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.
washingtoni_.
_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
reddish.
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
8.
=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
_stejnegeri_.
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
groups.
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
April.
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,
1891.
_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
_ruficaudus_.
_Measurements._--Five females from northeastern Washington
average: total length 346; length of tail 77; hind foot 48.6; ear
20.
_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_.