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Title: Taxonomy of the Chipmunks, Eutamias quadrivittatus and Eutamias
umbrinus

Author: John A. White

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*** START OF THE PROJECT GUTENBERG EBOOK TAXONOMY OF
THE CHIPMUNKS, EUTAMIAS QUADRIVITTATUS AND EUTAMIAS
UMBRINUS ***

Taxonomy of the Chipmunks,
Eutamias quadrivittatus and
Eutamias umbrinus
BY

JOHN A. WHITE

University of Kansas Publications
Museum of Natural History

Volume 5, No. 33, pp. 563-582, 6 figures in text
December 1, 1953

University of Kansas
LAWRENCE
1953

University of Kansas Publications, Museum of Natural History

Editors: E. Raymond Hall, Chairman, A. Byron Leonard,
and Robert W. Wilson

Volume 5, No. 33, pp. 563-582, 6 figures in text

December 1, 1953

University of Kansas
Lawrence, Kansas

PRINTED BY
FERD VOILAND, JR., STATE PRINTER
TOPEKA, KANSAS
1953

24-8966

Taxonomy of the Chipmunks,
Eutamias quadrivittatus and Eutamias umbrinus
By

JOHN A. WHITE

The differences in anatomy and color between many species of chipmunks
are subtle, and refined techniques are required to discover them. When
"measuring" chipmunks taxonomically, it is necessary to use a "chipmunk
scale" and not, for example, a "pocket-gopher scale." In explanation, some
species of pocket gophers closely allied to each other, and even some
subspecies of the same species, differ markedly in color and in size and
shape of parts of the skeleton; comparable differences are not so
pronounced among many species of chipmunks.

Historical Summary

Merriam (1905) was the first to show clearly that Eutamias quadrivittatus is
a distinct species, and pointed out that E. amoenus operarius (= E. minimus
operarius) is a small species which resembles, and is found in some areas
together with, E. quadrivittatus.
Howell (1929) placed under E. quadrivittatus the following subspecies: E.
q. quadrivittatus, E. q. hopiensis, E. q. inyoensis, E. q. frater, E. q.
sequoiensis, and E. q. speciosus.
Hardy (1945) placed E. adsitus under E. quadrivittatus as E. q. adsitus, and
Kelson (1951) placed E. umbrinus under E. quadrivittatus as E. q.
umbrinus.
Johnson (1943) re-established E. speciosus as a separate species, and in
California left only E. q. inyoensis in E. quadrivittatus.

Thus, since 1943 the recognized subspecies of E. quadrivittatus have been:
E. q. quadrivittatus, E. q. hopiensis, E. q. inyoensis, E. q. nevadensis, E. q.
umbrinus, and E. q. adsitus.

Methods, Materials, and Acknowledgments
Capitalized color terms, which are used in descriptions and comparisons, are of Ridgway, "Color
Standards and Color Nomenclature," Washington, D. C., 1912.
In the synonymy of each subspecies there appears only the first usage of a name, second the first
usage of the name combination now employed unless a new combination is proposed by me, and
third pure synonyms. The last is recognizable as such because the type locality is appended to each.
Unless otherwise specified, all specimens are in the Museum of Natural History, University of
Kansas. The various collections of institutions and of private persons are indicated by the following
symbols:

AM—American Museum of Natural History.
BS—United States Biological Surveys Collection.
CM—Colorado Museum of Natural History.
DC—Collection of Donald R. Dickey (now the collection of the University of
California at Los Angeles).
FC—Collection of James S. Findley.
KU—Museum of Natural History, University of Kansas.
MM—Museum of Zoology, University of Michigan.
NM—United States National Museum.
UU—Museum of Zoology, University of Utah.
WC—Collection of Edward R. Warren, Colorado College.
Of the external measurements, only the total length and the length of the tail are recorded in table 1.
Some field collectors measured the ear from the notch and others from the crown; most collectors
measured the length of the hind foot to the nearest millimeter rather than in tenths of a millimeter, as
would have been desired. Consequently, I decided against using the lengths of the ear and hind foot
in the study here reported on.
The measurements of the skull were made as shown in figure 1.

Fig. 1. Dorsal view of skull and a lateral and a medial view of the right
lower jaw to show points between which measurements of the skull
were taken. × 1¾. Based on Eutamias ruficaudus ruficaudus, from 6 mi.
S St. Mary, 6500 ft., Glacier Co., Montana. A to A'—greatest length of
skull; B to B'—length of nasals; C to C'—zygomatic breadth; D to D'—
least interorbital constriction; E to E'—cranial breadth; F to F'—inner
mandibular length; G to G'—condylo-alveolar length of mandible.
A total number of 434 specimens are listed as examined in this study, and additionally, numerous
other specimens were superficially examined in the United States Biological Surveys Collection.
Bacula of each of the named kinds of chipmunks in this paper, were examined.

Whenever two or more samples are stated to be significantly different, the meaning is that the
difference is statistically significant.
The geographic distribution of each subspecies and the localities of specimens or series of specimens
are plotted on the map (fig. 2).
When comparisons were made to ascertain specific and subspecific differences, only adults, or
animals in which the enamel was worn through on the permanent P4 and p4 were used. Within this
age range, only specimens in comparable pelage were used to ascertain differences in color.

Miss Viola S. Schantz of the United States Fish and Wildlife Service, Mr. Alfred Bailey of the
Colorado Museum of Natural History, Dr. W. H. Burt of the Museum of Zoology of the University of
Michigan, Dr. Stephen D. Durrant of the Museum of Zoology of the University of Utah, Dr. Robert
M. Stabler, curator of the Warren Collection of Colorado College, and Mr. James S. Findley,
generously loaned specimens for my use. Doctors E. Raymond Hall, Rollin H. Baker, Robert W.
Wilson, Keith R. Kelson, E. Lendell Cockrum, and other friends and associates have given valued
suggestions and assistance. My wife, Alice M. White, made the illustrations and helped me record
and analyze the data.
Assistance with field work is acknowledged from the Kansas University Endowment Association, the
National Science Foundation, and the United States Navy, Office of Naval Research, through
contract No. NR161 791.

Accounts of Species and Subspecies

Eutamias quadrivittatus (Say)

Diagnosis.—Size medium; general tone of upper parts tawny; cranial
breadth averaging between 16.0 and 16.8 mm.; baculum distinguishable
from that of any other species by the combination of width of base less than
¼ of length of shaft, shaft having a maximum diameter of more than ¼
mm., and height of keel ¼ of length of tip.
Eutamias quadrivittatus quadrivittatus (Say)
Sciurus quadrivittatus Say, in Jones, Long's Expedition to Rocky Mountains, 2:45,
1823.
Eutamias quadrivittatus, Miller and Rehn, Proc. Boston Soc. Nat. Hist., 30:43,
December 27, 1901.
Tamias quadrivittatus gracilis J. A. Allen, Bull. Amer. Mus. Nat. Hist., 3:99, June
1890, Type from San Pedro, Santa Fe Co., New Mexico.
Eutamias quadrivittatus animosus Warren, Proc. Biol. Soc. Washington, 22:105, June
25, 1909. Type from Irwin Ranch, Las Animas County, Colorado.
Type.—None designated; from along Arkansas River, about 26 mi. below Canon City, Fremont
County, Colorado; obtained on July 18, 1820.
Diagnosis.—Size medium; dorsal dark stripes blackish; sides Cinnamon to Clay Color; crown Light
Drab; baculum large.
Description.—Color pattern: Head Cinnamon, shaded on crown to Light Drab; ocular stripe Fuscous
Black, with Cinnamon along margins; other facial stripes Fuscous mixed with Cinnamon; ears
Fuscous Black, Ochraceous-Tawny on anterior margin, grayish white on posterior margin and on
postauricular patch; dark dorsal stripes black with Ochraceous-Tawny along margins; outer pair of
dark stripes often mainly Tawny; light dorsal stripes grayish white, outer pair usually creamy white;
sides Ochraceous-Tawny, shaded in the region of the shoulder with Cinnamon; rump and thighs

Cinnamon-Buff mixed with Smoke Gray; antipalmar surfaces of forefeet Cinnamon-Buff; antiplantar
surfaces of hind feet Pinkish Buff; dorsal surface of tail Fuscous Black, overlaid with Pinkish Buff;
ventral surface of tail Ochraceous-Tawny, Fuscous Black along margin, Pinkish Buff along outermost
edge; underparts creamy white. Skull: Large; braincase well inflated; zygomatic arches strong and
slightly appressed to skull. Baculum: Large; long and slender.
Comparisons.—From E. q. hopiensis, the only other subspecies in this species, E. q. quadrivittatus
differs in: Dorsal dark stripes blackish; crown grayer; rump and thighs grayer; general tone of upper
parts darker.

Remarks.—Specimens from the Chuska Mountains, Zuni Mountains, and
Blanco, New Mexico, are intergrades between E. q. quadrivittatus and E. q.
hopiensis, but are referable to E. q. quadrivittatus.
In north-central Colorado E. umbrinus occurs in the spruce and pine forests
at higher altitudes, while to the south and east of this area E. q.
quadrivittatus occurs in growths of piñon in lower, semiarid areas. In the
northern half of New Mexico and in south-central Colorado, E. q.
quadrivittatus occurs not only in semiarid habitats but also in the moist
habitats of the forests of higher altitudes. Ecologically, E. umbrinus thus
replaces E. q. quadrivittatus in north-central Colorado. This ecological
replacement is comparable to the ecological replacement of Thomomys
bottae by T. talpoides in Utah as shown by Durrant (1952:156).
Specimens examined.—Total number, 130.
Colorado: Larimer Co.: Arkins, 1 BS. Jefferson Co.: W spur Lookout Mountain, near Golden, 1 WC.
Gunnison Co.: Sapinero, 3 BS. Saguache Co.: 5 mi. N and 22 mi. W Saguache 10,000 ft., 1; 21 mi.
W and 3 mi. N Saguache, 1. Fremont Co.: 18 mi. S and 7 mi. W Colorado Springs, 1; Arkansas
River, "about" 26 mi. below Canon City, 15 BS. San Juan Co.: Silverton, 1 BS. Mineral Co.: 3 mi. E
Creede, 1. Alamosa Co.: Sangre de Cristo Range, 24 mi. E Hooper, 2 CM. La Plata Co.: 2 mi. NE
Bondad 6,100 ft., 1; Bondad, 15 mi. S Durango 6,050 ft., 1. Archuleta Co.: Chromo, 1 CM. Las
Animas Co.: Trinidad, 6 BS. Baca Co.: unspecified, 1.
New Mexico: San Juan Co.: Blanco, 1 BS; Chuska Mountains, 8 BS. Rio Arriba Co.: 8 mi. N El
Rito, 1; 4 mi. N El Rito, 5; Rim Rock, El Rito, 2; 2 mi. E El Rito, 7,000 ft., 1; 2 mi. SE El Rito, 1; 6
mi. E and ½ mi. S Truchas, 8,500 ft., 1; 2 mi. S and 4 mi. W Coyote, 8,100 ft., 1; unspecified, 2. Taos
Co.: 3 mi. N Taos Pueblo, 5 BS; 23 mi. S and 6 mi. E Taos, 8,750 ft., 2. Union Co.: Emery Peak, 1
BS; Folsom, 3 BS; Sierra Grande, 8 BS; unspecified, 2. McKinley Co.: Bear Ridge, Zuni Mountains,
9 BS. Sandoval Co.: Bear Canyon, W foothills, Sandia Mountains, 3 BS; W foothills, near S end,
Sandia Mountains, 7 BS. Santa Fe Co.: San Pedro, 7 BS. San Miguel Co.: Canadian River, 4 mi. NW
Tucumcari, 1 BS. Valencia Co.: Mount Taylor, San Mateo Mountains, 10 BS.
Oklahoma: Cimarron Co.: Kenton, 1 BS.

Eutamias quadrivittatus hopiensis Merriam
Eutamias hopiensis Merriam, Proc. Biol. Soc. Washington, 18:165, June 29, 1905.

Eutamias quadrivittatus hopiensis, Howell, Jour. Mamm. 3:184, August 4, 1922.
Type.—Female, adult, skull and skin, No. 67768 U. S. Nat. Mus.; from Keams Canyon, Painted
Desert, Arizona; obtained on July 27, 1894, by A. K. Fisher.
Diagnosis.—Size medium; dorsal dark stripes tawny; crown Drab-Gray; baculum of same
proportions as in E. q. quadrivittatus but smaller.

Fig. 2. Localities of specimens examined and probable geographic
ranges of the subspecies of Eutamias quadrivittatus and Eutamias
umbrinus. The symbols for locality records are as follows: circles,
precise localities; triangles, localities known only to county.

Guide to subspecies:
1. E. q. quadrivittatus
2. E. q. hopiensis
3. E. u. umbrinus

4. E. u. adsitus
5. E. u. sedulus
6. E. u. inyoensis
7. E. u. nevadensis
8. E. u. fremonti
9. E. u. montanus
Description.—Color pattern: Head Drab-Gray, with Snuff Brown around margin of crown; facial
stripes Sayal Brown with small blackish patches around eye; ears Ochraceous Tawny anteriorly and
Pinkish Buff posteriorly; dorsal stripes Tawny, median one sometimes blackish; median pair of dorsal
light stripes grayish white, outer pair creamy white; sides Ochraceous Tawny; rump and thighs
Cinnamon Buff washed with Pale Smoke Gray; antipalmar and antiplantar surfaces of feet Pinkish
Cinnamon; dorsal surface of tail Fuscous Black; ventral surface of tail Ochraceous Tawny, Fuscous
Black along margin, Cinnamon Buff along outermost edge; underparts creamy white. Skull: As in E.
q. quadrivittatus. Baculum: Same proportions as in E. q. quadrivittatus but smaller.
Comparisons.—See under the account of E. q. quadrivittatus.

Remarks.—Topotypes of this subspecies are intergrades between it and E.
q. quadrivittatus.
In a large part of the geographic range of E. q. hopiensis there are
numerous, massive outcrops of Mesozoic sandstones, which tend to form
cliffs, that are brightly colored with many shades of red. The color which is
characteristic of E. q. hopiensis seems to be helpful in adapting this
subspecies to this habitat of red sandstone, for these chipmunks are
generally found in the rubble and among the piñon at the base of the cliffs.
At many places in Utah above these cliffs of red sandstone there are forests
predominantly composed of yellow pine. Kelson (1951:42-43) states that
"these two habitats are in immediate juxtaposition, the transition from one
to the other often occurring in only a few feet ..." and again, "No one to my
knowledge, has found any evidence in specimens from Utah of
interbreeding of E. q. hopiensis with either E. q. adsitus [= E. umbrinus
adsitus] or E. q. umbrinus [= E. u. umbrinus]." Benson (1935:449) states,
"On Navajo Mountain these chipmunks [E. q. hopiensis] were most in
evidence on rock outcrops surrounded by brush at the lower edge of the
yellow pine zone. One was seen at about 9,500 feet in a south-facing rock
outcrop near the spruce-fir forest, but no chipmunk of any kind was seen in
the forest itself." This suggests that where only E. q. hopiensis occurs on a
mountain this subspecies goes higher than on a mountain where E. u.
adsitus also occurs. This same relationship between E. q. quadrivittatus and

the subspecies of E. umbrinus that occurs in north-central Colorado was
pointed out in the account of E. q. quadrivittatus.
Specimens examined.—Total number, 68.
Utah: Uintah Co.: E side of confluence of Green and White rivers, 1 mi. SE Ouray, 4,700 ft., 3 UU.
Grand Co.: Colorado River above Moab, 1 UU; side canyon of Colorado River above Moab, 1 UU;
Moab, up Colorado River, 1 UU; Moab, 4,500 ft., 4 UU; Moab Bridge over Colorado River, 3,995
ft., Moab, 1 UU; Colorado River, 5 mi. E Moab Bridge, 4,000 ft., 1 UU. Wayne Co.: Fruita, 1 UU.
Colorado: Moffat Co.: 11 mi. W and 11 mi. N Rangely, 6,000 ft., 3. Rio Blanco Co.: White River, 5
BS. Eagle Co.: McCoy, 2 BS. Mesa Co.: 1½ mi. S Loma, 4,600 ft., 1. Gunnison Co.: 1 mi. E
Somerset, 6,100 ft., 1. Montrose Co.: 1 mi. E Naturita, 5,900 ft., 1. Dolores Co.: 1 mi. N Cahone,
6,900 ft., 1. Montezuma Co.: 1 mi. S Cortez, 5,000 ft., 1; Mesa Verde, 25 mi. SW Mancos, 7,000 ft., 2
BS.
Arizona: Navajo Co.: Keams Canyon, 80 mi. N Holbrook, 15 BS. Apache Co.: Summit, 8,000 ft.,
Luka Chukai Mountains, 15 mi. E Luka Chukai Navajo School, 8 BS; Wheatfield Creek, W slope
Tunicha Mountains, 7,000 ft., 3 BS.

Eutamias umbrinus (J. A. Allen)

Diagnosis.—Size medium; pelage dark; sides dark; narrow cranial breadth;
baculum distinguishable from that of any other species (E. palmeri
excepted) by the combination of width of base more than ⅓ of length of
shaft, distal ½ of shaft laterally compressed, and keel ¼ of length of tip.
Eutamias umbrinus umbrinus (J. A. Allen)
Tamias umbrinus J. A. Allen, Bull. Amer. Mus. Nat. Hist. 3:96, June, 1890.
Eutamias umbrinus, Miller and Rehn, Proc. Boston Soc. Nat. Hist. 30:45, December
27, 1901.
Type.—Male, adult, skull and skin, No. 186463 U. S. Nat. Mus., Biol. Surv. Coll.; from Blacks Fork,
about 9,500 ft., Uinta Mountains, Utah; obtained on September 19, 1888, by Vernon Bailey; original
No. 228.
Diagnosis.—Size medium; general tone of upper parts dark and shadowy; skull relatively small.
Description.—Color pattern: Head Pale Smoke Gray; facial stripes Fuscous Black to Snuff Brown;
ear Fuscous Black; posterior margin of ear and postauricular patch grayish white; median dorsal dark
stripe black with Sayal Brown along margins; lateral pair of dorsal dark stripes Sayal Brown or
Fuscous Black mixed with Sayal Brown; outermost pair of dorsal dark stripes nearly absent; sides
Sayal Brown mixed with Cinnamon; rump and thighs Sayal Brown mixed with Smoke Gray;
antipalmar and antiplantar surfaces of feet Cinnamon-Buff; ventral surface of tail Ochraceous Tawny
or Sayal Brown, with Fuscous Black around margin and Pinkish Buff around outermost edge;
underparts creamy white with dark gray underfur. Skull: Large, with moderately inflated braincase
and well developed zygomata. Baculum: One of the largest in the species.

Comparisons.—From Eutamias umbrinus adsitus, the subspecies to the south on the Wasatch Range,
E. u. umbrinus differs in: Sides lighter; rump browner; hairs around outermost edge of tail tawnier (in
freshly molted tails); shorter inner mandibular length.
From E. u. inyoensis, the subspecies to the west in central and northeastern Nevada and in
northwestern Utah, E. u. umbrinus differs in: General tone of upper parts lighter; sides lighter; total
length more; interorbital region broader.
For comparisons with E. u. sedulus, E. u. fremonti, and E. u. montanus, see the accounts of those
subspecies.
Specimens examined.—Total number, 55.

Wyoming: Uinta Co.: 9 mi. S Robertson, 8,000 ft., 15; 10 mi. S and 1 mi. W Robertson, 8,700 ft., 5;
11½ mi. S and 2 mi. E Robertson, 9,200 ft., 1; 2 mi. E and 12 mi. S Robertson, Ashley Nat. For., 1;
13 mi. S and 2 mi. E Robertson, 9,200 ft., 1.
Utah: Rich Co.: Monte Cristo, 18 mi. W Woodruff, 8,000 ft., 2 UU. Summit Co.: 13½ mi. S and 2 mi.
E Robertson [Wyoming], 4; 1 mi. N Bridger Lake R. S., 9,400 ft., 4. Wasatch Co.: Snake Creek
Canyon, 3 mi. NW Midway, 6,000 ft., 1 UU. Uintah Co.: Paradise Park, 21 mi. W and 15 mi. N
Vernal, 10,050 ft., 20.

Eutamias umbrinus adsitus J. A. Allen
Eutamias adsitus J. A. Allen, Brooklyn Institute Mus. Sci. Bull. 1:118, March 31,
1905.
Type.—Unsexed adult, skull and skin, No. 28728 Amer. Mus. Nat. Hist.; from Briggs Meadow,
10,000 ft., Beaver Mountains, Utah; obtained on August 20, 1904, by George P. Engelhardt.
Diagnosis.—Size medium; sides dark; general tone of upper parts dark; dorsal light and dark stripes
strongly contrasting.
Description.—Color pattern: Head Cinnamon mixed with grayish white; stripe on margin of crown
Verona-Brown or Bister; ocular stripe Fuscous Black mixed with Sayal Brown; submalar stripe Sayal
Brown; ear Fuscous, Sayal Brown along anterior margin and Smoke Gray along posterior margin and
on postauricular patch; median dorsal stripe black; lateral dorsal dark stripes Fuscous Black mixed
with Russet; outermost dorsal dark stripes slightly darker or indistinguishable from sides in color;
dorsal light stripes grayish white with Mikado-Brown along margins; outermost pair of dorsal light
stripes nearly pure white; sides Russet mixed with Cinnamon or Ochraceous-Tawny; rump and thighs
Smoke Gray mixed with Cinnamon-Buff, with a larger or smaller number of Fuscous Black hairs;
antipalmar and antiplantar surfaces of feet Cinnamon-Buff; dorsal surface of tail black; ventral
surface of tail Sayal Brown to Tawny; underparts white with dark underfur. Skull and Baculum: As in
E. u. umbrinus.
Comparisons.—From E. u. inyoensis, the subspecies to the west, E. u. adsitus differs in: General tone
of upper parts darker; sides darker; interorbital region wider; skull significantly deeper.
For comparison with E. u. umbrinus, E. u. sedulus, and E. u. montanus, see the accounts of those
subspecies.

Remarks.—Specimens from West Rim, Zion National Park, 6,500 ft.,
Washington County, Utah, seem to be intergrades between E. u. adsitus and

E. u. inyoensis, and are referable to E. u. adsitus.
Specimens examined.—Total number, 34.
Utah: Beaver Co.: Britts Meadow, Beaver Range Mountains, 8,500 ft., 13 BS. Wayne Co.: Donkey
Lake, Boulder Mountain, 10,000 ft., 4 UU. Garfield Co.: Wildcat R. S., Boulder Mountain, 8,700 ft.,
5 UU.
Arizona: Coconino Co.: De Motte Park, Kaibab Plateau, 3 BS; Bright Angel, Kaibab Plateau, 9 BS.

Eutamias umbrinus sedulus new subspecies
Type.—Male, adult, skull, skin, and baculum, No. 158181 U. S. Nat. Mus. Biol. Surv. Coll.; from
Mount Ellen, Henry Mountains, Garfield County, Utah; obtained on October 13, 1908 by W. H.
Osgood; original No. 3667.
Diagnosis.—Size medium; general tone of upper parts dark reddish-brown; ventral surface of tail
Ochraceous-Orange; sides Mars Yellow.
Description.—Color pattern: Crown Drab-Gray mixed with Fuscous; upper facial stripe Fuscous
Black mixed with Sudan Brown; ocular stripe Sudan Brown mixed with black; submalar stripe Sudan
Brown slightly mixed with black; anterior margin of ear Sudan Brown slightly mixed with black;
hairs inside pinna, posteriorly, Warm Buff; posterior margin of ear and postauricular patch creamy
white; median dorsal dark stripe black with Antique Brown along margins; lateral dorsal dark stripes
black mixed with Antique Brown; outermost dorsal dark stripes Xanthine Orange slightly mixed with
black; median dorsal light stripes Pale Smoke Gray; outermost dorsal light stripes white slightly
mixed with gray; rump and thighs Smoke Gray; sides Mars Yellow; dorsal surface of tail black mixed
with Warm Buff; ventral surface of tail Ochraceous-Orange, with black around margin, and Warm
Buff around outermost edge; antipalmar and antiplantar surfaces of feet Ochraceous-Buff; underparts
creamy white with dark underfur. Skull: Large; braincase moderately inflated; zygomata strong.
Baculum: As in E. u. umbrinus.
Comparisons.—From E. u. umbrinus, the subspecies from the Uinta and northern Wasatch
Mountains of Utah, E. u. sedulus differs in: General tone of upper parts lighter; sides lighter.
From E. u. adsitus, the subspecies from the southern Wasatch Range in Utah and Kaibab Plateau in
Arizona, E. u. sedulus differs in: Sides lighter; general tone of upper parts markedly lighter.
From E. u. inyoensis, the subspecies from central and northeastern Nevada, and western and
northwestern Utah, E. u. sedulus differs in: Sides lighter (less grayish); general tone of upper parts
tawnier.
For comparison with E. u. montanus, see the account of that subspecies.
Specimens examined.—Total number, 7 BS, all from the type locality.

Eutamias umbrinus inyoensis Merriam
Eutamias speciosus inyoensis Merriam, Proc. Biol. Soc. Washington, 11:202, 208,
July 1, 1897.
Type.—Male, adult, skull and skin, No. 29387/41462 U. S. Nat. Mus. Biol. Surv. Coll.; from Black
Canyon, 8,200 ft., White Mountains, Inyo County, California; obtained on July 7, 1891, by E. W.

Nelson; original No. 1069.
Diagnosis.—Size medium; sides light; general tone of upper parts light; baculum one of largest in
species.
Description.—Color pattern: Head Smoke Gray mixed with Pink-Cinnamon; upper two pairs of
facial stripes Fuscous Black or black; submalar stripe Sayal Brown; ear Fuscous or Chaetura-Drab,
posterior margin and postauricular patch buffy white; median dorsal dark stripe black with Sayal
Brown along margins; lateral dorsal dark stripes black mixed with Sayal Brown or Mikado Brown;
outermost dorsal dark stripes Sayal Brown or Mikado Brown mixed with black; sides Ochraceous-
Tawny or Tawny; thighs Cinnamon-Buff mixed with Smoke Gray; antipalmar and antiplantar
surfaces of feet Cinnamon-Buff; ventral surface of tail Cinnamon-Buff or Ochraceous-Tawny with
Fuscous Black around margin and Pinkish Buff around outermost edge; underparts creamy white.
Skull: Large; zygomata strong; braincase moderately inflated. Baculum: One of largest in species.
Comparisons.—For comparisons with E. u. umbrinus, E. u. adsitus, E. u. sedulus, and E. u.
nevadensis, see the accounts of those subspecies.

Remarks.—The baculum in E. u. inyoensis is like that in E. palmeri.
Specimens examined.—Total number, 46.
Nevada: Elko Co.: Head Ackler Creek, N end Ruby Mountains, 1; Steels Creek, N end Ruby
Mountains, 1; Summit Secret Pass, 6,200 ft., Ruby Mountains, 2; Three Lakes, Ruby Mountains, 11;
Long Creek, S fork, Ruby Mountains, 4; Harrison Pass R. S., Green Mountain Canyon, 1; W side
Ruby Lake, 6 mi. N Elko Co. line, 3; W side Ruby Lake, 3 mi. N Elko Co. line, 8. White Pine Co.:
Willow Creek, 2 mi. S White Pine Co. line, Ruby Mountains, 6; W side Ruby Lake, 3 mi. S White
Pine Co. line, 5; Overland Pass, E slope Ruby Mountains, 8 mi. S White Pine Co. line, 2.
Utah: Boxelder Co.: Head of George Creek and Clear Creek, 5 mi. S Stanrod, Raft River Mountains,
8,500 ft., 2 UU.

Eutamias umbrinus nevadensis Burt
Eutamias quadrivittatus nevadensis Burt, Jour. Mamm. 12:299, August 24, 1931.
Type.—Male, adult, skull and skin, No. 15884 Donald R. Dickey Collection; from Hidden Forest,
Sheep Mountains, 8,500 ft., Clark County, Nevada; obtained on July 13, 1929, by W. H. Burt;
original No. 2337.
Diagnosis.—Size medium; general tone of upper parts grayish; baculum one of the largest of species.
Description.—"General tone of upperparts grayish; median dorsal stripe, extending from crown
between ears to rump, black faintly bordered with 'verona brown'; lateral dark dorsal stripes similar
to median stripe, but with anterior one-third deeply suffused with 'verona brown'; central light dorsal
stripes grayish, slightly lighter than head and rump; lateral stripes white; head and rump 'pale smoke
gray'; postauricular patch grayish white, a narrow margin extending up posterior border of ear;
anterior portion of ear 'fuscous black' mixed with 'verona brown' at base and bordered by light gray;
ocular stripe black grading into 'verona brown' in front of ear; submalar stripe nearly obsolete, 'sayal
brown'; sides of body grayish washed with 'verona brown'; feet grayish very faintly washed with
'pinkish buff'; dorsal surface of tail black overlaid with 'tilleul buff'; ventral surface of tail 'cinnamon
buff' narrowly bordered by black then by 'tilleul buff'; ventral surface of body white." (Burt
1931:299.) Skull similar to that of E. u. inyoensis but differing as indicated below.

Comparisons.—From E. u. inyoensis, the subspecies to the north, E. u. nevadensis differs in: Paler
and grayer throughout; tawny areas restricted; gray areas clearer and less suffused; dark facial
markings narrower and less distinct; ventral surface of tail distinctly paler; feet lighter, clearer gray;
nasals extend farther posteriorly with respect to premaxillae (Burt loc. cit.).
From E. u. adsitus, the subspecies to the northeast, E. u. nevadensis differs in: Narrower dorsal
stripes and facial markings; paler coloration of head, rump, sides, feet, and ventral surface of tail
(Burt op. cit.: 299-300).

Remarks.—The differences between E. umbrinus nevadensis and E.
palmeri, as shown by Burt (op. cit.) and Hall (1946), are such that one
might expect E. palmeri to be a subspecies of E. umbrinus. However,
having only the structure of the baculum as evidence additional to that
summarized by Hall (op. cit.), I follow him in according E. palmeri specific
status.
Specimens examined.—None.

Eutamias umbrinus fremonti new subspecies
Type.—Male, adult, skull, skin, and baculum, No. 41790 Univ. Kansas Mus. Nat. Hist.; from 31 mi.
N Pinedale, 8,025 ft., Sublette County, Wyoming; obtained on July 8, 1951, by Rollin H. Baker;
original No. 1596.
Diagnosis.—Size large; sides Capucine Yellow; antiplantar surface of hind feet Raw Sienna;
postauricular patch grayish white; baculum as in E. u. umbrinus.
Description.—Color pattern: Crown Cinnamon-Buff mixed with gray; upper facial stripe Sepia;
ocular stripe Chaetura-Drab; submalar stripe Fuscous Black mixed with Sayal Brown; ear black;
anterior margin of ear Mars-Yellow, posterior margin grayish white; hairs inside posterior portion of
pinna Dresden-Brown; postauricular patch Pale Smoke Gray; median dorsal dark stripe black; lateral
dorsal dark stripe black mixed with Sayal Brown; outermost dorsal dark stripe obsolete, Buckhorn-
Brown mixed with black; median pair of dorsal light stripes grayish mixed with Buckhorn-Brown;
outer pair of dorsal light stripes creamy white; sides Buckhorn-Brown; rump Pale Smoke Gray mixed
with Saccardo's Umber; dorsal surface of tail black mixed with Buckhorn-Brown; ventral surface of
tail Sayal Brown; outermost edge of tail Light Buff; antipalmar surface of forefeet Warm Buff;
antiplantar surface of hind foot Ochraceous-Tawny; underparts creamy white with dark underfur.
Skull: Large, with strong zygomata; braincase well inflated. Baculum: As in E. u. umbrinus.
Comparisons.—From E. u. umbrinus, the subspecies from the Uinta and northern Wasatch
Mountains in Utah, E. u. fremonti differs in: Sides darker; antiplantar surfaces of feet darker;
postauricular patch grayer; crown more grayish; skull slightly larger.
From E. ruficaudus ruficaudus, the species and subspecies from western Montana, E. u. fremonti
differs in: General tone of upper parts, sides, underside of tail, and feet, all darker in coloration;
baculum shorter and proportionally twice as wide at base.
For comparison with E. u. montanus, see the account of that subspecies.

Remarks.—The geographic ranges of E. umbrinus fremonti and E.
ruficaudus ruficaudus are allopatric and no specimens have ever been taken
in the intermediate area to indicate whether or not these two species
anywhere occur together. The bacula in the two species differ to the same
degree as those of E. quadrivittatus and E. umbrinus. The differences
between E. u. fremonti and E. r. ruficaudus are such that in my opinion, E.
ruficaudus is a distinct species.
Specimens examined.—Total number, 58.
Montana: Park Co.: Beartooth Mountains, 2 BS.
Idaho: Bonneville Co.: Big Hole Mountains, 9,000 ft., near Irwin, 1 BS.
Wyoming: Yellowstone Park, 2. Park Co.: 16¼ mi. N and 17 mi. W Cody, 5,625 ft., 2. Teton Co.: 1
mi. E and ¼ mi. N Togwotee Pass, 9,800 ft., 2; Amphitheatre Lake, Teton Park, 1 MM; Flat Creek, 4
MM; head of Cache Creek, 4 MM; Jackson, Upper Arizona Creek, 2 MM; Flat Creek-Granite Creek
divide, 6 MM; Flat Creek Pass, 1 MM; Flat Creek-Gravel Creek divide, 2 MM. Lincoln Co.: La
Barge Creek, 9,000 ft., 2 BS. Fremont Co.: Togwotee Pass, 12; 12 mi. N and 3 mi. W Shoshoni,
4,650 ft., 1; Mosquito Park R. S., 9,500 ft., 17½ mi. W and 2½ mi. N Lander, 1; 17 mi. S and 6½ mi.
W Lander, 8,450 ft., 3. Sublette Co.: 31 mi. N Pinedale, 8,025 ft., 2; W side Barbara Lake, 10,300 ft.,
8 mi. S and 3 mi. W Fremont Peak, 4; 19 mi. W and 2 mi. S Big Piney, 7,700 ft., 5.

Eutamias umbrinus montanus new subspecies
Type.—Male, adult, skull, skin, and baculum, No. 20105 Univ. Kansas Mus. Nat. Hist.; from ½ mi. E
and 3 mi. S Ward, 9,400 ft., Boulder County, Colorado; obtained on August 1, 1947, by E. L.
Cockrum; original No. 721.
Diagnosis.—Size large; sides Clay Color; antipalmar and antiplantar surfaces of feet Cinnamon-Buff;
baculum as in E. u. umbrinus.
Description.—Color pattern: Crown Raw Sienna mixed with gray; upper facial stripe and ocular
stripe black mixed with Sepia; submalar stripe Snuff Brown mixed with black; ear black or Sepia,
anterior margin Ochraceous-Tawny, posterior margin and postauricular patch grayish white; hairs
inside posterior part of pinna Cinnamon-Buff; median dorsal dark stripe black with Sayal Brown
along margins; lateral dark stripes black mixed with Sayal Brown; outermost dorsal dark stripes
obsolete, Sayal Brown mixed with black; median pair of dorsal light stripes Pale Smoke Gray mixed
with Clay Color; outer pair of dorsal light stripes creamy white; sides Clay Color; rump and thighs
Neutral Gray; dorsal surface of tail black mixed with Cinnamon-Buff; ventral surface of tail
Ochraceous-Tawny; hairs around margin of tail Cinnamon-Buff or Ochraceous-Tawny; antipalmar
and antiplantar surfaces of feet Cinnamon-Buff; underparts creamy white with dark underfur. Skull:
Large; zygomata strong; braincase well inflated. Baculum: As in E. u. umbrinus.
Comparisons.—From E. quadrivittatus quadrivittatus, the subspecies and species to the south, E. u.
montanus differs in: General tone of upper parts darker; braincase significantly narrower; baculum
shorter and markedly wider at base.
From E. u. umbrinus, the subspecies from the Uinta and northern Wasatch Mountains, E. u. montanus
differs in: General tone of upper parts brighter (less tawny); sides more tawny; skull slightly larger.

From E. u. sedulus, the subspecies from the Henry Mountains of Utah, E. u. montanus differs in:
Sides darker; general tone of upper parts darker.
From E. u. fremonti, the subspecies from the mountains of western and northwestern Wyoming, E. u.
montanus differs in: General tone of upper parts lighter; hairs around outermost edge of tail tawnier.

Remarks.—Howell (1929:83) stated that the specimens of E. quadrivittatus
quadrivittatus (= E. umbrinus montanus) from Estes Park, Long's Peak, and
Gold Hill, all in Colorado, "average somewhat darker on the back and sides
than typical quadrivittatus; the light dorsal stripes are also somewhat duller
and the dark stripes less blackish, thus showing an approach to the
characters of umbrinus." Now there are more specimens of E. u. montanus
from the mountains of north-central Colorado than were available to
Howell. He was not aware of the striking difference between the bacula of
E. quadrivittatus and E. umbrinus, and the constancy of this difference
between all the subspecies of one species and those of the other.
Although the geographic range of E. u. umbrinus is closer to the ranges of
E. u. fremonti and E. u. montanus than to the geographic range of E. u.
adsitus, E. u. umbrinus seems to be more closely related to E. u. adsitus
than to E. u. fremonti or E. u. montanus. This observation may be explained
by the presence of continuous habitat for E. umbrinus between the ranges of
E. u. umbrinus and E. u. adsitus, whereas E. u. fremonti and E. u. montanus
are each separated from E. u. umbrinus by areas unsuitable for occupancy
by E. umbrinus. It must be noted, however, that no actual intergrades
between E. u. umbrinus and E. u. adsitus are known.
Specimens examined.—Total number, 36.
Wyoming: Albany Co.: 3 mi. ESE Brown's Peak, 10,000 ft., 2; 3½ mi. S Wood's Landing, 1.
Utah: Uintah Co.: PR Springs, 7,950 ft., 43 mi. S Ouray, Uintah-Grand county line, 1 UU.
Colorado: Jackson Co.: Mount Zirkel, 10,000 ft., on trail, 2 WC; Buffalo Pass, 10,380 ft., 1 WC;
Buffalo Pass road, 10,130 ft., 1 WC. Larimer Co.: 2 mi. E Log Cabin, 7,450 ft., 1 WC; Estes Park,
7,600 ft., 1; 1½ mi. SW Estes Park, 1; 2½ mi. SW Estes Park, 2; 3½ mi. SW Estes Park, 1; 12 mi.
SW Estes Park, 1. Rio Blanco Co.: 1 mi. NW Pagoda Peak, 10,400 ft., 1. Boulder Co.: Long's Peak, 7
BS; 1 mi. NE Ward, 10,000 ft., 1; 3 mi. S Ward, 9,000 ft., 5; ½ mi. E and 3 mi. S Ward, 9,400 ft., 1; 1
mi. S Gold Hill, 8,200 ft., 1. Clear Creek Co.: Davidson Mine, 3 mi. SW Idaho Springs, 1;
Georgetown, 1 CM. Jefferson Co.: Silver Plume, 1 CM. Park Co.: Tarryall Creek Camp, 8,700 ft., 1
WC. Gunnison Co.: S side Crested Butte Mountain, 9,500 ft., 1 WC; mouth of Virginia Basin,
Gothic, 1 FC.

Discussion

The chipmunks that heretofore have been assigned to the species Eutamias
quadrivittatus are here assigned to two species, E. quadrivittatus and E.
umbrinus, for the following reasons:
1. The baculum of E. quadrivittatus differs from that of E. umbrinus in
having a narrow base (see figs. 3, 4). This difference permits any specimen
which has an associated baculum to be readily identified to species.
2. The cranial breadth in the subspecies of E. quadrivittatus is significantly
larger than in the subspecies of E. umbrinus.
3. Specimens of E. umbrinus are darker than any specimen of E.
quadrivittatus.
4. Where the geographic ranges of E. quadrivittatus and E. umbrinus come
close to one another (probably they meet at some places), E. umbrinus
occupies a higher position in terms of life-zones. Wherever either of these
two species, but not the other, occurs on a mountain the species occupies
both the higher and lower life-zones.

Figs. 3 and 4. Baculum of Eutamias quadrivittatus quadrivittatus, No.
35648/47919 BS; from Canon City, Fremont Co., Colorado. Figure 3,
lateral view of right side; figure 4, dorsal view.

Fig. 5. Baculum of Eutamias umbrinus umbrinus, No. 38062; from
Paradise Park, 21 mi. W and 15 mi. N Vernal, 10,050 ft., Uintah Co.,
Utah. Lateral view of right side.

Fig. 6. Baculum of Eutamias umbrinus montanus, No. 20105; from ½
mi. E and 3 mi. S Ward, 9,400 ft., Boulder Co., Colorado. Dorsal view.
The differences between E. quadrivittatus and E. umbrinus are as great as,
or greater than, between many species of chipmunks, such as between E.
minimus and E. amoenus, and between E. quadrivittatus and E.
cinereicollis.
Although I know of no ecological differences between E. umbrinus and E.
ruficaudus, the morphological differences, as for example, differences in
the structure of the baculum, and differences in color pattern, lead me to
maintain E. ruficaudus and E. umbrinus as separate species.
The present distribution of these two species is attributable to the uplift of
the Rocky Mountains in the Pleistocene. That the uplift of the Rocky
Mountains and the erosion which produced the present-day relief took place
in Pleistocene times is supported by the evidence found by several
geologists such as Hunt and Sokoloff (1950:109-123).
The present geographic distribution of E. umbrinus and E. quadrivittatus
conceivably came about as follows: E. umbrinus-like chipmunks were
present, before the uplift of the major chains of mountains, on isolated, low
mountain ranges that were not covered with glaciers (such as the laccolithic
mountains that occur in Utah) in Pleistocene time, while E. quadrivittatus-
like chipmunks were present in the central parts of New Mexico, Colorado,
and southern Wyoming. With the advent of uplift, the habitats in the central
parts of these states were changed from a plains-like habitat to a habitat that
resembled the forest habitats that exist today. E. umbrinus-like chipmunks
then invaded this newly formed habitat and displaced any E. quadrivittatus-
like chipmunks that were less well adapted to live there. The Colorado
River probably served as a barrier that kept the E. umbrinus-like chipmunks
and E. quadrivittatus-like chipmunks separated up to this time. Invasion of
the new forest-niche by E. umbrinus-like chipmunks may have taken place
through the Uinta Mountains of northeastern Utah, after the glaciers
disappeared from these mountains, since the Colorado River probably
prevented any eastward migration farther south.

Table 1

Average and Extreme Measurements in Millimeters of Adult
Eutamias quadrivittatus and E. umbrinus

Length
Condylo-
Greatest Length of
Zygomatic Cranial Total Length alveolar
length of lower
breadth breadth length of tail length of
of skull nasals tooth-
mandible
row

E. q. quadrivittatus, Canon City, Fremont Co., Colorado.
Mean (7) 35.7 19.3 16.2 11.0 222 99.4 5.40 18.98
Min ♂ 35.7 19.1 15.9 10.6 216 93.0 5.32 18.65
Max 35.8 19.8 16.5 11.6 230 104.0 5.49 19.41

Mean (3) 35.9 19.9 16.5 10.9 231 99.0 5.42 19.10
Min ♀ 35.6 19.9 16.5 10.7 200 98.0 5.39 18.85
Max 36.2 19.9 16.6 11.2 232 100.0 5.49 19.28

E. q. hopiensis, Moab, Grand Co., Utah.
Mean
34.4 19.4 16.3 10.0 212 90.7 5.19 18.36
(11)
33.5 19.2 15.9 9.3 208 85.0 4.92 17.80
Min ♂
35.4 20.0 16.8 10.5 220 96.0 5.38 18.96
Max
34.9 19.6 16.4 10.5 219 94.4 5.16 18.58
Mean (5)
34.2 19.3 15.9 10.1 210 85.0 5.13 18.00
Min ♀
35.7 20.1 16.6 10.7 228 104.0 5.20 19.19
Max

E. u. umbrinus, Mts. S Robertson, Uintah Co., Wyoming.
Mean 34.7 18.9 15.7 10.9 218 96.2 5.13 18.04
(11) 34.3 18.3 15.6 10.3 215 81.0 4.79 17.57
Min ♂ 35.2 19.4 16.0 11.7 228 112.0 5.42 18.59
Max
35.1 19.2 15.9 11.0 224 96.4 5.17 18.46
Mean (4) 34.9 18.6 15.7 10.3 204 90.0 5.11 18.31
35.4 20.0 16.2 11.8 234 100.0 5.22 18.98

Min ♀
Max

E. u. adsitus, Britts Meadow, Beaver Co., Utah.
Mean (6) 34.8 18.9 15.5 10.8 214 89.6 5.16 18.07
Min ♂ 34.3 18.5 15.3 10.4 203 73.0 4.64 17.69
Max 35.4 19.6 16.1 11.3 225 95.0 5.34 18.70

Mean (6) 35.1 19.5 16.0 11.0 228 96.5 5.11 18.75
Min ♀ 33.9 18.9 15.9 10.6 215 95.0 5.00 18.51
Max 36.2 20.0 16.3 11.8 233 98.0 5.33 19.40

E. u. sedulus, Mt. Ellen, Henry Mts., Garfield Co., Utah.
Mean (5) 34.7 18.7 15.6 10.7 218 93.0 5.21 18.74
Min ♂ 33.5 18.4 15.4 10.1 213 89.0 5.09 18.48
Max 35.5 19.1 15.9 11.2 224 97.0 5.28 19.38

Mean (2) 34.9 19.4 16.1 11.1 227 98.0 5.24 18.74
Min ♀ 34.9 19.3 16.1 11.0 224 96.0 5.07 19.24
Max 34.9 19.5 16.1 11.3 231 100.0 5.42 19.80

E. u. inyoensis, Ruby Mts., Elko and White Pine Cos., Nev.
Mean
34.4 19.0 15.7 10.5 208 89.5 5.15 18.12
(12)
33.5 18.6 15.2 10.0 196 85.0 5.01 17.32
Min ♂
35.4 19.6 16.1 11.5 220 100.0 5.37 18.81
Max
34.9 19.4 15.7 10.5 215 92.8 5.19 18.63
Mean (5)
34.4 19.1 15.4 10.4 204 86.0 5.04 18.50
Min ♀
35.2 19.7 16.0 10.7 226 102.0 5.33 18.80
Max

E. u. nevadensis, Measurements of the type (Burt 1931:300).
♂ 34.8 19.3 16.2 11.0 205 89.0 ... ...

E. u. fremonti, Togwotee Pass, Fremont Co., Wyoming.

Mean (8) 35.6 19.3 15.9 11.4 223 99.0 5.34 19.17
Min ♂ 35.2 18.9 15.8 11.1 216 95.0 5.22 18.72
Max 36.5 19.7 16.1 11.8 243 111.0 5.57 19.78

Mean (6) 35.3 19.6 15.9 11.3 229 101.0 5.40 19.02
Min ♀ 34.5 19.3 15.7 10.9 223 92.0 5.35 18.37
Max 36.0 20.0 16.5 12.0 239 110.0 5.44 19.51

E. u. montanus, Boulder Co., Colorado.
Mean (5) 35.2 18.8 15.5 10.8 226 96.0 5.20 18.29
Min ♂ 34.7 18.4 15.2 10.1 215 93.0 5.03 17.80
Max 36.8 19.4 16.2 11.5 232 115.0 5.53 19.36

Mean (6) 35.7 19.1 15.6 10.9 226 98.0 5.28 18.67
Min ♀ 35.1 18.8 15.1 10.3 215 89.0 5.06 18.09
Max 36.5 19.5 16.0 11.6 231 105.0 5.58 19.35

Literature Cited

Benson, S. B.
1935. A biological reconnaissance of Navajo Mountain, Utah. Univ. California Publ.
Zool., 40:439-455, December 31.

Burt, W. H.
1931. Three new subspecies of chipmunks of the genus Eutamias from Nevada. Jour.
Mamm., 12:298-301, August 24.

Durrant, S. D.
1952. Mammals of Utah, taxonomy and distribution. Univ. Kansas Publ. Mus. Nat.
Hist., 6:1-549, 91 figs., 30 tables, August 10.

Hall, E. R.
1946. Mammals of Nevada. Univ. California Press, Berkeley, California, pp. xi +
710, 11 pls., 485 figs., July 1.

Hardy, R.

1945. The taxonomic status of some chipmunks of the genus Eutamias in
southwestern Utah. Proc. Biol. Soc. Washington, 58:85-87, June 30.

Howell, A. H.
1929. Revision of the American chipmunks (genera Tamias and Eutamias). U. S.
Dept. Agric., Bur. Biol. Surv., N. Amer. Fauna, 52:1-157, 10 pls., 9 figs.,
November 30.

Hunt, C. B., and Sokoloff, V. P.
1950. Pre-Wisconsin soil in the Rocky Mountain region, a progress report. U. S.
Geol. Survey, Prof. Paper, 221-G:109-123.

Johnson, D. H.
1943. Systematic review of the chipmunks (genus Eutamias) of California. Univ.
California Publ. Zool., 48:63-148, 6 pls., December 24.

Kelson, K. R.
1951. Speciation in rodents of the Colorado River drainage. Univ. Utah Biol. Ser.,
11(3): vii + 125, 10 figs., February 15.

Merriam, C. H.
1905. Two new chipmunks from Colorado and Arizona. Proc. Biol. Soc. Washington,
18:163-166, June 29.

Transmitted June 26, 1953.

24-8966

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