proc. U. S. Nat. Mus., 7 (1884):592, 1885.

      _Oreamnos montanus_ Merriam, Science, n. s., 1:19, 1895.

      _Oreamnos montanus montanus_ Miller, U. S. Nat. Mus. Bull.,
      79:398, December 31, 1912.

      _Oreamnos americanus americanus_ Hollister, Proc. Biol. Soc.
      Washington, 25:186, December 24, 1912.

      _Type._--None. Descriptions all based on _Ovis montanus_ Ord who,
      in turn, based his description on the account of skins seen by
      Lewis and Clark on the Columbia River of Washington or Oregon. As
      the mountain goat is not known to have ever occurred in Oregon in
      Recent times, the type locality is probably near Mt. Adams,
      Washington, the point where goats come nearest the Columbia.

      _Measurements._--The measurements of a large male killed "west of
      North Yakima" were reported by A. S. Harmer as: 8 feet 3 inches
      from tip of nose to tip of tail; horns 10 inches; weight 507
      pounds (Outdoor Life, 1915: 459).

      _Distribution._--Throughout the higher Cascade Mountains.

=Oreamnos americanus missoulae= Allen

      _Oreamnos montanus missoulae_ Allen, Bull. Amer. Mus. Nat. Hist.,
      20:20, February 10, 1904.

      _Oreamnos americanus missoulae_ Hollister, Proc. Biol. Soc.
      Washington, 25:186, December 24, 1912.

      _Type._--Obtained at Missoula, Missoula County, Montana; type in
      American Museum of Natural History.

      _Distribution._--A single record for northeastern Washington.
      Seemingly a rare wanderer from outside the state.


Whales and porpoises

Because Victor B. Scheffer and John W. Slipp have in preparation a
detailed account of Cetaceans properly ascribable to Washington, members
of this order here are not treated in as much detail as are other native

=Berardius bairdii= Stejneger

Baird beaked whale

_Description._--Length about 40 feet; mandibles elongate, "beak-like";
one or two large teeth at tip of lower jaw; dorsal fin small and
situated posteriorly; color black but abdomen occasionally grayish.

_Remarks._--This rare whale is known from a few specimens recorded from
Alaska south to California.

=Mesoplodon stejnegeri= True

Stejneger beaked whale

_Description._--Similar to the previous species in proportions but
smaller; length about 17 feet. Differs in possessing one large flat tusk
in lower jaw.

_Remarks._--Known only from a specimen from Alaska and another from the
coast of Oregon.

=Delphinus bairdii= Dall


_Description._--A slender porpoise about 6 or 7 feet long; long, narrow
beak with 80-120 conical teeth; color above black tinged with greenish;
sides gray; belly and throat white.

_Remarks._--Recorded from California and probably ranges into Washington
in off-shore waters.

=Lissodelphis borealis= (Peale)

Right whale porpoise

_Description._--A small, beaked porpoise lacking a dorsal fin; color
black with narrow white area from breast to tail; length about 4 feet.

_Remarks._--A rare species described from waters 500 miles off the mouth
of the Columbia River.

=Lagenorhynchus obliquidens= Gill

Striped porpoise

_Description._--Length about 7 feet; beak not prominent; 44-46 conical
teeth in each jaw; anterior edge of dorsal fin curved; color of upper
parts greenish black; sides with one white or gray stripe posteriorly;
underparts white.

_Remarks._--This is the common porpoise in the Straits of Juan De Fuca
from about the vicinity of Port Townsend to the ocean and in immediate
off-shore waters. One specimen was obtained 100 miles off Grays Harbor.
Rarely seen in Puget Sound.

=Grampus rectipinna= (Cope)

Killer whale

_Description._--A large porpoise, 20-30 feet in length; dorsal fin high
and nearly straight; teeth large, conical, 12 above, 13 below; color
black above with white patch on side anteriorly.

_Remarks._--The killer is common in Puget Sound and off the ocean coast.
Both the killer and the true blackfish are locally termed "blackfish."

=Grampidelphis griseus= (Cuvier)


_Description._--A round-headed porpoise about 10 feet in length; dorsal
fin high and narrow; 4 to 6 teeth in each jaw; color black, occasionally
with white head or with body mottled with white and gray.

_Remarks._--An off-shore species, probably not uncommon off the coast of
Washington at times.

=Globicephalus scammonii= (Cope)


_Description._--A large, stout-bodied, round-headed porpoise about 20
feet in length. Teeth large; 8-10 in lower jaw; 10-12 in upper jaw.
Color black. Differs from the killer whale in possessing a low, long
dorsal fin rather than a high, erect one.

_Remarks._--The blackfish is a colonial species, often common in Puget
Sound. It is frequently confused with the killer whale.

=Phocoena vomerina= (Gill)

Harbor porpoise

_Description._--A small, stout-bodied porpoise with a rounded head.
Length 5 to 6 feet. Teeth small, slightly flattened; about 25 above, 24
below; color blackish or brownish to liver color.

_Remarks._--Small schools of harbor porpoises are common in Puget Sound,
among the San Juan Islands, and in the Straits of Juan De Fuca. This is
the commonest inshore porpoise but, because of its smaller size, is less
well known to most persons than are the blackfish and killer whale.

=Phocoenoides dalli= (True)

Dall porpoise

_Description._--A small porpoise, about 6 feet in length, with pointed
head and moderately stout body. Teeth small, 23 above and 27 below.
Color greenish black with flanks or posterior sides white.

_Remarks._--A rare off-shore species.

=Physeter catodon= Linnaeus

Sperm whale

_Description._--A toothed whale about 60 feet long; head enormous,
squarish; teeth conical, 50 or fewer in lower jaw.

_Remarks._--Occurs off the coast of Washington and rarely enters the
Straits of Juan De Fuca.

=Kogia breviceps= (Blainville)

Pigmy sperm whale

_Description._--Similar to the sperm whale in proportions but less than
10 feet in length.

_Remarks._--A rare species whose habits are little known. The available
information concerning this species in the Atlantic has been summarized
by Allen (1941).

=Rhachianectes glaucus= (Cope)

Gray whale

_Description._--A relatively slender whale, about 40 feet in length;
dorsal fin absent; baleen short, yellow in color; color of body mottled
gray or blackish.

_Remarks._--The gray whale was once abundant and occurred off the coast
of Washington in spring and summer migrations. Hunted until now rare.

=Balaenoptera physalus= (Linnaeus)

Finback whale

_Description._--A relatively slender whale, up to 82 feet in length;
dorsal fin far posterior, prominent; baleen in two slabs, well
developed, about two feet in length and gray in color; color of body
blackish or brownish above, white below.

_Remarks._--Recorded off-shore.

=Balaenoptera borealis= Lesson

Sei or Pollack whale

_Description._--Similar to the finback whale but dorsal fin larger;
baleen dark; body dark blue or brownish; belly with restricted white

_Remarks._--Recorded off-shore.

=Balaenoptera acutorostrata= Lacépède

Pike whale

_Description._--A small whalebone whale, about 30 feet in length; body
slender; head pointed; pectoral fin well developed and prominent; baleen
short, white; color black above, white beneath.

_Remarks._--A rare species that has been recorded from Washington.

=Sibbaldus musculus= (Linnaeus)

Blue whale or sulphur-bottom whale

_Description._--Largest of the whales; length up to 100 feet; body
slender; head flat above, rounded beneath; dorsal fin slender but
prominent; baleen in two series, heavy and black; belly with numerous
longitudinal ridges; color bluish black above, yellow beneath.

_Remarks._--Probably uncommon off-shore.

=Megaptera novaeangliae= (Borowski)

Humpback whale

_Description._--A medium-sized, stout-bodied whale up to 54 feet in
length; dorsal fin low, not prominent; head flat above, rounded beneath;
prominent "hump" at back of head; belly with numerous longitudinal
ridges; baleen small; color blackish or grayish above, paler below; body
characteristically blotched with patches of whitish barnacles.

_Remarks._--The humpback is the commonest whale off the coast of
Washington, often coming into northern Puget Sound.

=Eubalaena sieboldii= (Gray)

Pacific right whale

_Description._--A large whale, 60-70 feet in length, with enormous,
rounded head; dorsal fin absent; belly lacking longitudinal ridges;
baleen blackish, 8 feet in length; color uniformly blackish.

_Remarks._--A much hunted species now probably rare off shore.



      1. _Ursus canadensis_ Merriam, grizzly bear.--This species has been
      recorded from Kootenay Lake, British Columbia, 30 miles northeast
      from the northeastern corner of the state of Washington and may
      have occurred in northeastern Washington.

      2. _Ursus idahoensis_ Merriam, grizzly bear.--Recorded by Merriam
      (1918: 54) from the Wallowa Mountains, Oregon. This form may have
      occurred in the Blue Mountains of southeastern Washington.

      3. _Vulpes fulva macroura_ Baird, red fox.--Reported from the Blue
      Mountains and northeastern Washington; no specimen recorded.

      4. _Canis lupus columbianus_ Goldman, wolf.--Possibly occurred in
      northeastern Washington in historic time, and perhaps is
      occasionally still found there.

      5. _Canis lupus irremotus_ Goldman, wolf.--Perhaps once occurred
      in southeastern Washington.

      6. _Mirounga angustirostris_ (Gill), sea elephant.--May occur as a
      casual wanderer off the coast of Washington. The home of this
      species is Lower California but a dead specimen was washed upon
      the shore of Prince of Wales Island, Alaska (Willett, 1943: 500).

      7. _Perognathus parvus laingi_ Anderson, Great Basin pocket
      mouse.--Probably present in the mountains east of Lake Osoyoos in
      Washington, but no specimens have yet been collected.

      8. _Synaptomys borealis artemisiae_ Anderson, northern lemming
      mouse.--Probably present in the Cascades of northern Okanogan
      County but no specimens have yet been obtained.

      9. _Antilocapra americana_ (Ord) subsp?, pronghorned
      antelope.--This species is restricted to North America and once
      ranged over much of the plains region of the western part of the
      continent. Taylor and Shaw (1929: 31) included the antelope in
      their list of Washington mammals with the statement "Now
      extirpated within State; the former range of the pronghorn
      included much of the plains country of eastern Washington." So far
      as known to me, there is no record by any of the early explorers
      of antelope killed or seen in what is now Washington. No bones of
      antelope have been discovered in caves in eastern Washington.
      Ogden (1909: 339) mentions an antelope killed in Oregon a day's
      journey south of The Dalles. This record is fairly close to
      Washington and indicates that antelope might have occurred at
      least in southeastern Washington in historic times.


  1. The antelope (_Antilocapra americana_) has been introduced into the
  Yakima Valley by the state of Washington, Department of Game.

  2. The elephant seal (_Mirounga angustirostris_) has been discovered off
  the coast of Washington (Seattle _Times_, April 15, 1947, p. 3).

  3. The yellow-pine chipmunk of the Blue Mountains has been described as
  a new subspecies, _Eutamias [Tamias] amoenus albiventris_ Booth,
  Murrelet, 28 (no. 1):7, 1947. Type locality Wickiup Spring, 23 miles
  west of Anatone, Asotin-Garfield County boundary.


  ALLEN, G. M.

      1920. Dogs of the American aborigines. Harvard Col., Bull. Mus.
      Comp. Zoöl., 63:431-517, 12 pls.

      1941. Pigmy sperm whale in the Atlantic. Field Mus. Nat. Hist.,
      zoöl. ser., 27:17-36, 4 figs. in text.

  ALLEN, J. A.

      1893. On a collection of mammals from the San Pedro Martin region
      of Lower California, with notes on other species, particularly of
      the genus _Sitomys_. Bull. Amer. Mus. Nat. Hist., 5:181-202.


      1932. Five new mammals from British Columbia. Ann. Rept. Canadian
      Nat. Mus. for 1931, pp. 99-119, 1 pl.

  ANDERSON, R. M., and RAND, A. L.

      1943A. Variation in the porcupine (genus _Erethizon_) in Canada.
      Canadian Jour. Research, 21:292-309, 5 figs. in text.

      1943B. Status of the Richardson vole (_Microtus richardsoni_) in
      Canada. Canadian Field-Nat., 57:106-107.


      1900. Revision of the American voles of the genus _Microtus_. N.
      Amer. Fauna, 17:1-88, 5 pls., 17 figs. in text.

      1918. Wild animals of Glacier National Park: the mammals. U. S.
      Nat Park Serv. Bull., pp. 15-102, 21 pls., 18 figs. in text.

      1936. The mammals and life zones of Oregon. N. Amer. Fauna,
      55:1-416, 51 pls., 102 figs. in text.

  BAIRD. S. F.

      1857. Mammals. Gen. Rept., Zoölogy of the Several Pacific R. R.
      Routes, pp. xxv-xxxii, 1-737; xlviii + 757 pp., pls. xvii-lx, 35
      figs. in text.

  BENNETT, L. J., ENGLISH, P. F., and WATTS, R. L.

      1943. The food habits of the black bear in Pennsylvania. Jour.
      Mamm., 24:25-31, 1 fig. in text.

  BERRY, E. W.

      1931. A Miocene flora from Grand Coulee, Washington. _In_ U. S.
      Geol. Surv. Prof. Paper 170, pp. 31-42, pls. 11-13.


      1912. Some Common mammals of western Montana in relation to
      agriculture and spotted fever. U. S. Dept. Agric., Farm. Bull. 484,
      pp. 1-46, 34 figs. in text.


      1942. Records of harbor seals in lakes Washington and Union,
      Seattle. Murrelet, 23:76.

  BOOTH, E. S.

      1945. A new red-backed mouse from Washington state. Murrelet,

  BRETZ, J. H.

      1913. Glaciation of the Puget Sound region. Washington Geol. Surv.
      Bull. 8, pp. 1-244, 24 pls., 27 figs. in text.

      1923. Glacial drainage on the Columbian Plateau. Bull. Geol. Soc.
      Amer. vol. 34, pp. 573-608. 12 figs. in text.


      1939. Items of interest from Mount Rainier. Murrelet, 20:70-71.


      1939. Food habits of the vagrant shrew. Murrelet, 20:62-66.


      1927. The "Palouse soil" problem. _In_ U. S. Geol. Surv. Bull. 790,
      pp. 21-45, pls. 4-7.


      1945. Phylogeny of nearctic Sciuridae. Amer. Mid. Nat., vol. 33,
      pp. 257-390, 8 pls., 48 figs. in text.


      1943. Speed of some wild mammals. Jour. Mamm., 24:262.

  COUCH, L. K.

      1925. Storing habits of Microtus townsendii. Jour. Mamm.,

      1927. Migration of the Washington black-tailed jack rabbit. Jour.
      Mamm., 8:313-314.

      1930. Notes on the pallid yellow-bellied marmot. Murrelet, 11 (No.
      2):2-6, 3 figs.

  COWAN, I. M.

      1936A. Distribution and variation in deer (genus _Odocoileus_) of
      the Pacific coastal region of North America. California Fish and
      Game, 22:155-246. figs. 51-63.

      1936B. Nesting habits of the flying squirrel Glaucomys sabrinus.
      Jour. Mamm., 17:58-60.

      1937. A new race of _Peromyscus maniculatus_ from British
      Columbia. Proc. Biol. Soc. Washington, 50:215-216.

      1938. Geographic distribution of color phases of the red fox and
      black bear in the Pacific Northwest. Jour. Mamm., 19:202-206, 1

      1939. The vertebrate fauna of the Peace River district of British
      Columbia. Occ. Papers British Columbia Prov. Mus., No. 1, pp.

      1940. Distribution and variation in the native sheep of North
      America. Amer. Mid. Nat., 24:505-580, 4 pls., 1 map.

  COWAN, I. M., and HATTER, J.

      1940. Two mammals new to the known fauna of British Columbia.
      Murrelet, 21:9.

  CRABB, W. D.

      1944. Growth, development, and seasonal weights of spotted skunks.
      Jour. Mamm., vol. 25. pp. 213-221, 2 pls.


      1936. The geology of Washington: part 1, General features of
      Washington geology (with map). Washington State Div. Geol., Bull.
      32, pp. 1-70.

  DALE, F. H.

      1940. Geographic variation in the meadow mouse in British Columbia
      and southeastern Alaska. Jour. Mamm., 21:332-340.


      1938. Bats in the state of Washington. Jour. Mamm., 19:211-213.

      1940. Bats in the San Juan Islands, Washington. Murrelet, 21:4-5.

      1941A. Ecologic relationships of four small mammals in western
      Washington. Jour. Mamm., 22:170-173.

      1941B. Distribution of cottontail rabbits in Washington State.
      Jour. Wildlife Management, 5:408-411, 1 fig.

      1942. Geographic variation in northwestern snowshoe hares. Jour.
      Mamm., 23:166-183, 2 figs. in text.

      1943. Seasonal distribution of the hoary bat along the Pacific
      Coast. Murrelet, 24:20-24, 2 figs.

      1944. The moulting of the wandering shrew. Jour. Mamm.,


      1941. The shrew-mole of western Washington. Murrelet, 22:12-14.

  DALQUEST, W. W., and ORCUTT, D. R.

      1942. The biology of the least shrew-mole, Neurotrichus gibbsii
      minor. Amer. Mid. Nat., 27:387-401, 4 figs. in text.


      1942. The origin of the mima mounds of western Washington. Jour.
      Geol., 50:68-84, 8 figs. in text.

      1944. Distribution and variation in pocket gophers, _Thomomys
      talpoides_, in the state of Washington. Amer. Nat., part 1,
      78:308-333, part 2, 78:423-550, 8 figs. in text.

  DAVIS, W. B.

      1939. The Recent Mammals of Idaho. The Caxton Printers, Caldwell,
      Idaho, pp. 1-400.

  DICE, L. R.

      1919. The mammals of southeastern Washington. Jour. Mamm., 1:10-22,
      2 pls.

      1939. Variation in the deer-mouse (_Peromyscus maniculatus_) of
      the Columbia Basin of southeastern Washington and adjacent Idaho
      and Oregon. Contrib. Lab. Vert. Genetics, Univ. Michigan, 12:1-22,
      1 map.

      1940. Speciation in Peromyscus. Amer. Nat., 74:289-298.


      1937. Genetics and the origin of species. Columbia Univ. Press, New
      York, pp. xvi + 364.

  DOUTT, J. K.

      1942. A review of the genus _Phoca_. Ann. Carnegie Mus., 29:61-125,
      14 pls., 11 figs. in text.

  EDGE, E. R.

      1934. Burrows and burrowing habits of the Douglas ground squirrel.
      Jour. Mamm., 15:189-193, 1 pl., 1 fig. in text.

  EDSON, J. M.

      1916. Wild animals of Mount Baker. Mountaineer, 9:51-57.

      1933. A visitation of weasels. Murrelet, 14:76-77.

      1935. Yellow-bellied marmot out of bounds. Jour. Mamm., 16:68.


      1940-1941. The families and genera of living rodents. Jarrold and
      sons, Ltd., Norwich, vol. 1 (1940), xxvi + 689, 189 figs. in text;
      vol. 2 (1941) xii + 690, 49 figs, in text.


      1899. Catalogue of mammals from the Olympic Mountains, Washington,
      with descriptions of new species. Field Columb. Mus. Publ., 32,
      zoöl. ser., 1:241-276, pls. 41-62, numerous figs. in text.


      1936. Distribution of races of the brown bat (_Eptesicus_) in
      western North America. Amer. Mid. Nat., 17:653-660, 1 fig. in text.


      1943. Carnivorous activities of big brown and pallid bats. Jour.
      Mamm., 24:96-97.


      1930. Mammals of Austin Pass, Mount Baker. Mazama, 12:34-43,


      1919. With the birds and animals of Rainier. Mazama, 5:319-326,

      1925. Cougar kills a boy. Jour. Mamm., 6:197-199.


      1939. Habits of the southern sea otter. Jour. Mamm., 20:21-36.


      1939. Unusual location of hibernating jumping mice. Murrelet,
      20:17-18, 1 fig.

  FLINT, R. F.

      1935. Glacial features of the southern Okanogan region. Bull. Geol.
      Soc. Amer., vol. 46, pp. 169-194, pls. 13-18, 2 figs. in text.

      1937. Pleistocene drift border in eastern Washington. Bull. Geol.
      Soc. Amer., vol. 48, pp. 203-232, 5 pls., 1 fig. in text.

      1938. Summary of late-Cenozoic geology of southeastern Washington.
      Amer. Jour. Science, ser. 5, vol. 35, pp. 223-230.


      1922. The occurrence of tularaemia in nature as a disease of man.
      U. S. Publ. Health Serv., Hyg. Lab. Bull. 130:1-8.


      1906. The exploration of Samwel Cave. Amer. Jour. Sci., ser. 4,
      22:235-247, 3 figs. in text.


      See Suckley, G., and Gibbs, G. 1860.

  GIDLEY, J. W., and GAZIN, C. L.

      1933. New Mammalia in the Pleistocene fauna from Cumberland Cave.
      Jour. Mamm., 14:343-357.

      1938. The Pleistocene vertebrate fauna from Cumberland Cave,
      Maryland. U. S. Nat. Mus. Bull. 171, pp. i-vi + 1-99, pls. 1-10,
      50 figs.


      1941. Remarks on voles of the genus _Lemmiscus_, with one described
      as new. Proc. Biol. Soc. Washington, vol. 54, pp. 69-71.

      1943. Two new races of the puma. Jour. Mamm., 24:228-231.

  GRAY, J. A. JR.

      1943. Rodent populations in the sagebrush desert of the Yakima
      Valley, Washington. Jour. Mamm., 24:191-193.


      1918. Sea-lion question in British Columbia. Contr. Canadian Biol.,
      Sessional Paper No. 38a., pp. xv + 52, 36 photographs.


      1923. The burrowing rodents of California as agents in soil
      formation. Jour. Mamm., 4:137-149, pls. 13-15.


      1919. Natural history of the ground squirrels of California.
      California State Comm. Hort., Monthly Bull., 7:595-708, 5 col.
      pls., 30 figs. in text.


      1937. Fur-bearing mammals of California. Univ. California Press,
      Berkeley, 2 vols., pp. xii + xiv + 777, 13 col. pls., 345 figs. in


      1918. A synopsis of the bats of California. Univ. California Publs.
      Zoöl., 17:223-404, pls. 14-24, 24 figs. in text.

  HALL, E. R.

      1928. Notes on the life history of the sagebrush meadow mouse
      (_Lagurus_). Jour. Mamm., 9:201-204.

      1936. Mustelid mammals from the Pleistocene of North America with
      systematic notes on some Recent members of the genera Mustela,
      Taxidea and Mephitis. Carnegie Inst. Washington. Publ. 473, pp.
      41-119, 5 pls., 6 figs.

      1938A. Variation among insular mammals of Georgia Strait, British
      Columbia. Amer. Nat., 72:453-463, 2 figs.

      1938B. Gestation period in the long-tailed weasel. Jour. Mamm.,

      1944. A new genus of American Pliocene badger, with remarks on the
      relationships of badgers of the northern hemisphere. Carnegie
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      1945. Four new ermines from the Pacific Northwest. Jour. Mamm.,
      26:75-85, 1 fig.


      1934. The life history of the rufescent woodchuck, Marmota monax
      rufescens Howell. Carnegie Mus. Ann., 23:85-178, 6 pls., 9 figs.

      1940. The biology of the smoky shrew (_Sorex fumeus fumeus_
      Miller). Zoologica, 25:473-492, 4 pls.


      1939. Pollen analysis of a bog near Spokane, Washington. Bull.
      Torrey Bot. Club., 66:215-220, 1 fig. in text.

      1940. Paleoecology of a montane peat deposit at Bonaparte Lake,
      Washington. Northwest Science, 14:60-69.

      1941A. Paleoecology of a peat deposit in west central Oregon.
      Amer. Jour. Botany, 28:206-212. 1 fig. in text.

      1941B. Further studies of Post Pleistocene bogs in the Puget
      lowlands of Washington. Bull. Torrey Bot. Club., 68:133-148. 2
      figs. in text.

      1941C. A pollen study of Post Pleistocene lake sediments in the
      Upper Sonoran Life Zone of Washington. Amer. Jour. Sci.,
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      1923. Breeding habits, development, and birth of the opossum.
      Smithsonian Rept. for 1921, pp. 347-363, 10 pls.

  HATT, R. T.

      1927. Notes on the ground-squirrel, Callospermophilus. U. Michigan,
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  HAY, O. P.

      1921. Descriptions of species of Pleistocene Vertebrata, types or
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  HINTON, M. A. C.

      1926. Monograph of the voles and lemmings (Microtinae) living and
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      110 figs. in text.


      1927A. On the faunal position of the Pacific Coast of the United
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      1927B. Revision of the American lemming mice (Genus _Synaptomys_).
      N. Amer. Fauna, 50:1-38, 2 pls., 11 figs.


      1901. Revision of the skunks of the genus _Chincha_. N. Amer.
      Fauna, 20:1-62, 8 pls.

      1906. Revision of the skunks of the genus _Spilogale_. N. Amer.
      Fauna, 26:1-55, 10 pls.

      1914. Revision of the American harvest mice (Genus
      _Reithrodontomys_). N. Amer. Fauna, 36:1-97, 7 pls., 6 figs.

      1915. Revision of the American marmots. N. Amer. Fauna, 37:1-80,
      15 pls., 3 figs.

      1918. Revision of the American flying squirrels. N. Amer. Fauna,
      44:1-64, 7 pls., 4 figs.

      1924. Revision of the American pikas (Genus _Ochotona_). N. Amer.
      Fauna, 47:1-57, 6 pls., 4 figs.

      1929. Revision of the American chipmunks (Genera _Tamias_ and
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      1938. Revision of the North American ground squirrels with a
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      1939. |Review of "The Recent Mammals of Idaho" by William B.
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      1940. The new systematics. Oxford Univ. Press, pp. viii + 583, 55
      figs. in text.


      1915. A review of the American moles. N. Amer. Fauna, 38:1-100, 6
      pls., 27 figs.

      1928. A taxonomic review of the American long-tailed shrews
      (Genera _Sorex_ and _Microsorex_). N. Amer. Fauna, 51:i-vi +
      1-238, 13 pls., 24 figs.

      1944. Big-game resources of the United States, 1937-1942. U. S.
      Dept. Interior, Fish and Wildlife Research Rept. 8:1-56, 31 figs.
      in text.


      1943. Systematic review of the chipmunks (genus _Eutamias_) of
      California. Univ. California Publs. Zoöl., 48:63-148, 6 pls., 12

  JONES, G. N.

      1936. A botanical survey of the Olympic Peninsula, Washington.
      Univ. Washington Publs. Biol., 5:1-286, 9 pls.

      1938. The flowering plants and ferns of Mount Rainier. Univ.
      Washington Publs. Biol., 7:1-192, 9 pls.


      1912. Pleistocene rodents of California. Univ. California Publs.,
      Bull. Dept. Geol., 7:151-168, 16 figs. in text.


      1935. Rodent plague in California. Jour. Amer. Med. Assoc.


      1938. Microsorex hoyi washingtoni in Montana. Jour. Mamm., 19:372.


      1942. Pocket gophers and ecological succession in the Wenas region
      of Washington. Murrelet, 23:34-41, 2 figs.

      1943. Feral coypus in the Pacific Northwest. Murrelet, 24:3-9, 1

  LIVEZEY, R., and EVENDEN, F., Jr.

      1943. Notes on the western red fox. Jour. Mamm., 24:500-501.


      1902. List of the Pleistocene Fauna from Hay Springs, Nebraska.
      Bull. Amer. Mus. Nat. Hist., 16:317-322.

  MCCOY. G. W.

      1911. A plague-like disease of rodents. U. S. Publ. Health and
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      1940. Mink observations at Packwood Lake. Murrelet, 21:47.


      1892. The geographic distribution of life in North America. Ann.
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      1918. Review of the grizzly and big brown bears of North America
      (genus _Ursus_) with description of a new genus, Vetularctos. N.
      Amer. Fauna, 41:1-136, 16 pls.


      1911. The fauna of Rancho La Brea, Pt. 1, Occurrence. Mem. Univ.
      California, 1:197-213, pls. 19-23.


      1917. Age of strata referred to the Ellensburg Formation in the
      White Bluffs of the Columbia River. Univ. California Publs. Bull.
      Dept. Geol., 10:255-266, 1 pl.

  MEYER, K. F.

      1936. The sylvatic plague committee. Amer. Jour. Pub. Health,


      1942. Habitat selection among higher vertebrates and its relation
      to intraspecific variation. Amer. Nat., 76:25-35.

  MILLER, G. S., JR.

      1897. Revision of the North American bats of the family
      Vespertilionidae. N. Amer. Fauna, 13:1-140, 3 pls., 40 figs.

      1924. List of North American Recent mammals, 1923. U. S. Nat. Mus.
      Bull. 128, pp. xvi + 673.

  MILLER, G. S., JR., and ALLEN, G. M.

      1928. American bats of the genera Myotis and Pizonyx. U. S. Nat.
      Mus. Bull. 144, pp. 1-218, 1 pl., 1 fig., 13 maps.

  MOORE, A. W.

      1933. Food habits of the Townsend and coast moles. Jour. Mamm.,
      14:36-40, 1 pl.

      1940. Wild animal damage to seed and seedlings on cut-over Douglas
      fir lands of Oregon and Washington. U. S. Dept. Agri., Tech. Bull.
      706, pp. 1-28, 14 figs. in text.

      1942. Shrews as a check on Douglas fir regeneration. Jour. Mamm.,
      23:37-41, 1 pl.

      1943. Notes on the sage mouse in eastern Oregon. Jour. Mamm.,


      1932. The male reproductive tract of the Sciuridae. Amer. Jour.
      Anat., vol. 51, pp. 89-155, 7 pls., 16 figs.

  MURIE, O. J., and MURIE, A.

      1931. Travels of Peromyscus. Jour. Mamm., 12:200-209, 1 fig.

      1932. Further notes on travels of Peromyscus. Jour. Mamm.,


      1909. The rabbits of North America. N. Amer. Fauna, 29:1-314, 13
      pls., 19 figs.


      1944. Further consideration of American house mice. Jour. Mamm.,

  OGDEN, P. S.

      1909. The Peter Skene Ogden Journals. Quart. Oregon Hist. Soc.,

  ORR, R. T.

      1940. The rabbits of California. Occas. Papers, California Acad.
      Sci., 19:1-227, 10 pls., 30 figs. in text.


      1900. Revision of the pocket mice of the genus _Perognathus_. N.
      Amer. Fauna, 18:1-72, 4 pls., 15 figs.

      1909. Revision of the mice of the American genus _Peromyscus_. N.
      Amer. Fauna, 28:1-285, 8 pls., 12 figs.

      1943. The mammals of Chile. Field Mus. Nat. Hist., Zoöl. Ser.,
      vol. 30, publ. 542, pp. 1-268, 33 figs., 10 maps.

  PARDEE, J. T., and BRYAN, K.

      1926. Geology of the Latah formation in relation to the lavas of
      the Columbia Plateau near Spokane, Washington. _In_ U. S. Geol.
      Surv. Prof. Paper 140, pp. 1-16, 7 pls., 1 fig. in text.


      1920. Mountain beaver (Haplodontia or Aplodontia). Mountaineer,
      13:40-43, 1 illustration.

  PERRY, M. L.

      1939. Notes on a captive badger. Murrelet, 20:49-53, 1 fig.

  PIPER, C. V.

      1906. Flora of the state of Washington. Cont. U. S. Nat. Herb.,
      11:1-637, 22 pls., 1 map in pocket.

  POPE, C. H.

      1944. Attainment of sexual maturity in raccoons. Jour. Mamm.,

  RACEY, K., and COWAN, I. M.

      1935. Mammals of the Alta Lake region of southwestern British
      Columbia. Ann. Rept. Prov. Mus. British Columbia 1935, pp. H15-H27,
      5 pls., 1 fig. in text.

  RAND, A. L.

      1943. Canadian forms of the meadow mouse (_Microtus
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      1897. A revision of the west American flying squirrels. Proc. Acad.
      Nat. Sci. Philadelphia 1897, pp. 314-327.


      1893. A geological reconnaisance in central Washington. Bull. U. S.
      Geol. Surv. 108, pp. 1-108, 12 pls., 8 figs. in text.


      1906. Wild animals of the Mt. Rainier National Park. Sierra Club
      Bull., 6:32-38.


      1922. American moles as agriculture pests and as fur producers. U.
      S. Dept. Agri., Farm. Bull. 1247 (revised 1927), pp. 1-21, 18 figs.

      1928. Precarious status of the seal and sea-lion on our northwest
      coast. Jour. Mamm., 9:10-16.

      1929. Mountain beavers in the Pacific Northwest: their habits,
      economic status and control. U. S. Dept. Agric., Farm. Bull. 1598,
      pp. 1-18, 13 figs.

      1930. Bat matters. Murrelet, 11, (no. 2):11-13, 2 figs.

      1931. Habits and economic status of the pocket gophers. U. S.
      Dept. Agric., Tech. Bull. 224, pp. 1-26, 8 pls.

      1932. Weasels and snakes in gopher burrows. Murrelet, 13:54.

      1933. Breeding of the Washington varying hare. Murrelet, 14:77-78.

      1938A. Pocket mice of Oregon and Washington in relation to
      agriculture. U. S. Dept. Agric., Tech. Bull. 608, pp. 1-16, 6
      pls., 1 fig.

      1938B. Breeding records of Pacific Coast pocket gophers. Jour.
      Mamm., 19:220-224.

      1941. Ground squirrel studies in the Four-rivers Country,
      Washington. Jour. Mamm., 22:270-279, 2 pls.

  SCHEFFER, T. H., and SPERRY, C. C.

      1931. Food habits of the Pacific harbor seal, Phoca richardii.
      Jour. Mamm., 12:214-226.


      1938. Notes on the wolverine and fisher in the state of Washington.
      Murrelet, 19:8-10, 2 figs.

      1939. Fur seal in Willapa Harbor. Murrelet, 20:43, 1 fig.

      1940A. A newly located herd of Pacific white-tailed deer. Jour.
      Mamm., 21:271-282, 1 pl.

      1940B. The sea otter on the Washington coast. Pacific Northwest
      Quart., October, 1940, pp. 369-388, 5 figs.

      1941. Wolverine captured in Okanogan County, Washington. Murrelet,
      22:37, 1 fig.

      1942. A list of the marine mammals of the west coast of North
      America. Murrelet, 23:42-47.

      1943. The opossum settles in Washington State. Murrelet, 24:27-28.


      1939. Present distribution of the Douglas ground squirrel in
      Washington. Murrelet, 20:44.

  SCHEFFER, V. B., and SLIPP, J. W.

      1944. The harbor seal in Washington State. Amer. Mid. Nat.,
      32:373-416, 17 figs.

  SCHULTZ, L. P., and RAFN, A. M.

      1936. Stomach contents of fur seals taken off the coast of
      Washington. Jour. Mamm., 17:13-15.


      1943. The wild and commensal stocks of the house mouse, Mus
      musculus Linnaeus. Jour. Mamm., 24:59-72.

  SCOTT, W. B.

      1937. A history of land mammals in the western hemisphere.
      Macmillan Co., New York, pp. xiv + 786, 420 figs. in text.

  SHAW, W. T.

      1919. The Columbian ground squirrel. (_Citellus columbianus
      columbianus_). California State Comm. Hort., Monthly Bull. 7, pp.
      710-720, col. pl. vi. figs. 31-43.

      1924A. Alpine life of the heather vole (Phenacomys olympicus).
      Jour. Mamm., 5:12-15, pls. 2-4.

      1924B. The home life of the Columbian ground squirrel. Canadian
      Field Nat., 38:128-130, 4 figs.

      1925A. The seasonal differences of north and south slopes in
      controlling the activities of the Columbian ground squirrel.
      Ecology, 6:157-162, 2 figs. in text.

      1925B. Duration of the aestivation and hibernation of the
      Columbian ground squirrel (_Citellus columbianus_) and sex
      relations of the same. Ecology, 6:75-81, 2 figs.

      1925C. Breeding and development of the Columbian ground squirrel.
      Jour. Mamm., 6:106-113, pls. 11-14.

      1925D. The Columbian ground squirrel as a handler of earth. The
      Sci. Monthly, 20:483-490, 8 figs. in text.

      1925E. The food of ground squirrels. Amer. Nat., 59:250-264, 5
      figs. in text.

      1925F. A life history problem and a means for its solution. Jour.
      Mamm., 6:157-162, pls. 15-17.

      1925G. Observations on the hibernation of ground squirrels. Jour.
      Agric. Research, 31:761-769, 7 figs. in text.

      1925H. The hibernation of the Columbian ground squirrel. Canadian
      Field Nat., 39:56-61, 79-82, 11 figs.

      1925I. The marmots of Hannegan Pass. Nat. Hist., 25:169-177, 6
      unnumbered photographs.

      1926. Age of the animal and slope of the ground surface, factors
      modifying the structure of hibernation dens of ground squirrels.
      Jour. Mamm., 7:91-96, 1 pl., 3 figs.

      1930. The lemming mouse in North America and its occurrence in the
      state of Washington. Murrelet, 11 (No. 2):7-10, 2 figs.

      1944. Brood nests and young of two western chipmunks in the Olympic
      Mountains of Washington. Jour. Mamm., 25:274-284, 4 pls. 1 fig.


      1903. A preliminary account of the exploration of the Potter Creek
      Cave, Shasta county, California. Science, 17:708-712.

  SLIPP, J. W.

      1942. Nest and young of the Olympic dusky shrew. Jour. Mamm.,

  SMITH, G. O.

      1903. Ellensburg Folio (No. 86). U. S. Geol. Surv., Geol. Atlas of
      the U. S.


      1941. Food habits of the coyote. U. S. Dept. Interior, Wildlife
      Research Bull. 4, pp. 1-70, 3 pls., 3 figs. in text.


      1943. Notes on the resting place of Pipistrellus hesperus. Jour.
      Mamm. 24:266-267.

  ST. JOHN, H.

      1937. Flora of southeastern Washington. Student Book Corp.,
      Pullman, Washington, pp. xxv + 531, front (map) illus.

  ST. JOHN, H., and JONES, G. N.

      1928. An annotated catalogue of the vascular plants of Benton
      County, Washington. Northwest Science, 2:73-93, illustrated.


      1918. The Pleistocene fauna of Hawver Cave. Univ. California Publs.
      Bull. Dept. Geol., 10:461-515, 32 figs. in text.

      1930. Rancho La Brea: a record of Pleistocene life in California.
      Los Angeles Mus. Publ. 1, pp. 1-82, 27 figs. in text.

  SUCKLEY, G., and GIBBS, G.

      1860. Report upon the mammals collected on the survey. Repts.
      expls. and surveys ... route for a railroad, from the Mississippi
      River to the Pacific Ocean, ... 1853-1855, Pacific R. R. Rept.,
      vol. 12, book 2, pt. 3, zoöl. rept., No. 2, chap. 3, pp. 107-139, 6
      pls. (for chaps. 1-3).


      1917A. The role of isolation in the formation of a narrowly
      localized race of deer-mice (Peromyscus). Amer. Nat., 51:173-185.

      1917B. Several color "mutations" in mice of the genus Peromyscus.
      Genetics, 2:291-300, 1 fig. in text.

      1932. Genetic, distributional and evolutionary studies of the
      subspecies of deer mice (_Peromyscus_). Bibliographia Genetica,
      9:1-106, 24 figs. in text.


      1932. A comparative life history study of the mice of the genus
      _Peromyscus_. Univ. Michigan Mus. Zoöl., Misc. Publs. 24, pp. 1-39.

      1933. Notes on the deer-mouse. _Peromyscus maniculatus oreas_
      (Bangs). Murrelet, 14:13-14.

      1934. The mountain water shrew. Murrelet, 15:44-45.

      1936A. Development and growth of Peromyscus maniculatas oreas.
      Jour. Mamm., 17:132-137, 2 figs.

      1936B. Notes on the hibernation of a western chipmunk. Jour.
      Mamm., 17:289-290.

      1939. Breeding habits of Townsend's ground squirrel. Murrelet,

  SVIHLA, A., and SVIHLA, R. D.

      1931. Mink feeding on clams. Murrelet, 12:22.

      1933. Notes on the jumping mouse Zapus trinotatus trinotatus
      Rhoads. Jour. Mamm., 14:131-134.

      1940. Annotated list of the mammals of Whitman County, Washington.
      Murrelet, 21:53-58.


      1936. Breeding and young of the grasshopper mouse (Onychomys
      leucogaster fuscogriseus). Jour. Mamm., 17:172-173.

  TATE, G. H. H.

      1942. Review of the Vespertilionine bats, with special attention to
      genera and species of the Archbold collections. Bull. Amer. Mus.
      Nat. Hist., 80:221-297, 4 figs. in text.


      1918. Revision of the rodent genus _Aplodontia_. Univ. California
      Publs. Zoöl., 17:435-504, pls. 25-29, 16 figs. in text.

      1920A. The wood rat as a collector. Jour. Mamm., 1:91-92.

      1920B. A novel nesting place of the red-backed mouse. Jour. Mamm.,

      1921. Some birds and mammals of Mount Rainier. The Mountaineer,
      14:27-35, illustrated.

      1922. A distributional and ecological study of Mount Rainier,
      Washington. Ecology, 3:214-236, 4 figs. in text.

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      1927. Mammals and birds of Mount Rainier National Park. U. S. Dept.
      Interior, Nat. Park Service, U. S. Govt. Printing Office,
      Washington, D. C., pp. 1-249.

      1929. Provisional list of the land mammals of the state of
      Washington. Occ. Papers Charles R. Conner Mus. No. 2, pp. 1-32.


      1932. The genographical work with _Epilachna chrysomelina_, etc.
      Proc. 6th International Congress Genetics, 2:230.

      1940. Mutations and geographical variation, _in_ The New
      Systematics, ed. J. S. Huxley, Oxford Univ. Press, pp. 73-136, 38
      figs. in text.


      1887. Field-notes on the mammals, birds and reptiles of northern
      California. Proc. U. S. Nat. Mus., 10:159-241. 1 pl., 4 unnumbered
      figs. in text.


      1839. Narrative of a journey across the Rocky Mountains, to the
      Columbia River ... Reprint in Early Western Travels, 1748-1846, R.
      G. Thwaits, ed., A. H. Clark Co., Cleveland, 21:107-369, 1 pl.,


      1937. Tertiary stratigraphy of western-Washington and northwestern
      Oregon. Univ. Washington Publs. Geol., 4:1-266, 15 pls.

  WHITLOW, W. B., and HALL, E. R.

      1933. Mammals of the Pocatello Region of southeastern Idaho. Univ.
      California Publs. Zoöl., 40:235-275, 3 figs.

  WIGHT, H. M.

      1928. Food habits of the Townsend's mole, Scapanus townsendii
      (Bachman). Jour. Mamm., 9:19-33.


      1943. Elephant seal in southeastern Alaska. Jour. Mamm., 24:500.


      1933A. Pleistocene mammalian fauna from the Carpinteria asphalt.
      Carnegie Inst. Washington, Publ., 440:59-76.

      1933B. A rodent fauna from the later Cenozoic beds of southwestern
      Idaho. Carnegie Inst. Washington, Publ., 440:117-135, 2 pls., 8
      figs. in text.


      1932. The roles of mutation, interbreeding, crossbreeding and
      selection in evolution. Proc. 6th Internat. Congress Genetics,
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  YOUNG, S. P., and GOLDMAN, E. A.

      1944. The wolves of North America. Amer. Wildlife Inst.,
      Washington, D. C., xx + 636 pp., 131 pls., 15 figs. in text.


      1943. A coyote's speed and endurance. Jour. Mamm., 24:400.

      _Transmitted August 14, 1947._


  Abromys lordi, 299

  Accounts of species, 121

  acutorostrata, Balaenoptera, 413

  addenda, 416

  aequalidens, Thomomys talpoides, 310

    Eutamias amoenus, 257
    Tamias amoenus, 257

  akeleyi, Peromyscus, 330

  alascanus, Callorhinus, 246

  alascensis, Myotis lucifugus, 148

  Alaska fur seal, 246

    Neosorex bendirii, 144
    Sorex bendirii, 144

  albiventris, Tamias amoenus, 416

    americana, 403
    shirasi, 403

    americanus, 403
    shirasi, 403

    Mus, 364
    Rattus rattus, 364

  alpinus, Sciuropterus, 295

  alticola, Neotoma cinerea, 336

    Euarctos, 176
    Mustela frenata, 198
    Ursus, 176

    Alce, 403
    Antilocapra, 415, 416
    Martes, 186
    Mephitis, 217
    Rupicapra, 409

    Euarctos, 176
    Lepus, 382
    Oreamnos, 406
    Sitomys, 331
    Ursus, 171

    Eutamias, 255
    Tamias, 253

  angustirostris, Mirounga, 415, 416

  angustus, Microtus, 353

  Anisonyx rufa, 369

  antelope, 415, 416

  Antilocapra americana, 415, 416

    cantwelli, 169
    pallidus, 169

    columbiana, 369
    grisea, 369
    olympica, 369
    raineri, 369
    rainieri, 369
    rufa, 366

  Arctic-alpine Life-zone, 33

    Rangifer, 404
    Sorex, 132

    avarus, 263
    columbianus, 275
    douglasii, 276
    flaviventer, 263
    olympus, 267

  Arean arean, 137

  arizonensis, Mustela, 197

  artemisia, Lepus, 387

    Peromyscus maniculatus, 332
    Sitomys americanus, 332
    Synaptomys borealis, 415

  Artiodactyla, 118

    macropus, 356
    nanus, 348
    occidentalis, 351
    oregoni, 357
    pauperrimus, 359
    townsendii, 351

    Aulacomys, 356
    Microtus, 356

  asiaticus, Tamias, 262

  Atophyrax bendirii, 144

  Aulacomys arvicoloides, 356

    Hesperomys, 331
    Peromyscus, 331
    Sitomys americanus, 331

  avara, Marmota flaviventris, 263

  avarus, Arctomys flaviventer, 263

  badger, 220

  badius, Thomomys talpoides, 311

  Baird beaked whale, 410

    Lepus, 384
    Sorex obscurus, 140

    Berardius, 410
    Delphinus, 410

    acutorostrata, 413
    borealis, 413
    physalus, 413

  Balanus, 207

    Glaucomys sabrinus, 295
    Sciuropterus alpinus, 295

    big-brown, 165
    hoary, 168
    long-eared, 161
    pallid, 170
    silvery-haired, 159

  bats, mouse-eared, 146

  beaked whale,
    Baird, 410
    Stejneger, 410

    black, 170
    grizzly, 176, 415

  beaver, 315
    mountain, 366

  Beechey ground squirrel, 276

  Bendirei, Atophyrax, 144

    Atophyrax, 144
    Sorex, 144
    Neosorex, 144

  Berardius bairdii, 410

  bernardinus, Eptesicus fuscus, 165

  bibliography, 417

    brown bat, 417
    jumping mouse, 370
    myotis, 147

    bison, 404
    oregonus, 404

  bison, Bison, 404

  black bear, 171

  blackfish, 411

  black tailed,
    deer, 399
    jack rabbit, 385

  blue whale, 413

  bobcat, 241

    Balaenoptera, 413
    Lissodelphis, 410
    Lynx, 239
    Synaptomys, 337

  Brachylagus idahoensis, 390

  breviceps, Kogia, 412

  brown bat, big, 165

  brunnescens, Ochotona princeps, 380

  bushy-tailed wood rat, 333

    myotis, 156
    red-backed mouse, 344
    sea lion, 244

  californiana, Ovis, 406

    Otaria, 244
    Ovis, 406
    Zalophus, 244

    Clethrionomys, 344
    Lepus, 385
    Myotis, 156
    Vespertilio, 158

    alascanus, 246
    cynocephalus, 246
    ursinus, 246

    chrysodeirus, 280
    connectens, 280
    lateralis, 279
    saturatus, 281
    tescorum, 279

  Callotaria ursina cynocephala, 246

  campestris, Lepus, 380

    Castor, 315
    Cervus, 391
    Lutra, 205
    Lynx, 239
    Mustela, 187
    Ovis, 405

    Life-zone, 37
    lynx, 239

  Cancer productus, 180, 207

  canescens, Microtus montanus, 349

    Eutamias amoenus, 257
    Tamias amoenus, 257

    columbianus, 415
    familiaris, 234
    fuscus, 232
    gigas, 232
    incolatus, 231
    irremotus, 415
    latrans, 226
    lestes, 226
    lupus, 232
    lycaon, 232
    occidentalis, 232

    Antrozous pallidus, 169
    Microtus oregoni, 357

    hemionus, 403
    macrotis, 403

  caribou, 404

  carissima, Myotis lucifugus, 148

  Carnivora, 113

  carolinensis, Sciurus, 286

    Clethrionomys gapperi, 343
    Lepus americanus, 384
    Sciurus, 290
    Vulpes, 224

    canadensis, 315
    idoneus, 322
    pacificus, 322
    leucodonta, 322

  cat, civet, 212

  catodon, Physeter, 412

    Martes, 183
    Mustela, 186

    Eutamias, 255
    Myotis californicus, 157
    Tamias amoenus, 255

    canadensis, 391
    columbianus, 403
    hemionus, 403
    leucurus, 398
    macrotis, 403
    nelsoni, 394
    occidentalis, 394
    roosevelti, 394

  Cetacea, 410

  Check List, 112

  cheiragonus, Telmessus, 207

  chelan, Ursus, 176

    hudsonica, 217
    major, 218
    occidentalis, 219
    spissigrada, 219

    least, 251
    red-tailed, 258
    Townsend, 259
    Yellow-pine, 253, 416

  Chipmunks, 250

  Chiroptera, 112

  cicognanii, Mustela, 193

    Atalapha, 168
    Neotoma, 333

    Lasiurus, 168
    Sorex, 132
    Vespertilio, 168

    Euarctos, 176
    Ursus, 176

    beecheyi, 276
    columbianus, 272
    connectens, 280
    douglasii, 276
    lateralis, 278
    loringi, 271
    mollis, 268
    ruficaudus, 275
    saturatus, 281
    tescorum, 279
    townsendii, 268
    washingtoni, 271
    yakimensis, 268

  civet cat, 212

    californicus, 344
    cascadensis, 343
    gapperi, 341
    idahoensis, 342
    nivarius, 343
    occidentalis, 344
    saturatus, 342

  Climate and vegetation, 20

  clusius, Thomomys, 310

  Columbian ground squirrel, 272

    Arctomys, 275
    Canis lupus, 415
    Cariacus, 403
    Cervus, 403
    Citellus, 272
    Eucervus, 403
    Odocoileus, 403
    Perognathus, 299
    Spermophilus, 275
    Thomomys, 310

    Glaucomys sabrinus, 296
    Lepus americanus, 385

  concolor, Felis, 234

    Callospermophilus chrysodeirus, 280
    Citellus lateralis, 280

  cony, 377

    Eutamias, 262
    Tamias, 262

  coots, 209

    macrotis, 163
    megalotis, 163
    intermedius, 164
    rafinesquii, 161
    townsendii, 163

    Florida, 389
    Nuttall, 387

  couchi, Thomomys talpoides, 314

  cougar, 234

  coyote, 226

  coypu, 376

  crayfish, 209, 216

  creeping mouse, 357

    Lagurus, 359
    Lemmiscus, 359

  cuppes, Ochotona, 379

  cynocephala, Callotaria, 246

    Callorhinus, 246
    Siren, 246

  Dall porpoise, 412

  dalli, Phocenoides, 412

    black-tailed, 399
    mouse, 327
    mule, 399
    white-tailed, 395

  Delphinus bairdii, 410

  dermestes, 162

  deserticola, Lepus californicus, 385

  destructioni, Sorex trowbridgii, 136

  devexus, Thomomys talpoides, 309

  Didelphis virginiana, 121

    columbianus, 300
    ordii, 300

    areas, 20
    history, 68

  dog, 234

  dolphin, 410

  dorsata, Mazama, 409

  dorsatum, Erethizon, 374

  Douglas squirrel, 290

    Arctomys, 276
    Citellus, 276
    Geomys, 312
    Otospermophilus grammurus, 276
    Sciurus, 290
    Spermophilus, 276
    Tamiasciurus, 290
    Thomomys, 312

  drummondii, Microtus, 346

  Dytiscidae, 216

  Ecology, Life-zones and, 32

  edulis, Mytilus, 180, 207

  effera, Mustela frenata, 198

  elk, 391

  Emmigrational history, 54

    Mustela vison, 199
    Putorius vison, 199

  Enhydra lutris nereis, 211

  epixanthum, Erethizon, 376

    bernardinus, 165
    fuscus, 165
    pallidus, 165

    dorsatum, 374
    epixanthum, 376
    epixanthus, 376
    nigrescens, 376

  ericacus, Thomomys talpoides, 309

  eriomerus, Petrolisthes, 180, 207

  ermine, 191

  erminea, Mustela, 190

    altifrontalis, 176
    americanus, 176
    cinnamomum, 176

  Eubalaena sieboldii, 414

    jubata, 244
    stelleri, 244

    affinis, 257
    albiventris, 416
    amoenus, 255
    canicaudus, 257
    caurinus, 255
    cooperi, 262
    felix, 262
    grisescens, 253
    ludibundus, 257
    luteiventris, 258
    minimus, 252
    pictus, 252
    quadrivittatus, 256
    ruficaudus, 258
    scrutator, 252
    simulans, 258
    townsendii, 258

    Myotis, 151
    Vespertilio, 153

    gapperi, 342
    idahoensis, 342
    nivarius, 343
    occidentalis, 344
    pygmaeus, 344
    saturatus, 342

  excelsus, Procyon lotor, 182

  fallenda, Mustela erminea, 191

  familiaris, Canis, 234

  fasciatus, Lynx, 243

    Great Basin, 32
    Pacific Coastal, 52
    Rocky Mountain, 53

    concolor, 234
    hippolestes, 237
    olympus, 237
    oregonensis, 237
    missoulensis, 237

    Eutamias amoenus, 256
    Tamias amoenus, 256

  fenisex, Ochotona, 379

    occipitalis, 363
    osoyoosensis, 363
    zibethicus, 363

  finback whale, 413

  fisher, 187

  flaviventris, Marmota, 263

  Florida cottontail, 389

    Lepus sylvaticus, 389
    Sylvilagus, 389

  flying squirrel, northern, 292

  foetulenta, Mephitis, 219

    red, 224
    squirrel, 286

  frenata, Mustela, 194

  fringe-tailed myotis, 153

    Geomys, 312
    Glaucomys sabrinus, 296

  fulva, Vulpes, 224

  funebris, Microtus pennsylvanicus, 346

  fur seal, 246

  fuscogriseus, Onychomys leucogaster, 323

    Canis lupus, 232
    Eptesicus, 165
    Thomomys, 310

    Hesperomys, 331
    Peromyscus maniculatus, 331
    Sitomys americanus, 331

    Clethrionomys, 341
    Evotomys, 342

  Gapper red-backed mouse, 341

  Geologic history, 46

    douglasii, 312
    fuliginosus, 312

    Neurotrichus, 124, 122
    Urotrichus, 124

    Canis, 232
    Lupus, 232

  glacialis, Thomomys talpoides, 312

  Glaciation, 47

    bangsi, 295
    columbiensis, 296
    fuliginosus, 296
    latipes, 296
    olympicus, 295
    oregonensis, 295
    sabrinus, 292

  glaucus, Rhachianectes, 412

  Globicephalus scammonii, 411

  goat, mountain, 406

  Golden-mantled ground squirrel, 278, 281

  gopher, northern pocket, 302

  gracilis, Spilogale, 212

    Otospermophilus, 276
    Spermophilus, 276

  Grampidelphis griseus, 411

  Grampus rectipinna, 411

  grampus, 411

  grasshopper mouse, northern, 323

  gray squirrel,
    eastern, 286
    western, 284

  gray whale, 412

  Great Basin pocket mouse, 297

  grisea, Aplodontia rufa, 369

    Eutamias minimus, 253
    Tamias minimus, 253

    Grampidelphis, 411
    Sciurus, 284

  ground squirrel,
    Beechey, 276
    Columbian, 272
    golden-mantled, 278, 281
    Townsend, 268
    Washington, 271

  gryphus, Vespertilio, 148

  gulosa, Mustela erminea, 193

  hair seal, 247

  hairy-winged myotis, 154

  halli, Microtus longicaudus, 353

  Haplodon rufus, 369

    raineri, 369
    rufa, 369

    porpoise, 412
    seal, 247

  harvest mouse, western, 324

  heather vole, 338

  Helisoma occidentalis, 137

    oregonensis, 180
    nudus, 180, 207, 216

    Cervus, 403
    Odocoileus, 403

    austerus, 331
    gambelii, 331

    Pipistrellus, 165
    Scotophilus, 165
    Vespertilio, 165

  hippolestes, Felis, 237

  hoary marmot, 265

  hollisteri, Peromyscus maniculatus, 330

  house mouse, 365

  hoyi, Microsorex, 145

  Hudsonian Life-zone, 39

    Chincha, 217
    Lutra, 210
    Mephitis, 217

    Sciurus, 288
    Tamiasciurus, 286

  humpback whale, 413

  Hyla, 180
    regilla, 137

  hypophaeus, Sciurus carolinensis, 286

  hypothetical list, 415

    Brachylagus, 390
    Clethrionomys gapperi, 342
    Evotomys, 342
    Lepus, 390
    Sylvilagus, 390
    Zapus princeps, 373

  idoneus, Castor canadensis, 322

  immunis, Thomomys talpoides, 311

  imperator, Zapus, 372

  incolatus, Canis latrans, 231

  Insectivora, 112

  interior, Myotis volans, 155

    Corynorhinus rafinesquii, 164
    Phenacomys, 338

  invicta, Mustela erminea, 191

  irremotus, Canis lupus, 415

  jack rabbit,
    black-tailed, 385
    white-tailed, 380

    Eumetopias, 244
    Phoca, 244

  jumping mouse, big, 370

  kangaroo rat, Ord, 300

  Keen myotis, 151

    Myotis, 151
    Vespertilio, 151

  killer whale, 411

  kincaidi, Microtus pennsylvanicus, 347

  Kogia breviceps, 412

  kootenayensis, Zapus princeps, 373

  Lagenorhynchus obliquidens, 411

  Lagomorpha, 118

  Lagomys minimus, 379

    curtatus, 359
    pauperrimus, 359

  laingi, Perognathus parvus, 415

  Lasionycteris noctivagans, 159

  Latax lutris nereis, 211

    Callospermophilus, 279
    Citellus, 278
    Spermophilus, 281
    Tamias, 281

  latifrons, Spilogale, 215

  latipes, Glaucomys sabrinus, 296

  latrans, Canis, 226

  lemming mouse, northern, 337

    curtatus, 359
    pauperrimus, 359

  lepta, Mustela cicognanii, 193

    americanus, 382
    artemisia, 387
    bairdi, 384
    californicus, 385
    campestris, 380
    cascadensis, 384
    columbiensis, 385
    deserticola, 385
    floridanus, 389
    nuttallii, 387
    pineus, 384
    sylvaticus, 387
    texianus, 385
    townsendii, 380

  lestes, Canis, 230

  leucodonta, Castor canadensis, 322

  leucogaster, Onychomys, 323

    Cervus, 398
    Odocoileus, 398

  Life-zones and ecology, 32

  limosus, Thomomys, 312

  lion, mountain, 234

  Lissodelphis borealis, 410

  Littorina, 207

  long-eared myotis, 151

    meadow mouse, 352
    shrews, 131
    weasel, 194

  longicaudus, Microtus, 352

    Myotis, 155
    Vespertilio, 155

    Abromys, 299
    Perognathus, 299

  loringi, Citellus washingtoni, 271

  lotor, Procyon, 179

    Eutamias, 257
    Tamias amoenus, 257

    Gryphus, 148
    Myotis, 148
    Vespertilio, 148

  Lupus gigas, 232

  lupus, Canis, 232

    Eutamias amoenus, 258
    Tamias amoenus, 258

    canadensis, 205
    hudsonica, 210
    pacifica, 210
    vancouverensis, 210

    Enhydra, 211
    Latax, 211

  lycaon, Canis, 232

    borealis, 239
    canadensis, 239
    fasciatus, 243
    pallescens, 243
    rufus, 241
    uinta, 243

  lynx, Canadian, 239

  machetes, Ursus, 171

    Arvicola, 356
    Microtus, 356
    Mynomes, 356

  macroschisma, Pododesmus, 207

    Cariacus, 403
    Cervus, 403
    Corynorhinus, 163
    Odocoileus hemionus, 403

  macroura, Vulpes fulva, 415

  macrurus, Microtus, 354

    Chincha occidentalis, 218
    Mephitis mephitis, 218

  maniculatus, Peromyscus, 327

  Margarites, 207

    hoary, 265
    Olympic, 267
    yellow-bellied, 263

    avara, 263
    flaviventris, 263
    monax, 263
    olympus, 267
    petrensis, 263

  Marsupalia, 112

  marten, western, 183

    americana, 186
    caurina, 183
    origenes, 186
    pacifica, 187
    pennanti, 187

    dorsata, 409
    montana, 409

    Corynorhinus, 163
    Reithrodon, 324
    Reithrodontomys, 324

  Megaptera novaeangliae, 413

  melanops, Thomomys, 314

    Myotis, 158
    Vespertilio, 158

    americana, 217
    foetulenta, 219
    hudsonica, 217
    major, 218
    mephitis, 216
    notata, 219
    spissigrada, 219

  mephitis, Mephitis, 216

  merriami, Sorex, 134

  Mesoplodon stejnegeri, 410

    hoyi, 145
    washingtoni, 145

    angustus, 353
    arvicoloides, 356
    canescens, 349
    cantwelli, 357
    drummondii, 346
    funebris, 346
    halli, 353
    kincaidi, 347
    longicaudus, 352
    macropus, 356
    macrurus, 354
    modestus, 346
    montanus, 347
    mordax, 353
    morosus, 357
    nanus, 348
    oregoni, 357
    pauperrimus, 359
    pennsylvanicus, 345
    pugeti, 351
    richardsonii, 354
    townsendii, 349

    Eutamias, 252
    Ochotona, 379
    Lagomys, 379
    Tamias, 251

  minor, Neurotrichus, 124

  Mirounga angustirostris, 415, 416

  missoulae, Oreamnos americanus, 409

  missoulensis, Felis concolor, 237

  modestus, Microtus pennsylvanicus, 346

  mole, 125
    coast, 127
    Gibbs shrew-, 122
    shrew-, 122
    Townsend, 126

    Citellus, 268
    Spermophilus, 268

  monax, Marmota, 263

  montana, Mazama, 409

  Montane meadow mouse, 347

    Microtus, 347
    Oreamnos, 409
    Rangifer, 404

  monticola, Sorex vagrans, 138

  monticolus, Sorex, 138

  moose, 403

  Mopalia muscosa, 207

  mordax, Microtus, 353

  morosus, Microtus, 357

    beaver, 366
    goat, 406
    lion, 234
    sheep, 405

    big jumping, 370
    California red-backed, 344
    creeping, 357
    deer, 327
    Gapper red-backed, 341
    grasshopper, northern, 323
    Great Basin pocket, 297
    harvest, western, 324
    house, 365
    jumping, big, 370
    lemming, northern, 337
    long-tailed meadow, 352
    meadow, 345
    montane meadow, 347
    northern grasshopper, 323
    northern lemming, 337
    Pennsylvania meadow, 345
    pocket, Great Basin, 297
    red-backed, California, 344
    red-backed, Gapper, 341
    Townsend meadow, 349
    western harvest, 324

  mouse-eared bats, 146

  mule deer, 399

  murica, Mustela erminea, 193

    Mustela, 193
    Putorius, 193

    alexandrinus, 364
    musculus, 365
    norvegicus, 365
    rattus, 364

  muscosa, Mopalia, 207

    Mus, 365
    Sibbaldus, 413

  muskrat, 360

    altifrontalis, 198
    arizonensis, 197
    canadensis, 187
    caurina, 186
    cicognanii, 193
    effera, 190
    energumenos, 199
    erminea, 198
    fallenda, 191
    frenata, 194
    gulosa, 193
    invicta, 191
    lepta, 193
    murica, 193
    nevadensis, 197
    olympica, 192
    origenes, 186
    pacifica, 187
    pennanti, 187
    saturata, 198
    streatori, 193
    washingtoni, 198

    macropus, 356
    nanus, 348

  myops, Thomomys, 310

  Myotis, 146
    big, 147
    California, 156
    californicus, 156
    carissima, 148
    caurinus, 157
    evotis, 151
    fringe-tailed, 153
    hairy-winged, 154
    interior, 155
    Keen, 151
    keenii, 151
    long-eared, 151
    longicrus, 155
    lucifugus, 147
    melanorhinus, 158
    pacificus, 153
    saturatus, 150
    sociabilis, 150
    small-footed, 158
    subulatus, 158
    thysanodes, 153
    volans, 154
    Yuma, 149
    yumanensis, 149

  Mytilus edulis, 180, 207

    Arvicola, 348
    Microtus, 348

    Neosorex, 141
    Sorex palustris, 141

  neglecta, Taxidea taxus, 220

  nelsoni, Cervus canadensis, 394

    albiventer, 144
    bendirii, 144
    navigator, 141

    alticola, 336
    cinerea, 333
    occidentalis, 336

    Enhydra lutris, 211
    Latax lutris, 211

    gibbsii, 124
    minor, 124

  nevadensis, Mustela frenata, 197

  niger, Sciurus, 286

    Erethizon dorsatum, 376
    Reithrodontomys megalotis, 324

  nitidus, Vespertilio, 155, 157

    Clethrionomys gapperi, 343
    Evotomys, 343

    Lasionycteris, 159
    Vespertilio, 159
    Vesperugo, 159

    flying squirrel, 292
    grasshopper mouse, 323
    lemming mouse, 337
    pocket gopher, 302

    Mus, 365
    Rattus, 365

  Norway rat, 365

  notata, Mephitis mephitis, 219

  novaeangliae, Megaptera, 413

  nudus, Hemigrapsus, 180, 207, 216

  nutria, 376

  Nuttall cottontail, 387

    Lepus, 387
    Sylvilagus, 387

  obliquidens, Lagenorhynchus, 411

  obscurus, Sorex, 138

    Arvicola, 351
    Canis, 232
    Cervus canadensis, 394
    Chincha, 219
    Clethrionomys californicus, 344
    Evotomys, 344
    Helisoma, 137
    Neotoma, 336

    Fiber, 363
    Ondatra zibethicus, 363

    brunnescens, 380
    cuppes, 379
    fenisex, 379
    minimus, 379
    princeps, 377

  ochrourus, Odocoileus virginianus, 398

    columbianus, 403
    hemionus, 399
    leucurus, 398
    macrotis, 403
    ochrourus, 398
    virginianus, 398

  Olympic marmot, 267

    Aplodontia, 369
    Mustela erminea, 192
    Spilogale, 215

    Glaucomys sabrinus, 295
    Phenacomys, 340
    Sciuropterus alpinus, 295

    Felis, 237
    Marmota, 267

    occipitalis, 363
    osoyoosensis, 363
    zibethicus, 360

    fuscogriseus, 323
    leucogaster, 323

  opossum, 121

  oramontis, Phenacomys, 340

  orarius, Scapanus, 127

  Ord kangaroo rat, 300

    Dipodomys, 300
    Perodipus, 300

    americanus, 409
    missoulae, 409
    montanus, 409

  oreas, Peromyscus, 330

    Felis, 237
    Glaucomys sabrinus, 295
    Hemigrapsus, 180
    Pteromys, 295
    Sciuropterus alpinus, 295

    Arvicola, 357
    Microtus, 357

    Bison, 404
    Zapus princeps, 373

  origenes, Martes caurina, 186

    Fiber, 363
    Ondatra, 363

  Otaria californianus, 244

    river, 205
    sea, 211

    californiana, 406
    canadensis, 405

  Pacific right whale, 414

    Lutra hudsonica, 210
    Martes pennanti, 187
    Mustela canadensis, 187
    Procyon lotor, 181

    Castor canadensis, 322
    Myotis evotis, 153

  pallescens, Lynx, 243

    Antrozous, 169
    Eptesicus fuscus, 165

  palustris, Sorex, 140

  panther, 235

    Cricetodops, 298
    Perognathus, 297

    Arvicola, 359
    Lagurus, 359
    Lemmiscus, 359
    Microtus, 359

    Martes, 187
    Mustela, 187

  Pennsylvanian meadow mouse, 345

  pennsylvanicus, Microtus, 345

    columbianus, 300
    ordii, 300

    columbianus, 299
    laingi, 415
    lordi, 299
    parvus, 297

    akeleyi, 330
    americanus, 331
    artemisiae, 332
    austerus, 331
    gambelii, 331
    hollisteri, 330
    maniculatus, 327
    oreas, 330
    rubidus, 331
    texianus, 331

  personatus, Sorex, 133

  petrensis, Marmota monax, 263

  Petrolisthes eriomerus, 180, 207

    intermedius, 338
    oramontis, 340
    olympicus, 340

    jubata, 244
    richardii, 247
    vitulina, 247

  Phocena vomerina, 412

  Phocenoides dalli, 412

  physalus, Balaenoptera, 413

  Physeter catodon, 412

  Physiographic provinces, 16

  pictus, Eutamias minimus, 252

    rabbit, 390
    sperm whale, 412

  pika, 377

  pike whale, 413

  pineus, Lepus americanus, 384

  Pinnipedia, 114

  pipistrelle, western, 165

  Pipistrellus hesperus, 165

    gopher, northern, 302
    mouse, Great Basin, 297

  Podedesmus macroschisma, 207

  Plecotus townsendii, 163

  Plethodon vehiculum, 137

  Pollack whale, 413

  porcupine, 374

    Dall, 412
    harbor, 412
    right whale, 410
    striped, 411

  porpoises, 410

    Ochotona, 377
    Zapus, 371

    excelsus, 182
    lotor, 179
    pacifica, 181
    proteus, 181
    psora, 181

  productus, Cancer, 180, 207

  pronghorned antelope, 415

  proteus, Procyon, 181

  psora, Procyon, 181

  Pteromys oregonensis, 295

  pugetensis, Thomomys talpoides, 312

  pugeti, Microtus townsendii, 351

  puma, 235

  pygmacus, Evotomys, 344

  quadratus, Thomomys, 310

    Eutamias, 256
    Tamias, 256

    black-tailed jack, 385
    jack, 380
    pigmy, 390
    snowshoe, 382
    white-tailed jack, 380

  raccoon, 179

  rafinesquii, Carynorhinus, 161

    Aplodontia, 369
    Haplodontia, 369

  rainieri, Aplodontia rufa, 369

  Rana, 180

    arcticus, 404
    montanus, 404

    bushy-tailed wood, 333
    kangaroo, Ord, 300
    Norway, 365
    Ord kangaroo, 300
    roof, 364
    sage, 271
    water, 354
    wood, bushy-tailed, 333

    alexandrinus, 364
    norvegicus, 365
    rattus, 364

    Mus, 364
    Rattus, 364

  rectipinna, Grampus, 411

  red-backed mouse,
    California, 344
    Gapper, 341

    fox, 224
    squirrel, 286

  regilla, Hyla, 137

  Reithrodon megalotis, 324

    megalotis, 324
    nigrescens, 324

    Halicyon, 247
    Phoca, 247

    Microtus, 354
    Sciurus, 289
    Tamiasciurus, 289

    whale, Pacific, 414
    whale porpoise, 410

  river otter, 205

  Rhachianectes glaucus, 412

  Rodentia, 115

  roof rat, 364

  roosevelti, Cervus, 394

  rubidus, Peromyscus maniculatus, 331

    Aplodontia, 366
    Anisonyx, 369
    Haplodon, 369

    Citellus columbianus, 275
    Eutamias, 258
    Tamias, 258

    Haplodon, 369
    Lynx, 241

  Rupicapra americana, 409

  sabrinus, Glaucomys, 292

  sage rat, 271

  sagebrush vole, 359

  saturata, Mustela, 198

    Callospermophilus lateralis, 281
    Citellus, 281
    Clethrionomys gapperi, 342
    Evotomys gapperi, 342
    Myotis yumanensis, 150
    Spermophilus, 281
    Tamias, 281

  saxatilis, Spilogale, 215

  Scalops townsendii, 126

  scammonii, Globicephalus, 411

  Scapanus, 125
    orarius, 127
    schefferi, 130
    townsendii, 126
    yakimensis, 130

  schefferi, Scapanus orarius, 130

    alpinus, 295
    bangsi, 295
    olympicus, 295
    oregonensis, 295

    cascadensis, 290
    carolinensis, 286
    douglasii, 290
    griseus, 284
    hudsonicus, 288
    hypophaeus, 286
    niger, 286
    richardsonii, 289
    streatori, 290

  Scotophilus hesperus, 165

    Eutamias minimus, 252
    Tamias minimus, 252

  sea lion,
    California, 244
    Steller, 244

  sea otter, 211

    Alaska fur, 246
    fur, Alaska, 246
    hair, 247
    harbor, 247

  Sei whale, 413

  setosus, 140

  shawi, Thomomys talpoides, 311

  sheep, mountain, 405

  shirasi, Alce americanus, 403

    Bendire, 143
    cinereous, 132
    dusky, 138
    long-tailed, 131
    -mole, 122
    Merriam, 134
    mountain, 141
    pigmy, 145
    Trowbridge, 134
    wandering, 136
    water, 141, 143

  shrew-mole, Gibbs, 122

  shrews, long-tailed, 131

  Sibbaldus musculus, 413

  sieboldii, Eubalaena, 414

  silvery-haired bat, 159

  similis, Sorex vagrans, 140

    Eutamias ruficaudus, 258
    Tamias ruficaudus, 258

  Siren cynocephalus, 246

    americanus, 331
    artemisiae, 332
    austerus, 331
    gambelii, 331

    spotted, 212
    striped, 216

  small-footed myotis, 158

  snowshoe rabbit, 382

  sociabilis, Myotis yumanensis, 150

  sorex, 131,
    albiventer, 144
    areticus, 132
    bairdi, 140
    bendirei, 144
    bendirii, 144
    cinereus, 132
    destructioni, 136
    merriami, 134
    monticola, 138
    monticolus, 138
    navigator, 141
    obscurus, 138
    palustris, 141
    personatus, 133
    setosus, 140
    similis, 140
    streatori, 133
    suckleyi, 138
    trowbridgii, 134
    vagrans, 136

  sperm whale, 412

    columbianus, 275
    douglasii, 276
    grammurus, 276
    lateralis, 281
    mollis, 268
    saturatus, 281
    townsendii, 268

    gracilis, 212
    latifrons, 215
    olympica, 215
    saxatilis, 215

    Chincha occidentalis, 219
    Mephitis, 219

  spotted skunk, 212

    Beechey ground, 276
    Columbian ground, 272
    Douglas, 290
    eastern gray, 286
    flying, northern, 292
    fox, 286
    golden-mantled ground, 278, 281
    gray, eastern, 286
    gray, western, 284
    northern flying, 292
    red, 286
    Townsend ground, 268
    Washington ground, 271
    western gray, 284

  steelhead trout, 209

  Stejneger beaked whale, 410

  stejnegeri, Mesoplodon, 410

  Steller sea lion, 244

  Stenopalmatus, 170

    Mustela, 193
    Putorius, 193
    Sciurus, 288
    Sorex cinereus, 133
    Tamiasciurus, 288

    porpoise, 411
    skunk, 216

    Myotis, 151, 158
    Vespertilio, 151

  suckleyi, Sorex, 138

  sulphur-bottom whale, 413

  sylvaticus, Lepus, 387

    floridanus, 389
    idahoensis, 390
    nuttallii, 387

    artemisiae, 415
    borealis, 337
    truei, 337
    wrangeli, 337

  tacomensis, Thomomys talpoides, 312

  talpoides, Thomomys, 302

    affinis, 257
    albiventris, 416
    amoenus, 253
    asiaticus, 262
    canicaudus, 257
    caurinus, 255
    cooperi, 262
    felix, 256
    grisescens, 253
    lateralis, 281
    ludibundus, 257
    luteiventris, 258
    minimus, 251
    quadrivittatus, 256
    ruficaudus, 258
    saturatus, 281
    scrutator, 252
    simulans, 258
    townsendii, 258

    douglasii, 290
    hudsonicus, 286
    richardsonii, 289
    streatori, 290

    neglecta, 220
    taxus, 220

    Taxidea, 220
    Ursus, 220

  Telmessus cheiragonus, 207

    Callospermophilus lateralis, 279
    Citellus lateralis, 279

  texanus, Peromyscus, 331

  texianus, Lepus, 385

    aequalidens, 310
    badius, 311
    clusius, 310
    columbianus, 310
    couchi, 314
    devexus, 309
    douglasii, 312
    ericaeus, 309
    fuscus, 310
    glacialis, 312
    immunis, 311
    limosus, 312
    melanops, 314
    myops, 310
    pugetensis, 312
    quadratus, 310
    shawi, 311
    tacomensis, 312
    talpoides, 302
    tumuli, 313
    wallowa, 310
    yakimensis, 311
    yelmensis, 313

  thysanodes, Myotis, 153

  timber wolf, 232

    ground squirrel, 268
    meadow mouse, 349

    Arvicola, 351
    Citellus, 268
    Corynorhinus, 163
    Corynorhinus rafinesquii, 163
    Eutamias, 262
    Lepus, 380
    Microtus, 349
    Plecotus, 163
    Scalops, 126
    Scapanus, 125
    Spermophilus, 268
    Tamias, 259

  Transition Life-zone, 32

  trinotatus, Zapus, 372

  trout, steelhead, 209

  trowbridgii, Sorex, 134

  truei, Synaptomys, 337

  tumuli, Thomomys talpoides, 313

  uinta, Lynx, 243

  Upper Sonoran Life-zone, 36

  Urotrichus, gibbsii, 124

  ursina, Callotaria, 246

  ursinus, Callorhinus, 246

    altifrontalis, 176
    americanus, 176
    canadensis, 415
    cinnamomum, 176
    chelan, 176
    idahoensis, 415
    machetes, 171
    taxus, 220

  vagrans, Sorex, 136

  vancouverensis, Lutra, 210

  Vegetation, Climate and, 25

  vehiculum, Plethodon, 137

    californicus, 158
    cinereus, 168
    evotis, 153
    gryphus, 148
    hesperus, 165
    keenii, 151
    lucifugus, 155
    longicrus, 148
    nitidus, 155, 157
    noctivagans, 159
    subulatus, 151

    hesperus, 165
    noctivagans, 159

  virginiana, Didelphis, 121

  virginianus, Odocoileus, 398

  vitulina, Phoca, 247

  volans, Myotis, 154

    heather, 338
    sagebrush, 359

  vomerina, Phocena, 412

    cascadensis, 224
    fulva, 224
    macroura, 415

  wallowa, Thomomys talpoides, 310

  wallawalla, Lepus texianus, 385

  wapiti, 391

  Washington ground squirrel, 271

    Citellus, 271
    Microsorex hoyi, 145
    Mustela, 198
    Putorius, 198

  washingtonii, Lepus, 384

  water rat, 354

  weasel, long-tailed, 194

    gray squirrel, 284
    harvest mouse, 324
    pipistrelle, 165

    Baird beaked, 410
    blue, 413
    finback, 413
    gray, 412
    humpback, 413
    killer, 411
    Pacific right, 414
    pigmy sperm, 412
    pike, 413
    Pollack, 413
    Sei, 413
    sperm, 412
    sulphur-bottom, 413
    Stejneger beaked, 410

  whales, 410

    deer, 395
    jack rabbit, 380

  wolf, timber, 232

  woodchuck, 263

  wood rat, bushy-tailed, 333

  wrangeli, Synaptomys, 337

    Citellus mollis, 268
    Scapanus orarius, 130
    Thomomys talpoides, 311

  yellow-bellied marmot, 263

  yelmensis, Thomomys talpoides, 313

  Yuma myotis, 149

  yumanensis, Myotis, 149

  Zalophus californianus, 244

    idahoensis, 373
    imperator, 372
    kootenayensis, 373
    oregonus, 363
    princeps, 371
    trinotatus, 372

    Fiber, 363
    Ondatra, 360

Transcriber's notes:

Bold text marked as  = ... =

Italic text marked as  _ ... _

Legend for column headers (TABLE 1) inserted.

Obvious typographical errors have been silently corrected, but other
variations in spelling and punctuation remain unchanged.

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