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                      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.

    [Illustration: 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. × 1-3/4. 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 1/4 of length of shaft, shaft having a maximum diameter of
more than 1/4 mm., and height of keel 1/4 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 1/2 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.

    [Illustration: 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._: 1-1/2 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 1/3 of
length of shaft, distal 1/2 of shaft laterally compressed, and keel
1/4 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; 11-1/2 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._: 13-1/2 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.: 16-1/4 mi. N and
    17 mi. W Cody, 5,625 ft., 2. _Teton Co._: 1 mi. E and 1/4 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., 17-1/2 mi. W and
    2-1/2 mi. N Lander, 1; 17 mi. S and 6-1/2 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 1/2 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; 3-1/2 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; 1-1/2 mi. SW Estes
    Park, 1; 2-1/2 mi. SW Estes Park, 2; 3-1/2 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; 1/2 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.

    [Illustration: 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 1/2 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_

  KEY
  A: Greatest length of skull
  B: Zygomatic breadth
  C: Cranial breadth
  D: Length of nasals
  E: Total length
  F: Length of tail
  G: Length of lower tooth-row
  H: Condylo-alveolar length of mandible

======================================================================
             |  A   |  B   |  C   |  D   |  E  |   F   |  G   |  H
-------------+------+------+------+------+-----+-------+------+-------
             |
             | _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   [Male] | 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 [Female] | 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    (11) | 34.4 | 19.4 | 16.3 | 10.0 | 212 |  90.7 | 5.19 | 18.36
Min   [Male] | 33.5 | 19.2 | 15.9 |  9.3 | 208 |  85.0 | 4.92 | 17.80
Max          | 35.4 | 20.0 | 16.8 | 10.5 | 220 |  96.0 | 5.38 | 18.96
             |      |      |      |      |     |       |      |
Mean     (5) | 34.9 | 19.6 | 16.4 | 10.5 | 219 |  94.4 | 5.16 | 18.58
Min [Female] | 34.2 | 19.3 | 15.9 | 10.1 | 210 |  85.0 | 5.13 | 18.00
Max          | 35.7 | 20.1 | 16.6 | 10.7 | 228 | 104.0 | 5.20 | 19.19
             |      |      |      |      |     |       |      |
             +------+------+------+------+-----+-------+------+-------
             |
             | _E. u. umbrinus_, Mts. S Robertson, Uintah Co.,
             | Wyoming.
             |
Mean    (11) | 34.7 | 18.9 | 15.7 | 10.9 | 218 |  96.2 | 5.13 | 18.04
Min   [Male] | 34.3 | 18.3 | 15.6 | 10.3 | 215 |  81.0 | 4.79 | 17.57
Max          | 35.2 | 19.4 | 16.0 | 11.7 | 228 | 112.0 | 5.42 | 18.59
             |      |      |      |      |     |       |      |
Mean     (4) | 35.1 | 19.2 | 15.9 | 11.0 | 224 |  96.4 | 5.17 | 18.46
Min [Female] | 34.9 | 18.6 | 15.7 | 10.3 | 204 |  90.0 | 5.11 | 18.31
Max          | 35.4 | 20.0 | 16.2 | 11.8 | 234 | 100.0 | 5.22 | 18.98
             |      |      |      |      |     |       |      |
             +------+------+------+------+-----+-------+------+-------
             |
             | _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   [Male] | 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 [Female] | 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   [Male] | 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 [Female] | 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    (12) | 34.4 | 19.0 | 15.7 | 10.5 | 208 |  89.5 | 5.15 | 18.12
Min   [Male] | 33.5 | 18.6 | 15.2 | 10.0 | 196 |  85.0 | 5.01 | 17.32
Max          | 35.4 | 19.6 | 16.1 | 11.5 | 220 | 100.0 | 5.37 | 18.81
             |      |      |      |      |     |       |      |
Mean     (5) | 34.9 | 19.4 | 15.7 | 10.5 | 215 |  92.8 | 5.19 | 18.63
Min [Female] | 34.4 | 19.1 | 15.4 | 10.4 | 204 |  86.0 | 5.04 | 18.50
Max          | 35.2 | 19.7 | 16.0 | 10.7 | 226 | 102.0 | 5.33 | 18.80
             |      |      |      |      |     |       |      |
             +------+------+------+------+-----+-------+------+-------
             |
             | _E. u. nevadensis_, Measurements of the type
             | (Burt 1931:300).
             |
      [Male] | 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   [Male] | 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 [Female] | 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   [Male] | 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 [Female] | 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.
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    _Transmitted June 26, 1953._


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