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+The Project Gutenberg EBook of Subspeciation in the Meadow Mouse, Microtus
+montanus, in Wyoming and Colorado, by Sydney Anderson
+
+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: Subspeciation in the Meadow Mouse, Microtus montanus, in Wyoming and Colorado
+
+Author: Sydney Anderson
+
+Release Date: March 22, 2010 [EBook #31730]
+
+Language: English
+
+Character set encoding: UTF-8
+
+*** START OF THIS PROJECT GUTENBERG EBOOK MEADOW MOUSE ***
+
+
+
+
+Produced by Chris Curnow, Simon Gardner, Joseph Cooper and
+the Online Distributed Proofreading Team at
+http://www.pgdp.net
+
+
+
+
+
+
+
+UNIVERSITY OF KANSAS PUBLICATIONS
+
+MUSEUM OF NATURAL HISTORY
+
+Volume 7, No. 7, pp. 489-506, 2 figures in text
+
+July 23, 1954
+
+
+
+
+                    Subspeciation in the Meadow Mouse,
+                      Microtus montanus, in Wyoming
+                              and Colorado
+
+                                  BY
+
+                            SYDNEY ANDERSON
+
+
+UNIVERSITY OF KANSAS
+
+LAWRENCE
+
+1954
+
+
+UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY
+
+Editors: E. Raymond Hall, Chairman, A. Byron Leonard,
+Robert W. Wilson
+
+Volume 7, No. 7, pp. 489-506, 2 figures in text
+
+Published July 23, 1954
+
+
+UNIVERSITY OF KANSAS
+
+Lawrence, Kansas
+
+
+PRINTED BY
+
+FERD VOILAND, JR., STATE PRINTER
+
+TOPEKA, KANSAS
+
+1954
+
+25-3560
+
+
+
+
+Subspeciation in the Meadow Mouse, Microtus montanus, in Wyoming and
+Colorado
+
+BY
+
+SYDNEY ANDERSON
+
+
+_Microtus montanus_ reaches the eastern limits of its geographic
+distribution in Wyoming and Colorado. There the mountains, but in
+general not the lowlands, are occupied by this species. A certain
+minimum of moisture may be of direct importance to the mouse and
+certainly is indirectly important, because certain hydrophytic or
+mesophytic grasses used by the mouse for food, for protection from
+enemies, and for shelter from the elements are dependent on the
+moisture. Areas suitable for _Microtus montanus_ are separated by
+deserts that are dominated by sagebrush and other xerophytic plants or
+by forests or rocky exposures at higher altitudes. A relatively small
+percentage, probably less than ten per cent, of the total area even in
+the more favorable parts of the range of the species is suitable for
+occupancy. In these mice, as in other microtines (Elton, 1942; Piper,
+1909), there are seasonal, and irregularly multiannual fluctuations in
+population density, which sometimes are extreme. Consequently the mice
+at some times seem to be absent from suitable habitats, and at some
+other times occur there in amazingly large numbers.
+
+Because the species is broken up into partly isolated, or at times
+completely isolated, colonies or local populations it may be supposed
+that various evolutionary forces such as selection and random genetic
+drift operate to foster variation. The degree to which racial
+distinction is attained may depend upon these forces and the time
+available. In _Microtus montanus_ in the eastern Rocky Mountains the
+degree of subspecific distinction is not great.
+
+The study here reported upon is based on 1,187 specimens of _Microtus
+montanus_ from Wyoming, Colorado, Idaho and Montana, and on work in the
+field. I spent approximately four months in the field in this area, in
+the summers of 1950, 1951, and 1952. The specimens studied were arranged
+according to localities and the larger series were compared
+statistically. Each of two series, totaling 136 specimens, was studied
+intensively to ascertain the kind and range of variation within single
+populations. Twenty-seven measurements, various proportions based on
+these measurements, and differences in color were analyzed. Fifteen
+characters, judged to be most significant, were selected for use in
+comparing all series. In addition, certain characters that can not be
+expressed easily by measurements, such as inflation of the auditory
+bullae and the curvature of the zygomatic arch, were observed. The
+studies by A. B. Howell (1924) of variation in _Microtus montanus
+yosemite_ Grinnell in California and those by O. B. Goin (1943) of
+_Microtus pennsylvanicus pennsylvanicus_ (Ord) were useful. The external
+measurements are from the collectors' field labels. The measurements of
+the skull all were taken with dial calipers reading to a tenth of a
+millimeter. The anteroposterior measurements of the skull all were taken
+along the shortest line between the points specified below and are not
+necessarily along a line parallel to the long axis of the skull. These
+measurements were taken on the left side of the skull whenever possible.
+Some of the skulls are damaged and therefore some measurements could not
+be taken and are omitted in the computations. Measurements are in
+millimeters.
+
+The results of these studies were submitted to the Department of Zoology
+and the Graduate School of the University of Kansas in partial
+fulfillment of the requirements for the degree of Master of Arts (1952)
+and are available in manuscript form at the Museum of Natural History
+and the library of the University of Kansas.
+
+
+EXPLANATION OF MEASUREMENTS
+
+     Caudal index.--the length of the tail expressed as a percentage of
+     the length of the head and body. The length of the head and body is
+     the collector's measurement of total length less the length of the
+     tail.
+
+CRANIAL MEASUREMENTS OF LENGTH.
+
+     Condylobasilar length.--from the exoccipital condyle to the most
+     posterior point on the border of the alveolus of the upper incisor.
+
+     Alveolobasilar length.--from the posterior border of the alveolus
+     of the third upper molar to the posterior border of the alveolus of
+     the incisor.
+
+     Palatilar length.--from the anteriormost part of the posterior
+     border of the bony shelf of the palate to the posteriormost part of
+     the alveolus of the incisor.
+
+     Alveolar length of upper molar tooth-row.--from the most posterior
+     point of the alveolus of the third upper molar to the most anterior
+     point of the alveolus of the first upper molar.
+
+MEASUREMENTS OF BREADTH.
+
+     Zygomatic breadth.--greatest transverse width.
+
+     Interorbital breadth.--the breadth of the interorbital
+     constriction.
+
+     Lambdoidal breadth.--between the lateralmost points of the
+     lambdoidal crest.
+
+     Prelambdoidal breadth.--between the medialmost margins of the
+     prominent fenestrae in the posterodorsal parts of the squamosal
+     bones. To these fenestrae Howell (1924:995) applied the adjective
+     "prelambdoidal," but other authors have used other names (see Hill,
+     1935:127).
+
+     Depth of braincase.--shortest distance from the ventral side of the
+     cranium at the suture between the basioccipital and basisphenoid
+     bones to the dorsal surface of the cranium (usually not
+     perpendicular to the long axis of the skull).
+
+The history of our knowledge of _Microtus montanus_ in this area begins
+with the early work of the United States Bureau of Biological Survey
+directed by C. H. Merriam (1891), and participated in by Vernon Bailey
+(1900, 1917), Merritt Cary (1911, 1917), and others. The changes in
+nomenclature which grew out of increased understanding of these mice
+through additional collecting are expressed in the synonymies under the
+accounts of subspecies. As a result of my studies two of the three
+subspecific names previously proposed for mice from this area have been
+retained although changes are proposed in the ranges assigned to the two
+subspecies and two additional heretofore unrecognized subspecies are
+named and described. Furthermore the additional specimens and my studies
+of variation make modifications in the characterization of these
+subspecies necessary. Not all of the samples here assigned to a single
+subspecies are identical and I therefore list and discuss some of the
+local variants.
+
+     Numerous members of summer field parties from the Museum of Natural
+     History at the University of Kansas collected most of the specimens
+     studied and wrote field notes that have been helpful. I am grateful
+     to these persons and to Professor E. R. Hall and Dr. R. H. Baker
+     for their assistance and helpful suggestions. Specimens in the
+     following museums were made available by their respective curators:
+     Chicago Natural History Museum by Mr. Colin C. Sandborn, The Museum
+     of Zoology at the University of Michigan by Dr. E. T. Hooper, The
+     American Museum of Natural History by Mr. G. G. Goodwin, The United
+     States National Museum by Dr. David H. Johnson and the Biological
+     Surveys Collection by Miss Viola S. Schantz. A fellowship from the
+     National Science Foundation made possible the studies at the
+     museums other than at the University of Kansas.
+
+
+
+=Microtus montanus nanus= (Merriam)
+
+     _Arvicola (Mynomes) nanus_ Merriam, N. Amer. Fauna, 5:62, July 30,
+     1891.
+
+     _Microtus montanus nanus_, Hall, Proc. Biol. Soc. Washington,
+     51:131, August 23, 1938.
+
+     _Microtus nanus_, Bailey, N. Amer. Fauna, 17:30, June 6, 1900
+     (part).
+
+     _Microtus montanus caryi_ Bailey, Proc. Biol. Soc. Washington,
+     30:29, February 21, 1917.
+
+     _Microtus nanus nanus_, Cary, N. Amer. Fauna, 42:43, October 3,
+     1917 (part).
+
+     _Type._--Adult male No. 23853/31253, U. S. National Museum,
+     Department of Agriculture collection, from Pahsimeroi Mountains,
+     Custer County, Idaho; obtained by C. Hart Merriam and Vernon
+     Bailey, September 16, 1890.
+
+     _Range._--Idaho; southwestern Montana; most of the southwestern
+     half of Wyoming; southward to central Colorado. See figure 1.
+
+     _Comparisons._--Comparisons with subspecies named as new in this
+     paper will be found in the accounts of those subspecies beyond.
+     From _Microtus montanus fusus_ Hall, the subspecies to the south,
+     _M. m. nanus_ from Idaho differs as follows: averages smaller;
+     slightly darker and less reddish and less yellowish in color;
+     slightly wider braincase (see measurement of prelambdoidal
+     breadth); larger bullae.
+
+     _Measurements._--Average (= arithmetical mean) measurements of 34
+     specimens, both male and female, from several localities in eastern
+     Idaho are: total length, 151; length of tail, 39; hind foot, 19.2;
+     condylobasilar length of the skull, 25.0; zygomatic breadth, 15.0;
+     alveolar length of upper molar tooth-row, 6.4; prelambdoidal
+     breadth, 8.9; and lambdoidal breadth, 11.7.
+
+     Average and extreme measurements of six adult males from near
+     Pocatello, Bannock County, Idaho, and nine adult males from near
+     Afton, Lincoln County, Wyoming, are, respectively, as follows:
+     total length, 143(135-150), 163(143-179); length of tail,
+     35.1(33-38), 42.8(36-49); caudal index, 32.0(28.0-33.1),
+     35.7(30.6-41.9); hind foot, 18.9(18-20), 18.8(17-20);
+     condylobasilar length of skull, 24.4(24.0-26.0), 25.6(24.5-26.2);
+     alveolobasilar length, 14.1(13.7-14.5), 14.6(13.8-15.0); palatilar
+     length, 13.2(12.9-13.6), 13.8(13.2-14.5); alveolar length of upper
+     molar tooth-row, 6.3(6.1-6.5), 6.3(6.0-6.6); depth of braincase,
+     7.7(7.5-7.9), 8.0(7.7-8.3); lambdoidal breadth, 11.4(11.0-11.7),
+     12.0(11.3-12.7); prelambdoidal breadth, 9.1(8.6-9.4), 8.7(8.0-9.4);
+     zygomatic breadth, 14.3(13.8-14.7), 15.3(14.4-16.3); interorbital
+     breadth, 3.6(3.5-3.7), 3.5(3.3-3.7). The average length of the
+     nasal bones in the series from Pocatello is 7.1 mm. The averages,
+     which have not been included in Table 1, for three measurements of
+     the series from Carbon County, Wyoming, are as follows (Encampment,
+     males; Encampment, females; Savery, males; and Savery, females,
+     respectively): alveolobasilar length, 14.4, 14.3, 14.5, 14.3;
+     interorbital breadth, 3.5, 3.4, 3.5, 3.4; depth of braincase, 7.8,
+     7.6, 7.9, 7.6. Additional measurements are included in Table 1 for
+     other series.
+
+_Discussion._--The name _Microtus montanus caryi_ Bailey is here placed
+in synonymy under _M. m. nanus_ (Merriam). Vernon Bailey (1917) in his
+description of _caryi_ made four assumptions that have been found to be
+entirely or partly invalid. First, he assumed that this is an "extreme
+variant which gradually changes in characters across Nevada and Utah,
+and reaches its maximum variation in Wyoming." The differences pointed
+out in subsequent descriptions of subspecies found in the above area do
+not show a gradual change in any character, or in the number of
+characters, nor is _caryi_ an extreme when compared with the other
+subspecies. Second, _Microtus nanus_ was not, as Bailey assumed, a
+different species than _Microtus montanus_. Third, he assumed that the
+characteristics of adults of _nanus_ were adequately ascertainable from
+the thirteen topotypes available to him. Subsequent sampling from Idaho
+shows that the series of specimens available to Bailey was made up
+mostly of young and subadult animals. Finally, _caryi_ does not occupy
+as Bailey stated "the meadows along streams in the arid sagebrush
+country of the Bear River, Green River, and Wind River valleys"
+exclusively, or characteristically. When the localities from which the
+species actually is known are plotted, it seems that the arid basin
+serves as a barrier and that the species is more commonly and abundantly
+found in montane meadows in the Transition and Canadian life-zones.
+
+Certain samples, here assigned to _M. m. nanus_, that vary from the
+average of the subspecies deserve comment. For example, mice from the
+area in Wyoming southwest of the Green River (in the Uinta Mountains)
+have relatively smaller feet, but are larger in both total length and
+size of skull. Specimens from near Afton, Lincoln County, Wyoming, are
+relatively large in both total length and size of skull. This series and
+specimens from Teton County, Wyoming, are intermediate between _nanus_
+from Idaho and the newly named subspecies from near Cody, Park County,
+Wyoming, described below, in terms of both darkness and the amount of
+reddish color. Mice from Laramie County are more richly reddish-brown.
+The specimens from near Savery, in Carbon County, Wyoming, are darker.
+The alveolobasilar length relative to the condylobasilar length is
+smaller in the series from along Deer Creek, 16 mi. S, 11 mi. W Waltman,
+Natrona County, Wyoming. The series from the southern tier of counties
+in Wyoming and some of the specimens from Colorado have relatively wider
+zygomatic arches. The specimens from southern Sweetwater County,
+Wyoming, are relatively paler, have a relatively longer tail and longer
+hindfeet, lesser condylobasilar length, and wider braincase. Most of
+these variations are of questionable significance; they may be chance
+variations owing to errors in sampling.
+
+Much of the south-central part of the state is relatively low and
+relatively arid. This area includes the arid basin of the Green River
+and its major tributaries and the arid Red Desert along the continental
+divide in Sweetwater County. This area might have acted as a barrier to
+the mice; gene flow might have been prevented between the populations of
+the western part of the state and those farther east in the Medicine Bow
+Mountains and Laramie Mountains. Nevertheless geographic variations of
+subspecific worth have not taken place. The barrier has either not been
+of as long duration, or has not been so complete and effective, as the
+other barriers in the state, namely the Absaroka Range, the Big Horn
+Basin, the Shoshone Basin, and the valley of the North Platte River.
+These four barriers presumably have led to the differentiation of the
+two subspecies that are newly named beyond. Each of the two areas which
+is set apart by these barriers and in which one of the newly named
+subspecies has evolved is small; therefore there is a lesser amount of
+suitable habitat available for each of the newly named mice than there
+is for _M. m. nanus_. It is conceivable, therefore, that in periods of
+adverse conditions in each of the small areas the size of the effective
+breeding population may have been so small that random genetic drift
+could have operated effectively, or that selection was more critical
+than in a larger, more stable population. It is difficult to test these
+possibilities because the selective value of the taxonomic characters is
+unknown. The observed pattern of variation and facts of distribution
+are, however, not contradictory to the above possibilities.
+
+     _Specimens examined._--Total, 993, distributed as follows: All
+     specimens unless otherwise indicated are in the University of
+     Kansas Museum of Natural History. Specimens in other museums are
+     labeled as follows: Chicago Natural History Museum (Chi);
+     University of Michigan, Museum of Zoology (Mich); American Museum
+     of Natural History (AMNH); United States National Museum (USNM);
+     Biological Surveys Collection (USBS). Localities that are not
+     represented in Fig. 1 because overlapping or crowding of the
+     symbols would result are Italicized. Localities are arranged from
+     north to south by states, within a state from northwest to
+     southeast by counties, and within a county from north to south.
+
+     WYOMING: _Yellowstone Park_: Canyon Camp, 1 (USBS); Lower Geyser
+     Basin, 1 (USBS); Upper Yellowstone River, 2 (AMNH); _North end of
+     Lake_, _Yellowstone National Park_, 2 (AMNH). _Teton Co._: Pacific
+     Creek, 1 (USBS); _Big Game Ridge_, 3 (USBS 1, Mich 2); _Whetstone
+     Creek_, 7 (Mich); Moran and environs (6 localities within a 5 mile
+     radius), 28 (USBS 2, Mich 5); _S fork Buffalo River_, 7 (AMNH); 2
+     mi. W pass, Black Rock Creek, 1 (USBS); _Jenny Lake_, 5 (Mich); Bar
+     BC Ranch, 2½ mi. NE Moose, 6500 ft., 2; Teton Pass above Fish
+     Creek, 1 (USBS); Jackson and environs, 142 (Mich 141); _Sheep
+     Creek_, 2 (Mich). _Lincoln Co.: 13 mi. N, 2 mi. W Afton_, 2; _10
+     mi. N, 2 mi. W Afton_, 4; _9½ mi. N, 2 mi. W Afton_, 3; 9 mi. N,
+     2 mi. W Afton, 9; _7 mi. N, 1 mi. W Afton_, 12; Afton, 1 (USBS);
+     Labarge Creek, 1 (USBS); Border, 6 (USBS); _Cokeville_, 2 (USBS); 6
+     mi. N, 2 mi. E Sage, 1; Cumberland, 5 (USBS). _Sublette Co._: 34
+     mi. N, 4 mi. W Pinedale, 1; _33 mi. N, 2 mi. W Pinedale_, 6; _32
+     mi. N, 1 mi. W Pinedale_, 1; _31 mi. N Pinedale_, 4; Dell Creek, on
+     Ferris Ranch, 7 (Mich); Horse Creek, 7800 ft., Merna, 4 (USBS); Big
+     Piney, 1 (USBS). _Fremont Co._: 17½ mi. W, 2½ mi. N Lander,
+     9500 ft., 3; _17 mi. W, 2 mi. N Lander, 9300 ft._, 4; Milford and
+     environs (5 localities within a 1 mile radius), 23 (USBS 4); 15½
+     mi. S, 7½ mi. W Lander, 9200 ft., 1; South Pass City, 8000 ft, 8
+     (USBS); _23½ mi. S, 5 mi. W Lander, 8600 ft._, 7. _Natrona Co._:
+     Deer Creek, 16 mi. S, 11 mi. W Waltman, 6950 ft., 44; 6 mi. S, 2
+     mi. W Casper, 5900 ft., 4; _6-4/5 mi. S, 2 mi. W Casper, 6100 ft._,
+     1; _7 mi. S, 2 mi. W Casper, 6370 ft._, 3; _10 mi. S Casper, 7750
+     ft._, 33; Sun, 2 (USBS); _5 mi. W Independence Rock, 6000 ft._, 4;
+     _5 mi. W, 1 mi. S Independence Rock_, 2. _Converse Co._: Beaver, 1
+     (USBS). _Uinta Co.: 1½ mi. W, ½ mi. S Cumberland_, 6; _16 mi.
+     S, 2 mi. W Kemmerer, 6700 ft._, 3; 10 mi. SW Granger, 3 (Mich);
+     Fort Bridger, 6650 ft., 25 (USNM 6); 9 mi. S Robertson, 8000 ft.,
+     9; _9½ mi. S, ½ mi. W Robertson, 8600 ft._, 1; _10 mi. S, 1
+     mi. W Robertson, 8700 ft._, 25; _14 mi. S, 2 mi. E Robertson, 9000
+     ft._, 5; 4 mi. S Lonetree, 1 (USBS). _Sweetwater Co._: Farson, 3;
+     Bitter Creek, 3 (AMNH); Kinney Ranch, 21 mi. S Bitter Creek, 6800
+     ft., 9 (USNM 1, AMNH 2); 32 mi. S, 22 mi. E Rock Springs, 7025 ft.,
+     on Vermillion Creek, 15. _Carbon Co._: 18 mi. NNE Sinclair, 6500
+     ft., 10; Bridgers Pass, 18 mi. SW Rawlins, 7500 ft., 7; Saratoga, 1
+     (USBS); _6 mi. S, 13 mi. E Saratoga, 8500 ft._, 5; _6 mi. S, 14 mi.
+     E Saratoga, 8800 ft._, 1; Lake Marie, 10,440 ft., 2; _1 mi. S Lake
+     Marie_, 2; _½ mi. S, 2 mi. E Medicine Bow Peak, 10,800 ft._, 1;
+     Encampment (12 localities from 10 mi. N, 14 mi. E to 9 mi. N, 3 mi.
+     E Encampment and from 6500 to 8400 ft.), 63; ¼ mi. N Riverside,
+     7380 ft., 2; S base Bridger Peak, 8800 ft., Sierra Madre Mountains,
+     1; _2 mi. S Bridger Peak, 9300 ft._, 2; Savery (10 localities from
+     8 mi. N, 21 mi. E to 4 mi. N, 8 mi. E Savery and from 7300 to 8800
+     ft.), 80. _Albany Co._: _30 mi. N, 10 mi. E Laramie, 6760 ft._, 6;
+     _29¾ mi. N, 9½ mi. E Laramie, 6350 ft._, 1; 26 mi. N, 4½
+     mi. E Laramie, 6960 ft., 8; _26¾ mi. N, 6½ mi. E Laramie,
+     6700 ft._, 3; _3 mi. N, 13 mi. E Laramie, 7500 ft._, 1; _7 mi. N, 2
+     mi. E Laramie_, 1 (Chi); 5 mi. N Laramie, 7400 ft., 15; _Laramie_,
+     4 (AMNH); _1 mi. E Laramie, 7160 ft._, 4; 7-7/10 mi. SSW Laramie,
+     7200 ft., 4; 6½ mi. S, 8¾ mi. E Laramie, 8200 ft., 1;
+     _Headquarters Park, 10,200 ft., Medicine Bow Mountains_, 3 (USBS);
+     Centennial, 8120 ft., 1; _2¼ mi. ESE Brown's Peak, 10,300 ft._,
+     3; _3 mi. ESE Brown's Peak, 10,000 ft._, 12; _2 mi. S Brown's Peak,
+     10,600 ft._, 1; _Pole Mountain, 15 mi. SE Laramie_, 4 (USBS 3); _1
+     mi. SSE Pole Mountain, 8350 ft._, 4; _2 mi. SW Pole Mountain, 8300
+     ft._, 13; _3 mi. S Pole Mountain, 8100 ft._, 1; Sherman, 2 (AMNH).
+     _Laramie Co._: 5 mi. N, 1 mi. W Horse Creek P. O., 7200 ft., 1;
+     Meadow, 2 (USBS); 11 mi. N, 5½ mi. E Cheyenne, 5450 ft., 7; _7
+     mi. W Cheyenne, 6500 ft._, 10; Cheyenne, 3 (USNM).
+
+     COLORADO: _Moffat Co._: Lay, 6160 ft., 1 (AMNH). _Routt Co._:
+     Wright's Ranch, Yampa, 7700 ft., 2; Gore Range, 8 mi. E Toponas,
+     8000 ft., 2 (USBS). _Larimer Co._: _12½ mi. W, 1½ mi. S
+     Rustic_, 1; 11 mi. W, 1 mi. S Rustic, 1; Cache La Poudre River, 1
+     (Chi); _Estes Park_, 3 (USBS 1, AMNH 2); 19½ mi. W, 2½ mi. S
+     Loveland, 7280 ft., 6; _16 mi. W Loveland, 6840 ft._, 1; 6 mi. W,
+     ½ mi. S Loveland, 5200 ft., 1. _Rio Blanco Co._: Meeker, 1
+     (USBS); _9½ mi. SW Pagoda Peak_, 7700 ft., 3; 5 mi. S Pagoda
+     Peak, 9100 ft., 2. _Eagle Co._: Eagle, 1 (USBS); Pando, 2 (USBS).
+     _Grand Co._: Mt. Whiteley, 2 (USBS); Arapahoe Pass, Rabbit Ear
+     Mountains, 2 (USBS); Coulter (near Granby), 5 (USBS); _Arrowhead_
+     (near Dale), 1 (USBS). _Boulder Co._: ¾ mi. N, 2 mi. W
+     Allenspark, 8400 ft., 4; _3 mi. S Ward_, 9000 ft., 3; Nederland, 16
+     (Chi). _Clear Creek Co._: Mt. McLellan, 2 (USBS); Berthoud Pass, 4.
+     _Park Co._: Trout Creek Ranch, 2 mi. N Garo, 1 (USBS).
+
+     Specimens examined of _M. m. nanus_ from eastern Idaho and Montana
+     are as follows: IDAHO: _Custer Co._: Challis, 7 (USBS); Mill Creek,
+     Challis Nat. Forest, 1 (USBS); Pahsimeroi Mts., 12 (USBS); Lost
+     River Mts., 1 (USBS). _Fremont Co._: N fork Snake River, 10 mi. SW
+     Island Park, 6200 ft., 2 (AMNH); Black Springs Creek, 4 mi. W
+     Ashton, 5200 ft., 1 (AMNH); 5 mi. W St. Anthony, 5000 ft., 1
+     (AMNH). _Camas Co._: Camas Prairie, Corral, 5100 ft., 2 (USBS).
+     _Blaine Co._: Alturas Lake, 3 (USBS); _Sawtooth Lake_, 2 (USBS);
+     Craters of the Moon, Laidlow Park, 2 (Mich); Ticura, 10 mi. S
+     Picabo, 1 (USBS); 19 mi. NE Carey (Lava Lake), 8 (Mich). _Butte
+     Co._: _26 mi. SW Arco_, 12 (Mich). _Bingham Co._: Shelley, 6
+     (USBS). _Bonneville Co._: 10 mi. SE Irwin, 4 (USBS). _Owyhee Co._:
+     Three Creeks, 3 (USBS). _Twin Falls Co._: Castleford Fenced Plot,
+     11 mi. W, 9 mi. S Twin Falls, 1. _Minidoka Co._: _Heyburn_, 2
+     (USBS). _Cassia Co._: 2 mi. S, 2 mi. W Burley, 5. _Bannock Co._:
+     Pocatello, 23 (USBS 4); Swan Lake, 1 (USBS). _Bear Lake Co._:
+     Montpelier Creek, 6700 ft., 3 (USBS). MONTANA: _Gallatin Co._: W.
+     Fork of W. Fork, Gallatin River, 1 (USBS). _Park Co._: Lamar River,
+     7000 ft., 1 (USBS); Gardiner, 1 (USBS). _Sweet Grass Co._: 14 mi. S
+     Big Timber, 1 (USBS); _McLeod_, 1 (USBS); West Boulder Creek, 18
+     mi. SE Livingston, 2 (USBS).
+
+
+=Microtus montanus codiensis=, new subspecies
+
+     _Type._--Female, adult, skin and skull; No. 27578, Museum of
+     Natural History, University of Kansas, from 3â…• mi. E and â…— mi.
+     S Cody, 5020 ft., Park Co., Wyoming; obtained on August 11, 1948,
+     by James W. Bee, original number 18-8-11-48.
+
+     _Range._--In northwestern Wyoming eastward from the Absaroka and
+     Wind River ranges into the western part of the Big Horn Basin.
+
+     _Diagnosis._--A relatively large _Microtus montanus_; tail actually
+     and relatively long; hind foot actually but not relatively large;
+     skull large; zygomatic expanse actually and relatively large;
+     alveolobasilar length relatively large; upper molar tooth-row
+     relatively long; color relatively light, not reddish.
+
+[Illustration: FIG. 1. Geographic range of _Microtus montanus_ in
+Wyoming, Colorado, and adjacent areas. The solid circles represent
+localities from which specimens have been examined; the hollow circles
+represent type localities. The ranges of subspecies in Utah are after
+Durrant, 1952.
+
+Guide to subspecies
+
+1. _M. m. nanus_
+2. _M. m. codiensis_
+3. _M. m. zygomaticus_
+4. _M. m. fusus_
+5. _M. m. micropus_
+6. _M. m. nexus_
+7. _M. m. amosus_
+8. _M. m. rivularis_
+]
+
+     _Comparisons._--As compared with the specimens of _M. m. nanus_
+     from Idaho, the size is larger (see diagnosis and measurements).
+     Certain proportions which differ from those of _nanus_ and which
+     are not in close agreement with the observed differences with age
+     in specimens of _nanus_ of a size comparable to _codiensis_ are
+     relatively large alveolobasilar length, relatively long alveolar
+     length of upper molar tooth-row, relatively wide-spreading
+     zygomatic arches, and relatively long tail. The color in
+     _codiensis_ is lighter than in _nanus_. As compared to the new
+     subspecies named below from the Big Horn Mountains to the east,
+     _codiensis_ is of similar size in head-body length, but has a
+     relatively as well as actually longer tail; the hind foot averages
+     longer; the upper molar tooth-row is relatively longer; the color
+     is slightly paler and less grizzled; the bullae are larger and less
+     flattened; the angle formed at the suture between the basioccipital
+     and basisphenoid bones is less acute; and the region of the suture
+     is less prominently elevated between the bullae when viewed from
+     the ventral aspect. The pterygoid plates mesial and posterodorsal
+     to the posterior end of the last upper molar are less fenestrated,
+     and the incisive foramina are less constricted posteriorly.
+
+[Illustration: FIG. 2. Map showing the major barriers to _Microtus
+montanus_ in Wyoming and Colorado; the barriers are the low areas named
+on the map (the name "Black Hills" is on the map for another reason;
+these hills are not a barrier). The major mountainous areas higher than
+approximately 8000 feet in elevation in Wyoming, Colorado and Utah are
+stippled. These mountainous areas include the habitat that is most
+suitable for the montane meadow mouse. The Black Hills are unoccupied by
+this species but these hills seem to be ecologically suitable for the
+species.]
+
+     _Measurements._--The average and the extremes for some measurements
+     of 34 males and females, 27 from the type locality and 7 from other
+     localities in the range assigned to this subspecies, are as
+     follows: total length, 165 (146-186); length of tail, 44.2 (35-55);
+     hind foot, 19.6 (17-21); condylobasilar length of the skull, 25.5
+     (24.0-27.5); zygomatic breadth, 15.6 (14.7-16.6); alveolar length
+     of upper molar tooth-row, 6.6 (6.2-7.0); prelambdoidal breadth, 8.8
+     (8.1-9.5); lambdoidal breadth, 12.0 (11.2-12.8). As an indication
+     of variability and for comparison with other series the coefficient
+     of variability and two times the standard error of the mean for
+     each measurement in this series are included in Table 1. The
+     averages for some measurements of 27 topotypes are as follows:
+     total length, 162; length of tail, 45.5; hind foot, 19.9;
+     condylobasilar length, 25.6; palatilar length, 14.0; molar series,
+     6.6; alveolobasilar length, 14.9; zygomatic breadth, 15.6;
+     interorbital breadth, 3.5; lambdoidal breadth, 12.1; prelambdoidal
+     breadth, 8.9; depth of braincase, 7.8.
+
+_Discussion._--Three species of _Microtus_ were collected by James W.
+Bee at the type locality. _Microtus montanus codiensis_, _Microtus
+longicaudus mordax_, and _Microtus pennsylvanicus modestus_ were taken
+in the same runways in the same meadow, at the same time. _Microtus
+ochrogaster haydeni_, although not taken at this locality, occurs in the
+Big Horn Basin. These four species differ in their geographic ranges,
+being largely allopatric, except _M. montanus_ and _M. longicaudus_
+which are sympatric. Although the different species have ecological
+preferences and habits which differ, several species of _Microtus_ may
+occur together in local areas such as the above. Certain of the
+characteristics of _M. m. codiensis_ are intermediate between those of
+the species _M. montanus_ on one hand and those of the other three
+species on the other hand. Could interspecific hybridization between
+"good species" of _Microtus_ take place in nature and possibly alter the
+characteristics of a local population?
+
+     _Specimens examined._--Total, 50, distributed as follows
+     (abbreviations for collections are given in the account of _M. m.
+     nanus_; localities that are not represented in Fig. 1 because
+     overlapping or crowding of the symbols would result are
+     Italicized):
+
+     MONTANA: _Carbon Co._: Beartooth Mountains, 2 (USBS); _Beartooth
+     Lake_, 1 (USBS).
+
+     WYOMING: _Park Co._: Black Mountain, head of Pat O'Hara Creek, 3
+     (USBS); 13 mi. N, 1 mi. E Cody, 5200 ft., 1; SW slope Whirlwind
+     Peak, 9000 ft., 1; _5 mi. N Cody, 6300 ft._, 1 (USBS); 3â…• mi. E,
+     â…— mi. S Cody, 31; Ishawooa Creek, 6300 ft., 2 (USBS); _Valley_, 1
+     (USBS); Needle Mountain, 10,500 ft., 4 (USBS). _Hot Springs Co._: 3
+     mi. N, 10 mi. W Thermopolis, 4950 ft., 3.
+
+
+=Microtus montanus zygomaticus=, new subspecies
+
+     _Type._--Male, adult, skin and skull, No. 32761, Museum of Natural
+     History, University of Kansas, from Medicine Wheel Ranch, 9000 ft.,
+     28 mi. E Lovell, Big Horn County, Wyoming; obtained by R. Freiburg,
+     original number 105.
+
+     _Range._--The Big Horn Mountains of north-central Wyoming.
+
+     _Diagnosis._--A large _Microtus montanus_ with a relatively short
+     tail; short molar series; broad zygomatic arches well rounded in
+     lateral outline when viewed from above; small and flattened bullae;
+     raised basioccipito-basisphenoid suture.
+
+     _Comparisons._--For comparison with _M. m. codiensis_ from the
+     west, on the other side of the Big Horn Basin, see the account of
+     that subspecies. In comparison with _nanus_ this subspecies is
+     slightly paler, in this respect showing more resemblance to
+     _codiensis_ although not so pale, and more grizzled or unevenly
+     colored. This difference in color between _zygomaticus_ and
+     _codiensis_ may not be of taxonomic significance. From both the
+     topotypes of _nanus_, and the series of it from Wyoming,
+     _zygomaticus_ differs on the average in having a relatively shorter
+     tail, a relatively shorter upper molar tooth-row, relatively more
+     rounded and relatively more wide-spread zygomatic arches, and
+     smaller more flattened bullae.
+
+     _Measurements._--Average and extreme measurements of 24 adult males
+     and females from several localities here referred to _M. m.
+     zygomaticus_ are as follows: total length, 159(150-175); length of
+     tail, 37.6(31-46); hind foot, 18.6(17-20); condylobasilar length of
+     the skull, 25.3(24.2-26.7); zygomatic breadth, 15.3(14.1-16.7);
+     alveolar length of upper molar tooth-row, 6.2 (5.7-6.8);
+     prelambdoidal breadth, 8.7(8.3-9.4); lambdoidal breadth,
+     11.9(11.0-12.5). Average and extreme measurements of a series of 12
+     adult male topotypes are as follows: total length, 159(144-174);
+     length of tail, 36.4 (30-41); hind foot, 18.2(16-20);
+     condylobasilar length of skull, 25.8(24.7-26.7); alveolobasilar
+     length, 14.8(13.8-15.3); palatilar length, 13.8 (12.7-14.2);
+     alveolar length of upper molar tooth-row, 6.4(5.9-6.6); zygomatic
+     breadth, 15.9 (15.0-16.7); interorbital breadth, 3.6(3.4-3.7);
+     lambdoidal breadth, 12.1 (11.5-12.5); prelambdoidal breadth,
+     8.6(8.3-8.9); depth of braincase, 8.0 (7.6-8.3).
+
+_Discussion._--This subspecies is separated from _M. m. codiensis_ to
+the west by the Big Horn Basin. A series from along Buffalo Creek, 27
+mi. N, 1 mi. E Powder River, 6075 ft., in Natrona County, Wyoming, is
+intermediate between the topotypes of _zygomaticus_ and _nanus_ in the
+characters cited above as distinguishing the two, but shows greater
+resemblance to _zygomaticus_ in the shape of the zygomatic arch, in
+color which is paler than in topotypes of _zygomaticus_, and in the
+short hind foot. On these and on geographic grounds this population is
+referred to _zygomaticus_. Unfortunately we cannot be certain in many
+cases that an intermediate condition in a certain character indicates a
+genetically intermediate population and therefore true intergradation
+between the two subspecies to which the population is geographically
+intermediate. The topotypes of this subspecies are the most distinct of
+all the series which I have studied from the eastern Rocky Mountains, in
+terms of the degree of morphological departure from the norm for the
+species. After _zygomaticus_ the following populations are arranged
+according to their degree of deviation from this norm (_codiensis_
+deviates most): topotypes of _codiensis_, _fusus_ and a population from
+southern Sweetwater County, Wyoming, and lastly the _nanus-caryi_
+complex. Within the latter group, as I have mentioned, there are a
+number of local variants most of which do not differ significantly and
+do not conform to any geographic pattern.
+
+     _Specimens examined._--Total, 55, distributed as follows
+     (abbreviations for collections are given in the account of _M. m.
+     nanus_; localities that are not represented in Fig. 1 because
+     overlapping or crowding of the symbols would result are
+     Italicized): WYOMING: _Big Horn Co._: Medicine Wheel Ranch, 9000
+     ft., 28 mi. E Lovell, 30; W slope, head of Trappers Creek, 9500
+     ft., 2 (USBS). _Washakie Co._: 9 mi. E, 5 mi. N Tensleep, 7400 ft.,
+     1. _Johnson Co._: 7½ mi. W, 1 mi. S Buffalo, 6500 ft., 3; Big
+     Horn Mountains, 3 (USBS). _Natrona Co._: Buffalo Creek, 27 mi. N, 1
+     mi. E Powder River, 6075 ft., 16.
+
+
+=Microtus montanus fusus= Hall
+
+     _Microtus nanus_, Bailey, N. Amer. Fauna, 17:30, June 6, 1900
+     (part); Cary, N. Amer. Fauna 33:123, August 17, 1911.
+
+     _Microtus montanus fusus_ Hall, Proc. Biol. Soc. Washington,
+     51:131-134, August 23, 1938; Warren, The Mammals of Colorado, Univ.
+     of Okla. Press, p. 229, 1942.
+
+     _Type._--Male, adult, skin and skull; No. 61281, Museum of
+     Vertebrate Zoology; 2½ miles east of summit of Cochetopa Pass,
+     Saguache County, Colorado; Sept. 21, 1933; collected by Annie M.
+     Alexander; original number 2568. Type not seen by me.
+
+     _Range._--Southern Colorado and northern New Mexico.
+
+     _Comparisons._--For comparison with _M. m. nanus_, the subspecies
+     to the northward, see the preceding account of that subspecies. For
+     comparison with _M. m. amosus_ the subspecies to the west see Hall
+     (1938) and Durrant (1952). I have not examined specimens of
+     _amosus_.
+
+     _Measurements._--Average and extreme measurements for 17 adults
+     including both males and females from several localities in
+     southern Colorado are as follows: total length, 160 (136-179);
+     length of tail, 42 (35-55); hind foot, 19.2 (17-23); condylobasilar
+     length of the skull, 25.2 (24.0-26.0); zygomatic breadth, 15.0
+     (14.1-15.5); alveolar length of upper molar tooth-row, 6.4
+     (6.0-6.7); prelambdoidal breadth, 8.7 (8.3-9.2); lambdoidal
+     breadth, 11.7 (11.1-12.6).
+
+     Average and extreme measurements of 4 adults (2 males and 2
+     females) from the type locality and 11 adults (4 males and 7
+     females) from other localities in southern Colorado are as follows:
+     total length, 162 (157-168), 157 (137-169); length of tail (means
+     only), 44.5, 40.5; hind foot, 18.8 (18-19), 18.6 (18-23);
+     condylobasilar length of skull, 24.5 (24.0-24.7), 25.2 (24.3-26.1);
+     alveolobasilar length, 14.2 (13.9-14.5), 14.6 (14.1-15.1);
+     palatilar length, 13.2 (13.0-13.4), 13.5 (13.1-14.2); alveolar
+     length of upper molar tooth-row, 6.3 (6.0-6.6), 6.4 (6.3-6.7);
+     zygomatic breadth, 15.0 (14.3-15.5), 14.9 (14.1-15.5); interorbital
+     breadth, 3.5 (3.3-3.6), 3.5 (3.3-3.7); lambdoidal breadth, 11.8
+     (11.1-12.6), 11.7 (11.2-12.3); prelambdoidal breadth, 8.6
+     (8.3-9.2), 8.8 (8.3-9.0); depth of braincase, 7.5 (7.2-7.8), 7.6
+     (7.1-7.9).
+
+_Discussion._--There is no sharp boundary between _M. m. fusus_ of
+southern Colorado and the subspecies to the north, _M. m. nanus_.
+Although the line separating these two subspecies is drawn somewhat
+arbitrarily, on the whole the samples from north of this line more
+closely resemble _nanus_. All of the means for total length given above
+are larger than the maximum given in Hall's description of _fusus_. The
+caudal index (38 and 35% in two series) is slightly larger than that
+cited by Hall (33.3%) and is not significantly different from that in
+_nanus_ (35.2%). The color in both young and old mice is variable, but
+in general is more yellowish, and less grayish, than in any other series
+studied.
+
+There is a large area in western Colorado and eastern Utah, between the
+known ranges of _M. m. fusus_ and _M. m. amosus_ from which there are no
+specimens. Probably the species occurs only at certain places in this
+arid region which seems to be a partial barrier to the species.
+
+Specimens of _M. montanus_ from northern New Mexico have been referred
+previously to _M. m. arizonensis_. When he named _M. m. fusus_, Hall
+mentioned its resemblance to _arizonensis_ in reddish coloration, but
+pointed out that _fusus_ is less reddish. Of six specimens from Valle
+Santa Rosa, Jemez Mountains (USBS), 8500 ft., Rio Arriba County, New
+Mexico, three are immature, and the skulls of the remaining specimens
+are damaged. In reddish color and relatively large size these few
+specimens resemble _arizonensis_ more than _fusus_ although the locality
+of occurrence is closer to the geographic range of the northern _fusus_
+than to that of _arizonensis_. The identification of these specimens as
+_arizonensis_ is provisional; additional specimens are needed from the
+area, 200 miles wide, which separates the ranges as now known of
+_arizonensis_ in Arizona from the occurrence in New Mexico. There is a
+single specimen from this area, the damaged skull of which prevents
+conclusive identification. The specimen is either _M. montanus_ or _M.
+mexicanus_, and is from Nutria, on the southern edge of the Zuni
+Mountains (USBS). Detailed comparison of _fusus_ and _arizonensis_ is
+not attempted here although it may be stated that in several characters
+_fusus_ is intermediate between _arizonensis_ to the south and _nanus_
+to the north.
+
+     _Specimens examined._--Total, 89, distributed as follows
+     (abbreviations for collections are given in the account of _M. m.
+     nanus_; localities that are not represented in Fig. 1 because
+     overlapping or crowding of the symbols would result are
+     Italicized):
+
+     COLORADO: _Pitkin Co._: 5 mi. W Independence Pass, 11,000 ft., 1
+     (Chi). _Lake Co._: _Independence Pass, 12,095 ft._, 2 (Chi).
+     _Gunnison Co._: _Gothic_, 2 (USBS); Decker's Ranch, Crested Butte,
+     2 (AMNH); Almont, 3 (USBS). _Montrose Co._: Coventry, 5 (USBS 4,
+     AMNH 1). _Saguache Co._: Cochetopa Pass and environs, 44 (USBS
+     22). _Hinsdale Co._: Ruby Lake, 1 (USBS). _Mineral Co._: 3 mi. E
+     Creede, 1; 23 mi. S, 11 mi. E Creede, 9300 ft., 7. _La Plata Co._:
+     Florida, 6800 ft., 1. _Conejos Co._: 1 mi. S, 19 mi. W Antonito,
+     10,200 ft., 3; _4 mi. S, 23 mi. W Antonito_, 1; _5 mi. S, 24 mi. W
+     Antonito, 9600 ft._, 9.
+
+     NEW MEXICO: _Rio Arriba Co._: 6 mi. W Hopewell, 9900 ft., 6 (USBS);
+     _Tusas River, 8700 ft._, 1 (USBS).
+
+TABLE 1. AVERAGE MEASUREMENTS, IN MILLIMETERS, OF ADULTS OF MICROTUS
+MONTANUS.
+
+Key to column headings:
+
+A: No. of individuals averaged
+B: Total length
+C: Length of tail
+D: Length of hind foot
+E: Condylobasilar length
+F: Alveolar length of upper molar tooth-row
+G: Zygomatic breadth
+H: Lambdoidal breadth
+I: Prelambdoidal breadth
+
+=========================================================================
+       Locality   |   A  |  B  |  C  |  D  |  E  |  F  |  G  |  H  |  I
+------------------+------+-----+-----+-----+-----+-----+-----+-----+-----
+                  |               _M. m. codiensis_, all
+                  |
+Average           | 34   |165.3|44.2 |19.6 |25.47| 6.56|15.55|12.05|8.76
+2 × stand. error  |      | 3.56|1.84 |.395 | .308| .067| .198| .144|.129
+Coeff. variab     |      |  6.0|11.6 | 5.6 |  3.5|  3.0| 3.65| 1.20|1.47
+------------------+------+-----+-----+-----+-----+-----+-----+-----+-----
+                  |            _M. m. nanus_, Eastern Idaho
+                  |
+Average           | 21[1]|151.1|39.4 |19.2 |25.00| 6.44|14.99|11.74|8.94
+2 × stand. error  |      | 3.20|2.89 |.293 | .286| 1.15| .295| .210|.182
+Coeff. variab.    |      |  6.1|21.1 |4.38 | 2.49| 3.99| 4.10| 3.79|4.31
+------------------+------+-----+-----+-----+-----+-----+-----+-----+-----
+                  |            _M. m. nanus_, Wyoming
+                  |
+Teton Co.         | 35   |160.5|40.7 |18.6 |25.16| 6.51|15.17|11.86|8.77
+Fremont Co.       | 26   |157.0|41.4 |19.6 |25.23| 6.25|15.05|11.88|8.91
+Lincoln Co.       | 24   |159.9|41.8 |18.9 |25.08| 6.26|15.10|11.82|8.75
+Uinta Co.         | 26   |162.4|41.3 |19.0 |25.33| 6.42|15.31|12.16|8.89
+Sweetwater Co.    | 12   |159.8|43.7 |20.1 |24.98| 6.31|15.00|11.84|9.02
+Natrona Co.       | 40   |159.6|41.0 |19.6 |25.04| 6.40|15.00|11.84|8.93
+Carbon Co.        |108   |158.7|40.0 |19.1 |24.96| 6.27|15.05|11.83|8.72
+ Encampment ♂     | 27   |161  |41.7 |18.9 |25.1 | 6.16|15.2 |11.9 |8.7
+ Encampment ♀     | 11   |159  |41.1 |19.4 |24.9 | 6.18|14.9 |11.6 |8.6
+ Savery ♂         | 23   |159  |41.0 |19.2 |25.3 | 6.32|15.2 |12.2 |8.8
+ Savery ♀         | 25   |155  |37.1 |18.8 |24.7 | 6.33|14.8 |11.6 |8.6
+------------------+------+-----+-----+-----+-----+-----+-----+-----+-----
+                  |                     _M. m. nanus_
+                  |
+Northern Colo.    |  8   |163.1|42.4 |19.6 |25.20| 6.44|14.86|11.70|8.56
+------------------+------+-----+-----+-----+-----+-----+-----+-----+-----
+                  |                     _M. m. fusus_
+                  |
+Southern Colo.    | 17[2]|159.8|42.4 |19.2 |24.97| 6.43|14.98|11.73|8.69
+------------------+------+-----+-----+-----+-----+-----+-----+-----+-----
+
+[Footnote 1: For external parts, 34 individuals were used.]
+
+[Footnote 2: For external parts, 29 individuals were used.]
+
+Some measurements not given above are included in Table 1, together with
+the number of specimens and the sex if restricted to one sex. So that
+the variability can be evaluated more adequately, the coefficient of
+variability and 2 times the standard error of the mean are included for
+the measurements in two series. The series consist of all the adult
+specimens (with a condylobasilar length of 24.0 mm. or more) of both
+sexes from the areas specified. Various barriers are shown in Fig. 2 for
+comparison with the distributions of the subspecies and the localities
+of known occurrence shown in Fig. 1. _Microtus montanus_ has not been
+taken in the Black Hills area of extreme northeastern Wyoming. Suitable
+montane habitat is present and both _Microtus pennsylvanicus insperatus_
+and _Microtus longicaudus longicaudus_ occur there. The arid basin of
+the Powder River presumably is a barrier that has prevented _M.
+montanus_ from reaching this area.
+
+
+REFERENCES CITED
+
+BAILEY, V.
+
+     1900. Revision of American voles of the genus _Microtus_. N. Amer.
+     Fauna, 17:1-88, June 6.
+
+     1917. A new subspecies of meadow mouse from Wyoming. Proc. Biol.
+     Soc. Washington, 30:29-30, February 21.
+
+CARY, M.
+
+     1911. A biological survey of Colorado. N. Amer. Fauna, 33:1-256,
+     August 17.
+
+     1917. Life zone investigations in Wyoming. N. Amer. Fauna, 42:1-95,
+     October 3.
+
+DAVIS, W. B.
+
+     1939. The Recent mammals of Idaho. 400 p., front., illus., maps,
+     diagrs., Caldwell, Id., The Caxton Printers Ltd., April 5.
+
+DURRANT, S. D.
+
+     1952. Mammals of Utah, taxonomy and distribution. Univ. Kans.
+     Publ., Mus. Nat. Hist., 6:1-549, 91 figs, in text, 30 tables,
+     August 10.
+
+ELTON, C.
+
+     1942. Voles, mice and lemmings; problems in population dynamics.
+     496 p., illus. (maps), tables, Oxford, The Clarendon Press, London.
+
+GOIN, O. B.
+
+     1943. A study of individual variation in _Microtus pennsylvanicus
+     pennsylvanicus_. Jour. Mamm., 24:212-224, June 7.
+
+HALL, E. R.
+
+     1938. Notes on the meadow mice _Microtus montanus_ and _Microtus
+     nanus_ with descriptions of a new subspecies from Colorado. Proc.
+     Biol. Soc. Washington, 51:131-134, August 23.
+
+HILL, E. A.
+
+     1935. Cranial foramina in rodents. Jour. Mamm. 16(2):121-129.
+
+HOWELL, A. B.
+
+     1924. Individual and age variation in _Microtus montanus yosemite_.
+     Jour. Agr. Res., 28(10):977-1015, June 7.
+
+KELLOGG, R.
+
+     1922. A study of the California forms of the _Microtus montanus_
+     group of meadow mice. Univ. California Publ. Zool., 21:245-274,
+     April 18.
+
+MERRIAM, C. H.
+
+     1891. Results of a biological reconnaissance of south-central
+     Idaho. N. Amer. Fauna, 5:1-113, July 30.
+
+PIPER, S. E.
+
+     1909. The Nevada mouse plague of 1907-8. U. S. Dept. Agr., Farmers'
+     Bul. 352, pp. 1-23, 9 figs.
+
+WARREN, E. R.
+
+     1942. The mammals of Colorado, their habits and distribution.
+     Second (revised) edition, Univ. Oklahoma Press, Norman, xviii-330
+     p., front., 50 plates.
+
+
+_Transmitted March 22, 1954._
+
+
+
+â–¡
+
+25-3560
+
+
+       *       *       *       *       *
+
+Transcriber's Notes
+
+Italic typeface in the original is indicated by by _underscores_.
+
+Bold typeface in the original is indicated by =equals=.
+
+One typographical error was corrected: "Castlford" was corrected to
+"Castleford" in "Twin Falls Co.: Castleford Fenced Plot".
+
+
+
+
+
+End of the Project Gutenberg EBook of Subspeciation in the Meadow Mouse,
+Microtus montanus, in Wyoming and , by Sydney Anderson
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+The Project Gutenberg EBook of Subspeciation in the Meadow Mouse, Microtus
+montanus, in Wyoming and Colorado, by Sydney Anderson
+
+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: Subspeciation in the Meadow Mouse, Microtus montanus, in Wyoming and Colorado
+
+Author: Sydney Anderson
+
+Release Date: March 22, 2010 [EBook #31730]
+
+Language: English
+
+Character set encoding: ISO-8859-1
+
+*** START OF THIS PROJECT GUTENBERG EBOOK MEADOW MOUSE ***
+
+
+
+
+Produced by Chris Curnow, Simon Gardner, Joseph Cooper and
+the Online Distributed Proofreading Team at
+http://www.pgdp.net
+
+
+
+
+
+
+
+UNIVERSITY OF KANSAS PUBLICATIONS
+
+MUSEUM OF NATURAL HISTORY
+
+Volume 7, No. 7, pp. 489-506, 2 figures in text
+
+July 23, 1954
+
+
+
+
+                    Subspeciation in the Meadow Mouse,
+                      Microtus montanus, in Wyoming
+                              and Colorado
+
+                                  BY
+
+                            SYDNEY ANDERSON
+
+
+UNIVERSITY OF KANSAS
+
+LAWRENCE
+
+1954
+
+
+UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY
+
+Editors: E. Raymond Hall, Chairman, A. Byron Leonard,
+Robert W. Wilson
+
+Volume 7, No. 7, pp. 489-506, 2 figures in text
+
+Published July 23, 1954
+
+
+UNIVERSITY OF KANSAS
+
+Lawrence, Kansas
+
+
+PRINTED BY
+
+FERD VOILAND, JR., STATE PRINTER
+
+TOPEKA, KANSAS
+
+1954
+
+25-3560
+
+
+
+
+Subspeciation in the Meadow Mouse, Microtus montanus, in Wyoming and
+Colorado
+
+BY
+
+SYDNEY ANDERSON
+
+
+_Microtus montanus_ reaches the eastern limits of its geographic
+distribution in Wyoming and Colorado. There the mountains, but in
+general not the lowlands, are occupied by this species. A certain
+minimum of moisture may be of direct importance to the mouse and
+certainly is indirectly important, because certain hydrophytic or
+mesophytic grasses used by the mouse for food, for protection from
+enemies, and for shelter from the elements are dependent on the
+moisture. Areas suitable for _Microtus montanus_ are separated by
+deserts that are dominated by sagebrush and other xerophytic plants or
+by forests or rocky exposures at higher altitudes. A relatively small
+percentage, probably less than ten per cent, of the total area even in
+the more favorable parts of the range of the species is suitable for
+occupancy. In these mice, as in other microtines (Elton, 1942; Piper,
+1909), there are seasonal, and irregularly multiannual fluctuations in
+population density, which sometimes are extreme. Consequently the mice
+at some times seem to be absent from suitable habitats, and at some
+other times occur there in amazingly large numbers.
+
+Because the species is broken up into partly isolated, or at times
+completely isolated, colonies or local populations it may be supposed
+that various evolutionary forces such as selection and random genetic
+drift operate to foster variation. The degree to which racial
+distinction is attained may depend upon these forces and the time
+available. In _Microtus montanus_ in the eastern Rocky Mountains the
+degree of subspecific distinction is not great.
+
+The study here reported upon is based on 1,187 specimens of _Microtus
+montanus_ from Wyoming, Colorado, Idaho and Montana, and on work in the
+field. I spent approximately four months in the field in this area, in
+the summers of 1950, 1951, and 1952. The specimens studied were arranged
+according to localities and the larger series were compared
+statistically. Each of two series, totaling 136 specimens, was studied
+intensively to ascertain the kind and range of variation within single
+populations. Twenty-seven measurements, various proportions based on
+these measurements, and differences in color were analyzed. Fifteen
+characters, judged to be most significant, were selected for use in
+comparing all series. In addition, certain characters that can not be
+expressed easily by measurements, such as inflation of the auditory
+bullae and the curvature of the zygomatic arch, were observed. The
+studies by A. B. Howell (1924) of variation in _Microtus montanus
+yosemite_ Grinnell in California and those by O. B. Goin (1943) of
+_Microtus pennsylvanicus pennsylvanicus_ (Ord) were useful. The external
+measurements are from the collectors' field labels. The measurements of
+the skull all were taken with dial calipers reading to a tenth of a
+millimeter. The anteroposterior measurements of the skull all were taken
+along the shortest line between the points specified below and are not
+necessarily along a line parallel to the long axis of the skull. These
+measurements were taken on the left side of the skull whenever possible.
+Some of the skulls are damaged and therefore some measurements could not
+be taken and are omitted in the computations. Measurements are in
+millimeters.
+
+The results of these studies were submitted to the Department of Zoology
+and the Graduate School of the University of Kansas in partial
+fulfillment of the requirements for the degree of Master of Arts (1952)
+and are available in manuscript form at the Museum of Natural History
+and the library of the University of Kansas.
+
+
+EXPLANATION OF MEASUREMENTS
+
+     Caudal index.--the length of the tail expressed as a percentage of
+     the length of the head and body. The length of the head and body is
+     the collector's measurement of total length less the length of the
+     tail.
+
+CRANIAL MEASUREMENTS OF LENGTH.
+
+     Condylobasilar length.--from the exoccipital condyle to the most
+     posterior point on the border of the alveolus of the upper incisor.
+
+     Alveolobasilar length.--from the posterior border of the alveolus
+     of the third upper molar to the posterior border of the alveolus of
+     the incisor.
+
+     Palatilar length.--from the anteriormost part of the posterior
+     border of the bony shelf of the palate to the posteriormost part of
+     the alveolus of the incisor.
+
+     Alveolar length of upper molar tooth-row.--from the most posterior
+     point of the alveolus of the third upper molar to the most anterior
+     point of the alveolus of the first upper molar.
+
+MEASUREMENTS OF BREADTH.
+
+     Zygomatic breadth.--greatest transverse width.
+
+     Interorbital breadth.--the breadth of the interorbital
+     constriction.
+
+     Lambdoidal breadth.--between the lateralmost points of the
+     lambdoidal crest.
+
+     Prelambdoidal breadth.--between the medialmost margins of the
+     prominent fenestrae in the posterodorsal parts of the squamosal
+     bones. To these fenestrae Howell (1924:995) applied the adjective
+     "prelambdoidal," but other authors have used other names (see Hill,
+     1935:127).
+
+     Depth of braincase.--shortest distance from the ventral side of the
+     cranium at the suture between the basioccipital and basisphenoid
+     bones to the dorsal surface of the cranium (usually not
+     perpendicular to the long axis of the skull).
+
+The history of our knowledge of _Microtus montanus_ in this area begins
+with the early work of the United States Bureau of Biological Survey
+directed by C. H. Merriam (1891), and participated in by Vernon Bailey
+(1900, 1917), Merritt Cary (1911, 1917), and others. The changes in
+nomenclature which grew out of increased understanding of these mice
+through additional collecting are expressed in the synonymies under the
+accounts of subspecies. As a result of my studies two of the three
+subspecific names previously proposed for mice from this area have been
+retained although changes are proposed in the ranges assigned to the two
+subspecies and two additional heretofore unrecognized subspecies are
+named and described. Furthermore the additional specimens and my studies
+of variation make modifications in the characterization of these
+subspecies necessary. Not all of the samples here assigned to a single
+subspecies are identical and I therefore list and discuss some of the
+local variants.
+
+     Numerous members of summer field parties from the Museum of Natural
+     History at the University of Kansas collected most of the specimens
+     studied and wrote field notes that have been helpful. I am grateful
+     to these persons and to Professor E. R. Hall and Dr. R. H. Baker
+     for their assistance and helpful suggestions. Specimens in the
+     following museums were made available by their respective curators:
+     Chicago Natural History Museum by Mr. Colin C. Sandborn, The Museum
+     of Zoology at the University of Michigan by Dr. E. T. Hooper, The
+     American Museum of Natural History by Mr. G. G. Goodwin, The United
+     States National Museum by Dr. David H. Johnson and the Biological
+     Surveys Collection by Miss Viola S. Schantz. A fellowship from the
+     National Science Foundation made possible the studies at the
+     museums other than at the University of Kansas.
+
+
+
+=Microtus montanus nanus= (Merriam)
+
+     _Arvicola (Mynomes) nanus_ Merriam, N. Amer. Fauna, 5:62, July 30,
+     1891.
+
+     _Microtus montanus nanus_, Hall, Proc. Biol. Soc. Washington,
+     51:131, August 23, 1938.
+
+     _Microtus nanus_, Bailey, N. Amer. Fauna, 17:30, June 6, 1900
+     (part).
+
+     _Microtus montanus caryi_ Bailey, Proc. Biol. Soc. Washington,
+     30:29, February 21, 1917.
+
+     _Microtus nanus nanus_, Cary, N. Amer. Fauna, 42:43, October 3,
+     1917 (part).
+
+     _Type._--Adult male No. 23853/31253, U. S. National Museum,
+     Department of Agriculture collection, from Pahsimeroi Mountains,
+     Custer County, Idaho; obtained by C. Hart Merriam and Vernon
+     Bailey, September 16, 1890.
+
+     _Range._--Idaho; southwestern Montana; most of the southwestern
+     half of Wyoming; southward to central Colorado. See figure 1.
+
+     _Comparisons._--Comparisons with subspecies named as new in this
+     paper will be found in the accounts of those subspecies beyond.
+     From _Microtus montanus fusus_ Hall, the subspecies to the south,
+     _M. m. nanus_ from Idaho differs as follows: averages smaller;
+     slightly darker and less reddish and less yellowish in color;
+     slightly wider braincase (see measurement of prelambdoidal
+     breadth); larger bullae.
+
+     _Measurements._--Average (= arithmetical mean) measurements of 34
+     specimens, both male and female, from several localities in eastern
+     Idaho are: total length, 151; length of tail, 39; hind foot, 19.2;
+     condylobasilar length of the skull, 25.0; zygomatic breadth, 15.0;
+     alveolar length of upper molar tooth-row, 6.4; prelambdoidal
+     breadth, 8.9; and lambdoidal breadth, 11.7.
+
+     Average and extreme measurements of six adult males from near
+     Pocatello, Bannock County, Idaho, and nine adult males from near
+     Afton, Lincoln County, Wyoming, are, respectively, as follows:
+     total length, 143(135-150), 163(143-179); length of tail,
+     35.1(33-38), 42.8(36-49); caudal index, 32.0(28.0-33.1),
+     35.7(30.6-41.9); hind foot, 18.9(18-20), 18.8(17-20);
+     condylobasilar length of skull, 24.4(24.0-26.0), 25.6(24.5-26.2);
+     alveolobasilar length, 14.1(13.7-14.5), 14.6(13.8-15.0); palatilar
+     length, 13.2(12.9-13.6), 13.8(13.2-14.5); alveolar length of upper
+     molar tooth-row, 6.3(6.1-6.5), 6.3(6.0-6.6); depth of braincase,
+     7.7(7.5-7.9), 8.0(7.7-8.3); lambdoidal breadth, 11.4(11.0-11.7),
+     12.0(11.3-12.7); prelambdoidal breadth, 9.1(8.6-9.4), 8.7(8.0-9.4);
+     zygomatic breadth, 14.3(13.8-14.7), 15.3(14.4-16.3); interorbital
+     breadth, 3.6(3.5-3.7), 3.5(3.3-3.7). The average length of the
+     nasal bones in the series from Pocatello is 7.1 mm. The averages,
+     which have not been included in Table 1, for three measurements of
+     the series from Carbon County, Wyoming, are as follows (Encampment,
+     males; Encampment, females; Savery, males; and Savery, females,
+     respectively): alveolobasilar length, 14.4, 14.3, 14.5, 14.3;
+     interorbital breadth, 3.5, 3.4, 3.5, 3.4; depth of braincase, 7.8,
+     7.6, 7.9, 7.6. Additional measurements are included in Table 1 for
+     other series.
+
+_Discussion._--The name _Microtus montanus caryi_ Bailey is here placed
+in synonymy under _M. m. nanus_ (Merriam). Vernon Bailey (1917) in his
+description of _caryi_ made four assumptions that have been found to be
+entirely or partly invalid. First, he assumed that this is an "extreme
+variant which gradually changes in characters across Nevada and Utah,
+and reaches its maximum variation in Wyoming." The differences pointed
+out in subsequent descriptions of subspecies found in the above area do
+not show a gradual change in any character, or in the number of
+characters, nor is _caryi_ an extreme when compared with the other
+subspecies. Second, _Microtus nanus_ was not, as Bailey assumed, a
+different species than _Microtus montanus_. Third, he assumed that the
+characteristics of adults of _nanus_ were adequately ascertainable from
+the thirteen topotypes available to him. Subsequent sampling from Idaho
+shows that the series of specimens available to Bailey was made up
+mostly of young and subadult animals. Finally, _caryi_ does not occupy
+as Bailey stated "the meadows along streams in the arid sagebrush
+country of the Bear River, Green River, and Wind River valleys"
+exclusively, or characteristically. When the localities from which the
+species actually is known are plotted, it seems that the arid basin
+serves as a barrier and that the species is more commonly and abundantly
+found in montane meadows in the Transition and Canadian life-zones.
+
+Certain samples, here assigned to _M. m. nanus_, that vary from the
+average of the subspecies deserve comment. For example, mice from the
+area in Wyoming southwest of the Green River (in the Uinta Mountains)
+have relatively smaller feet, but are larger in both total length and
+size of skull. Specimens from near Afton, Lincoln County, Wyoming, are
+relatively large in both total length and size of skull. This series and
+specimens from Teton County, Wyoming, are intermediate between _nanus_
+from Idaho and the newly named subspecies from near Cody, Park County,
+Wyoming, described below, in terms of both darkness and the amount of
+reddish color. Mice from Laramie County are more richly reddish-brown.
+The specimens from near Savery, in Carbon County, Wyoming, are darker.
+The alveolobasilar length relative to the condylobasilar length is
+smaller in the series from along Deer Creek, 16 mi. S, 11 mi. W Waltman,
+Natrona County, Wyoming. The series from the southern tier of counties
+in Wyoming and some of the specimens from Colorado have relatively wider
+zygomatic arches. The specimens from southern Sweetwater County,
+Wyoming, are relatively paler, have a relatively longer tail and longer
+hindfeet, lesser condylobasilar length, and wider braincase. Most of
+these variations are of questionable significance; they may be chance
+variations owing to errors in sampling.
+
+Much of the south-central part of the state is relatively low and
+relatively arid. This area includes the arid basin of the Green River
+and its major tributaries and the arid Red Desert along the continental
+divide in Sweetwater County. This area might have acted as a barrier to
+the mice; gene flow might have been prevented between the populations of
+the western part of the state and those farther east in the Medicine Bow
+Mountains and Laramie Mountains. Nevertheless geographic variations of
+subspecific worth have not taken place. The barrier has either not been
+of as long duration, or has not been so complete and effective, as the
+other barriers in the state, namely the Absaroka Range, the Big Horn
+Basin, the Shoshone Basin, and the valley of the North Platte River.
+These four barriers presumably have led to the differentiation of the
+two subspecies that are newly named beyond. Each of the two areas which
+is set apart by these barriers and in which one of the newly named
+subspecies has evolved is small; therefore there is a lesser amount of
+suitable habitat available for each of the newly named mice than there
+is for _M. m. nanus_. It is conceivable, therefore, that in periods of
+adverse conditions in each of the small areas the size of the effective
+breeding population may have been so small that random genetic drift
+could have operated effectively, or that selection was more critical
+than in a larger, more stable population. It is difficult to test these
+possibilities because the selective value of the taxonomic characters is
+unknown. The observed pattern of variation and facts of distribution
+are, however, not contradictory to the above possibilities.
+
+     _Specimens examined._--Total, 993, distributed as follows: All
+     specimens unless otherwise indicated are in the University of
+     Kansas Museum of Natural History. Specimens in other museums are
+     labeled as follows: Chicago Natural History Museum (Chi);
+     University of Michigan, Museum of Zoology (Mich); American Museum
+     of Natural History (AMNH); United States National Museum (USNM);
+     Biological Surveys Collection (USBS). Localities that are not
+     represented in Fig. 1 because overlapping or crowding of the
+     symbols would result are Italicized. Localities are arranged from
+     north to south by states, within a state from northwest to
+     southeast by counties, and within a county from north to south.
+
+     WYOMING: _Yellowstone Park_: Canyon Camp, 1 (USBS); Lower Geyser
+     Basin, 1 (USBS); Upper Yellowstone River, 2 (AMNH); _North end of
+     Lake_, _Yellowstone National Park_, 2 (AMNH). _Teton Co._: Pacific
+     Creek, 1 (USBS); _Big Game Ridge_, 3 (USBS 1, Mich 2); _Whetstone
+     Creek_, 7 (Mich); Moran and environs (6 localities within a 5 mile
+     radius), 28 (USBS 2, Mich 5); _S fork Buffalo River_, 7 (AMNH); 2
+     mi. W pass, Black Rock Creek, 1 (USBS); _Jenny Lake_, 5 (Mich); Bar
+     BC Ranch, 2½ mi. NE Moose, 6500 ft., 2; Teton Pass above Fish
+     Creek, 1 (USBS); Jackson and environs, 142 (Mich 141); _Sheep
+     Creek_, 2 (Mich). _Lincoln Co.: 13 mi. N, 2 mi. W Afton_, 2; _10
+     mi. N, 2 mi. W Afton_, 4; _9½ mi. N, 2 mi. W Afton_, 3; 9 mi. N,
+     2 mi. W Afton, 9; _7 mi. N, 1 mi. W Afton_, 12; Afton, 1 (USBS);
+     Labarge Creek, 1 (USBS); Border, 6 (USBS); _Cokeville_, 2 (USBS); 6
+     mi. N, 2 mi. E Sage, 1; Cumberland, 5 (USBS). _Sublette Co._: 34
+     mi. N, 4 mi. W Pinedale, 1; _33 mi. N, 2 mi. W Pinedale_, 6; _32
+     mi. N, 1 mi. W Pinedale_, 1; _31 mi. N Pinedale_, 4; Dell Creek, on
+     Ferris Ranch, 7 (Mich); Horse Creek, 7800 ft., Merna, 4 (USBS); Big
+     Piney, 1 (USBS). _Fremont Co._: 17½ mi. W, 2½ mi. N Lander,
+     9500 ft., 3; _17 mi. W, 2 mi. N Lander, 9300 ft._, 4; Milford and
+     environs (5 localities within a 1 mile radius), 23 (USBS 4); 15½
+     mi. S, 7½ mi. W Lander, 9200 ft., 1; South Pass City, 8000 ft, 8
+     (USBS); _23½ mi. S, 5 mi. W Lander, 8600 ft._, 7. _Natrona Co._:
+     Deer Creek, 16 mi. S, 11 mi. W Waltman, 6950 ft., 44; 6 mi. S, 2
+     mi. W Casper, 5900 ft., 4; _6-4/5 mi. S, 2 mi. W Casper, 6100 ft._,
+     1; _7 mi. S, 2 mi. W Casper, 6370 ft._, 3; _10 mi. S Casper, 7750
+     ft._, 33; Sun, 2 (USBS); _5 mi. W Independence Rock, 6000 ft._, 4;
+     _5 mi. W, 1 mi. S Independence Rock_, 2. _Converse Co._: Beaver, 1
+     (USBS). _Uinta Co.: 1½ mi. W, ½ mi. S Cumberland_, 6; _16 mi.
+     S, 2 mi. W Kemmerer, 6700 ft._, 3; 10 mi. SW Granger, 3 (Mich);
+     Fort Bridger, 6650 ft., 25 (USNM 6); 9 mi. S Robertson, 8000 ft.,
+     9; _9½ mi. S, ½ mi. W Robertson, 8600 ft._, 1; _10 mi. S, 1
+     mi. W Robertson, 8700 ft._, 25; _14 mi. S, 2 mi. E Robertson, 9000
+     ft._, 5; 4 mi. S Lonetree, 1 (USBS). _Sweetwater Co._: Farson, 3;
+     Bitter Creek, 3 (AMNH); Kinney Ranch, 21 mi. S Bitter Creek, 6800
+     ft., 9 (USNM 1, AMNH 2); 32 mi. S, 22 mi. E Rock Springs, 7025 ft.,
+     on Vermillion Creek, 15. _Carbon Co._: 18 mi. NNE Sinclair, 6500
+     ft., 10; Bridgers Pass, 18 mi. SW Rawlins, 7500 ft., 7; Saratoga, 1
+     (USBS); _6 mi. S, 13 mi. E Saratoga, 8500 ft._, 5; _6 mi. S, 14 mi.
+     E Saratoga, 8800 ft._, 1; Lake Marie, 10,440 ft., 2; _1 mi. S Lake
+     Marie_, 2; _½ mi. S, 2 mi. E Medicine Bow Peak, 10,800 ft._, 1;
+     Encampment (12 localities from 10 mi. N, 14 mi. E to 9 mi. N, 3 mi.
+     E Encampment and from 6500 to 8400 ft.), 63; 1/4 mi. N Riverside,
+     7380 ft., 2; S base Bridger Peak, 8800 ft., Sierra Madre Mountains,
+     1; _2 mi. S Bridger Peak, 9300 ft._, 2; Savery (10 localities from
+     8 mi. N, 21 mi. E to 4 mi. N, 8 mi. E Savery and from 7300 to 8800
+     ft.), 80. _Albany Co._: _30 mi. N, 10 mi. E Laramie, 6760 ft._, 6;
+     _29¾ mi. N, 9½ mi. E Laramie, 6350 ft._, 1; 26 mi. N, 4½
+     mi. E Laramie, 6960 ft., 8; _26¾ mi. N, 6½ mi. E Laramie,
+     6700 ft._, 3; _3 mi. N, 13 mi. E Laramie, 7500 ft._, 1; _7 mi. N, 2
+     mi. E Laramie_, 1 (Chi); 5 mi. N Laramie, 7400 ft., 15; _Laramie_,
+     4 (AMNH); _1 mi. E Laramie, 7160 ft._, 4; 7-7/10 mi. SSW Laramie,
+     7200 ft., 4; 6½ mi. S, 8¾ mi. E Laramie, 8200 ft., 1;
+     _Headquarters Park, 10,200 ft., Medicine Bow Mountains_, 3 (USBS);
+     Centennial, 8120 ft., 1; _2-1/4 mi. ESE Brown's Peak, 10,300 ft._,
+     3; _3 mi. ESE Brown's Peak, 10,000 ft._, 12; _2 mi. S Brown's Peak,
+     10,600 ft._, 1; _Pole Mountain, 15 mi. SE Laramie_, 4 (USBS 3); _1
+     mi. SSE Pole Mountain, 8350 ft._, 4; _2 mi. SW Pole Mountain, 8300
+     ft._, 13; _3 mi. S Pole Mountain, 8100 ft._, 1; Sherman, 2 (AMNH).
+     _Laramie Co._: 5 mi. N, 1 mi. W Horse Creek P. O., 7200 ft., 1;
+     Meadow, 2 (USBS); 11 mi. N, 5½ mi. E Cheyenne, 5450 ft., 7; _7
+     mi. W Cheyenne, 6500 ft._, 10; Cheyenne, 3 (USNM).
+
+     COLORADO: _Moffat Co._: Lay, 6160 ft., 1 (AMNH). _Routt Co._:
+     Wright's Ranch, Yampa, 7700 ft., 2; Gore Range, 8 mi. E Toponas,
+     8000 ft., 2 (USBS). _Larimer Co._: _12½ mi. W, 1½ mi. S
+     Rustic_, 1; 11 mi. W, 1 mi. S Rustic, 1; Cache La Poudre River, 1
+     (Chi); _Estes Park_, 3 (USBS 1, AMNH 2); 19½ mi. W, 2½ mi. S
+     Loveland, 7280 ft., 6; _16 mi. W Loveland, 6840 ft._, 1; 6 mi. W,
+     ½ mi. S Loveland, 5200 ft., 1. _Rio Blanco Co._: Meeker, 1
+     (USBS); _9½ mi. SW Pagoda Peak_, 7700 ft., 3; 5 mi. S Pagoda
+     Peak, 9100 ft., 2. _Eagle Co._: Eagle, 1 (USBS); Pando, 2 (USBS).
+     _Grand Co._: Mt. Whiteley, 2 (USBS); Arapahoe Pass, Rabbit Ear
+     Mountains, 2 (USBS); Coulter (near Granby), 5 (USBS); _Arrowhead_
+     (near Dale), 1 (USBS). _Boulder Co._: ¾ mi. N, 2 mi. W
+     Allenspark, 8400 ft., 4; _3 mi. S Ward_, 9000 ft., 3; Nederland, 16
+     (Chi). _Clear Creek Co._: Mt. McLellan, 2 (USBS); Berthoud Pass, 4.
+     _Park Co._: Trout Creek Ranch, 2 mi. N Garo, 1 (USBS).
+
+     Specimens examined of _M. m. nanus_ from eastern Idaho and Montana
+     are as follows: IDAHO: _Custer Co._: Challis, 7 (USBS); Mill Creek,
+     Challis Nat. Forest, 1 (USBS); Pahsimeroi Mts., 12 (USBS); Lost
+     River Mts., 1 (USBS). _Fremont Co._: N fork Snake River, 10 mi. SW
+     Island Park, 6200 ft., 2 (AMNH); Black Springs Creek, 4 mi. W
+     Ashton, 5200 ft., 1 (AMNH); 5 mi. W St. Anthony, 5000 ft., 1
+     (AMNH). _Camas Co._: Camas Prairie, Corral, 5100 ft., 2 (USBS).
+     _Blaine Co._: Alturas Lake, 3 (USBS); _Sawtooth Lake_, 2 (USBS);
+     Craters of the Moon, Laidlow Park, 2 (Mich); Ticura, 10 mi. S
+     Picabo, 1 (USBS); 19 mi. NE Carey (Lava Lake), 8 (Mich). _Butte
+     Co._: _26 mi. SW Arco_, 12 (Mich). _Bingham Co._: Shelley, 6
+     (USBS). _Bonneville Co._: 10 mi. SE Irwin, 4 (USBS). _Owyhee Co._:
+     Three Creeks, 3 (USBS). _Twin Falls Co._: Castleford Fenced Plot,
+     11 mi. W, 9 mi. S Twin Falls, 1. _Minidoka Co._: _Heyburn_, 2
+     (USBS). _Cassia Co._: 2 mi. S, 2 mi. W Burley, 5. _Bannock Co._:
+     Pocatello, 23 (USBS 4); Swan Lake, 1 (USBS). _Bear Lake Co._:
+     Montpelier Creek, 6700 ft., 3 (USBS). MONTANA: _Gallatin Co._: W.
+     Fork of W. Fork, Gallatin River, 1 (USBS). _Park Co._: Lamar River,
+     7000 ft., 1 (USBS); Gardiner, 1 (USBS). _Sweet Grass Co._: 14 mi. S
+     Big Timber, 1 (USBS); _McLeod_, 1 (USBS); West Boulder Creek, 18
+     mi. SE Livingston, 2 (USBS).
+
+
+=Microtus montanus codiensis=, new subspecies
+
+     _Type._--Female, adult, skin and skull; No. 27578, Museum of
+     Natural History, University of Kansas, from 3-1/5 mi. E and 3/5 mi.
+     S Cody, 5020 ft., Park Co., Wyoming; obtained on August 11, 1948,
+     by James W. Bee, original number 18-8-11-48.
+
+     _Range._--In northwestern Wyoming eastward from the Absaroka and
+     Wind River ranges into the western part of the Big Horn Basin.
+
+     _Diagnosis._--A relatively large _Microtus montanus_; tail actually
+     and relatively long; hind foot actually but not relatively large;
+     skull large; zygomatic expanse actually and relatively large;
+     alveolobasilar length relatively large; upper molar tooth-row
+     relatively long; color relatively light, not reddish.
+
+[Illustration: FIG. 1. Geographic range of _Microtus montanus_ in
+Wyoming, Colorado, and adjacent areas. The solid circles represent
+localities from which specimens have been examined; the hollow circles
+represent type localities. The ranges of subspecies in Utah are after
+Durrant, 1952.
+
+Guide to subspecies
+
+1. _M. m. nanus_
+2. _M. m. codiensis_
+3. _M. m. zygomaticus_
+4. _M. m. fusus_
+5. _M. m. micropus_
+6. _M. m. nexus_
+7. _M. m. amosus_
+8. _M. m. rivularis_
+]
+
+     _Comparisons._--As compared with the specimens of _M. m. nanus_
+     from Idaho, the size is larger (see diagnosis and measurements).
+     Certain proportions which differ from those of _nanus_ and which
+     are not in close agreement with the observed differences with age
+     in specimens of _nanus_ of a size comparable to _codiensis_ are
+     relatively large alveolobasilar length, relatively long alveolar
+     length of upper molar tooth-row, relatively wide-spreading
+     zygomatic arches, and relatively long tail. The color in
+     _codiensis_ is lighter than in _nanus_. As compared to the new
+     subspecies named below from the Big Horn Mountains to the east,
+     _codiensis_ is of similar size in head-body length, but has a
+     relatively as well as actually longer tail; the hind foot averages
+     longer; the upper molar tooth-row is relatively longer; the color
+     is slightly paler and less grizzled; the bullae are larger and less
+     flattened; the angle formed at the suture between the basioccipital
+     and basisphenoid bones is less acute; and the region of the suture
+     is less prominently elevated between the bullae when viewed from
+     the ventral aspect. The pterygoid plates mesial and posterodorsal
+     to the posterior end of the last upper molar are less fenestrated,
+     and the incisive foramina are less constricted posteriorly.
+
+[Illustration: FIG. 2. Map showing the major barriers to _Microtus
+montanus_ in Wyoming and Colorado; the barriers are the low areas named
+on the map (the name "Black Hills" is on the map for another reason;
+these hills are not a barrier). The major mountainous areas higher than
+approximately 8000 feet in elevation in Wyoming, Colorado and Utah are
+stippled. These mountainous areas include the habitat that is most
+suitable for the montane meadow mouse. The Black Hills are unoccupied by
+this species but these hills seem to be ecologically suitable for the
+species.]
+
+     _Measurements._--The average and the extremes for some measurements
+     of 34 males and females, 27 from the type locality and 7 from other
+     localities in the range assigned to this subspecies, are as
+     follows: total length, 165 (146-186); length of tail, 44.2 (35-55);
+     hind foot, 19.6 (17-21); condylobasilar length of the skull, 25.5
+     (24.0-27.5); zygomatic breadth, 15.6 (14.7-16.6); alveolar length
+     of upper molar tooth-row, 6.6 (6.2-7.0); prelambdoidal breadth, 8.8
+     (8.1-9.5); lambdoidal breadth, 12.0 (11.2-12.8). As an indication
+     of variability and for comparison with other series the coefficient
+     of variability and two times the standard error of the mean for
+     each measurement in this series are included in Table 1. The
+     averages for some measurements of 27 topotypes are as follows:
+     total length, 162; length of tail, 45.5; hind foot, 19.9;
+     condylobasilar length, 25.6; palatilar length, 14.0; molar series,
+     6.6; alveolobasilar length, 14.9; zygomatic breadth, 15.6;
+     interorbital breadth, 3.5; lambdoidal breadth, 12.1; prelambdoidal
+     breadth, 8.9; depth of braincase, 7.8.
+
+_Discussion._--Three species of _Microtus_ were collected by James W.
+Bee at the type locality. _Microtus montanus codiensis_, _Microtus
+longicaudus mordax_, and _Microtus pennsylvanicus modestus_ were taken
+in the same runways in the same meadow, at the same time. _Microtus
+ochrogaster haydeni_, although not taken at this locality, occurs in the
+Big Horn Basin. These four species differ in their geographic ranges,
+being largely allopatric, except _M. montanus_ and _M. longicaudus_
+which are sympatric. Although the different species have ecological
+preferences and habits which differ, several species of _Microtus_ may
+occur together in local areas such as the above. Certain of the
+characteristics of _M. m. codiensis_ are intermediate between those of
+the species _M. montanus_ on one hand and those of the other three
+species on the other hand. Could interspecific hybridization between
+"good species" of _Microtus_ take place in nature and possibly alter the
+characteristics of a local population?
+
+     _Specimens examined._--Total, 50, distributed as follows
+     (abbreviations for collections are given in the account of _M. m.
+     nanus_; localities that are not represented in Fig. 1 because
+     overlapping or crowding of the symbols would result are
+     Italicized):
+
+     MONTANA: _Carbon Co._: Beartooth Mountains, 2 (USBS); _Beartooth
+     Lake_, 1 (USBS).
+
+     WYOMING: _Park Co._: Black Mountain, head of Pat O'Hara Creek, 3
+     (USBS); 13 mi. N, 1 mi. E Cody, 5200 ft., 1; SW slope Whirlwind
+     Peak, 9000 ft., 1; _5 mi. N Cody, 6300 ft._, 1 (USBS); 3-1/5 mi. E,
+     3/5 mi. S Cody, 31; Ishawooa Creek, 6300 ft., 2 (USBS); _Valley_, 1
+     (USBS); Needle Mountain, 10,500 ft., 4 (USBS). _Hot Springs Co._: 3
+     mi. N, 10 mi. W Thermopolis, 4950 ft., 3.
+
+
+=Microtus montanus zygomaticus=, new subspecies
+
+     _Type._--Male, adult, skin and skull, No. 32761, Museum of Natural
+     History, University of Kansas, from Medicine Wheel Ranch, 9000 ft.,
+     28 mi. E Lovell, Big Horn County, Wyoming; obtained by R. Freiburg,
+     original number 105.
+
+     _Range._--The Big Horn Mountains of north-central Wyoming.
+
+     _Diagnosis._--A large _Microtus montanus_ with a relatively short
+     tail; short molar series; broad zygomatic arches well rounded in
+     lateral outline when viewed from above; small and flattened bullae;
+     raised basioccipito-basisphenoid suture.
+
+     _Comparisons._--For comparison with _M. m. codiensis_ from the
+     west, on the other side of the Big Horn Basin, see the account of
+     that subspecies. In comparison with _nanus_ this subspecies is
+     slightly paler, in this respect showing more resemblance to
+     _codiensis_ although not so pale, and more grizzled or unevenly
+     colored. This difference in color between _zygomaticus_ and
+     _codiensis_ may not be of taxonomic significance. From both the
+     topotypes of _nanus_, and the series of it from Wyoming,
+     _zygomaticus_ differs on the average in having a relatively shorter
+     tail, a relatively shorter upper molar tooth-row, relatively more
+     rounded and relatively more wide-spread zygomatic arches, and
+     smaller more flattened bullae.
+
+     _Measurements._--Average and extreme measurements of 24 adult males
+     and females from several localities here referred to _M. m.
+     zygomaticus_ are as follows: total length, 159(150-175); length of
+     tail, 37.6(31-46); hind foot, 18.6(17-20); condylobasilar length of
+     the skull, 25.3(24.2-26.7); zygomatic breadth, 15.3(14.1-16.7);
+     alveolar length of upper molar tooth-row, 6.2 (5.7-6.8);
+     prelambdoidal breadth, 8.7(8.3-9.4); lambdoidal breadth,
+     11.9(11.0-12.5). Average and extreme measurements of a series of 12
+     adult male topotypes are as follows: total length, 159(144-174);
+     length of tail, 36.4 (30-41); hind foot, 18.2(16-20);
+     condylobasilar length of skull, 25.8(24.7-26.7); alveolobasilar
+     length, 14.8(13.8-15.3); palatilar length, 13.8 (12.7-14.2);
+     alveolar length of upper molar tooth-row, 6.4(5.9-6.6); zygomatic
+     breadth, 15.9 (15.0-16.7); interorbital breadth, 3.6(3.4-3.7);
+     lambdoidal breadth, 12.1 (11.5-12.5); prelambdoidal breadth,
+     8.6(8.3-8.9); depth of braincase, 8.0 (7.6-8.3).
+
+_Discussion._--This subspecies is separated from _M. m. codiensis_ to
+the west by the Big Horn Basin. A series from along Buffalo Creek, 27
+mi. N, 1 mi. E Powder River, 6075 ft., in Natrona County, Wyoming, is
+intermediate between the topotypes of _zygomaticus_ and _nanus_ in the
+characters cited above as distinguishing the two, but shows greater
+resemblance to _zygomaticus_ in the shape of the zygomatic arch, in
+color which is paler than in topotypes of _zygomaticus_, and in the
+short hind foot. On these and on geographic grounds this population is
+referred to _zygomaticus_. Unfortunately we cannot be certain in many
+cases that an intermediate condition in a certain character indicates a
+genetically intermediate population and therefore true intergradation
+between the two subspecies to which the population is geographically
+intermediate. The topotypes of this subspecies are the most distinct of
+all the series which I have studied from the eastern Rocky Mountains, in
+terms of the degree of morphological departure from the norm for the
+species. After _zygomaticus_ the following populations are arranged
+according to their degree of deviation from this norm (_codiensis_
+deviates most): topotypes of _codiensis_, _fusus_ and a population from
+southern Sweetwater County, Wyoming, and lastly the _nanus-caryi_
+complex. Within the latter group, as I have mentioned, there are a
+number of local variants most of which do not differ significantly and
+do not conform to any geographic pattern.
+
+     _Specimens examined._--Total, 55, distributed as follows
+     (abbreviations for collections are given in the account of _M. m.
+     nanus_; localities that are not represented in Fig. 1 because
+     overlapping or crowding of the symbols would result are
+     Italicized): WYOMING: _Big Horn Co._: Medicine Wheel Ranch, 9000
+     ft., 28 mi. E Lovell, 30; W slope, head of Trappers Creek, 9500
+     ft., 2 (USBS). _Washakie Co._: 9 mi. E, 5 mi. N Tensleep, 7400 ft.,
+     1. _Johnson Co._: 7½ mi. W, 1 mi. S Buffalo, 6500 ft., 3; Big
+     Horn Mountains, 3 (USBS). _Natrona Co._: Buffalo Creek, 27 mi. N, 1
+     mi. E Powder River, 6075 ft., 16.
+
+
+=Microtus montanus fusus= Hall
+
+     _Microtus nanus_, Bailey, N. Amer. Fauna, 17:30, June 6, 1900
+     (part); Cary, N. Amer. Fauna 33:123, August 17, 1911.
+
+     _Microtus montanus fusus_ Hall, Proc. Biol. Soc. Washington,
+     51:131-134, August 23, 1938; Warren, The Mammals of Colorado, Univ.
+     of Okla. Press, p. 229, 1942.
+
+     _Type._--Male, adult, skin and skull; No. 61281, Museum of
+     Vertebrate Zoology; 2½ miles east of summit of Cochetopa Pass,
+     Saguache County, Colorado; Sept. 21, 1933; collected by Annie M.
+     Alexander; original number 2568. Type not seen by me.
+
+     _Range._--Southern Colorado and northern New Mexico.
+
+     _Comparisons._--For comparison with _M. m. nanus_, the subspecies
+     to the northward, see the preceding account of that subspecies. For
+     comparison with _M. m. amosus_ the subspecies to the west see Hall
+     (1938) and Durrant (1952). I have not examined specimens of
+     _amosus_.
+
+     _Measurements._--Average and extreme measurements for 17 adults
+     including both males and females from several localities in
+     southern Colorado are as follows: total length, 160 (136-179);
+     length of tail, 42 (35-55); hind foot, 19.2 (17-23); condylobasilar
+     length of the skull, 25.2 (24.0-26.0); zygomatic breadth, 15.0
+     (14.1-15.5); alveolar length of upper molar tooth-row, 6.4
+     (6.0-6.7); prelambdoidal breadth, 8.7 (8.3-9.2); lambdoidal
+     breadth, 11.7 (11.1-12.6).
+
+     Average and extreme measurements of 4 adults (2 males and 2
+     females) from the type locality and 11 adults (4 males and 7
+     females) from other localities in southern Colorado are as follows:
+     total length, 162 (157-168), 157 (137-169); length of tail (means
+     only), 44.5, 40.5; hind foot, 18.8 (18-19), 18.6 (18-23);
+     condylobasilar length of skull, 24.5 (24.0-24.7), 25.2 (24.3-26.1);
+     alveolobasilar length, 14.2 (13.9-14.5), 14.6 (14.1-15.1);
+     palatilar length, 13.2 (13.0-13.4), 13.5 (13.1-14.2); alveolar
+     length of upper molar tooth-row, 6.3 (6.0-6.6), 6.4 (6.3-6.7);
+     zygomatic breadth, 15.0 (14.3-15.5), 14.9 (14.1-15.5); interorbital
+     breadth, 3.5 (3.3-3.6), 3.5 (3.3-3.7); lambdoidal breadth, 11.8
+     (11.1-12.6), 11.7 (11.2-12.3); prelambdoidal breadth, 8.6
+     (8.3-9.2), 8.8 (8.3-9.0); depth of braincase, 7.5 (7.2-7.8), 7.6
+     (7.1-7.9).
+
+_Discussion._--There is no sharp boundary between _M. m. fusus_ of
+southern Colorado and the subspecies to the north, _M. m. nanus_.
+Although the line separating these two subspecies is drawn somewhat
+arbitrarily, on the whole the samples from north of this line more
+closely resemble _nanus_. All of the means for total length given above
+are larger than the maximum given in Hall's description of _fusus_. The
+caudal index (38 and 35% in two series) is slightly larger than that
+cited by Hall (33.3%) and is not significantly different from that in
+_nanus_ (35.2%). The color in both young and old mice is variable, but
+in general is more yellowish, and less grayish, than in any other series
+studied.
+
+There is a large area in western Colorado and eastern Utah, between the
+known ranges of _M. m. fusus_ and _M. m. amosus_ from which there are no
+specimens. Probably the species occurs only at certain places in this
+arid region which seems to be a partial barrier to the species.
+
+Specimens of _M. montanus_ from northern New Mexico have been referred
+previously to _M. m. arizonensis_. When he named _M. m. fusus_, Hall
+mentioned its resemblance to _arizonensis_ in reddish coloration, but
+pointed out that _fusus_ is less reddish. Of six specimens from Valle
+Santa Rosa, Jemez Mountains (USBS), 8500 ft., Rio Arriba County, New
+Mexico, three are immature, and the skulls of the remaining specimens
+are damaged. In reddish color and relatively large size these few
+specimens resemble _arizonensis_ more than _fusus_ although the locality
+of occurrence is closer to the geographic range of the northern _fusus_
+than to that of _arizonensis_. The identification of these specimens as
+_arizonensis_ is provisional; additional specimens are needed from the
+area, 200 miles wide, which separates the ranges as now known of
+_arizonensis_ in Arizona from the occurrence in New Mexico. There is a
+single specimen from this area, the damaged skull of which prevents
+conclusive identification. The specimen is either _M. montanus_ or _M.
+mexicanus_, and is from Nutria, on the southern edge of the Zuni
+Mountains (USBS). Detailed comparison of _fusus_ and _arizonensis_ is
+not attempted here although it may be stated that in several characters
+_fusus_ is intermediate between _arizonensis_ to the south and _nanus_
+to the north.
+
+     _Specimens examined._--Total, 89, distributed as follows
+     (abbreviations for collections are given in the account of _M. m.
+     nanus_; localities that are not represented in Fig. 1 because
+     overlapping or crowding of the symbols would result are
+     Italicized):
+
+     COLORADO: _Pitkin Co._: 5 mi. W Independence Pass, 11,000 ft., 1
+     (Chi). _Lake Co._: _Independence Pass, 12,095 ft._, 2 (Chi).
+     _Gunnison Co._: _Gothic_, 2 (USBS); Decker's Ranch, Crested Butte,
+     2 (AMNH); Almont, 3 (USBS). _Montrose Co._: Coventry, 5 (USBS 4,
+     AMNH 1). _Saguache Co._: Cochetopa Pass and environs, 44 (USBS
+     22). _Hinsdale Co._: Ruby Lake, 1 (USBS). _Mineral Co._: 3 mi. E
+     Creede, 1; 23 mi. S, 11 mi. E Creede, 9300 ft., 7. _La Plata Co._:
+     Florida, 6800 ft., 1. _Conejos Co._: 1 mi. S, 19 mi. W Antonito,
+     10,200 ft., 3; _4 mi. S, 23 mi. W Antonito_, 1; _5 mi. S, 24 mi. W
+     Antonito, 9600 ft._, 9.
+
+     NEW MEXICO: _Rio Arriba Co._: 6 mi. W Hopewell, 9900 ft., 6 (USBS);
+     _Tusas River, 8700 ft._, 1 (USBS).
+
+TABLE 1. AVERAGE MEASUREMENTS, IN MILLIMETERS, OF ADULTS OF MICROTUS
+MONTANUS.
+
+Key to column headings:
+
+A: No. of individuals averaged
+B: Total length
+C: Length of tail
+D: Length of hind foot
+E: Condylobasilar length
+F: Alveolar length of upper molar tooth-row
+G: Zygomatic breadth
+H: Lambdoidal breadth
+I: Prelambdoidal breadth
+
+=========================================================================
+       Locality   |   A  |  B  |  C  |  D  |  E  |  F  |  G  |  H  |  I
+------------------+------+-----+-----+-----+-----+-----+-----+-----+-----
+                  |               _M. m. codiensis_, all
+                  |
+Average           | 34   |165.3|44.2 |19.6 |25.47| 6.56|15.55|12.05|8.76
+2 × stand. error  |      | 3.56|1.84 |.395 | .308| .067| .198| .144|.129
+Coeff. variab     |      |  6.0|11.6 | 5.6 |  3.5|  3.0| 3.65| 1.20|1.47
+------------------+------+-----+-----+-----+-----+-----+-----+-----+-----
+                  |            _M. m. nanus_, Eastern Idaho
+                  |
+Average           | 21[1]|151.1|39.4 |19.2 |25.00| 6.44|14.99|11.74|8.94
+2 × stand. error  |      | 3.20|2.89 |.293 | .286| 1.15| .295| .210| .182
+Coeff. variab.    |      |  6.1|21.1 |4.38 | 2.49| 3.99| 4.10| 3.79|4.31
+------------------+------+-----+-----+-----+-----+-----+-----+-----+-----
+                  |            _M. m. nanus_, Wyoming
+                  |
+Teton Co.         | 35   |160.5|40.7 |18.6 |25.16| 6.51|15.17|11.86|8.77
+Fremont Co.       | 26   |157.0|41.4 |19.6 |25.23| 6.25|15.05|11.88|8.91
+Lincoln Co.       | 24   |159.9|41.8 |18.9 |25.08| 6.26|15.10|11.82|8.75
+Uinta Co.         | 26   |162.4|41.3 |19.0 |25.33| 6.42|15.31|12.16|8.89
+Sweetwater Co.    | 12   |159.8|43.7 |20.1 |24.98| 6.31|15.00|11.84|9.02
+Natrona Co.       | 40   |159.6|41.0 |19.6 |25.04| 6.40|15.00|11.84|8.93
+Carbon Co.        |108   |158.7|40.0 |19.1 |24.96| 6.27|15.05|11.83|8.72
+ Encampment [M]   | 27   |161  |41.7 |18.9 |25.1 | 6.16|15.2 |11.9 |8.7
+ Encampment [F]   | 11   |159  |41.1 |19.4 |24.9 | 6.18|14.9 |11.6 |8.6
+ Savery [M]       | 23   |159  |41.0 |19.2 |25.3 | 6.32|15.2 |12.2 |8.8
+ Savery [F]       | 25   |155  |37.1 |18.8 |24.7 | 6.33|14.8 |11.6 |8.6
+------------------+------+-----+-----+-----+-----+-----+-----+-----+-----
+                  |                     _M. m. nanus_
+                  |
+Northern Colo.    |  8   |163.1|42.4 |19.6 |25.20| 6.44|14.86|11.70|8.56
+------------------+------+-----+-----+-----+-----+-----+-----+-----+-----
+                  |                     _M. m. fusus_
+                  |
+Southern Colo.    | 17[2]|159.8|42.4 |19.2 |24.97| 6.43|14.98|11.73|8.69
+------------------+------+-----+-----+-----+-----+-----+-----+-----+-----
+
+[Footnote 1: For external parts, 34 individuals were used.]
+
+[Footnote 2: For external parts, 29 individuals were used.]
+
+Some measurements not given above are included in Table 1, together with
+the number of specimens and the sex if restricted to one sex. So that
+the variability can be evaluated more adequately, the coefficient of
+variability and 2 times the standard error of the mean are included for
+the measurements in two series. The series consist of all the adult
+specimens (with a condylobasilar length of 24.0 mm. or more) of both
+sexes from the areas specified. Various barriers are shown in Fig. 2 for
+comparison with the distributions of the subspecies and the localities
+of known occurrence shown in Fig. 1. _Microtus montanus_ has not been
+taken in the Black Hills area of extreme northeastern Wyoming. Suitable
+montane habitat is present and both _Microtus pennsylvanicus insperatus_
+and _Microtus longicaudus longicaudus_ occur there. The arid basin of
+the Powder River presumably is a barrier that has prevented _M.
+montanus_ from reaching this area.
+
+
+REFERENCES CITED
+
+BAILEY, V.
+
+     1900. Revision of American voles of the genus _Microtus_. N. Amer.
+     Fauna, 17:1-88, June 6.
+
+     1917. A new subspecies of meadow mouse from Wyoming. Proc. Biol.
+     Soc. Washington, 30:29-30, February 21.
+
+CARY, M.
+
+     1911. A biological survey of Colorado. N. Amer. Fauna, 33:1-256,
+     August 17.
+
+     1917. Life zone investigations in Wyoming. N. Amer. Fauna, 42:1-95,
+     October 3.
+
+DAVIS, W. B.
+
+     1939. The Recent mammals of Idaho. 400 p., front., illus., maps,
+     diagrs., Caldwell, Id., The Caxton Printers Ltd., April 5.
+
+DURRANT, S. D.
+
+     1952. Mammals of Utah, taxonomy and distribution. Univ. Kans.
+     Publ., Mus. Nat. Hist., 6:1-549, 91 figs, in text, 30 tables,
+     August 10.
+
+ELTON, C.
+
+     1942. Voles, mice and lemmings; problems in population dynamics.
+     496 p., illus. (maps), tables, Oxford, The Clarendon Press, London.
+
+GOIN, O. B.
+
+     1943. A study of individual variation in _Microtus pennsylvanicus
+     pennsylvanicus_. Jour. Mamm., 24:212-224, June 7.
+
+HALL, E. R.
+
+     1938. Notes on the meadow mice _Microtus montanus_ and _Microtus
+     nanus_ with descriptions of a new subspecies from Colorado. Proc.
+     Biol. Soc. Washington, 51:131-134, August 23.
+
+HILL, E. A.
+
+     1935. Cranial foramina in rodents. Jour. Mamm. 16(2):121-129.
+
+HOWELL, A. B.
+
+     1924. Individual and age variation in _Microtus montanus yosemite_.
+     Jour. Agr. Res., 28(10):977-1015, June 7.
+
+KELLOGG, R.
+
+     1922. A study of the California forms of the _Microtus montanus_
+     group of meadow mice. Univ. California Publ. Zool., 21:245-274,
+     April 18.
+
+MERRIAM, C. H.
+
+     1891. Results of a biological reconnaissance of south-central
+     Idaho. N. Amer. Fauna, 5:1-113, July 30.
+
+PIPER, S. E.
+
+     1909. The Nevada mouse plague of 1907-8. U. S. Dept. Agr., Farmers'
+     Bul. 352, pp. 1-23, 9 figs.
+
+WARREN, E. R.
+
+     1942. The mammals of Colorado, their habits and distribution.
+     Second (revised) edition, Univ. Oklahoma Press, Norman, xviii-330
+     p., front., 50 plates.
+
+
+_Transmitted March 22, 1954._
+
+
+
+[SQUARE]
+
+25-3560
+
+
+       *       *       *       *       *
+
+Transcriber's Notes
+
+Italic typeface in the original is indicated by by _underscores_.
+
+Bold typeface in the original is indicated by =equals=.
+
+The symbols for [Male] and [Female] are indicated by [M] and [F].
+
+One typographical error was corrected: "Castlford" was corrected to
+"Castleford" in "Twin Falls Co.: Castleford Fenced Plot".
+
+
+
+
+
+End of the Project Gutenberg EBook of Subspeciation in the Meadow Mouse,
+Microtus montanus, in Wyoming and Colorado, by Sydney Anderson
+
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+Microtus montanus, in Wyoming
+and Colorado, by Sydney Anderson.
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+<pre>
+
+The Project Gutenberg EBook of Subspeciation in the Meadow Mouse, Microtus
+montanus, in Wyoming and Colorado, by Sydney Anderson
+
+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: Subspeciation in the Meadow Mouse, Microtus montanus, in Wyoming and Colorado
+
+Author: Sydney Anderson
+
+Release Date: March 22, 2010 [EBook #31730]
+
+Language: English
+
+Character set encoding: UTF-8
+
+*** START OF THIS PROJECT GUTENBERG EBOOK MEADOW MOUSE ***
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+Produced by Chris Curnow, Simon Gardner, Joseph Cooper and
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+
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+</pre>
+
+
+
+
+
+<p class="center"><span class="smcap">University of Kansas Publications</span></p>
+
+<p class="center"><span class="smcap">Museum of Natural History</span></p>
+
+<p class="center">Volume 7, No. 7, pp. 489-506, 2 figures in text</p>
+
+<p class="center">July 23, 1954</p>
+
+
+
+
+<h1>Subspeciation in the Meadow Mouse,
+Microtus montanus, in Wyoming
+and Colorado</h1>
+
+<h3>BY</h3>
+
+<h2>SYDNEY ANDERSON</h2>
+
+
+<p class="center"><span class="smcap">University of Kansas</span>
+<br />
+<span class="smcap">Lawrence</span>
+<br />
+1954</p>
+
+
+<p class="center"><span class="smcap">University of Kansas Publications, Museum of Natural History</span></p>
+
+<p class="center">Editors: E. Raymond Hall, Chairman, A. Byron Leonard,
+Robert W. Wilson</p>
+
+<p class="center">Volume 7, No. 7, pp. 489-506, 2 figures in text
+<br />
+Published July 23, 1954</p>
+<hr style="width: 45%;" />
+
+<p class="center"><span class="smcap">University of Kansas</span>
+<br />
+Lawrence, Kansas</p>
+
+
+<p class="center">PRINTED BY
+<br />
+FERD VOILAND, JR., STATE PRINTER
+<br />
+TOPEKA, KANSAS
+<br />
+1954</p>
+
+<p class="center">25-3560
+</p>
+<hr style="width: 45%;" />
+
+
+<h1>Subspeciation in the Meadow Mouse,
+Microtus montanus, in Wyoming
+and Colorado</h1>
+
+<h3>BY</h3>
+
+<h2>SYDNEY ANDERSON</h2>
+
+
+<p><i>Microtus montanus</i> reaches the eastern limits of its geographic
+distribution in Wyoming and Colorado. There the mountains, but
+in general not the lowlands, are occupied by this species. A certain
+minimum of moisture may be of direct importance to the mouse and
+certainly is indirectly important, because certain hydrophytic or
+mesophytic grasses used by the mouse for food, for protection from
+enemies, and for shelter from the elements are dependent on the
+moisture. Areas suitable for <i>Microtus montanus</i> are separated by
+deserts that are dominated by sagebrush and other xerophytic plants
+or by forests or rocky exposures at higher altitudes. A relatively
+small percentage, probably less than ten per cent, of the total area
+even in the more favorable parts of the range of the species is suitable
+for occupancy. In these mice, as in other microtines (Elton,
+1942; Piper, 1909), there are seasonal, and irregularly multiannual
+fluctuations in population density, which sometimes are extreme.
+Consequently the mice at some times seem to be absent from suitable
+habitats, and at some other times occur there in amazingly
+large numbers.</p>
+
+<p>Because the species is broken up into partly isolated, or at times
+completely isolated, colonies or local populations it may be supposed
+that various evolutionary forces such as selection and random genetic
+drift operate to foster variation. The degree to which racial distinction
+is attained may depend upon these forces and the time available.
+In <i>Microtus montanus</i> in the eastern Rocky Mountains the
+degree of subspecific distinction is not great.</p>
+
+<p>The study here reported upon is based on 1,187 specimens of
+<i>Microtus montanus</i> from Wyoming, Colorado, Idaho and Montana,
+and on work in the field. I spent approximately four months in the
+field in this area, in the summers of 1950, 1951, and 1952. The specimens
+studied were arranged according to localities and the larger
+series were compared statistically. Each of two series, totaling 136
+specimens, was studied intensively to ascertain the kind and range
+of variation within single populations. Twenty-seven measurements,
+various proportions based on these measurements, and differences
+in color were analyzed. Fifteen characters, judged to be most significant,
+were selected for use in comparing all series. In addition,
+certain characters that can not be expressed easily by measurements,
+such as inflation of the auditory bullae and the curvature of the
+zygomatic arch, were observed. The studies by A. B. Howell
+(1924) of variation in <i>Microtus montanus yosemite</i> Grinnell in California
+and those by O. B. Goin (1943) of <i>Microtus pennsylvanicus
+pennsylvanicus</i> (Ord) were useful. The external measurements
+are from the collectors' field labels. The measurements of the skull
+all were taken with dial calipers reading to a tenth of a millimeter.
+The anteroposterior measurements of the skull all were taken along
+the shortest line between the points specified below and are not
+necessarily along a line parallel to the long axis of the skull. These
+measurements were taken on the left side of the skull whenever
+possible. Some of the skulls are damaged and therefore some
+measurements could not be taken and are omitted in the computations.
+Measurements are in millimeters.</p>
+
+<p>The results of these studies were submitted to the Department of
+Zoology and the Graduate School of the University of Kansas in
+partial fulfillment of the requirements for the degree of Master of
+Arts (1952) and are available in manuscript form at the Museum of
+Natural History and the library of the University of Kansas.</p>
+
+
+<h3><span class="smcap">Explanation of Measurements</span></h3>
+
+<blockquote><p>Caudal index.&mdash;the length of the tail expressed as a percentage of the length
+of the head and body. The length of the head and body is the collector's
+measurement of total length less the length of the tail.</p></blockquote>
+
+<h4><span class="smcap">Cranial Measurements of Length.</span></h4>
+
+<blockquote><p>Condylobasilar length.&mdash;from the exoccipital condyle to the most posterior
+point on the border of the alveolus of the upper incisor.</p>
+
+<p>Alveolobasilar length.&mdash;from the posterior border of the alveolus of the third
+upper molar to the posterior border of the alveolus of the incisor.</p>
+
+<p>Palatilar length.&mdash;from the anteriormost part of the posterior border of the
+bony shelf of the palate to the posteriormost part of the alveolus of the
+incisor.</p>
+
+<p>Alveolar length of upper molar tooth-row.&mdash;from the most posterior point of
+the alveolus of the third upper molar to the most anterior point of the
+alveolus of the first upper molar.</p></blockquote>
+
+<h4><span class="smcap">Measurements of Breadth.</span></h4>
+
+<blockquote><p>Zygomatic breadth.&mdash;greatest transverse width.</p>
+
+<p>Interorbital breadth.&mdash;the breadth of the interorbital constriction.</p>
+
+<p>Lambdoidal breadth.&mdash;between the lateralmost points of the lambdoidal crest.</p>
+
+<p>Prelambdoidal breadth.&mdash;between the medialmost margins of the prominent
+fenestrae in the posterodorsal parts of the squamosal bones. To these
+fenestrae Howell (1924:995) applied the adjective "prelambdoidal," but other
+authors have used other names (see Hill, 1935:127).</p>
+
+<p>Depth of braincase.&mdash;shortest distance from the ventral side of the cranium at
+the suture between the basioccipital and basisphenoid bones to the dorsal
+surface of the cranium (usually not perpendicular to the long axis of the
+skull).</p></blockquote>
+
+<p>The history of our knowledge of <i>Microtus montanus</i> in this area
+begins with the early work of the United States Bureau of Biological
+Survey directed by C. H. Merriam (1891), and participated in by
+Vernon Bailey (1900, 1917), Merritt Cary (1911, 1917), and others.
+The changes in nomenclature which grew out of increased understanding
+of these mice through additional collecting are expressed
+in the synonymies under the accounts of subspecies. As a result
+of my studies two of the three subspecific names previously proposed
+for mice from this area have been retained although changes are
+proposed in the ranges assigned to the two subspecies and two additional
+heretofore unrecognized subspecies are named and described.
+Furthermore the additional specimens and my studies of
+variation make modifications in the characterization of these subspecies
+necessary. Not all of the samples here assigned to a single
+subspecies are identical and I therefore list and discuss some of the
+local variants.</p>
+
+<blockquote><p>Numerous members of summer field parties from the Museum of Natural
+History at the University of Kansas collected most of the specimens studied
+and wrote field notes that have been helpful. I am grateful to these persons
+and to Professor E. R. Hall and Dr. R. H. Baker for their assistance and helpful
+suggestions. Specimens in the following museums were made available by
+their respective curators: Chicago Natural History Museum by Mr. Colin C.
+Sandborn, The Museum of Zoology at the University of Michigan by Dr. E. T.
+Hooper, The American Museum of Natural History by Mr. G. G. Goodwin,
+The United States National Museum by Dr. David H. Johnson and the Biological
+Surveys Collection by Miss Viola S. Schantz. A fellowship from the
+National Science Foundation made possible the studies at the museums other
+than at the University of Kansas.</p></blockquote>
+
+
+
+<p class="center"><b>Microtus montanus nanus</b> (Merriam)</p>
+
+<blockquote><p><i>Arvicola (Mynomes) nanus</i> Merriam, N. Amer. Fauna, 5:62, July 30, 1891.</p>
+
+<p><i>Microtus montanus nanus</i>, Hall, Proc. Biol. Soc. Washington, 51:131, August
+23, 1938.</p>
+
+<p><i>Microtus nanus</i>, Bailey, N. Amer. Fauna, 17:30, June 6, 1900 (part).</p>
+
+<p><i>Microtus montanus caryi</i> Bailey, Proc. Biol. Soc. Washington, 30:29, February
+21, 1917.</p>
+
+<p><i>Microtus nanus nanus</i>, Cary, N. Amer. Fauna, 42:43, October 3, 1917 (part).</p></blockquote>
+
+<blockquote><p><i>Type.</i>&mdash;Adult male No. 23853/31253, U. S. National Museum, Department of
+Agriculture collection, from Pahsimeroi Mountains, Custer County, Idaho; obtained
+by C. Hart Merriam and Vernon Bailey, September 16, 1890.</p>
+
+<p><i>Range.</i>&mdash;Idaho; southwestern Montana; most of the southwestern half of
+Wyoming; southward to central Colorado. See <a href="#fig_1">figure 1</a>.</p>
+
+<p><i>Comparisons.</i>&mdash;Comparisons with subspecies named as new in this paper
+will be found in the accounts of those subspecies beyond. From <i>Microtus
+montanus fusus</i> Hall, the subspecies to the south, <i>M. m. nanus</i> from Idaho
+differs as follows: averages smaller; slightly darker and less reddish and less
+yellowish in color; slightly wider braincase (see measurement of prelambdoidal
+breadth); larger bullae.</p></blockquote>
+
+<blockquote><p><i>Measurements.</i>&mdash;Average (= arithmetical mean) measurements of 34 specimens,
+both male and female, from several localities in eastern Idaho are: total
+length, 151; length of tail, 39; hind foot, 19.2; condylobasilar length of the
+skull, 25.0; zygomatic breadth, 15.0; alveolar length of upper molar tooth-row,
+6.4; prelambdoidal breadth, 8.9; and lambdoidal breadth, 11.7.</p>
+
+<p>Average and extreme measurements of six adult males from near Pocatello,
+Bannock County, Idaho, and nine adult males from near Afton, Lincoln County,
+Wyoming, are, respectively, as follows: total length, 143(135-150), 163(143-179);
+length of tail, 35.1(33-38), 42.8(36-49); caudal index, 32.0(28.0-33.1),
+35.7(30.6-41.9); hind foot, 18.9(18-20), 18.8(17-20); condylobasilar length
+of skull, 24.4(24.0-26.0), 25.6(24.5-26.2); alveolobasilar length, 14.1(13.7-14.5),
+14.6(13.8-15.0); palatilar length, 13.2(12.9-13.6), 13.8(13.2-14.5);
+alveolar length of upper molar tooth-row, 6.3(6.1-6.5), 6.3(6.0-6.6); depth
+of braincase, 7.7(7.5-7.9), 8.0(7.7-8.3); lambdoidal breadth, 11.4(11.0-11.7),
+12.0(11.3-12.7); prelambdoidal breadth, 9.1(8.6-9.4), 8.7(8.0-9.4); zygomatic
+breadth, 14.3(13.8-14.7), 15.3(14.4-16.3); interorbital breadth, 3.6(3.5-3.7),
+3.5(3.3-3.7). The average length of the nasal bones in the series from Pocatello
+is 7.1 mm. The averages, which have not been included in <a href="#table_1">Table 1</a>, for
+three measurements of the series from Carbon County, Wyoming, are as follows
+(Encampment, males; Encampment, females; Savery, males; and Savery, females,
+respectively): alveolobasilar length, 14.4, 14.3, 14.5, 14.3; interorbital
+breadth, 3.5, 3.4, 3.5, 3.4; depth of braincase, 7.8, 7.6, 7.9, 7.6. Additional
+measurements are included in<a href="#table_1"> Table 1</a> for other series.</p></blockquote>
+
+<p><i>Discussion.</i>&mdash;The name <i>Microtus montanus caryi</i> Bailey is here
+placed in synonymy under <i>M. m. nanus</i> (Merriam). Vernon Bailey
+(1917) in his description of <i>caryi</i> made four assumptions that have
+been found to be entirely or partly invalid. First, he assumed that
+this is an "extreme variant which gradually changes in characters
+across Nevada and Utah, and reaches its maximum variation in
+Wyoming." The differences pointed out in subsequent descriptions
+of subspecies found in the above area do not show a gradual change
+in any character, or in the number of characters, nor is <i>caryi</i> an
+extreme when compared with the other subspecies. Second,
+<i>Microtus nanus</i> was not, as Bailey assumed, a different species than
+<i>Microtus montanus</i>. Third, he assumed that the characteristics of
+adults of <i>nanus</i> were adequately ascertainable from the thirteen
+topotypes available to him. Subsequent sampling from Idaho shows
+that the series of specimens available to Bailey was made up mostly
+of young and subadult animals. Finally, <i>caryi</i> does not occupy as
+Bailey stated "the meadows along streams in the arid sagebrush
+country of the Bear River, Green River, and Wind River valleys"
+exclusively, or characteristically. When the localities from which
+the species actually is known are plotted, it seems that the arid
+basin serves as a barrier and that the species is more commonly and
+abundantly found in montane meadows in the Transition and
+Canadian life-zones.</p>
+
+<p>Certain samples, here assigned to <i>M. m. nanus</i>, that vary from the
+average of the subspecies deserve comment. For example, mice
+from the area in Wyoming southwest of the Green River (in the
+Uinta Mountains) have relatively smaller feet, but are larger in both
+total length and size of skull. Specimens from near Afton, Lincoln
+County, Wyoming, are relatively large in both total length and size
+of skull. This series and specimens from Teton County, Wyoming,
+are intermediate between <i>nanus</i> from Idaho and the newly named
+subspecies from near Cody, Park County, Wyoming, described
+below, in terms of both darkness and the amount of reddish color.
+Mice from Laramie County are more richly reddish-brown. The
+specimens from near Savery, in Carbon County, Wyoming, are
+darker. The alveolobasilar length relative to the condylobasilar
+length is smaller in the series from along Deer Creek, 16 mi. S, 11
+mi. W Waltman, Natrona County, Wyoming. The series from the
+southern tier of counties in Wyoming and some of the specimens
+from Colorado have relatively wider zygomatic arches. The specimens
+from southern Sweetwater County, Wyoming, are relatively
+paler, have a relatively longer tail and longer hindfeet, lesser
+condylobasilar length, and wider braincase. Most of these variations
+are of questionable significance; they may be chance variations
+owing to errors in sampling.</p>
+
+<p>Much of the south-central part of the state is relatively low and
+relatively arid. This area includes the arid basin of the Green
+River and its major tributaries and the arid Red Desert along the
+continental divide in Sweetwater County. This area might have
+acted as a barrier to the mice; gene flow might have been prevented
+between the populations of the western part of the state and those
+farther east in the Medicine Bow Mountains and Laramie Mountains.
+Nevertheless geographic variations of subspecific worth have
+not taken place. The barrier has either not been of as long duration,
+or has not been so complete and effective, as the other barriers
+in the state, namely the Absaroka Range, the Big Horn Basin, the
+Shoshone Basin, and the valley of the North Platte River. These
+four barriers presumably have led to the differentiation of the two
+subspecies that are newly named beyond. Each of the two areas
+which is set apart by these barriers and in which one of the newly
+named subspecies has evolved is small; therefore there is a lesser
+amount of suitable habitat available for each of the newly named
+mice than there is for <i>M. m. nanus</i>. It is conceivable, therefore,
+that in periods of adverse conditions in each of the small areas the
+size of the effective breeding population may have been so small
+that random genetic drift could have operated effectively, or that
+selection was more critical than in a larger, more stable population.
+It is difficult to test these possibilities because the selective value of
+the taxonomic characters is unknown. The observed pattern of
+variation and facts of distribution are, however, not contradictory
+to the above possibilities.</p>
+
+<blockquote><p><i>Specimens examined.</i>&mdash;Total, 993, distributed as follows: All specimens
+unless otherwise indicated are in the University of Kansas Museum of Natural
+History. Specimens in other museums are labeled as follows: Chicago Natural
+History Museum (Chi); University of Michigan, Museum of Zoology (Mich);
+American Museum of Natural History (AMNH); United States National Museum
+(USNM); Biological Surveys Collection (USBS). Localities that are not
+represented in <a href="#fig_1">Fig. 1</a> because overlapping or crowding of the symbols would
+result are Italicized. Localities are arranged from north to south by states,
+within a state from northwest to southeast by counties, and within a county from
+north to south.</p>
+
+<p><span class="smcap">Wyoming</span>: <i>Yellowstone Park</i>: Canyon Camp, 1 (USBS); Lower Geyser
+Basin, 1 (USBS); Upper Yellowstone River, 2 (AMNH); <i>North end of Lake</i>,
+<i>Yellowstone National Park</i>, 2 (AMNH). <i>Teton Co.</i>: Pacific Creek, 1 (USBS);
+<i>Big Game Ridge</i>, 3 (USBS 1, Mich 2); <i>Whetstone Creek</i>, 7 (Mich); Moran
+and environs (6 localities within a 5 mile radius), 28 (USBS 2, Mich 5);
+<i>S fork Buffalo River</i>, 7 (AMNH); 2 mi. W pass, Black Rock Creek, 1 (USBS);
+<i>Jenny Lake</i>, 5 (Mich); Bar BC Ranch, 2½ mi. NE Moose, 6500 ft., 2; Teton
+Pass above Fish Creek, 1 (USBS); Jackson and environs, 142 (Mich 141);
+<i>Sheep Creek</i>, 2 (Mich). <i>Lincoln Co.: 13 mi. N, 2 mi. W Afton</i>, 2; <i>10 mi. N,
+2 mi. W Afton</i>, 4; <i>9½ mi. N, 2 mi. W Afton</i>, 3; 9 mi. N, 2 mi. W Afton, 9;
+<i>7 mi. N, 1 mi. W Afton</i>, 12; Afton, 1 (USBS); Labarge Creek, 1 (USBS);
+Border, 6 (USBS); <i>Cokeville</i>, 2 (USBS); 6 mi. N, 2 mi. E Sage, 1; Cumberland,
+5 (USBS). <i>Sublette Co.</i>: 34 mi. N, 4 mi. W Pinedale, 1; <i>33 mi. N, 2
+mi. W Pinedale</i>, 6; <i>32 mi. N, 1 mi. W Pinedale</i>, 1; <i>31 mi. N Pinedale</i>, 4; Dell
+Creek, on Ferris Ranch, 7 (Mich); Horse Creek, 7800 ft., Merna, 4 (USBS);
+Big Piney, 1 (USBS). <i>Fremont Co.</i>: 17½ mi. W, 2½ mi. N Lander, 9500 ft.,
+3; <i>17 mi. W, 2 mi. N Lander, 9300 ft.</i>, 4; Milford and environs (5 localities
+within a 1 mile radius), 23 (USBS 4); 15½ mi. S, 7½ mi. W Lander, 9200 ft.,
+1; South Pass City, 8000 ft, 8 (USBS); <i>23½ mi. S, 5 mi. W Lander, 8600 ft.</i>,
+7. <i>Natrona Co.</i>: Deer Creek, 16 mi. S, 11 mi. W Waltman, 6950 ft., 44;
+6 mi. S, 2 mi. W Casper, 5900 ft., 4; <i>6-4/5 mi. S, 2 mi. W Casper, 6100 ft.</i>, 1;
+<i>7 mi. S, 2 mi. W Casper, 6370 ft.</i>, 3; <i>10 mi. S Casper, 7750 ft.</i>, 33; Sun, 2
+(USBS); <i>5 mi. W Independence Rock, 6000 ft.</i>, 4; <i>5 mi. W, 1 mi. S Independence
+Rock</i>, 2. <i>Converse Co.</i>: Beaver, 1 (USBS). <i>Uinta Co.: 1½ mi. W,
+½ mi. S Cumberland</i>, 6; <i>16 mi. S, 2 mi. W Kemmerer, 6700 ft.</i>, 3; 10 mi. SW
+Granger, 3 (Mich); Fort Bridger, 6650 ft., 25 (USNM 6); 9 mi. S Robertson,
+8000 ft., 9; <i>9½ mi. S, ½ mi. W Robertson, 8600 ft.</i>, 1; <i>10 mi. S, 1 mi. W Robertson,
+8700 ft.</i>, 25; <i>14 mi. S, 2 mi. E Robertson, 9000 ft.</i>, 5; 4 mi. S Lonetree, 1
+(USBS). <i>Sweetwater Co.</i>: Farson, 3; Bitter Creek, 3 (AMNH); Kinney
+Ranch, 21 mi. S Bitter Creek, 6800 ft., 9 (USNM 1, AMNH 2); 32 mi. S, 22
+mi. E Rock Springs, 7025 ft., on Vermillion Creek, 15. <i>Carbon Co.</i>: 18 mi.
+NNE Sinclair, 6500 ft., 10; Bridgers Pass, 18 mi. SW Rawlins, 7500 ft., 7;
+Saratoga, 1 (USBS); <i>6 mi. S, 13 mi. E Saratoga, 8500 ft.</i>, 5; <i>6 mi. S, 14 mi.
+E Saratoga, 8800 ft.</i>, 1; Lake Marie, 10,440 ft., 2; <i>1 mi. S Lake Marie</i>, 2; <i>½
+mi. S, 2 mi. E Medicine Bow Peak, 10,800 ft.</i>, 1; Encampment (12 localities
+from 10 mi. N, 14 mi. E to 9 mi. N, 3 mi. E Encampment and from 6500 to
+8400 ft.), 63; 1/4 mi. N Riverside, 7380 ft., 2; S base Bridger Peak, 8800 ft.,
+Sierra Madre Mountains, 1; <i>2 mi. S Bridger Peak, 9300 ft.</i>, 2; Savery (10
+localities from 8 mi. N, 21 mi. E to 4 mi. N, 8 mi. E Savery and from 7300 to
+8800 ft.), 80. <i>Albany Co.</i>: <i>30 mi. N, 10 mi. E Laramie, 6760 ft.</i>, 6; <i>29¾ mi.
+N, 9½ mi. E Laramie, 6350 ft.</i>, 1; 26 mi. N, 4½ mi. E Laramie, 6960 ft., 8;
+<i>26¾ mi. N, 6½ mi. E Laramie, 6700 ft.</i>, 3; <i>3 mi. N, 13 mi. E Laramie, 7500
+ft.</i>, 1; <i>7 mi. N, 2 mi. E Laramie</i>, 1 (Chi); 5 mi. N Laramie, 7400 ft., 15;
+<i>Laramie</i>, 4 (AMNH); <i>1 mi. E Laramie, 7160 ft.</i>, 4; 7-7/10 mi. SSW Laramie,
+7200 ft., 4; 6½ mi. S, 8¾ mi. E Laramie, 8200 ft., 1; <i>Headquarters Park, 10,200
+ft., Medicine Bow Mountains</i>, 3 (USBS); Centennial, 8120 ft., 1; <i>2-1/4 mi. ESE
+Brown's Peak, 10,300 ft.</i>, 3; <i>3 mi. ESE Brown's Peak, 10,000 ft.</i>, 12; <i>2 mi. S
+Brown's Peak, 10,600 ft.</i>, 1; <i>Pole Mountain, 15 mi. SE Laramie</i>, 4 (USBS 3);
+<i>1 mi. SSE Pole Mountain, 8350 ft.</i>, 4; <i>2 mi. SW Pole Mountain, 8300 ft.</i>, 13;
+<i>3 mi. S Pole Mountain, 8100 ft.</i>, 1; Sherman, 2 (AMNH). <i>Laramie Co.</i>: 5
+mi. N, 1 mi. W Horse Creek P. O., 7200 ft., 1; Meadow, 2 (USBS); 11 mi.
+N, 5½ mi. E Cheyenne, 5450 ft., 7; <i>7 mi. W Cheyenne, 6500 ft.</i>, 10; Cheyenne,
+3 (USNM).</p>
+
+<p><span class="smcap">Colorado</span>: <i>Moffat Co.</i>: Lay, 6160 ft., 1 (AMNH). <i>Routt Co.</i>: Wright's
+Ranch, Yampa, 7700 ft., 2; Gore Range, 8 mi. E Toponas, 8000 ft., 2 (USBS).
+<i>Larimer Co.</i>: <i>12½ mi. W, 1½ mi. S Rustic</i>, 1; 11 mi. W, 1 mi. S Rustic, 1;
+Cache La Poudre River, 1 (Chi); <i>Estes Park</i>, 3 (USBS 1, AMNH 2); 19½ mi.
+W, 2½ mi. S Loveland, 7280 ft., 6; <i>16 mi. W Loveland, 6840 ft.</i>, 1; 6 mi. W,
+½ mi. S Loveland, 5200 ft., 1. <i>Rio Blanco Co.</i>: Meeker, 1 (USBS); <i>9½ mi.
+SW Pagoda Peak</i>, 7700 ft., 3; 5 mi. S Pagoda Peak, 9100 ft., 2. <i>Eagle Co.</i>:
+Eagle, 1 (USBS); Pando, 2 (USBS). <i>Grand Co.</i>: Mt. Whiteley, 2 (USBS);
+Arapahoe Pass, Rabbit Ear Mountains, 2 (USBS); Coulter (near Granby), 5
+(USBS); <i>Arrowhead</i> (near Dale), 1 (USBS). <i>Boulder Co.</i>: ¾ mi. N, 2 mi.
+W Allenspark, 8400 ft., 4; <i>3 mi. S Ward</i>, 9000 ft., 3; Nederland, 16 (Chi).
+<i>Clear Creek Co.</i>: Mt. McLellan, 2 (USBS); Berthoud Pass, 4. <i>Park Co.</i>:
+Trout Creek Ranch, 2 mi. N Garo, 1 (USBS).</p>
+
+<p>Specimens examined of <i>M. m. nanus</i> from eastern Idaho and Montana are
+as follows: <span class="smcap">Idaho</span>: <i>Custer Co.</i>: Challis, 7 (USBS); Mill Creek, Challis Nat.
+Forest, 1 (USBS); Pahsimeroi Mts., 12 (USBS); Lost River Mts., 1 (USBS).
+<i>Fremont Co.</i>: N fork Snake River, 10 mi. SW Island Park, 6200 ft., 2 (AMNH);
+Black Springs Creek, 4 mi. W Ashton, 5200 ft., 1 (AMNH); 5 mi. W St.
+Anthony, 5000 ft., 1 (AMNH). <i>Camas Co.</i>: Camas Prairie, Corral, 5100 ft.,
+2 (USBS). <i>Blaine Co.</i>: Alturas Lake, 3 (USBS); <i>Sawtooth Lake</i>, 2 (USBS);
+Craters of the Moon, Laidlow Park, 2 (Mich); Ticura, 10 mi. S Picabo, 1
+(USBS); 19 mi. NE Carey (Lava Lake), 8 (Mich). <i>Butte Co.</i>: <i>26 mi. SW
+Arco</i>, 12 (Mich). <i>Bingham Co.</i>: Shelley, 6 (USBS). <i>Bonneville Co.</i>: 10
+mi. SE Irwin, 4 (USBS). <i>Owyhee Co.</i>: Three Creeks, 3 (USBS). <i>Twin
+Falls Co.</i>: Castleford Fenced Plot, 11 mi. W, 9 mi. S Twin Falls, 1. <i>Minidoka
+Co.</i>: <i>Heyburn</i>, 2 (USBS). <i>Cassia Co.</i>: 2 mi. S, 2 mi. W Burley, 5. <i>Bannock
+Co.</i>: Pocatello, 23 (USBS 4); Swan Lake, 1 (USBS). <i>Bear Lake Co.</i>: Montpelier
+Creek, 6700 ft., 3 (USBS). <span class="smcap">Montana</span>: <i>Gallatin Co.</i>: W. Fork of W.
+Fork, Gallatin River, 1 (USBS). <i>Park Co.</i>: Lamar River, 7000 ft., 1 (USBS);
+Gardiner, 1 (USBS). <i>Sweet Grass Co.</i>: 14 mi. S Big Timber, 1 (USBS);
+<i>McLeod</i>, 1 (USBS); West Boulder Creek, 18 mi. SE Livingston, 2 (USBS).</p></blockquote>
+
+
+<p class="center"><b>Microtus montanus codiensis</b>, new subspecies</p>
+
+<blockquote><p><i>Type.</i>&mdash;Female, adult, skin and skull; No. 27578, Museum of Natural History,
+University of Kansas, from 3â…• mi. E and â…— mi. S Cody, 5020 ft., Park
+Co., Wyoming; obtained on August 11, 1948, by James W. Bee, original number
+18-8-11-48.</p></blockquote>
+
+<blockquote><p><i>Range.</i>&mdash;In northwestern Wyoming eastward from the Absaroka and Wind
+River ranges into the western part of the Big Horn Basin.</p>
+
+<p><i>Diagnosis.</i>&mdash;A relatively large <i>Microtus montanus</i>; tail actually and relatively
+long; hind foot actually but not relatively large; skull large; zygomatic expanse
+actually and relatively large; alveolobasilar length relatively large; upper molar
+tooth-row relatively long; color relatively light, not reddish.</p></blockquote>
+
+<hr style="width: 45%;" />
+
+<div class="figcenter" style="width: 560px;">
+<a name="fig_1" id="fig_1"></a>
+<a href="images/i010.png">
+<img src="images/i010_tn.png" width="560" height="600" alt="Fig. 1. Geographic range of Microtus montanus in Wyoming, Colorado,
+and adjacent areas." title="Fig. 1. Geographic range of Microtus montanus in Wyoming, Colorado, and adjacent areas." />
+</a>
+<span class="caption">Fig. 1. Geographic range of Microtus montanus in Wyoming, Colorado,
+and adjacent areas. The solid circles represent localities from which specimens
+have been examined; the hollow circles represent type localities. The ranges
+of subspecies in Utah are after Durrant, 1952.
+
+</span>
+</div>
+
+<blockquote>
+<p>Guide to subspecies</p>
+<ol>
+<li>M. m. nanus</li>
+<li>M. m. codiensis</li>
+<li>M. m. zygomaticus</li>
+<li>M. m. fusus</li>
+<li>M. m. micropus</li>
+<li>M. m. nexus</li>
+<li>M. m. amosus</li>
+<li>M. m. rivularis</li></ol>
+</blockquote>
+
+<hr style="width: 45%;" />
+
+<blockquote><p><i>Comparisons.</i>&mdash;As compared with the specimens of <i>M. m. nanus</i> from Idaho,
+the size is larger (see diagnosis and measurements). Certain proportions
+which differ from those of <i>nanus</i> and which are not in close agreement with
+the observed differences with age in specimens of <i>nanus</i> of a size comparable
+to <i>codiensis</i> are relatively large alveolobasilar length, relatively long alveolar
+length of upper molar tooth-row, relatively wide-spreading zygomatic arches,
+and relatively long tail. The color in <i>codiensis</i> is lighter than in <i>nanus</i>. As
+compared to the new subspecies named below from the Big Horn Mountains
+to the east, <i>codiensis</i> is of similar size in head-body length, but has a relatively
+as well as actually longer tail; the hind foot averages longer; the upper molar
+tooth-row is relatively longer; the color is slightly paler and less grizzled; the
+bullae are larger and less flattened; the angle formed at the suture between
+the basioccipital and basisphenoid bones is less acute; and the region of the
+suture is less prominently elevated between the bullae when viewed from the
+ventral aspect. The pterygoid plates mesial and posterodorsal to the posterior
+end of the last upper molar are less fenestrated, and the incisive foramina are
+less constricted posteriorly.</p></blockquote>
+
+<hr style="width: 45%;" />
+
+<div class="figcenter" style="width: 559px;">
+<a name="fig_2" id="fig_2"></a>
+<a href="images/i011.png">
+<img src="images/i011_tn.png" width="559" height="600" alt="Fig. 2. Map showing the major barriers to Microtus montanus in Wyoming
+and Colorado; the barriers are the low areas named on the map (the name
+&quot;Black Hills&quot; is on the map for another reason; these hills are not a barrier).
+The major mountainous areas higher than approximately 8000 feet in elevation
+in Wyoming, Colorado and Utah are stippled. These mountainous areas
+include the habitat that is most suitable for the montane meadow mouse. The
+Black Hills are unoccupied by this species but these hills seem to be ecologically
+suitable for the species." title="Fig. 2. Map showing the major barriers to Microtus montanus in Wyoming and Colorado." />
+</a>
+<span class="caption">Fig. 2. Map showing the major barriers to Microtus montanus in Wyoming
+and Colorado; the barriers are the low areas named on the map (the name
+&quot;Black Hills&quot; is on the map for another reason; these hills are not a barrier).
+The major mountainous areas higher than approximately 8000 feet in elevation
+in Wyoming, Colorado and Utah are stippled. These mountainous areas
+include the habitat that is most suitable for the montane meadow mouse. The
+Black Hills are unoccupied by this species but these hills seem to be ecologically
+suitable for the species.</span>
+</div>
+
+<hr style="width: 45%;" />
+
+<blockquote><p><i>Measurements.</i>&mdash;The average and the extremes for some measurements of
+34 males and females, 27 from the type locality and 7 from other localities in
+the range assigned to this subspecies, are as follows: total length, 165 (146-186);
+length of tail, 44.2 (35-55); hind foot, 19.6 (17-21); condylobasilar length of
+the skull, 25.5 (24.0-27.5); zygomatic breadth, 15.6 (14.7-16.6); alveolar length
+of upper molar tooth-row, 6.6 (6.2-7.0); prelambdoidal breadth, 8.8 (8.1-9.5);
+lambdoidal breadth, 12.0 (11.2-12.8). As an indication of variability and for
+comparison with other series the coefficient of variability and two times the
+standard error of the mean for each measurement in this series are included
+in <a href="#table_1">Table 1</a>. The averages for some measurements of 27 topotypes are as follows:
+total length, 162; length of tail, 45.5; hind foot, 19.9; condylobasilar
+length, 25.6; palatilar length, 14.0; molar series, 6.6; alveolobasilar length,
+14.9; zygomatic breadth, 15.6; interorbital breadth, 3.5; lambdoidal breadth,
+12.1; prelambdoidal breadth, 8.9; depth of braincase, 7.8.</p></blockquote>
+
+<p><i>Discussion.</i>&mdash;Three species of <i>Microtus</i> were collected by James
+W. Bee at the type locality. <i>Microtus montanus codiensis</i>, <i>Microtus
+longicaudus mordax</i>, and <i>Microtus pennsylvanicus modestus</i> were
+taken in the same runways in the same meadow, at the same time.
+<i>Microtus ochrogaster haydeni</i>, although not taken at this locality,
+occurs in the Big Horn Basin. These four species differ in their
+geographic ranges, being largely allopatric, except <i>M. montanus</i> and
+<i>M. longicaudus</i> which are sympatric. Although the different species
+have ecological preferences and habits which differ, several species
+of <i>Microtus</i> may occur together in local areas such as the above.
+Certain of the characteristics of <i>M. m. codiensis</i> are intermediate
+between those of the species <i>M. montanus</i> on one hand and those
+of the other three species on the other hand. Could interspecific
+hybridization between "good species" of <i>Microtus</i> take place in
+nature and possibly alter the characteristics of a local population?</p>
+
+<blockquote><p><i>Specimens examined.</i>&mdash;Total, 50, distributed as follows (abbreviations for
+collections are given in the account of <i>M. m. nanus</i>; localities that are not
+represented in <a href="#fig_1">Fig. 1</a> because overlapping or crowding of the symbols would
+result are Italicized):</p>
+
+<p><span class="smcap">Montana</span>: <i>Carbon Co.</i>: Beartooth Mountains, 2 (USBS); <i>Beartooth Lake</i>,
+1 (USBS).</p>
+
+<p><span class="smcap">Wyoming</span>: <i>Park Co.</i>: Black Mountain, head of Pat O'Hara Creek, 3 (USBS);
+13 mi. N, 1 mi. E Cody, 5200 ft., 1; SW slope Whirlwind Peak, 9000 ft., 1;
+<i>5 mi. N Cody, 6300 ft.</i>, 1 (USBS); 3â…• mi. E, â…— mi. S Cody, 31; Ishawooa
+Creek, 6300 ft., 2 (USBS); <i>Valley</i>, 1 (USBS); Needle Mountain, 10,500 ft.,
+4 (USBS). <i>Hot Springs Co.</i>: 3 mi. N, 10 mi. W Thermopolis, 4950 ft., 3.</p></blockquote>
+
+
+<p class="center"><b>Microtus montanus zygomaticus</b>, new subspecies</p>
+
+<blockquote><p><i>Type.</i>&mdash;Male, adult, skin and skull, No. 32761, Museum of Natural History,
+University of Kansas, from Medicine Wheel Ranch, 9000 ft., 28 mi. E Lovell,
+Big Horn County, Wyoming; obtained by R. Freiburg, original number 105.</p>
+
+<p><i>Range.</i>&mdash;The Big Horn Mountains of north-central Wyoming.</p>
+
+<p><i>Diagnosis.</i>&mdash;A large <i>Microtus montanus</i> with a relatively short tail; short
+molar series; broad zygomatic arches well rounded in lateral outline when
+viewed from above; small and flattened bullae; raised basioccipito-basisphenoid
+suture.</p>
+
+<p><i>Comparisons.</i>&mdash;For comparison with <i>M. m. codiensis</i> from the west, on the
+other side of the Big Horn Basin, see the account of that subspecies. In comparison
+with <i>nanus</i> this subspecies is slightly paler, in this respect showing
+more resemblance to <i>codiensis</i> although not so pale, and more grizzled or
+unevenly colored. This difference in color between <i>zygomaticus</i> and <i>codiensis</i>
+may not be of taxonomic significance. From both the topotypes of <i>nanus</i>, and
+the series of it from Wyoming, <i>zygomaticus</i> differs on the average in having a
+relatively shorter tail, a relatively shorter upper molar tooth-row, relatively
+more rounded and relatively more wide-spread zygomatic arches, and smaller
+more flattened bullae.</p>
+
+<p><i>Measurements.</i>&mdash;Average and extreme measurements of 24 adult males and
+females from several localities here referred to <i>M. m. zygomaticus</i> are as follows:
+total length, 159(150-175); length of tail, 37.6(31-46); hind foot,
+18.6(17-20); condylobasilar length of the skull, 25.3(24.2-26.7); zygomatic
+breadth, 15.3(14.1-16.7); alveolar length of upper molar tooth-row, 6.2
+(5.7-6.8); prelambdoidal breadth, 8.7(8.3-9.4); lambdoidal breadth, 11.9(11.0-12.5).
+Average and extreme measurements of a series of 12 adult male
+topotypes are as follows: total length, 159(144-174); length of tail, 36.4
+(30-41); hind foot, 18.2(16-20); condylobasilar length of skull, 25.8(24.7-26.7);
+alveolobasilar length, 14.8(13.8-15.3); palatilar length, 13.8 (12.7-14.2); alveolar
+length of upper molar tooth-row, 6.4(5.9-6.6); zygomatic breadth, 15.9
+(15.0-16.7); interorbital breadth, 3.6(3.4-3.7); lambdoidal breadth, 12.1
+(11.5-12.5); prelambdoidal breadth, 8.6(8.3-8.9); depth of braincase, 8.0
+(7.6-8.3).</p></blockquote>
+
+<p><i>Discussion.</i>&mdash;This subspecies is separated from <i>M. m. codiensis</i> to
+the west by the Big Horn Basin. A series from along Buffalo Creek,
+27 mi. N, 1 mi. E Powder River, 6075 ft., in Natrona County, Wyoming,
+is intermediate between the topotypes of <i>zygomaticus</i> and
+<i>nanus</i> in the characters cited above as distinguishing the two, but
+shows greater resemblance to <i>zygomaticus</i> in the shape of the
+zygomatic arch, in color which is paler than in topotypes of
+<i>zygomaticus</i>, and in the short hind foot. On these and on geographic
+grounds this population is referred to <i>zygomaticus</i>. Unfortunately
+we cannot be certain in many cases that an intermediate condition
+in a certain character indicates a genetically intermediate population
+and therefore true intergradation between the two subspecies
+to which the population is geographically intermediate. The topotypes
+of this subspecies are the most distinct of all the series which
+I have studied from the eastern Rocky Mountains, in terms of the
+degree of morphological departure from the norm for the species.
+After <i>zygomaticus</i> the following populations are arranged according
+to their degree of deviation from this norm (<i>codiensis</i> deviates
+most): topotypes of <i>codiensis</i>, <i>fusus</i> and a population from southern
+Sweetwater County, Wyoming, and lastly the <i>nanus-caryi</i> complex.
+Within the latter group, as I have mentioned, there are a number
+of local variants most of which do not differ significantly and do
+not conform to any geographic pattern.</p>
+
+<blockquote><p><i>Specimens examined.</i>&mdash;Total, 55, distributed as follows (abbreviations for
+collections are given in the account of <i>M. m. nanus</i>; localities that are not
+represented in <a href="#fig_1">Fig. 1</a> because overlapping or crowding of the symbols would
+result are Italicized): <span class="smcap">Wyoming</span>: <i>Big Horn Co.</i>: Medicine Wheel Ranch,
+9000 ft., 28 mi. E Lovell, 30; W slope, head of Trappers Creek, 9500 ft., 2
+(USBS). <i>Washakie Co.</i>: 9 mi. E, 5 mi. N Tensleep, 7400 ft., 1. <i>Johnson
+Co.</i>: 7½ mi. W, 1 mi. S Buffalo, 6500 ft., 3; Big Horn Mountains, 3 (USBS).
+<i>Natrona Co.</i>: Buffalo Creek, 27 mi. N, 1 mi. E Powder River, 6075 ft., 16.</p></blockquote>
+
+
+<p class="center"><b>Microtus montanus fusus</b> Hall</p>
+
+<blockquote><p><i>Microtus nanus</i>, Bailey, N. Amer. Fauna, 17:30, June 6, 1900 (part); Cary,
+N. Amer. Fauna 33:123, August 17, 1911.</p>
+
+<p><i>Microtus montanus fusus</i> Hall, Proc. Biol. Soc. Washington, 51:131-134,
+August 23, 1938; Warren, The Mammals of Colorado, Univ. of Okla.
+Press, p. 229, 1942.</p></blockquote>
+
+<blockquote><p><i>Type.</i>&mdash;Male, adult, skin and skull; No. 61281, Museum of Vertebrate
+Zoology; 2½ miles east of summit of Cochetopa Pass, Saguache County, Colorado;
+Sept. 21, 1933; collected by Annie M. Alexander; original number 2568.
+Type not seen by me.</p>
+
+<p><i>Range.</i>&mdash;Southern Colorado and northern New Mexico.</p>
+
+<p><i>Comparisons.</i>&mdash;For comparison with <i>M. m. nanus</i>, the subspecies to the
+northward, see the preceding account of that subspecies. For comparison
+with <i>M. m. amosus</i> the subspecies to the west see Hall (1938) and Durrant
+(1952). I have not examined specimens of <i>amosus</i>.</p>
+
+<p><i>Measurements.</i>&mdash;Average and extreme measurements for 17 adults including
+both males and females from several localities in southern Colorado are as
+follows: total length, 160 (136-179); length of tail, 42 (35-55); hind foot,
+19.2 (17-23); condylobasilar length of the skull, 25.2 (24.0-26.0); zygomatic
+breadth, 15.0 (14.1-15.5); alveolar length of upper molar tooth-row, 6.4
+(6.0-6.7); prelambdoidal breadth, 8.7 (8.3-9.2); lambdoidal breadth, 11.7 (11.1-12.6).</p>
+
+<p>Average and extreme measurements of 4 adults (2 males and 2 females)
+from the type locality and 11 adults (4 males and 7 females) from other
+localities in southern Colorado are as follows: total length, 162 (157-168),
+157 (137-169); length of tail (means only), 44.5, 40.5; hind foot, 18.8 (18-19),
+18.6 (18-23); condylobasilar length of skull, 24.5 (24.0-24.7), 25.2 (24.3-26.1);
+alveolobasilar length, 14.2 (13.9-14.5), 14.6 (14.1-15.1); palatilar length, 13.2
+(13.0-13.4), 13.5 (13.1-14.2); alveolar length of upper molar tooth-row, 6.3
+(6.0-6.6), 6.4 (6.3-6.7); zygomatic breadth, 15.0 (14.3-15.5), 14.9 (14.1-15.5);
+interorbital breadth, 3.5 (3.3-3.6), 3.5 (3.3-3.7); lambdoidal breadth, 11.8 (11.1-12.6),
+11.7 (11.2-12.3); prelambdoidal breadth, 8.6 (8.3-9.2), 8.8 (8.3-9.0);
+depth of braincase, 7.5 (7.2-7.8), 7.6 (7.1-7.9).</p></blockquote>
+
+<p><i>Discussion.</i>&mdash;There is no sharp boundary between <i>M. m. fusus</i> of
+southern Colorado and the subspecies to the north, <i>M. m. nanus</i>.
+Although the line separating these two subspecies is drawn somewhat
+arbitrarily, on the whole the samples from north of this line
+more closely resemble <i>nanus</i>. All of the means for total length
+given above are larger than the maximum given in Hall's description
+of <i>fusus</i>. The caudal index (38 and 35% in two series) is
+slightly larger than that cited by Hall (33.3%) and is not significantly
+different from that in <i>nanus</i> (35.2%). The color in both young and
+old mice is variable, but in general is more yellowish, and less
+grayish, than in any other series studied.</p>
+
+<p>There is a large area in western Colorado and eastern Utah,
+between the known ranges of <i>M. m. fusus</i> and <i>M. m. amosus</i> from
+which there are no specimens. Probably the species occurs only at
+certain places in this arid region which seems to be a partial barrier
+to the species.</p>
+
+<p>Specimens of <i>M. montanus</i> from northern New Mexico have been
+referred previously to <i>M. m. arizonensis</i>. When he named <i>M. m.
+fusus</i>, Hall mentioned its resemblance to <i>arizonensis</i> in reddish
+coloration, but pointed out that <i>fusus</i> is less reddish. Of six specimens
+from Valle Santa Rosa, Jemez Mountains (USBS), 8500 ft.,
+Rio Arriba County, New Mexico, three are immature, and the skulls
+of the remaining specimens are damaged. In reddish color and
+relatively large size these few specimens resemble <i>arizonensis</i> more
+than <i>fusus</i> although the locality of occurrence is closer to the geographic
+range of the northern <i>fusus</i> than to that of <i>arizonensis</i>. The
+identification of these specimens as <i>arizonensis</i> is provisional; additional
+specimens are needed from the area, 200 miles wide, which
+separates the ranges as now known of <i>arizonensis</i> in Arizona from
+the occurrence in New Mexico. There is a single specimen from
+this area, the damaged skull of which prevents conclusive identification.
+The specimen is either <i>M. montanus</i> or <i>M. mexicanus</i>, and
+is from Nutria, on the southern edge of the Zuni Mountains (USBS).
+Detailed comparison of <i>fusus</i> and <i>arizonensis</i> is not attempted here
+although it may be stated that in several characters <i>fusus</i> is intermediate
+between <i>arizonensis</i> to the south and <i>nanus</i> to the north.</p>
+
+<blockquote><p><i>Specimens examined.</i>&mdash;Total, 89, distributed as follows (abbreviations for
+collections are given in the account of <i>M. m. nanus</i>; localities that are not
+represented in <a href="#fig_1">Fig. 1</a> because overlapping or crowding of the symbols would
+result are Italicized):</p>
+
+<p><span class="smcap">Colorado</span>: <i>Pitkin Co.</i>: 5 mi. W Independence Pass, 11,000 ft., 1 (Chi).
+<i>Lake Co.</i>: <i>Independence Pass, 12,095 ft.</i>, 2 (Chi). <i>Gunnison Co.</i>: <i>Gothic</i>, 2
+(USBS); Decker's Ranch, Crested Butte, 2 (AMNH); Almont, 3 (USBS).
+<i>Montrose Co.</i>: Coventry, 5 (USBS 4, AMNH 1). <i>Saguache Co.</i>: Cochetopa
+Pass and environs, 44 (USBS 22). <i>Hinsdale Co.</i>: Ruby Lake, 1 (USBS).
+<i>Mineral Co.</i>: 3 mi. E Creede, 1; 23 mi. S, 11 mi. E Creede, 9300 ft., 7. <i>La
+Plata Co.</i>: Florida, 6800 ft., 1. <i>Conejos Co.</i>: 1 mi. S, 19 mi. W Antonito,
+10,200 ft., 3; <i>4 mi. S, 23 mi. W Antonito</i>, 1; <i>5 mi. S, 24 mi. W Antonito, 9600
+ft.</i>, 9.</p>
+
+<p><span class="smcap">New Mexico</span>: <i>Rio Arriba Co.</i>: 6 mi. W Hopewell, 9900 ft., 6 (USBS);
+<i>Tusas River, 8700 ft.</i>, 1 (USBS).</p></blockquote>
+
+<p>Some measurements not given above are included in <a href="#table_1">Table 1</a>,
+together with the number of specimens and the sex if restricted to
+one sex. So that the variability can be evaluated more adequately,
+the coefficient of variability and 2 times the standard error of the
+mean are included for the measurements in two series. The series
+consist of all the adult specimens (with a condylobasilar length of
+24.0 mm. or more) of both sexes from the areas specified. Various
+barriers are shown in <a href="#fig_2">Fig. 2</a> for comparison with the distributions
+of the subspecies and the localities of known occurrence shown in
+<a href="#fig_1">Fig. 1</a>. <i>Microtus montanus</i> has not been taken in the Black Hills
+area of extreme northeastern Wyoming. Suitable montane habitat
+is present and both <i>Microtus pennsylvanicus insperatus</i> and <i>Microtus
+longicaudus longicaudus</i> occur there. The arid basin of the Powder
+River presumably is a barrier that has prevented <i>M. montanus</i> from
+reaching this area.</p>
+
+<hr style="width: 45%;" />
+
+<p class="center"><a name="table_1" id="table_1"></a><span class="smcap">Table 1. Average Measurements, in Millimeters, of Adults of
+Microtus montanus.</span></p>
+
+<p>
+Key to column headings:<br />
+<br />
+A: No. of individuals averaged<br />
+B: Total length<br />
+C: Length of tail<br />
+D: Length of hind foot<br />
+E: Condylobasilar length<br />
+F: Alveolar length of upper molar tooth-row<br />
+G: Zygomatic breadth<br />
+H: Lambdoidal breadth<br />
+I: Prelambdoidal breadth<br />
+<br />
+</p>
+
+<table border="0" cellpadding="4" cellspacing="0" summary="Average Measurements of Adults of Microtus montanus.">
+<tr>
+<th align="center"><span class="smcap">Locality</span></th>
+<th align="center">A</th>
+<th align="center">B</th>
+<th align="center">C</th>
+<th align="center">D</th>
+<th align="center">E</th>
+<th align="center">F</th>
+<th align="center">G</th>
+<th align="center">H</th>
+<th align="center">I</th>
+</tr>
+<tr><td align="center" colspan="10"><i>M. m. codiensis</i>, all</td></tr>
+<tr><td align="left">Average</td><td align="right">34</td><td align="right">165.3</td><td align="right">44.2</td><td align="right">19.6</td><td align="right">25.47</td><td align="right">6.56</td><td align="right">15.55</td><td align="right">12.05</td><td align="right">8.76</td></tr>
+<tr><td align="left">2 × stand. error</td><td align="right">&nbsp;</td><td align="right">3.56</td><td align="right">1.84</td><td align="right">.395</td><td align="right">.308</td><td align="right">.067</td><td align="right">.198</td><td align="right">.144</td><td align="right">.129</td></tr>
+<tr><td align="left">Coeff. variab</td><td align="right">&nbsp;</td><td align="right">6.0</td><td align="right">11.6</td><td align="right">5.6</td><td align="right">3.5</td><td align="right">3.0</td><td align="right">3.65</td><td align="right">1.20</td><td align="right">1.47</td></tr>
+<tr><td align="center" colspan="10"><i>M. m. nanus</i>, Eastern Idaho</td></tr>
+<tr><td align="left">Average</td><td align="right">21<a name="FNanchor_1_1" id="FNanchor_1_1"></a><a href="#Footnote_1_1" class="fnanchor">[1]</a></td><td align="right">151.1</td><td align="right">39.4</td><td align="right">19.2</td><td align="right">25.00</td><td align="right">6.44</td><td align="right">14.99</td><td align="right">11.74</td><td align="right">8.94</td></tr>
+<tr><td align="left">2 × stand. error</td><td align="right">&nbsp;</td><td align="right">3.20</td><td align="right">2.89</td><td align="right">.293</td><td align="right">.286</td><td align="right">1.15</td><td align="right">.295</td><td align="right">.210</td><td align="right">.182</td></tr>
+<tr><td align="left">Coeff. variab.</td><td align="right">&nbsp;</td><td align="right">6.1</td><td align="right">21.1</td><td align="right">4.38</td><td align="right">2.49</td><td align="right">3.99</td><td align="right">4.10</td><td align="right">3.79</td><td align="right">4.31</td></tr>
+<tr><td align="center" colspan="10"><i>M. m. nanus</i>, Wyoming</td></tr>
+<tr><td align="left">Teton Co.</td><td align="right">35</td><td align="right">160.5</td><td align="right">40.7</td><td align="right">18.6</td><td align="right">25.16</td><td align="right">6.51</td><td align="right">15.17</td><td align="right">11.86</td><td align="right">8.77</td></tr>
+<tr><td align="left">Fremont Co.</td><td align="right">26</td><td align="right">157.0</td><td align="right">41.4</td><td align="right">19.6</td><td align="right">25.23</td><td align="right">6.25</td><td align="right">15.05</td><td align="right">11.88</td><td align="right">8.91</td></tr>
+<tr><td align="left">Lincoln Co.</td><td align="right">24</td><td align="right">159.9</td><td align="right">41.8</td><td align="right">18.9</td><td align="right">25.08</td><td align="right">6.26</td><td align="right">15.10</td><td align="right">11.82</td><td align="right">8.75</td></tr>
+<tr><td align="left">Uinta Co.</td><td align="right">26</td><td align="right">162.4</td><td align="right">41.3</td><td align="right">19.0</td><td align="right">25.33</td><td align="right">6.42</td><td align="right">15.31</td><td align="right">12.16</td><td align="right">8.89</td></tr>
+<tr><td align="left">Sweetwater Co.</td><td align="right">12</td><td align="right">159.8</td><td align="right">43.7</td><td align="right">20.1</td><td align="right">24.98</td><td align="right">6.31</td><td align="right">15.00</td><td align="right">11.84</td><td align="right">9.02</td></tr>
+<tr><td align="left">Natrona Co.</td><td align="right">40</td><td align="right">159.6</td><td align="right">41.0</td><td align="right">19.6</td><td align="right">25.04</td><td align="right">6.40</td><td align="right">15.00</td><td align="right">11.84</td><td align="right">8.93</td></tr>
+<tr><td align="left">Carbon Co.</td><td align="right">108</td><td align="right">158.7</td><td align="right">40.0</td><td align="right">19.1</td><td align="right">24.96</td><td align="right">6.27</td><td align="right">15.05</td><td align="right">11.83</td><td align="right">8.72</td></tr>
+<tr><td align="left">&nbsp;&nbsp; Encampment ♂</td><td align="right">27</td><td align="left">161</td><td align="right">41.7</td><td align="right">18.9</td><td align="left">25.1</td><td align="right">6.16</td><td align="left">15.2</td><td align="left">11.9</td><td align="left">8.7</td></tr>
+<tr><td align="left">&nbsp;&nbsp; Encampment ♀</td><td align="right">11</td><td align="left">159</td><td align="right">41.1</td><td align="right">19.4</td><td align="left">24.9</td><td align="right">6.18</td><td align="left">14.9</td><td align="left">11.6</td><td align="left">8.6</td></tr>
+<tr><td align="left">&nbsp;&nbsp; Savery ♂</td><td align="right">23</td><td align="left">159</td><td align="right">41.0</td><td align="right">19.2</td><td align="left">25.3</td><td align="left">6.3</td><td align="left">15.2</td><td align="left">12.2</td><td align="left">8.8</td></tr>
+<tr><td align="left">&nbsp;&nbsp; Savery ♀</td><td align="right">25</td><td align="left">155</td><td align="right">37.1</td><td align="right">18.8</td><td align="left">24.7</td><td align="left">6.3</td><td align="left">14.8</td><td align="left">11.6</td><td align="left">8.6</td></tr>
+<tr><td align="center" colspan="10"><i>M. m. nanus</i></td></tr>
+<tr><td align="left">Northern Colo.</td><td align="right">8</td><td align="right">163.1</td><td align="right">42.4</td><td align="right">19.6</td><td align="right">25.20</td><td align="right">6.44</td><td align="right">14.86</td><td align="right">11.70</td><td align="right">8.56</td></tr>
+<tr><td align="center" colspan="10"><i>M. m. fusus</i></td></tr>
+<tr><td align="left">Southern Colo.</td><td align="right">17<a name="FNanchor_2_2" id="FNanchor_2_2"></a><a href="#Footnote_2_2" class="fnanchor">[2]</a></td><td align="right">159.8</td><td align="right">42.4</td><td align="right">19.2</td><td align="right">24.97</td><td align="right">6.43</td><td align="right">14.98</td><td align="right">11.73</td><td align="right">8.69</td></tr>
+</table>
+
+
+<div class="footnote"><p><a name="Footnote_1_1" id="Footnote_1_1"></a><a href="#FNanchor_1_1"><span class="label">[1]</span></a> For external parts, 34 individuals were used.</p></div>
+
+<div class="footnote"><p><a name="Footnote_2_2" id="Footnote_2_2"></a><a href="#FNanchor_2_2"><span class="label">[2]</span></a> For external parts, 29 individuals were used.</p></div>
+
+<hr style="width: 45%;" />
+
+<h2>REFERENCES CITED</h2>
+
+<p><span class="smcap">Bailey, V.</span></p>
+
+<blockquote><p>1900. Revision of American voles of the genus <i>Microtus</i>. N. Amer. Fauna,
+17:1-88, June 6.</p>
+
+<p>1917. A new subspecies of meadow mouse from Wyoming. Proc. Biol.
+Soc. Washington, 30:29-30, February 21.</p></blockquote>
+
+<p><span class="smcap">Cary, M.</span></p>
+
+<blockquote><p>1911. A biological survey of Colorado.  N. Amer. Fauna, 33:1-256,
+August 17.</p>
+
+<p>1917. Life zone investigations in Wyoming. N. Amer. Fauna, 42:1-95,
+October 3.</p></blockquote>
+
+<p><span class="smcap">Davis, W. B.</span></p>
+
+<blockquote><p>1939. The Recent mammals of Idaho. 400 p., front., illus., maps, diagrs.,
+Caldwell, Id., The Caxton Printers Ltd., April 5.</p></blockquote>
+
+<p><span class="smcap">Durrant, S. D.</span></p>
+
+<blockquote><p>1952. Mammals of Utah, taxonomy and distribution. Univ. Kans. Publ.,
+Mus. Nat. Hist., 6:1-549, 91 figs, in text, 30 tables, August 10.</p></blockquote>
+
+<p><span class="smcap">Elton, C.</span></p>
+
+<blockquote><p>1942. Voles, mice and lemmings; problems in population dynamics. 496
+p., illus. (maps), tables, Oxford, The Clarendon Press, London.</p></blockquote>
+
+<p><span class="smcap">Goin, O. B.</span></p>
+
+<blockquote><p>1943. A study of individual variation in <i>Microtus pennsylvanicus pennsylvanicus</i>.
+Jour. Mamm., 24:212-224, June 7.</p></blockquote>
+
+<p><span class="smcap">Hall, E. R.</span></p>
+
+<blockquote><p>1938. Notes on the meadow mice <i>Microtus montanus</i> and <i>Microtus nanus</i>
+with descriptions of a new subspecies from Colorado. Proc. Biol.
+Soc. Washington, 51:131-134, August 23.</p></blockquote>
+
+<p><span class="smcap">Hill, E. A.</span></p>
+
+<blockquote><p>1935. Cranial foramina in rodents. Jour. Mamm. 16(2):121-129.</p></blockquote>
+
+<p><span class="smcap">Howell, A. B.</span></p>
+
+<blockquote><p>1924. Individual and age variation in <i>Microtus montanus yosemite</i>. Jour.
+Agr. Res., 28(10):977-1015, June 7.</p></blockquote>
+
+<p><span class="smcap">Kellogg, R.</span></p>
+
+<blockquote><p>1922. A study of the California forms of the <i>Microtus montanus</i> group of
+meadow mice. Univ. California Publ. Zool., 21:245-274, April 18.</p></blockquote>
+
+<p><span class="smcap">Merriam, C. H.</span></p>
+
+<blockquote><p>1891. Results of a biological reconnaissance of south-central Idaho. N.
+Amer. Fauna, 5:1-113, July 30.</p></blockquote>
+
+<p><span class="smcap">Piper, S. E.</span></p>
+
+<blockquote><p>1909. The Nevada mouse plague of 1907-8. U. S. Dept. Agr., Farmers'
+Bul. 352, pp. 1-23, 9 figs.</p></blockquote>
+
+<p><span class="smcap">Warren, E. R.</span></p>
+
+<blockquote><p>1942. The mammals of Colorado, their habits and distribution. Second
+(revised) edition, Univ. Oklahoma Press, Norman, xviii-330 p.,
+front., 50 plates.</p></blockquote>
+
+
+<p><i>Transmitted March 22, 1954.</i></p>
+
+
+<p class="center">
+25-3560
+</p>
+
+<hr style="width: 45%;" />
+
+<h4>Transcriber's Notes</h4>
+
+<p>One typographical error was corrected:
+"Castlford" was corrected to "Castleford" in "Twin Falls Co.: Castleford Fenced Plot".</p>
+
+
+
+
+
+
+
+
+<pre>
+
+
+
+
+
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+The Project Gutenberg EBook of Subspeciation in the Meadow Mouse, Microtus
+montanus, in Wyoming and Colorado, by Sydney Anderson
+
+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: Subspeciation in the Meadow Mouse, Microtus montanus, in Wyoming and Colorado
+
+Author: Sydney Anderson
+
+Release Date: March 22, 2010 [EBook #31730]
+
+Language: English
+
+Character set encoding: ASCII
+
+*** START OF THIS PROJECT GUTENBERG EBOOK MEADOW MOUSE ***
+
+
+
+
+Produced by Chris Curnow, Simon Gardner, Joseph Cooper and
+the Online Distributed Proofreading Team at
+http://www.pgdp.net
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+
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+
+
+
+UNIVERSITY OF KANSAS PUBLICATIONS
+
+MUSEUM OF NATURAL HISTORY
+
+Volume 7, No. 7, pp. 489-506, 2 figures in text
+
+July 23, 1954
+
+
+
+
+                    Subspeciation in the Meadow Mouse,
+                      Microtus montanus, in Wyoming
+                              and Colorado
+
+                                  BY
+
+                            SYDNEY ANDERSON
+
+
+UNIVERSITY OF KANSAS
+
+LAWRENCE
+
+1954
+
+
+UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY
+
+Editors: E. Raymond Hall, Chairman, A. Byron Leonard,
+Robert W. Wilson
+
+Volume 7, No. 7, pp. 489-506, 2 figures in text
+
+Published July 23, 1954
+
+
+UNIVERSITY OF KANSAS
+
+Lawrence, Kansas
+
+
+PRINTED BY
+
+FERD VOILAND, JR., STATE PRINTER
+
+TOPEKA, KANSAS
+
+1954
+
+25-3560
+
+
+
+
+Subspeciation in the Meadow Mouse, Microtus montanus, in Wyoming and
+Colorado
+
+BY
+
+SYDNEY ANDERSON
+
+
+_Microtus montanus_ reaches the eastern limits of its geographic
+distribution in Wyoming and Colorado. There the mountains, but in
+general not the lowlands, are occupied by this species. A certain
+minimum of moisture may be of direct importance to the mouse and
+certainly is indirectly important, because certain hydrophytic or
+mesophytic grasses used by the mouse for food, for protection from
+enemies, and for shelter from the elements are dependent on the
+moisture. Areas suitable for _Microtus montanus_ are separated by
+deserts that are dominated by sagebrush and other xerophytic plants or
+by forests or rocky exposures at higher altitudes. A relatively small
+percentage, probably less than ten per cent, of the total area even in
+the more favorable parts of the range of the species is suitable for
+occupancy. In these mice, as in other microtines (Elton, 1942; Piper,
+1909), there are seasonal, and irregularly multiannual fluctuations in
+population density, which sometimes are extreme. Consequently the mice
+at some times seem to be absent from suitable habitats, and at some
+other times occur there in amazingly large numbers.
+
+Because the species is broken up into partly isolated, or at times
+completely isolated, colonies or local populations it may be supposed
+that various evolutionary forces such as selection and random genetic
+drift operate to foster variation. The degree to which racial
+distinction is attained may depend upon these forces and the time
+available. In _Microtus montanus_ in the eastern Rocky Mountains the
+degree of subspecific distinction is not great.
+
+The study here reported upon is based on 1,187 specimens of _Microtus
+montanus_ from Wyoming, Colorado, Idaho and Montana, and on work in the
+field. I spent approximately four months in the field in this area, in
+the summers of 1950, 1951, and 1952. The specimens studied were arranged
+according to localities and the larger series were compared
+statistically. Each of two series, totaling 136 specimens, was studied
+intensively to ascertain the kind and range of variation within single
+populations. Twenty-seven measurements, various proportions based on
+these measurements, and differences in color were analyzed. Fifteen
+characters, judged to be most significant, were selected for use in
+comparing all series. In addition, certain characters that can not be
+expressed easily by measurements, such as inflation of the auditory
+bullae and the curvature of the zygomatic arch, were observed. The
+studies by A. B. Howell (1924) of variation in _Microtus montanus
+yosemite_ Grinnell in California and those by O. B. Goin (1943) of
+_Microtus pennsylvanicus pennsylvanicus_ (Ord) were useful. The external
+measurements are from the collectors' field labels. The measurements of
+the skull all were taken with dial calipers reading to a tenth of a
+millimeter. The anteroposterior measurements of the skull all were taken
+along the shortest line between the points specified below and are not
+necessarily along a line parallel to the long axis of the skull. These
+measurements were taken on the left side of the skull whenever possible.
+Some of the skulls are damaged and therefore some measurements could not
+be taken and are omitted in the computations. Measurements are in
+millimeters.
+
+The results of these studies were submitted to the Department of Zoology
+and the Graduate School of the University of Kansas in partial
+fulfillment of the requirements for the degree of Master of Arts (1952)
+and are available in manuscript form at the Museum of Natural History
+and the library of the University of Kansas.
+
+
+EXPLANATION OF MEASUREMENTS
+
+     Caudal index.--the length of the tail expressed as a percentage of
+     the length of the head and body. The length of the head and body is
+     the collector's measurement of total length less the length of the
+     tail.
+
+CRANIAL MEASUREMENTS OF LENGTH.
+
+     Condylobasilar length.--from the exoccipital condyle to the most
+     posterior point on the border of the alveolus of the upper incisor.
+
+     Alveolobasilar length.--from the posterior border of the alveolus
+     of the third upper molar to the posterior border of the alveolus of
+     the incisor.
+
+     Palatilar length.--from the anteriormost part of the posterior
+     border of the bony shelf of the palate to the posteriormost part of
+     the alveolus of the incisor.
+
+     Alveolar length of upper molar tooth-row.--from the most posterior
+     point of the alveolus of the third upper molar to the most anterior
+     point of the alveolus of the first upper molar.
+
+MEASUREMENTS OF BREADTH.
+
+     Zygomatic breadth.--greatest transverse width.
+
+     Interorbital breadth.--the breadth of the interorbital
+     constriction.
+
+     Lambdoidal breadth.--between the lateralmost points of the
+     lambdoidal crest.
+
+     Prelambdoidal breadth.--between the medialmost margins of the
+     prominent fenestrae in the posterodorsal parts of the squamosal
+     bones. To these fenestrae Howell (1924:995) applied the adjective
+     "prelambdoidal," but other authors have used other names (see Hill,
+     1935:127).
+
+     Depth of braincase.--shortest distance from the ventral side of the
+     cranium at the suture between the basioccipital and basisphenoid
+     bones to the dorsal surface of the cranium (usually not
+     perpendicular to the long axis of the skull).
+
+The history of our knowledge of _Microtus montanus_ in this area begins
+with the early work of the United States Bureau of Biological Survey
+directed by C. H. Merriam (1891), and participated in by Vernon Bailey
+(1900, 1917), Merritt Cary (1911, 1917), and others. The changes in
+nomenclature which grew out of increased understanding of these mice
+through additional collecting are expressed in the synonymies under the
+accounts of subspecies. As a result of my studies two of the three
+subspecific names previously proposed for mice from this area have been
+retained although changes are proposed in the ranges assigned to the two
+subspecies and two additional heretofore unrecognized subspecies are
+named and described. Furthermore the additional specimens and my studies
+of variation make modifications in the characterization of these
+subspecies necessary. Not all of the samples here assigned to a single
+subspecies are identical and I therefore list and discuss some of the
+local variants.
+
+     Numerous members of summer field parties from the Museum of Natural
+     History at the University of Kansas collected most of the specimens
+     studied and wrote field notes that have been helpful. I am grateful
+     to these persons and to Professor E. R. Hall and Dr. R. H. Baker
+     for their assistance and helpful suggestions. Specimens in the
+     following museums were made available by their respective curators:
+     Chicago Natural History Museum by Mr. Colin C. Sandborn, The Museum
+     of Zoology at the University of Michigan by Dr. E. T. Hooper, The
+     American Museum of Natural History by Mr. G. G. Goodwin, The United
+     States National Museum by Dr. David H. Johnson and the Biological
+     Surveys Collection by Miss Viola S. Schantz. A fellowship from the
+     National Science Foundation made possible the studies at the
+     museums other than at the University of Kansas.
+
+
+
+=Microtus montanus nanus= (Merriam)
+
+     _Arvicola (Mynomes) nanus_ Merriam, N. Amer. Fauna, 5:62, July 30,
+     1891.
+
+     _Microtus montanus nanus_, Hall, Proc. Biol. Soc. Washington,
+     51:131, August 23, 1938.
+
+     _Microtus nanus_, Bailey, N. Amer. Fauna, 17:30, June 6, 1900
+     (part).
+
+     _Microtus montanus caryi_ Bailey, Proc. Biol. Soc. Washington,
+     30:29, February 21, 1917.
+
+     _Microtus nanus nanus_, Cary, N. Amer. Fauna, 42:43, October 3,
+     1917 (part).
+
+     _Type._--Adult male No. 23853/31253, U. S. National Museum,
+     Department of Agriculture collection, from Pahsimeroi Mountains,
+     Custer County, Idaho; obtained by C. Hart Merriam and Vernon
+     Bailey, September 16, 1890.
+
+     _Range._--Idaho; southwestern Montana; most of the southwestern
+     half of Wyoming; southward to central Colorado. See figure 1.
+
+     _Comparisons._--Comparisons with subspecies named as new in this
+     paper will be found in the accounts of those subspecies beyond.
+     From _Microtus montanus fusus_ Hall, the subspecies to the south,
+     _M. m. nanus_ from Idaho differs as follows: averages smaller;
+     slightly darker and less reddish and less yellowish in color;
+     slightly wider braincase (see measurement of prelambdoidal
+     breadth); larger bullae.
+
+     _Measurements._--Average (= arithmetical mean) measurements of 34
+     specimens, both male and female, from several localities in eastern
+     Idaho are: total length, 151; length of tail, 39; hind foot, 19.2;
+     condylobasilar length of the skull, 25.0; zygomatic breadth, 15.0;
+     alveolar length of upper molar tooth-row, 6.4; prelambdoidal
+     breadth, 8.9; and lambdoidal breadth, 11.7.
+
+     Average and extreme measurements of six adult males from near
+     Pocatello, Bannock County, Idaho, and nine adult males from near
+     Afton, Lincoln County, Wyoming, are, respectively, as follows:
+     total length, 143(135-150), 163(143-179); length of tail,
+     35.1(33-38), 42.8(36-49); caudal index, 32.0(28.0-33.1),
+     35.7(30.6-41.9); hind foot, 18.9(18-20), 18.8(17-20);
+     condylobasilar length of skull, 24.4(24.0-26.0), 25.6(24.5-26.2);
+     alveolobasilar length, 14.1(13.7-14.5), 14.6(13.8-15.0); palatilar
+     length, 13.2(12.9-13.6), 13.8(13.2-14.5); alveolar length of upper
+     molar tooth-row, 6.3(6.1-6.5), 6.3(6.0-6.6); depth of braincase,
+     7.7(7.5-7.9), 8.0(7.7-8.3); lambdoidal breadth, 11.4(11.0-11.7),
+     12.0(11.3-12.7); prelambdoidal breadth, 9.1(8.6-9.4), 8.7(8.0-9.4);
+     zygomatic breadth, 14.3(13.8-14.7), 15.3(14.4-16.3); interorbital
+     breadth, 3.6(3.5-3.7), 3.5(3.3-3.7). The average length of the
+     nasal bones in the series from Pocatello is 7.1 mm. The averages,
+     which have not been included in Table 1, for three measurements of
+     the series from Carbon County, Wyoming, are as follows (Encampment,
+     males; Encampment, females; Savery, males; and Savery, females,
+     respectively): alveolobasilar length, 14.4, 14.3, 14.5, 14.3;
+     interorbital breadth, 3.5, 3.4, 3.5, 3.4; depth of braincase, 7.8,
+     7.6, 7.9, 7.6. Additional measurements are included in Table 1 for
+     other series.
+
+_Discussion._--The name _Microtus montanus caryi_ Bailey is here placed
+in synonymy under _M. m. nanus_ (Merriam). Vernon Bailey (1917) in his
+description of _caryi_ made four assumptions that have been found to be
+entirely or partly invalid. First, he assumed that this is an "extreme
+variant which gradually changes in characters across Nevada and Utah,
+and reaches its maximum variation in Wyoming." The differences pointed
+out in subsequent descriptions of subspecies found in the above area do
+not show a gradual change in any character, or in the number of
+characters, nor is _caryi_ an extreme when compared with the other
+subspecies. Second, _Microtus nanus_ was not, as Bailey assumed, a
+different species than _Microtus montanus_. Third, he assumed that the
+characteristics of adults of _nanus_ were adequately ascertainable from
+the thirteen topotypes available to him. Subsequent sampling from Idaho
+shows that the series of specimens available to Bailey was made up
+mostly of young and subadult animals. Finally, _caryi_ does not occupy
+as Bailey stated "the meadows along streams in the arid sagebrush
+country of the Bear River, Green River, and Wind River valleys"
+exclusively, or characteristically. When the localities from which the
+species actually is known are plotted, it seems that the arid basin
+serves as a barrier and that the species is more commonly and abundantly
+found in montane meadows in the Transition and Canadian life-zones.
+
+Certain samples, here assigned to _M. m. nanus_, that vary from the
+average of the subspecies deserve comment. For example, mice from the
+area in Wyoming southwest of the Green River (in the Uinta Mountains)
+have relatively smaller feet, but are larger in both total length and
+size of skull. Specimens from near Afton, Lincoln County, Wyoming, are
+relatively large in both total length and size of skull. This series and
+specimens from Teton County, Wyoming, are intermediate between _nanus_
+from Idaho and the newly named subspecies from near Cody, Park County,
+Wyoming, described below, in terms of both darkness and the amount of
+reddish color. Mice from Laramie County are more richly reddish-brown.
+The specimens from near Savery, in Carbon County, Wyoming, are darker.
+The alveolobasilar length relative to the condylobasilar length is
+smaller in the series from along Deer Creek, 16 mi. S, 11 mi. W Waltman,
+Natrona County, Wyoming. The series from the southern tier of counties
+in Wyoming and some of the specimens from Colorado have relatively wider
+zygomatic arches. The specimens from southern Sweetwater County,
+Wyoming, are relatively paler, have a relatively longer tail and longer
+hindfeet, lesser condylobasilar length, and wider braincase. Most of
+these variations are of questionable significance; they may be chance
+variations owing to errors in sampling.
+
+Much of the south-central part of the state is relatively low and
+relatively arid. This area includes the arid basin of the Green River
+and its major tributaries and the arid Red Desert along the continental
+divide in Sweetwater County. This area might have acted as a barrier to
+the mice; gene flow might have been prevented between the populations of
+the western part of the state and those farther east in the Medicine Bow
+Mountains and Laramie Mountains. Nevertheless geographic variations of
+subspecific worth have not taken place. The barrier has either not been
+of as long duration, or has not been so complete and effective, as the
+other barriers in the state, namely the Absaroka Range, the Big Horn
+Basin, the Shoshone Basin, and the valley of the North Platte River.
+These four barriers presumably have led to the differentiation of the
+two subspecies that are newly named beyond. Each of the two areas which
+is set apart by these barriers and in which one of the newly named
+subspecies has evolved is small; therefore there is a lesser amount of
+suitable habitat available for each of the newly named mice than there
+is for _M. m. nanus_. It is conceivable, therefore, that in periods of
+adverse conditions in each of the small areas the size of the effective
+breeding population may have been so small that random genetic drift
+could have operated effectively, or that selection was more critical
+than in a larger, more stable population. It is difficult to test these
+possibilities because the selective value of the taxonomic characters is
+unknown. The observed pattern of variation and facts of distribution
+are, however, not contradictory to the above possibilities.
+
+     _Specimens examined._--Total, 993, distributed as follows: All
+     specimens unless otherwise indicated are in the University of
+     Kansas Museum of Natural History. Specimens in other museums are
+     labeled as follows: Chicago Natural History Museum (Chi);
+     University of Michigan, Museum of Zoology (Mich); American Museum
+     of Natural History (AMNH); United States National Museum (USNM);
+     Biological Surveys Collection (USBS). Localities that are not
+     represented in Fig. 1 because overlapping or crowding of the
+     symbols would result are Italicized. Localities are arranged from
+     north to south by states, within a state from northwest to
+     southeast by counties, and within a county from north to south.
+
+     WYOMING: _Yellowstone Park_: Canyon Camp, 1 (USBS); Lower Geyser
+     Basin, 1 (USBS); Upper Yellowstone River, 2 (AMNH); _North end of
+     Lake_, _Yellowstone National Park_, 2 (AMNH). _Teton Co._: Pacific
+     Creek, 1 (USBS); _Big Game Ridge_, 3 (USBS 1, Mich 2); _Whetstone
+     Creek_, 7 (Mich); Moran and environs (6 localities within a 5 mile
+     radius), 28 (USBS 2, Mich 5); _S fork Buffalo River_, 7 (AMNH); 2
+     mi. W pass, Black Rock Creek, 1 (USBS); _Jenny Lake_, 5 (Mich); Bar
+     BC Ranch, 2-1/2 mi. NE Moose, 6500 ft., 2; Teton Pass above Fish
+     Creek, 1 (USBS); Jackson and environs, 142 (Mich 141); _Sheep
+     Creek_, 2 (Mich). _Lincoln Co.: 13 mi. N, 2 mi. W Afton_, 2; _10
+     mi. N, 2 mi. W Afton_, 4; _9-1/2 mi. N, 2 mi. W Afton_, 3; 9 mi. N,
+     2 mi. W Afton, 9; _7 mi. N, 1 mi. W Afton_, 12; Afton, 1 (USBS);
+     Labarge Creek, 1 (USBS); Border, 6 (USBS); _Cokeville_, 2 (USBS); 6
+     mi. N, 2 mi. E Sage, 1; Cumberland, 5 (USBS). _Sublette Co._: 34
+     mi. N, 4 mi. W Pinedale, 1; _33 mi. N, 2 mi. W Pinedale_, 6; _32
+     mi. N, 1 mi. W Pinedale_, 1; _31 mi. N Pinedale_, 4; Dell Creek, on
+     Ferris Ranch, 7 (Mich); Horse Creek, 7800 ft., Merna, 4 (USBS); Big
+     Piney, 1 (USBS). _Fremont Co._: 17-1/2 mi. W, 2-1/2 mi. N Lander,
+     9500 ft., 3; _17 mi. W, 2 mi. N Lander, 9300 ft._, 4; Milford and
+     environs (5 localities within a 1 mile radius), 23 (USBS 4); 15-1/2
+     mi. S, 7-1/2 mi. W Lander, 9200 ft., 1; South Pass City, 8000 ft, 8
+     (USBS); _23-1/2 mi. S, 5 mi. W Lander, 8600 ft._, 7. _Natrona Co._:
+     Deer Creek, 16 mi. S, 11 mi. W Waltman, 6950 ft., 44; 6 mi. S, 2
+     mi. W Casper, 5900 ft., 4; _6-4/5 mi. S, 2 mi. W Casper, 6100 ft._,
+     1; _7 mi. S, 2 mi. W Casper, 6370 ft._, 3; _10 mi. S Casper, 7750
+     ft._, 33; Sun, 2 (USBS); _5 mi. W Independence Rock, 6000 ft._, 4;
+     _5 mi. W, 1 mi. S Independence Rock_, 2. _Converse Co._: Beaver, 1
+     (USBS). _Uinta Co.: 1-1/2 mi. W, 1/2 mi. S Cumberland_, 6; _16 mi.
+     S, 2 mi. W Kemmerer, 6700 ft._, 3; 10 mi. SW Granger, 3 (Mich);
+     Fort Bridger, 6650 ft., 25 (USNM 6); 9 mi. S Robertson, 8000 ft.,
+     9; _9-1/2 mi. S, 1/2 mi. W Robertson, 8600 ft._, 1; _10 mi. S, 1
+     mi. W Robertson, 8700 ft._, 25; _14 mi. S, 2 mi. E Robertson, 9000
+     ft._, 5; 4 mi. S Lonetree, 1 (USBS). _Sweetwater Co._: Farson, 3;
+     Bitter Creek, 3 (AMNH); Kinney Ranch, 21 mi. S Bitter Creek, 6800
+     ft., 9 (USNM 1, AMNH 2); 32 mi. S, 22 mi. E Rock Springs, 7025 ft.,
+     on Vermillion Creek, 15. _Carbon Co._: 18 mi. NNE Sinclair, 6500
+     ft., 10; Bridgers Pass, 18 mi. SW Rawlins, 7500 ft., 7; Saratoga, 1
+     (USBS); _6 mi. S, 13 mi. E Saratoga, 8500 ft._, 5; _6 mi. S, 14 mi.
+     E Saratoga, 8800 ft._, 1; Lake Marie, 10,440 ft., 2; _1 mi. S Lake
+     Marie_, 2; _1/2 mi. S, 2 mi. E Medicine Bow Peak, 10,800 ft._, 1;
+     Encampment (12 localities from 10 mi. N, 14 mi. E to 9 mi. N, 3 mi.
+     E Encampment and from 6500 to 8400 ft.), 63; 1/4 mi. N Riverside,
+     7380 ft., 2; S base Bridger Peak, 8800 ft., Sierra Madre Mountains,
+     1; _2 mi. S Bridger Peak, 9300 ft._, 2; Savery (10 localities from
+     8 mi. N, 21 mi. E to 4 mi. N, 8 mi. E Savery and from 7300 to 8800
+     ft.), 80. _Albany Co._: _30 mi. N, 10 mi. E Laramie, 6760 ft._, 6;
+     _29-3/4 mi. N, 9-1/2 mi. E Laramie, 6350 ft._, 1; 26 mi. N, 4-1/2
+     mi. E Laramie, 6960 ft., 8; _26-3/4 mi. N, 6-1/2 mi. E Laramie,
+     6700 ft._, 3; _3 mi. N, 13 mi. E Laramie, 7500 ft._, 1; _7 mi. N, 2
+     mi. E Laramie_, 1 (Chi); 5 mi. N Laramie, 7400 ft., 15; _Laramie_,
+     4 (AMNH); _1 mi. E Laramie, 7160 ft._, 4; 7-7/10 mi. SSW Laramie,
+     7200 ft., 4; 6-1/2 mi. S, 8-3/4 mi. E Laramie, 8200 ft., 1;
+     _Headquarters Park, 10,200 ft., Medicine Bow Mountains_, 3 (USBS);
+     Centennial, 8120 ft., 1; _2-1/4 mi. ESE Brown's Peak, 10,300 ft._,
+     3; _3 mi. ESE Brown's Peak, 10,000 ft._, 12; _2 mi. S Brown's Peak,
+     10,600 ft._, 1; _Pole Mountain, 15 mi. SE Laramie_, 4 (USBS 3); _1
+     mi. SSE Pole Mountain, 8350 ft._, 4; _2 mi. SW Pole Mountain, 8300
+     ft._, 13; _3 mi. S Pole Mountain, 8100 ft._, 1; Sherman, 2 (AMNH).
+     _Laramie Co._: 5 mi. N, 1 mi. W Horse Creek P. O., 7200 ft., 1;
+     Meadow, 2 (USBS); 11 mi. N, 5-1/2 mi. E Cheyenne, 5450 ft., 7; _7
+     mi. W Cheyenne, 6500 ft._, 10; Cheyenne, 3 (USNM).
+
+     COLORADO: _Moffat Co._: Lay, 6160 ft., 1 (AMNH). _Routt Co._:
+     Wright's Ranch, Yampa, 7700 ft., 2; Gore Range, 8 mi. E Toponas,
+     8000 ft., 2 (USBS). _Larimer Co._: _12-1/2 mi. W, 1-1/2 mi. S
+     Rustic_, 1; 11 mi. W, 1 mi. S Rustic, 1; Cache La Poudre River, 1
+     (Chi); _Estes Park_, 3 (USBS 1, AMNH 2); 19-1/2 mi. W, 2-1/2 mi. S
+     Loveland, 7280 ft., 6; _16 mi. W Loveland, 6840 ft._, 1; 6 mi. W,
+     1/2 mi. S Loveland, 5200 ft., 1. _Rio Blanco Co._: Meeker, 1
+     (USBS); _9-1/2 mi. SW Pagoda Peak_, 7700 ft., 3; 5 mi. S Pagoda
+     Peak, 9100 ft., 2. _Eagle Co._: Eagle, 1 (USBS); Pando, 2 (USBS).
+     _Grand Co._: Mt. Whiteley, 2 (USBS); Arapahoe Pass, Rabbit Ear
+     Mountains, 2 (USBS); Coulter (near Granby), 5 (USBS); _Arrowhead_
+     (near Dale), 1 (USBS). _Boulder Co._: 3/4 mi. N, 2 mi. W
+     Allenspark, 8400 ft., 4; _3 mi. S Ward_, 9000 ft., 3; Nederland, 16
+     (Chi). _Clear Creek Co._: Mt. McLellan, 2 (USBS); Berthoud Pass, 4.
+     _Park Co._: Trout Creek Ranch, 2 mi. N Garo, 1 (USBS).
+
+     Specimens examined of _M. m. nanus_ from eastern Idaho and Montana
+     are as follows: IDAHO: _Custer Co._: Challis, 7 (USBS); Mill Creek,
+     Challis Nat. Forest, 1 (USBS); Pahsimeroi Mts., 12 (USBS); Lost
+     River Mts., 1 (USBS). _Fremont Co._: N fork Snake River, 10 mi. SW
+     Island Park, 6200 ft., 2 (AMNH); Black Springs Creek, 4 mi. W
+     Ashton, 5200 ft., 1 (AMNH); 5 mi. W St. Anthony, 5000 ft., 1
+     (AMNH). _Camas Co._: Camas Prairie, Corral, 5100 ft., 2 (USBS).
+     _Blaine Co._: Alturas Lake, 3 (USBS); _Sawtooth Lake_, 2 (USBS);
+     Craters of the Moon, Laidlow Park, 2 (Mich); Ticura, 10 mi. S
+     Picabo, 1 (USBS); 19 mi. NE Carey (Lava Lake), 8 (Mich). _Butte
+     Co._: _26 mi. SW Arco_, 12 (Mich). _Bingham Co._: Shelley, 6
+     (USBS). _Bonneville Co._: 10 mi. SE Irwin, 4 (USBS). _Owyhee Co._:
+     Three Creeks, 3 (USBS). _Twin Falls Co._: Castleford Fenced Plot,
+     11 mi. W, 9 mi. S Twin Falls, 1. _Minidoka Co._: _Heyburn_, 2
+     (USBS). _Cassia Co._: 2 mi. S, 2 mi. W Burley, 5. _Bannock Co._:
+     Pocatello, 23 (USBS 4); Swan Lake, 1 (USBS). _Bear Lake Co._:
+     Montpelier Creek, 6700 ft., 3 (USBS). MONTANA: _Gallatin Co._: W.
+     Fork of W. Fork, Gallatin River, 1 (USBS). _Park Co._: Lamar River,
+     7000 ft., 1 (USBS); Gardiner, 1 (USBS). _Sweet Grass Co._: 14 mi. S
+     Big Timber, 1 (USBS); _McLeod_, 1 (USBS); West Boulder Creek, 18
+     mi. SE Livingston, 2 (USBS).
+
+
+=Microtus montanus codiensis=, new subspecies
+
+     _Type._--Female, adult, skin and skull; No. 27578, Museum of
+     Natural History, University of Kansas, from 3-1/5 mi. E and 3/5 mi.
+     S Cody, 5020 ft., Park Co., Wyoming; obtained on August 11, 1948,
+     by James W. Bee, original number 18-8-11-48.
+
+     _Range._--In northwestern Wyoming eastward from the Absaroka and
+     Wind River ranges into the western part of the Big Horn Basin.
+
+     _Diagnosis._--A relatively large _Microtus montanus_; tail actually
+     and relatively long; hind foot actually but not relatively large;
+     skull large; zygomatic expanse actually and relatively large;
+     alveolobasilar length relatively large; upper molar tooth-row
+     relatively long; color relatively light, not reddish.
+
+[Illustration: FIG. 1. Geographic range of _Microtus montanus_ in
+Wyoming, Colorado, and adjacent areas. The solid circles represent
+localities from which specimens have been examined; the hollow circles
+represent type localities. The ranges of subspecies in Utah are after
+Durrant, 1952.
+
+Guide to subspecies
+
+1. _M. m. nanus_
+2. _M. m. codiensis_
+3. _M. m. zygomaticus_
+4. _M. m. fusus_
+5. _M. m. micropus_
+6. _M. m. nexus_
+7. _M. m. amosus_
+8. _M. m. rivularis_
+]
+
+     _Comparisons._--As compared with the specimens of _M. m. nanus_
+     from Idaho, the size is larger (see diagnosis and measurements).
+     Certain proportions which differ from those of _nanus_ and which
+     are not in close agreement with the observed differences with age
+     in specimens of _nanus_ of a size comparable to _codiensis_ are
+     relatively large alveolobasilar length, relatively long alveolar
+     length of upper molar tooth-row, relatively wide-spreading
+     zygomatic arches, and relatively long tail. The color in
+     _codiensis_ is lighter than in _nanus_. As compared to the new
+     subspecies named below from the Big Horn Mountains to the east,
+     _codiensis_ is of similar size in head-body length, but has a
+     relatively as well as actually longer tail; the hind foot averages
+     longer; the upper molar tooth-row is relatively longer; the color
+     is slightly paler and less grizzled; the bullae are larger and less
+     flattened; the angle formed at the suture between the basioccipital
+     and basisphenoid bones is less acute; and the region of the suture
+     is less prominently elevated between the bullae when viewed from
+     the ventral aspect. The pterygoid plates mesial and posterodorsal
+     to the posterior end of the last upper molar are less fenestrated,
+     and the incisive foramina are less constricted posteriorly.
+
+[Illustration: FIG. 2. Map showing the major barriers to _Microtus
+montanus_ in Wyoming and Colorado; the barriers are the low areas named
+on the map (the name "Black Hills" is on the map for another reason;
+these hills are not a barrier). The major mountainous areas higher than
+approximately 8000 feet in elevation in Wyoming, Colorado and Utah are
+stippled. These mountainous areas include the habitat that is most
+suitable for the montane meadow mouse. The Black Hills are unoccupied by
+this species but these hills seem to be ecologically suitable for the
+species.]
+
+     _Measurements._--The average and the extremes for some measurements
+     of 34 males and females, 27 from the type locality and 7 from other
+     localities in the range assigned to this subspecies, are as
+     follows: total length, 165 (146-186); length of tail, 44.2 (35-55);
+     hind foot, 19.6 (17-21); condylobasilar length of the skull, 25.5
+     (24.0-27.5); zygomatic breadth, 15.6 (14.7-16.6); alveolar length
+     of upper molar tooth-row, 6.6 (6.2-7.0); prelambdoidal breadth, 8.8
+     (8.1-9.5); lambdoidal breadth, 12.0 (11.2-12.8). As an indication
+     of variability and for comparison with other series the coefficient
+     of variability and two times the standard error of the mean for
+     each measurement in this series are included in Table 1. The
+     averages for some measurements of 27 topotypes are as follows:
+     total length, 162; length of tail, 45.5; hind foot, 19.9;
+     condylobasilar length, 25.6; palatilar length, 14.0; molar series,
+     6.6; alveolobasilar length, 14.9; zygomatic breadth, 15.6;
+     interorbital breadth, 3.5; lambdoidal breadth, 12.1; prelambdoidal
+     breadth, 8.9; depth of braincase, 7.8.
+
+_Discussion._--Three species of _Microtus_ were collected by James W.
+Bee at the type locality. _Microtus montanus codiensis_, _Microtus
+longicaudus mordax_, and _Microtus pennsylvanicus modestus_ were taken
+in the same runways in the same meadow, at the same time. _Microtus
+ochrogaster haydeni_, although not taken at this locality, occurs in the
+Big Horn Basin. These four species differ in their geographic ranges,
+being largely allopatric, except _M. montanus_ and _M. longicaudus_
+which are sympatric. Although the different species have ecological
+preferences and habits which differ, several species of _Microtus_ may
+occur together in local areas such as the above. Certain of the
+characteristics of _M. m. codiensis_ are intermediate between those of
+the species _M. montanus_ on one hand and those of the other three
+species on the other hand. Could interspecific hybridization between
+"good species" of _Microtus_ take place in nature and possibly alter the
+characteristics of a local population?
+
+     _Specimens examined._--Total, 50, distributed as follows
+     (abbreviations for collections are given in the account of _M. m.
+     nanus_; localities that are not represented in Fig. 1 because
+     overlapping or crowding of the symbols would result are
+     Italicized):
+
+     MONTANA: _Carbon Co._: Beartooth Mountains, 2 (USBS); _Beartooth
+     Lake_, 1 (USBS).
+
+     WYOMING: _Park Co._: Black Mountain, head of Pat O'Hara Creek, 3
+     (USBS); 13 mi. N, 1 mi. E Cody, 5200 ft., 1; SW slope Whirlwind
+     Peak, 9000 ft., 1; _5 mi. N Cody, 6300 ft._, 1 (USBS); 3-1/5 mi. E,
+     3/5 mi. S Cody, 31; Ishawooa Creek, 6300 ft., 2 (USBS); _Valley_, 1
+     (USBS); Needle Mountain, 10,500 ft., 4 (USBS). _Hot Springs Co._: 3
+     mi. N, 10 mi. W Thermopolis, 4950 ft., 3.
+
+
+=Microtus montanus zygomaticus=, new subspecies
+
+     _Type._--Male, adult, skin and skull, No. 32761, Museum of Natural
+     History, University of Kansas, from Medicine Wheel Ranch, 9000 ft.,
+     28 mi. E Lovell, Big Horn County, Wyoming; obtained by R. Freiburg,
+     original number 105.
+
+     _Range._--The Big Horn Mountains of north-central Wyoming.
+
+     _Diagnosis._--A large _Microtus montanus_ with a relatively short
+     tail; short molar series; broad zygomatic arches well rounded in
+     lateral outline when viewed from above; small and flattened bullae;
+     raised basioccipito-basisphenoid suture.
+
+     _Comparisons._--For comparison with _M. m. codiensis_ from the
+     west, on the other side of the Big Horn Basin, see the account of
+     that subspecies. In comparison with _nanus_ this subspecies is
+     slightly paler, in this respect showing more resemblance to
+     _codiensis_ although not so pale, and more grizzled or unevenly
+     colored. This difference in color between _zygomaticus_ and
+     _codiensis_ may not be of taxonomic significance. From both the
+     topotypes of _nanus_, and the series of it from Wyoming,
+     _zygomaticus_ differs on the average in having a relatively shorter
+     tail, a relatively shorter upper molar tooth-row, relatively more
+     rounded and relatively more wide-spread zygomatic arches, and
+     smaller more flattened bullae.
+
+     _Measurements._--Average and extreme measurements of 24 adult males
+     and females from several localities here referred to _M. m.
+     zygomaticus_ are as follows: total length, 159(150-175); length of
+     tail, 37.6(31-46); hind foot, 18.6(17-20); condylobasilar length of
+     the skull, 25.3(24.2-26.7); zygomatic breadth, 15.3(14.1-16.7);
+     alveolar length of upper molar tooth-row, 6.2 (5.7-6.8);
+     prelambdoidal breadth, 8.7(8.3-9.4); lambdoidal breadth,
+     11.9(11.0-12.5). Average and extreme measurements of a series of 12
+     adult male topotypes are as follows: total length, 159(144-174);
+     length of tail, 36.4 (30-41); hind foot, 18.2(16-20);
+     condylobasilar length of skull, 25.8(24.7-26.7); alveolobasilar
+     length, 14.8(13.8-15.3); palatilar length, 13.8 (12.7-14.2);
+     alveolar length of upper molar tooth-row, 6.4(5.9-6.6); zygomatic
+     breadth, 15.9 (15.0-16.7); interorbital breadth, 3.6(3.4-3.7);
+     lambdoidal breadth, 12.1 (11.5-12.5); prelambdoidal breadth,
+     8.6(8.3-8.9); depth of braincase, 8.0 (7.6-8.3).
+
+_Discussion._--This subspecies is separated from _M. m. codiensis_ to
+the west by the Big Horn Basin. A series from along Buffalo Creek, 27
+mi. N, 1 mi. E Powder River, 6075 ft., in Natrona County, Wyoming, is
+intermediate between the topotypes of _zygomaticus_ and _nanus_ in the
+characters cited above as distinguishing the two, but shows greater
+resemblance to _zygomaticus_ in the shape of the zygomatic arch, in
+color which is paler than in topotypes of _zygomaticus_, and in the
+short hind foot. On these and on geographic grounds this population is
+referred to _zygomaticus_. Unfortunately we cannot be certain in many
+cases that an intermediate condition in a certain character indicates a
+genetically intermediate population and therefore true intergradation
+between the two subspecies to which the population is geographically
+intermediate. The topotypes of this subspecies are the most distinct of
+all the series which I have studied from the eastern Rocky Mountains, in
+terms of the degree of morphological departure from the norm for the
+species. After _zygomaticus_ the following populations are arranged
+according to their degree of deviation from this norm (_codiensis_
+deviates most): topotypes of _codiensis_, _fusus_ and a population from
+southern Sweetwater County, Wyoming, and lastly the _nanus-caryi_
+complex. Within the latter group, as I have mentioned, there are a
+number of local variants most of which do not differ significantly and
+do not conform to any geographic pattern.
+
+     _Specimens examined._--Total, 55, distributed as follows
+     (abbreviations for collections are given in the account of _M. m.
+     nanus_; localities that are not represented in Fig. 1 because
+     overlapping or crowding of the symbols would result are
+     Italicized): WYOMING: _Big Horn Co._: Medicine Wheel Ranch, 9000
+     ft., 28 mi. E Lovell, 30; W slope, head of Trappers Creek, 9500
+     ft., 2 (USBS). _Washakie Co._: 9 mi. E, 5 mi. N Tensleep, 7400 ft.,
+     1. _Johnson Co._: 7-1/2 mi. W, 1 mi. S Buffalo, 6500 ft., 3; Big
+     Horn Mountains, 3 (USBS). _Natrona Co._: Buffalo Creek, 27 mi. N, 1
+     mi. E Powder River, 6075 ft., 16.
+
+
+=Microtus montanus fusus= Hall
+
+     _Microtus nanus_, Bailey, N. Amer. Fauna, 17:30, June 6, 1900
+     (part); Cary, N. Amer. Fauna 33:123, August 17, 1911.
+
+     _Microtus montanus fusus_ Hall, Proc. Biol. Soc. Washington,
+     51:131-134, August 23, 1938; Warren, The Mammals of Colorado, Univ.
+     of Okla. Press, p. 229, 1942.
+
+     _Type._--Male, adult, skin and skull; No. 61281, Museum of
+     Vertebrate Zoology; 2-1/2 miles east of summit of Cochetopa Pass,
+     Saguache County, Colorado; Sept. 21, 1933; collected by Annie M.
+     Alexander; original number 2568. Type not seen by me.
+
+     _Range._--Southern Colorado and northern New Mexico.
+
+     _Comparisons._--For comparison with _M. m. nanus_, the subspecies
+     to the northward, see the preceding account of that subspecies. For
+     comparison with _M. m. amosus_ the subspecies to the west see Hall
+     (1938) and Durrant (1952). I have not examined specimens of
+     _amosus_.
+
+     _Measurements._--Average and extreme measurements for 17 adults
+     including both males and females from several localities in
+     southern Colorado are as follows: total length, 160 (136-179);
+     length of tail, 42 (35-55); hind foot, 19.2 (17-23); condylobasilar
+     length of the skull, 25.2 (24.0-26.0); zygomatic breadth, 15.0
+     (14.1-15.5); alveolar length of upper molar tooth-row, 6.4
+     (6.0-6.7); prelambdoidal breadth, 8.7 (8.3-9.2); lambdoidal
+     breadth, 11.7 (11.1-12.6).
+
+     Average and extreme measurements of 4 adults (2 males and 2
+     females) from the type locality and 11 adults (4 males and 7
+     females) from other localities in southern Colorado are as follows:
+     total length, 162 (157-168), 157 (137-169); length of tail (means
+     only), 44.5, 40.5; hind foot, 18.8 (18-19), 18.6 (18-23);
+     condylobasilar length of skull, 24.5 (24.0-24.7), 25.2 (24.3-26.1);
+     alveolobasilar length, 14.2 (13.9-14.5), 14.6 (14.1-15.1);
+     palatilar length, 13.2 (13.0-13.4), 13.5 (13.1-14.2); alveolar
+     length of upper molar tooth-row, 6.3 (6.0-6.6), 6.4 (6.3-6.7);
+     zygomatic breadth, 15.0 (14.3-15.5), 14.9 (14.1-15.5); interorbital
+     breadth, 3.5 (3.3-3.6), 3.5 (3.3-3.7); lambdoidal breadth, 11.8
+     (11.1-12.6), 11.7 (11.2-12.3); prelambdoidal breadth, 8.6
+     (8.3-9.2), 8.8 (8.3-9.0); depth of braincase, 7.5 (7.2-7.8), 7.6
+     (7.1-7.9).
+
+_Discussion._--There is no sharp boundary between _M. m. fusus_ of
+southern Colorado and the subspecies to the north, _M. m. nanus_.
+Although the line separating these two subspecies is drawn somewhat
+arbitrarily, on the whole the samples from north of this line more
+closely resemble _nanus_. All of the means for total length given above
+are larger than the maximum given in Hall's description of _fusus_. The
+caudal index (38 and 35% in two series) is slightly larger than that
+cited by Hall (33.3%) and is not significantly different from that in
+_nanus_ (35.2%). The color in both young and old mice is variable, but
+in general is more yellowish, and less grayish, than in any other series
+studied.
+
+There is a large area in western Colorado and eastern Utah, between the
+known ranges of _M. m. fusus_ and _M. m. amosus_ from which there are no
+specimens. Probably the species occurs only at certain places in this
+arid region which seems to be a partial barrier to the species.
+
+Specimens of _M. montanus_ from northern New Mexico have been referred
+previously to _M. m. arizonensis_. When he named _M. m. fusus_, Hall
+mentioned its resemblance to _arizonensis_ in reddish coloration, but
+pointed out that _fusus_ is less reddish. Of six specimens from Valle
+Santa Rosa, Jemez Mountains (USBS), 8500 ft., Rio Arriba County, New
+Mexico, three are immature, and the skulls of the remaining specimens
+are damaged. In reddish color and relatively large size these few
+specimens resemble _arizonensis_ more than _fusus_ although the locality
+of occurrence is closer to the geographic range of the northern _fusus_
+than to that of _arizonensis_. The identification of these specimens as
+_arizonensis_ is provisional; additional specimens are needed from the
+area, 200 miles wide, which separates the ranges as now known of
+_arizonensis_ in Arizona from the occurrence in New Mexico. There is a
+single specimen from this area, the damaged skull of which prevents
+conclusive identification. The specimen is either _M. montanus_ or _M.
+mexicanus_, and is from Nutria, on the southern edge of the Zuni
+Mountains (USBS). Detailed comparison of _fusus_ and _arizonensis_ is
+not attempted here although it may be stated that in several characters
+_fusus_ is intermediate between _arizonensis_ to the south and _nanus_
+to the north.
+
+     _Specimens examined._--Total, 89, distributed as follows
+     (abbreviations for collections are given in the account of _M. m.
+     nanus_; localities that are not represented in Fig. 1 because
+     overlapping or crowding of the symbols would result are
+     Italicized):
+
+     COLORADO: _Pitkin Co._: 5 mi. W Independence Pass, 11,000 ft., 1
+     (Chi). _Lake Co._: _Independence Pass, 12,095 ft._, 2 (Chi).
+     _Gunnison Co._: _Gothic_, 2 (USBS); Decker's Ranch, Crested Butte,
+     2 (AMNH); Almont, 3 (USBS). _Montrose Co._: Coventry, 5 (USBS 4,
+     AMNH 1). _Saguache Co._: Cochetopa Pass and environs, 44 (USBS
+     22). _Hinsdale Co._: Ruby Lake, 1 (USBS). _Mineral Co._: 3 mi. E
+     Creede, 1; 23 mi. S, 11 mi. E Creede, 9300 ft., 7. _La Plata Co._:
+     Florida, 6800 ft., 1. _Conejos Co._: 1 mi. S, 19 mi. W Antonito,
+     10,200 ft., 3; _4 mi. S, 23 mi. W Antonito_, 1; _5 mi. S, 24 mi. W
+     Antonito, 9600 ft._, 9.
+
+     NEW MEXICO: _Rio Arriba Co._: 6 mi. W Hopewell, 9900 ft., 6 (USBS);
+     _Tusas River, 8700 ft._, 1 (USBS).
+
+TABLE 1. AVERAGE MEASUREMENTS, IN MILLIMETERS, OF ADULTS OF MICROTUS
+MONTANUS.
+
+Key to column headings:
+
+A: No. of individuals averaged
+B: Total length
+C: Length of tail
+D: Length of hind foot
+E: Condylobasilar length
+F: Alveolar length of upper molar tooth-row
+G: Zygomatic breadth
+H: Lambdoidal breadth
+I: Prelambdoidal breadth
+
+=========================================================================
+       Locality   |   A  |  B  |  C  |  D  |  E  |  F  |  G  |  H  |  I
+------------------+------+-----+-----+-----+-----+-----+-----+-----+-----
+                  |               _M. m. codiensis_, all
+                  |
+Average           | 34   |165.3|44.2 |19.6 |25.47| 6.56|15.55|12.05|8.76
+2 x stand. error  |      | 3.56| 1.84| .395| .308| .067| .198| .144|.129
+Coeff. variab     |      |  6.0|11.6 | 5.6 | 3.5 | 3.0 | 3.65| 1.20|1.47
+------------------+------+-----+-----+-----+-----+-----+-----+-----+-----
+                  |            _M. m. nanus_, Eastern Idaho
+                  |
+Average           | 21[1]|151.1|39.4 |19.2 |25.00| 6.44|14.99|11.74|8.94
+2 x stand. error  |      | 3.20| 2.89| .293| .286| 1.15| .295| .210|.182
+Coeff. variab.    |      |  6.1|21.1 | 4.38| 2.49| 3.99| 4.10| 3.79|4.31
+------------------+------+-----+-----+-----+-----+-----+-----+-----+-----
+                  |            _M. m. nanus_, Wyoming
+                  |
+Teton Co.         | 35   |160.5|40.7 |18.6 |25.16| 6.51|15.17|11.86|8.77
+Fremont Co.       | 26   |157.0|41.4 |19.6 |25.23| 6.25|15.05|11.88|8.91
+Lincoln Co.       | 24   |159.9|41.8 |18.9 |25.08| 6.26|15.10|11.82|8.75
+Uinta Co.         | 26   |162.4|41.3 |19.0 |25.33| 6.42|15.31|12.16|8.89
+Sweetwater Co.    | 12   |159.8|43.7 |20.1 |24.98| 6.31|15.00|11.84|9.02
+Natrona Co.       | 40   |159.6|41.0 |19.6 |25.04| 6.40|15.00|11.84|8.93
+Carbon Co.        |108   |158.7|40.0 |19.1 |24.96| 6.27|15.05|11.83|8.72
+ Encampment [M]   | 27   |161  |41.7 |18.9 |25.1 | 6.16|15.2 |11.9 |8.7
+ Encampment [F]   | 11   |159  |41.1 |19.4 |24.9 | 6.18|14.9 |11.6 |8.6
+ Savery [M]       | 23   |159  |41.0 |19.2 |25.3 | 6.32|15.2 |12.2 |8.8
+ Savery [F]       | 25   |155  |37.1 |18.8 |24.7 | 6.33|14.8 |11.6 |8.6
+------------------+------+-----+-----+-----+-----+-----+-----+-----+-----
+                  |                     _M. m. nanus_
+                  |
+Northern Colo.    |  8   |163.1|42.4 |19.6 |25.20| 6.44|14.86|11.70|8.56
+------------------+------+-----+-----+-----+-----+-----+-----+-----+-----
+                  |                     _M. m. fusus_
+                  |
+Southern Colo.    | 17[2]|159.8|42.4 |19.2 |24.97| 6.43|14.98|11.73|8.69
+------------------+------+-----+-----+-----+-----+-----+-----+-----+-----
+
+[Footnote 1: For external parts, 34 individuals were used.]
+
+[Footnote 2: For external parts, 29 individuals were used.]
+
+Some measurements not given above are included in Table 1, together with
+the number of specimens and the sex if restricted to one sex. So that
+the variability can be evaluated more adequately, the coefficient of
+variability and 2 times the standard error of the mean are included for
+the measurements in two series. The series consist of all the adult
+specimens (with a condylobasilar length of 24.0 mm. or more) of both
+sexes from the areas specified. Various barriers are shown in Fig. 2 for
+comparison with the distributions of the subspecies and the localities
+of known occurrence shown in Fig. 1. _Microtus montanus_ has not been
+taken in the Black Hills area of extreme northeastern Wyoming. Suitable
+montane habitat is present and both _Microtus pennsylvanicus insperatus_
+and _Microtus longicaudus longicaudus_ occur there. The arid basin of
+the Powder River presumably is a barrier that has prevented _M.
+montanus_ from reaching this area.
+
+
+REFERENCES CITED
+
+BAILEY, V.
+
+     1900. Revision of American voles of the genus _Microtus_. N. Amer.
+     Fauna, 17:1-88, June 6.
+
+     1917. A new subspecies of meadow mouse from Wyoming. Proc. Biol.
+     Soc. Washington, 30:29-30, February 21.
+
+CARY, M.
+
+     1911. A biological survey of Colorado. N. Amer. Fauna, 33:1-256,
+     August 17.
+
+     1917. Life zone investigations in Wyoming. N. Amer. Fauna, 42:1-95,
+     October 3.
+
+DAVIS, W. B.
+
+     1939. The Recent mammals of Idaho. 400 p., front., illus., maps,
+     diagrs., Caldwell, Id., The Caxton Printers Ltd., April 5.
+
+DURRANT, S. D.
+
+     1952. Mammals of Utah, taxonomy and distribution. Univ. Kans.
+     Publ., Mus. Nat. Hist., 6:1-549, 91 figs, in text, 30 tables,
+     August 10.
+
+ELTON, C.
+
+     1942. Voles, mice and lemmings; problems in population dynamics.
+     496 p., illus. (maps), tables, Oxford, The Clarendon Press, London.
+
+GOIN, O. B.
+
+     1943. A study of individual variation in _Microtus pennsylvanicus
+     pennsylvanicus_. Jour. Mamm., 24:212-224, June 7.
+
+HALL, E. R.
+
+     1938. Notes on the meadow mice _Microtus montanus_ and _Microtus
+     nanus_ with descriptions of a new subspecies from Colorado. Proc.
+     Biol. Soc. Washington, 51:131-134, August 23.
+
+HILL, E. A.
+
+     1935. Cranial foramina in rodents. Jour. Mamm. 16(2):121-129.
+
+HOWELL, A. B.
+
+     1924. Individual and age variation in _Microtus montanus yosemite_.
+     Jour. Agr. Res., 28(10):977-1015, June 7.
+
+KELLOGG, R.
+
+     1922. A study of the California forms of the _Microtus montanus_
+     group of meadow mice. Univ. California Publ. Zool., 21:245-274,
+     April 18.
+
+MERRIAM, C. H.
+
+     1891. Results of a biological reconnaissance of south-central
+     Idaho. N. Amer. Fauna, 5:1-113, July 30.
+
+PIPER, S. E.
+
+     1909. The Nevada mouse plague of 1907-8. U. S. Dept. Agr., Farmers'
+     Bul. 352, pp. 1-23, 9 figs.
+
+WARREN, E. R.
+
+     1942. The mammals of Colorado, their habits and distribution.
+     Second (revised) edition, Univ. Oklahoma Press, Norman, xviii-330
+     p., front., 50 plates.
+
+
+_Transmitted March 22, 1954._
+
+
+
+[SQUARE]
+
+25-3560
+
+
+       *       *       *       *       *
+
+Transcriber's Notes
+
+Italic typeface in the original is indicated by by _underscores_.
+
+Bold typeface in the original is indicated by =equals=.
+
+The symbols for [Male] and [Female] are indicated by [M] and [F].
+
+One typographical error was corrected: "Castlford" was corrected to
+"Castleford" in "Twin Falls Co.: Castleford Fenced Plot".
+
+
+
+
+
+End of the Project Gutenberg EBook of Subspeciation in the Meadow Mouse,
+Microtus montanus, in Wyoming and Colorado, by Sydney Anderson
+
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