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+The Project Gutenberg EBook of Taxonomy of the Chipmunks, Eutamias
+quadrivittatus and Eutamias umbrinus, by John A. White
+
+This eBook is for the use of anyone anywhere at no cost and with
+almost no restrictions whatsoever. You may copy it, give it away or
+re-use it under the terms of the Project Gutenberg License included
+with this eBook or online at www.gutenberg.org
+
+
+Title: Taxonomy of the Chipmunks, Eutamias quadrivittatus and Eutamias umbrinus
+
+Author: John A. White
+
+Release Date: February 12, 2010 [EBook #31267]
+
+Language: English
+
+Character set encoding: ISO-8859-1
+
+*** START OF THIS PROJECT GUTENBERG EBOOK TAXONOMY--CHIPMUNKS EUTAMIAS ***
+
+
+
+
+Produced by Chris Curnow, Joseph Cooper and the Online
+Distributed Proofreading Team at https://www.pgdp.net
+
+
+
+
+
+
+
+
+
+ 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 piņon 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 piņon 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.
+ 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
+
+
+
+
+
+End of the Project Gutenberg EBook of Taxonomy of the Chipmunks, Eutamias
+quadrivittatus and Eutamias umbrinus, by John A. White
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+The Project Gutenberg EBook of Taxonomy of the Chipmunks, Eutamias
+quadrivittatus and Eutamias umbrinus, by John A. White
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+Title: Taxonomy of the Chipmunks, Eutamias quadrivittatus and Eutamias umbrinus
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+*** START OF THIS PROJECT GUTENBERG EBOOK TAXONOMY--CHIPMUNKS EUTAMIAS ***
+
+
+
+
+Produced by Chris Curnow, Joseph Cooper and the Online
+Distributed Proofreading Team at https://www.pgdp.net
+
+
+
+
+
+
+</pre>
+
+
+
+<h1>Taxonomy of the Chipmunks,<br />
+Eutamias quadrivittatus and Eutamias umbrinus</h1>
+
+<h4>BY</h4>
+
+<h3>JOHN A. WHITE<br /><br /></h3>
+
+
+
+<h3>University of Kansas Publications<br />
+Museum of Natural History<br /><br />
+
+<small>Volume 5, No. 33, pp. 563-582, 6 figures in text</small><br />
+
+<small>December 1, 1953</small></h3>
+
+
+<h4>University of Kansas<br />
+LAWRENCE<br />
+1953</h4>
+
+<hr style="width: 65%;" />
+<h4><span class="smcap">University of Kansas Publications, Museum of Natural History</span><br /><br />
+Editors: E. Raymond Hall, Chairman, A. Byron Leonard,<br />
+and Robert W. Wilson<br />
+<br />
+Volume 5, No. 33, pp. 563-582, 6 figures in text<br />
+<br />
+December 1, 1953<br /><br /></h4>
+
+<h3><span class="smcap">University of Kansas</span><br />
+Lawrence, Kansas<br /><br /></h3>
+
+<h5><small>PRINTED BY<br />
+FERD VOILAND, JR., STATE PRINTER<br />
+TOPEKA, KANSAS<br />
+1953</small><br />
+<br />
+24-8966</h5>
+
+
+
+
+<hr style="width: 65%;" />
+<p><span class='pagenum'><a name="Page_565" id="Page_565">[Pg 565]</a></span></p>
+<h2>Taxonomy of the Chipmunks,<br />
+Eutamias quadrivittatus and Eutamias umbrinus</h2>
+
+<h4>By</h4>
+<h4>JOHN A. WHITE</h4>
+
+
+<p>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.</p>
+
+
+<h3><span class="smcap">Historical Summary</span></h3>
+
+<p>Merriam (1905) was the first to show clearly that <i>Eutamias
+quadrivittatus</i> is a distinct species, and pointed out that <i>E. amoenus
+operarius</i> (= <i>E. minimus operarius</i>) is a small species which resembles,
+and is found in some areas together with, <i>E. quadrivittatus</i>.</p>
+
+<p>Howell (1929) placed under <i>E. quadrivittatus</i> the following subspecies:
+<i>E. q. quadrivittatus</i>, <i>E. q. hopiensis</i>, <i>E. q. inyoensis</i>, <i>E. q.
+frater</i>, <i>E. q. sequoiensis</i>, and <i>E. q. speciosus</i>.</p>
+
+<p>Hardy (1945) placed <i>E. adsitus</i> under <i>E. quadrivittatus</i> as <i>E. q.
+adsitus</i>, and Kelson (1951) placed <i>E. umbrinus</i> under <i>E. quadrivittatus</i>
+as <i>E. q. umbrinus</i>.</p>
+
+<p>Johnson (1943) re-established <i>E. speciosus</i> as a separate species,
+and in California left only <i>E. q. inyoensis</i> in <i>E. quadrivittatus</i>.</p>
+
+<p>Thus, since 1943 the recognized subspecies of <i>E. quadrivittatus</i>
+have been: <i>E. q. quadrivittatus</i>, <i>E. q. hopiensis</i>, <i>E. q. inyoensis</i>,
+<i>E. q. nevadensis</i>, <i>E. q. umbrinus</i>, and <i>E. q. adsitus</i>.</p>
+
+
+<h3><span class="smcap">Methods, Materials, and Acknowledgments</span></h3>
+
+<div class="pblockquot"><p>Capitalized color terms, which are used in descriptions and
+comparisons, are of Ridgway, "Color Standards and Color Nomenclature,"
+Washington, D. C., 1912.</p>
+
+<p>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.</p>
+
+<p>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:</p></div>
+
+<p><span class='pagenum'><a name="Page_566" id="Page_566">[Pg 566]</a></span></p>
+
+<div class="pblockquot">
+<div class="blockquot"><p>
+AM&mdash;American Museum of Natural History.<br />
+BS&mdash;United States Biological Surveys Collection.<br />
+CM&mdash;Colorado Museum of Natural History.<br />
+DC&mdash;Collection of Donald R. Dickey (now the collection of the University of California at Los Angeles).<br />
+FC&mdash;Collection of James S. Findley.<br />
+KU&mdash;Museum of Natural History, University of Kansas.<br />
+MM&mdash;Museum of Zoology, University of Michigan.<br />
+NM&mdash;United States National Museum.<br />
+UU&mdash;Museum of Zoology, University of Utah.<br />
+WC&mdash;Collection of Edward R. Warren, Colorado College.</p></div>
+
+<p>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.</p>
+
+<p>The measurements of the skull were made as shown in figure 1.</p></div>
+
+<div class="figcenter" style="width: 100%;">
+<img src="images/fig1.jpg" width="100%" alt="FIG. 1" title="FIG. 1" />
+<p><b><span class="smcap">Fig. 1.</span> 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. &times; 1&frac34;. Based on <i>Eutamias ruficaudus
+ruficaudus</i>, from 6 mi. S St. Mary, 6500 ft., Glacier Co., Montana. A
+to A'&mdash;greatest length of skull; B to B'&mdash;length of nasals; C to
+C'&mdash;zygomatic breadth; D to D'&mdash;least interorbital constriction; E to
+E'&mdash;cranial breadth; F to F'&mdash;inner mandibular length; G to
+G'&mdash;condylo-alveolar length of mandible.</b></p>
+</div>
+
+<div class="pblockquot"><p>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.</p></div>
+<p><span class='pagenum'><a name="Page_567" id="Page_567">[Pg 567]</a></span></p>
+<div class="pblockquot"><p>Whenever two or more samples are stated to be significantly different, the
+meaning is that the difference is statistically significant.</p>
+
+<p>The geographic distribution of each subspecies and the localities of
+specimens or series of specimens are plotted on the map (fig. 2).</p>
+
+<p>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.</p>
+
+<p>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.</p>
+
+<p>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.</p></div>
+
+
+<h3><span class="smcap">Accounts of Species and Subspecies</span></h3>
+
+<h3><b>Eutamias quadrivittatus</b> (Say)</h3>
+
+<p><i>Diagnosis.</i>&mdash;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 &frac14; of length of shaft, shaft having a maximum
+diameter of more than &frac14; mm., and height of keel &frac14; of length
+of tip.</p>
+
+<p style='text-align:center;'><b>Eutamias quadrivittatus quadrivittatus</b> (Say)</p>
+
+<div class="pblockquot">
+<div class="blockquot"><p><i>Sciurus quadrivittatus</i> Say, in Jones, Long's Expedition to Rocky Mountains,
+2:45, 1823.</p>
+
+<p><i>Eutamias quadrivittatus</i>, Miller and Rehn, Proc. Boston Soc. Nat. Hist.,
+30:43, December 27, 1901.</p>
+
+<p><i>Tamias quadrivittatus gracilis</i> J. A. Allen, Bull. Amer. Mus. Nat.
+Hist., 3:99, June 1890, Type from San Pedro, Santa Fe Co., New Mexico.</p>
+
+<p><i>Eutamias quadrivittatus animosus</i> Warren, Proc. Biol. Soc.
+Washington, 22:105, June 25, 1909. Type from Irwin Ranch, Las Animas
+County, Colorado.</p></div>
+
+<p><i>Type.</i>&mdash;None designated; from along Arkansas River, about 26 mi. below
+Canon City, Fremont County, Colorado; obtained on July 18, 1820.</p>
+
+<p><i>Diagnosis.</i>&mdash;Size medium; dorsal dark stripes blackish; sides Cinnamon to
+Clay Color; crown Light Drab; baculum large.</p>
+
+<p><i>Description.</i>&mdash;<i>Color pattern</i>: 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
+<span class='pagenum'><a name="Page_568" id="Page_568">[Pg 568]</a></span>
+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. <i>Skull</i>: Large; braincase
+well inflated; zygomatic arches strong and slightly appressed to
+skull. <i>Baculum</i>: Large; long and slender.</p>
+
+<p><i>Comparisons.</i>&mdash;From <i>E. q. hopiensis</i>, the only other subspecies in
+this species, <i>E. q. quadrivittatus</i> differs in: Dorsal dark stripes
+blackish; crown grayer; rump and thighs grayer; general tone of upper
+parts darker.</p></div>
+
+<p><i>Remarks.</i>&mdash;Specimens from the Chuska Mountains, Zuni Mountains, and
+Blanco, New Mexico, are intergrades between <i>E. q. quadrivittatus</i> and
+<i>E. q. hopiensis</i>, but are referable to <i>E. q. quadrivittatus</i>.</p>
+
+<p>In north-central Colorado <i>E. umbrinus</i> occurs in the spruce and pine
+forests at higher altitudes, while to the south and east of this area
+<i>E. q. quadrivittatus</i> occurs in growths of pi&ntilde;on in lower, semiarid
+areas. In the northern half of New Mexico and in south-central
+Colorado, <i>E. q. quadrivittatus</i> occurs not only in semiarid habitats
+but also in the moist habitats of the forests of higher altitudes.
+Ecologically, <i>E. umbrinus</i> thus replaces <i>E. q. quadrivittatus</i> in
+north-central Colorado. This ecological replacement is comparable to
+the ecological replacement of <i>Thomomys bottae</i> by <i>T. talpoides</i> in
+Utah as shown by Durrant (1952:156).</p>
+
+<div class="pblockquot"><p><i>Specimens examined.</i>&mdash;Total number, 130.</p>
+
+<p><b>Colorado</b>: <i>Larimer Co.</i>: Arkins, 1 BS. <i>Jefferson Co.</i>: W spur Lookout
+Mountain, near Golden, 1 WC. <i>Gunnison Co.</i>: Sapinero, 3 BS. <i>Saguache
+Co.</i>: 5 mi. N and 22 mi. W Saguache 10,000 ft., 1; 21 mi. W and 3 mi.
+N Saguache, 1. <i>Fremont Co.</i>: 18 mi. S and 7 mi. W Colorado Springs,
+1; Arkansas River, "about" 26 mi. below Canon City, 15 BS. <i>San Juan
+Co.</i>: Silverton, 1 BS. <i>Mineral Co.</i>: 3 mi. E Creede, 1. <i>Alamosa
+Co.</i>: Sangre de Cristo Range, 24 mi. E Hooper, 2 CM. <i>La Plata Co.</i>: 2
+mi. NE Bondad 6,100 ft., 1; Bondad, 15 mi. S Durango 6,050 ft., 1.
+<i>Archuleta Co.</i>: Chromo, 1 CM. <i>Las Animas Co.</i>: Trinidad, 6 BS. <i>Baca
+Co.</i>: unspecified, 1.</p>
+
+<p><b>New Mexico</b>: <i>San Juan Co.</i>: Blanco, 1 BS; Chuska Mountains, 8 BS. <i>Rio
+Arriba Co.</i>: 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 &frac12; mi. S Truchas, 8,500 ft., 1; 2 mi. S and 4 mi. W Coyote, 8,100
+ft., 1; unspecified, 2. <i>Taos Co.</i>: 3 mi. N Taos Pueblo, 5 BS; 23 mi.
+S and 6 mi. E Taos, 8,750 ft., 2. <i>Union Co.</i>: Emery Peak, 1 BS;
+Folsom, 3 BS; Sierra Grande, 8 BS; unspecified, 2. <i>McKinley Co.</i>:
+Bear Ridge, Zuni Mountains, 9 BS. <i>Sandoval Co.</i>: Bear Canyon, W
+foothills, Sandia Mountains, 3 BS; W foothills, near S end, Sandia
+Mountains, 7 BS. <i>Santa Fe Co.</i>: San Pedro, 7 BS. <i>San Miguel Co.</i>:
+Canadian River, 4 mi. NW Tucumcari, 1 BS. <i>Valencia Co.</i>: Mount
+Taylor, San Mateo Mountains, 10 BS.</p>
+
+<p><b>Oklahoma</b>: <i>Cimarron Co.</i>: Kenton, 1 BS.</p></div>
+
+<p><span class='pagenum'><a name="Page_569" id="Page_569">[Pg 569]</a></span></p>
+
+<p style='text-align:center;'><b>Eutamias quadrivittatus hopiensis</b> Merriam</p>
+
+<div class="pblockquot">
+<div class="blockquot"><p><i>Eutamias hopiensis</i> Merriam, Proc. Biol. Soc. Washington, 18:165, June 29,
+1905.</p>
+
+<p><i>Eutamias quadrivittatus hopiensis</i>, Howell, Jour. Mamm. 3:184, August 4,
+1922.</p></div>
+
+<p><i>Type.</i>&mdash;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.</p>
+
+<p><i>Diagnosis.</i>&mdash;Size medium; dorsal dark stripes tawny; crown Drab-Gray;
+baculum of same proportions as in <i>E. q. quadrivittatus</i> but smaller.</p></div>
+
+<div class="figcenter" style="width: 100%;">
+<img src="images/fig2.jpg" width="100%" alt="FIG. 1" title="FIG. 1" />
+<p><b><span class="smcap">Fig. 2.</span> Localities of specimens examined and probable
+geographic ranges of the subspecies of <i>Eutamias quadrivittatus</i> and
+<i>Eutamias umbrinus</i>. The symbols for locality records are as follows:
+circles, precise localities; triangles, localities known only to
+county.<br /><br />
+Guide to subspecies:<br />
+1. <i>E. q. quadrivittatus</i><br />
+2. <i>E. q. hopiensis</i><br />
+3. <i>E. u. umbrinus</i><br />
+4. <i>E. u. adsitus</i><br />
+5. <i>E. u. sedulus</i><br />
+6. <i>E. u. inyoensis</i><br />
+7. <i>E. u. nevadensis</i><br />
+8. <i>E. u. fremonti</i><br />
+9. <i>E. u. montanus</i>
+</b></p>
+</div>
+
+<div class="pblockquot"><p><i>Description.</i>&mdash;<i>Color pattern</i>: 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
+<span class='pagenum'><a name="Page_570" id="Page_570">[Pg 570]</a></span>
+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. <i>Skull</i>: As in <i>E. q. quadrivittatus</i>.
+<i>Baculum</i>: Same proportions as in <i>E. q. quadrivittatus</i> but smaller.</p>
+
+<p><i>Comparisons.</i>&mdash;See under the account of <i>E. q. quadrivittatus</i>.</p></div>
+
+<p><i>Remarks.</i>&mdash;Topotypes of this subspecies are intergrades between
+it and <i>E. q. quadrivittatus</i>.</p>
+
+<p>In a large part of the geographic range of <i>E. q. hopiensis</i> 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 <i>E. q. hopiensis</i> 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 pi&ntilde;on 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 <i>E. q. hopiensis</i>
+with either <i>E. q. adsitus</i> [= <i>E. umbrinus adsitus</i>] or <i>E. q.
+umbrinus</i> [= <i>E. u. umbrinus</i>]." Benson (1935:449) states, "On Navajo
+Mountain these chipmunks [<i>E. q. hopiensis</i>] 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 <i>E. q. hopiensis</i>
+occurs on a mountain this subspecies goes higher than on a mountain
+where <i>E. u. adsitus</i> also occurs. This same relationship between <i>E.
+q. quadrivittatus</i> and the subspecies of <i>E. umbrinus</i> that occurs in
+north-central Colorado was pointed out in the account of <i>E. q.
+quadrivittatus</i>.</p>
+
+<div class="pblockquot"><p><i>Specimens examined.</i>&mdash;Total number, 68.</p>
+
+<p><b>Utah</b>: <i>Uintah Co.</i>: E side of confluence of Green and White rivers, 1 mi.
+SE Ouray, 4,700 ft., 3 UU. <i>Grand Co.</i>: 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. <i>Wayne
+Co.</i>: Fruita, 1 UU.</p>
+<p><span class='pagenum'><a name="Page_571" id="Page_571">[Pg 571]</a></span>
+<b>Colorado</b>: <i>Moffat Co.</i>: 11 mi. W and 11 mi. N Rangely, 6,000 ft., 3.
+<i>Rio Blanco Co.</i>: White River, 5 BS. <i>Eagle Co.</i>: McCoy, 2 BS. <i>Mesa
+Co.</i>: 1&frac12; mi. S Loma, 4,600 ft., 1. <i>Gunnison Co.</i>: 1 mi. E
+Somerset, 6,100 ft., 1. <i>Montrose Co.</i>: 1 mi. E Naturita, 5,900 ft.,
+1. <i>Dolores Co.</i>: 1 mi. N Cahone, 6,900 ft., 1. <i>Montezuma Co.</i>: 1 mi.
+S Cortez, 5,000 ft., 1; Mesa Verde, 25 mi. SW Mancos, 7,000 ft., 2 BS.</p>
+
+<p><b>Arizona</b>: <i>Navajo Co.</i>: Keams Canyon, 80 mi. N Holbrook, 15 BS. <i>Apache
+Co.</i>: 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.</p></div>
+
+
+<h3><b>Eutamias umbrinus</b> (J. A. Allen)</h3>
+
+
+<p><i>Diagnosis.</i>&mdash;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 &#8531; of length of shaft, distal &frac12; of shaft laterally
+compressed, and keel &frac14; of length of tip.</p>
+
+<p style='text-align:center;'><b>Eutamias umbrinus umbrinus</b> (J. A. Allen)</p>
+
+<div class="pblockquot">
+<div class="blockquot"><p><i>Tamias umbrinus</i> J. A. Allen, Bull. Amer. Mus. Nat. Hist. 3:96, June, 1890.</p>
+
+<p><i>Eutamias umbrinus</i>, Miller and Rehn, Proc. Boston Soc. Nat. Hist. 30:45,
+December 27, 1901.</p></div>
+
+<p><i>Type.</i>&mdash;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.</p>
+
+<p><i>Diagnosis.</i>&mdash;Size medium; general tone of upper parts dark and shadowy;
+skull relatively small.</p>
+
+<p><i>Description.</i>&mdash;<i>Color pattern</i>: 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. <i>Skull</i>: Large, with moderately
+inflated braincase and well developed zygomata. <i>Baculum</i>: One of the
+largest in the species.</p>
+
+<p><i>Comparisons.</i>&mdash;From <i>Eutamias umbrinus adsitus</i>, the subspecies to
+the south on the Wasatch Range, <i>E. u. umbrinus</i> differs in: Sides
+lighter; rump browner; hairs around outermost edge of tail tawnier (in
+freshly molted tails); shorter inner mandibular length.</p>
+
+<p>From <i>E. u. inyoensis</i>, the subspecies to the west in central and
+northeastern Nevada and in northwestern Utah, <i>E. u. umbrinus</i> differs
+in: General tone of upper parts lighter; sides lighter; total length
+more; interorbital region broader.</p>
+
+<p>For comparisons with <i>E. u. sedulus</i>, <i>E. u. fremonti</i>, and <i>E. u.
+montanus</i>, see the accounts of those subspecies.</p>
+
+<p><i>Specimens examined.</i>&mdash;Total number, 55.</p></div>
+<p><span class='pagenum'><a name="Page_572" id="Page_572">[Pg 572]</a></span></p>
+<div class="pblockquot"><p><b>Wyoming</b>: <i>Uinta Co.</i>: 9 mi. S Robertson, 8,000 ft., 15; 10 mi. S and 1
+mi. W Robertson, 8,700 ft., 5; 11&frac12; 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.</p>
+
+<p><b>Utah</b>: <i>Rich Co.</i>: Monte Cristo, 18 mi. W Woodruff, 8,000 ft., 2 UU.
+<i>Summit Co.</i>: 13&frac12; mi. S and 2 mi. E Robertson [Wyoming], 4; 1 mi. N
+Bridger Lake R. S., 9,400 ft., 4. <i>Wasatch Co.</i>: Snake Creek Canyon, 3
+mi. NW Midway, 6,000 ft., 1 UU. <i>Uintah Co.</i>: Paradise Park, 21 mi. W
+and 15 mi. N Vernal, 10,050 ft., 20.</p></div>
+
+
+<p style='text-align:center;'><b>Eutamias umbrinus adsitus</b> J. A. Allen</p>
+
+<div class="pblockquot">
+<div class="blockquot"><p><i>Eutamias adsitus</i> J. A. Allen, Brooklyn Institute Mus. Sci. Bull.
+1:118, March 31, 1905.</p></div>
+
+<p><i>Type.</i>&mdash;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.</p>
+
+<p><i>Diagnosis.</i>&mdash;Size medium; sides dark; general tone of upper parts
+dark; dorsal light and dark stripes strongly contrasting.</p>
+
+<p><i>Description.</i>&mdash;<i>Color pattern</i>: 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. <i>Skull</i> and <i>Baculum</i>: As in <i>E. u. umbrinus</i>.</p>
+
+<p><i>Comparisons.</i>&mdash;From <i>E. u. inyoensis</i>, the subspecies to the west,
+<i>E. u. adsitus</i> differs in: General tone of upper parts darker; sides
+darker; interorbital region wider; skull significantly deeper.</p>
+
+<p>For comparison with <i>E. u. umbrinus</i>, <i>E. u. sedulus</i>, and <i>E. u.
+montanus</i>, see the accounts of those subspecies.</p></div>
+
+<p><i>Remarks.</i>&mdash;Specimens from West Rim, Zion National Park, 6,500
+ft., Washington County, Utah, seem to be intergrades between <i>E. u.
+adsitus</i> and <i>E. u. inyoensis</i>, and are referable to <i>E. u. adsitus</i>.</p>
+
+<div class="pblockquot"><p><i>Specimens examined.</i>&mdash;Total number, 34.</p>
+
+<p><b>Utah</b>: <i>Beaver Co.</i>: Britts Meadow, Beaver Range Mountains, 8,500 ft., 13
+BS. Wayne Co.: Donkey Lake, Boulder Mountain, 10,000 ft., 4 UU. <i>Garfield
+Co.</i>: Wildcat R. S., Boulder Mountain, 8,700 ft., 5 UU.</p>
+
+<p><b>Arizona</b>: <i>Coconino Co.</i>: De Motte Park, Kaibab Plateau, 3 BS; Bright Angel,
+Kaibab Plateau, 9 BS.</p></div>
+
+<p><span class='pagenum'><a name="Page_573" id="Page_573">[Pg 573]</a></span></p>
+
+<p style='text-align:center;'><b>Eutamias umbrinus sedulus</b> new subspecies</p>
+
+<div class="pblockquot"><p><i>Type.</i>&mdash;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.</p>
+
+<p><i>Diagnosis.</i>&mdash;Size medium; general tone of upper parts dark
+reddish-brown; ventral surface of tail Ochraceous-Orange; sides Mars
+Yellow.</p>
+
+<p><i>Description.</i>&mdash;<i>Color pattern</i>: 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.
+<i>Skull</i>: Large; braincase moderately inflated; zygomata strong.
+<i>Baculum</i>: As in <i>E. u. umbrinus</i>.</p>
+
+<p><i>Comparisons.</i>&mdash;From <i>E. u. umbrinus</i>, the subspecies from the Uinta
+and northern Wasatch Mountains of Utah, <i>E. u. sedulus</i> differs in:
+General tone of upper parts lighter; sides lighter.</p>
+
+<p>From <i>E. u. adsitus</i>, the subspecies from the southern Wasatch Range
+in Utah and Kaibab Plateau in Arizona, <i>E. u. sedulus</i> differs in:
+Sides lighter; general tone of upper parts markedly lighter.</p>
+
+<p>From <i>E. u. inyoensis</i>, the subspecies from central and northeastern
+Nevada, and western and northwestern Utah, <i>E. u. sedulus</i> differs in:
+Sides lighter (less grayish); general tone of upper parts tawnier.</p>
+
+<p>For comparison with <i>E. u. montanus</i>, see the account of that
+subspecies.</p>
+
+<p><i>Specimens examined.</i>&mdash;Total number, 7 BS, all from the type locality.</p></div>
+
+
+<p style='text-align:center;'><b>Eutamias umbrinus inyoensis</b> Merriam</p>
+
+<div class="pblockquot">
+<div class="blockquot"><p><i>Eutamias speciosus inyoensis</i> Merriam, Proc. Biol. Soc. Washington, 11:202,
+208, July 1, 1897.</p></div>
+
+<p><i>Type.</i>&mdash;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.</p>
+
+<p><i>Diagnosis.</i>&mdash;Size medium; sides light; general tone of upper parts light;
+baculum one of largest in species.</p>
+
+<p><i>Description.</i>&mdash;<i>Color pattern</i>: 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
+<span class='pagenum'><a name="Page_574" id="Page_574">[Pg 574]</a></span>
+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. <i>Skull</i>: Large;
+zygomata strong; braincase moderately inflated. <i>Baculum</i>: One of
+largest in species.</p>
+
+<p><i>Comparisons.</i>&mdash;For comparisons with <i>E. u. umbrinus</i>, <i>E. u. adsitus</i>, <i>E. u.
+sedulus</i>, and <i>E. u. nevadensis</i>, see the accounts of those subspecies.</p></div>
+
+<p><i>Remarks.</i>&mdash;The baculum in <i>E. u. inyoensis</i> is like that in <i>E. palmeri</i>.</p>
+
+<div class="pblockquot"><p><i>Specimens examined.</i>&mdash;Total number, 46.</p>
+
+<p><b>Nevada</b>: <i>Elko Co.</i>: 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. <i>White Pine Co.</i>: 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.</p>
+
+<p><b>Utah</b>: <i>Boxelder Co.</i>: Head of George Creek and Clear Creek, 5 mi. S
+Stanrod, Raft River Mountains, 8,500 ft., 2 UU.</p></div>
+
+
+<p style='text-align:center;'><b>Eutamias umbrinus nevadensis</b> Burt</p>
+
+<div class="pblockquot">
+<div class="blockquot"><p><i>Eutamias quadrivittatus nevadensis</i> Burt, Jour. Mamm. 12:299, August 24,
+1931.</p></div>
+
+<p><i>Type.</i>&mdash;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.</p>
+
+<p><i>Diagnosis.</i>&mdash;Size medium; general tone of upper parts grayish;
+baculum one of the largest of species.</p>
+
+<p><i>Description.</i>&mdash;"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 <i>E. u. inyoensis</i> but differing as indicated below.</p>
+
+<p><i>Comparisons.</i>&mdash;From <i>E. u. inyoensis</i>, the subspecies to the north,
+<i>E. u. nevadensis</i> differs in: Paler and grayer throughout; tawny areas
+restricted; gray areas clearer and less suffused; dark facial markings narrower and less
+<span class='pagenum'><a name="Page_575" id="Page_575">[Pg 575]</a></span>
+distinct; ventral surface of tail distinctly paler; feet lighter, clearer gray; nasals extend farther
+posteriorly with respect to premaxillae (Burt <i>loc. cit.</i>).</p>
+
+<p>From <i>E. u. adsitus</i>, the subspecies to the northeast, <i>E. u.
+nevadensis</i> differs in: Narrower dorsal stripes and facial markings;
+paler coloration of head, rump, sides, feet, and ventral surface of
+tail (Burt <i>op. cit.</i>: 299-300).</p></div>
+
+<p><i>Remarks.</i>&mdash;The differences between <i>E. umbrinus nevadensis</i> and
+<i>E. palmeri</i>, as shown by Burt (<i>op. cit.</i>) and Hall (1946), are such
+that one might expect <i>E. palmeri</i> to be a subspecies of <i>E. umbrinus</i>.
+However, having only the structure of the baculum as evidence
+additional to that summarized by Hall (<i>op. cit.</i>), I follow him in
+according <i>E. palmeri</i> specific status.</p>
+
+<div class="pblockquot"><p><i>Specimens examined.</i>&mdash;None.</p></div>
+
+
+<p style='text-align:center;'><b>Eutamias umbrinus fremonti</b> new subspecies</p>
+
+<div class="pblockquot"><p><i>Type.</i>&mdash;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.</p>
+
+<p><i>Diagnosis.</i>&mdash;Size large; sides Capucine Yellow; antiplantar surface of hind
+feet Raw Sienna; postauricular patch grayish white; baculum as in <i>E. u.
+umbrinus</i>.</p>
+
+<p><i>Description.</i>&mdash;<i>Color pattern</i>: 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. <i>Skull</i>: Large, with strong zygomata; braincase well
+inflated. <i>Baculum</i>: As in <i>E. u. umbrinus</i>.</p>
+
+<p><i>Comparisons.</i>&mdash;From <i>E. u. umbrinus</i>, the subspecies from the Uinta
+and northern Wasatch Mountains in Utah, <i>E. u. fremonti</i> differs in:
+Sides darker; antiplantar surfaces of feet darker; postauricular patch
+grayer; crown more grayish; skull slightly larger.</p>
+
+<p>From <i>E. ruficaudus ruficaudus</i>, the species and subspecies from
+western Montana, <i>E. u. fremonti</i> 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.</p>
+
+<p>For comparison with <i>E. u. montanus</i>, see the account of that
+subspecies.</p></div>
+
+<p><i>Remarks.</i>&mdash;The geographic ranges of <i>E. umbrinus fremonti</i> and
+<i>E. ruficaudus ruficaudus</i> are allopatric and no specimens have ever
+been taken in the intermediate area to indicate whether or not
+<span class='pagenum'><a name="Page_576" id="Page_576">[Pg 576]</a></span>
+these two species anywhere occur together. The bacula in the two
+species differ to the same degree as those of <i>E. quadrivittatus</i> and
+<i>E. umbrinus</i>. The differences between <i>E. u. fremonti</i> and <i>E. r.
+ruficaudus</i> are such that in my opinion, <i>E. ruficaudus</i> is a distinct
+species.</p>
+
+<div class="pblockquot"><p><i>Specimens examined.</i>&mdash;Total number, 58.</p>
+
+<p><b>Montana</b>: <i>Park Co.</i>: Beartooth Mountains, 2 BS.</p>
+
+<p><b>Idaho</b>: <i>Bonneville Co.</i>: Big Hole Mountains, 9,000 ft., near Irwin, 1 BS.</p>
+
+<p><b>Wyoming</b>: <i>Yellowstone Park</i>, 2. Park Co.: 16&frac14; mi. N and 17 mi. W
+Cody, 5,625 ft., 2. <i>Teton Co.</i>: 1 mi. E and &frac14; 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. <i>Lincoln Co.</i>: La Barge Creek, 9,000
+ft., 2 BS. <i>Fremont Co.</i>: Togwotee Pass, 12; 12 mi. N and 3 mi. W
+Shoshoni, 4,650 ft., 1; Mosquito Park R. S., 9,500 ft., 17&frac12; mi. W
+and 2&frac12; mi. N Lander, 1; 17 mi. S and 6&frac12; mi. W Lander, 8,450 ft.,
+3. <i>Sublette Co.</i>: 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.</p></div>
+
+
+<p style='text-align:center;'><b>Eutamias umbrinus montanus</b> new subspecies</p>
+
+<div class="pblockquot"><p><i>Type.</i>&mdash;Male, adult, skull, skin, and baculum, No. 20105 Univ. Kansas
+Mus. Nat. Hist.; from &frac12; 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.</p>
+
+<p><i>Diagnosis.</i>&mdash;Size large; sides Clay Color; antipalmar and antiplantar
+surfaces of feet Cinnamon-Buff; baculum as in E. u. umbrinus.</p>
+
+<p><i>Description.</i>&mdash;<i>Color pattern</i>: 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. <i>Skull</i>: Large; zygomata strong; braincase well
+inflated. <i>Baculum</i>: As in <i>E. u. umbrinus</i>.</p>
+
+<p><i>Comparisons.</i>&mdash;From <i>E. quadrivittatus quadrivittatus</i>, the
+subspecies and species to the south, <i>E. u. montanus</i> differs in:
+General tone of upper parts darker; braincase significantly narrower;
+baculum shorter and markedly wider at base.</p>
+
+<p>From <i>E. u. umbrinus</i>, the subspecies from the Uinta and northern
+Wasatch Mountains, <i>E. u. montanus</i> differs in: General tone of upper
+parts brighter (less tawny); sides more tawny; skull slightly larger.</p></div>
+
+<p><span class='pagenum'><a name="Page_577" id="Page_577">[Pg 577]</a></span></p>
+
+<div class="pblockquot"><p>From <i>E. u. sedulus</i>, the subspecies from the Henry Mountains of Utah,
+<i>E. u. montanus</i> differs in: Sides darker; general tone of upper parts
+darker.</p>
+
+<p>From <i>E. u. fremonti</i>, the subspecies from the mountains of western
+and northwestern Wyoming, <i>E. u. montanus</i> differs in: General tone of
+upper parts lighter; hairs around outermost edge of tail tawnier.</p></div>
+
+<p><i>Remarks.</i>&mdash;Howell (1929:83) stated that the specimens of <i>E.
+quadrivittatus quadrivittatus</i> (= <i>E. umbrinus montanus</i>) from Estes
+Park, Long's Peak, and Gold Hill, all in Colorado, "average somewhat
+darker on the back and sides than typical <i>quadrivittatus</i>; the light
+dorsal stripes are also somewhat duller and the dark stripes less
+blackish, thus showing an approach to the characters of <i>umbrinus</i>."
+Now there are more specimens of <i>E. u. montanus</i> from the mountains of
+north-central Colorado than were available to Howell. He was not aware
+of the striking difference between the bacula of <i>E. quadrivittatus</i>
+and <i>E. umbrinus</i>, and the constancy of this difference between all
+the subspecies of one species and those of the other.</p>
+
+<p>Although the geographic range of <i>E. u. umbrinus</i> is closer to the
+ranges of <i>E. u. fremonti</i> and <i>E. u. montanus</i> than to the geographic
+range of <i>E. u. adsitus</i>, <i>E. u. umbrinus</i> seems to be more closely
+related to <i>E. u. adsitus</i> than to <i>E. u. fremonti</i> or <i>E. u.
+montanus</i>. This observation may be explained by the presence of
+continuous habitat for <i>E. umbrinus</i> between the ranges of <i>E. u.
+umbrinus</i> and <i>E. u. adsitus</i>, whereas <i>E. u. fremonti</i> and <i>E. u.
+montanus</i> are each separated from <i>E. u. umbrinus</i> by areas unsuitable
+for occupancy by <i>E. umbrinus</i>. It must be noted, however, that no
+actual intergrades between <i>E. u. umbrinus</i> and <i>E. u. adsitus</i> are
+known.</p>
+
+<div class="pblockquot"><p><i>Specimens examined.</i>&mdash;Total number, 36.</p>
+
+<p><b>Wyoming</b>: <i>Albany Co.</i>: 3 mi. ESE Brown's Peak, 10,000 ft., 2; 3&frac12; mi. S
+Wood's Landing, 1.</p>
+
+<p><b>Utah</b>: <i>Uintah Co.</i>: PR Springs, 7,950 ft., 43 mi. S Ouray, Uintah-Grand
+county line, 1 UU.</p>
+
+<p><b>Colorado</b>: <i>Jackson Co.</i>: Mount Zirkel, 10,000 ft., on trail, 2 WC;
+Buffalo Pass, 10,380 ft., 1 WC; Buffalo Pass road, 10,130 ft., 1 WC.
+<i>Larimer Co.</i>: 2 mi. E Log Cabin, 7,450 ft., 1 WC; Estes Park, 7,600
+ft., 1; 1&frac12; mi. SW Estes Park, 1; 2&frac12; mi. SW Estes Park, 2; 3&frac12;
+mi. SW Estes Park, 1; 12 mi. SW Estes Park, 1. <i>Rio Blanco Co.</i>: 1 mi.
+NW Pagoda Peak, 10,400 ft., 1. <i>Boulder Co.</i>: Long's Peak, 7 BS; 1 mi.
+NE Ward, 10,000 ft., 1; 3 mi. S Ward, 9,000 ft., 5; &frac12; mi. E and 3
+mi. S Ward, 9,400 ft., 1; 1 mi. S Gold Hill, 8,200 ft., 1. <i>Clear
+Creek Co.</i>: Davidson Mine, 3 mi. SW Idaho Springs, 1; Georgetown, 1
+CM. <i>Jefferson Co.</i>: Silver Plume, 1 CM. <i>Park Co.</i>: Tarryall Creek
+Camp, 8,700 ft., 1 WC. <i>Gunnison Co.</i>: S side Crested Butte Mountain,
+9,500 ft., 1 WC; mouth of Virginia Basin, Gothic, 1 FC.</p></div>
+
+<p><span class='pagenum'><a name="Page_578" id="Page_578">[Pg 578]</a></span></p>
+
+<h3><span class="smcap">Discussion</span></h3>
+
+<p>The chipmunks that heretofore have been assigned to the species
+<i>Eutamias quadrivittatus</i> are here assigned to two species, <i>E.
+quadrivittatus</i> and <i>E. umbrinus,</i> for the following reasons:</p>
+
+<p>1. The baculum of <i>E. quadrivittatus</i> differs from that of <i>E.
+umbrinus</i> 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.</p>
+
+<p>2. The cranial breadth in the subspecies of <i>E. quadrivittatus</i> is
+significantly larger than in the subspecies of <i>E. umbrinus</i>.</p>
+
+<p>3. Specimens of <i>E. umbrinus</i> are darker than any specimen of <i>E.
+quadrivittatus</i>.</p>
+
+<p>4. Where the geographic ranges of <i>E. quadrivittatus</i> and <i>E.
+umbrinus</i> come close to one another (probably they meet at some
+places), <i>E. umbrinus</i> 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.</p>
+
+<div class="figcenter" style="width: 100%;">
+<img src="images/fig3.jpg" width="100%" alt="FIGS. 3, 4, 5, 6" title="FIGS. 3, 4, 5, 6" />
+<p><b><span class="smcap">Figs.</span> 3 and 4. Baculum of <i>Eutamias quadrivittatus
+quadrivittatus</i>, No. 35648/47919 BS; from Canon City, Fremont Co.,
+Colorado. Figure 3, lateral view of right side; figure 4, dorsal view.<br /><br />
+
+<span class="smcap">Fig. 5.</span> Baculum of <i>Eutamias umbrinus umbrinus</i>, No. 38062; from
+Paradise Park, 21 mi. W and 15 mi. N Vernal, 10,050 ft., Uintah Co.,
+Utah. Lateral view of right side.<br /><br />
+
+<span class="smcap">Fig. 6.</span> Baculum of <i>Eutamias umbrinus montanus</i>, No. 20105; from &frac12;
+mi. E and 3 mi. S Ward, 9,400 ft., Boulder Co., Colorado. Dorsal view.</b></p>
+</div>
+
+<p><span class='pagenum'><a name="Page_579" id="Page_579">[Pg 579]</a></span>
+The differences between <i>E. quadrivittatus</i> and <i>E. umbrinus</i> are as
+great as, or greater than, between many species of chipmunks, such as
+between <i>E. minimus</i> and <i>E. amoenus</i>, and between <i>E. quadrivittatus</i>
+and <i>E. cinereicollis</i>.</p>
+
+<p>Although I know of no ecological differences between <i>E. umbrinus</i> and
+<i>E. ruficaudus</i>, the morphological differences, as for example,
+differences in the structure of the baculum, and differences in color
+pattern, lead me to maintain <i>E. ruficaudus</i> and <i>E. umbrinus</i> as
+separate species.</p>
+
+<p>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).</p>
+
+<p>The present geographic distribution of <i>E. umbrinus</i> and <i>E.
+quadrivittatus</i> conceivably came about as follows: <i>E. umbrinus</i>-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 <i>E. quadrivittatus</i>-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. <i>E.
+umbrinus</i>-like chipmunks then invaded this newly formed habitat and
+displaced any <i>E. quadrivittatus</i>-like chipmunks that were less
+well adapted to live there. The Colorado River probably served as a
+barrier that kept the <i>E. umbrinus</i>-like chipmunks and <i>E.
+quadrivittatus</i>-like chipmunks separated up to this time. Invasion of
+the new forest-niche by <i>E. umbrinus</i>-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.</p>
+
+<p><span class='pagenum'><a name="Page_580" id="Page_580">[Pg 580]</a></span></p>
+
+<h4><span class="smcap">Table 1</span></h4>
+
+<h4>Average and Extreme Measurements in Millimeters of Adult<br />
+<i>Eutamias quadrivittatus</i> and <i>E. umbrinus</i></h4>
+
+<table border="1" cellpadding="5" cellspacing="0" summary="Table of Contents">
+<colgroup><col width="36%" /><col width="8%" /><col width="8%" />
+<col width="8%" /><col width="8%" /><col width="8%" />
+<col width="8%" /><col width="8%" /><col width="8%" /></colgroup>
+<tr align="center">
+ <td>&nbsp;</td>
+ <td>Greatest length of skull</td>
+ <td>Zygomatic breadth</td>
+ <td>Cranial breadth</td>
+ <td>Length of nasals</td>
+ <td>Total length</td>
+ <td>Length of tail</td>
+ <td>Length of lower tooth-row</td>
+ <td>Condylo-alveolar length of mandible</td>
+</tr>
+<tr>
+ <td>&nbsp;</td>
+ <td colspan="8" align="center"><br /><i>E. q. quadrivittatus</i>, Canon City, Fremont Co., Colorado.</td>
+</tr>
+<tr>
+ <td>Mean (7)<br />Min &#9794;<br />Max<br /><br />Mean (3)<br />Min &#9792;<br />Max</td>
+ <td align="center">35.7<br />35.7<br />35.8<br /><br />35.9<br />35.6<br />36.2</td>
+ <td align="center">19.3<br />19.1<br />19.8<br /><br />19.9<br />19.9<br />19.9</td>
+ <td align="center">16.2<br />15.9<br />16.5<br /><br />16.5<br />16.5<br />16.6</td>
+ <td align="center">11.0<br />10.6<br />11.6<br /><br />10.9<br />10.7<br />11.2</td>
+ <td align="center">222<br />216<br />230<br /><br />231<br />200<br />232</td>
+ <td align="center">99.4<br />93.0<br />104.0<br /><br />99.0<br />98.0<br />100.0</td>
+ <td align="center">5.40<br />5.32<br />5.49<br /><br />5.42<br />5.39<br />5.49</td>
+ <td align="center">18.98<br />18.65<br />19.41<br /><br />19.10<br />18.85<br />19.28</td>
+</tr>
+<tr>
+ <td>&nbsp;</td>
+ <td colspan="8" align="center"><br /><i>E. q. hopiensis</i>, Moab, Grand Co., Utah.</td>
+</tr>
+<tr>
+ <td>Mean (11)<br />Min &#9794;<br />Max<br /><br />Mean (5)<br />Min &#9792;<br />Max</td>
+ <td align="center">34.4<br />33.5<br />35.4<br /><br />34.9<br />34.2<br />35.7</td>
+ <td align="center">19.4<br />19.2<br />20.0<br /><br />19.6<br />19.3<br />20.1</td>
+ <td align="center">16.3<br />15.9<br />16.8<br /><br />16.4<br />15.9<br />16.6</td>
+ <td align="center">10.0<br />9.3<br />10.5<br /><br />10.5<br />10.1<br />10.7</td>
+ <td align="center">212<br />208<br />220<br /><br />219<br />210<br />228</td>
+ <td align="center">90.7<br />85.0<br />96.0<br /><br />94.4<br />85.0<br />104.0</td>
+ <td align="center">5.19<br />4.92<br />5.38<br /><br />5.16<br />5.13<br />5.20</td>
+ <td align="center">18.36<br />17.80<br />18.96<br /><br />18.58<br />18.00<br />19.19</td>
+</tr>
+<tr>
+ <td>&nbsp;</td>
+ <td colspan="8" align="center"><br /><i>E. u. umbrinus</i>, Mts. S Robertson, Uintah Co., Wyoming.</td>
+</tr>
+<tr>
+ <td>Mean (11)<br />Min &#9794;<br />Max<br /><br />Mean (4)<br />Min &#9792;<br />Max</td>
+ <td align="center">34.7<br />34.3<br />35.2<br /><br />35.1<br />34.9<br />35.4</td>
+ <td align="center">18.9<br />18.3<br />19.4<br /><br />19.2<br />18.6<br />20.0</td>
+ <td align="center">15.7<br />15.6<br />16.0<br /><br />15.9<br />15.7<br />16.2</td>
+ <td align="center">10.9<br />10.3<br />11.7<br /><br />11.0<br />10.3<br />11.8</td>
+ <td align="center">218<br />215<br />228<br /><br />224<br />204<br />234</td>
+ <td align="center">96.2<br />81.0<br />112.0<br /><br />96.4<br />90.0<br />100.0</td>
+ <td align="center">5.13<br />4.79<br />5.42<br /><br />5.17<br />5.11<br />5.22</td>
+ <td align="center">18.04<br />17.57<br />18.59<br /><br />18.46<br />18.31<br />18.98</td>
+</tr>
+<tr>
+ <td>&nbsp;</td>
+ <td colspan="8" align="center"><br /><i>E. u. adsitus</i>, Britts Meadow, Beaver Co., Utah.</td>
+</tr>
+<tr>
+ <td>Mean (6)<br />Min &#9794;<br />Max<br /><br />Mean (6)<br />Min &#9792;<br />Max</td>
+ <td align="center">34.8<br />34.3<br />35.4<br /><br />35.1<br />33.9<br />36.2</td>
+ <td align="center">18.9<br />18.5<br />19.6<br /><br />19.5<br />18.9<br />20.0</td>
+ <td align="center">15.5<br />15.3<br />16.1<br /><br />16.0<br />15.9<br />16.3</td>
+ <td align="center">10.8<br />10.4<br />11.3<br /><br />11.0<br />10.6<br />11.8</td>
+ <td align="center">214<br />203<br />225<br /><br />228<br />215<br />233</td>
+ <td align="center">89.6<br />73.0<br />95.0<br /><br />96.5<br />95.0<br />98.0</td>
+ <td align="center">5.16<br />4.64<br />5.34<br /><br />5.11<br />5.00<br />5.33</td>
+ <td align="center">18.07<br />17.69<br />18.70<br /><br />18.75<br />18.51<br />19.40</td>
+</tr>
+<tr>
+ <td><span class='pagenum'><a name="Page_581" id="Page_581">[Pg 581]</a></span></td>
+ <td colspan="8" align="center"><br /><i>E. u. sedulus</i>,&nbsp; Mt. Ellen, Henry Mts., Garfield Co., Utah.</td>
+</tr>
+<tr>
+ <td>Mean (5)<br />Min &#9794;<br />Max<br /><br />Mean (2)<br />Min &#9792;<br />Max</td>
+ <td align="center">34.7<br />33.5<br />35.5<br /><br />34.9<br />34.9<br />34.9</td>
+ <td align="center">18.7<br />18.4<br />19.1<br /><br />19.4<br />19.3<br />19.5</td>
+ <td align="center">15.6<br />15.4<br />15.9<br /><br />16.1<br />16.1<br />16.1</td>
+ <td align="center">10.7<br />10.1<br />11.2<br /><br />11.1<br />11.0<br />11.3</td>
+ <td align="center">218<br />213<br />224<br /><br />227<br />224<br />231</td>
+ <td align="center">93.0<br />89.0<br />97.0<br /><br />98.0<br />96.0<br />100.0</td>
+ <td align="center">5.21<br />5.09<br />5.28<br /><br />5.24<br />5.07<br />5.42</td>
+ <td align="center">18.74<br />18.48<br />19.38<br /><br />18.74<br />19.24<br />19.80</td>
+</tr>
+<tr>
+ <td>&nbsp;</td>
+ <td colspan="8" align="center"><br /><i>E. u. inyoensis</i>, Ruby Mts., Elko and White Pine Cos., Nev.</td>
+</tr>
+<tr>
+ <td>Mean (12)<br />Min &#9794;<br />Max<br /><br />Mean (5)<br />Min &#9792;<br />Max</td>
+ <td align="center">34.4<br />33.5<br />35.4<br /><br />34.9<br />34.4<br />35.2</td>
+ <td align="center">19.0<br />18.6<br />19.6<br /><br />19.4<br />19.1<br />19.7</td>
+ <td align="center">15.7<br />15.2<br />16.1<br /><br />15.7<br />15.4<br />16.0</td>
+ <td align="center">10.5<br />10.0<br />11.5<br /><br />10.5<br />10.4<br />10.7</td>
+ <td align="center">208<br />196<br />220<br /><br />215<br />204<br />226</td>
+ <td align="center">89.5<br />85.0<br />100.0<br /><br />92.8<br />86.0<br />102.0</td>
+ <td align="center">5.15<br />5.01<br />5.37<br /><br />5.19<br />5.04<br />5.33</td>
+ <td align="center">18.12<br />17.32<br />18.81<br /><br />18.63<br />18.50<br />18.80</td>
+</tr>
+<tr>
+ <td>&nbsp;</td>
+ <td colspan="8" align="center"><br /><i>E. u. nevadensis</i>, Measurements of the type (Burt 1931:300).</td>
+</tr>
+<tr>
+ <td>&#9794;</td>
+ <td align="center">34.8</td>
+ <td align="center">19.3</td>
+ <td align="center">16.2</td>
+ <td align="center">11.0</td>
+ <td align="center">205</td>
+ <td align="center">89.0</td>
+ <td align="center">...</td>
+ <td align="center">...</td>
+</tr>
+<tr>
+ <td>&nbsp;</td>
+ <td colspan="8" align="center"><br /><i>E. u. fremonti</i>, Togwotee Pass, Fremont Co., Wyoming.</td>
+</tr>
+<tr>
+ <td>Mean (8)<br />Min &#9794;<br />Max<br /><br />Mean (6)<br />Min &#9792;<br />Max</td>
+ <td align="center">35.6<br />35.2<br />36.5<br /><br />35.3<br />34.5<br />36.0</td>
+ <td align="center">19.3<br />18.9<br />19.7<br /><br />19.6<br />19.3<br />20.0</td>
+ <td align="center">15.9<br />15.8<br />16.1<br /><br />15.9<br />15.7<br />16.5</td>
+ <td align="center">11.4<br />11.1<br />11.8<br /><br />11.3<br />10.9<br />12.0</td>
+ <td align="center">223<br />216<br />243<br /><br />229<br />223<br />239</td>
+ <td align="center">99.0<br />95.0<br />111.0<br /><br />101.0<br />92.0<br />110.0</td>
+ <td align="center">5.34<br />5.22<br />5.57<br /><br />5.40<br />5.35<br />5.44</td>
+ <td align="center">19.17<br />18.72<br />19.78<br /><br />19.02<br />18.37<br />19.51</td>
+</tr>
+<tr>
+ <td>&nbsp;</td>
+ <td colspan="8" align="center"><br /><i>E. u. montanus</i>, Boulder Co., Colorado.</td>
+</tr>
+<tr>
+ <td>Mean (5)<br />Min &#9794;<br />Max<br /><br />Mean (6)<br />Min &#9792;<br />Max</td>
+ <td align="center">35.2<br />34.7<br />36.8<br /><br />35.7<br />35.1<br />36.5</td>
+ <td align="center">18.8<br />18.4<br />19.4<br /><br />19.1<br />18.8<br />19.5</td>
+ <td align="center">15.5<br />15.2<br />16.2<br /><br />15.6<br />15.1<br />16.0</td>
+ <td align="center">10.8<br />10.1<br />11.5<br /><br />10.9<br />10.3<br />11.6</td>
+ <td align="center">226<br />215<br />232<br /><br />226<br />215<br />231</td>
+ <td align="center">96.0<br />93.0<br />115.0<br /><br />98.0<br />89.0<br />105.0</td>
+ <td align="center">5.20<br />5.03<br />5.53<br /><br />5.28<br />5.06<br />5.58</td>
+ <td align="center">18.29<br />17.80<br />19.36<br /><br />18.67<br />18.09<br />19.35</td>
+</tr>
+</table>
+
+<p><span class='pagenum'><a name="Page_582" id="Page_582">[Pg 582]</a></span></p>
+
+<h3><span class="smcap">Literature Cited</span></h3>
+
+<p><span class="smcap">Benson, S. B.</span></p>
+<div class="litblockquot"><p>1935.&nbsp; A biological reconnaissance of Navajo Mountain, Utah. Univ.
+California Publ. Zool., 40:439-455, December 31.</p></div>
+
+<p><span class="smcap">Burt, W. H.</span></p>
+<div class="litblockquot"><p>1931.&nbsp; Three new subspecies of chipmunks of the genus Eutamias from
+Nevada. Jour. Mamm., 12:298-301, August 24.</p></div>
+
+<p><span class="smcap">Durrant, S. D.</span></p>
+<div class="litblockquot"><p>1952.&nbsp; Mammals of Utah, taxonomy and distribution. Univ. Kansas
+Publ. Mus. Nat. Hist., 6:1-549, 91 figs., 30 tables, August 10.</p></div>
+
+<p><span class="smcap">Hall, E. R.</span></p>
+<div class="litblockquot"><p>1946.&nbsp; Mammals of Nevada. Univ. California Press, Berkeley,
+California, pp. xi + 710, 11 pls., 485 figs., July 1.</p></div>
+
+<p><span class="smcap">Hardy, R.</span></p>
+<div class="litblockquot"><p>1945.&nbsp; The taxonomic status of some chipmunks of the genus Eutamias
+in southwestern Utah. Proc. Biol. Soc. Washington, 58:85-87, June 30.</p></div>
+
+<p><span class="smcap">Howell, A. H.</span></p>
+<div class="litblockquot"><p>1929.&nbsp; Revision of the American chipmunks (genera <i>Tamias</i> and <i>Eutamias</i>).
+U. S. Dept. Agric., Bur. Biol. Surv., N. Amer. Fauna, 52:1-157, 10 pls., 9 figs., November 30.</p></div>
+
+<p><span class="smcap">Hunt, C. B.</span>, and <span class="smcap">Sokoloff, V. P.</span></p>
+<div class="litblockquot"><p>1950.&nbsp; Pre-Wisconsin soil in the Rocky Mountain region, a progress
+report. U. S. Geol. Survey, Prof. Paper, 221-G:109-123.</p></div>
+
+<p><span class="smcap">Johnson, D. H.</span></p>
+<div class="litblockquot"><p>1943.&nbsp; Systematic review of the chipmunks (genus Eutamias) of California.
+Univ. California Publ. Zool., 48:63-148, 6 pls., December 24.</p></div>
+
+<p><span class="smcap">Kelson, K. R.</span></p>
+<div class="litblockquot"><p>1951.&nbsp; Speciation in rodents of the Colorado River drainage. Univ. Utah
+Biol. Ser., 11(3): vii + 125, 10 figs., February 15.</p></div>
+
+<p><span class="smcap">Merriam, C. H.</span></p>
+<div class="litblockquot"><p>1905.&nbsp; Two new chipmunks from Colorado and Arizona. Proc. Biol. Soc.
+Washington, 18:163-166, June 29.</p></div>
+
+<p><span style="margin-left: 2em;"><i>Transmitted June 26, 1953.</i></span></p>
+
+<h5>24-8966</h5>
+
+
+
+
+
+
+
+
+<pre>
+
+
+
+
+
+End of the Project Gutenberg EBook of Taxonomy of the Chipmunks, Eutamias
+quadrivittatus and Eutamias umbrinus, by John A. White
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+The Project Gutenberg EBook of Taxonomy of the Chipmunks, Eutamias
+quadrivittatus and Eutamias umbrinus, by John A. White
+
+This eBook is for the use of anyone anywhere at no cost and with
+almost no restrictions whatsoever. You may copy it, give it away or
+re-use it under the terms of the Project Gutenberg License included
+with this eBook or online at www.gutenberg.org
+
+
+Title: Taxonomy of the Chipmunks, Eutamias quadrivittatus and Eutamias umbrinus
+
+Author: John A. White
+
+Release Date: February 12, 2010 [EBook #31267]
+
+Language: English
+
+Character set encoding: ASCII
+
+*** START OF THIS PROJECT GUTENBERG EBOOK TAXONOMY--CHIPMUNKS EUTAMIAS ***
+
+
+
+
+Produced by Chris Curnow, Joseph Cooper and the Online
+Distributed Proofreading Team at https://www.pgdp.net
+
+
+
+
+
+
+
+
+
+ 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. x 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 pinon 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 pinon 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.
+ 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
+
+
+
+
+
+End of the Project Gutenberg EBook of Taxonomy of the Chipmunks, Eutamias
+quadrivittatus and Eutamias umbrinus, by John A. White
+
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