vy fv -u o HARVARD UNIVERSITY M R LIBRARY OF THE Museum of Comparative Zoology '^ ri" L,fcH/tM^^'*t^^J^ University of Kansas Publications MUSEUM OF natural HISTORY /-J VOLUME 9 • 1955-1960 EDITORS E. Raymond Hall, Chairman Henry S. Fitch A. Byron Leonard Robert W. Wilson Museum of Natural History UNIVERSITY OF KANSAS LAWRENCE 1960 Museum of Natural History university of kansas lawrence '"'" "'1'*? ZOBl k.LiurtK! V ^^c; -7]960 {fvARS PRINTED IN THE STATE PRINTING PLANT TOPEKA, KANSAS I960 28-4057 Volume 9 CONTENTS 1. Speciation of the wandering shrew. By James S. Findley. Pp. 1-68, 18 figures in text. December 10, 1955. 2. Additional records and extensions of ranges of mammals from Utah. By Stephen D. Durrant, M. Raymond Lee, and Richard M. Hansen. Pp. 69-80. December 10, 1955. 3. A new long-eared myotis (Myotis evotis) from northeastern Mexico. By Rollin H. Baker and Howard J. Stains. Pp. 81-84. December 10, 1955. 4. Subspeciation in the meadow mouse, Microtus pennsylvanicus, in Wyoming, Colorado, and adjacent areas. By Sydney Anderson. Pp. 85-104, 2 fig- ures in text. May 10, 1956. 5. The condylarth genus Elfipsodon. By Robert W. Wilson. Pp. 105-116, 6 figures in text. May 19, 1956. 6. Additional remains of the multituberculate genus Eucosmodon. By Robert W. Wilson. Pp. 117-123, 3 figures in text. May 19, 1956. 7. Mammals of Coahuila, Mexico. By Rollin H. Baker. Pp. 125-335, 75 figures in text. June 15, 1956. 8. Comments on the taxonomic status of Apodemus peninsulae, with descrip- tion of a new subspecies from North China. By J. Knox Jones, Jr. Pp. 337-346, 1 figure in text, 1 table. August 15, 1956. 9. Extensions of known ranges of Mexican bats. By Sydney Anderson. Pp. 347-351. August 15, 1956. 10. A new bat (Genus Leptonycteris ) from Coahuila. By Howard J. Stains. Pp. 353-356. January 21, 1957. 11. A new species of pocket gopher (Genus Pappogeomys) from Jalisco, Me- xico. By Robert J. Russell. Pp. 357-361. January 21, 1957. 12. Geographic variation in the pocket gopher, Thomomys bottae, in Colorado. By Phillip M. Youngman. Pp. 363-384, 7 figures in text, 1 table. Feb- ruary 21, 1958. 13. A new bog lemming (Genus Synaptomys) from Nebraska. By J. Knox Jones, Jr. Pp. 385-388. May 12, 1958. 14. Pleistocene bats from San Josecito Cave, Nuevo Leon, Mexico. By J. Knox Jones, Jr. Pp. 389-396. December 19, 1958. 15. New subspecies of the rodent Baiomys from Central America. By Robert L. Packard. Pp. 397-404. December 19, 1958. 16. Mammals of the Grand Mesa, Colorado. By Sydney Anderson. Pp. 405- 414, 1 figure in text. May 20, 1959. 17. Distribution, variation, and relationships of the montane vole, Microtus montanus. By Sidney Anderson. Pp. 415-511, 12 figures in text, 2 tables. August 1, 1959. 18. Conspecificity of two pocket mice, Perognathus goldmani and P. artus. By E. Ravmond Hall and Marilyn Bailev Ogilvie. Pp. 513-518, 1 map. January 14, 1960. 19. Records of harvest mice, Reithrodontomys, from Central America, with description of a new subspecies from Nicaragua. By Sydney Anderson and J. Knox Jones, Jr. Pp. 519-529. January 14, 1960. 20. Small carnivores from San Josecito Cave (Pleistocene), Nuevo Leon, Me- xico. By E. Raymond Hall. Pp. 531-538, 1 figure in text. January 14, 1960. 21. Pleistocene pocket gophers from San Josecito Cave, Nuevo Leon, Mexico. By Robert J. Russell. Pp. 539-548, 1 figure in text. January 14, 1960. 22. Review of the insectivores of Korea. By J. Knox Jones, Jr., and David H. Johnson. Pp. 549-578. February 23, 1960. 23. Speciation and evolution of the pygmv mice, genus Baiomys. By Robert L. Packard. Pp. 579-670, 4 plates, 12 figures in text. June 16, 1960. Index. Pp. 671-690. ' *T' ' (,■ University of Kansas Publications Museum of Natural History Volume 9, No. 1, pp. 1-68, figures 1-18 December 10, 1955 Speciation of the Wandering Shrew BY JAMES S. FINDLEY FEB, 5 1956 ...,...i„ , University of Kansas Lawrence 1955 UNIVERSITY OF KANSAS PUBLICATIONS. MUSEUM OF NATURAL HISTORY Institutional libraries interested in publications exchange may obtain this series by addressing the Exchange Librarian, University of Kansas Library, Lawrence, Kansas. Copies for individuals, persons working in a particular field of study, may be obtained by addressing instead the Museum of Natural History, University of Kansas, Lawrence, Kansas. There is no provision for sale of this series by the University Library which meets institutional requests, or by the Museum of Natural History which meets the requests of individuals. However, when individuals request copies from the Museum, 25 cents should be included, for each separate number that is 100 pages or more in length, for the purpose of defraying the costs of wrapping and mailing. * An asterisk designates those numbers of which the Museum's supply (not the Li- brary's supply) is exhausted. Numbers published to date, in this series, are as follows: Vol. 1, Nos. 1-26 and index. Pp. 1-638, 1946-1950. *Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140 figures in text. April 9, 1948. Vol. 3. *1, The avifauna of Micronesia, its origin, evolution, and distribution. By Rol- lin H. Baker. Pp. 1-359, 16 figures in text. June 12, 1951. *2. A quantitative study of the nocturnal migration of birds. By George H. Lowery, Jr. Pp. 361-472, 47 figures in text. June 29, 1951. 3. Phylogeny of the waxwings and allied birds. By M. Dale Arvey. Pp. 473- 530, 49 figures in text, 13 tables. October 10, 1951. 4. Birds from the state of Veracruz, Mexico. By George H. Lowery, Jr. and Walter W. Dalquest. Pp. 531-649, 7 figures in text, 2 tables. October 10. 1951. Index. Pp. 651-681. •Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466. 41 plates, 31 figures in text. December 27, 1951. Vol. 5. 1. Preliminary survey of a Pal eocene faunule from the Angels Peak area. New Mexico. By Robert W. Wilson. Pp. 1-11, 1 figure in text. February 24, 1951. 2. Two new moles (Genus Scalopus) from Mexico and Texas. By RoUin H. Baker. Pp. 17-24. February 28, 1951. 3. Two new pocket gophers from Wyoming and Colorado. By E. Raymond Hall and H. Gordon Montague. Pp. 25-32. February 28, 1951. 4. Mammals obtained by Dr. Curt von Wedel from the barrier beach of Tamaulipas, Mexico. By E. Raymond Hall. Pp. 33-47, 1 figure in text. October 1. 1951. 5. Comments on the taxonomy and geographic distribution of some North Ameri- can rnbbits. By E. Raymond Hall and Keith R. Kelson. Pp. 49-58. October 1, 195L 6. Two new subspecies of Thomomys bottae from New Mexico and Colorado. By Keith R. Kelson. Pp. 59-71, 1 figure in text. October 1, 1951. 7. , A new subspecies of Microtus montanus from Montana and comments/on Microtus canicaudus Miller. By E. Raymond Hall and Keith R. Kelson. Pp. 73-79. '^October 1, 1951. 8. A new pocket gopher (Genus Thomomys) from eastern Colorado. By E. Raymond HaU. Pp. 81-85. October 1, 1951. 9. Mammals taken along the Alaskan Highway. By RoUin H. Baker. Pp. 87- 117, 1 figure in text. November 28, 1951. *10. A synopsis of the North American Lagomorpha. By E. Raymond Hall. Pp. 119-202, 68 figru-es in text. December 15, 1951. 11. A new pocket mouse (Genus Perognathus) from Kansas. By E. Lendell Cockrum. Pp. 203-206. December 15. 1951. 12. Mammals from Tamaulipas, Mexico. By Rollin H. Baker. Pp. 207-218. December 15. 1951. ' 13. A new pocket gopher (Genus Thomomys) from Wyoming and Colorado. By E. Raymond Hall. Pp. 219-222. December 15, 1951. 14. A new name for the Mexican red bat. By E. Raymond Hall. Pp. St23-226. December 15, 1951. 15. Taxonomic notes on Mexican bats of the Genus RhogeSssa. By E. Raymond Hall. Pp. 227-232. AprU 10, 1952. 16. Comments on the taxonomy and geographic distribution of some North Ameri- can woodrats (Genus Neotoma). By Keith R. Kekon. Pp. 233-242. April 10. 1952. (Continued on inside of back cover) University of Kansas Publications Museum of Natural History Volume 9, No. 1, pp. 1-68, figures 1-18 December 10, 1955 Speciation of the Wandering Shrew BY JAMES S. FINDLEY University of Kansas Law^rence 1955 University of Kansas Publications, Museum of Natural History Editors: E. Raymond Hall, Chairman, A. Byron Leonard, Robert W. Wilson Volume 9, No. 1, pp. 1-68, figures 1-18 Published December 10, 1955 University of Kansas Lawrence, Kansas MOS. CO:,!P. ZOOL i: ' P£B 9 1955 ?••• J PRINTED BY FERD VOILAND. JR. . STATE PRINTER TOPEKA, KANSAS 1955 25-7903 Speciation of the Wandering Shrew BY JAMES S. FINDLEY CONTENTS PAGE Introduction 4 Materials Methods and Acknowledgments 4 Non-geographic Variation 7 Characters of Taxonomic Worth 8 Pelage Change 9 Geographic Distribution and Variation 9 Pacific Coastal Section 9 Inland Montane Section 11 Great Basin and Columbia Plateau Section 12 Summary of Geographic Variation 13 Origin of the Sorex vagrans Rassenkreis 16 Relationships With Other Species 26 Conclusions 60 Table of Measurements 62 Literature Cited 66 FIGURES Figs. 1-2. — Cranial Measurements 5 Fig. 3. — Graph Illustrating Wear of Teeth 8 Fig. 4. — Graph Illustrating Heterogonic Growth of Rostrum 10 Fig. 5. — Present Geographic Distribution of Sorex vagrans 15 Fig. 6. — Skulls of Sorex vagrans 17 Figs. 7-10. — Past Geographic Distribution of Shrews 19-20-22- 27 Figs. 11, 12. — Medial View of Lower Jaws of Two Shrews 30 Figs. 13, 14. — Second Unicuspid Teeth of Shrews 30 Fig. 15. — Diagram of Probable Phylogeny of Shrews 32 Figs. 16-18. — Geograpihc Distribution of Subspecies 33-40- 53. (3) 4 University of Kansas Publs., Mus. Nat. Hist. INTRODUCTION The purpose of this report is to make clear the biological rela- tionships between the shrews of the Sorex vagrans-obscurus "species group." This group as defined by H. H. T. Jackson (1928:101) included the species Sorex vagrans, S. obscurus, S. pacificus, S. yaquinae, and S. durangae. The last mentioned species has been shown (Findley, 1955:617) to belong to another species group. Sorex milleri, also assigned to this group by Jackson (1947:131), seems to have its afiinities with the cinereus group as will be ex- plained beyond. The position of the vagrans group in relationship to other members of the genus will be discussed. Of this group, the species that was named first was Sorex vagrans Baird, 1858. Subsequently many other names were based on mem- bers of the group and these names were excellently organized by Jackson in his 1928 revision of the genus. Subsequent students of western mammals, nevertheless, have been puzzled by such prob- lems as the relationship of (1) Sorex vagrans monticola to Sorex obscurus obscurus in the Rocky Mountains, (2) Sorex pacificus, S. yaquinae, and S. obscurus to one another on the Pacific Coast, and (3) S. o. obscurus to S. t;. amoenus in California. Few studies have been made of these relationships. Clothier (1950) studied S. v. monticola and S. o. obscurus in western Montana and concluded that the two supposed kinds actually were not separable in that area. Durrant (1952:33) was able to separate the two kinds in Utah as was Hall (1946:119, 122) in Nevada. Other mammalogists who worked within the range of the vagrans-obscurus groups have avoided the problems in one way or another. Recently Rudd ( 1953 ) has examined the relationships of S. vagrans to S. ornatus. MATERIALS METHODS AND ACKNOWLEDGMENTS Approximately 3,465 museum study skins and skulls were studied. Most of these were assembled at the University of Kansas Museum of Natural History, tut some were examined in other institutions. Specimens were grouped by geographic origin, age, and sex. Studies of the role of age and sex in variation were made. Because it was discovered that secondary sexual variation was negligible, both males and females, if of like age and pelage, were used in comparisons designed to reveal geographic variation. Ex-temal measurements used were total length, length of tail, and length of hind foot. After studying a number of cranial dimensions I chose those listed below as the most useful in showing differences in size and proportions of the skull. Figures 1 and 2 show the points between which those measurements ■were taken. Speciation of the Wandering Shrew 5 Condylobasal length. — From anteriormost projection of the premaxillae to pos- teriormost projection of the occipital condyles (a to a')- Maxillary tooth-row. — From posteriormost extension of M3 to anteriormost ex- tension of first unicuspid (b to b'). , Palatal length. — From anteriormost projection of premaxillae to posteriormost part of bony palate (c to c'). Cranial breadth. — Greatest lateral diameter of braincase (d to d')- Least interorhital breadth. — Distance between medialmost superior edges of orbital fossae, measured between points immediately above and behind posterior openings of infraorbital foramina (e to e'). Maxillary breadth. — Distance between lateral tips of maxillary processes (f to f). r^f f'-^ Figs. 1 and 2. Showing where certain cranial measurements were taken. X 3/2. ( Based on Sorex vagrans obscurus, from Stonehouse Creek, 5/2 mi., W junction of Stonehouse Creek and Kelsall River, British Columbia, 9 , 28545 KU. ) In descriptions of color, capitalized terms refer to those in Ridgway (1912). In addition the numerical and alphabetical designations of these terms are given since a knowledge of the arrangements of these designations enables one quickly to evaluate differences between stated colors. Color terms which are not capitalized do not refer to any precise standard of color nomenclature. In the accounts of subspecies, descriptions, unless otherwise noted, are of first year animals as herein defined. Descriptions of color are based on fresh pelages. 6 University of Kansas Publs., Mus, Nat. Hist. Unless otherwise indicated, specimens are in the University of Kansas Museum of Natural History. Those in other collections are identified by tlie following abbreviations : AMNH American Museum of Natural History CM Carnegie Museum ChM Chicago Museum of Natural History CMNH . . Cleveland Museum of Natural History FC Collection of James S. Findley HC Collection of Robert Holdenreid SGJ Collection of Stanley G. Jewett CDS Collection of Charles D. Snow AW Collection of Alex Walker NMC National Museum of Canada OSC Oregon State College PMBC British Columbia Provincial Museum of Natural History SD San Diego Natural History Museum BS United States Biological Surveys Collection USNM United States National Museum UM . ... University of Michigan Museum of Zoology OU .... University of Oregon Museum of Natural History UU University of Utah Museum of Zoology WSC Washington State College, Charles R. Conner Museum In nature, the subspecies of Sorex vagrans form a cline and are distributed geographically in a chain which is bent back upon itself. The subspecies in the following accounts are listed in order from the southwestern end of the chain clockwise back to the zone of overlap. The synonymy of each subspecies includes the earliest available name and other names in chronological order. These include the first usage of the name combination employed by me and other name combinations that have been applied to the subspecies concerned. In the lists of specimens examined, localities are arranged first by state or province. These are listed in tiers from north to south and in any given tier from west to east. Within a given state, locahties are grouped by counties, which are listed in the same geographic sequence as were the states and provinces ( N to S and W to E ) . Within a given county, locahties are arrangr^d from north to south. If two or more localities are at the same latitude the westernmost is Hsted first. Marginal localities are fisted in a separate paragraph at the end of each account. The northernmost marginal locality is listed first and the rest follow in clockwise order. Those records followed by a citation to an authority are of specimens which I have not personally examined. Marginal records are shown by dots on the range maps. Marginal records which cannot be shown on the maps because of undue crowding are listed in Italic type. To persons in charge of the collections listed above I am deeply indebted. Without their generous cooperation in allowing me to examine specimens in their care this study would not have been possible. Appreciated suggestions in the course of the work have been received from Professors Rollin H. Baker, A. Byron Leonard, R. C. Moore, Robert W. Wilson, and H. B. TordoflF, and many of my fellow students. Mr. Victor Hogg gave helpful suggestions on the preparation of the illustrations. My wife, Muriel Findley, devoted many hours Speciation of the Wandering Shrew 7 to secretarial work and typing of manuscript. Finally I am grateful to Professor E. Raymond Hall for guidance in the study and for assistance in preparing the manuscript. During the course of the study I received support from the University of Kansas Endowment Association, from the Office of Naval Research, and from the National Science Foundation. NON-GEOGRAPHIC VARIATION Non-geographic variation, that is to say, variation within a single population of shrews, consists of variation owing to age and normal individual variation. In Sorex I have detected no significant sec- ondary sexual differences between males and females; accordingly the two sexes are here considered together. Variation with age must be considered in order to assemble comparable samples of these shrews. Increased age results in wear on all teeth and in particularly striking changes in the size and shape of the first incisors. Skulls of older shrews develop sagittal and lambdoidal ridges, and further differ from skulls of young animals in being slightly broader and shorter, and in developing thicker bone, particularly on the rostrum which thus seems to be, but is not always in fact, more robust. Pruitt has recently (1954) noted these same cranial differences in specimens of Sorex cinereiis of different ages. Several students of American shrews, notably Pearson (1945) on Blarina, Hamilton (1940) on Sorex ftimeus, and Conaway (1952) on Sorex palustris, have shown that young are born in spring and summer, usually reach sexual maturity the following spring, and rarely survive through, or even to, a second winter. The result is that collections made, as most of them are, in spring and summer, contain two age classes, first year and second year animals. These two age classes are readily separable on the basis of differences in the skull as well as on the decreased pubescence of the tail and the increased weight of second year animals. My own examination of hundreds of museum specimens confirms this for the Sorex vagrans group. Separation of the two age classes in an August-taken series of Sorex vagrans from coastal Washington is shown in figure 3, in which two tooth-measurements that are dependent upon wear are plotted against one another. First year animals are more abundant in collections than are second year animals. Within the first year, that is to say from spring to late fall, animals vary but little. Dental characters are best studied in first year shews. For this reason I have used them as the basis for the study of geographic variation, and descriptions are based on first year animals unless otherwise noted. 8 University of Kansas Publs., Mus. Nat. Hist. CHARACTERS OF TAXONOMIC WORTH Within the Sorex vagrans complex, the only characters of taxo- nomic significance that I have detected are in size and color. It is true that cranial proportions, such as relative size of rostrum, may change from population to population, but these proportions seem to me to be dependent upon actual size of the individual shrew as I shall elsewhere point out. Of the cranial measurements here Protoconid Height of ml Fig. 3. Two measurements (in millimeters) reflecting tooth-wear plotted against one another. First year and second year individuals of Sorex vagrans vagrans, all taken in August at Willapa Bay, Washington, are com- pletely separated. Open circles represent teetli of second year shrews; solid circles represent teeth of first year shews. employed, palatal length and least interorbital breadth are the most significant and useful. Color in the S. vagrans group seems to be in Orange and Cadmium Yellow, colors 15 and 17 of Ridgway (1912). No specimens actually possess these pure colors, but most colors in these shrews are seen to be derived from the two mentioned by admixture of black and/or neutral gray. In color designations an increase in neutral gray is indicated by an increased number of prime signs ('), whereas increase in black is indicated by progressive characters of the Roman alphabet (i, k, m). Thus, IT'k is grayer than 17'k and 17"m is blacker than 17"k. In subspecific diagnoses in this report, color and size, and sometimes relative size, are the characters usually mentioned. Speciation of the Wandering Shrew 9 PELAGE CHANGE In general, winter pelage is darker than summer pelage in these shrews. Winter pelage comes in first on the rump and spreads caudad and ventrad. The growth line of incoming hair is easily detected on the fur side of the skin. Throughout the winter the color of the pelage changes, often becoming somewhat browner, although no actual molt takes place. This was noted by Dalquest (1944) who assumed that the color change resulted from molt although he was unable to detect actual replacement of hairs. Summer pelage usually comes in first on the back or head and moves posteriorly and laterally. Time of molt depends on latitude and altitude. Summer pelage may appear fairly late in the season and may account for the anomalous midsummer molt noted by Dalquest. Fresh pelages of summer and winter are best seen in first year animals and are less variable than are worn pelages and hence are used as the basis of color descriptions. GEOGRAPHIC DISTRIBUTION AND VARIATION Pacific Coastal Section The largest shrews of the vagrans group ( large in all dimensions ) occur in the coastal forests of northern California and of Oregon. Those shrews are reddish, large-skulled, large-toothed, and have rostra that are large in proportion to the size of the skull as a whole. The very largest of these shrews live along the coast of northwestern California. To the southward they are somewhat smaller, and at suc- cessively more northern localities, to as far as southwestern British Columbia, they are likewise progressively smaller and also some- what less reddish. The relative size of the rostrum decreases with the decrease in size of the skull; consequently smaller shrews have relatively smaller rostra (see fig. 4). In addition the zygomatic ridge of the squamosal decreases in relative size with decrease in actual size of the skull. Thus, these features change in a clitial fashion as one proceeds from, say, Humboldt County, California, northward to Astoria, Oregon. Turning our attention now farther inland to the Cascade Moun- tains of northern Oregon, the shrews there also are smaller and less reddish (more brownish) than in northwestern California, and the trend to smaller and darker shrews culminates in the northern Cascades of Washington. Shrews from there, and from the south- western coast of British Columbia, compared with those from northwestern California, are much smaller and have so great a suf- fusion of black that they appear brown rather than red. At places 10 University of Kansas Publs., Mus. Nat. Hist. 44 •c c» £43 >J "5 ■ pacificus #- # yoQuinae • bairdi • longicauda • setosus • vagrans 41 2 (£ 40 16 17 18 19 20 21 22 Condylobasal Length Fig. 4. Condylobasal length (in millimeters) plotted against palatal index ( palatal length/condylobasal length X 100 ) in several subspecies of Sorex vagrans to show relative increase in size of rostrum with actual increase in size of skull. along the coast successively farther north of southwestern British Columbia the shrews become larger again, the largest individuals being those from near Wrangell, Alaska. From that place north- westerly along the coast of Alaska, size decreases again. The shrews so far discussed inhabit forests in a region of high rainfall and a minimum of seasonal fluctuation in temperature. Such a habitat seems to be the optimum for shrews of the vagrans group since the largest individuals are found there. In addition, shrews seem to be as common, or commoner, in this coastal belt, than they are in other places. The large shrews of the vagrans group on the Pacific coast were divided into three species by H. H. T. Jackson in his revision of the North American Sorex in 1928. The large reddish shrews of the coast of California and southern Oregon were called S. pacificus. The somewhat smaller ones from the coast of central Oregon were called S. yaquinae. Still smaller shrews from northwestern Oregon and from the rest of the Pacific coast north into Alaska were called S. obscurus. I find these kinds to intergrade continuously one with the next in the manner described and conclude that all are of a single species. Speciation of the Wandering Shrew 11 Inland Montane Section Inland from the coasts of British Columbia and Alaska the size of the vagrans shrew decreases rapidly. Specimens from western Alaska, central Alaska, and the interior of British Columbia are uniformly smaller than coastal specimens. In addition the red of the hair is masked more by neutral gray than by black with the result that the pelage is grayish rather than brownish or reddish. Shrews of this general appearance are found southward through the Rocky Mountain chain to Colorado and New Mexico. On the more or less isolated mountain ranges of Montana east of the continental divide the vagrans shrew is somewhat smaller still. On the Sacra- mento Mountains of southeastern New Mexico the shrew is some- what larger and slightly darker. Southwestward from the Colorado Rockies this shrew becomes smaller and slightly more reddish (less grayish ) . All of these montane populations of the vagrans shrew are com- monest in hydrosere communities, that is to say, streamsides and marshy areas where the predominant vegetation is grass, sedges, willows, and alders. Since these animals are less common within the montane forests, hydrosere communities, rather than the actual forest, seem to be the positive feature important for the shrews. The shrews of the montane region just described were regarded by Jackson as belonging to two species: Sorex ohscurus, occupying all the Rocky Mountains south to, and including, the Sacramento Mountains; S. vagrans, made up of small individuals from various places in Wyoming, Montana, and Colorado, and all the shrews of western New Mexico and all of Arizona. My study of these animals has led me to conclude that the smaller shrews of Arizona and New Mexico intergrade in a clinal fashion with the shrews of Colorado and in fact represent but one species. Since some individuals from Colorado are as small as larger individuals from this southwestern population of small animals, I conclude that such specimens are the basis for reports of S. vagrans from Colorado. The shrews of the Sacramento Mountains resemble those of the Colorado Rockies more than they do the smaller shrews of western New Mexico and Arizona, possibly because the climate is similar in the Sacramento Mountains and the higher Colorado Rockies. There is less precipita- tion in the more western mountain ranges in New Mexico and in Arizona in April, May, and June than in the Colorado Rockies. These months are critical for the reproduction and growth of shrews. 12 University of Kansas Publs., Mus. Nat. Hist. As mentioned above, the shrews from east of the continental divide in Montana are smaller than those of the other mountains of the state, and it is upon such small animals that the name Sorex vagrans has been based in this area. It is clear, however, that these smaller animals intergrade with the larger shrews of the more western mountains. The small size might be an adaptation to the lesser precipitation and harsher continental climate east of the continental divide in Montana. Great Basin and Columbia Plateau Section The vagrant shrews of the Great Basin and adjoining Columbia Plateau and Snake River Plains are smaller than their relatives in the Rocky Mountains and, by virtue of less gray in their pelage, are reddish in summer and blackish rather than grayish in winter. There is little significant geographic variation in shrews throughout this region, although owing to their restriction to the vicinity of water, the populations of shrews are more or less isolated from one another and each is somewhat different from the next. Those from nearest the Rockies are sometimes slightly larger and those from some places in Nevada are slightly paler than the average. This small reddish shrew is found all the way to the Pacific coast of California, Oregon, and Washington. In these coastal areas it is somewhat darker and sometimes a trifle larger than elsewhere. It intergrades with a somewhat larger, grayer shrew in the Sierra Nevada of California. Along the Wasatch front in Utah, this Great Basin shrew intergrades with the larger, grayer shrew of the Rockies. Owing to the abrupt change in elevation, the zone of intergradation is rather narrow horizontally. In the latitude of Salt Lake City, populations of intergrades occur at between 8,700 and 9,000 feet elevation. The lowland shrew occurs in the eastern part of the Snake River Plains, and along the valleys of the Bear and Salt rivers into Wyoming. Along the northern edge of the Snake River Plains and on the western edge of the mountains of central Idaho the transition from lowland to montane habitats is abrupt and in consequence the zone of contact between small and large shrews is narrow. In northern Idaho and northwestern Montana the transition from low- land to highland is more gradual. Tributaries of the Columbia River system, especially the Clark Fork, provide a path for move- ment of lowland forms into intermontane basins of western Montana. In addition, the vegetational zones are found at lower elevations, and there are boreal forests in the lowlands rather than only in Speciation of the Wandering Shrew 13 the mountains as is the case in Utah and Colorado. In this area, therefore, the zone of intergradation between the smaller lowland shrew and the larger montane shrew is more gradual and gradually intergrading populations are found over a relatively large area. This has been well demonstrated for northwestern Montana by Clothier ( 1950 ) . In southern British Columbia and northern Wash- ington this shrew in the mountains is large and in the intermontane valleys is small. There is extensive interdigitation of valleys and mountain ranges, and, consequently, of Hfe-zones in this region. In a few places, recognizably distinct populations of the vagrant shrew occur within a few miles of one another, but in other places there are populations of intergrades. West of the Cascades no evidence of intergradation has been found and the two kinds occur almost side by side and maintain their distinctness. These Great Basin shrews dwell in hydrosere communities as do their Rocky Mountain counterparts. In this arid region such a habitat obviously is the only one habitable for a shrew of the vagrans group. These shrews often maintain their predeliction for such habitats when they reach the Pacific coast, and are commonly found in such places as coastal marshes, marshy meadows, and streamsides, while the woodlands are inhabited by other species. These small shrews of the Great Basin and the small vagrant shrews of the Pacific Coast were called Sorex vagrans by Jackson. Summary of Geographic Variation Large reddish shrews of the coast of California and southwestern Oregon become smaller and darker to the north. From southwest- ern British Columbia they again become larger as one proceeds northward along the coast to Wrangell, Alaska, and north of that they again become smaller. Moving inland from the coast the shrews become markedly smaller in Alaska and British Columbia. The smaller inland and montane form occurs south through the Rocky Mountains, becoming slightly smaller in central Montana, slightly larger in southeastern New Mexico, and slightly smaller in western New Mexico and in Arizona. This montane form inter- grades with a smaller more reddish Great Basin shrew, the zone of intergradation roughly following the western slope of the Rocky Mountains. The Great Basin shrew occurs westward to the Pacific Coast; there the Great Basin shrew occurs with, although in part it is ecologically separated from, the large reddish coastal shrews. There seems to be an intergrading chain of subspecies of one 14 University of Kansas Publs., Mus. Nat. Hist. species, the end members of which (the small Great Basin form and the large coastal form) are so different in size and ecological niche that they are able to coexist without interbreeding. In south- ern British Columbia the morphological differences are not so marked as farther south along the Pacific Coast. There, in British Columbia, reproductive isolation is not complete and occasional populations of intergrades occur. In Montana extensive intergrada- tion occurs in a broad zone of transitional habitat. Along the west- ern edge of the Rockies from Idaho south to Utah the zone of transition from montane to basin habitat is sharp and the zone of intergradation, although present, is fairly narrow, perhaps because there is little intermediate habitat which logically might be expected to be most suitable for intergrading populations. The oldest name applied to a shrew of the group under consid- eration is Sorex vagrans Baird, 1858, the type locality of which is Willapa Bay, Pacific County, Washington. The name apphes to the small vagrant shrew of this area, rather than to the larger forest dweller which has been known as Sorex ohscunis. The name S. vagrans, in the specific sense, must therefore apply to all the shrews discussed which have heretofore been known by the names S. pacif- iciis, S. yaqiiinae, S. ohscunis, and S. vagrans. A situation such as the one here described where well differenti- ated end members of a chain of subspecies overlap over an extensive geographic range throughout the year without interbreeding — thus reacting toward one another as do full species — so far as I know has not previously been found to exist in mammals. The overlap- ping end-members of the chain of subspecies of Sorex vagrans really do coexist; specimens of the overlapping subspecies have been taken together at the same localities from California to British Columbia. I have taken a specimen of S. t;. vagrans and several of S. V. setostis in the same woodlot at Fort Lewis, Pierce County, Washington. Two subspecies of deer, Odocoileus hemionus, in the Sierra Nevada of California, occur together over a sizeable area but for only a part of each year that does not include the breeding season (Cowan, 1936:156-157). In the deer mouse, Peromyscus maniculatus, the geographic ranges of several pairs of subspecies meet at certain places without intergradation of the two kinds. In these instances well marked ecological differences exist between the subspecies involved. In western Washington, for example, the geographic range of the lowland subspecies, P. m. austeriis, inter- digitates to the east and west with the range of tlie montane and coniferous forest-inhabiting subspecies, P. m. oreas, and the two Speciation of the Wandering Shrew 15 120 100 Other kinds of S.vagrans S.v.vagrans and derivaiives scale of miles 120 100 Fig. 5. Probable present geographic distribution of Sorex vagrans. The range of S. v. vagrans and its derivatives S. v. vancouverensis, S. v. hali- coetes, and S. v. paludivagus, is shown by hnes slanting in a different direc- tion than those vi^hich mark the range of all the other subspecies of S. vagrans. The region in which S. v. vagrans occurs together v^ath other subspecies of S. vagrans is shown by the superposition of one pattern upon the other. 16 University of Kansas Publs., Mus. Nat. Hist. kinds have not been shown to intergrade. Peromyscus maniculatus artemesiae and P. m. osgoodi come together without interbreeding in Glacier National Park, Montana. P. m. artemesiae is almost en- tirely a forest-dwelling subspecies, whereas osgoodi is an inhabitant of open country. The two kinds do not actually occur together ecologically although they occur together in buildings at the edge of the woods (A. Murie, 1933:4-5). Cases of sympatric existence of two subspecies of one species are known in birds and in reptiles. Notable examples are in the gull, Larus argentatus (Mayr, 1940), in the Old World warbler, Phyl- loscopus trochiloides (Ticehurst, 1938), and in the great titmouse, Parus major ( Rensch, 1933 ) , of the Old World. In the first species the two end-members, the herring gull and the lesser black-backed gull, occur together over an extensive region from northern Europe and the British Isles throughout Fennoscandia. Fitch (1940) de- scribed a rassenkreis with overlapping subspecies in the garter snake Thamnophis ordinoides. The geographic distribution of the species Sorex vagrans is shown in figure 5. The geographic range of the Great Basin subspecies is shown by a different pattern of lines than the other subspecies of S. vagrans. In the region in which the greographic range of the Great Basin subspecies overlaps those of the subspecies of the Pacific Coast, the pattern of shading for the Great Basin subspecies is superimposed on the patterns for the other subspecies. ORIGIN OF THE SOREX VAGRANS RASSENKREIS The distribution of the species Sorex vagrans and that of its im- mediate ancestors obviously has not always been tlie same; during glacial ages much of the present range of the species in Canada and in some of the higher mountains of tlie United States was covered with ice and not available to the shrew. Furthermore, large areas that are now too hot and dry to permit the existence of S. vagrans were at one time habitable. If we "are to speculate on the manner in which the Sorex vagrans rassenkreis originated we must inquire into the nature and extent of these climatic changes. The most recent epoch of geological time, the Pleistocene, is known to have been divided into a series of alternating glacial and interglacial ages. During the glacial ages continental and montane glaciers are judged to have covered much of Canada and the north- ern United States. Concurrently the major storm tracks of the west probably were shifted southward; in any event much of the now arid intermontane west was much better watered than it is today. Speciation of the Wandering Shrew 17 Figs. 6a-6f. Fig. 6a. Sorex vagrans pacificus, 1 mi. N Trinidad, Humboldt Co., California, FC 1442. Fig. 6b. S. v. yaquinae, Newport, Lincoln Co., Oregon, AW 707. Fig. 6c. S. v. yaquinae (near hairdi), McKenzie Bridge, Lane Co., Oregon, AW 82. Fig. 6d. S. v. setosus. Reflection Lake, Jeffer- son Co., Washington, CMNH 4275. Fig. 6e. S. v. obscurus, 10 mi. SSW Leadore, Lemhi Co., Idaho, FC 1499. Fig. 6/. S. v. vagrans. Baker Creek, White Pine Co., Nevada, 88042 (after Hall, 1946:113). The increased precipitation, and probally glacial meltwater, formed large lakes in the closed basins of the Great Basin. There were boreal forests at lower elevations than there are today in comparable latitudes and continuous boreal habitat probably connected many of the isolated mountain ranges of the southwest. That probability is supported by the presence of boreal animals and plants on many 2—7903 18 UNivERSiTi' OF Kansas Publs., Mus. Nat. Hist. of these isolated ranges today. A boreal tree squirrel, such as Tamiasciurus, could hardly be suspected of crossing a treeless, inter- montane desert valley, miles wide. Interglacial ages were characterized by warmth and aridity as compared to the glacial ages. Glaciers retreated or disappeared, boreal forests became montane in much of the United States, and tlie lakes in the Great Basin were reduced or disappeared. One can envision that during such times boreal mammals were isolated, theii- geographic ranges were restricted, and Sonoran mammals ex- panded their ranges. Evidence is more extensive concerning the number and extent of glacial ages in the eastern than in the western part of North America. This evidence suggests a division of the Pleistocene into four glacial ages and four interglacial ages, the fourth interglacial age corre- sponding to the present time. More information is available about the Wisconsinan, or last, glacial age, than about the earlier ones, because the last glaciation in many montane areas destroyed evi- dence of earlier glaciations. The names of currently recognized glacial and interglacial ages of the Pleistocene are listed below. The names of interglacial ages are in Italic type. Wisconsinan Sangamonian lUinoian Yarmouthian Kansan Aftonian Nebraskan We may think of these ages as an alternating series of cool moist and warm dry periods during which boreal mammals, and other organisms, alternately moved southward ( disapjDearing in the glaci- ated regions) and northward into previously glaciated areas (while disappearing from southern areas except on isolated mountain ranges). Sorex vagrans probably followed this pattern of move- ment and now is restricted to forested or well-watered places. One possible series of events culminating in the formation of the Sorex vagrans rassenkreis may be thought of as having begun during the Illinoian age. With much of Canada, and perhaps also many areas in the Rockies, Cascades, and the Sierra Nevada covered with glacial ice, the shrew-stock ancestral to Sorex vagrans may well have occupied a more or less continuous range over the Colorado Plateau, the Columbian Plateau, the Great Basin, and in the forests of the Pacific Coast ( as well as over part of eastern United States, as will Speciation of the Wandering Shrew 19 be explained beyond; see fig. 7 ) . At that time the species probably was a continuously interbreeding unit. In the ensuing Sangamonian interglacial age all glaciers retreated or disappeared thereby opening up extensive areas in the north and in the higher mountains which were occupied by a boreal fauna^ including S. vagrans. Concurrently the Great Basin, and probably also much of the Columbian Plateau, became dry, and desert con- ditions developed, perhaps much as they are today. Increasing aridity eliminated shrew habitat in most places between the Rocky Mountains and the Sierra Nevada-Cascade mountain chain with the result that the geographic range of the species resembled an inverted "U", one arm lying along the Rocky Mountains and the other along the Cascade-Sierra Nevada axis; the connection between the two arms was in British Columbia (see fig. 7). At present Sorex vagrans: .??==q>^ Fig. 7. Possible distribution in lUinoian (inset) and Sangamonian times of the ancestor of the Sorex vagrans-ornatus-longirostris-veraepacis complex. Approxi- mate southern boundary of Illinoian glaciation marked by heavy line. 20 University of Kansas Publs., Mus. Nat. Hist. does occur in isolated places in the Great Basin, but its existence there is tenuous and seemingly dependent upon the occurrence of permanent water such as Ruby Lake and Reese River. With such an arrangement as this it can readily be seen that gene flow between the eastern and western arms of the "U" would be greatly reduced by distance; consequently differentiation between the two might be expected. Wisconsinan glaciation again rendered Canada uninhabitable, and it is quite possible that extensive areas in the Rocky Mountains, the Cascades and the Sierra Nevada were heavily glaciated. With the elimination of the northern part of the "U", the eastern and Fig. 8. Possible distribution of Sorex vagrans at two different times in the Wisconsinan Age. Left, early Wisconsinan; right, mid-Wisconsinan. Speciation of the Wandering Shrew 21 western arms became isolated, if not by the width of the Columbian Plateau at least by the glaciated Cascade Mountains. At the same time extensive areas on the Colorado Plateau and much of the area south to the Mexican highlands were again occupied by the species. Finally the Great Basin, again being well-watered, provided suit- able habitat for, and was reoccupied by, Sorex vagrans ( see fig. 8 ) . This reoccupation of the Great Basin took place probably from the Colorado Plateau and mountains of Arizona and Utah, since the present day shrews of the species S. vagrans in the Great Basin closely resemble Rocky Mountain shrews but differ markedly from the large endemic subspecies of the Pacific Coast. Finally, with the waning of Wisconsinan ice, the species again was able to occupy northern and montane areas as it had during Sangamonian times. Again dessication of the Great Basin caused drastic restriction of shrew habitat. The small, marsh-dwelling kind of wandering shrew which had developed there around the lakes of Wisconsinan time occupied suitable habitat all the way to the Pacific coast where its range came into contact with that of the west- ern arm of the Sangamonian "U."-pattern of shrew distribution (see fig. 9). The animals of this western segment and the new arrivals from the east were by this time so different from one another that the two kinds lived in the same areas without interbreeding. The descendants of the original western arm now are known as Sorex vagrans sonomae, S. v. pacificus, S. v. yaqiiinae, and S. t;. bairdi. The newcomers from the east are known as S. v. vagrans, S. v. halicoetes, S. v. paliidivagus and S. v. Vancouver ensis. In addition to occupying the Pacific Coast from San Francisco Bay north to the Fraser Delta, the Great Basin subspecies populated the Columbia Plateau and the western foothills of the central and northern Rockies. By so doing that subspecies came into secondary contact with its own parent stock with which it was still in repro- ductive continuity in Utah. In some places in British Columbia dif- ferentiation between the two kinds had proceeded to such an extent that some reproductive isolation was effected, but in many other places the two interbred. The Rocky Mountain form spread north and west and occupied the Cascades and coastal lowlands in south- western British Columbia and in Washington. Here the differentia- tion between the Rocky Mountain subspecies and the Great Basin subspecies was great enough to cause complete reproductive isola- tion. Deglaciation of the Sierra Nevada opened it up for reoccupation 22 University of Kansas Publs., Mus. Nat. Hist. 120 ICXD 30 600 =1 scole of miles Fig. 9. Probable changes in the distribution of Sorex vagrans concurrent with and following the dissipation of Wisconsinan ice. Dark arrows in Washington, Idaho, Oregon, and California, shows S. v. vagrans. Speciation of the Wandering Shrew 23 from the east by Sorex vagrans of the Great Basin. In response to the montane environment the subspecies ohscuroides, resembling the subspecies ohscurus of the Rockies, developed. Desiccation of the intermontane parts of New Mexico, Arizona, and Chihuahua, left "marooned" populations of Sorex vagrans on suitable mountain ranges. In this way Sorex vagrans orizabae may have been isolated in southern Mexico. The isolated populations of Arizona and New Mexico differentiated in situ into the subspecies monticola and neomexicanus. Western Canada and Alaska were populated by shrews which originated in the habitable parts of the Rocky Mountains and Colorado Plateau during Wisconsinan time (as opposed to shrews originating, as subspecies, in the Great Basin or on the Pacific Coast). These shrews differentiated into the currently recognized subspecies of the west coast and coastal islands of British Columbia and Alaska in response to the different environments in these places, many of which were isolated; the subspecies isolatus, mixtus, setosus, longicauda, elassodon, prevostensis, malitiosiis, and alaskensis are thought to have originated in this fashion after the areas now oc- cupied by them were freed of Wisconsinan ice. This group of shrews from the Rocky Mountains probably came into contact with the Pacific coastal segment of the species some- where in northwestern Oregon. The clinal decrease in size from S. V. pacificus to S. v. setosus seems steepest in this area. Upon the establishment of this contact reproductive continuity was resumed, probably because the temporal separation of the two stocks in- volved was not so great as, say, that between S. v. vagrans and S. v. pacificus, and in addition the morphological differentiation was not so great. On the eastern side of the Rockies the montane stock moved northeastward, occupying suitable territory opened up by the dis- solution of the Laurentide ice sheet. Still later changes in the character of the northern plains owing to desiccation divided the range of the species and isolated S. v. soperi in Manitoba and central Saskatchewan and a population of S. v. ohscurus in the Cypress Hills. A number of semi-isolated stocks in central Montana became differentiated as a recognizable subspecies there. A number of other boreal mammals have geographic ranges which resemble that of Sorex vagrans, except that the geographic ranges of subspecies do not overlap. Because of the general simi- 24 University of Kansas Publs., Mus. Nat. Hist. larities of these geographic ranges, it is pertinent to examine the reasons suggested by students to account for the present geographic distributions of some of these other boreal species. The red squirrel genus, Tamiasciurus, has a Rocky Mountain ( and northern coniferous forest) species, T. hudsonicus, that occurs all along the Rocky Mountain chain and northward into Alaska. In the Cascade Mountains of Washington and British Columbia this species meets the range of a well marked western species, T. dotig- lasii, with no evidence of intergradation. Dalquest (1948:86) attributes the divergence of the two species to separation in a glacial age but feels that the degree of difiFerence bet^veen the two is too great to have all taken place during the Wisconsinan. Perhaps he has overemphasized the importance of the differences between the two, but, be that as it may, it seems that the two kinds differen- tiated during a glacial age when they were isolated, perhaps by ice on the Cascades into a coastal population and an inland population. One difference between the distribution of the red squirrels and vagrant shrew is that the squirrel of the Sierra Nevada is the species of the Pacific Coast, whereas the vagrant shrew of the Sierra Nevada was derived from the Great Basin population, which in turn was derived from tlie Rocky Mountain kind. Red squirrels do not occur on any of the boreal montane "islands" of Nevada. During the pluvial periods when hydrosere-loving shrews populated the Great Basin, that region may have been a treeless grassland. Vagrant shrews, then as now, probably depended on hydrosere communities, while red squirrels required trees. Therefore the shrews were able to traverse the Great Basin, while the Sierran red squirrels were of necessity derived from the coastal population. The ecological requirements of jumping mice, genus Zapus, and the subspecies of Sorex vagrans that dwell in hydroseres are essen- tially similar. The species Zapus princeps lives in the Rocky Mountains, the Great Basin, the Sierra Nevada, and north to Yukon (Krutzsch, 1954:395). Its geographic range is similar to that of the montane and basin segments of S. vagrans. The species Z. tri- notatus occurs along the Pacific coast and in the Cascades north to southwestern British Columbia. Its distribution thus coincides in general with that of the large red coastal subspecies of S. vagrans. Krutzsch (1954:368-369) thought that these two kinds of jumping mice were first separated by the formation of the Cascade Moun- tains and the Sierra Nevada and finally by Pleistocene glaciation. The Sierran jumping mouse (Zapus princeps), as is the Sierran vagrant shrew, is more closely related to the jumping mouse of the Speciation of the Wandering Shrew 25 Great Basin and of the Rocky Mountains than it is to the jumping mouse ( Z. trinotatus ) of the Pacific Coast, just as the Sierran vagrant shrew is related to the shrew of the Great Basin and Rocky Moun- tains. The jumping mouse also is limited in its distribution by hydrosere communities, not by forests. In western North America there are two species of water or marsh shrews: Sorex palustris and S. hendiri. They have been placed in separate subgenera, but, as pointed out beyond, are closely related and here are placed in the same subgenus. The species palustris is found throughout the Rocky Mountains, north into Alaska, across the Great Basin into the Sierra Nevada, and west to the Pacific coast in Washington. The species hendiri is found from northwest- ern California north along the Pacific coast to southwestern British Columbia and east to the Cascades. Where the ranges of the two species overlap in western Washington they do not interbreed so far as is known, and are somewhat diflFerent in their ecology, hendiri being a lowland, and palustris being a montane, species. The two species probably were separated in a glacial period as seems to have been the case with the wandering shrews. Also, the water shrew of the Sierra Nevada is derived from that of the Great Basin and Rocky Mountains. Sorex palustris is tied closely in its distribution to hydro- sere communities and is not dependent upon the presence of forests. Red-backed mice, genus Clethrionomys, occur throughout the Rocky Mountains and west to the Cascades in Washington as the species C. gapperi. The species C. californicus is found along the Pacific Coast from California north to the Olympic Peninsula. Where the ranges of the two species meet in Washington they seem not to intergrade. In some glacial interval these two species may have evolved in the same manner as has been described for the species of Zapus and those of Tamiasciurus. No Clethrionomys are found in the Sierra Nevada, nor are red-backed mice found in the boreal islands of the Great Basin. It is not known why Clethrionomys californicus does not occur in the Sierra Nevada. Some boreal birds have distributional patterns similar to those of the mammalian examples cited above. One kind of sapsucker, Sphyrapicus varius nuchalis, occurs in the Rocky Mountains north into British Columbia and west to the Cascades and Sierra Nevada. A related kind, S. varius ruher, occurs along the Pacific Coast from California north into British Columbia. Recently Howell (1952) has shown that some intergradation takes place between ruher and nuchalis in Washington and British Columbia, although they do not intergrade 26 University of Kansas Publs., Mus. Nat. Hist. freely. Previously the two kinds were thought not to intergrade and were regarded as two species. The two kinds intergrade also in northeastern California, although in that state S. v. daggeti, rather than S. t;. ruber, is involved in the intergradation. Howell considered the two kinds to be conspecific with one another as well as with the eastern S. variiis. He attributed a measure of the distinctness of nuchalis and ruber to their separation during a glacial period, but felt that the separation was much older than Wisconsinan. What- ever the time of separation, the pattern seems clear: nuchalis and ruber (as well as varius) were separated into montane, coastal, and eastern segments respectively, probably by glaciation (it seems to me in the Pleistocene), and have since re-established contact with one another. The grouse genus Dendrogapus is divided into a Great Basin species, D. obscurus, which extends northward into British Colum- bia, and a Rocky Mountain species, D. fuliginosus, that is found in the Sierra Nevada and northward along the coast and Cascades into British Columbia. Although the two kinds have at times been considered conspecific, they diflFer in voice, hooting mechanism, and characters of the downy young, and so far no actual intergradation between the two has been shown (Grinnell and Miller, 1944:113). These grouse thus seem to offer additional evi- dence for a Pleistocene, possibly Wisconsinan, separation of the boreal fauna into a Rocky Mountain and a Pacific coastal segment. A notable sidelight on these data is the frequency with which species in the Sierra Nevada have their closest relatives in the Rocky Mountains, rather than in the geographically nearer Cascades or coastal areas. This similarity in fauna of the Sierra Nevada and the Rockies was noted long ago by Merriam (1899:86). RELATIONSHIPS WITH OTHER SPECIES During the Sangamonian interval, isolated segments of the once wide-spread ancestral Sorex vagrans quite possibly persisted in such places as the Sierra Nevada, coastal southern California, the moun- tains of Arizona, New Mexico, and southern Mexico, and in the Black Hills (see fig. 6). One might expect that by Wisconsinan time these populations would have become reproductively isolated from their parent stock. They would therefore have remained specifically distinct when Wisconsinan Sorex vagrans, reoccupied these outlying areas, and may still be found isolated in places peripheral to the range of the ancestral species. In fact, we do find species closely related to Sorex vagrans in just Speciation of the Wandering Shrew 27 such places today (fig. 10). Probably Sorex ornatus, including members of tlie ornatus group such as S. trigonirostris, S. sintiosus, S. willeti, S. tcnellus, and S. nanus, and also S. veraepacis, arose by separation from the ancestral vagrans stock in Sangamonian time. Probably the eastern S. longirostris arose in a like manner. The ancestor of S. ornatus may have been isolated in southwestern Cali- fornia during Sangamonian time, spread north and south during the Wisconsinan age, and afterward given rise to S. trigonirostris and Fig. 10. Probable distribution of S. veraepacis, S. longirostris, and the S. ornatus group (stipple) and of their Wisconsinan ancestors (lines). Heavy line indicates limits of Wisconsinan glaciation. the modern S, ornatus complex of California and Baja California. In at least one place reproductive isolation between ornatus and the invading S. vagrans has broken down (Rudd, 1953); the place is a salt marsh along San Pablo Bay, where a hybrid population between S. vagrans and S. sinuosus, an ornatus derivative, has formed. Sorex tenellus may have been isolated in the Sierra Nevada in the Sanga- 28 University of Kansas Publs., Mus. Nat. Hist. monian interval, moved into the valleys east of the mountains during the Wisconsinan age, and become restricted to its present range since the retreat of the last ice. Sorex nanus may have occurred in the Black Hills and isolated mountains of Arizona and New Mexico during the Sangamonian interval and remained in these general areas during the Wisconsinan age. Its present range is peripheral to the main body of the Rockies and the Colorado Plateau. The eastern species Sorex longirostris has many similarities with shrews of the ornattis-va grans stock. S. /. longirostris is close in many ways to S. nanus. Indeed, the differences between the species S. nanus, S. ornatus, and S. longirostris seem to me to be of the same magnitude and indicate a similar period of differentiation from a common ancestor. The ancestor of S. longirostris may have gained access to the eastern United States in the Illinoian Age via the north- ern Great Plains south of the glacial boundary (fig. 7). The an- cestor of Sorex veraepacis of southern Mexico probably reached that area in Illinoian time as part of the ancestral vagrans stock and prob- ably attained its differentiation during the Sangamonian interval. All the kinds of shrews so far discussed, including the S. vagrans complex, might thus be thought of as having had a common ancestor in the Illinoian Age. This entire group of shrews has the third uni- cuspid smaller than the fourth, a pigmented ridge from the apex to the cingulum of each upper unicuspid, and, in most individuals, lacks a post-mandibular foramen in the lower jaw (Findley, 1953: 636-637). The pigment is not always prominent in S. longirostris. Two other species of North American shrews, Sorex palustris, the water shrew, and Sorex bendiri, the marsh shrew, show these three characters to a greater or lesser degree, and it seems that these two species and the vagrans-ornatus-veraepacis group had a common ancestor, probably before Illinoian time for reasons stated beyond. I judge, however, that far from being subgenerically distinct as they have been considered to be, S. palustris and S. bendiri are actually closely related species of the same subgenus and may have differen- tiated from one another because of separation into eastern (palus- tris) and western (bendiri) segments in the Sangamonian interval, much as has been postulated concerning the eastern and western stocks of Sorex vagrans. Indeed, Jackson (1928:192) has noted that in the Pacific northwest the characters of the two kinds approach one another and become differences of degree only. The widespread species Sorex cinereus resembles all the foregoing species in the ridges on the unicuspid teeth and in the lack of a post-mandibular foramen, but differs from those other species in Speciation of the Wandering Shrew 29 having the third upper unicuspid larger than the fourth. The sub- species S. cinereus ohionensis, however, often has the sizes of these teeth reversed. With S. cinereus I inckide S. preblei (eastern Ore- gon) and S. lijelli (Sierra Nevada), both obviously closely related to cinereus as Jackson (1928:37) recognized when he included them in the cinereus group. Sorex milleri ( Coahuila and central western Nuevo Leon) seems to me to resemble S. cinereus more than it does other species of North American Sorex, and I judge tliat it also belongs to tlie cinereus group. Sorex cinereus and its close relatives seem more closely related to the species which have thus far been discussed than they do to such other North American species as S. arcticus, S. fumeus, S. trowbridgi, S. merriami, and the members of the S. saussurei group; most of these five species last mentioned possess a post-mandibular foramen, lack pigmented unicuspid ridges, and have the third unicuspid larger than the fourth. Be- cause of the morphological resemblences mentioned above, it seems likely to me that S. cinereus and the vagrans-ornatus-veraepacis-pa- lustris complex had a common ancestor in early Pleistocene time. Sorex cinereus has recently been considered to be conspecific with the Old World S. caecutiens Laxmann (Van den Brink, 1953) which name, being the older, would apply to the circumpolar species. Hibbard (1944:719) recovered S. cinereus and a species of Neosorex (a name formerly applied to the water shrew) from the Pleistocene (late Kansan) Cudahy Fauna. This indicates that the ancestors of the modern S. cinereus and of the water shrew had diverged from one another before that time. Brown (1908:172) recorded S. cinereus and S. ohscurus from the Conard Fissure in Arkansas. These materials were deposited probably at a later time than was the Cudahy Fauna. The S, ohscurus from Conard Fissure probably represents the ancestral S. va grans stock which I think reached eastern United States in Ilhnoian time and gave rise to S. longirostris. The Conard Fissure material was deposited at a time (Illinoian?) when northern faunas extended farther south than they do today. All of the species mentioned as having structural characters in common with S. vagrans seem to have arisen from a common ances- tor which had already differentiated from the ancestor of such species as S. arcticus, S. saussurei, and others. Consequently all are here included in a single subgenus. The oldest generic name apphed to a shrew of this group, other than the name Sorex, is Otisorex DeKay, 1842, type species Otisorex platyrhinus DeKay, a 30 University of Kansas Publs., Mus. Nat, Hist. synonym of Sorex cinereus. The subgenus can be characterized as follows. Subgenus Otisorex DeKay 1842. Otisorex DeKay, Zoology of New York, pt. 1, Mammalia, p. 22, and pi. 5, fig. 1. Type, Otisorex platyrhinus DeKay (= Sorex cinereus Kerr). Third unicuspid usually smaller than fourth; upper unicuspids usually with pigmented ridge extending from apices medially to cingula, uninterrupted by antero-posterior groove; post-mandibular foramen usually absent. Includes the species S. cinereus, S. longi- rostris, S. vagrans, S. ornatus, S. tenellus, S. trigonirostris, S. nanus, S. juncensis, S. willeti, S. sinuosus, S. veraepacis, S. palustris, S. bendiri, S. alaskanus, and S. pribilofensis. 11 12 13 14 Figs. 11-14. Characters of the svibgenera Sorex and Otisorex. Fig. 11. Medial view of right ramus of Sorex (Otisorex) vagrans. X 14. Fig. 12. Medial view of right ramus of Sorex {Sorex) arcticus. X 14. Fig. 13. Anterior view of left second upper unicuspid of Sorex {Otisorex) vagrans. X 45. Fig. 14. Anterior view of left second upper unicuspid of Sorex {Sorex) arcticus. X 45. Speciation of the Wandering Shrew 31 Other species of Sorex now occurring in North America difiFer from Otisorex in having the 3rd imicuspid usually larger than 4th, in lack- ing a pigmented ridge from the apices to the cingula of the upper unicuspids, and in usually possessing a well-developed post-mandi- bular foramen. Exceptions to the last mentioned character aie S. fwnens and S. clispar. The subgenus Sorex in North America should include only the following species: S. jacksoni, S. tundrensis, S. arcticus, S. gaspensis, S. dispar, S. fumeiis, S. troivhridgii, S. merriami, and all tlie members of the Mexican S. saussurei group. The subgenera Otisorex and Sorex probably separated in early Pleistocene or late Pliocene. Sorex is unknown in North America earlier than tlie late Pliocene (Simpson, 1945:51). In the genus Microsorex the characters of tlie subgenus Otisorex are carried to an extreme; the unicuspid ridges are prominent and end in distinct cusplets, and the 3rd unicuspid is not merely smaller than the 4tli, but is reduced almost to the vanishing point. In addi- tion, the post-mandibular foramen is absent. Although it is closer structurally to Otisorex than to Sorex, the recognition of Microsorex as a distinct genus seems warranted. Figure 15 is intended to represent graphically some of the rela- tionships discussed above. It must be re-emphasized that much of it is purely speculative, especially as regards actual time when various separations took place. It will be noted that I have indi- cated most separations as having taken place in interglacial ages. They are generally regarded as periods of warmth and aridity and, therefore, probably are times of segmentation of the ranges of boreal mammals and hence times exceptionally favorable to the process of speciation. Glacial ages, characterized by extensive and continuous areas of boreal habitat, probably were times of relatively unre- stricted gene flow between many populations of boreal mammals and hence not favorable to rapid speciation. Sorex vagrans Wandering Shrew The size of the v;^andering shrew varies from small in the subspecies monticola and vagrans to large in the subspecies pacificiis. The tail makes up from a little more than a third to almost half of the total length. The color pattern ranges from tricolored through bicolored to almost monocolored. Color ranges from reddish ( Sayal or Snuff Brown) to grayish in summer pelage and from black to light gray in winter. Diagnostic dental characters include: 3rd upper unicuspid smaller than 4th, and unicuspids, except 5th, with a pigmented ridge extending from near apex of each tooth medially to cingulum and sometimes ending as internal cusplet. S. vagrans differs from mem- 32 University of Kansas Publs., Mus. Nat. Hist. Fig. 15. Diagrammatic representation of the probable phylogcny of Sorex vagrans and its near relatives. bers of the ornatus group in less flattened skull, and in more ven- trally situated foramen magnum that encroaches more on the basi- occipital and less on the supraoccipital. The wandering shrew dif- fers from S. trowbridgii and S. saussurei in the dental characters mentioned above. These dental characters also serve to distinguish S. vagrans readily from S. cinereus, S. merriami, and S. arcticus which may occur with vagrans. The large marsh shrew and water shrew, S. palustris and S. bendiri, can be distinguished at a glance from S. vagrans by larger size and darker color. In the following treatment of the 29 subspecies of Sorex vagrans, the subspecies are arranged in geographic sequence, beginning with the southernmost large subspecies on the California coast and pro- ceeding clockwise, north, east, south, and then west back to the starting point. Sorex vagrans sonomae Jackson Sorex pacificus sonomae Jackson, Jour. Mamm., 2:162, August 19, 1921. Type. — Adult female, skin and skull; No. 19658, Mus. Vert. Zool.; obtained on July 2, 1913, by Alfred C. Shelton, from Gualala, on the Sonoma County side of the Gualala River, Sonoma Co., California. Range. — Coastal California from Point Reyes north to Point Arena. Diagnosis. — Size large; average and extreme measurements of 3 topotypes Speciation of the Wandering Shrew 33 scale of miles 130 Fig. 16. Probable geographic ranges of 16 subspecies of Sorex vagrans. Guide to 5. S. V. isolatus 11. S. V. souomae subspecies 6. S. V. setosus 12. S. V. longiquus 1. S. V. shumaginensis 7. S. V. bairdi 13. s. V. parvidens 2. S. V. obscurus 8. S. V. permiliensis 14. s. V. monticola 3. S. V. alascensis 9. S. V. yaquinae 15. s. V. neomexicanus 4. S. V. soperi 10. s. V. pacificus 16. s. V. orizabae S— 7903 34 University of ICansas Publs., Mus. Nat, Hist. are: total length, 141.7 (141-143); tail, 59 (54-63); hind foot, 17 (17-17). Color reddish in summer, somewhat grayer in winter. Comparisons. — Differs from S. v. pacificus, with which it intergrades to the north, in average smaller size and somewhat darker color; differs from the sympatric S. v. vagrans in much larger size and more reddish color in both summer and winter. Remarks. — This subspecies inhabits the Transition Life-zone below 300 feet, and occurs on moist ground in forests and beneath dense vegetation. Marginal records. — Californl\: Point Arena (Grinnell, 1933:82); Monte Rio (Jackson, 1928:144); Inverness (Grinnell, 1933:82). Sorex vagrans pacificus Coues Sorex pacificus Coues, Bull. U. S. Geol. and Geog. Surv. Terr., 3 (3):650, May 15, 1877. Sorex pacificus pacificus, Jackson, Jour. Mamm., 2:162, August 19, 1921. Type. — Adult, sex unknown, skin and skull; No. 3266 U. S. Nat. Mus.; date of capture unknown; received from E. P. Vollum and catalogued on March 8, 1858; obtained at Ft. Umpqua, mouth of Umpqua River, Douglas Co., Oregon. Range. — Coast of California and Oregon from Mendocino north to Gardiner. Diagnosis. — Size large, largest of the species; average and extreme measure- ments of 8 specimens from Orick, Humboldt Co., California, are: total length, 143.1 (134-154); tail, 65.5 (59-72); hind foot, 17.5 (16-19). Color reddish in summer, browner or grayer in winter. Comparisons. — See account of S. v. sonomae for comparison with that sub- species; averaging larger in all dimensions than S. v. yaquinae with which it intergrades to the north; much larger and has more reddish than the sympatric S. V. vagrans. Remarks. — This subspecies occurs in the Canadian and Transition life-zones below 1500 ft. where there is found moist ground in or adjacent to heavy forests. Specimens examined. — Total number, 76. Oregon: Douglas Co.: Umpqua, 1 BS. Coos Co.: Marshfield, 1 BS; Myrtle Point, 1 BS. Josephine Co.: Bolan Lake, 1 SGJ. Cajlifornia: Del Norte Co.: Smith River, 2 BS; Gasquet, 4 BS; Crescent City, 17 BS. Humboldt Co.: Orick, 13 BS; 1 mi. N Trinidad, 18 FC; Trinidad Head, 1 BS; Carson's Camp, Mad River, Humboldt Bay, 5 BS; Areata, 3 BS; Cape Mendocino, 2 BS; 5 mi. S Dyerville, 1 BS. Mendocino Co.: Mendocino, 6 BS. Marginal Records. — Oregon: Marshfield; Umpqua. California: Gasquet; 5 mi. S Dyerville; Mendocino, thence up coast to point of beginning. Sorex vagrans yaquinae Jackson Sorex yaquinae Jackson, Proc. Biol. Soc. Washington, 31:127, November 29, 1918. Sorex pacificus yaquinae, V. Bailey, N. Amer. Fauna, 55:364, August 29, 1936. Type.— Adult female, skin and skull; No. 73051 U. S. Biol. Surv. Coll., obtained on July 18, 1895, by B. J. Bretherton, from Yaquina Bay, Lincoln Co., Oregon. Diagnosis. — Size large for the species; average and extreme external measure- Speciation of the Wandering Shrew 35 ments of 11 specimens from Oakridge, Lane Co., Oregon, are: total length, 125.3 (11-136); tail, 55.1 (49-61); hind foot, 14.9 (14-16). Color reddish in summer, browner or grayer in winter. Comparisons. — See account of S. v. pacificus for comparison with that subspecies. Larger and more reddish than S. v. bairdi with which it intergrades to the north and east. Much larger and more reddish than the sympatric S. V. vagrans. Remarks. — The name yaquinae actually applies to a population of inter- grades between pacificus and bairdi. There is much variation over the range of the subspecies, and individuals from the western and southern parts are larger than those from the west slope of the Cascades. Specimens from Vida and McKenzie Bridge are smaller than those from Mapleton, Mercer, and the type locality but still seem closer to yaquinae than to topotypes of bairdi. Between Marshfield and Umpqua on the one hand, and the Columbia River and the Cascade Mountains on the other, the size of Sorex vagrans decreases quite rapidly from the large pacificus to the smaller permiliensis. Size decreases less rapidly northward along the coast than it does eastward toward the moun- tains; consequently, at any given latitude, coastal shrews are larger than mountain shrews. In this area of rapid change in size it is difficult to draw subspecific boundaries between pacificus, yaquinae, and bairdi, and this must be done somewhat arbitrarily. Jackson (1928:141) remarked upon the possibility that intergradation between pacificus and yaquinae took place. He noted also the close resemblance between yaquinae and bairdi, and stated {loc. cit.) that specific affinity be- tween the two might be demonstrated with more specimens. He had a series of eight specimens from Vida, Oregon, seven of which he assigned to S. o. bairdi and one to yaquinae. I have examined these specimens and find no more variation between the largest and the smallest than would be expected in any normally variable series of shrews. Vernon Bailey (1936:364) arranged yaquinae as a subspecies of pacificus without giving his reasons for so doing. Specimens examined. — Total number, 65. Oregon: Lincoln Co.: type lo- cahty, 2 AW. Benton Co.: Philomath, 2 BS. Lane Co.: Mable, 1 OU; Vida, 4 BS, 1 OSC, 3 OU; McKenzie Bridge, 8 OSC, 3 AW, 17 OU, 2 SGJ; Mercer, 1 OSC, 1 OU; Mapleton, 3 BS; Oakridge, 11 OU. Douglas Co.: Gardiner, 2 BS; Elkhead, 1 BS, Klamath Co.: Crescent Lake, 3 OU. Marginal Records. — Oregon: Yaquina Bay; Philomath; McKenzie Bridge; Prospect (Jackson, 1928:140); Crescent Lake; Gardiner. Sorex vagrans bairdi Merriam Sorex bairdi Merriam, N. Amer. Fauna, 10:77, December 31, 1895. Sorex obscurus bairdi, Jackson, Proc. Biol. Soc. Washington, 31:127, No- vember 29, 1918. Tt/pe.— Adult female, skin and skull; No. 17414/24318, U. S. Biol. Surv. Coll.; obtained on August 2, 1889, by T. S. Palmer, from Astoria, Clatsop Co., Oregon. Range. — Northwestern Oregon, south to Otis and east to Portland. Diagnosis. — Size medium for the species; average and extreme external measurements of 6 specimens from the type locality are: total length, 126.3^ (124-130); tail, 55.0 (52-57); hind foot, 15.0 (14-15). Color Fuscous to Sepia in summer, darker in winter, underparts buffy. 36 University of Kansas Publs., Mus. Nat. Hist. Comparisons. — For comparisons with yaquinae see account of that subspecies. More reddish and larger than permiliensis with which bairdi intergrades to the east; specimens from Portland show evidence of such intergradation. Some specimens from southern Tillamook County show an approach to yaquinae. Remarks. — S. v. bairdi hves primarily in forests as do yaquinae and pacificus. Specimens examined. — Total number, 39. Oregon: Clatsop Co.: tvpe lo- cality, 12 BS; Seaside, 3 BS. Tillamook Co.: Netarts, 1 OU; Tillamook, 2 OSC; Blaine, 1 AW; Hebo Lake, 1 SGJ; 5 mi. SW Cloverdale, 1 AW. Multnomah Co.: Portland, 6 USNM. Lincoln Co.: Otis, 7 USNM; Delake, 1 KU. Lane Co.: north slope Three Sisters, 6000 ft., 4 BS. Marginal Records. — Oregon: type locality; Portland; north slope Three Sisters; Taft (Macnab and Dirks, 1941:178). Sorex vagrans permiliensis Jackson Sorex obscurus permiliensis Jackson, Proc. Biol. Soc. Washington, 31:128, November 29, 1918. Type.— Adult male, skin and skull; No. 91048, U. S. Biol. Surv. Coll.; obtained on October 2, 1897, by J. A. Loring from Permilia Lake, W base Mt. Jefferson, Cascade Range, Marion Co., Oregon. Range. — The Cascade Mountains of Oregon from Mt. Jefferson north to the Columbia River. Diagnosis. — Size medium for the species; average and extreme measurements of 14 specimens from the type locahty are: total length, 117.7 (110-124); tail, 51.9 (45-58); hind foot, 14.0 (14-15). Pale reddish in summer, darker and brownish in winter. Comparisons. — For comparison with S. v. bairdi see account of that sub- species. Larger than S. v. setosus except tail relatively shorter. More reddish in summer pelage than setosus. Remarks. — S. v. bairdi is larger in the southern part of its range than else- where. Specimens from McKenzie Bridge, herein referred to yaquinae, are intermediate in character between yaquinae and bairdi or between yaquinae and permiliensis. The transition between yaquinae and bairdi is much more gradual than between yaquinae and permiliensis. Specimens examined. — Total number, 21. Oregon: Hood River Co.: Mt. Hood, 2 BS. Wasco Co.: Camas Prairie, E base Cascade Mts., SE Mt. Hood, 1 BS. Marion Co.: Detroit, 1 BS; type locahty, 17 BS. Marginal Records. — Oregon: Mt. Hood; type locality; Detroit. Sorex vagrans setosus Elliott Sorex setosus Elliott, Field Columb. Mus. Publ. 32, zool. ser. 1:274, May 19, 1899. Sorex obscurus setosus, Jackson, Proc. Biol. Soc. Washington, 31:127, No- vember 29, 1918. Type. — Adult male, skin and skull; No. 6213/238, Chicago Nat. Hist. Mus.; obtained on August 18, 1898, by D. G. Elliott from Happy Lake, Olympic Mts., Clallum Co., Washington. Range. — Washington from the Cascades west; southwestern British Columbia west of 120° W Longitude north to Lund. Diagnosis. — Size medium for the species; average and extreme measurements •of 20 specimens from the Olympic Mountains, Washington, are: total length. Speciation of the Wandering Shrew 37 117.3 (107-125); tail, 49.8 (41-54); hind foot, 13.4 (12-14). Color dark in both summer and winter. Comparisons. — For comparison with permiliensis see account of that sub- species. Darker, longer-tailed, and somewhat larger cranially than S. v. ob- scunis with which it intergrades in southwestern British Columbia. Smaller in all dimensions, but much the same color as S. v. longicauda with which it intergrades along the British Columbian coast north of Lund. Larger, darker, less reddish, and longer-tailed than the sympatric S. v. vagrans. Remarks. — S. v. setosus lives mostly in forests. According to Dalquest (1948:139) it is commonest at high altitudes in western Washington. In the Iludsonian Life-zone where shrew habitat is more restricted and marginal than it is at lower altitudes in the humid part of Washington, setosus might be ex- pected to compete with S. v. vagrans and to supplant it. Records of occur- rence in the Olympic Mountains suggest a degree of such separation there. Specimens examined. — Total number, 135. British Columbia: Lund, Malaspina Inlet, 4 BS; Gibson's Landing, 10 BS; Port Moody, 19 BS; Langley, 2 BS; Chiliwack, 1 BS; Manning Park, 2 PMBC. Washington: Whatcom Co.: Mt. Baker, 6 WSC; Barron, 1 BS. Chelan Co.: Clovay Pass, 1 WSC; Stehekin, 6 (4 WSC, 2 BS); Cascade Tunnel, 1 WSC. King Co.: Scenic, 1 WSC. Kittitas Co.: Lake Kachess, 1 WSC; Easton, 10 BS. Clallam Co.: 8 mi. W Sekin River, 1 WSC; mouth of Sekin River, 1 WSC; Clallam Bay, 1 CMNH; 7 mi. W Port Angeles, 1 WSC; Ozette Lake, 1 CMNH; 12 mi. S Port Angeles, 4 WSC; Forks, 1 CMNH; Deer Lake, 7 CMNH; Hoh Lake, 1 CMNH; Bogachiel Peak, 1 CMNH; Sol Due Hot Springs, 3 CMNH; Sol Due Park, 1 CMNH; Canyon Creek, 1 WSC; Sol Due Divide, 2 WSC; Cat Creek, 2 WSC. Jefferson Co.: Jackson Ranger Station, 1 CMNH; Mt. Kimta, 2 CMNH; Reflection Lake, 6 CMNH; Blue Glacier, 3 CMNH. Grai/s Harbor Co.: Westport, 1 WSC. Pierce Co.: Fort Lewis, 1 FC; Mt. Ranier, 19 (16 BS, 3 WSC). Pacific Co.: Tokeland, 2 BS. Yakima Co.: Gotchen Creek, 3 WSC; Mt. Adams, 1 WSC. Skamania Co.: Mt. St. Helens, 1. Oregon: Hood River Co.: 2 mi. W Parkdale, 2 BS. Marginal Records. — British Columbia: Rivers Inlet (Anderson, 1947:20); Agassiz (Jackson, 1928:136); Chilliwack Lake. Washington: Barron; Lvman Lake (Jackson, 1928:137); Mt. Stuart (Dalquest, 1948:141); Mt. Adams. Oregon: 2 mi. W Parkdale. Washington: Ilwaco (Jackson, 1928:137); Lund, Malaspina Inlet. Sorex vagrans longicauda Merriam Sorex obscurus longicauda Merriam, N. Amer. Fauna, 10:74, December 31, 1895. Type. — Adult male, skin and skull; No. 74711, U. S. Biol. Surv. Coll.; ob- tained on September 9, 1895 by C. P. Streator, from Wrangell, Alaska. Range. — The British Columbian and Alaskan coasts from Rivers Inlet north to near Juneau and also certain islands including Etolin, Gravina, Revillagigedo, Sergeif, and Wrangell. Diagnosis. — Size medium for the species, tail relatively long; average and extreme measurements of 17 specimens from the type locaHty are: total length, 128.4 (122-138); tail, 57.8 (53-66); hind foot, 15.1 (14-16). Color dark in summer and winter. Comparisons. — For comparison with S. v. setosus see account of that sub- species. Larger and darker than S. v. obscurus with which it intergrades east of the humid coastal region; larger and darker than S. v. alascensis with which 38 University of Kansas Publs., Mus. Nat. Hist, it intergrades in the Lynn Canal area; larger and darker than S. v. calvertensis which occurs on Calvert Island and Banks Island, British Columbia; differs from S. V. insularis of Smythe, Townsend, and Reginald islands in larger size and blackish ratlier than brown winter pelage; larger and relatively longer-tailed than S. V. elassodon wliich occurs on most of the islands west of the range of longicauda; larger and relatively longer-tailed than S. v. isolatiis. Specimens examined. — Total number, 151. Alaska: Wrangell, 54 BS; 8 AMNH; Crittenden Creek, 1 BS; Ketchikan, 2 BS; Loring, 11 BS. British Columbia: Port Simpson, 25 BS; Inverness, 15 BS; head of Rivers Inlet, 35 BS. Marginal Records. — British Columbia: Great Glacier, Stikine River (Jack- son, 1928:133). Alaska: Burroughs Bay (ibid.). British Columbia: Bella Coola region (Anderson, 1947:19); head of Rivers Inlet; Spider Island (Cowan, 1941:101); Goose Island (Cowan, 1941:99); Princess Royal Island (Cowan, 1941:98); Pitt Island (ibid.); Metlakatla (Jrxkson, 1928:133); Port Simpson. Alaska: Gravina Island (ibid.); Helm Bay (ibid.); Etolin Island (ibid.); Sergeif Island, mouth of Stikine River (ibid.); Sumdum Village (ibid.); Port Snettisham (ibid.). Sorex vagrans mixtus Hall Sorex obscurus mixtus Hall, American Nat., 72:462, September 10, 1938. Type. — Adult male, skin and skull; No. 70376, Mus. Vert. Zool; obtained on May 4, 1936, by R. A. Gumming, from Vanada, Texada Island, Georgia Strait, British Columbia. Range. — Known only from the type locality. Diagnosis. — Size medium; average and extreme measurements of 5 specimens from the type locality are: total length. 111 (108-117); tail, 48 (44-49); hind foot, 12 (12-13) (Hall, 1938:463). Comparisons. — Color much as in S. v. setosus or S. v. isolatus; palate longer than that of isolatus or setosus; hind foot shorter than either; smaller than S. V. longicauda. Sorex vagrans isolatus Jackson Sorex obscurus isolatus Jackson, Jour. Washington Acad. Sci., 12:263, June 14. 1922. Type. — Adult male, skin and skull; No. 177719, U. S. Biol. Surv. Coll.; obtained on May 21, 1911, by Alexander Wetmore from mouth of Millstone Creek, Nanaimo, Vancouver Island, British Columbia, Range. — Vancouver Island. Diagnosis. — Size medium; measurements of two from the type locality are: total length, 113, 118; tail, 48, 49; hind foot, 14, 14. Dark in summer and winter, underparts brownish. Comparisons. — Smaller than S. v. setosus but color much the same; resembles S. V. obscurus in size and cranial characters but darker in all pelages; similar in color to S. v. vancouverensis with which isolatus is sympatric but with longer tail, longer hind feet, broader rostrum and larger teeth. For comparsion with S. V. mixtus see account of that subspecies. Remarks. — S. v. isolatus and S. v. vancouverensis seemingly approach one Speciation of the Wandering Shrew 39 another morphologically more closely than do any other pair of sympatric sub- species of Sorex vagrans. The exceptions may be S. v. vagrans and S. v. obscurus which are geographically sympatric in a few places although they may be ecologically separated. Specimens examined. — Total number, 9. British Columbia, Vancouver Island: Nanaimo, 3 BS; Barclay Sound, 1 AMNH; Coldstream, 5 BS. Marginal Records. — British Columbia, Vancouver Island. (Anderson, 1947:19): Cape Scott; Victoria. Sorex vagrans insularis Cowan Sorex obscurus insularis Cowan, Proc. Biol. Soc. Washington, 54:103, July 31, 1941. Type. — Adult female, skin and skull; No. 3110, Prov. Mus. British Columbia; obtained on August 24, 1938, by T. T. and E. B. McCabe from Smythe Island, Bardswell Group, British Columbia. Range. — Smythe, Townsend, and Reginald islands, British Columbia. Diagnosis. — Size medium; average and extreme measurements of 50 speci- mens from within the range of the subspecies are: total length, 122.3 (111- 134); tail 52.6 (46-58); hind foot, 14.6 (13-15) (Cowan, 1941:107). Comparisons. — Smaller externally and cranially than S. v. longicauda and brown instead of blackish or grayish in winter pelage. Skull broader than that of S. V. calvertensis and color brown rather than blackish or grayish in winter pelage. Remarks. — S. v. insularis occurs together with S. cinereus on Townsend and Smythe islands. S. vagrans far outnumbered the cinereus shrew (Cowan, 1941:96). Records of occurrence. — British Columbia (Cowan, 1941:104): Smythe Island, Towsend Island, Reginald Island, Sorex vagrans calvertensis Cowan Sorex obscurus calvertensis Cowan, Proc. Biol. Soc. Washington, 54:103, July 31, 1941. Type. — Adult male, skin and skull; No. 1947, Prov. Mus. British Columbia; obtained on July 14, 1937, by T. T. and E. T. McCabe from Safety Cove, Calvert Island, British Columbia. Diagnosis. — Size medium; average and extreme measurements of 13 speci- mens from Calvert Island are: total length, 121.6 (109-129); tail, 54.0 (52-58); hind foot, 14.7 (13-15) (Cowan, 1941:106). Blackish or grayish in winter pelage. Comparisons. — Smaller externally and cranially and paler in winter and summer than S. v. longicauda; for comparisons with S. v. insularis see account of that subspecies. Remarks. — S. v. calvertensis seems to be the only shrew on Calvert and Banks islands. Records of occurrence. — British Columbia (Cowan, 1941:103): Safety Cove, Calvert Island; Larson Harbor, Banks Island. Marginal Records. — British Columbia: Larson Harbor, Banks Island; type locality. 40 University of Kansas Publs., Mus. Nat. Hist. yr^. 41i V ^ ^^4' '^^' 0\ K?^A^- Sorex vagrans malitiosus Jackson Sorcx obscurus malitiosus Jackson, Proc. Biol. Soc. Washington, 32:23, ' * Type. — Adult female, skin and skull; No. 8401; Mus. Vert. Zool.; ob- tained on May 21, 1909, by H. S. Swarth from east side of Warren Is- land, Alaska. Range. — Warren and Coronation islands, Alaska. Diagnosis. — Size medium; average and extreme measurements of 5 topo- types are: total length, 129.8 (126- 135); tail, 56.4 (53-61); hind foot, 15.4 (15-16). Color brownsh in sum- mer, brownish rather than blackish in winter. Comparisons. — Somewhat more brownish than S. v. longicauda but resembling it in size; skull slightly more flattened and rostrum broader. Larger than S. v. elassodon. Larger and relatively longer-tailed than S. v. alascensis. Records of occurrence. — Alaska ( Jackson, 1928 : 130 ) : Warren Island; Coronation Island. Sorex vagrans elassodon Osgood Sorex longicauda elassodon Osgood, N. Amer. Fauna, 21:35, Septem- ber 26, 1901. Sorex obscurus elassodon, Elliott, Field Columb. Mus. Publ. 105, zool. ser. 6:450, 1905. Type. — Adult male, skin and skull; No. 100597, U. S. Biol. Surv. Coll.; obtained on June 13, 1900, by W. II. Osgood from Cumshewa Inlet, near old Indian village of Clew, Moresby Island, Queen Charlotte Islands, Brit- ish Columbia. Range. — Alaskan and British Co- lumbian islands from Admiralty Island south to Moresby Island. Diagnosis. — Size medium; average and extreme measurements of 4 from the type locality are: total length, 126, (119-131); tail, 53.5 (52-55); hind foot, 13.8 (13-14). Color dark. Comparisons. — Smaller with rela- tively smaller tail and hind feet than Fig. 17. Probable geographic ranges of the subspecies of Sorex vagrans on the coast of British Columbia and southeastern Alaska. 1. Sorex vagrans malitiosus 2. Sorex vagrans elassodon 3. Sorex vagrans prevostensis 4. Sorex vagrans calvertensis 5. Sorex vagrans insularis 6. Sorex vagrans longicauda 7. Sorex vagrans obscurus Speciation of the Wandering Shrew 41 S. V. longicauda, but resembling it in color. Smaller and paler than S. v. pre- vostensis with relatively narrower rostrum. Larger, darker, and with relatively longer tail than S. v. ohscurns. Resembles S. v. alascensis but hind foot smaller and skull relatively narrower. Smaller than S. v. malitiosus. Remarks. — In the northern part of its range S. v. elassodon occurs with Sorex cinereus. In the southern part it is the only shrew present. Specimens examined. — Total number 93. Alaska: near Killisnoo, Admiraltry Island, 2 BS; Kupreanof Island, 15 BS; Petersburg, Mitkof Island, 10 BS; Woewodski Island, 4 AMNH; Kasaan Bay, Prince of Wales Island, 18 BS. British Columbia: Cumshewa Inlet, Moresby Island, 25 BS; Massett, Graham Island, 6 BS; Queen Charlotte Islands, 13 AMNH. Marginal Records. — Alaska: Hawk Inlet, Admiralty Island (Jackson, 1928:131); Kupreanof Island; Mitkof Island; St. John Harbor, Zarembo Island (Jackson, 1928:131); Kasaan Bay, Prince of Wales Island; Duke Island (Jack- son, 1928:131). British Columbia: Massett, Graham Island, Queen Char- lotte Islands; tvpe localitv; Langara Island, Queen Charlotte Islands (Jackson, 1928:131). Alaska: Forrester Island (ibid.)- Rocky Bay, Dall Island (ibid.); Shakan (really on Kosciusko Island) (ibid.); Point Baker (ibid.); Kuiu Island ( ibid. ) ; Port Conclusion, Baranof Island ( ibid. ) . Sorex vagrans prevostensis Osgood Sorcx longicauda prevostensis Osgood, N. Amer. Fauna, 21:35, September 26, 1901. Sorex obscurus prevostensis, Elliott, Field Columb. Mus. Publ. 105, zool. ser. 6:450, 1905. Type.— Adult male, skin and skull; No. 100618, U. S. Biol. Surv. Coll.; obtained on July 3, 1900, by W. H. Osgood from north end of Prevost Island (Kunghit Island on some maps) on coast of Houston Stewart Channel, Queen Charlotte Islands, British Columbia. Range. — Known only from the type locality. Diagnosis. — Size medium; measurements of two specimens from the type locality are: total length, 132, 142; tail, 53, 59; hind foot, 14, 15. Color dark. Comparisons. — Larger and darker than S. v. elassodon. Resembles S. v. longicauda but darker, tail relatively somewhat shorter on the average and rostrum relatively shghtly broader. Specimens examined. — Total number, 14. British Columbia: Prevost Is- land, Queen Charlotte Group, 14 BS. Sorex vagrans alascensis Merriam Sorex obscurus alascensis Merriam, N. Amer. Fauna, 10:76, December 31, 1895. Sorex glacialis Merriam, Proc. Washington Acad. Sci., 2:16, March 14, 1900, type from Point Gustavus, east side of entrance to Glacier Bay, Alaska. S[orex]. alascensis, Merriam, Proc. Washington Acad. Sci., 2:18, March 14, 1900. [Sorex glacialis] alascensis, Elliott, Field Columb. Mus. Publ. 45, zool. ser. 2:372, 1901. Sorex alascensis alascensis. Miller, U. S. Nat. Mus. Bull., 79:16, December 31, 1912. Type.— Adult female, skin and skull; No. 73539, U. S. Biol. Surv. Coll.; obtained on July 10, 1895, by C. P. Streator from Yakutat, Alaska. Range. — The coast of southern Alaska from the vicinity of Juneau west to include eastern part of the Kenai Peninsula, 42 University of Kansas Publs., Mus. Nat, Hist. Diagnosis. — Size medium for the species; average and extreme measure- ments of 9 specimens from 9 mi. W and 4 mi. N of Haines, Alaska, are: total length, 110 (104-128); tail, 45.4 (41-52); hind foot, 14 (14-14). Color gray- ish brown. Comparisons. — For comparison with S. v. longicauda and S. v. elassodon see accounts of those subspecies. Resembles S. v. obscurus in color but differs in larger skull, longer hind foot and in somewhat darker color. Larger and darker than S. v. shumaginensis; the two intergrade near the base of the Kenai Peninsula. Remarks. — This subspecies is transitional between the large, usually dark subspecies of the southeastern Alaskan and British Columbian coast and islands, and the smaller, paler subspecies of western and interior Alaska. There seem to be no sharp breaks between alascensis and shumaginesis. North of Haines, Alaska, size of shrews decreases in a short distance across a narrow intergrada- tional zone between alascensis and obscurus. Throughout most of its range S. V. alascensis occurs with Sorex cinereus. Specimens examined. — Total number, 88. Alaska: Orca, 1 BS; Montague Island, Prince William Sound, 2 BS; Yakutat, 8 BS; north shore Yakutat Bay, 2 BS; Yakutat Bay, 1 BS; E side Chilkat River. 100 ft., 9 mi. W and 4 mi. N Haines, 12 KU; 1 mi. S Haines, 5 ft., 10 KU; 7 mi. SSE Haines, 10 ft, 2 KU; N end Sullivan Island, 10 ft., 6 KU; SE end SulUvan Island, 10 ft., 2 KU; Glacier Bay, 3 BS; xMendenhall River, 1 BS; Juneau, 36 BS. British Columbia: Sheslay River, 1 AMNH; headwaters Sheslay River, 1 AMNH. Marginal Records. — Alaska: Valdez Narrows, Prince William Sound (Jack- son, 1928:128); north shore Yakutat Bay; east side Chilkat River, 100 ft., 9 mi. W and 4 mi. N Haines. British Columbia: Sheslay River (Jackson, 1928: 128). Alaska: Juneau; Glacier Bay; Montague Island, Prince William Sound (ibid.); Port Nell Juan (ibid.). Sorex vagrans shumaginensis Merriam Sorex alascensis shumaginensis Merriam, Proc. Washington Acad. Sci., 2:18, March 14, 1900. [Sorex glacialis] shumaginensis, ElUott, Field Coluinb. Mus. Publ. 45, zool. ser. 2:373, 1901. Sorex obscurus shumaginensis, Allen, Bull. American Mus. Nat. Hist., 16:228, July 12, 1902. Type.— Adult male, skin and skull; No. 97993, U. S. Biol. Surv. Coll.; ob- tained on July 17, 1899, by De Alton Saunders from Popof Island, Shumagin Islands, Alaska. (Measured by C. Hart Merriam and numbered 2210 in A. K. Fisher's catalog.) Range. — Southwestern Alaska from Seward Peninsula southeasterly to west- ern part of Kenai Peninsula and southwesterly to the southwestern end of the Alaskan Peninsula. Diagnosis. — Size medium to small for the species; average and extreme measurements of 6 specimens from King Cove, Alaska, are: total length, 112.7 (107-118); tail, 48.3 (45-52); hind foot, 13.8 (13-14). Tending toward the development of a tricolor pattern, the back darkest, the sides buffy, and the venter paler. Comparisons. — Paler and more definitely tricolored than S. t>. obscurus; also with relatively shorter palate, narrower rostrum and smaller teeth. For com- parison with S. V. alascensis see account of that subspecies. Speciation of the Wandering Shrew 43 Remarks. — S. v. shumaginensis occurs together with Sorex cinereus over much of southwestern Alaska. Part of the range of shumaginensis falls within the tundra of the Arctic Life-zone. Tliis may be a partial explanation of the tricolored pattern of the animal. Sorex tundrensis, S. cinereus ugyunak, and S. cinereus haydcni, shrews which dwell mostly in treeless areas, are markedly tricolored, or bicolored. Sorex arcticus, however, although tricolored, is found in forested areas. Specimens examined. — Total number, 340. Alaska: Sawtooth Mts., Nome River, 2 AMNH; Nulato, 5 BS; St. Michaels, 1 BS; Bethel, 7 BS; Aniak, 1 BS; Skeventna River, 1 BS; 6 mi. WSW Snowshoe Lake, 1 KU; 1 mi. NE Anchor- age, 1 KU; Tyonek River, 48 BS; Hope, 15 BS; Hope, Mts. near, 13 BS; Moose Camp, 3 AMNH; Kenai Peninsula, 24 AMNH; Kakwok River, 80 mi. up, 1 BS; Kakhtul River, 5 BS; Kakwok, 3 BS; Goodnews Bay, 1 BS; Lake Aleknagik, 6 BS; Nushagak River, 25 mi. above Nushagak, 1 BS; Dillingham, 1 BS; Nush- agak Village, 15 BS; Homer, 1 AMNH; Kenai Mts., 37 AMNH; Seldovia, 24 AMNH; Barabor, 1 AMNH; Caribou Camp, 7 AMNH; Ugagik River, 3 BS; Becharof Lake, 8 BS; Cold Bay, 14 BS; Kanatak, Portgage Bay, 4 BS; Chignik, 6 BS; Moller Bay, 1 BS; Alaska Peninsula, near Popof Island, 6 AMNH; Frosty Peak, 15 BS; Alorzhovoi Bay, 7 BS; Ungu Island, 3 BS; Sand Point, Popof Island, 45 AMNH; Popof Island, 3 BS. Marginal Records. — Alaska: Nome River; Nulato; Kuskokwim River, 200 mi. above Bethel, Crooked Creek (Jackson, 1928:126); 6 mi. WSW Snowshoe Lake; Seldovia; mts. near Hope; Morhzovoi Bay; thence along coast to St. Michael. Sorex vagrans obscurus Merriam Sorex vagrans similis Merriam, N. Amer. Fauna, 5:34, July 31, 1891, nee. S. similis Hensel, Zeitschr. der Deutsch. Geolog. Gesellsch., 7:459, 1855 ( = Neomys similis ) . Sorex obscurus Merriam, N. Amer. Fauna, 10:72, December 31, 1895, new name for Sorex vagrans similis Merriam. Sorex obscurus obscurus. Miller, Bull. U. S. Nat. Mus., 79:15, December 31, 1912. Type.— Adult female, skin and skull; No. 23525/30943, U. S. Biol. Surv. Coll.; obtained on August 26, 1890, by Vernon Bailey and B. H. Dutcher from near Timber Creek, 8200 ft., Lemhi Mts., 10 mi. SSW Junction (now Leadore), Lemhi Co., Idaho. Range. — Mountainous interior of western North America from central Alaska east across Yukon and southwestern Northwest Territories to northeastern Alberta, south in the mountains through north-central and western Washington, Idaho, western Montana, Wyoming, Utah, and Colorado, into northern New Mexico. Diagnosis. — Size medium to small for the species; average and extreme meas- urements of 9 topotypes are: total length, 110.3 (105-117); tail, 46.4 (42-50); hind foot, 13.1 (12.5-13.5). Color grayish or brownish gray in summer, hght grayish in winter. Comparisons. — For comparisons with S. v. setosus, S. v. longicauda, S. v. alascensis and S. v. shumaginensis see accounts of those subspecies. Paler and slightly larger than S. v. soperi. Larger than the subspecies from central Mon- tana herein described as new. Smaller than S. v. neomexicanus. Averaging larger in all dimensions than S. v. monticola with which obscurus intergrades in northern New Mexico and northern Arizona. Larger than S. v. vagrans with more grayish rather than reddish fresh summer pelage and light gray rather than dark grayish-black fresh winter pelage. 44 University of Kansas Publs., Mus. Nat. Hist. Remarks. — Intergradation of S. v. ohscurus with S. v. setostis, S. t;. longicauda, S. v. alascensis, and the new subspecies from Montana takes place in the usual way with specimens from intermediate lo- calities being intermediate in size and color. However the relation- ship of S. V. ohscurus and S. v. vagrans (as the latter subspecies is defined in this study) is rather complicated. In southern British Columbia where the two subspecies come together a situation of remarkable complexity prevails. Series from some localities seem to represent intergrades between ohscurus and vagrans; from other localities some specimens seem to be referable to one and some to the other subspecies; from other localities all specimens seem refer- able to one subspecies. A similar situation is seen in specimens from northeastern Washington, northern and central Idalio, and extreme western Montana. The region mentioned is one of extensive inter- fingering of fife-zones. In southern British Columbia the main axes of the rivers, valleys and mountain ranges are north and south. Most of the valleys are in the Transition Life-zone; the forests are rather dry and of pine with more or less isolated hydrosere com- munities about streams and ponds. These hydrosere situations are the habitat of Sorex vagrans. Shrews from these situations are usually referable to vagrans. The high ridges and mountain ranges are usually in the Canadian Life-zone or higher and most of the shrews referable to ohscurus come from such places. Marginal localities with regard to life-zone produce most of the populations which seem to represent intergrades between the two subspecies. Isolated areas of Canadian Life-zone, even tliough surrounded with Transition Life-zone, often harbor a population of ohscurus, whereas the streams in the nearby dry valleys harbor populations of vagrans. Farther south in the Rocky Mountain chain, ohscurus seemingly intergrades regularly with vagrans. This intergradation is seen in populations from several localities in Utah. There the lower elevations west of the Wasatch and Uintah mountains are inhabited by S. v. vagrans, the higher elevations by ohscurus and where the ranges of the two abut intergrading populations occur. In these series of intergrades there are specimens which, using size as a subspecific criterion, would unhesitatingly be assigned, as indi- viduals, to ohscurus, and others would be assigned to vagrans, but these individuals represent extremes of a normally variable popula- tion. At Cuddy Mountain, Idaho, the two subspecies seemingly abut without intergradation; anyhow the available specimens from this locality are referable to one or the other subspecies and none Speciation of the Wandering Shrew 45 is intermediate. The situation just described understandably has been the source of much anguish to students who sought to identify shrews from the Rocky Mountains. The reason for the relationship just described has been discussed at length in a previous section. In the Rocky Mountains of Wyoming and Colorado the subspecies S. V. obscurus ranges almost uninterruptedly over relatively large areas, but southward in New Mexico and southwestward into Utah and Arizona, suitable boreal habitat becomes insular in nature and obscurus there is confined to the higher mountains. With one ex- ception, once the shrev/ populations become 'insular' in this region they become smaller and show intergradation with Sorex vagrans monticola. The exception is the population in the Sacramento Mountains of southeastern New Mexico which is larger than ob- scurus and has been rightly recognized as a distinct subspecies, neomexicanus. Almost without exception the range of typical Sorex vagrans obscurus is sympatric with that of Sorex cinereus, usually the sub- species S. c. cinereus. So close is this correspondence that the pres- ence of S. cinereus comes near to being a useful aid in identifying S. V. obscurus. In areas where individuals of obscurus show inter- gradation with vagrans, Sorex cinereus is absent or rare. The impli- cation is that as the species S. vagrans approaches the size of the species S. cinereus, competition between the two increases with resultant displacement of cinereus. Specimens examined. — Total number, 982. Alaska: Wahoo Lake. 69°08'N, 146°58'W, 2350 ft., 2 KU; Chandler Lake, 68°12'N, 152°45'W, 2900 ft., 1 KU; Bettles, 1 KU, 5 BS; Alatna, 1 BS; Yukon River, 20 mi. above Circle, 1 BS; Tanana, 1 BS; Mountains near Eagle, 18 BS; Richardson, 8 BS; head of Toklat River, 11 BS; Savage River, 8 BS. Yukon: MacMillan Pass, Mile 282, Canol Road, 1 NMC; MacMillan River, Mile 249, Canol Road, 1 NMC; S. fork MacMillan River, Mile 249, Canol Road, 2 NMC; Sheldon Lake, Mile 222, Canol Road, 5 NMC; Rose River, Mile 95, Canol Road, 1 NMC; Mclntyre Creek, 3 mi. NW Whitehorse, 2250 ft. 1 KU; Nisuthn River, Mile 40, Canol Road, 6 NMC; SW end Dezadeash Lake, 2 KU; 3 mi. E and Hi mi. S Dalton Post, 2500 ft., 1 KU. Mackenzie: Nahanni River Mtns., Mackenzie River, 1 BS; Fort Simpson, 3 BS; Fort Resolution, Mission Island, 1 BS. British Columbia: W. side Mt. Clave, 4000 ft., 14 mi. S and 2 mi. E Kelsall Lake, 1 KU; Stonehouse Creek, 532 mi. W jet. Stonehouse Creek and Kelsall River, 4 KU; Bennett City, 6 BS; Wilson Creek, Atlin, 1 PMBC; Mc- Dame Post, Dease River, 6 BS; McDanie Creek, 3 BS; Hot Springs, 3 mi. WNW jet. Trout River and Liard River, 1 KU; NW side Muncho Lake, 1 KU; Little Tahtlan River, 1 AMNH; Junction (4 mi. N Telegraph Creek), 7 BS; Raspberry Creek, 16 AMNH; Klappan River Valley, 1 BS; Chapa-atan River, 4 BS; Fort Grahame, 3 BS; Kispiox Valley, 23 mi. N Hazleton, 1 BS; Bear Lake, site of Fort Connully, 2 BS; Tetana Lake, 1 PMBC; Hudson Hope, 2 BS; Charlie Lake, 3 PMBC; Babine Mts., 6 mi. N Babine Trail, 5200 ft., 1 BS; Big Sahnon River (S branch near Canyon), 1 BS; Ootsa Lake, 2 PMBC; Indianpoint Lake, 4 PMBC; Barkerville, 7 BS; Yellowhead Lake, 2 NMC, 1 46 University of Kansas Publs., Mus. Nat. Hist. PMBC; N. fork Moose River, 1 BS; Moose Lake, 2 BS; Moose Pass. 1 BS; Glacier, 7 AMNH, 12 BS; Golden, 1 BS; Field, 2 BS; Caribou Lake, near Kamloops, 2 BS; Sicamous, 1 BS; Monashee Pass, 4 PJvlBC; Paradise Mine, 3 PMBC; Level Mtn., 4 AMNH; 6 mi. S Nelson, 6 BS; Morrissey, 5 NMC; Wall Lake, 1 BS. Alberta: Hays Camp, Slave River, Wood BuflFalo Park, 1 NMC; Kinuso, Assineau River, 1920 ft., 2 KU; Athabaska River, 30 mi. above Athabaska Landing, 8 BS; Smokey Valley, 50 mi. N Jasper House, 1 BS; Sulfur Prairie, Grande Cache River, 3 BS; Stoney River, 35 mi. N Jasper House, 1 BS; Moose Mtn., 1 NMC; Rodent Valley, 25 mi. W Henry House, 1 BS; Henry House, 3 BS; Jasper, 2 NMC; Shovel Pass, 4 NMC; mouth of Cavell Creek, Jasper Park, 1 NMC; 11 mi. S Henry House, 2 BS; 15 mi. S Henry House, 1 BS; Red Deer River, 1 AMNH; 27 mi. W Banff, 3 NMC; 12 mi. WNW Banff, 4500 ft., 1 NMC; N. Fork Saskatchewan River, 5000 ft., 1 NMC; Cypress Hills, 1 NMC; Waterton Lakes Park, 53 NMC. Saskatchewan: Cypress Hills, 21 NMC. Washington: Okanogan Co.: Pasayten River, 1 BS; Bauerman Ridge, 1 BS- Conconully, 2 BS. Pend Oreille Co.: 2 mi, N Gypsy Meadows, 2 WSC; Round Top Mtn., 1 WSC; head Pass Creek, 1. Chelan Co.: Stehekin, 4 BS; head Lake Chelan, 4 BS; Wenatchee, 1 BS. Kittitas Co.: Easton, 10 BS. Idaho: Boundary Co.: Cabinet Mtns., E Priest Lake, 2 BS. Adams Co.: ^ mi. E Black Lake. 1 KU; 1 mi. N Bear Creek R. S., SW slope Smith Mtn., 2 KU. Washington Co.: 1 mi. NE Heath, SW slope Cuddy Mtn., 4000 ft., 4 KU. Lemhi Co.: 10 mi. SSW Leadore (type locaUty), 4 BS; 5 FC. Fremont Co.: 7 mi. W West Yellowstone, 4 KU. Custer Co.: head Pahsimeroi River, Pahsimeroi Mtns., 1 BS. Blaine Co.: Perkins Lake, 1 KU. Bear Lake Co.: — Caribou Co. hne: Preuss Mts., 1 BS. Montana: Glacier Co.: Sherburne Lake, 3 UM; 2)2 mi. W and IM mi. S Babb, 1 KU; St. Mary's, 6 UM; St. Mary Lakes, 9 BS; Fish Creek, 2 BS; Gun- sight Lake, 2 BS. Flathead Co.: Nyack, 3 UM, 1 BS; 1 mi. W and 2 mi. 5 Summit, 1 KU. Ravalli Co.: 8 mi. NE Stevensville, 3 BS; Sula, 1 BS. Meagher Co.: Big Belt Mtns., Camas Creek, 4 mi. S Fort Logan, 7 BS. Gallatin Co.: West Gallatin River, 4 BS. Park Co.: Emmigrant Gulch, 3 mi. SE Chico, 2 BS; Beartooth Mtns., 2 BS; Carbon Co.: Pryor Mtns., 2 BS. Wyoming: Yellowstone Nat'l Park: Mammoth Hot Springs, 11 BS; Tower Falls, 1 BS; Astringent Creek, 1 BS; Flat Mtn., 1 BS; Yellowstone Park, 1 UM. Park Co.: Beartooth Lake, 15 BS; SW slope Whirlwind Peak, 1 KU; Pahaska Tepee, 6300 ft., 8 BS; Pahaska, mouth Grinnell Creek, 15 BS; Pahaska, Grinnell Creek, 7000-7500 ft., 18 BS; 25 mi. S and 28 mi. W Cody, 1 KU; Valley, Absaroka Mts., 14 BS; Needle Mtn., 2 BS. Big Horn Co.: 28 mi. E Lovell, 9000 ft., 12 KU; head Trapper's Creek, W slope Bighorn Mtns., 7 BS; ITA mi. E and 4% mi. S Shell, 1 KU. Teton Co.: Two Ocean Lake, 6 FC; Emma Matilda Lake, 2 BS; 1 mi. N Moran, 1 FC; 2)i mi. E and 'A mi. N Moran, 6230 ft., 2 KU; Moran, 7 FC, 1 KU; 25^ mi. E Moran, 6220 ft., 1 KU; 1 mi. S Moran, 1 FC; S% mi. E and 1 mi. S Moran, 9 KU; 7 mi. S Moran, 3 FC; Timbered Lsland, 14 mi. N Moose, 6750 ft., 3 KU; Bar BC Ranch. 2)^ mi. NE Mnnsts 6500 ft., 1 KU; Beaver Dick Lake, 1 UM; Teton Mtns., Moose Creek, 6800 ft., 9 BS; Teton Mtns., S Moose Creek, 10,000 ft, 3 BS; Teton Pass, above Fish Creek, 7200 ft., 15 BS; Whetstone Creek, 4 UM; Flat Creek-Gravel Creek Divide, 2 UM; Flat Creek-Granite Creek Divide, 1 UM; Jackson. 3 KU, 2 UM. Fremont Co.: Togwotee Pass, 5 FC; Jackey's Creek, 3 mi. S Dubois, 1 BS; Milford, 5400 ft., 2 KU; Mosquito Park R. S. 175^ mi. W and 2J^ mi. N Lander, 1 KU; 17 mi. S and 6)i mi. W Lander, 9300 ft., 1 KU; Mocassin Lake, 19 mi. W and 4 mi. N Lander. 10,000 ft., 1 KU; 23M mi. S and 5 mi. W Lander, 8600 ft., 1 KU; Green Mts., 8 mi. E Rongis, 8000 ft., 4 BS. Waslwkic Co.: 9 mi. E and 5 mi. N Tcnsleep, 7400 ft., 2 KU; 9 mi. E ond 4 mi. N Tf^nsleep, 7000 ft., 2 KU. Lincoln Co.: Salt River Mtns., 10 mi. SE Afton, 5 BS; Labarge Creek, 9000 ft., 1 BS. Sublette Co.: 31 mi. N Pinedale, 8025 ft., 3 KU; Sur- veyor's Park, 12 mi. NE Pinedale, 8000 ft.. 2 BS; N. side Half Moon Lake. 7900 ft., 1 KU; 2)i mi. NE Pinedale, 7500 ft., 2 KU. Natrona Co.: Rattlesnake Mts., 7000-7500 ft., 18 BS; Casper Mts., 7 mi. S Casper, 6 BS. Converse Co.: 2Hi mi. S and 24M mi. W Douglas, 7600 ft., 7 KU; 22 mi. S and 245^ mi. W Speciation of the Wandering Shrew 47 Douglas, 7600 ft., 4 KU; 22)^ mi. S and 24M mi. W Douglas, 7600 ft.. 2 KU. Uinta Co.: 1 mi. N Fort Bridger, 6650 ft., 1 KU; Fort Bridger, 3 KU; Evanston, 1 BS; 9 mi. S Robertson, 8000 ft., 6 KU; 9 mi. S and 2'A mi. E Robertson, 8600 ft., 1 KU; 10 mi. S and 1 mi. W Robertson, 8700 ft., 3 KU; lOYi mi. S and 2 mi. E Robertson, 8900 ft., 1 KU; 13 mi. S and 1 mi. E Robertson, 9000 ft., 1 KU; 13 mi. S and 2 mi E Robertson, 9200 ft., 1 KU. Carbon Co.: Ferris Mts., 7800 to 8500 ft., 13 BS; Shirley Mts., 7600 ft., 7 BS; Bridger's Pass, 18 mi. SW Rawlins, 7500 ft., 2 KU; 10 mi. N and 12 mi. E Encampment, 7200 ft., 1 KU; 10 mi. N and 14 mi. E Encampment, 8000 ft., 6 KU; 95^ mi. N and IVA mi. E Encampment, 7200 ft., 2 KU; 9 mi. N and 3 mi. E Encampment. 6500 ft., 1 KU; 9 mi. N and 8 mi. E Encampment, 7000 ft., 1 KU; 8 mi. N and 14 mi. E Encampment, 8400 ft., 3 KU; 8 mi. N and 143^ mi. E Encampment, 8100 ft., 2 KU; 8 mi. N and 16 mi, E Encampment, 4 KU; 8 mi. N and 21M mi. E Encampment, 9400 ft., 2 KU; S. base Bridger's Peak, 8800 ft.. Sierra Madre Mts., 3 BS; 8 mi. N and 193»: mi. E Savery, 8800 ft, 2 KU; 7 mi. N and 17 mi. E Saver>', 8300 ft., 1 KU; 6)^ mi. N and 16 mi. E Savery, 8300 ft., 1 KU; 6 mi. N and 15 mi. E Savery, 8500 ft., 1 KU; 5 mi. N and 105^ mi. E Savery, 8000 ft., 2 KU; 14 mi. E and 6 mi. S Saratoga, 8800 ft., 1 KU. Albany Co.: Spring- hill, 12 mi. N Laramie Peak, 6300 ft, 10 BS; Laramie Peak, N. slope, 8000 to 8800 ft., 7 BS; Bear Creek, 3 mi. SW Laramie Peak, 7500 ft., 6 BS; 2M mi. ESE Brown's Peak, 10,500 ft, 2 KU; 3 mi. ESE Brown's Peak, 10,000 ft., 1 KU; 27 mi. N and 5 mi. E Laramie, 6960 ft., 2 KU; 1 mi. SSE Pole Mtn., 8350 ft., 3 KU; 2 mi. SW Pole Mtn., 3 KU; 3 mi. S Pole Mtn., 8100 ft., 2 KU; 8% mi. E and 6M mi. S Laramie, 8200 ft., 2 KU; Woods P. O., 1 BS. Laramie Co.: 5 mi. W and 1 mi. N Horse Creek P. O., 7200 ft., 2 KU. Utah: Weber Co.: Mt. Willard, Weber-Box Elder Co. line, 9768 ft., 2 UU. Salt Lake Co.: Butterfield Canyon, 7000 ft., 1 UU; Brighton, Silver Lake P. O., 8700 ft., 2 UU; Brighton, Silver Lake P. O., 8750 ft., 8 UU; Brighton, Silver Lake P.O., 9000 ft., 2 UU; Brighton, Silver Lake P.O., 9500 ft., 1 UU. Summit Co.: Jet. Bear River and East Fork, 2 CM; Smith and Morehouse Canyon, 7000 ft., 1 UU; Mirror Lake, 10,000 ft., 1 UU. Daggett Co.: Jet. Deep and Carter creeks, 7900 ft., 1 UU. Utah Co.: Nebo Mtn., 1 mi. E Payson Lake, 8300 ft., 1 UU; Nebo Mts., 12 mi. SE Payson Lake, 1 UU. Wasatch Co.: Current Creek, Uinta Mts., 1 BS; Wasatch Mts., 1 BS. Uintah Co.: Paradise Park, 21 mi. W and 15 mi. N Vernal, Uinta Mts., 10,050 ft., 2 CM, 3 KU; Paradise Park, Uinta Mts., 10,100 ft., 6 UU. Sanpete Co.: Manti, 3 BS. Sevier Co.: 7 mi. Creek, 20 mi. SE Salina, 5 CM; Fish Lake Plateau, 2 BS. Emery Co.: Lake Creek, 11 mi. E Mt. Pleasant, 4 CM. Grand Co.: Warner R. S., La Sal Mts., 9750 ft., 2 UU; La Sal Mts., 11,000 ft., 1 BS. Beaver Co.: PuflFer Lake, Beaver Mts., 2 BS. Wayne Co.: Elkhom G. S., Fish Lake Plateau, 14 mi. N Torrey, 9400 ft., 3 UU. Garfield Co.: Wildcat R. S., Boulder Mtn., 8700 ft., 6 UU; 18 mi. N Escalante, 9500 ft., 1 UU. Washington Co.: Pine Valley Mts., 7 BS. San Juan Co.: Geyer Pass, 18 mi. SSE Moab, 3 CM; Cooley, 8 mi. W Monticello, 3 CM. Colorado: Larim.er Co.: Poudre River, 1 KU. Rio Blanco Co.: 9M mi. SW Pagoda Peak, 2 KU. Grand Co.: Arapaho Pass, Rabbit Ears Mts., 2 BS. Boulder Co.: Willow Park, Rocky Mtn. Nat'l Park, 8 UM; Longs Peak, 1 BS; % mi. N and 2 mi. W Allenspark, 8400 ft., 5 KU; Ward, 9500 ft., 1 BS; Bu- chanan Pass, 1 BS; 3 mi. S Ward, 1 KU; 7 mi. NW Nederland, 1 KU; 5 mi. W Boulder, 3 BS; Boulder, 3 BS, 1 ChM; Nederland, 6 BS, 4 ChM; Eldora, 1 BS. Garfield Co.: Baxter Pass, 8500 ft., 2 BS. Eagle Co.: Gores Range, 1 BS. Gilpen Co.: Black Hawk, 1 BS. Lake Co.: 3 mi. W Twin Lakes, 2 KU; 12 mi. S and 1 mi. W Leadville, 1 KU. Gunnison Co.: 2 mi. W Gothic, 2 FC; Copper Lake, 2 FC; Gothic, 1 FC. Chaffee Co.: St. Elmo, 10,100 ft., 2 BS; E side Monarch Pass, 7 mi. W Salida, 2 ChM. Teller Co.: Glen Core, Pikes Peak, 2 UM. El Paso Co.: Hunters Creek, a tributary of Bear Creek, 7250- 7400 ft., 1 AMNH. Montrose Co.: Uncomphagre Plateau, 8500 ft., 3 BS. Saguache Co.: 3 mi. N and 16 mi. W Saguache, 8500 ft., 2 KU; Cochetopa Pass, 10,000 ft., 4 KU; Monshower Meadows, 27 mi. W Saguache, 2 BS. San Juan Co.: Silverton, 4 BS. Mineral Co.: 23 mi. S and 11 mi. E Creede, 1 KU. Costilla Co.: Fort Garland, 2 BS. Huerfano Co.: 5 mi. S and 1 mi. W Cuchara Camps, 8 KU. 48 University of Kansas Publs., Mus. Nat. Hist. New Mexico: Taos Co.: 3 mi. N Red River, 2 BS; Taos, 1 BS. Colfax Co.: 1 mi. S and 2 mi. E Eagle Nest, 8100 ft., 2 KU. Sandoval Co.: Jemez Mts., 3 BS. Santa Fe Co.: Hyde Park, 5 mi. NE Santa Fe, 2 HC; Santa Fe Field Station, 1 HC; Santa Fe Ski Basin, 1 KU; Pecos Baldy, 4 BS. Torrance Co.: Manzano Mts., 2 BS. Marginal Records.— Alaska: Chandler Lake, 68° 12' N, 152°45' W; Yukon River, 20 mi. above Circle; Mountains near Eagle. Mackenzie: Nahanni River Mts.; Fort Simpson; Fort Resolution, Mission Island. Alberta: Wood Buffalo Park; Athabaska River, 30 mi. above Athabaska Landing. Saskat- chewan: Cypress Hills. Montana: St. Mary; 4 mi. S Fort Logan; Prvor Mts. Wyoming: 1 mi. W and 1 mi. S Buffalo, 27424 KU; Springhill, 12 mi. N Laramie Peak; 5 mi. W and 1 mi. N Horse Creek PO. Colorado: Boulder; Hunters Creek; 5 mi. S and 1 mi. W Cuchara Camps. New Mexico: 3 mi. N Red River, 10,700 ft.; Pecos Baldy; Manzano Mts.; Jemez Mts. Colorado: Navajo River (Jackson, 1928:120); Silverton. Utah: La Sal Mts., 11,000 ft, Colorado: Baxter Pass. Utah: junction Trout and Ashley Creeks, 9700 ft. (Durrant, 1952:35); Mirror Lake, 10,000 ft.; Mt. Baldy R. S. (Durrant, 1952: 53); Wildcat R. S.; Pine Valley Mts.; Puffer Lake; Butterfield Canyon. Idaho: Preuss Mts.; 4 mi. S Trude (Davis, 1939:104); head Pahsimeroi River, Pah- simeroi Mts.; Perkins Lake; 1 mi. NE Heath; ^ mi. E Black Lake. Montana: Sula; 8 mi. NE Stevensville. Washington: head Pass Creek; Conconully; Wenatchee; Easton; Stehekin; Pasayten River. British Columbia: Second Summit, Skagit River, 5000 ft., (Jackson, 1928:120); Babine Mts., 6 mi. N Babine Trail, 5200 ft.; Hazleton (Jackson, 1928:120); 23 mi. N Hazleton; Flood Glacier, Stikine River (Jackson, 1928:120); Cheonee Mts. (ibid.); Level Mtn.; west side Mt. Clave, 4000 ft., 14 mi. S and 2 mi. E Kelsall Lake. Alaska: head Toklat River; Tanana; Alatna; Bettles. Sorex vagrans soperi Anderson and Rand Sorex obscurus soperi Anderson and Rand, Canadian Field-Nat., 59:47, October 16, 1945. Type. — Adult male, skin and skull; No. 18249, Nat. Mus. Canada; obtained on September 21, 1940, by J. Dewey Soper, from 2/2 mi. NW Lake Audy, Riding Mtn. Nat'l Park, Manitoba. Range. — Southwestern Manitoba to central Saskatchewan. Diagnosis. — Size medium to small for the species; measurements of type and two topotypes are: total length, 107, 108, 117; tail, 45, 45, 45; hind foot, 12.1, 12.3, 12.5. Color dark brownish or fuscous in summer pelage; winter pelage unknown. Comparison. — Resembles S. v. obscurus in size; darker than obscurus in summer pelage; cranium slightly higher and top more nearly Hat; larger and darker in summer pelage than the new subspecies from central Montana. Remarks. — In their description of this subspecies Anderson and Rand pointed out that specimens from the type locahty and from central Saskatchewan represent the dark extreme in a color cline which begins in south-central British Columbia with "pale, brownish-tinged animals." These authors referred shrews from Cypress Hills, southwestern Saskatchewan and southeastern Alberta to S. o. soperi, although they noted that these specimens, taken by themselves, are not strikingly different from S. o. obscurus from the Rocky Mountains. The specimens from tlie Cypress Hills were included in soperi because the authors felt that the subspecific boundary should be drawn "where specimens average about half way between the extremes (of the cline) in characters." It is true, as Anderson and Rand say, that the shrews from C>'press Hills are hardly separable from those from, say, Waterton Lakes Park. The speci- Speciation of the Wandering Shrew 49 mens from the Cypress Hills are noticeably diflFerent from specimens from the Okanagon area, but some of the latter, in my opinion may represent intergrades between S. v. obscurus and the more reddish S. v. vagrans and are not, at any rate, typical obscurus. In view of the similarity of shrews from Cypress Hills to typical S. v. obscurus and since the Cypress Hills are much nearer to the range of S. v. obscurus than to the record-stations of occurrence in central Saskatchewan and Manitoba, I have chosen to restrict tlie name soperi to shrews from these latter two localities. Seemingly S. vagrans is absent from the plains separating the Cypress Hills from the Rocky Mountains and from Riding Mountain National Park. Specimens examined. — none. Marginal records. — Saskatchewan: Prince Albert National Park, 1700 ft. (Anderson and Rand, 1945:48). Manitoba: Riding Mountain National Park, 2)2 mi. NW Audy Lake (ibid.). Sorex vagrans longiquus new subspecies Type. — First year male, skin and skull; No. 87332, Univ. Michigan Mus. Zool.; obtained on July 21, 1942, by Emmet T. Hooper from 25 mi. ESE Big Sandy, Eagle Creek, Chouteau Co., Montana, original no. 2184. Range. — Central Montana; marginal localities are: Bearpaw Mts., Zortman, Big Snowy Mts., Buffalo, Little Belt Mts. Diagnosis. — Size small for the species; measurements of three topotypes are: total length, 101, 105, 108; tail, 39, 40, 42; hind foot, 11.5, 11.5, 12. Color pale; summer pelage: back near ( 17"'k) OHve Brown but hairs of dorsum with a pale, buffy band proximal from the tips which imparts a pale over-all appear- ance; flanks near Wood Brown; underparts Pale Smoke Gray, usually not with a buffy wash; color of underparts often extending along margin of upper lip. Skull small for species; rostrum relatively broad and heavy; relatively broad interorbitally. Comparisons. — From S. v. obscurus, S. v. longiquus differs as follows: size smaller; skull smaller in all dimensions although similar in proportion. From S. V. soperi, S. v. longiquus differs in: size smaller; color paler in summer pelage. From S. v. vagrans, S. v. longiquus differs in: color paler in summer pelage, less brownish; color of venter extending higher on flanks; venter Pale Smoke Gray, rarely tinged with buffy rather than usually tinged with buffy. From S. v. monticola, S. v. longiquus differs in: summer pelage slightly paler, venter Pale Smoke Gray rather than suffused with buffy. Remarks. — The subspecies longiquus is obviously derived from the neighbor- ing S. V. obscurus and differs from it mainly in size. Some specimens of obscurus from western Montana show evidences of intergradation with S. v. vagrans in possessing a somewhat buffy belly and these are thus more strikingly different from longiquus than are other specimens of obscurus. Many speci- mens of obscurus from the eastern slope of the Lewis and Clark Range in Mon- tana show the tricolored pattern seen in many specimens of longiquus. The smallest individuals of longiquus are found on the Big Snowy Mountains. In- tergradation with obscurus is seen in specimens here referred to S. v. obscurus from the Big Belt Mountains. Specimens examined. — Total number, 45. Montana: Hill Co.: Bearpaw Mts., 5 UM, 2 BS. Phillips Co.: Zortman, 1 BS. Chouteau Co.: type locality, 4—7903 50 University of Kansas Publs., Mus. Nat. Hist. 3 UM; Highwood Mts., 13 BS. Cascade Co.: Neihart, Little Belt Mts., 1 BS. Judith Basin Co.: 3 mi. W Geyser, 4100 ft., 1 KU; Otter Creek, 10 mi. SW Geyser, 1 BS; Dry Wolf Creek, 20 mi. SW Stanford, 1 BS. Buffalo, 13 mi. W Buffalo Canyon, 2 BS. Fergus Co.: Moccasin Mts., 15 mi. NW Hilger, 3 BS; Judith Mts., 17 mi. NE Lewiston, 1 BS; 15 mi. S Heath, N. fork Flat Willow Creek, Big Snowy Mts., 1 BS; Timber Creek, Big Snowy Mts., 1 BS; Crystal Lake, 6000 ft., Big Snowy Mts., 2 UM; Rocky Creek, 5600 ft.. Big Snowy Mts., 3 UM; Big Snowy Mts., 3 BS. Meagher Co.: Sheep Creek, 16 mi. N White Sulphur Springs, Little Belt Mts., 1 BS. Marginal records. — Montana: Bearpaw Mts.; Zortman; Big Snowy Mts.; 16 mi. N White Sulphur Springs; Highwood Mts. Sorex vagrans neomexicanus Bailey Sorex obscurus neomexicanus Bailey, Proc. Biol. Soc. Washington, 26:133, May 21, 1913. Type.— Adult male, skin and skull; No. 100440, U. S. Biol. Surv. Coll.; obtained on May 29, 1900, by Vernon Bailey, from Cloudcroft, 9000 ft., Otero Co., New Mexico. Range. — Sacramento and Capitan Mountains of New Mexico. Diagnosis. — Size medium for the species; average and extreme measurements of 4 topotypes are: total length, 105.2 (103-107); tail, 41.0 (39-42); hind foot, 13.1 (12.5-14). Color near Olive Brown in summer; winter pelage unknown; skull large and relatively broad; teeth relatively large. Comparisons. — Skull larger than that of S. v. obscurus and relatively some- what broader; much larger in all cranial dimensions than S. v. monticola. Remarks. — S. v. neomexicanus is a well-marked subspecies seemingly limited to the mountains of southeastern New Mexico. It is the only species of Sorex thus far recorded from that area. Specimens examined. — Total number, 12. New Mexico: Otero Co.: SW slope Capitan Mts., 2 BS; 10 mi. NE Cloudcroft, 2 BS; Cloudcroft, 7 BS. 1 UM. Marginal records. — New Mexico: SW slope Capitan Mts.; 10 mi. NE Cloudcroft; type locahty. Sorex vagrans monticola Merriam Sorex monticolus Merriam, N. Amer. Fauna, 3:43, September 11, 1890. Sorex vagrans monticola, Merriam, N. Amer. Fauna, 10:69, December 31, 1895. Sorex melanogenys Hall, Jour. Mamm., 13:260, August 9, 1932, type from Marijilda Canyon, 8600 ft., Graham Mts. [= Pinaleno Alts.] Graham Co., Arizona. Type.— Adult male, skin and skull; No. 17599/24535, U. S. Biol. Surv. Coll.; obtained on August 28, 1899, by C. Hart Merriam and Vernon Bailey from San Francisco Mtn., 11,500 ft., Coconino Co., Arizona. Range. — Mountains of western New Mexico, eastern Arizona, and the north- em Sierra Madre Occidental of Mexico. Diagnosis. — Size small for the species; average and extreme measurements of 12 specimens from the White Mountains, Arizona, are: total length, 104.3 (98-112); tail, 41.2 (37-45); hind foot, 12.0 (11-13). Summer pelage between (15'm) Proutts Brown and (15"7n) Bister, venter tinged with (15'/) Pale Ochraceous Buff; winter pelage near {I7"'k) Olive Brown; skull relatively broad. SpECIATION of the WA>fDERING ShREW 51 Comparisons. — For comparisons with S. v. obscurus and S. v. neomexicanus see accounts of those subspecies. Skull slightly larger and relatively broader than that of S. v. orizabae, and color slightly paler. Differs from S. v. vagrans in: winter pelage grayish (near 17"'k Olive Brown) rather than blackish {17""k or 17""m Chaetura Drab or Chaetura Black); summer pelage slightly grayer; skull relatively slightly broader rostrally and interorbitally. Remarks. — S. v. monticola intergrades gradually with S. v. obscurus to the north and east; indeed the type locality is actually in this area of intergrada- tion. So far as I know, monticola is not in reproductive continuity with any other subspecies of Sorex vagrans. Specimens from southeastern Arizona are the smallest and seem to be the most "typical" in the sense that they are most different from S. t>. obscurus. Some specimens from the whole length of the Rocky Mountain chain in the United States have for years been referred to monticola. Some of these, as I have pointed out, belong to S. v. longiquus, and others are intergrades between S. v. obscurus and S. v. vagrans. Since vagrans and monticola resemble one another somewhat, and since topotypes of S. v. monticola actually show the influence of intergradation with obscurus, it is easy to understand how intergrades between obscurus and vagrans could have been assigned to monticola. Throughout most of its range, S. v. monticola is the only Sorex present. In some places monticola may occur with S. ruinus or S. merriami. S. v. monticola occurs with the water shrew in southeastern Arizona. Specimens examined. — Total number, 80, Arizona: Coconino Co.: San Francisco Mtn., 3 BS, 6 CMNH. Apache Co.: Spruce Creek, Tunitcha Mts., 7 BS; Springerville, 1 BS; North Fork White River, White Mts., 12 SD; White River, Horseshoe Cienega, 8300 ft.. White Mts., 5 BS; Mt. Thomas, 9500 to 11,000 ft.. White Mts., 12 BS; Little Colorado River, White Mts., 4 BS; near head Burro Creek, 9000 ft.. White Mts., 1 BS. Graham Co.: Graham Mts., 9200 ft., 2 BS. Greenlee Co.: Prieto Plateau, 9000 ft., S. end Blue Range, 1 BS. Pima Co.: Summerhaven, 7500 ft., Santa Catalina Mts., 3 BS, 1 SD. Cochise Co.: Fly Park, Chiricahua Mts., 4 BS; Rustler Park, Chiricahua Mts., 1 SD; Long Park, Chiricahua Mts., 1 UM; Huachuca Mts., 1 BS. Santa Cruz Co.: Stone Cabin Canyon, 8500 ft., Santa Rita Mts., 1 BS. New Mexico. — San Juan Co.: Chusca Mts., 1 BS. Catron Co.: MogoUon Mts., 3 BS; 10 mi. E Mogollon, 1 KU. Socorro Co.: Copper Canyon, Magdalena Mts., 3 BS. Sierra Co.: Mimbres Mts., near Kingston, 1 BS. Chihuahua. — Sierra Madre, near Guadalupe y Calvo, 5 BS. Marginal records. — Arizona: Tunitcha Mts. New Mexico: Chusca Mts.; Copper Canyon, Magdalena Mts.; Mimbres Mts., near Kingston. Chihuahua: Guadalupe y Calvo. Arizona: Huachuca Mts.; Santa Catalina Mts.; White River, Horseshoe Cienega, 8300 ft., White Mts.; San Francisco Mtn. Sorex vagrans orizabae Merriam Sorex orizabae Merriam, N. Amer. Fauna, 10:71, December 31, 1895. Sorex vagrans orizabae, Jackson, N. Amer. Fauna, 51:113, July 24, 1928, Type. — Adult female, skin and skull; No. 53633, U. S. Biol. Surv. Coll.; obtained on April 24, 1893, by E. W. Nelson from W slope of Mt. Orizaba, 9,500 ft., Puebla. Range. — Transverse volcanic belt of mountains at the southern end of the Mexican Plateau. 52 University of ICansas Publs., Mus. Nat. Hist. Diagnosis. — Size small for the species; measurements of 3 specimens from Volcan Toluca, Mexico, are: total length, 98, 100, 108; tail, 35, 39, 40; hind foot, 13, 13, 14. Summer pelage Mummy Brown tending toward Olive Brown; Fuscous to Fuscous-Black in winter; skull and teeth relatively narrow. Comparisons. — For comparison with S. v. monticola see account of that subspecies. Remarks. — The range of S. v. orizabae probably is not now in contact with that of any other subspecies of S. vagrans, although judging by the shght degree of difference between orizabae and monticola the separation bewteen the two has not been of great duration. Sorex vagrans orizabae occurs with S. saussurei saussurei throughout the transverse volcanic belt. Specimens examined. — Total number, 23. Michoacan: Patamban, 1 BS; Nahuatzin, 3 BS; Mt. Tancitaro, 4 BS. Mexico: Salazar, 2 BS, 1 KU; Volcan de Toluca, 3 BS. Tlaxcala: Mt. Malinche, 2 BS. Puebla: Mt. Orizaba, 6 BS. Veracruz: Cofre de Perote, 1 BS. Marginal records. — Michoacan: Patamban. Veracri^: Cofre de Perote. Puebla: Mt. Orizaba. Michoacan: Mt. Tancitaro. Sorex vagrans vagrans Baird Sorex vagrans Baird, Rep't Pacific R. R. Survey 8: pt. 1, Mammals, p. 15, July 14, 1858. Sorex suckleyi Baird, Rep't Pacific R. R. Survey 8: pt. 1, Mammals, p. 18, July 14, 1858, type from Steilacoom, Pierce Co., Washington. Sorex dobsoni Merriam, N. Amer. Fauna, 5:33, July 30, 1891, type from Alturas or Sawtooth Lake, altitude about 7200 ft., E base Sawtooth Mts., Blaine Co., Idaho. Sorex amoenus Merriam, N. Amer. Fauna, 10:69, December 31, 1895, type from near Mammoth, 8000 ft., head Owens River, E slope Sierra Nevada, Mono Co., CaUfomia. Sorex nevadensis Merriam, N. Amer. Fauna, 10:71, December 31, 1895, type from Reese River, 6000 ft., Nye-Lander Co. hne, Nevada. Sorex shastensis Merriam, N. Amer. Fauna, 16:87, October 28, 1899, type from Wagon Camp, Mt. Shasta, 5700 ft., Siskiyou Co., California. Type. — Adult male, alcoholic; No. 1675, U. S. Nat. Mus.; obtained at Shoalwater (Willapa) Bay, Pacific Co., Washington; received from J. G. Cooper, and entered in Museum catalog on October 23, 1856. Range. — The Great Basin, and Columbia Plateau west across the mountains to the Pacific coast of northern California, Oregon, Washington and southwest- ern British Columbia. Diagnosis. — Size small for the species; average and extreme measurements of 8 topotypes are: total length, 104.1 (99-109); tail, 43.3 (42-45); hind foot, 12.9 (12-14). Summer pelage ranging from (15'/c) Cinnamon Brown through (15'm) Proutt's Brown to (17'm) Mummy Brown. Winter pelage (13""m) Fuscous Black to (17""m) Chaetura Black. Comparisons. — For comparison with S. v. monticola see account of that subspecies. Differs from S. v. halicoetes in relatively narrower and more at- tenuate rostrum and in less brownish underparts in winter pelage; smaller and more brownish (less grayish) than Sorex vagrans from the southern Sierra Nevada. Speciation of the Wandering Shrew 53 122 112 \4^^|)'^ ®^B 46 - ^B ^B« - 46 1 ^^^B ^^M <.x ^^B ^^L/-^ ^^l ^^^^^ 40 ~ 0 ■ 9 t 40 1 20 •y «^x i" \ 1 1 scale of miles ^^ \; , \ 122 112 Fig. 18. Probable geographic ranges of Sorex vagrans vagrans, its deriva- tive subspecies, and S. v. mixttis. 1. S. V. vancouverensis 4. S. v. paludivagus 2. S. V. vagrans 5. S. v. obscuroides 3. S. V. halicoetes 6. S. u. mixtus Remarks. — Restriction of the range of S. v. monticola to Arizona and New Mexico leaves shrews that were formerly assigned to this subspecies from Utah, Idaho, Washington and southern British Columbia unassigned. Several names are available for considera- tion. The name Sorex vagrans dobsoni Merriam, 1891, type locality Alturas Lake, Blaine Co., Idaho, was once applied to small shrews from Idaho, Montana, Wyoming, and Utah, but was considered by Jackson to be synonymous with S. v. monticola. The name Sorex vagrans amoenus Merriam, 1895, type locality near Mammoth, Mono Co., California, has been applied to wandering shrews from western Nevada, northeastern California and southern Oregon. Sorex vag- rans nevadensis Merriam, 1895, type locahty Reese River on Nye- Lander Co. line, Nevada was considered by Hall (1946:119) to be synonymous with S. v. amoenus. Specimens of Sorex vagrans west 54 University of Kansas Publs., Mus. Nat. Hist. of the Cascade Mountains have long been referred to the nominate subspecies which has its type locahty at Willapa Bay, Pacific Co., Washington. Over so wide an area it is only to be expected that some geographic variation is to be found. Thus specimens from central Nevada average slightly paler in summer pelage than those from the Pacific Coast or from the foothills of the Rocky Mountains. In addition there are slight average differences in size from place to place. Topotypes of S, v. vagrans, however, show a fair degree of variability and some are nearly as pale as the paler Great Basin stocks. Furthermore topotypical individuals of vagrans can be lost in series of S. v. amoenus, although amoeniis is shorter-tailed on the average. Specimens from the western foot-hills of the Rocky Moun- tains show an amazing series of relationships with the montane S. v. obscurus. In Utah, as previously pointed out, complete intergrada- tion occurs. At 1 mi. N Heath, Washington Co., Idaho, the lowland and tlie highland forms approach each other within a short distance and still maintain a degree of distinctness, especially in size. In northwestern Montana intergradation is extensive (Clothier, 1950). In northeastern Washington distinctly separable populations occur within a few miles of one another. In southern British Columbia some populations are clearly intergrades while at 6 mi. S Yahk intergradation seemingly has not taken place. Where some inter- gradation has occurred the result often has been increased size of the lowland shrews, although they usually retain the reddish summer pelage rather than acquiring the more grayish pelage of obscurus. The name dobsoni was based upon shrews from a place where low- land and highland forms occur almost together with only a slight amount of intergradation. Examples of "dobsoni" may not with certainty be distinguished from typical vagrans except that they are, as Merriam (1895:68-69) points out, somewhat larger. Mer- riam {loc. cit.) further notes that dobsoni is "intermediate in size and cranial characters between S. vagrans and obscurus" a state- ment which hits very close to the heart of the matter. I consider the name dobsoni to apply to intergrades. To attempt to apply the name to the highly variable populations of intergrades from British Columbia to southern Idaho seems inadvisable. I have examined the possibility of using the name amoenus for the animals from this region. The characters which set amoenus apart from vagrans, slightly shorter tail and slightly darker summer pelage, however, are not universally found in shrews from the Columbian Plateau and eastern Great Basin and furthermore these differences between amoenus and vagrans do not seem to me to be of great enough Speciation of the Wandering Shrew 55 magnitude to warrant subspecific recognition of the former. Thus the name S. v. vagrans may apply to shrews in the region under consideration. The subspecies, as thus thought of, embraces several incipient subspecies, namely (1) the populations on the isolated mountain ranges of Nevada, (2) the coastal rain forest population and possibly (3) the population on the Columbian Plateau. In western British Columbia, Washington, and Oregon no evi- dences of intergradation between S. v. vagrans and the races setosus, permiliensis, bairdi, ijaqtiinae, or pacificus are seen. In this region S. V. vagrans occurs sympatrically with one or the other of these sub- species. Different degrees of differentiation thus obtain between the subspecies vagrans as here defined and the surrounding sub- species of Sorex vagrans to wit: complete intergradation and allopatry in Utah with S. v. ohscurus; partial intergradation and partial sympatry with S. v. ohscurus in the foothill region from Idaho to British Columbia; no intergradation and complete sympatry with all the other races of Sorex vagrans from the Cascades to the coast and south to San Francisco Bay. The relationship of S. v. vagrans to the wandering shrews of the high Sierra is discussed on page 58. Throughout most of the Great Basin and Columbian Plateau Sorex vagrans is, with the exception of the rare S. merriami and S. preblei, the only small shrew. In the Cascades and in the coastal lowlands it is the only small shrew except for S. cinereus and S. trigonirostris, both extremely rare and local in this region. S. vagrans seemingly competes to a certain extent with the larger S. trowhridgi in western Washington and seems to be partially dominant to trowhridgi, at least in marshy habitats (Dalquest, 1941:171). Specimens examined. — Total number, 1197. British Columbia: Osoyoos District: Okanagan, 20 PMBC; Okanagan Landing, 2 PMBC; Nahun Plateau, 2 PMBC. Vaiicouver District: Vancouver, 2 PMBC. New Westminister District: Port Moody, 16 BS; Westminster Jet., 4 AMNH; Langley, 1 BS; Vedder Crossing, 1 PMBC; Huntingdon, 69 NMC; Sumas, 16 BS; Cultus Lake, 1 NMC. SimiJkameen District: Princeton, 6 Mile Creek, 1 NMC. Hedley, Stirling Creek, 7 NMC; Fairview-Keremeos Summit, 5 NMC; Oliver. 1 NMC; Westbridge, 6 NMC; Osoyoos, 1 PMBC; Osoyoos- Bridesville Summit, 4 NMC; Cascade, 7 NMC. Nelson District: Kuskonook 1 PMBC; Rossland, 14 NMC; Trail, 2 NMC; Creston, 4 PMBC, 4 NMC; neai Creston, 7 NMC. Cranbrook District: Cranbrook, 5 BS; Yahk, 2 NMC; Yahk Camp 6, 2 NMC; Goatfell, 2 NMC. Fernie District: Newgate, 3 NMC. Washington: Whatcom Co.: Blaine, 1 BS; Beaver Creek, 5 WSC; Glacier, 1 BS; Mt. Baker Lodge, 1 WSC; Lake Whatcom, 1 BS; Barron, 2 BS. Okanogan Co.: Sheep Mtn., 3 BS; E. end Bauerman Ridge, 1 BS; Oroville, 1 BS; Hidden Lakes, 1 BS; Loomis, 1 BS; Conconully, 1 BS; Twisp, 1 BS. Ferry Co.: 5 mi. W Curlevi', 2 BS. Stevens Co.: Marcus, 1 BS. Pend Oreille Co.: Canyon, 1 WSC; Metaline, 2 BS; Sullivan Lake, 1 BS. San Juan Co.: East Sound, Orcas Island, 3 BS; Friday Harbor, San Juan Island, 1 BS; San Juan Park, 2 WSC; Blakely Island, 1 KU; Richardson, 6 BS. Skagit Co.: Cypress Island, 1 KU; Hamilton, 1 BS; Sauk. 1 BS; Avon, 3 BS; Mt. Vernon, 2 BS; La Conner, 5 BS. 56 University of Kansas Publs., Mus. Nat. Hist. Island Co.: San de Fuca, Whidby Island, 3 BS; Greenbank, Whidby Island, 2 BS; 3 mi. N Clinton, Whidby Island, 1 BS. Snohomish Co.: Oso, 2 BS; Hermosa Point, Tulalip Indian Reservation, 7 mi. W and Ja mi. N Marysville, 3 KU. Chelan Co.: Entiat, 2 BS. Lincoln Co.: 6 mi. E Odessa, 4 BS. Spokane Co.: Marshall, 7 BS. Clallam Co.: Neah Bay, 29 BS; 8 mi. W Sekin Eiver, 1 WSC; mouth Sekin River, 1 WSC; Dungeness, 1 BS; Port Townsend, 3 BS; Ozette Indian Reservation, 1 CMNH; Sequim, 4 BS; Tivoli Island, Ozette Lake, 1 CMNH; Garden Island, Ozette Lake, 3 CMNH; Elwah, 1 WSC; Blyn, 1 BS; Soleduck River, 1 BS; 12 mi. S Port Angeles, 1 WSC; Forks, 9 CMNH, 1 BS; Cat Creek, 1 WSC; Lapush, 5 BS. Jefferson Co.: Jefferson Ranger Sta- tion, N Fork Hoh River, 5 CMNH; Duckabush, 6 BS. Kitsap Co: Vashon Island, 2 BS. King Co.: Redniont, 2 BS; Kirkland, 20 BS; Seattle, 1 WSC, 3 KU; North- bend, 2 BS; Lake Washington, near Renton, 2 BS; Kent, 1 BS; Enumclaw, 1 BS. Grays Harbor Co.: Lake Quinault, 9 BS; Aberdeen, 20 BS; Westport, 5 BS, 2 WSC; Oakville, 1 BS. Mason Co.: Lake Cushman, 11 BS; Hoodsport, 1 BS; North Fork Skokomish River, 1 BS; Shelton, 2 BS. Pierce Co.: Puyallup, 6 BS; Steilacoom, 1 BS; 6 mi. S Tacoma, 2 BS; Roy, 3 BS; Bear Prairie, Mt. Rainier, 1 BS; Reflection Lake, Mt. Rainier, 1 WSC. KHtitas Co.: Blew.ett Pass, 3 BS; Easton, 3 BS; 2 mi. E Cle Elum, 4 FC; Ellensburg, 2 BS. Grant Co.: Moses Lake, 1 BS; 9 mi. S and 1 mi. W Neppel, 1 UM. Whitman Co.: Hang- man Creek, Tekoa, 1 WSC; 4 mi. ENE Pullman, 1 KU; 2 mi. N Pullman, 2 WSC; 2 mi. NW Pullman, 1 WSC; 2 mi. W Pullman, 1 WSC; Pullman, 5 WSC; Armstrong, 1 WSC; 5 mi. NE Wawawai, 1 BS; Wawavi^ai, 5 WSC. Thurston Co.: Nisqually Flats, 2 BS; Nisqually, 1 BS; 4 mi. S Olympia, 1 BS; Tenino, 4 BS. Pacific Co.: Tokeland, 4 BS; 1 mi. S Nemah, 2 FC; 1 mi. N Bear River, Willapa Bay, 8 FC; }i mi. N Bear River, 3 FC; SVz mi. E Seavievi', 6 FC; Ilwaco, 1 BS. Lewis Co.: 8 mi. W Chehallis, 2 BS; Chehallis, 2 BS; Toledo, 1 BS. Yakima Co.: Selah, 7 KU; Wiley City, 4 BS. Wahkiakum Co.: Cathlamet, 1 BS. Skamania Co.: 45 mi. SE Toledo, 2 BS; Carson, 1 BS; Stevenson, 1 BS; 15 mi. NW White Salmon, 1 BS. Klickitat Co.: Trout Lake, 15 mi. S Mt. Adams, 2 BS; 15 mi. N Goldendale, 1 WSC; Goldendale, 1 BS. Walla Walla Co.: College Place, 1 KU. Columbia Co.: Starbuck, 3 BS. Garfield Co.: 1 mi. E Pomeroy, 1 SGJ. Asotin Co.: 21 mi. SE Dayton, 1 BS; Rogersburg, 1 BS. Idaho: Bonner Co.: 4 mi. S Sandpoint, 1 UM. Kootenai Co.: Coeur d'Alene, 2 BS. Shoshone Co.: Osburry, 1 BS; MuUan, 2 BS. Latah Co.: Felton's Mills, 1 WSC; Cedar Mtn., 5 WSC. Lewis Co.: Nezperce, 2 BS. Idaho Co.: Seven Devils Mtn., 1 BS. Adams Co.: Summit of Smith Mtn., 7500 ft., 5 KU; New Meadows, 1 BS; Tamarack, 1 BS. Washington Co.: 1 mi. NE Heath, SW slope Cuddy Mtn., 4000 ft., 7 KU. Boise Co.: Bald Mtn. R. S., 10 mi. S Idaho City, 1 BS. Elmore Co.: Cayuse Creek, 10 mi. N Featherville, 1 BS. Canyon Co.: Nampa, 5 BS. Blaine Co.: Sawtooth City, 5 BS; Alturas Lake, 1 BS. Bonneville Co.: 10 mi. SE Irwin, 5 BS. Bannock Co.: Pocatello, 1 BS, 1 KU; 1 mi. W Bancroft, 1 KU; Swan Lake, 1 BS. Owyhee Co.: Gras- mere, 1 SGJ. Cassia Co.: 10 mi. S Albion, Mt. Harrison, 1 BS. Montana: Sanders Co.: Prospect Creek, near Thomson Falls, 4 BS. Lake Co.: Flathead Lake, 5 BS. Ravalli Co.: Bass Creek, NW Stevensville. 2 BS; 2 mi. NE Stevensville, 1 UM; Corvallis, 4 BS; 6 mi. E Hamilton, 1 KU. Oregon: Clatsop Co.: Seaside, 1 BS. Washington Co.: 5 mi. SE Hillsboro, 1 BS; Beaverton, 1 BS. Multnomah Co.: Portland, 20 BS; Portland, Switzler Lake, 5 BS. Hood River Co.: 2 mi. W Parkdale, 1 BS; north slope Mt. Hood, 2 BS. Umatilla Co.: 10 mi. W Meacham, 2 BS; Meacham, 3 BS. Union Co.: Elgin, 2 BS; Kamela, 2 BS; Hot Lake, 2 BS. Wallowa Co.: 25 mi. N. Enter- prise, 4 BS; Wallowa Lake, 23 BS; S Wallowa Lake, 1 BS. Clackamas Co.: Estacada, 1 KU. Marion Co.: Salem, 8 BS; Permilia Lake, 2 BS. Benton Co.: Corvallis, 2 BS; 5 mi. SW Philomath, 5 BS. Linn Co.: Shelbum, 1 BS. Jeffer- son Co.: 20 mi. W Warm Springs, 2 BS. Grant Co.: Beech Creek. 6 BS; Austin, 1 BS; Strawberry Butte, 1 BS; Strawberry Mts., 12 BS. Baker Co.: Homestead, 1 BS; Cornucopia, 11 BS; Rock Creek. 1 BS; Bourne, 7 BS; Mc- Ewen, 1 BS; Huntington. 1 BS; Anthony, 42 AMNH. Lane Co.: north slope Three Sisters, 3 BS; Vida, 1 BS; Mapleton. 1 BS; Eugene, 2 BS; 10 mi. S Speciation of the Wandering Shrew 57 McKenzie Bridge, 1 BS; Florence, 1 BS. Deschutes Co.: Paulina Lake, 7 BS; Lapine, 8 BS. Crook Co.: 1 SGJ. Douglas Co.: Winchester Bay, 1 SGJ; Scottsburg, 3 BS; Drain, 5 BS; Lookinglass, 1 BS; Diamond Lake, 6 BS. Coos Co.: Empire, 5 BS. Curry Co.: Port Orford, 1 BS; Gold Beach, 4 BS. Klamath Co.: Anna Creek, Mt. Mazama, 1 BS; Crater Lake, 14 BS; Upper Klamath Marsh, 2 BS; Ft. Klamath, 35 BS; Klamath Falls, 6 BS. Lake Co.: 10 mi. SW Silver Lake, 3 BS; west fork Silver Creek, Yamsay Mts., 4 BS; Plush, 1 BS; Warner Creek, Warner Mts., 1 BS; Warner Mts., 3 BS; Gearhart Mts., 17 SGJ; Harney Co.: Diamond, 2 BS; Keiger Gorge, Steens Mts., 3 BS. Malheur Co.: 8 mi. W Jordon Valley, 1 BS. Wyoming: Lincoln Co.: 13 mi. N and 2 mi. W Afton, 6 KU; 10 mi. N Afton, Salt River, 2 BS; 9 mi. N and 2 mi. W Afton, 6 KU; 7 mi. N and 1 mi. W Alton, 4 KU; Cokeville, 1 BS; 12 mi. N and 2 mi. E Sage, 2 KU; 6 mi. N and 2 mi. E Sage, 1 KU. California: Del Norte Co.: Smith River, 2 BS; Crescent City, 20 BS. Siskiyou Co.: Beswick, 1 BS; Hombrook, 3 BS; Brownell, Klamath Lake, 1 BS; Mayten, 2 BS; Squaws Creek, Mt. Shasta, 5 BS; Upper Ash Creek, Mt. Shasta, 1 BS; upper Mud Creek, Mt. Shasta, 8 BS; Wagon Camp, Mt. Shasta, 5 BS; Warmcastle Soda Springs, Squaw Creek Valley, 2 BS; Sisson, 7 BS. Modoc Co.: Davis Creek, Goose Lake, 1 BS. Humboldt Co.: Humboldt Bay, 10 BS. Trinity Co.: Canyon Creek, 2 BS. Shasta Co.: Fort Crook, 11 BS; Dana, 17 BS; Fall Lake, Fall River Valley, 3 BS; Cassel, 2 BS; 12 mi. E Bumey, 1 BS; Lassen Peak, 13 BS; Kellys, Warner Creek, 1 KU; Drakes Hot Springs, Warner Creek, 2 BS. Mendocino Co.: Russian Gulch State Park, 2 FC. Plumas Co.: 12 mi. NE Prattville, 2 BS; Spring Garden Ranch, Grizzly Mts., 3 BS; Sierra Valley, 1 BS. Sierra Co.: Lincoln Creek, 1 BS. Sonoma Co.: Petaluma, 3 BS; Point Reyes, 7 BS. Placer Co.: Donner, 3 BS. El Dorado Co.: Tallac, 3 BS. Mono Co.: Mt. Conness, 1-BS; Mono Lake, 1 BS; near Mammoth, 8000 ft., head of Owens River, 2 BS. Inyo Co.: Alvord, 1 BS. Nevada: Elko Co.: Mountain City, 1 BS; Three Lakes, 1 KU; west side Ruby Lake, 3 mi. N White Pine Co. line, 8 KU; Ruby Mts., 9 BS; W side Ruby Lake, 3 BS. White Pine Co.: W side Ruby Lake, 3 mi. S Elko Co. hne, 1 KU. Nye Co.: Cloverdale, Reese River, 3 BS. Utah: Weber Co.: Beaver Creek, S Fork Ogden River, 2 UU; Huntsville, 10 mi. E Ogden, 1 UU; Hooper Bay Refuge, 4200 ft., 1 UU; Riverdale, 4200 ft., 3 UU; Riverdale, 4250 ft., 1 UU; 3 mi. SE Ogden, 2 UU; Snow Basin, 2 UU; Snow Basin, S part Wheeler Canyon, 1 UU; Uinta, 2 mi. W Weber Canyon entrance, 4 UU; 2 mi. W Uinta, 1 UU. Salt Lake Co.: City Creek Canyon, 6 mi. NE Salt Lake City, 4700 ft., 2 UU; 1 mi. up City Creek Canyon, 4600 ft., 1 UU; %. mi. above Forks, City Creek Canyon, 1 UU; The Firs, Mill- creek Canyon, 1 UU; Olympus Water Box, 1 UU; Salamander Lake, Lamb's Canyon, 9000 ft., 3 UU (near obscurus); Salt Lake City, 7500 ft., 1 UU; 1 mi. W Draper, 4500 ft., 6 UU; Draper, 4500 ft., 5 UU; IJ^ mi. SW Draper, 4500 ft., 1 UU; 3 mi. SW Draper, 4400 ft., 2 UU; 3 mi. S Draper, 4400 ft., 2 UU; 1 mi. S Draper, 4500 ft., 1 UU. Juab Co.: W side Deep Creek Mts., Queen of Sheba Canyon, 8000 ft., 3 UU. Wasatch Co.: Midway Fish Hatchery, 5450 ft., 1 UU. Marginal records. — British Columbia: Okanagan; Westbridge; Kuskonook; Cranbrook. Montana: Flathead Lake; 6 mi. E Hamilton; Prospect Creek. Idaho: Cedar Mtn.; New Meadows; Alturas Lake; 10 mi. SE Irwin. Wyoming: 13 mi. N and 2 mi. W Afton; 6 mi. N and 2 mi. E Sage. Idaho: 1 mi. W Bancroft; Swan Lake. Utah: Beaver Creek, South Fork, Ogden River; Mid- way Fish Hatchery; west side Deep Creek Mts., Queen of Sheba Canyon, 8000 ft. Nevada: Baker Creek (Hall, 1946:120); Reese River (ibid.); 2 mi. S Hinds Hot Springs (ibid.). California: Mono Lake (Jackson, 1928:110); near Mammoth; Alvord; Mount Conness; Donner; Buck Ranch (Jackson, 1928:110); Warner Creek, Drake Hot Springs (ibid.); Canyon Creek; Cuddeback (Jackson, 1928:105); Novato Point (ibid.), thence northward along the coast to Wash- ington: Friday Harbor, San Juan Island. British Columbia: Port Moody. 58 University of Kansas Publs., Mus. Nat. Hist. Sorex vagrans obscuroides new subspecies Type.— First year female, sldn and skull; No. 30064/42074, U. S. Biol. Surv. Coll.; obtained on August 9, 1891, by Frank Stephens from Bishop Creek, 6600 ft., Inyo Co., California, original no. 811. Range. — The Sierra Nevada of California, north at least to El Dorado County, intergrading northerly with S. v. vagrans. Diagnosis. — Size medium for the species; average and extreme measurements of 5 topotypes are: total length, 107 (103-112); tail, 47 (45-50); hind foot, 12.8 (12-13.5). Skull relatively broad interorbitally; color of dorsum in sum- mer pelage nearest ( 17"'/c ) Olive Brown. Comparisons. — Differs from S. v. vagrans, with which it intergrades to the north, in: longer tail and total length; skull larger and relatively broader interorbitally; color in summer grayer (less reddish), the lighter subterminal color bands of the hair often showing through the darker tips and imparting a grizzled appearance to the dorsum. Differs from S. v. parvidens to the south in: skull relatively broader interorbitally and less flattened; teeth slightly larger. Remarks. — S. v. obscuroides has long been called S. o. obscurus. In fact, obscuroides is separated from the range of obscurus by the intervening, smaller subspecies S. v. vagrans. S. v. obscuroides resembles S. v. obscurus in color and size but the skull is smaller, although relatively slightly broader. The re- semblance in color is possibly due to the fact that obscuroides, like obscurus, is a high mountain form. S. v. obscuroides intergrades with S. v. vagrans along the crest of the Sierra between Yosemite National Park and Lassen Peak and on the eastern slope of the Sierra from approximately Mammoth north- ward. Specimens from Donner are intergrades but are closest to S. v. vagrans. Although all specimens from Lassen Peak are referable to S. v. vagrans, some show cranial characters of obscuroides. Specimens examined. — Total number, 76. Californi.\: Mono Co.: Mt. Dana, 6 BS; Mt. Lyell, 11 BS. Mariposa Co.: Tuolumne Meadows, Muir Meadow, 9300 ft., 1 BS; Tuolumne Meadows, Mt. Unicorn, 1 BS; Tuolumne Meadows, N base Mt. Lyell, 8 BS; Tuolumne Meadows, Soda Springs, 4 BS; Lake Tenaya, 5 BS. Madera Co.: San Joaquin River, 8000 ft., 4 BS. Fresno Co.: Horse Corral Meadows, 3 BS. Mono Co.: head of Owens River near Mammoth, 2 BS. Inyo Co.: Bishop Creek, 5 BS; Round Valley, 1 BS. Tulare Co.: E. Fork Kaweah River, 7 BS; Mt. Whitney, 5 BS; Whitney Creek, Mt. Whitney, 4 ChM; Whitney Meadows, 9700 ft., 1 BS; Mineralking, 2 BS; N. Fork Kern River, 9600 ft., 1 BS; S. Fork Kern River, 4 BS; Kern Lakes, 1 BS. Marginal records. — California: Pyramid Peak; near Mammoth; Round Valley; Bishop Creek; Mt. Whitney; Kern Lakes; Halstead Meadows; Morse Corral Meadows; east fork Indian Canyon (Jackson, 1928:121). Sorex vagrans parvidens Jackson Sorex obscurus parvidens Jackson, Jour. Mamm., 2:161, August 19, 1921. Type. — Adult male, skin and skull; No. 56561, U. S. Biol. Surv. Coll.; ob- tained on October 3, 1893, by J. E. McLellan from Thurmans Camp, Bluff Lake, 7500 ft., San Bernardino Mts., California. Range. — Confined, so far as known, to the San Bernardino and San Gabriel mountains, San Bernardino Co., California. Diagnosis. — Size medium for the species; measurements of two specimens from the San Bernardino Mountains are: total length, 105, 106; tail, 41, 48; hind foot, 12, 14. Upper parts in summer Olive-Brown to Buffy-Brown; Speciation of the Wandering Shrew 59 cranium flattened and relatively narrow; unicuspids and incisors relatively small. Comparisons. — For comparison with S. v. obscuroides, the only adjacent subspecies, see the account of that subspecies. Remarks. — S. v. parvidens is seemingly an uncommon mammal. I have been informed by Terry Vaughan that repeated attempts by him to obtain it in suitable habitat in the San Gabriel Mountains failed. This shrew is probably no longer in reproductive continuity with Sorex vagrans of the Sierra Nevada. Specimens examined. — Total number, 4. California: San Bernardino Co.: type locality, 4 BS. Marginal records. — California: Camp Baldy, San Antonio Canyon (Jack- son, 1928:124); type locality. Sorex vagrans halicoetes Grinnell Sorex halicoetes Grinnell, Univ. California Publ. Zool., 10:183, March 20, 1913. Sorex vagrans halicoetes, Jackson, N. Amer. Fauna, 51:108, July 24, 1928. Type. — Young adult male, skin and skull; No. 3638, Mus. Vert. Zool.; ob- tained on May 6, 1908, by Joseph Dixon from salt marsh near Palo Alto, Santa Clara Co., California. Range. — Marshes in the southern part of San Francisco Bay, California. Diagnosis. — Size small for the species; measurements of two topotypes are: total length, 105, 106; tail, 39, 40; hind foot, 12, 13. Upper parts in winter Chaetura Black or near Fuscous-Black; underparts brownish; upper parts in summer near ( I7'm ) Mummy Brown; underparts with a decided buffy wash, near (IS'd) Light Ochraceous BufiF; rostrum relatively large; maxillary tooth-row relatively long; teeth relatively large. Comparisons. — Darker ventrally, both summer and winter, than S. v. vagrans; slightly more reddish dorsally in summer pelage than S. v. vagrans, rostrum and teeth relatively larger; smaller externally than S. v. paludivagus, paler; skull longer, narrower cranially and broader rostrally. Remarks. — This subspecies seems to be restricted to salt marshes where it occurs with Sorex ornatus. Specimens examined. — Total number, 12. California: San Francisco Co. San Francisco, 4 BS. Alameda Co.: West Berkeley, 1 BS; Berkeley, 1 BS Dumbarton Point, 1 KU. San Mateo Co.: San Mateo, 2 BS. Santa Clara Co. Palo Alto, 3 BS. Marginal records. — California: Berkley, Elmhurst; Palo Alto; San Mateo. Sorex vagrans paludivagus von Bloeker Sorer vagrans paludivagus von Bloeker, Proc. Biol. Soc. Washington, 52:93, June 5, 1939. Type. — Adult male, skin and skull; No. 5053, Los Angeles Museum of History, Science and Art, obtained on November 3, 1938, by Jack C. von Bloeker, Jr., from salt marsh at mouth of Elkhom Slough, Moss Landing, Monterey Co., California, original no. 9456. Diagnosis. — Size medium for the species; average and extreme measurements of 6 topotypes are: total length, 115 (113-118); tail, 46.5 (42-48); hind foot, 14.5 (14-15) (von Bloeker, 1939:94). In winter nearly black dorsally, deep mouse gray ventrally; in summer nearly as dark dorsally as in winter, hairs of 60 University of Kansas Publs., Mus. Nat. Hist. venter tipped with Clove Brown; skull short, relatively broad cranially and relatively narrow rostrally. Comparisons. — For comparison with S. v. halicoetes see account of that subspecies. Remarks. — This subspecies, occurring at the limits of the range of the species, is uncommon in most collections. Seven specimens were available for the original description. The summer pelage is not completely described in the original description, but is stated to be darker than the winter pelage of S. v. vagrans, and must thus be considerably darker than the summer pelage of S. V. halicoetes. Two specimens in the Museum of Vertebrate Zoology, from San Gregario, referred by Jackson to S. v. halicoetes, were included in the present subspecies by von Blocker. Specimens examined. — None. Records of occurrence (von Blocker, 1939:94). — California: San Mateo Co.: San Gregario. Monterey Co.: Seaside; mouth of Salinas River; Moss Landing. Marginal records. — California: San Gregario; Seaside. Sorex vagrans vancouverensis Merriam Sorex vancouverensis Merriam, N. Amer. Fauna, 10:70, December 31, 1895. Sorex vagrans vancouverensis, Jackson, N. Amer. Fauna, 51:106, July, 1928. Type.— Adult male, skin and skull; No. 71913, U. S. Biol. Surv. Coll.; obtained on May 10, 1895, by Clark P. Streator, from Coldstream, Vancouver Island, British Columbia. Range. — Vancouver Island from Sayward south, and Bowen Island. Diagnosis. — Size small for the species; average and extreme measurements of 6 specimens from Albemi Valley, Vancouver Island, are: total length, 106.5 (97-115); tail, 41.7 (40-43); hind foot, 12 (11-13) (Jackson, 1928:107). Ventral parts brownish, winter pelage reddish browm rather than grayish. Comparisons. — Differs from S. v. vagrans in more brownish ventral parts and more brownish, rather than grayish, winter pelage; differs from the sym- patric S. v. isolatus in shorter tail, shorter hind foot, more narrow skull, and smaller teeth. Remarks. — This is a poorly differentiated subspecies which is closely related to S. V. vagrans. The differences in color noted are average ones. Some indi- viduals of this shrew might be difficult to separate from S. v. isolatus. The shght degree of morphological divergence is such that intergrades might be expected to occur. Possibly some habitat separation occurs, but such has not been reported. Specimens examined. — Total number, 3. British Columbia: Vancouver Is- land: Mt. Washington, 1 KU; Nanaimo, 1 BS; type locahty, 1 BS. Marginal records. — British Columbla: Sayward (Anderson, 1947:18); Bowen Island (Hall, 1938:463); Albemi (Jackson. 1928:107). CONCLUSIONS 1. Sorex vagrans, S. obscurus, S. pacificus, and S. yaqitinae are conspecific with one another. Each is a vaHd subspecies but all should bear the specific name Sorex vagrans Baird, 1858. Speciation of the Wandering Shrew 61 2. The subspecies of Sorex vagrans form a cline from large ( pacif- icus) to small {vagrans). The cline is bent in such a manner that the terminal subspecies occur together. Where the two subspecies occur together, individuals of one subspecies do not crossbreed with individuals of the odier subspecies and therefore react toward one another as do full species. Sorex vagrans vagrans occurs sympatrically with S. v. sonomae, S. v. pacificiis, S. v. yaquinae, S. v. bairdi, S. v. permiliensis, and S. v. setosus. S. V. vancouverensis occurs sympatrically with S. v. isolatus. 3. The sympatric existence of the terminal subspecies of the Sorex vagrans rassenkreis is made possible by marked differences be- tween them in size and in ecological preference. 4. The west-coast subspecies, sonomae, pacificiis, yaquinae, bairdi, and permiliensis probably differentiated from the Great Basin and Rocky Mountain subspecies, vagrans, obsctirus and monti- cola, during a separation caused first by aridity in the Great Basin, and secondly by glaciation of the Cascade Mountains and the Sierra Nevada, possibly in the Sangamonian and Wis- consinan ages respectively. 5. Sorex v. vagrans originated in the Great Basin and arrived on the Pacific Coast after the last deglaciation of the Cascades and Sierra Nevada. 6. In S. vagrans, heterogonic growth is illustrated; the larger the skull, the larger the rostrum in proportion to the skull as a whole. 7. In the species S. vagrans, size and color vary geographically more than do other features. 8. The S. ornatus group, S. longirostris, and S. veraepacis had a common ancestor with S. vagrans, possibly in the Illinoian Age. 9. S. vagrans, the S. ornatus group, S. veraepacis, S. longirostris, S. palustris, S. bendiri, and the S. cinereus group, because of structural resemblances, should be placed in a single subgenus, Otisorex. S. trowbridgi, the S. arcticus group, the S. saussurei group, S. merriami, S. fumeus, and S. dispar, should be included in the subgenus Sorex. 10. Sorex cinereus occurs with the medium-sized and large-sized S. vagrans in the Rocky Mountains and in Canada, but does not occur with the smaller subspecies of S. vagrans, probably be- cause competition between two shrews of hke size excludes S. cinereus. 62 University of Kansas Publs., Mus. Nat. Hist. Table 1 — Cranial Measurements of Sorex vagrans Catalog number or number of individuals averaged 8 av. Max Min. 707 AW. 706 AW. 205273 USES. 205270 USES. 205272 USES. 4 av. Max, Min. 6 av. Max Min. 6 av. Max Min. 14 av Max. Min.. c3 21.8 22.8 21.3 20.1 19.3 20.6 20.4 19.5 20.3 19.3 18.9 19.5 18.7 18.5 19.2 18.0 1=5 £.0 , rt .*.3 J2 u 0 (-. OJS 1 .^ _C -1.3 0 CO ^ ci .a QJ ►J Sorex vagrans pacificus, Orick, California. 9.6 8.6 10.4 4.1 10.2 9.0 11.1 4.3 9.3 8.4 10.2 4.1 6.6 6.8 6.4 Sorex vagrans yaquinae, Newport, Oregon. 8.9 8.8 7.6 7.3 9.3 9.3 3.7 4.0 5.7 5.8 Mapleton, Oregon. 9.0 8.9 9.2 8.2 7.9 8.3 9.9 9.3 4.2 3.7 4.0 6.0 6.0 6.1 Vida, Oregon. 8.4 8.8 8.2 7.5 7.9 7.3 9.2 9.5 8.9 3.6 3.7 3.5 5.5 5.7 McKenzie Bridge, Oregon. 8.2 8.6 8.0 7.1 7.6 6.8 9.1(5) 9.4 8.4 3.7 3.8 3.6 5.6 5.7 5.4 Sorex vagrans bairdi, Astoria, Oregon. 7.8 8.1 7.6 7.0 7.4 6.9 8.9(4) 3.4 9.0 3.5 8.9 3.2 5.3 5.5 5.2 Sorex vagrans permiliensis, Mt. Jefferson, Oregon. 18.0 18.9 17.2 7.5 6.8 9.0 3.5 7.9 7.2 9.4 3.7 7.2 6.5 8.6 3.3 5.2 5.3 4.8 Speciation of the Wandering Shrew 63 Table 1 — Continued Catalog number or number of individuals averaged 12 av, Max. Min. . 16 av Max. Min.. 15 av. Max. Min.. 10 av, Max. Min. . 18 av Max. Min.. 20550 AMNH . 20553 AMNH . 10 av Max. Min.. c3 en a -a a o— ' O -2 -73 -4-5 (U a Sorex vagrans setosus, Olympic Mts., Washington. 17.2(9) 17.9 16.7 7.1 7.4 6.9 6.5 8.5(8) 6.8 8.7 6.2 8.3 3.3 3.4 3.0 5.0 5.3 4.8 Mt. Rainier, Washington. 17.2 17.6 16.4 7.1 7.3 6.5 6.5 8.4(14) 3.3 6.7 8.7 3.5 6.1 8.1 3.2 5.0 5.2 4.7 Sorex vagrans longicauda, head Rivers Inlet, B.C. 18.0 18.4 17.6 7.4 7.6 7.2 6.7 6.9 6.4 8.7 8.9 8.4 3.2 3.3 3.1 5.0 5.2 4.8 Port Simpson, British Columbia. 18.1(9) 18.8 17.2 7.6 7.8 7.2 7.0 7.2 6.6 8.9 9.2 8.5 3.4 3.6 3.3 5.1 5.4 4.9 Fort WrangeU, Alaska. 18.5 18.9 17.8 7.8(15) 8.0 7.5 7.1 7.3 6.7 9.0(15) 3.3 9.2 3.5 8.6 3.2 5.1 5.3 5.0 Sorex vagrans elassodon, Woewodsky Is., Alaska. 18.0 17.5 7.5 7.1 6.7 6.3 8.7 8.3 3.3 3.2 5.0 4.7 Sorex vagrans alascensis, 9 mi. W and 4 mi. N Haines, Alaska. 17.2 7.2 6.7 8.5 3.1 4.9 17.6 7.4 6.9 8.8 3.3 5.0 16.9 6.9 6.5 8.2 3.0 4.7 64 University of Kansas Publs., Mus. Nat. Table 1 — Continued Hist. Catalog number or number of individuals averaged ^1 cS-D 73543 USES 73536 USES 73541 USES 18.0 18.0 17.9 7.5 7.6 7.4 Yakutat Bay, Alaska. 6.8 6.8 6.7 5.0 5.3 5.2 Sorex vagrans shumaginensis, Sandpoint, Popoff Is., Alaska. 9 av. Max Min. 5 av. Max. Min. 7 av. Max, Min. 20 av Max. Min.. 17.2(5) 17.6 16.8 7.0 7.2 6.8 6.3 6.6 6.1 8.3(7) 8.5 8.0 3.1 3.2 3.0 4.8 5.1 4.7 Sorex vagrans obscurus, Barkerville, Eritish Columbia. 17.1 17.3 16.6 7.1 7.3 6.7 6.5 6.6 6.4 8.5(4) 8.6 8.2 3.3 3.3 3.2 4.8 5.0 4.6 10 mi. SSW Leadore. Idaho. 17.2(4) 17.3 17.0 7.3(9) 7.5 7.1 6.6 6.8 6.4 8.6(4) 8.9 8.3 3.3 3.4 3.2 5.0 5.1 4.7 Albany Co., Wyoming (several localities ). 17.3 17.9 16.7 7.3 7.6 6.9 6.8 6.9 6.5 8.7(19) 9.0 8.4 3.2 3.4 3.1 5.2 5.5 5.0 Sorex vagrans longiqinis, 25 mi. ESE Eig Sandy, Montana. 87332 UM 87334 UM 87335 UM 10 av Max. Min.. 16.4 6.8 6.2 8.2 3.0 4.8 16.8 7.1 6.3 8.1 3.2 4.7 15.8 6.7 6.0 8.4 3.1 4.9 Ilighwood Mts., Montana. 16.3(9) 6.7 6.2 8.0(9) 3.1 4.7 16.9 6.9 6.4 8.3 3.3 5.0 15.6 6.5 6.0 7.8 3.0 4.5 Speciation of the Wandering Shrew 65 Table 1 — Continued Catalog number or number of individuals averaged 4 av. Max Min. 12 av Max. Min. . 55900 USES 55898 USES 55897 USES 7 av. Max, Min. 17 av, Max., Min. . 9 av. Max, Min. 4 av. Max, Min. fin . O t- I ax: c3 ^ Sorex vagrans neomexicanus, Cloudcroft, New Mexico. 17.6(3) 17.7 17.4 7.6 7.7 7.4 7.0 7.1 7.0 8.7 8.8 8.5 3.3 3.4 3.2 5.2 5.4 5.1 Sorex vagrans monticola, White Mts., Arizona. 16.1(9) 16.6 15.5 6.6 7.0 6.5 5.9(9) 6.1 5.6 8.2 8.5 8.1 3.1(11) 3.3 3.0 4.7 4.9 4.6 Sorex vagrans orizabae, Volcan Toluca, Mexico. 17.1 17.1 16.8 6.9 6.8 6.9 6.2 6.1 6.1 7.8 2.9 8.0 3.0 7.9 2.9 4.5 4.8 4.6 Sorex vagrans vagrans, Lincoln Co., Wyoming. 16.5(6) 17.1 16.0 6.6 7.0 6.4 6.1 6.4 5.9 8.2 8.5 7.9 2.9 3.1 2.9 4.7 4.9 4.5 Gearhart Mtn., Lake Co., Oregon. 16.5(15) 6.6 5.9 17.1 7.0 6.6 16.1 6.2 5.7 8.1 8.5 7.8 2.9 3.1 2.8 4.6 4.9 4.4 Willapa Eay, Washington. 16.6 17.2 16.2 6.8 7.1 6.6 6.2 6.4 5.9 8.1 8.3 7.9 2.8 3.1 2.7 4.7 5.0 4.6 Sorex vagrans obscuroides, Eishop Creek, California. 16.7 16.8 16.6 7.0 7.1 6.9 6.3 6.4 6.2 8.2 8.3 8.1 3.2 3.4 3.1 4.8 4.9 4.7 5—7903 66 University of Kansas Publs., Mus. Nat. Table 1 — Concluded Hist. Catalog number or number of individuals averaged 4 av Max Min 56559 USES 56558 USES '. bC c — 1— < ci ->J t- & O >-.s. JS QJJ3 JZ C3 fc£ 'z:* -^ c5 c3 rt ^ ^ • •— " s ■^ (» X o 03 JD 1— ( r< u h-3 Mt. Whitney, California. 16.7(3) 6.9 6.3 8.4 3.3 16.7 7.0 6.4 8.5 3.4 16.7 6.7 6.1 8.4 3.1 4.8 5.0 4.7 Sorex vagrans parvidens, San Eernardino Peak, California. 17.1 16.4 7.0 6.8 6.1 6.1 8.0 8.0 2.9 3.0 4.8 4.8 Anderson, 1947. Anderson, 1945. Bailey, V. 1936. Brown, B. 1908. Clothier, 1950. Conaway, 1952. Cowan, I. 1936. 1941. LITERATURE CITED R. M. Catalogue of Canadian Recent mammals, Nat. Mus. Canada, Bull. 102, Biol. set. 31:i-v + 1-238, January 24. R. M. and A. L. Rand A new form of dusky shrew from the prairie provinces of Canada. Canadian Field-Nat., 59:47-48, March- April. The mammals and life zones of Oregon. N. Amer. Fauna, 55:1-416, 52 pis., 102 figs, in text, August 29. The Conard Fissure, a Pleistocene bone deposit in northern Arkan- sas: with descriptions of two new genera and twenty new species of mammals. Mem. Amer. Mus. Nat. Hist., 9:157-208, pis. 14-25, February. R. R. Contribution to the taxonomy and life historj' of Sorex vagrans monticola and Sorex ohscurus obscurus. Master's thesis, Mon- tana State University, Missoula, Montana — a manuscript. C. H. Life history of the water shrew (Sorex palustris navigator). Am. Midi. Nat., 48:219-248, 6 tables, 9 figs, in text, July. McT. Distribution and variation in deer (Genus Odocoileus) of the Pacific coastal region of North America. California Fish and Game, 22(3):155-246, 13 figs., 3 graphs, 8 tables, July. Insularity in the genus Sorex on the north cost of British Columbia. Proc. Biol. Soc. Washington, 54:95-108, July 31. Speciation of the Wandering Shrew 67 Dalquest, W. W. 1941. Ecologic relationships of four small mammals in western Washing- ton. Jour. Mamm., 22:170-173, May 14. 1944. The molting of the wandering shrew. Jour. Mamm., 25:146-148, one fig. in text, May 25. 1948. Mammals of Washington. Univ. Kansas Publ., Mus. Nat. Hist., 2:1-444, 140 figs, in text, April 9. Davis, W. B. 1939. The Recent Mammals of Idaho. The Caxton Printers, Ltd., Cald- well, Idaho. Pp. 1-400, 33 figs, in text, 2 pis., April 5. DURRANT, S. D. 1952. Mammals of Utah, taxonomy and distribution. Univ. Kansas Publ., Mus. Nat. Hist., 6:1-549, 91 figs, in text, 30 tables, August 10. FiNDLEY, J. S. 1953. Pleistocene Soricidae from San Josecito Cave, Nuevo Leon, Mexico. Univ. Kansas Publ., Mus. Nat. Hist., 5:633-639, December 1. 1955. Taxonomy and distribution of some American shrews. Univ. Kan- sas Publ., Mus. Nat. Hist., 7:613-618, June 10. Fitch, H. S. 1940. A biogeographical study of the ordinoides artenkreis of garter snakes (genus Thamnophis). Univ. California Publ. Zool., 44: 1-150, October 31. Grinnell, J. 1933. Review of the Recent mammal fauna of California. Univ. Cali- fornia Publ. Zool., 40:71-234, September 26. Grinnell, J., and A. H. Miller. 1944. The distribution of tlie birds of California. Pacific Coast Avifauna, 27:1-608, 57 figs, in text, December 30. Hall, E. R. 1938. Variation among insular mammals of Georgia Strait, British Co- lumbia. Amer. Nat., 72:453-463. 1946. Mammals of Nevada. University of California Press, Berkeley and Los Angeles, pp. i-xi -f- 1-710, 11 pis., 485 figs, in text, July 1. Hamilton, W. J., Jr. 1940. The biology of the smoky shrew {Sorex fumeus fumeus Miller). Zoologica, 25:473-492, 4 pis., 1 fig. in text, 2 tables. Hibbard, C. 1944. Stratigraphy and vertebrate paleontology of Pleistocene deposits of southwestern Kansas. Geol. Soc. America, Bull. 55:707-754, 3 pis., 20 figs, in text, June. HovraLL, T. R. 1952. Natural history and differentiation in the yellow-beUied sapsucker. Condor, 54:237-282, September 22. Jackson, H. H. T. 1928. A taxonomic review of the American long-tailed shrews (genera Sorex and Microsorex). N. Amer. Fauna, 51:i-vi-|- 1-238, 13 pis., 24 figs, in text, July 24. 1947. A new shrew (genus Sorex) from Coahuila. Proc. Biol. Soc. Washington, 60:131-132, October 9. 68 University of Kansas Publs., Mus. Nat. Hist. KRxrrzscH, P. H. 1954. North American jumping mice ( genus Zapus ) . Univ. Kansas Publ., Mus. Nat. Hist., 7:349-472, 47 figs, in text, 4 tables, April 21. Macnab, J. A., and J. C. Dirks. 1941. The Cahfomia red-backed mouse in the Oregon Coast Range. Jour. Mamm., 22:174-180, May 14. Mayr, E. 1940. Speciation phenomena in birds. Amer. Nat., 74:249-278. Merriam, C. H. 1895. Synopsis of the American shrews of the genus Sorex. in N. Amer. Fauna, 10:57-100, December 31. 1899. Results of a biological survey of Mt. Shasta, Cahfomia. N. Amer. Fauna, 16:1-179, 46 figs, in text, 5 pis., October 28. MURIE, A. 1933. The ecological relationship of two species of Peromyscus in the Glacier Park region, Montana. Occ. Pap. Mus. Zool., Univ. Mich- igan, 270:1-17, 2 figs., 3 tables, July 1. Pearson, O. P. 1945. Longevity of the short-tailed shrew. Amer. Midi. Nat., 34:531-546, 2 tables, 4 figs, in text, September. Prihtt, W. O., Jr. 1954. Aging in the masked shrew, Sorex cinereus cinereus Kerr. Jour. Mamm., 35:35-39, February 10. Rensch, B. 1933. Zoologische systematik und artbildungsproblem. Ver. deutsch. zool. Gesellschaft, 1933:19-83. RlDGWAY, R. 1912. Color standards and color nomenclature. Washington, D. C, pri- vately printed, i-iv -f- 1-44, 53 pis. RuDD, R. L. 1953. Difi^erentiation in shrews of the tidal marshes of the San Francisco Bay region. Summary of the dissertation for the degree of Doctor of Philosophy, University of California Graduate Division, 4 pages, unnumbered, June. Simpson, G. G. 1945. The principles of classification and a classification of mammals. Bull. American Mus. Nat. Hist., 85:i-xvi + 1-350, October 5. TlCEHtTRST, Cl. B. 1938. A systematic review of the genus Phylloscopus. British Mus., London, i-viii + 1-193, 8 maps, 2 pis., November 26. Van Den Brink, F. H. 1953. La musaraigne masquee, espece circum-boreale. Mammaha, 17: 96-125, 1 map, June. D 26-7903 (Continued from inside of front cover) 17. The subspecies of the Mexican red-bellied squirrel, Sciurus aureogaster. By Keith R. Kelson. Pp. 243-250, 1 figure in text. April 10, 1952. 18. Geographic range of Peromyscus melanophrys, with description of new sub- species. By Rollin H. Baker. Pp. 251-258, 1 figure in text. May 10, 1952. 19. A new chipmunk (Genus Eutamias) from the Black Hills. By John A. White. Pp. 259-262. April 10, 1952. 20. A new pifion mouse (Peromyscus truei) from Durango, Mexico. By Robert B. Finley, Jr. Pp. 263-267. May 23, 1952. 21. An annotated checklist of Nebraskan bats. By Olin L. Webb and J. Knox Jones, Jr. Pp. 269-279. May 31, 1952. 22. Geographic variation in red-backed mice (Genus Clethrionomys ) of the south- em Rocky Mountain region. By E. Lendell Cockrum and Kenneth L. Fitch. Pp. 281-292, 1 figure in text. November 15, 1952. 23. Comments on the taxonomy and geographic distribution of North American microtines. By E. Raymond Hall and E. Lendell Cockrum. Pp. 293-312. November 17. 1952. ^ 24. The subspecific status of two Central American sloths. By E. Raymond Hall and Keith R. Kelson. Pp. 313-317. November 21, 1952. 25. Comments on the taxonomy and geographic distribution of some North Ameri- can marsupials, insectivores, and carnivores. By E. Raymond Hall and Keith R. Kelson. Pp. 319-341. December 5, 1952. 26. Comments on the taxonomy and geographic distribution of some North Ameri- can rodents. By E. Raymond Hall and Keith R. Kelson. Pp. 343-371. December 15, 1952. 27. A synopsis of the North American microtine rodents. By E. Raymond Hall and E. Lendell Cockrum. Pp. 373-498, 149 figures in text. January 15, , 1953. 28. The pocket gophers (Genus Thomomys) of Coahuila, Mexico. By Rollin H. Baker. Pp. 499-514, 1 figure in text. June 1, 1953. 29. Geographic distribution of the pocket mouse, Perognathus fasciatus. By J. Knox Jones, Jr. Pp. 515-526, 7 figures in text. August 1, 1953. 30. A new subspecies of wood rat (Neotoma mexicana)- from Colorado. By Robert B. Finley, Jr. Pp. 527-534, 2 figures in text. August 15, 1953. 31. Four new pocket gophers of the genus Cratogeomys from Jalisco, Mexico. By Robert J. Russell. Pp. 535-542. October 15, 1953. 32. Genera and subgenera of chipmunks. By John A. White. Pp. 543-561, 12 figvu-es in text. Deceniber 1, 1953. 33. Taxonomy of the chipmunks, Eutamias quadrivittatus and Eutamias um- brinus. By John A. White. Pp. 563-582, 6 figures in text. December 1, 1953. 34. Geographic distribution and taxonomy of the chipmunks of Wyoming. By John A. MTiite. Pp. 584-610, 3 figures in text. December 1, 1953. 35. The baculum of the chipmmiks of western North America. By John A. White. Pp. 611-631, 19 figures in text. December 1, 1953. 36. Pleistocene Soricidae from San Josecito Cave, Nuevo Leon, Mexico. By James S. Findley. Pp. 633-639. December 1, 1953. 37. Seventeen species of bats recorded from Barro Colorado Island, Panama Canal Zone. By E. Raymond Hall and William B. Jackson. Pp. 641-646. De- cember 1, 1953. ' Index. Pp. 647-676. •Vol. 8. (Complete) Mammals of Utah, taxonomy and distribution. By Stephen D. Dur- rant. Pp. 1-549, 91 figures in text, 30 tables. August 10, 1952. Vd. 7. *1. Mammals of Kansas. By E. Lendell Cockrum. Pp. 1-303, 73 figures in text, 37 tables. August 25, 1952. 2. Ecology of the opossum on a natural area in northeastern Kansas. By Henry S. Fitch and Lewis L. Sandidge. Pp. 305-338, 5 figures in text. August 24. 1953. ^ 3. The silky pocket mice (Perognathus flavus) of Mexico. By Rollin H. Baker. Pp. 339-347, 1 figure in text. February 15, 1954. 4. North American jumping mice (Genus Zapus). By Philip H. Krutzsch. Pp. 349-472, 47 figures in text, 4 tables. April 21, 1954. 5. Mammals from Southeastern Alaska. By Rollin H. Baker and James S. Find- ley. Pp. 473-477. April 21. 1954, 6. Distribution of some Nebraskan Mammals. By J. Knox Jones. Jr. Pp. 479- 487. April 21. 1954. 7. Subspeciation in the montane meadow mouse, Microtus montantis. in Wyoming and Colorado. By Sydney Anderson. Pp. 489-506, 2 figures in text. July 23, 1954. (Continued on outside of back cover) (Continued from inside of back cover) 8. A new subspecies of bat (Myotis velifer) from southeastern California and Arizona. By Terry A. Vaughn. Pp. 507-512. July 23, 1954. 9. Mammals of the San Gabriel Mountains of CaKfomia. By Terry A. Vaughn. Pp. 513-582, 1 figure in text, 12 tables. November 15, 1954. 10. A new bat (Genus Pipistrellus) from Northeastern Mexico. By RoUin H. Baker. Pp. 583-586. November 15, 1954. 11. A new subspecies of pocket mouse from Kansas. By E. Raymond Hall. Pp. 587-590. November 15, 1954. ^ J2. Geographic variation in the pocket gopher, Cratogeomys castanops, in Coa- huila, Mexico. By Robert J. Russell and Rollin H. Baker. Pp. 591-608. March 15, 1955. 13. A new cottontail (Sylvilagus floridanus) from northeastern Mexico. By Rollin H. Baker. Pp. 609-612. April 8, 1955. 14. Taxonomy and distribution of some American shrews. By James S. Findley. Pp. 613-618. June 10, 1955. 15. The pigmy woodrat, Neotoma goldmani, its distribution and systematic posi- tion. By Dennis G. Rainey and Rollin H. Baker. Pp. 619-624, 2 figs, in text. June 10, 1955. Index. Pp. 625-651. Vol. 8. 1. Life history and ecology of the five-lined skink, Eumeces fasciatus. By Henry S. Fitch. Pp. 1-156, 26 figvires in text. September 1, 1954. 2. Myology and serology of the avian family Fringillidae, a taxonomic study. By WilUam B. Stallcup. Pp. 157-211, 23 figures in text, 4 tables. November 15, 1954. More numbers will appear in volume 8. Vol. 9. 1. Speciation of the wandering shrew. By James S. Findley. Pp. 1-68, 18 figures ' in text. December 10, 1955. More nimibers will appear in volume 9. • A-Cd li//ftv» Ci^Aj University of Kansas Publications Museum of Natural History Volume 9, No. 2, pp. 69-80 — December 10, 1955 — Additional Records and Extensions of Known Ranges of Mammals from Utah BY STEPHEN D. DURRANT, M. RAYMOND LEE, AND RICHARD M. HANSEN m. CO?;!P. ZOCL FEB 9195P University of Kansas Lawrence 1955 University of Kansas Piiblications, Museijm of Natural, History Editors: E. Raymond Hall, Chairman, A. Byron Leonard, Robert W. Wilson Volume 9, No. 2, pp. 69-80 Published December 10, 1955 Untveesity of Kansas Lawrence, Kansas PRINTED BY FERD VOILAND. JR.. STATE PRINTER TOPEKA. KANSAS 1955 25-8618 Additional Records and Extensions of Known Ranges of Mammals From Utah FEB 91958 I' » "»",'( ■ 1 BY STEPHEN D. DURRANT, M, RAYMOND LEE, AND RICHARD M. HANSEN The Museum of Zoology, University of Utah, contains approxi- mately 5000 specimens in addition to those available to Durrant (1952) when he prepared his account of the "Mammals of Utah, Taxonomy and Distribution." Study of this material discloses two kinds of mammals not heretofore known to occur in Utah, and ex- tends the known limits of occurrence of many others as is set forth below in what may be thought of as a supplement to the aforemen- tioned report of 1952. Our study was financed in part by a grant from the National Science Foundation. Sorex vagrans obscurus Merriam. Dusky Shrew. — ^Twelve speci- mens are available from the Abajo Mountains and Elk Ridge, San Juan County, Utah, as follows: North Creek, 6 mi. W Monticello, 8300 ft.; 1 mi. S Twin Peaks, 9500 ft.; Kigaha R. S., 8000 ft.; and Gooseberry R. S., 8250 ft. Previously, the only known specimens from east of the Colorado River in Utah were from the La Sal Moun- tains in extreme eastern Grand County and extreme northern San Juan County. These twelve specimens extend the known area of occurrence of the species in Utah approximately 80 miles to the south, and indicate that this shrew occurs throughout the state in favorable habitats. Sorex palustris navigator (Baird). Water Shrew. — An individual was observed by M. Raymond Lee at North Creek, seven miles west of Monticello, Abajo Mountains, 8000 feet, San Juan County, on July 10, 1954. Usually we are extremely reluctant to record sight records, but do so in this instance because the water shrew is so distinctive that it can be readily recognized and because the occurrence extends the known range approximately 80 miles south- ward in Utah. This individual was observed at close range while swimming and foraging in North Creek, and there can be no doubt of its identity. Myotis yumanensis yumanensis (H. Allen). Yuma Myotis. — Dur- (71) 72 University of Kansas Publs., Mus. Nat. Hist. rant (1952:43) reported this subspecies from Utah on the authority of Hardy (1941:289) who had specimens from two locaHties in extreme southwestern Utah. Durrant (1952:41) referred speci- men Number 6784, from Willow Creek, 25 miles south of Ouray, Uintah County, to Myotis lucifugus carissima. Restudy of this specimen reveals that it is Myotis ywnanensis yumanensis. This identification has been corroborated by Dr. Philip H. Krutzsch, University of Pittsburgh, Pennsylvania, and extends the known range of the subspecies M. y. ywnanensis approximately 300 miles northeastward in Utah. See also Krutzsch and Heppenstall (1955: 126 ) who record specimens from 2 mi. SW Jensen. Myotis subulatus melanorhinus ( Merriam ) . Small-footed Myotis. — This bat previously was known from only seven localities in Utah, which indicated that it occurred in only the western and southern areas of the state. Four additional records are now available from the following localities: Logan Canyon Cave, 15 miles north of Logan, Cache County; Weber College Campus, Ogden, Weber County; University of Utah Campus, Salt Lake City, Salt Lake County; Six Mile Canyon, 3/2 miles east of Sterling, Sanpete County. These occurrences extend the known range to the eastward in Utah, and indicate a state-wide distribution. Specimens of the subspecies Myotis s. melanorhinus are recorded also from as far north as Double Springs, Custer County, Idaho (Davis, 1939:117). Pipestrellus hesperus hesperus (H. Allen). Western Pipistrelle. — Heretofore, the northermost known specimens of this bat from Utah were from Old Lincoln Highway, 18 miles southwest of Orr's Ranch, Tooele County. Specimen Number 7531 is now available from cliffs NE [3 mi.] Ogden, Weber County, and extends the known range of this species in Utah approximately 100 miles north- eastward. This pipistrelle probably inhabits all of northern and northwestern Utah in suitable habitats. This probability is sup- ported by Davis' (1939:120) report of a specimen from Salmon Creek, eight miles west of Rogerson, Twin Falls County, Idaho. See also Krutzsch and Heppenstall (1955:127) who record a speci- men from, eastern Utah as far north as, Desert Springs which is 10 mi. SW Ouray, Uintah County. Corynorhinus rafinesquii pallescens Miller. Long-eared Bat. — Formerly, the northernmost record of the long-eared bat in Utah was from east of Springville, Utah County. Specimens are now available from Goldhill, Tooele County, and from South Fork, Ogden River, Weber County. Professor J. S. Stanford, Department Mammals from Utah 73 of Zoology, Utah State Agricultural College, infoiined us (by letter) that this bat is the common cave bat in Logan Canyon, Cache County. This northern extension of known area of occur- rence of approximately 100 miles indicates that it probably is state- wide in distribution in suitable habitats. It can be inferred from Hall (1946:161) that the range of C. r. intermedius in Nevada ex- tended northeastward into northwestern Utah, and Davis (1939: 124) reported specimens from Bingham and Bannock counties, Idaho, that he referred to the above mentioned subspecies. This led Stanford to comment (in litt.) that bats of this species from northern Utah in Cache County might be C. r. intermedius. Insofar as we are aware, C. r. pallescens differs from C. r. intermedius only in being slightly paler. Our specimens from Goldhill and South Fork of the Ogden River are not beyond tlie range of color of speci- mens from elsewhere in the state that are referable to C r. pal- lescens. Inasmuch as specimens are not available from Logan Canyon, we deem it best pending the acquisition of specimens from that locality to refer all members of this species from Utah to the subspecies C. r. pallescens. Marmota flaviventer nosophora Howell. Yellow-bellied Marmot. — Durrant (1952:101) did not indicate that any species of the genus Marmota occurred on the mountains within the basin of Pleistocene Lake Bonneville. Furthermore, he commented (op. cit. :502) upon the dearth of sciurids within this basin. One speci- men. No. 10,905, of the subspecies M. /. nosophora has been taken from South Willow Canyon, 10,000 feet, base of Deseret Peak, Stansbury Mountains, Tooele County. This specimen is noteworthy not only in that it extends the known range of this kind of mammal 50 miles to the west in Utah, but in that it is well within the basin of the ancient lake. The marmot is common in the Wasatch Moun- tains on the eastern mainland of Lake Bonneville, but to date has not been found on the Oquirrh Mountains immediately to the west. The Oquirrh Mountains are interposed between the Stansbury and Wasatch mountains. The presence of the marmot on the Stans- bury Mountains indicates that it probably occurs also on the Oquirrh Mountains. Citellus beldingi crebrus Hall. Belding Ground Squirrel. — Dur- rant (1952:113) had only two specimens of this ground squirrel from Standrod, Boxelder County. Additional specimens have been obtained from the following localities in northwestern Boxelder County: Grouse Creek, Park Valley, Grouse Creek Mountains, 74 University of Kansas Publs., Mus. Nat. Hist. 12 miles northwest of Grouse Creek, and Goose Creek. C. b. cre- brus now is known to inhabit all the major drainages of the Raft River, Goose Creek, and Grouse Creek mountains. In addition to extending the known area of occurrence of this animal in Utah, these specimens prove also that this species is not restricted to the Snake River Drainage as Durrant (1952:113) supposed, but occurs also in the Great Basin Drainage. Citellus richardsonii elegans (Kennicott). Richardson Ground Squirrel. — Recently, Hansen (1953:132) reported on specimens of this species from Rich and Summit counties. Additional specimens are now available from Highway 165 [2 mi. E Summit — Daggett Co. Line], 2 miles south of Utah-Wyoming State Line; 5 miles west of Manila, and one mile northeast of Manila (Carnegie Museum). These localities are in Daggett County. The occurrence of these ground squirrels in Rich, Summit and Daggett counties suggests that they occur along the entire northern piedmont of the Uinta Mountains. Citellus lateralis trepidus (Taylor). Golden-mantled Ground Squirrel. — Durrant (1952:126) estimated that practically all of the area in Utah that is within the Great Basin might be included in the range of this subspecies. Actually, he had specimens from only the Raft River Mountains in northwestern Boxelder County. He included sight records from the Deep Creek Mountains and from the Oquirrh Mountains. Subsequently t\vo specimens, numbers 7469A and 7470A, were obtained from the Deep Creek Mountains. To date neither specimens nor subsequent sight records have been obtained from the Oquirrh Mountains, and we are of the opinion that Durrant erred, and that the golden-mantled ground squirrel does not inhabit these mountains. Therefore, it seems at this writ- ing that this subspecies, in Utah, occurs only in the extreme western and northwestern parts of the state in the Deep Creek and Raft River mountains, and not within the basin of Pleistocene Lake Bonneville as formerly supposed. Eutamias umbrinus umbrinus ( J. A. Allen ) . Uinta Chipmunk. — Two specimens, numbers 10,236 and 10,237, from the junction of Argyle and Minnie Maud creeks. Carbon County, prove that mem- bers of this subspecies occur on the West Tavaputs Plateau, which is outside the range ascribed to this subspecies by White (1953:572) and by Durrant (1952:142). The grayish color of these specimens and the reduction of ochraceous pigments constitute basis for re- ferring fhe specimens to E. u. umbrinus, and not to E. u. adsitus or Mammals from Utah 75 E. u. montanus. E. u. umbrinus on the West Tavaputs Plateau is separated from E. u. montanus on the East Tavaputs Plateau by the Green River and its deep chasm. Perognathus formosiis incolatus Hall. Long-tailed Pocket Mouse. — Prior to the description of this subspecies by Hall (1941:56), animals of this species had not been reported from within the basin of the Pleistocene Lake Bonneville. When Durrant (1952) prepai-ed his manuscript he had but a single specimen from western Millard County and one nearby record (Fautin, 1946:280). Additional specimens are known from the following localities: junction of Trout Creek and Birch Creek, Deep Creek Mountains, Tooele County; six miles north of Ibapah, Tooele County; five miles south of Timpie, Tooele County; north end of Newfoundland Mountains, Boxelder County; and Groome, Boxelder County. These occur- rences show that the species is not restricted to the extreme western part of the state, but occurs in suitable habitats throughout the basin of Pleistocene Lake Bonneville. The known range is extended approximately 150 miles north and 40 miles east. Perognathus parvus trumbullensis Benson. Great Basin Pocket Mouse. — Durrant (1952:477), suspecting that this mouse occurred in Utah, included the subspecies P. p. trumbullensis in his hypo- thetical list. Numerous specimens are now available from the fol- lowing localities: Pine Valley Mountains, Enterprise Reservoir, and 19 miles west of Enterprise, Washington County; Bown's Reser- voir, Snow Ranch, Hall Ranch, Steep Creek, Garfield County; Aquarius Guard Station, Aquarius Plateau, Wayne County. Inso- far as we are aware, these occurrences are the first to be recorded from Utah, and extend the known range of this subspecies 150 miles northward. The specimens from Washington County are paler than those from Garfield County, and this pallor indicates intergradation with the subspecies P. p. olivaceus. Of animals from the Aquarius Pla- teau, those from the eastern and southern localities are pale and have a marked suffusion of ochraceous in the upper parts, whereas those from the western and northern localities are extremely dark owing to a heavy suffusion of black in the upper parts. The skulls of animals from the Aquarius Plateau resemble those of P. p. trum- bullensis in the majority of diagnostic characters. In some few characters, nevertheless, the skulls resemble those of P. p. olivaceus, and in other characters are intermediate between these two named subspecies. In shape and size of the interparietal, in slightly longer 76 University of Kansas Publs., Mus. Nat. Hist. nasals, and in slightly greater alveolar length of upper molariform teeth, animals from the Aquarius Plateau differ from either of the aforementioned subspecies. All characters considered, we deem it best to refer these specimens to the subspecies P. p. trumbullensis. Thomomys talpoides bridgeri Merriam. Northern Pocket Gopher. — An adult female. Catalogue No. 25667 of the Museum of Natural History of the University of Kansas, skin w^ith skull, was trapped, on 30 June 1948, 14 miles south and 2 miles east of Robertson, 9,300 feet, in Summit County, Utah, by James O. Lonnquist (original number 146). This is the first record of this subspecies from Utah, and raises to 37 the named kinds of pocket gophers known from Utah. Reithrodontomys megalotis megalotis ( Baird ) . Western Harvest Mouse. — Durrant (1952:295) reported no harvest mice from the Uinta Basin in northeastern Utah. One specimen. No. 10,239, was obtained from two miles east of Duchesne, Duchesne County. This specimen extends the known range in Utah 50 miles northward, and indicates that the harvest mouse of the subspecies R. m. mega- lotis occurs throughout the Uinta Basin. Onychomys leucogaster pallescens Merriam. Northern Grasshop- per Mouse. — Hansen obtained specimens (in alcohol) from Ken- nedys Hole, junction of the White and Green rivers, Uintah County. The northernmost specimens available to Durrant (1952:328) were from one mile east of Greenriver, Grand County. These specimens from Uintah County extend the known range 80 miles to the north, and substantiate Durrant's conclusion that this subspecies occurs east of the Green and Colorado rivers. Clethrionomys gapperi uintaensis Doutt. Red-backed Mouse. — Previously, the red-backed mouse in Utah was known only from the Uinta and Wasatch mountains. The southernmost localities from which specimens were available were in northern Wasatch County and southern Salt Lake County. Durrant (1952:355) supposed that the species ranged southward to Mount Timpanogos in Utah County. One specimen. No. 10,075, from the summit, 18 miles east of Mayfield, Sanpete County, and 4 from Ephraim Canyon, 15 miles east of Ephraim, Sanpete County, show that this subspecies occurs also on the Wasatch Plateau of central Utah. These latter specimens extend the known range of the red-backed mouse in Utah approximately 100 miles southward. Owing to the prac- tically continuous nature of the central mountain ranges of Utah, students of mammals of Utah usually suspect that most montane Mammals from Utah 77 mammals occur throughout these mountain ranges. The red-backed mouse has been sought for in vain in the mountains south of the Wasatch Plateau. Suitable habitats for this mouse occur through- out the Fishlake Mountains, Thousand Lake Mountains and the Aquarius Plateau, but despite intensive collecting, none has been obtained from these areas. Phenacomys intermedins intermedius Merriam. Heather Vole. — The heather vole, while not rare, is uncommon in Utah. Durrant (1952:360) had but eight specimens from various localities in Sum- mit, Wasatch, Salt Lake and Utah counties and supposed that the species was restricted to the western Uinta Mountains and southern Wasatch Mountains. In the summers of 1952 and 1953, intensive collecting of mammals was carried out on Boulder Mountain and the Aquarius Plateau, in Wayne and Garfield counties. Two specimens, nos. 8956 and 9074, were obtained from Spectacle Lake, Boulder Mountain, Garfield County. These specimens extend the known area of occurrence 175 miles southward in Utah. No specimens are known from the areas between Mount Timpanogos in Utah County, and Boulder Mountain in Garfield County. We suspect, however, that when this intervening area has been thoroughly studied, the heather vole, like many other montane mammals, will be found throughout the entire length of the central mountain ranges. Microtus pennsylvanicus modestus (Baird). Pennsylvanian Meadow Mouse. — In Utah this mouse was known only from wet meadows in valleys immediately west of the Wasatch Mountains, as far south as a place 2 miles south of Provo ( Hall and Cockrum, 1953:410). Norman V. Chamberlain collected several specimens "near" Koosharem Reservoir, Sevier County. These extend the known range of this subspecies 110 miles southward, and suggest that Pennsylvanian meadow mice occur, in suitable habitat, all along the eastern margin of the Great Basin in Utah, at least as far south as Sevier County. All northern specimens are from the drainages of Utah Lake and Great Salt Lake, but these specimens from Sevier County are from the Sevier River Drainage. This species requires a fairly moist environment, and such habitat exists between the aforementioned drainages which are practically inter- connected by Mona Reservoir and its adjacent areas of springs. Zapus princeps utahensis Hall. Big Jumping Mouse. — Durrant (1952:388) reported a specimen from Puffer Lake, Beaver Moun- tains, Beaver County. He supposed that this mouse occurred also at high elevations still farther south in Utah. Subsequently, two 78 University of Kansas Publs., Mus. Nat. Hist. specimens were obtained from Garfield County; No. 9006 from Steep Creek, 12 miles north of Boulder and No. 9071 from East Fork of Boulder Creek, 10 miles north of Boulder. These two speci- mens extend the known range of jumping mice in Utah 75 miles southeastward. Several specimens have been obtained also from the Fishlake Plateau, and further bear out Durrant's supposition that these mammals occur on all of the high mountains of central Utah, at least as far south as the Aquarius Plateau. Urocyon cinereoargenteus scottii Meams. Gray Fox. — One skull, No. 10,240, from mouth of Birch Creek, Deep Creek Mountains, Juab County, extends the known geographic range 50 miles east from Cherry Creek Canyon, Nevada (see Hall, 1946:241). This record indicates that the species occurs in the mountainous areas on the western margins of Pleistocene Lake Bonneville, and extends the known range in Utah approximately 150 miles northward. Fur- thermore, this record proves that the gray fox occurs as far north in western Utah as it does in eastern Nevada, but to date none has been obtained from the mountains within the basin of the ancient lake, even though some of them are not far removed from the Deep Creek Mountains. Lutra canadensis nexa Goldman. River Otter. — River otters are rare in a semi-arid state like Utah, and few have been preserved as scientific specimens. Durrant (1952:436) had access to but one skull from an immature animal from the Raft River Mountain area in northwestern Boxelder County. At present there are two com- plete specimens (skins, skulls and skeletons) in the collection of the University of Utah. They were trapped by an employee of the Utah State Fish and Game Department, and were donated to the University of Utah by J. Perry Egan, Director of the above men- tioned department. They are nos. 8854 and 8855, and are from the Raft River, 2 miles south of the Utah-Idaho border, Boxelder County. Alces americanus shirasi Nelson. Moose. — The moose is rare in Utah, and to date records of its occurrence have been based solely upon sight records. There are, nevertheless, two specimens pre- served. One is a young bull (skull only) from Farmington Canyon, Davis County, in the collection of Weber College, Ogden, Utah. The other is one antler (No. 10,745) of a young bull from Henrys Fork, 16 miles south of the Utah-Wyoming border, Summit County, and it is in the collection of the University of Utah. This large cervid apparently is increasing in numbers in the state. Dale Jones of the Mammals from Utah 79 Utah State Fish and Game Department reported to us that a herd of 25 animals was observed in 1954, in the vicinity of Haydens Peak, Bear River Drainage, Summit County. A cow and a calf were seen in the vicinity of Strawberry Reservoir, Wasatch County, in 1951, by employees of the same department. This latter locality is the most southern and eastern point of their known occurrence in Utah. Ovis canadensis canadensis Shaw. Mountain Sheep. — Formerly, the mountain sheep was not known to occur in the La Sal Moun- tains in Grand and San Juan counties. On October 23, 1954, a two year old ram, No. 10,906, was killed by a deer hunter at a locality 1/2 miles north of La Sal, La Sal Mountains, San Juan County. This constitutes the first complete specimen ( skin and skull ) of a moun- tain sheep from Utah. According to Harold Crane, of the Utah State Fish and Game Department, this ram was running with a herd of mule deer, and was the only mountain sheep that was seen. The ram was confiscated and given to the Department of Zoology, Uni- versity of Utah, for preservation as a scientific specimen. 80 University of Kansas Publs., Mus. Nat. Hist. LITERATURE CITED Davis, W. B. 1939. The Recent Mammals of Idaho. The Caxton Printers, Ltd., Cald- well, Idaho, 400 pp., 2 full-page half tones, 33 figs, in text, April 5. DURRANT, S. D. 1952. Mammals of Utah, taxonomy and distribution. Univ. Kansas Publ., Mus. Nat. Hist., 6:1-549, 91 figs, in text, 30 tables, August 10. Fautin, R. W. 1946. Biotic communities of the northern desert slirub biome in Western Utah. Ecol. Monogr., 16:251-310, 19 figs, in text, 33 tables, Octo- ber. Hall, E. R. 1941. New heteromyid rodents from Nevada. Proc. Biol. Soc. Washing- ton, 54:55-61, May 20. 1946. Mammals of Nevada. Univ. Cafifornia Press, Berkeley, California, xi 4- 710 pp., 11 pis., 485 figs, in text, July 1. Hall, E. R., and Cockrum, E. L. 1953. A synopsis of the North American microtine rodents. Univ. Kansas Publ., Mus. Nat. Hist., 5:373-498, January 15. Hansen, R. M. 1953. Richardson ground squirrel in Utah. Jour. Mamm., 34:131-132, February 9. Hardy, R. 1941. Some notes on Utah bats. Jour. Mamm., 22:289-295, August 14. Krutzsch, p. H., and Heppenstall, C. A. 1955. Additional distributional records of bats in Utah. Jour. Mamm., 36:126-127, February. White, J. A. 1953. Taxonomy of the chipmunks, Eutamias quadrivittatus and Euta- mias umbrinus. Univ. Kansas Publ., Mus. Nat. Hist, 5:563- 582, 6 figs, in text, December 1. Transmitted April 16, 1955. D 25-8618 t-f (3 wre y,e. ^-Jf University of Kansas Publications Museum of Natural History Vol. 9, No. 3, pp. 81-84 - December 10, 1955 - A New Long-eared Myotis (Myotis evotis) from Northeastern Mexico BY ROLLIN H. BAKER AND HOWARD J. STAINS i ..-/ 'EB 91956 I ^ University of Kansas Law^rence 1955 University of JCansas Publications, Museum of Natural History Editors: E. Raymond Hall, Chairman, A. Byron Leonard, Robert W. Wilson Volume 9, No. 3, pp. 81-84 Published December 10, 1955 Univebsity of Kansas Lawrence, Kansas PRINTED BY FERD VOILAND. JR.. STATE PRINTBR TOPEKA. KANSAS 1955 25-8617 L FEB 9 m^ A New Long-eared Myotis (Myotis Evotis) From Northeastern Mexico BY ROLLIN H. BAKER AND HOWARD ]. STAINS Long-eared bats obtained by field parties from the University of Kansas in the Mexican states of Coahuila, Nuevo Leon, and Tamauli- pas, are found to belong to the species, Myotis evotis, but are not referable to any named subspecies. They are named and described as follows: Myotis evotis auriculus new subspecies Type. — Female, adult, skin and skull; No. 55110, Univ. Kansas Mus. Nit. Hist.; 10 mi. W and 2 mi. S Piedra, 1200 ft., Sierra de Tamaulipas, Tamaulipas; 9 June 1953; obtained by Gerd H. Heinrich, original number 7061. Distribution. — Coastal foothills and adjacent mountains of northeastern Mexico from central Coahuila south and east to central Veracruz. Diagnosis. — Size medium (see measurements), ears small for the species; color dark, upper parts (/14) Ochraceous-Tawny (color terms are after Ridg- way, Color Standards and Color Nomenclature, Washington, D. C, 1912), underparts Wann Buff, ears pale; skull large, teeth large, mandible long. Comparison. — From Myotis evotis evotis ( H. Allen ) , from Colorado, Wyo- ming, and Montana, M. e. auriculus differs in: Ears averaging shorter; color darker and richer; ears paler and contrasting less, in color, with pelage; skull larger in all measurements taken except that of least interorbital constriction; forehead, when viewed laterally, rising more abruptly, because frontal region is more inflated; teeth larger. Remarks. — Myotis evotis auriculus, although no larger external y than M. e. evotis, has a larger skull, which in lateral view has a more abruptly rising forehead. The teeth, especially the first upper pre- molars, of auriculus are noticeably larger than those of evotis. The first two lower premolars are sub-equal in auriculus whereas in evotis the first lower premolar usually is larger. The mandible, in relation to the greatest length of the skull, is longer in auriculus (ratio, 71-74) than in evotis ( ratio, 67-71 ) . Coahuilan specimens, although assigned to auriculus, are slightly paler (upper parts (16) Ochraceous-Tawny; underparts (e) Light Buff) and have less abruptly rising foreheads than do the bats from Tamaulipas. In these features, the Coahuilan animals are somewhat intermediate between typical auriculus and evotis. The bat from (83) 84 University of Kansas Publs., Mus. Nat. Hist. Nuevo Leon, in both color and degree of slope of forehead, is inter- mediate between those from Coahuila and those from Tamaulipas. A bat from Perote, Veracruz, identified as Mijotis evotis chryso- nottis (J. A. Allen) [= M. e. evotis] by Miller and Allen (U. S. Nat. Mus., Bull. 144:118 and 120-121, May 25, 1928) is here assigned to M. e. auriciiliis. Measurements given by these authors indicate that this bat has a large skull, which is characteristic of this subspecies. Another specimen, similarly assigned by these authors and from the San Luis Mountains in northwestern Chihuahua, seems to be M. e. evotis, although the published measurements (loc. cit.) show that this bat tends toward auriculus in size of skull and mandible. All specimens were taken in mist nets stretched over water. Those from Coahuila were snared over a concrete water tank situated near the base of low hills in mixed mesquite and chaparral. In Nuevo Leon, one bat was netted over a small pond around which grew some low trees in an intermontane valley in the Sierra Madre Oriental. In Tamaulipas two bats were caught in a mist net stretched across a narrow, brush-bordered arroyo in the Sierra de Tamaulipas. One adult male weighed 7.0 grams; average and extreme weights of 7 adult, non-pregnant females were 6.8 ( 5.2-8.0 ) . Females taken on March 25 and 26 were not pregnant; one obtained on June .20 was lactating. Funds for financing field work were made available by the Kansas University Endowment Association and the National Science Foundation. Measurements. — Measurements, in millimeters, of the holotype and another female (No. 55111 KU) from the type locality and 3 females (Nos. 44726, 44729-30 KU) from Coahuila, respectively, are: total length, 94, 93, 97, 86, 96; length of tail vertebrae, 43, 42, 41, 39, 45; length of hind foot, 9.5, 9, 10, 10, 8; height of ear from notch, 20, 20, 20, 18, 20; length of forearm, 37.9, 38.4, 40.2, 37.3, 38.5; greatest length of skull, 16.1, 16.4, 16.2, 15.8, 16.1; condylobasal length, 15.4, 15.4, 15.6, 15.0, 15.4; zygomatic breadth, 9.7, 9.9, 10.1, 9.4, 9.9; least interorbital constriction, 3.9, 3.8, 3.9, 3.7, 3.7; breadth of braincase, 7.5, 7.6, 7.5, 7.5, 7.6; occipital depth, 5.9, 5.9, 5.5, 5.7, 5.6; palatal length, 8.5, 8.7, 8.7, 8.7, 8.9; mastoid breadth, 8.2, 8.4, 8.3, 8.2, 8.3; breadth across third upper molars, 6.1, 6.2, 6.3, 6.1, 6.1; length of maxillary tooth- row, 6.5, 6.5, 6.7, 6.6, 6.6; length of mandible, 11.5, 11.8, 11.9, 11.2, 11.7; length of mandibular tooth-row, 8.0, 8.0, 8.1, 8.1, 8.1. Specimens examined. — Total, 8, all in the University of Kansas Museum of Natural History. Coahuila: 4 mi. W Hacienda La Mariposa, 2300 ft., 5 (2 alcoholics). Nuevo Leon: Jturbide, 5000 ft., Sierra Madre Oriental, 1. Tamaulipas: 10 mi. W and 2 mi. S Piedra, 1200 ft.. Sierra de Tamaulipas, 2, Transmitted April 16, 1955. 2.'5-8617 rt" Lc^ vvvdic e._j University of Kansas Publications Museum of Natural History Volume 9, No. 4, pp. 85-104, 2 figs, in text May 10, 1956 Subspeciation in the Meadow Mouse, Microtus pennsylvanicus, in Wyoming, Colorado, and Adjacent Areas BY SYDNEY ANDERSON yys. coMP. Z0OL University of Kansas Lav^rence 1956 Universitv of Kansas Publications, Museum of Natural History Editors: E. Raymond Hall, Chairman, A. Byron Leonard, Robert W. Wilson Volume 9, No. 4, pp. 85-104, 2 figures in text Published May 10, 1956 University of Kansas Lawrence, Kansas PRINTED BY HERD VOILAND. JR.. STATE PRINTER TOPEKA. KANSAS 1956 26-422 ii.H^/li Subspeciation in the Meadow Mouse, I — "EWUIIY^ Micro tus pennsylvanicus, in Wyoming, Colorado, and Adjacent Areas BY SYDNEY ANDERSON INTRODUCTION In the region including Wyoming and Colorado, Microtus penn- sylvanicus has been divided into two subspecies: the pale M. p. insperattis (J. A. Allen) inhabits the Black Hills of the northeastern- most part of Wyoming; the dark Af. p. modestus (Baird) inhabits extensive areas in both Wyoming and Colorado. Initial examina- tion of Microtus pennsylvanicus revealed that specimens from the Big Horn Mountains of north-central Wyoming (within the range of modestus as mapped by Hall and Cockrum 1952:407), in color at least, resemble the subspecies insperatus more than they do modestus, and that specimens from southwestern Wyoming are notably dark. Durrant (1952:363) noted that specimens from Utah are dark, and Davis (1939:315) did the same for specimens from near Pocatello, Idaho. It seemed, therefore, that dark color might characterize populations of a wide geographic region and distinguish them from modestus named from southern Colorado. Also, there seemed to be a hiatus of at least 180 miles between the ranges of modestus in northern Colorado and modestus in eastern Wyoming, and an even greater distance separating populations of modestus in northern Colorado from those in western Wyoming. Microtus pennsylvanicus has not been taken in central or south- eastern Wyoming despite extensive collecting there, which yielded numerous records of other kinds of Microtus (M. longicaudus, M. montanus, and M. ochrogaster) . Subsequent study revealed a pat- tern of geographic variation within the range now ascribed to modestus which, in my opinion, can be described best by the recognition of three new subspecies. MATERIALS, METHODS, AND ACKNOWLEDGMENTS To study geographic variation in color a method was devised as follows: A single skin (KU 42407, from VA miles east of Buckhorn in Weston County, Wyoming) was selected as a representative of the paler mice and arbitrarily given the number 2. A single skin (87) 88 University of Kansas Publs., Mus. Nat. Hist. ( KU 17491, from 3 miles east of Moran in Teton County, Wyoming ) was selected as a representative of the darker mice from the western part of Wyoming and arbitrarily given the number 4. These mice were selected so that they were respectively paler and darker than the estimated average of the total variation within the populations to be studied, but the two mice were not at the extremes of paleness and darkness. Comparisons were based on visual inspection of the dorsal pelage as a whole. Skins were compared with these two mice and given whole numbers from one to five. If paler than the standard for 2, the skin was numbered one; if not distinguishably paler or darker, it was given the number two; if intermediate in color to the standards for 2 and 4 and not definitely more nearly referable to one than the other, it was given the number three; if it resembled the standard for 4, it was numbered four; and if darker, it was given the number five. In this manner skins from a given locality could be evaluated one by one and the results plotted, averaged, and treated statistically. On Figure 1 the aver- age values for color of 32 series are mapped to show the geographic variation of color. The following series of adults are the basis for Figure 1 (abbreviations for collections other than at the University of Kansas are included in parentheses): Each locality is followed by the month (or months) of capture, the number of specimens, and the average value for color. Montana: Glacier County, August, 6, 1.8; Hill and Chouteau counties com- bined (Mich), July, 24, 1.5; Malta, Philips County, August, 14, 1.5; Sheridan County, August, 6, 1.5; Fergus County (USES), August, 5, 2.4; RavalH County (KU and USES), August, 12, 2.8; Silver Eow County, August, 7, 3.0; Sweet Grass County (Mich), June and July, 7, 2.7; Park County, August, 10, 2.6. Idaho: Pocatello and vicinity, November and December, 5, 3.4. Wyoming: Park County, August, 6, 2.8; Sheridan County, September, 9, 1.2; Johnson County, August, 12, 1.5; Campbell and Crook counties, July, 11, 1.4; Weston County, July, 7, 1.6; Teton County, September, 8, 3.4; Teton County (Mich), June, 17, 3.1; Afton and vicinity, Lincoln County, July, 10, 4.2; Sage, Lincoln County, July, 5, 5.0. South Dakota: Pennington County (Chi), June, 14, 2.1; Pennington County (Mich), December and January, 8, 1.1; Walworth County, July, 4, 3.7; Buffalo County, July, 6, 3.2. Colorado: Loveland and vicinity, Larimer County (KU and USES), July, 13, 2.8; Eoulder County (Chi), Sep- tember, 34, 2.6; Park County (Denv), March, 8, 1.9; Colorado Springs (ERW), March, April, and May, 5, 2.8; Saguache County (USES), August, 46, 3.0; Conejos County, June, 4, 3.0; Wray, Yuma County (USES), 3, 4.7. Nebraska: Dundy County, August and November, 14, 4.6. New Mexico: Colfa.x County, June, 8, 3.2. Variation in color is discussed in the accounts of the subspecies concerned. SUBSPECIATION IN THE MeADOW MoUSE 89 114 108 102 114 108 102 FiGUFiE 1. Geographic variation in color in Microtus pennsylvanicus in the Rocky Mountains. Paler colors are represented by smaller numbers. Niun- bers are derived from the series of specimens listed in the text by the method described there. The subspecies that occur in the region studied are as follows: a. M. p. pullatus e. b. M. p. insperatus f. c. M. p. uligocola g. d. M. p. finitus h. M. p. modestus M. p. aztecus M. p. drummondi M. p. pennsylvanicus For each of the series Hsted in Table 1 all adult mice having skulls that measured more than 24.0 mm. in condylobasilar length were studied. Total length, length of tail, and length of hind foot were taken from the collector's field labels. The measurements of the skulls listed below were taken by means of dial calipers reading to one-tenth of a millimeter, and in the same fashion as described previously (Anderson, 1954:492). 90 University of Kansas Publs., Mus. Nat. Hist. Measurements of specimens in each series were averaged (the arithmetic means were computed ) . If the averages differed notice- ably the significance of the difference was tested statistically. Av- erages referred to in the text as significantly different differ by as much as, or more than, the sum of two times the standard error of each of the two averages. Linear measurements are in millimeters; color values are in the arbitrary units described in a preceding paragraph. Measurements taken of the skulls are: condylobasilar length, zygomatic breadth, interorbital breadth, lambdoidal breadth, prelambdoidal breadth, depth of braincase, and alveolar length of upper molar tooth-row. Secondary sexual variation was not detected in the material studied. Variation with age is important to the taxonomist even among specimens designated as "adults", because growth and changes in various proportions continue throughout the life of the mice. The possibility that differences detected in the statistical treatment or observed directly could be the result of differences in average age within the samples of "adults" was considered in each case. In order to study certain variations, the following "method of pairs" was used. Skulls of two series to be compared were matched in pairs so that they corresponded in size and ontogenetic stage of development. Then the two skulls of each pair were examined for differences in each of the following features: size of circle in- scribed by the upper incisor teeth, width of nasal bones relative to their length, curvatiue of the zygomatic arch, elongation of the braincase relative to its width when viewed from the dorsal aspect, degree of indentation in the anterior edge of the zygomatic arch near the rostrum, degree of depression of the nasal bones when viewed from the side, width in the vertical plane of the zygomatic arch at the suture between the maxillary and jugal bones, length relative to width of the prominent fenestra in the posterodorsal part of the squamosal bone, size of the meatus of the auditory canal, distance between the internal margin of an occipital condyle at its posteriormost point and the tip of the paraoccipital process of the same side of the skull, size of the foramen magnum, vertical height of the supraoccipital bone from the dorsalmost point on the margin of the foramen magnum to the midpoint of the lambdoidal crest, constriction posteriorly or narrowness of the incisive foramen relative to its length, distance across the premaxillary bone from the anteriormost point of the incisive foramen to the posteriormost point of the margin of the alveolus of the upper incisor, area of the maxillary septum (Howell 1926:112, or "zygomatic plate" of Eller- man 1941:1), acuminateness of the anterior border of the palatine SXJBSPECIATION IN THE MeADOW MoUSE 91 opening (internal nares), size of auditory bullae, size of foramen ovale, acuteness of the angle between the basioccipital and basi- sphenoidal bones at the suture between them (degree to which the area of the suture is raised between the bullae when viewed from the ventral aspect), width of first upper molar tooth, least distance between alveoli of first upper molars. Any difi^erential feature present in more than 75 per cent of the pairs of animals is reported in the discussion of the subspecies concerned. The sig- nificance of each difference reported was calculated by the Chi- square test and the confidence limit is given in each case. The probability used in the Chi-square formula is one-half of the per- centage of all pairs compared in which the skulls were different in regard to the character being considered. For example, in 68 per cent of the total number of pairs of skulls compared in this study a difference in the size of the auditory bullae was noted. Therefore the probability that a specified skull of a pair will have larger bullae than the other skull was taken as 34 per cent. A different probability for each feature compared was derived in like manner. This study is concerned primarily with mice from Wyoming and Colorado; I realize, however, that the physiographic and ecological conditions important to the distribution and subspeciation of Micro- tus pennsylvanicus do not correspond to political boundaries. Geo- graphic variation within these two states can be seen in proper perspective only when related to the neighboring areas and to previous studies. I have attempted to do this in the accounts of the subspecies. Approximately five months in the field in Wyoming and Colorado in the summers of 1950, 1951, 1952, and 1953 gave me a familiarity with the region that has helped to clarify the pattern of distribution. My study was based, in addition, on 762 specimens that are listed under "specimens examined" in the accounts of subspecies, and on comparative material from other states. Most of these specimens are sldns with skulls but some are skins only and others are skulls only. Some localities are represented by too few aduft individuals to permit significant comparisons. Owing to damaged skulls, certain measurements of some specimens were omitted from the calculations. If it seemed that the damaged skull was exceptionally large or small or a deviant in any other regard it was not used, in order not to bias the computed averages, which might be used in comparing proportions of the skulls. In the lists of specimens examined, localities that are omitted from Figure 2 because overlapping or undue crowding of the symbols would have resulted are italicized. I am grateful to Professor E. Raymond Hall for critical reading of the manuscript and helpful suggestions, to Dr. Rollin H. Baker and various of my fellow students at the Museum of Natural History for stimulating comments 92 University of Kansas Publs., Mus. Nat. Hist. pertinent to the problems involved in this study, to my wife, Justine Anderson, for assistance in the preparation of the manuscript, to numerous members of field parties from the Museum of Natural History, who collected much of the material studied, and to the curators and other persons, at the museums hsted below, who courteously made specimens available for study. The field work of the Museum of Natural History was assisted by the Kansas University 0 50 ■■■ I I 1 scale of miles 109 103 Figure 2. Distribution of the subspecies of Microtus pennsylvanicus in Wyoming and Colorado. Sohd dots represent localities from which speci- mens have been examined, and triangles represent localities reported in the literature from which I have not examined specimens. The question mark in southern Colorado denotes a questionable record discussed in tlie text. A. M. p. pullatus D. M. p. finitus B. M. p. in,fperatus E. M. p. modestus C. M. p. uligocula F. M. p. aztecus SUBSPECIATION IN THE MeADOW MoUSE 93 Endowment Association. A National Science Foundation Fellowship made it possible for me to visit the museums listed below. An honorarium awarded by the American Society of Mammalogists enabled me to present this paper at the 34th Annual Meeting of the Society, in June of 1954. Unless otherwise indicated, specimens are in the University of Kansas Museum of Natural History. Specimens in other museums are designated beyond as follows: American Museum of Natural History (AMNH); Museum of Zoology, Uni- versity of Michigan (Mich); Chicago Natural History Museum (Chi); United States National Museum (USNM); Biological Surveys Collection (USES); Museum of Vertebrate Zoology, Berkeley, California (MVZ); Colorado Mu- seum of Natural History, Denver (Denv); E. R. Warren collection, Colorado College, Colorado Springs (ERW); University of Colorado Museum, Boulder (UC). ACCOUNTS OF SUBSPECIES Microtus pennsylvanicus modestus (Baird) Arvicola modesta Baird, Repts. Expl. and Surv. . . ., pt. 1, Mammals, p. 535, July 14, 1858. Microttis pennsylvanicus modestus, Bailey, N. Amer. Fauna, 17:20, June 6, 1900. Type. — Immature specimen (sex not specified), skin and skull, number of skin 594, number of skull 1717, deposited in the collections of the Smithsonian Institution, obtained by F. Kreutzfeldt, Sawatch Pass, Rocky Moimtains [=Sa- guache Pass or Cochetopa Pass, Saguache County, Colorado], exact date unknown. I have not examined the type specimen. Range. — Northern New Mexico, and southern Colorado (see list of speci- mens and Fig. 2). Comparisons. — For comparison with the subspecies newly described below from northern Colorado see the account of that subspecies. The subspecies M. p. aztecus has been compared with M. p. modestus by Hall and Cockrvun (1952:308) who reduced aztecus to subspecific rank. Although aztecus is separated by approximately 100 miles from modestus, and although no proof of intergradation is available, my studies of variation in this species lead me to agree with Hall and Cockrum that "the morphological diflFerences between the two kinds of animals are of the degree and kind that separate subspecies, rather than species." A more adequate series of adults of aztecus is needed to clarify even the subspecific differences between aztecus and modestus. Measurements. — Averages, extremes, and standard deviations of a number of series are included in Table 1 in order to facilitate comparisons between different subspecies. Remarks. — The dividing line between M. p. modestus and the subspecies to the north on Figure 2 is drawn somewhat arbitrarily because few specimens are available from this area. Actual inter- gradation, in the form of a geographically intermediate population also morphologically intermediate between these two subspecies, is lacking. However, in most populations of both subspecies some individuals are intermediate between the two subspecies or even 94 University of Kansas Publs., Mus. Nat. Hist, more like the other subspecies than the one to which they are referred. Warren (1942:226) states that modestus has been re- corded from Lake County, although no reference to a specimen is given. Bailey (1900:21) cites Twin Lakes, in Lake County. That county is near the dividing line as I have drawn it, and therefore specimens from Lake County would be of special interest. An isolated colony of modestus occurs at San Rafael in Valencia County, New Mexico (Bailey, 1932:201). A hiatus of approxi- mately 150 miles separates that colony from the southernmost locality shown in Figure 2. A single specimen, the skin of an immature Microtus without skull, from Trinchera, Colorado, taken by L. R. Hersey in 1912, is in the Colorado Museum of Natural History. No species of Microtine has been recorded from within 50 miles of this locality. The specimen is seemingly more like M. pennsylvanicus than any other species of Microtus. This locality is represented in Figure 2 by a question mark. Ecologically M. montanus fusus Hall and M. longicaudus mordax (Merriam) in the same region occupied by modestus seem to be more montane than modestus. It favors lush grass on the wet floors of alluvial valleys, and also irrigated areas such as that at Manassa. Specimens examined. — Total 130. Colorado: Chaffee Co.: Salida, 3 (ERW). Saguache Co.: Monshower Meadows, 27 mi. NW Saguache [=3 mi. E Cochetopa Pass], 8 (USES); Tevebaugh's Ranch, 20 mi. W Saguache, 46 (USES, there are additional specimens not examined in detail by me); Cochetopa Pass, 33 mi. W Saguache, 1; 3 mi. N, 16 mi. W Saguache, 8500 ft., 5; 5 mi. NW Hooper, 1 (AMNH); Medano Ranch, 15 mi. NE Mosca, 2 (1 ERW, 1 USES). Custer Co.: Westchffe, 7800 ft., 1 (ERW). Alamosa Co.: Hooper, 10 (2 AMNH, 8 Denv); Mosca, 3 (ERW). Conejos Co.: IJi mi. E Manassa, 11. Costilla Co.: Alamosa, 3 (Mich); 2 mi. S Blanca, 7800 ft., 6 (MVZ). New Mexico: Taos Co.: Arroyo Hondo. 7600 ft., 6 (USES); Taos, Pueblo, 1 (USNM). Colfax Co.: 1 mi. S, 2 mi. E Eagle Nest, 8100 ft., 21; Taos Mountains, east slope, 8800 ft., 1 (USES); Coyote Creek, 1 (USES). Microtus pennsylvanicus uligocola new subspecies Type. — Adult male, skin and skull, number 26898, University of Kansas, Museum of Natural History, obtained by James O. Lounquist, original number 349, 6 miles west and Ji mile south of Loveland, 5200 ft., Larimer Co., Colo- rado, on July 26, 1948. Range. — Northern Colorado. See Figure 2 and list of specimens examined. Diagnosis. — Entire animal and skull large; color average for the species, neither extremely pale or dark in summer pelage; molar tooth-row long; nasals narrow; maxillary septum large; first upper molar wide; anterior margin of zygomatic arch above infraorbital foramen not deeply indented; fenestrae in posterodorsal parts of squamosal bones relatively long; braincase not elongate; auditory bullae and meatus large. Comparisons. — From M. p. modestus, M. p. uligocola differs as follows: averages paler; prelambdoidal breadth and alveolar length of molar tooth-row SUBSPECIATION IN THE MeADOW MoUSE 95 significantly greater. Six pairs of skulls were compared. Of the features listed above under the "method of pairs" only two features differed in more than 75 per cent of the pairs; in 5 of 6 pairs uligocola had a less distinctly indented anterior margin of the zygomatic arch (Confidence Limit .95) and a more elongate posterodorsal squamosal fenestra (C. L. .85). Seven pairs of skulls from Boulder, Colorado, representing uligocola and from Colfax Coimty, New Mexico, representing modestus differed in more than 75 per cent ol the pairs in three features. Only one of these differences, the elongation of the posterodorsal squamosal fenestra, was the same as a difference noted above between topotypes of uligocola and modestus. A comparison of ten pairs of skulls of uligocola from Boulder, Colorado, and topotypes of uligocola revealed no significant differences. These observations are indicative of 1 ) the differences between samples and populations which may be assigned to a single subspecies, and 2) the fact that in general these local differences are less than the differences between subspecies. From insperatus, the subspecies to the north, uligocola differs as follows: darker in both summer and vsanter pelage; averaging larger in most measurements of the skull; significantly longer molar tooth-row; hind foot averaging longer. For comparisons with the sub- species to the east and the northwest see the accounts of those below. Remarks. — M. p. uligocola is more closely restricted to wet situa- tions than M. ochrogaster haydeni (Baird) whose general range lies to the eastward. The numerous lakes, the continuous supply of water from the mountains, and the irrigation systems at lower altitudes along the eastern base of the mountains provide the con- ditions to which uligocola is suited. It is named for its predilection for water. The variability in color is relatively greater in topotypes of both uligocola and of modestus than in many of the other series studied. Specimens from Denver and Colorado Springs taken in late autumn and winter (October to February) are paler, more reddish and less blackish, than specimens taken in June and July at Loveland. This reddishness results from longer, and more intensely reddish tips of the hair. The entire hairs also are longer. The average weight of 16 adults (12 males and 4 non-pregnant females) from near Loveland is 49.7 gms. The average length of the ear is 13.4. Specimens examined. — Total 228. Colorado: Larimer Co.: 6 mi. W, % mi. S Loveland, 5200 ft., 18; 3 mi. N Loveland, 3; Loveland, 4 (USES). Morgan Co.: 4 mi. W Orchard, 4 (Mich); Orchard, 1 (Mich). Boxjlder Co.: Boulder, 91 (USNM 19, UC 12, Chi 60 examined, additional specimens in the collection ) ; Valmont, 4 ( UC ) . Clear Creek Co. : Clear Creek, N side of Idaho Springs, 1. Jefferson Co.: Olivet, 10 (Denv). Adams Co.: Crook's Lake, 8 (Denv); Barr, 16 (Denv 15, ERW 1). Arapahoe Co.: Denver, 21 (USES 3, AMNH 8, Denv 10). Park Co.: Wilham's Ranch, near Tarryall, 11 (Denv). Teller Co.: Divide, 2 (ERW). El Paso Co.: Colorado Springs, 31 (ERW 22, AMNH 7, MVZ 2); 12 mi. S Colorado Springs, 3 (MVZ). 96 University of Kansas Publs., Mus. Nat. Hist. Microtus pennsylvanicus finitus new subspecies Type. — Adult female, skin and skull, number 50204, University of Kansas, Museum of Natural History, obtained by J. K. Jones, Jr., original number 906, 5 miles north and 2 miles west of Parks in Dundy County, Nebraska, on August 16, 1952. Range. — The valley of the north fork of the Republican River in eastern Colorado and southwestern Nebraska. Diagnosis. — Entire animal and skull large; color dark for the species; zygo- matic breadth large; upper molars large and upper molar tooth-row relatively long; braincase elongate; auditory meatus relatively small; bullae large; in- cisors relatively procumbent. Comparisons. — From M. p. uligocola, the subspecies to the west, M. p. finitus diflFers in darker color. These subspecies resemble each other in large size and large molar teeth. A comparison of nine pairs of skulls by the "method of pairs" shows three features in which finitus (from at, or near, the type locahty) differs from uligocola (from at, or near, the type locality); in 7 of 9 pairs finitus had a relatively more elongate braincase (Confidence Limit .60); in 9 of 9 pairs uligocola had larger auditory meatuses (C. L. .99); in 7 of 9 pairs uligocola had relatively larger bullae ( C. L. .80 ) . From M. p. penn- sylvanicus (Ord) from eastern Nebraska and eastern South Dakota finitus differs in larger size; darker color; larger molar teeth and longer upper molar tooth-row. Five pairs of skulls were compared; in all 5 pairs, finitus had more procumbent incisor teeth ( C. L. .97 ) and wider first upper molars ( C. L. .97 ) ; and in 4 of 5 pairs finitus had a relatively more elongate braincase (C. L. .60). From M. p. insperatus, the subspecies to the north, finitus differs as follows: color darker, size larger, molar teeth relatively larger and alveolar length of the upper molar tooth-row greater. Five pairs of skulls were compared and in all 5 pairs finitus had more procvunbent upper incisors (C. L. .97) than insperatus; in 4 of 5 pairs insperatus had a relatively more elongate braincase (C. L. .60), narrower first upper molariform tooth (C. L. .75), and shorter distance between the alveoli of the first upper molars ( C. L. .88 ) . Remarks. — The species, Microtus pennsylvanicus, in Pleistocene time ranged onto the plains of Kansas as far southward as Meade County, Kansas (Hibbard, 1940:421). This occurrence indicates a cooler more humid climate then than now in southwestern Kansas. M. p. finitus is more closely associated with water than Microtus ochrogaster, the only other species of Microtus now occupying the same region, although both species have been captured at certain places in the same runways. In Nebraska, a marginal part of the range of the species, M. pennsylvanicus has been taken at scattered localities. This scattered and localized distribution of suitable habitat undoubtedly limits gene-flow between these relict popula- tions. Presumably as a result of this isolation finitus has accumu- lated and maintained its distinctive characteristics. The subspecies is so named because of its limited range. The average weight of SUBSPECIATION IN THE MeADOW MoUSE 97 eight specimens ( 4 males and 4 non-pregnant females ) from Dundy County is 57.2 grams. Specimens examined. — Total 26. Colorado: Yuma Co.: Wray, 3 (USES); 1 mi. W Laird, 2. Nebraska: Dijndy Co.: 5 mi. N, 2 mi. W Parks (Rock Creek State Fish Hatchery), 19; Haigler, 2 (USES). Microtus pennsylvanicus pullatus new subspecies Type. — Adult male, skin and skull, number 37873, University of Kansas, Museum of Natural History, obtained by RolHn H. Baker, original number 1343, 12 miles north and 2 miles east of Sage, 6100 ft., in Lincohi County, Wyoming, on July 19, 1950. Range. — North-central Utah, eastern Idaho, western Wyoming, and south- western Montana. See Figures 1 and 2. Diagnosis. — Size average; color dark, especially in southern part of range; tail relatively long; molar teeth small; nasals relatively broad; maxillary septum relatively small. Comparisons. — From M. p. uligocola, the subspecies to the southeast, M. p. pullatus differs as follows: relatively darker in southern part of its range (see Figure 1 ) ; smaller, tail relatively longer. In 6 of 7 pairs of skulls compared of pullatus (from Lincoln Co., Wyoming) and uligocola (from Larimer Co., Colorado), pullatus had relatively broader nasals (Confidence Limit .85); uligocola had larger maxillary septa ( C. L. .97 ) and larger molar teeth ( C. L. .90). From insperatus, the subspecies to the east, pullatus differs as follows: both summer and winter pelage darker; tail longer both actually and relatively; upper molar tooth-row shorter. Ten pairs of skulls of specimens from near Afton, Wyoming, representing pullatus, and from northeastern Wyoming, representing insperatus, revealed no significant differences in the features observed by the "method of pairs". Although not compared in detail with the subspecies to the north, M. p. drummondi (Audubon and Bachman), examination of specimens from western Montana and the accounts of other authors indicate that topotypes of pullatus are darker, longer-tailed, slightly larger-skulled and perhaps longer over all. Remarks. — In this subspecies there is a cline in color from dark in extreme southwestern Wyoming to pale in north-central Wyo- ming and Montana as the range of M. p. insperatus is approached. There is thus a broad zone of intergradation in color and the line separating the subspecies must be drawn somewhat arbitrarily. In Wyoming the most distinct break in this chne is in the Big Horn Basin and if a detailed study of the species were made in Montana the break would probably be found where the mountains meet the plains, roughly as shown in Figure 1. There is a similar chne in western Montana in color. The mice are paler farther north as one approaches the Canadian border although they do not become so pale as insperatus. Darkness is a characteristic of several non- adjacent subspecies of Microtus pennsylvanicus, for example M. p. 98 University of Kansas Publs., Mus. Nat. Hist. kincaidi Dalquest in central Washington (Dalquest, 1948:347), M. p. finitus, and M. p. nigrans Rhoads in eastern Virginia, but these subspecies presumably differ in other characters. Some morphological features of the same kind and degree that differentiate subspecies in one place may not vary geographically in another place. Furthermore the geographic variation in one feature may be only partly correlated with the variation in another feature. The variation in M. p. puUatus is an example: Specimens from near Pocatello, Idaho, are darker than topotypes of modestus but specimens from Fremont County, Idaho, are indistinguishable from topotypes of modestus (Davis, 1939:315). I have examined a number of mice from the Bitterroot Valley in western Montana and the color value for 12 adults is 2.7. They are slightly but not significantly paler than topotypes of modestus. This is a result of the cline mentioned above and does not indicate relationship with modestus. Some average measurements of 10 skulls from this series are as follows : condylobasilar length, 24.9; zygomatic breadth, 14.4; interorbital breadth, 3.4; lambdoidal breadth, 11.5; prelamb- doidal breadth, 9.1; alveolar length of upper molar teeth, 6.5; and depth of braincase, 7.6. Average external measurements of 9 specimens are as follows: total length, 157; length of tail, 36; length of hind feet, 19.4. Mice from the Bitterroot Valley were compared with topotypes of modestus by the "method of pairs," and modestus had a larger foramen magnum in 6 of 6 pairs (Confidence Limit .97) and larger first upper molar teeth in 5 of 6 pairs (C. L. .75). A comparison of topotypes of pullatus with modestus shows a similar difference in the teeth, modestus being larger, but in the size of the foramen magnum there is no difference. A comparison of the measurements of pullatus ( Bitterroot Valley ) , pullatus ( near topotypes), and modestus (topotypes) shows that the two series of topotypes differ significantly in condylobasilar length ( of border- hne significance), zygomatic breadth and lambdoidal breadth (both of which vary greatly with age), and length of molar series; the specimens from the Bitterroot Valley agree with pullatus rather than modestus in all of these characters. The specimens from the Bitterroot Valley are smaller than pullatus (topotypes) in total length; they more closely resemble mx)destus in length of tail, and the hind foot is shorter than in either modestus or pullatus (which do not differ significantly). Specimens from western Montana re- semble modestus in certain respects but in most respects resemble topotypes of pullatus and are referred to pullatus. Specimens from SUBSPECIATION IN THE MeADOW MoUSE 99 Coeur d'Alene, Idaho, and from Blackfoot, Montana (marginal records of modestus. Hall and Cockrum, 1953:410) may be referred to drummondi. Marginal records of pullatus in Montana to my knowledge are: Florence, Ravalli Co.; Highwood Mtns., Chouteau Co.; 7 mi. NE Hilger, Fergus Co.; 10 mi. NW Park City, Stillwater Co. ( all represented by specimens in the USES ) . The line separat- ing pullatus and drummondi is tentatively drawn as shown in Figure 1. Specimens examined. — Total 256. Wyoming: Yellowstone National Park: Mammoth Hot Springs, 17 (USNM). Park Co.: 3i mi. E, i, mi. S Cody, 5020 ft., 15. Teton Co.: Whetstone Creek, 4 (Mich); 5 mi. N Moran, 13 (Mich); Moran and environs (4 localities within 4 miles of Moran), 6200 ft., 54; Teton Park, Trappers Lake, 3 ( Mich ) ; Teton Park, Jenny Lake, 1 ( Mich ) ; Teton Park, String Lake, 1 (Mich); Sheep Creek, 1 (Mich); Jackson and environs, 115 (Mich 113, USES 1). Sublette Co.: 34 mi. N, 4 mi. W Pine- dale, 7950 ft., 2; Kendall, 5 (Mich). Lincoln Co.: 95-i mi. N, 2 mi. W Afton, 6100 ft., 1; 9 mi. N, 2 mi. W Afton, 3; 7 mi. N, 1 mi. W Afton, II; 15 mi. N, 3 mi. E Sage, 6100 ft., 1; 12 mi. N, 2 mi. E Sage, 4; 6 mi. N, 2 mi. E Sage, 2. Microtus pennsylvanicus insperatus (Allen) Arvicola insperatus Allen, Bull. Amer. Mus. Nat. Hist., 6:347, December 7, 1894. Microtus pennsylvanicus insperatus, Anderson, Canadian Field-Nat., 57:92, October 17, 1943. Microtus pennsylvanicus wahema Bailey, Jour. Mamm., 1:72, March 2, 1920. Type. — Adult male, skin and skull, number |iff, American Museum of Natural History, obtained by W. W. Granger, at Custer, Black Hills, South Dakota, August 9, 1894. Range. — Western South Dakota, southwestern North Dakota, eastern Mon- tana, southern Alberta and Saskatchewan, eastern Montana, and northeastern Wyoming. Comparisons. — Microtus pennsylvanicus insperatus is paler than any adjacent subspecies. It has been compared with pullatus in the preceding account. Bailey's studies of M. p. wahema [= insperatus] and his comparisons with M. p. pennsylvanicus and M. p. drummondi to the east and north, in North Dakota (1920, 1927), are the basis for the northern and eastern boundaries of the range of insperatus in Figure 1. Comparison of insperatus with finitus to the south is made in the account of the latter. Remarks. — Bailey (1900:20) had only 7 specimens from north- eastern Wyoming and western South Dakota, of M. pennsylvanicus and thought that Arvicola insperatus Allen (1894:347) was not subspecifically distinct from modestus. Subsequently Bailey (1920: 72) had adequate numbers of specimens and described Microtus pennsylvanicus wahema from eastern Montana and western North Dakota. Anderson (1943:92) concluded that wahema was not dis- tinct from insperatus and therefore the name M. p. insperatus ( Allen ) is applicable to this subspecies. On the basis of specimens 100 University of Kansas Publs., Mus. Nat. Hist. Table 1. Average MEAstrREMENTs (in Millimeters) of Adult Microtus pennsylvanicus Locality (or area) O ^ c" rt bC C 0) o H C -a t^ _> iS "T IS c3 CO o -O - o-a •— ' -*j •5*^ -b ^ bC 4> "TJ C c i. F» f • Molar paraconids internal, but usually nearly if not entirely indistinguishable. Entoconids indistinguish- able from hypoconulids, reduced to a low, bordering rim. Reduc- tion of M3/m3 extreme. Molar trigonid cusps (ml-m2) closely appressed, turgid. Lower jaws short and stout. Premolars short and wide, crowded (obliquely set in rami), simple with basins of heels shallow and poorly defined but wide. Species: E. inaequidens (type), E. grangeri, £.? sternbergi. Ellipsodon? sternbergi of the Dragon fauna seems to be older geologically than either E. inaequidens or E. grangeri. The two last-mentioned are nearly of the same age, but probably are not con- temporaneous. £.? sternbergi agrees in size with E. grangeri, but is less advanced in reduction of paraconid, stage of adpression of trigonid cusps, and reduction of inner rim of heel in the molars. E.? sternbergi probably is a somewhat primitive species of Ellip- sodon although as Gazin indicates a small species of Mioclaenus possibly is represented (he also considers a possible relationship to ]epsenia). A large variant of Ellipsodon grangeri or an unnamed species of Ellipsodon or Mioclaenus is represented by KU No. 9521. This specimen was obtained with a relatively large collection of other mammals in Kimbetoh Arroyo, and is representative of the Delta- therium fauna of Osborn. It consists of a left ramus with m2 and incomplete ml. Certain features, such as the presence of a para- conid, suggest that this specimen represents a small species of Mioclaenus rather than a species of Ellipsodon. USNM No. 15781, a lower jaw from the Dragon, was named Ellipsodon? species (a) CONDYLARTII GeNUS ElLIPSODON 113 by Gazin, and he states that it resembles somewhat the Torrejon species referred to E. inaequidens. It probably is related to it. Finally, USNM No. 16333, an upper molar, has been identified by Gazin as possibly pertaining to E.? sternbergi. It is slightly smaller than our only upper molar, the metaconule is more prominent, and the external cingulum more developed although the posteroexternal cingulum is absent in both. From E. inaequidens, Ellipsodon grangeri differs chiefly in larger size. It is possibly slightly more advanced otherwise. Interpreta- tion of an ancestor-descendant relationship turns on unknown strati- graphic relationships. If the Gallegos beds are older, then E. grangeri has progressed slightly in regard to size, but if a reverse stratigraphic relation holds, E. inaequidens can be viewed as a slightly dwarfed descendant. E. grangeri is hardly a geographic variant of E. inaequidens — the known localities are too close to- gether— unless geographic distribution has become changed with time. The differences which separate Ellipsodon inaequidens and E. grangeri are so slight (an average size difference of 17 per cent is not in itself definitive) that it might have been advisable to treat E. grangeri as only a temporal subspecies. The decision to regard them as fully distinct was made because no overlap in size is re- corded in the thirteen known specimens of the two species. Classification of Mioclaeninae. — Simpson (1945:123) has classi- fied as Mioclaeninae the genera Tiznatzinia, Choeroclaenus, Mio- claenus, Ellipsodon, Litaletes, and Jepsenia. Of these, Mioclaenus, Ellipsodon, Litaletes, and Jepsenia are of Torrejonian age. It has never been too satisfactory an arrangement to include in Ellipsodon as many and varied species as has been done. With in- creased information concerning the morphology of Ellipsodon as a result of the finding and naming of E. grangeri, it becomes evi- dent that several of the species can no longer be assigned to the genus. These are t^. shephardi, E. aquilonius, E. acolytus, E. lemu- roides, and presumably also the poorly known E. aequidens. (E. priscus of the Puercan has been previously assigned to Tiznatzinia by Simpson, 1936:8.) E. shephardi, E. aquilonius and E. acolytus form a group of closely related species. E. lemur oides is somewhat more removed, but evidently allied. Review of the opinions ex- pressed by Matthew, Simpson, and Gazin make it seem unlikely that this group of four species can be referred to any genus named in the preceding paragraph or that the species can be distributed among the several listed genera. 114 University of Kansas Fuels., Mus. Nat. Hist. Simpson, who has presented us with the most thorough review of the Mioc\a.enina.e states (lQS7:2So)," Ellipsodon aquilonius . , . closely resembles E. acohjtus. The latter is fairly close to E. lemu- roides, which in turn approaches E. inaequidens. There is no logical or convenient separation, probably of more than specific rank, in this series of four species, yet E. aquilonius is markedly unlike E. inaequidens, which is the type of the genus." Farther on he says (op. cit. -.234), "In general aspect, E. aquilonius resembles some of its associates such as Litaletes disjunctus more than it does Ellipso- don inaequidens, but in structural detail it seems closer to Ellipsodon acolytus and is conservatively associated with that species generic- ally ( rather than definitely with the t>'pe of Ellipsodon ) ." Thus it seems that the removal of E. inaequidens from this group does not alter the relationship of the remaining species sujBBciently to warrant referring all three (or only E. aquilonius and E. acolytus) to Lita- letes although this remains a possibiHty. Gazin indicates that E. shephardi is closer to the cluster of these species than to Litaletes (1941:29). An important and additional reason for not referring E. aquilo- nius, acolytus, lemuroides, and shephardi to Litaletes is the presence of a Litaletes-like species in the Puercan (early Paleocene). This species (represented by KU Nos. 9446, 9447) is smaller than L. disjunctus, but in the upper dentition does not otherwise differ — at least not at a generic level. In the lower dentition, the premolars are narrower ( partly the result of crushing? ) , paraconids and meta- conids of p3-p4 are of lesser development; the molars have larger hypoconulids, and are more distinctly separate from entoconids. The presence of this species contemporaneous with a species ( Tiz- natzinia priscus), which is close to the ancestry of E. acolytus sug- gests a dichotomy sufficiently important to be recognized by generic separation of the Torrejonian species. Jepsenia mantiensis is said by Gazin (1941:32) to be closer to Litaletes disjunctus than to any of the above mentioned species. Further, Ellipsodon shephardi is regarded by him as generically distinct from species of Jepsenia, and hence by inference the latter is distinct from all those obviously closely related to E. shephardi. Thus his taxonomy suggests as does that of Simpson the inadvisabil- ity of referring the species previously assigned to Ellipsodon to either of the generic groups named Jepsenia by Gazin and Litaletes by Simpson. Nevertheless, it is possible that such may eventually prove to be the correct procedure. I propose, however, that the species E. lemuroides, E. acolytus, E. aquilonius, and E. shephardi CONDYLARTH GeNTJS ElLIPSODON 115 be grouped together as a generic assemblage distinct from these groups. For this assemblage, the name Promiocluenus Trouessart, 1904, seems to be available although never mentioned by classifiers of the Mioclaeninae. Trouessart included in his nominal genus only P. acolytus and P. lemiiroides. P. acolytus was listed first, but he did not select a typical species. To remedy this omission, I select Mioclaenus lemuroides as type of the genus Promioclaenus. The type species is somewhat aberrant with respect to the others. It is selected rather than the otherwise taxonomically preferable P. aco- lytus because P. lemuroides is less likely, at some future date, to have a type species of another and later-named genus (such as Litaletes or Jepsenia) associated with it — a reason which I think is of some practical worth in the present instance. The diagnosis of EUipsodon given by Simpson (1937:226-227) is essentially of Promioclaenus. As modified by the removal of E. irmequidens, it may be restated as follows: Promioclaenus Trouessart, 1904 Type. — Mioclaenus lemuroides Matthew 1897. Distribution. — Torrejonian of Nacimiento formation. New Mexico; Lebo formation, Montana; Joes Valley member of North Horn formation, Utah. Diagnosis. — Dentition ^, \, |, f . PI caniniform. pi cuspidate, more or less enlarged, but not really inflated. The p4 without distinct paraconid, meta- conid absent or rudimentary, protoconid inflated rather than bladelike; talonid basined and relatively large. Molar hypocones weak (or absent?), meta- conules present. Molar trigonids with paraconids internal, fusing with meta- conids; moderately separated metaconids and protoconids. Molar talonids with relatively wide basins, entoconids generally indistinct and fusing with hypoconulids (especially on m2). m| moderately reduced. M3 with reduced metacone. Rami relatively long and slender. 116 University of Kansas Publs., Mus. Nat. Hist. LITERATURE CITED Gazin, C. L. 1941. The mammalian faunas of the Paleocene of central Utah, with notes on the geology. Proc. U. S. Nat. Mus., 91:1-53, pis. 1-3, 29 figs, in text. Matthew, W. D. 1897. A revision of the Puerco fauna. Bull. Amer. Mus. Nat. Hist., 9:259- 323, 20 figs, in text. 1937. Paleocene faunas of tlie San Juan Basin, New Mexico. Trans. Amer. Philos. Soc, n. s. 30:i-viii, 1-510, pis. 1-64, 85 figs, in text. Simpson, G. G. 1936. Additions to the Puerco fauna, lower Paleocene. Amer. Mus. Novit. 849:1-11, 6 figs, in text. 1937. The Fort Union of the Crazy Mountain Field, Montana, and its mammalian faunas. Bull. U. S. Nat. Mus., 169:i-x, 1-287, pis. 1-10, 80 figs, in text. 1945. The principles of classification and a classification of mammals. Bull. Amer. Mus. Nat. Hist., 85:i-xvi, 1-350. Trouessart, E. L. 1904. Catalogus mammalium tam viventium quam fossihum. Supple- ment, pts. 1-2:1-546. Wilson, R. W. 1951. Preliminary survey of a Paleocene faunule from the Angels Peak area, New Mexico. Univ. Kansas Publ., Mus. Nat. Hist., 5:1-11, 1 fig. in text. Transmitted August 22, 1955. n 26-1037 n " Z-/:.^ tv-re-yictej University of Kansas Publications Museum of Natural History Volume 9, No. 6, pp. 117-123, 3 figures in text May 19, 1956 Additional Remains of the Multituberculate Genus Eucosmodon BY ROBERT W. WILSON mm m 6t956 University of Kansas Law^rence 1956 University of Kansas Publications, Museum of Natxtoal History Editors: E. Raymond Hall, Chairman, A. Byron Leonard, Robert W. Wilson Volume 9, No. 6, pp. 117-123, 3 figures in text Published May 19, 1956 University of Kansas Lawrence, Kansas PRINTED BY FERD VOILAND. JR.. STATE PRINTER TOPEKA, KANSAS 1956 26-1038 J(/« 6 1956 mm Additional Remains of the MultitubercuM^IiMDL. Genus Eucosmodon BY ROBERT W. WILSON Eucosmodon americanus is the type of the multituberculate genus Eucosmodon Matthew and Granger. This species is recorded only from the Puercan (early Paleocene) of the San Juan Basin of New Mexico ( Taeniolabis zone of Ojo Alamo and Barrel Spring arroyos, and in the Ectoconiis zone three miles southeast of Kimbetoh ) . A number of specimens, perhaps six in all, have been secured, but most of these are exceedingly fragmentary ( Granger and Simpson, 1929:648-649 ) . No specimens have been recorded from the Taenio- labis zone, the type level, since David Baldwin obtained the original material from there for E. D. Cope in the Eighties. It is therefore of importance to describe additional specimens of Eucosmodon americanus obtained by parties of the Museum of Natural History, University of Kansas, from the Taeniolabis zone at Barrel Spring Arroyo (from whence the Baldwin material probably came if it did not come from beds a few miles farther west in Ojo Alamo Arroyo ) . In addition, a probable P3 of Eucosmodon obtained at the head of Kimbetoh Arroyo and associated with a Torrejonian ( middle Paleo- cene) faunule is figured and described briefly. In the summer field season of 1950, we found in Barrel Spring Arroyo ( KU Loc. 3; W/2 sec. 11, T. 24 N, R. IIW) in the Taeniolabis zone four specimens referable, with greater or lesser certainty, to Eucosmodon americanus. These are: ( 1 ) KU No. 9401, left upper jaw with P2, P4, incomplete Ml, and M2; (2) KU No. 9402, left ml and associated fragment of lower incisor; (3) KU No. 9403, fragment of lower jaw with stump of incisor; and KU No. 9404, fragment of incisor. Except that No. 9402 confirms the presence of Eucosmodon americanus in the Barrel Spring Arroyo, principal interest attaches to the upper jaw, No. 9401, the first such identifiable specimen to be found in the Puercan of the San Juan Basin. No. 9401 in addition to the teeth listed has what may be a socket for an additional anterior tooth which is separated from P2 by a short diastema. If this is a tooth socket then it is for a reduced PI, a tooth supposed to be absent in Eucosmodon. It cannot well be for an incisor inasmuch as the socket almost certainly lies in the maxillary bone bounded anteriorly by a depression which seemingly represents the palatal fossa and anterior palatal foramen of ptilo- dontid multituberculates (Simpson, 1937:742, fig. 6). A vague outline anterior to the possible alveolus suggests a root of an incisor in the premaxilla. Nothing is definite enough to warrant, however, changing the accepted dental formula of Eucosmodon. (119) 120 University of Kansas Publs,, Mus. Nat. Hist. P2 is a single-rooted tooth with three cusps, each with grooved tip and sides much as in AMNH No. 16534, Eucosmodon sp. from the younger Torrejon fauna (Granger and Simpson, 1929:636, fig. 17). Our specimen has an an- terior cusp that is more internal and perhaps larger than the one in No. 16534. The two posterior cusps are more nearly transversely placed (less alternating) than in the latter. Some of this difference may be the result of rotation in the socket of P2 in one or the other of the two specimens. P3 is not preserved in KU No. 9401, but the single large alveolus present shows tliat Uke P2, the tooth is single-rooted. P4 is heavily worn and the resulting truncation of the crown prevents a de- termination of the exact number of cusps present in the species. Nevertheless, it is, seemingly, the only complete upper fourth premolar of Eucosmodon known. Three cusps are present in the external row. The internal row is partly obliterated by wear, but three anterior cusps are present, and the most posterior part has two, possibly three, cusps. There seems to have been nine or ten cusps in all. P4 of Eucosmodon americanus has the same pattern as that shown in the preserved anterior half of P4 in AMNH No. 16534 of the Tor- rejon fauna. Only the posterior half of Ml is preserved. In this fragment four or five cusps are present internally; four cusps are present in the median row; and three cusps are preserved externally. Figures 1-2. Eucosmodon americanus (Cope), Puercan (early Paleocene), Barrel Spring Arroyo, San Juan County, New Mexico. Fig. 1. No. 9401, occlusal view in outline of left upper jaw fragment with P2 and P4. la. No. 9401, occlusal view of left upper dentition (P2, P4, incom- plete Ml, and M2). Fig. 2. No. 9402, occlusal view of left ml. Drawings by Jane S. Mangel, all X 4. MULTITUBERCULATE GeNUS EuCOSMODON 121 The second upper molar is nearly perfectly preserved. It is the only terminal upper molar so far known in specimens of the genus Eucosmodon. Three cusps form the inner row; the posterior cusp is slightly damaged and could represent more than a single cusp, but I think only one. Two large cusps form the me- dian row and, except for extension into this row of a cingular ridge from the external row of cusps, there is no indication of any more. This formula of 3:2 for these two rows of cusps is a primitive one found in the Late Jurassic multi- tuberculates, but seemingly rarely in the Late Cretaceous and Early Tertiary species (Simpson, 1928, 1929). Essentially the same count is present, how- ever, in other Paleocene genera and species, as for example, Prionesstts lucifer (Granger and Simpson, 1929:663); and Ectypodus laytoni (Jepsen, 1940 :pl. Ill, fig. 2). In the outer row three cusps are present with possible traces of two others, one anterior, the other posterior. Additional specimens, representing other individuals, were obtained by us from the same locahty as No. 9401, only a matter of yards away. No. 9402 is an ml with an associated incisor. This ml has four cusps in its inner row and five(?) in the outer one. The pattern thus developed is close to that in ml of AMNH No. 3028 (cotype of E. americanus), but the two posteroexternal cusps are more nearly separate from each other, and the most posteroexternal one is relatively larger than in No. 3028 (Granger and Simpson, 1929:648, fig. 27). The fragments of the lower incisors represented in our collection add little to knowledge of the species. No. 9404 is too much distorted to yield measure- ments of value; those of No. 9403 are given in the table of measurements. These newly described specimens of Eucosmodon americanus furnish im- portant data as follows: ( 1 ) No. 9401 is the first upper jaw which may be assigned to the type species, £. americanus, with high probability. (2) No. 9401 demonstrates that the upper dentition (AMNH No. 16534) from the Torrejon previously assigned to Eucosmodon by Granger and Simpson does in fact pertain to that genus. (3) No. 9401 reveals the structure of M2 in this genus for the first time. ( 4 ) These specimens are the first certain records of the genus and species from Barrel Spring Arroyo, and hence the first to demonstrate conclusively that £. americanus occurs at the type locahty of the Puerco fauna. To a cer- tain extent they reinforce the highly probable, but unproved, contention that Baldwin's specimens came from here or from a continuation of these beds to the west in Ojo Alamo Arroyo (Sinclair and Granger, 1914:310; Granger and Simpson, 1929:648). (5) The material indicates that the character of size which in part allegedly separates £. a. americanus of the Taeniolabis zone from £. a. primus of the Ectoconus zone (Granger and Simpson, 1929:649) may not be valid be- cause our specimens correspond more closely in size with material from the Ectoconus zone. 122 University of Kansas Publs., Mus. Nat. Hist. Figure 3. Eucosmodon? species. Tor- rejonian (middle Paleocene), head of Kimbetoh Arroyo, San Juan County, New Mexico. No. 7910, occlusal view of P3?, X 5M. Drawing by Jane S. Mengel. In addition to the Puercan material, a single tooth. No. 7910, from Tor- rejonian strata exposed at the head of Kimbetoh Arroyo (KU Loc. 9, NWM sec. 2, T. 23N, R. 9W) may represent P3 of Eucosmodon. If it is this tooth, it differs from that in AMNH No. 16534 in its slightly smaller size, and the absence of a tiny anteromedian cusp. An alternative to a determina- tion of No. 7910 as P3 of Eucosmodon is that it is an exceptionally large P2 of Ptilodus mediaevus. In favor of the latter assignment is the angulation and separation of cusps which seem more nearly like the condition in Ptilodus than in Eucosmodon. The root por- tion of the tooth is mostly gone, and what is left suggests a vague separation, but I guess that there was a single root as in Eucosmodon rather than a divided one as in Ptilodus. The proportions of the crown, nearly quadrate, also favor an assignment to Eucosmodon as does the large size. I am recording this specimen of Eucosmodon, if such it be, because the head of Kimbetoh Arroyo, from whence it comes, was one of Baldwin's main collecting areas for Torrejoii fossils (Sinclair and Granger, 1914:310). Table 1. Measitrements (in millimeters) of Eucosmodon, University of Kansas Collections. Transverse diameter 3.2 3.3 ?3.4 3.6 2.6 1.8 Anteroposterior diameter E. americanus No. 9401 P2 3.1 P4 7.25 Ml M2 3.2 No. 9402 ml 5.2— No. 9403 i 4.1 Eucosmodon? sp. No. 7910 P3 3.5+ 3.5 MULTITUBERCULATE GeNUS EuCOSMODON 123 LITERATURE CITED Granger, W., and G. G. Simpson. 1929. A revision of the Tertiary Multituberculata. Bull. Amer. Mus. Nat. Hist., 56:601-676, 43 figs, in text. Jepsen, G. L. 1940. Paleocene faunas of the Polecat Bench formation. Park County, Wyoming. Proc. Amer. Philos. Soc, 83:217-340, 5 pis., 22 figs, in text. Simpson, G. G. 1928. A catalogue of the Mesozoic Mammalia in the geological depart- ment of the British Musevun. x -f- 215 pp., 12 pis., 56 figs, in text. 1929. American Mesozoic Mammalia. Mem. Peabody Mus. Yale Univ., Vol. Ill, pt. I, XV -f 235 pp., 32 pis., 62 figs, in text. 1937. Skull structure of the Multituberculata. Bull. Amer. Mus. Nat. Hist., 73:727-763, 9 figs, in text. Sinclair, W. J., and W. Granger. 1914. Paleocene deposits of the San Juan Basin, New Mexico. Bull. Amer. Mus. Nat. Hist., 33:297-316, 8 pis., 2 figs, in text. Transmitted August 22, 1955. D 26-1038 OCT 1 =#= University of Kansas Publication! Museum of Natural History Volume 9, No. 7, pp. 125-335, 75 figs, in text June 15, 1956 -^ Mammals of Coahuila, Mexico BY ROLLIN H. BAKER University of Kansas Lav^rence 1956 UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY Institutional libraries interested in publications exchange may obtain this series by addressing the Exchange Librarian, University of Kansas Library, Lawrence, Kansas. Copies for individuals, persons working in a particular field of study, may be obtained by addressing instead the Museum of Natural History, University of Kansas, Lawrence, Kansas. There is no provision for sale of this series by the University Library which meets institutional requests, 01 by the Museum of Natural History which meets the requests of individuals. However, when individuals request copies from the Museum, 25 cents should be included, for each separate number that is 100 pages or more in length, for the purpose of defraying the costs of wrapping and mailing. * An asterisk designates those numbers of which the Museum's supply (not the Li- brary's supply) is exhausted. Numbers published to date, in this series, are as follows: Vol. 1, Nos. 1-26 and index. Pp. 1-638, 1946-1930. *Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140 figures in text. April 9, 1948. Vol. S. *1« iThe avifauna of Micronesia, its origin, evolution, and distribution. By Rol- lin H. Baker. Pp. 1-359, 16 figures in text. June 12, 1951. *2. A quantitative study of the tioctumal migration of birds. By George H. Lowery, Jr. Pp. 361-472, 47 figures in text. June 29, 1951. 3. Phylogeny of the waxwings and allied birds. By M. Dale Arvey. Pp. 473- 530, 49 figures in text, 13 tables. October 10, 1951. 4. Birds from the state of Veracruz, Mexico. By George H. Lowery, Jr. and Walter W. Dalquest. Pp. 531-649, 7 figures in text, 2 tables. October 10. 1951. Index. Pp. 651-681. •Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466. 41 plates, 31 figures in text. December 27, 1951. Vol. 5. 1. Preliminary siu-vey of a Paleocene faunule from the Angels Peak area. New Mexico. By Robert W. Wilson. Pp. 1-11, 1 figure in text. February 24, 1951. 2. Two new moles (Genus Scalopus) from Mexico and Texas. By RoUin H. Baker. Pp. 17-24. February 28. 1951. 3. Two new pocket gophers from Wyoming and Colorado. By E. Raymond Hall and H. Gordon Montague. Pp. 25-32. February 28. 1951. 4. Mammals obtained by Dr. Ciu-t von Wedel from the barrier beach of Tamaulipas, Mexico. By E. Raymond Hall. Pp. 33-47, 1 figure in text. October 1, 1951. 5. Comments on the taxonomy and geographic distribution of some North Ameri- can rabbits. By E. Raymond Hall and Keith R. Kelson. Pp. 49-58. October 1, 1951. 6. Two new subspecies of Thomomys bottae from New Mexico and Colorado. By Keith R. Kelson. Pp. 59-71. 1 figure in text. October 1, 1951. 7. A new subspecies of Microtus montanus from Montana and comments on Microtus canicaudus Miller. By E. Raymond Hall and Keith R. Kelson. Pp. 73-79. October 1. 1951. 8. A new pocket gopher (Genus Thomomys) from eastern Colorado. By E. Raymond Hall. Pp. 81-85. October 1. 1951. 9. Mammals taken along the Alaskan Highway. By RoUln H. Baker. Pp. 87- 117, 1 figure in text. November 28, 1951. *10. A synopsis of the North American Lagomorpha. By E. Raymond Hall. Pp. 119-202, 68 figures in text. December 15, 1951. 11. A new pocket mouse (Genus Perognathus) from Kansas. By E. Lendell Cockrum. Pp. 203-206. December 15, 1951. 12. Mammals from Tamaulipas, Mexico. By Rollin H. Baker. Pp. 207-218. Docembcr 15, 1951. 13. A new pocket gopher (Genus Thomomys) from Wyoming and Colorado. By E. Raymond Hall. Pp. 219-222. December 15, 1951. 14. A new name for the Mexican red bat. By E. Raymond Hall. Pp. 223-226. December 15, 1951. 15. Taxonomic notes on Mexican bats of the Genus Rhogeessa. By E. Raymond Hall. Pp. 227-232. AprQ 10. 1952. 16. Comments on the taxonomy and geographic distribution of some North Ameri- can woodrats (Genus Neotoma). By Keith R. Kelson. Pp. 233-242. April 10. 1952. (Continued on inside of back cover) University of Kansas Publications Museum of Natural History Volume 9, No. 7, pp. 125-335, 75 figs, in text June 15, 1956 Mammals of Coahuila, Mexico BY ROLLIN H. BAKER University of Kansas Law^rence 1956 University of Kansas Publications, Museum of Natural History Editors: E. Raymond Hall, Chairman, A. Byron Leonard, Robert W. Wilson Volume 9, No. 7, pp. 125-335, 75 figs, in text Published June 15, 1956 fas. COMF. ZQOl 0CT16i95S IHHYERSmr University of Kansas Lawrence, Kansas PRINTED BY FERD VOILAND. JR.. STATE PRINTER TOPEKA. KANSAS 19S6 26-1204 Mammals of Coahuila, Mexico BY ROLLIN H. BAKER CONTENTS PAGE Introduction 127 Topography and Climate 128 Plant-mammal Relationships 132 Lower Sonoran Life-zone 134 Upper Sonoran Life-zone 136 Transition Life-zone 138 Canadian Life-zone 139 Plant-mammal Relationships in the Sierra de la Madera 140 Characteristics of the Mammalian Fauna 142 Composition and Derivation of the Fauna 142 Topography and Speciation 147 Endemic Species 151 Mammalogical Exploration in Coahuila 152 Conservation of Mammals in Coahuila 154 Methods and Acknowledgments 156 Gazetteer of Localities in Coahuila 158 Check-bst of the Mammals of Coahuila 160 Accounts of Species and Subspecies 165 Literature Cited 329 INTRODUCTION Most of the 144 kinds of mammals, belonging to 103 species, that live in Coahuila must have the ability to live their entire life spans without the opportimity to drink surface water. This is because most of the State's 58,052 square miles are characterized by pro- nounced aridity. Even so, small mammals are notably abundant in some places where edaphic factors are favorable. In most places, however, mammals are uncommon — so scarce that the collector of mammals often is discouraged by the poor yields obtained from his trap-lines. Irregularly spaced mountains rise from the hot arid lowlands and have cool, mesic caiions where grasses are shaded by pines and oaks. These isolated mountains lure the naturalist to ascend them — usually done with considerable difficulty — to learn what mammals live there. My interest in Coahuila first was aroused in 1936, when from the Chisos Mountains of Brewster County, Texas, I viewed the higher Sierra del Carmen across the Rio Grande. The following summer. (127) 128 University of Kansas Publs., Mus, Nat. Hist. as a student wildlife technician with the National Park Service in the then proposed Big Bend National Park, my enthusiasm for a closer inspection of Coahuila increased as the result of a proposal to develop the Big Bend International Peace Park including both the Chisos and the Carmens. My aim in this report is to place on record our accumulated knowl- edge of the kinds and the distribution of the mammals of Coahuila. It is hoped that this information will be used by biologists as a basis for additional study of the fauna of the State and will stimulate in- terest in the proposed Big Bend International Peace Park and help in its establishment. TOPOGRAPHY AND CLIMATE Coahuila lies just east of the center of the broad, northern part of Mexico. The State has an irregular topography made up of mountains and arid plains. Mountains extending in a north-south direction divide the lower lands into two areas, a larger one, a part of the Central Plateau, to the westward and a smaller one, a part of the Gulf Coastal Plain, to the northeastward. The mountains of Coahuila are eastern extensions of the Cor- dillera forming a part of tlie Rocky Mountain- Sierra Madre Oriental Axis as well as a segment of the eastern escarpment of the Mesa del Norte, which is the northern part of the Central Plateau of Mexico (see McBride, 1923). The mountains are either anticlines or broad domes of middle Cretaceous limestones separated by downwarped areas of Upper Cretaceous marls, chalks and sand- stones. Rocks of Permian and Jurassic ages also are exposed in the State. The highest mountains, including the summits of the Sierra Madre Oriental and the Sierra del Ciirmen, are composed of volcanic rocks. Volcanic rocks in smaller quantities occur in scattered places throughout the mountains and plains. In the extreme southern part of Coahuila, highlands extend from the Sierra Madre Oriental, southeast of Saltillo, in a westerly direc- tion. Although discontinuous, this chain of mountains links the higher lands of Nuevo Leon with those of the Sierra Madre Occi- dental of Durango to the west. Most of the mountains of northern and central Coahuila are elon- gated in a northwest-southeast direction. Their principal axis, in Coahuila, begins at the Rio Grande, east of Boquillas, where the river is deeply intrenched in the highlands of the Sierra del Carmen. From these highlands bordering the river, a broad series of more or less continuous ridges and massive uplifts extend southeastward. Mammals of Coahuila, Mexico 129 to the extreme southeastern part of the State. This irregular moun- tain chain is divided, poHtically, into numerous segments; the higher parts, from north-northwest to south-southeast, include the Sierra del Carmen, the Sierra Hermosa de Santa Rosa, the Sierra de la Madera, the Sierra de San Marcos and the Sierra Madre Oriental. This topography has some important breaks that affect mammalian distribution. One is along the watershed of the Rio Salado that drains parts of central Coahuila and extends in a broad valley as far west as Cuatro Cienegas and cuts northwestward of that city through a narrow cafion to head on the Llano de Ocampo. Other breaks, less important as means for mammalian dispersal, are the Puerto San Lazaro, the Rio Salinas, which bisects, in an east-west direction, the escarpment north of Saltillo, and the low pass east of Saltillo. To the westward of this major montane axis are other north-south trending ranges. One series of discontinuous highlands extends from the Sierra de San Vicente, just south of the Rio Grande, through the Sierra del Pino and Sierra del Fuste to the western edge of the Sierra de la Madera. Another series, along the western boundary of Coahuila, extends southward from the Sierra de los Hecheros through the Sierra de Almagre to the Sierra Mojada. Other scat- tered mountainous areas, mostly lower in elevation than those men- tioned above, also help account for the irregular topography of Coahuila. Most mountains in the State do not exceed 6000 feet in elevation, but the altitudes of most higher places have not been measured accurately. Estimations of elevations were obtained usually with an aneroid altimeter in the hands of one member of our field parties or from reports of local ranchers who had approximated the eleva- tion of summits of the mountains using the altimeters in their pri- vate airplanes. Peaks more than 10,000 feet in elevation occur in the Sierra Madre Oriental near the southeastern border. The high- est peaks in the Sierra del Carmen, Sierra del Pino, Sierra de la Madera, Sierra Encarnacion, and the Sierra de Guadalupe are more than 9000 feet. The Sierra de la Encantada, and the Sierra de San Marcos and perhaps other ranges contain peaks more than 8000 feet in elevation. The Serranias del Burro are no higher than 7500 feet. The nortlieastern lowlands are a part of the Coastal Plain of the Gulf of Mexico. Much of this area is flat and composed of Upper Cretaceous rocks overlain by gravels and aeolian deposits, and al- though well-drained by way of the Rio Grande, is not severely 130 University of Kansas Publs., Mus. Nat. Hist. eroded. Westward towards the mountains and upstream along the Rio Grande this lowland resembles in topography the Mesa del Norte in western Coahuila. Elevations of the Gulf Coastal Plain range from as low as 700 feet to as high as 1800 feet. The desert plains of the Mesa del Norte to the westward of the Sierra del Carmen-Sierra Madre Oriental Axis are higher and more dissected than tliose of the Coastal Plain and marked by scattered desert ranges, buttes, low hills with massive rimrock, and igneous knobs. Drainage in the north is into the Rio Grande. South of the Sierra del Pino are a series of undrained basins. The most prom- inent one, surrounding the Laguna de Mayran and called the Bolson de Mapimi, is west of Saltillo. This laguna is now principally a cotton-producing area and irrigated, in part, by the Rio Nazas and the Rio Aguanaval which flow into the area from the south and west. Elevations of the Mesa del Norte in Coahuila range from as low as 1700 feet near Boquillas at the Rio Grande to more than 4000 feet in the southern part of the State. According to Muller (1947:36), four classes of soils occur in Coahuila; these are stony immature-soils, light desert-soils, alkaline desert-soils, and dark loam-soils. Sand dunes, such as are found in the vicinity of Boquillas, are only of local importance. Stony im- mature-soils blanket many areas, especially the foothills of the desert ranges. Light desert-soils are widespread on the Mesa del Norte, west of the mountains. Alkaline soils are present in the undrained basins and result from extreme aridity. Near Cuatro Cienegas a vast intermontane valley is overlain with glistening gypsum (anhydrite) derived from Upper Jurassic and Upper Cretaceous rocks. Dark loam-soils, characteristic of more humid situations, are generally confined to the higher mountains and the Gulf Coastal Plain, east of the mountains. As pointed out by Muller ( loc. cit. ) soils of igneous origin often produce well-developed grasslands adjacent to soils originating from limestone which bear desert shrub. Most of Coahuila is arid; the little rain which falls comes in the warmer months. On the Coastal Plain rainfall is moderate whereas west of the central mountains the rainfall is low; this low rainfall re- flects the continental position of most of the State. At the Rancho La Mariposa, east of the mountains, Mr. David McKellar recorded, over a period of several years, an average annual rainfall of 23 inches. He said that on a ranch west of the mountains less than 17 inches annually were recorded. Probably much of western Coahuila has an annual rainfall of less than 10 inches (see Webb, 1931:93). Ranchers in the lowlands in the vicinity of Castillon, in northwestern Mammals of Coahuila, Mexico 131 Coahuila, told me in 1953 that no appreciable amount of rain had fallen in their area for seven years. Muller (1947:38) writes that Coahuila owes much of its aridity to its latitudinal position ". . . within the subtropical high-pressure belt or the belt of subtropical calms." Northeastern Mexico receives for several months of the year the moisture-laden air from the east and southeast from the Atlantic Ocean and the Gulf of Mexico. When moist air reaches the north- south trending mountain barriers, it is cooled below the saturation point and the characteristic summer rains in the mountains occur as a consequence. The highest and most easterly mountains receive most of this rainfall; mountains to the westward receive propor- tionately less precipitation. The lowlands of the Mesa del Norte, west of the mountains, receive the least rain. Furthermore, the Sierra Madre Occidental, west of the Mesa del Norte, acts as a barrier to divert air moving eastward from the Pacific Ocean. The Sierra Madre Oriental in extreme southeastern Coahuila receives the highest rainfall; this heavy precipitation is reflected by the luxuriant growth of boreal plants living in the higher places there. Dry farming is practiced in the narrow montane valleys in that area; in most other parts of Coahuila crops must be irrigated. The winters have cool to cold temperatures. Northerly winds sweep down between the north-south trending mountains producing the chilling "nortes." Snow is reported from the higher mountains; Robert Dickerman observed a heavy snowfall in the Sierra de la Madera in December, 1953. Evaporation is highest on the south-facing slopes which are ex- posed to a maximum degree to the direct rays of the sun. Usually the most luxuriant growths are on the northeastern slopes, where the rainfall is near the maximum and exposure to direct sunlight is near the minimum. Springs and other surface water are either scarce or absent west of the mountains. The only permanent streams in this part of Coahuila are the Rio Grande, at the northern border, and the Rio Nazas and Rio Aguanaval, in southwestern Coahuila. The Rio Grande, in many places flowing through steep-walled canons, is the only streamway which bars the passage of certain species. In eastern Coahuila springs, flowing streams and other surface waters are more generally distributed. A few seep-springs occur in the foothills of mountains composed of massive limestone, but higher in the mountains usually no water is present except where an occasional hole (tinaja), in the floor of a narrow canon, contains 132 University of Kansas Publs., Mus. Nat. Hist. rain water. Grasslands have been spared from intensive grazing by domestic livestock because of the absence of adequate water for the animals on the Sierra del Pino and surrounding lowlands, the Sierra de la Encantada, the Sierra de los Hecheros, the Sierra de San Marcos, the Sierra de la Madera and other mountains. In 1952, ranchers were building earth dams in the vicinity of the Sierra del Pino to impound water so that livestock may utilize these grazing lands. PLANT-MAMMAL RELATIONSHIPS Climatic and edaphic factors interact to influence the distri- bution and abundance of vegetation in Coahuila. Mammals gen- erally are confined to specific kinds of plant associations from which they derive either food or shelter or both. Once the in- vestigator has learned the ecological preferences of a given kind of mammal, he can map the occurrence of that mammal by noting the occurrence of the plants. The biotic communities of Coahuila might be divided in ac- cordance with the three physiographic areas of the State: the Gulf Coastal Plain, the mountains, and the desert plains of the Mesa del Norte. However, the irregular topography and the inter- action of various environmental influences, producing numerous micro-habitats, make a more detailed appraisal desirable. In the past, workers concerned chiefly with vegetation have analyzed the biotic communities of Coahuila. Goldman and Moore (1945:349) who were interested in large areas containing natural groupings — chiefly plants, mammals and birds — consider that Coahuila con- tains parts of three biotic provinces. These are: the Chihuahua- Zacatecas Biotic Province, in the western half of the State; the Tamaulipas Biotic Province, in the northeastern part of the State; and the Sierra Madre Oriental Biotic Province in the southeastern part of the State. Leopold (1952) mapped the Vegetational-Zones (Zonas de Vegetacion) of Mexico and assigned parts of five such areas to Coahuila. Beginning at the lowest elevations he defines the Mesquite-Grassland (Mezquite Pastizal), the Desert (Desierto), the Chaparral (Chaparral), the Pine-Oak Forest (Bosque Pino Encino), and the Boreal Forest (Bosque Boreal). Merriam (1898) mapped, in a preliminary way, the life-zones of Coahuila. Using the concept which he proposed, definable portions of the Lower Sonoran Life-zone, the Upper Sonoran Life-zone, the Transition Life-zone, and the Canadian Life-zone can be distinguished (see fig. 1). Most of the State lies within the Lower Sonoran; less Mammals of CoAHtriLA, Mexico 133 than 5 per cent of the area is within the Transition and Canadian Hfe-zones. The most detailed analysis of the distribution of vege- tation in Coahuila is that of Muller (1947), who recognized seven vegetation-types . Fig. 1. Life-zones of Coahuila. For the study of mammalian distribution, I find that Merriam's life-zones are the most satisfactory major divisions, and Muller 's vegetation-types provide the most useful minor divisions. These two arrangements, beginning at the lowest elevations, are con- 134 University of Kansas Publs., Mus. Nat. Hist, sidered together in the following discussion of plant-mammal relationships. Those mammals, listed as characteristic of one life- zone (or vegetation-type) but also found in one or more other life-zones (or vegetation-types) are marked with an asterisk. Lower Sonoran Life-zone More than three-fourths of the total area of Coahuila is included in the Lower Sonoran Life-Zone (see fig. 1). This expanse in- cludes areas ranging in elevation from less than 800 feet in the northeast to more than 5000 feet in the south and southwest. Using Muller's floral divisions, two vegetation-types are distinguishable: the Tamaulipan Thorn Shrub and the Chihuahuan Desert Shrub. Characteristic mammals of the Lower Sonoran Life-zone are: Didelphis marsupialis Spcrmophilus mexicanus Notiosorex crawfordi SpermopJiilus spilosoma* Pipistrellus hes^^erus Thomomys bottae* Pipistrellus sitbflavus Thomomys umbrinus Sylvilagus audubonii Cratogeomys castanops* Sylvilagus floridanus* Ferognathus hispidus* Lepus californicus* Ferognathus merriami Ferognathus nelsoni Feromyscus maniculatus* Ferognathus penicillatus Feromyscus melanophrys Dipodomys merriami Sigmodon hispidus Dipodomys ordii* Neotoma albigula* Reithrodontomys fulvescens* Neotoma goldmani Reithrodontomys megalotis* Neotoma micropus Onychomys leucogaster Vulpes macrotis Onychomys torridus* Fecari tajacu Baiomys taylori Odocoileus hemionus* Feromyscus eremicus* Odocoileus virginianus* Feromyscus leucopus Ovis canadensis* * Denotes occurrence also in at least one other life-zone. Tamaulipan Thorn Shrub. — East of the Sierra del Carmen- Sierra Madre Oriental Axis, the foothills and, to the northward, the Upper Rio Grande Plain give way to the Gulf Coastal Plain, which is characterized by the Tamaulipan Thorn Shrub Vegetation-type. Here, the vegetation is most nearly like that of southern Texas, northern Nuevo Leon, and northern Tamaulipas. Deep loam- soils offer suitable places for a rich grass flora interspersed with shrubs and small trees; in some areas, where grazing is not severe, this floral complex has the appearance of a savannah. Muller (1947:42) lists several dominant shrubs including three species Mammals of Coahuila, Mexico 135 of catclaw (Acacia spp.), mesquite (Prosopis glandulosa), soap- bush (Porlieria angustifolia) , anacahuita (Cordia Boissieri), and blue paloverde (Cercidittm foridum). Grasses include grama ( Bouteloua trifida ) and curly mesquite ( Hilaria Belangeri ) . Thick- ets of live oak (Qiicrcus sp.) occur east of Nava, especially along the flood plain of the Rio Grande. Parklike stands of willow (Salix), pecan (Carya), hackberry (Celtis), cypress (Taxodium) and otlier trees occur along the Rio de San Diego, Rio Salado, Rio Sabinas, and other permanent streams. Mammals characteristic of the Tamaulipan Thorn Shrub are: Stjlvilagus floridanus chapmani Ontjchomys leucogaster Spermophilus mexicanus * Peromyscus leucopus texanus Perognathus hispidus hispidus Baiomys taylori Perognathus mcrriami merriami Neotoma micropus * Reithrodontomys fulvescens intermedius * Odocoileus virginianus texanus Along streams, in the riparian growth occur: Pipistrellus subflavus Castor canadensis Nycticcius humeralis Dasypus novemcinctus Sciurus niger * * Denotes occurrence also in at least one other vegetation-type. Chihuahuan Desert Shrub. — The plains and basins of western and southern Coahuila are covered with this vegetation-type, which is characterized by low, sparse perennials and annuals scattered among low shrubs, of which creosote ( Larrea tridentata ) is often the most common species. Associated with creosote are blackbrush (Flourensia cernus), catclaw (Acacia vernicosa), ocotillo (Fou- quieria splendens), mesquite (Prosopis velutina), various cacti (in- cluding Opuntia spp.) and yuccas (Yucca spp.). Low limestone hills and rocky slopes bear succulent desert plants such as Agave lecheguilla, Euphorbia antisyphilitica, Jatropha dioica, and various species of Opuntia, Echinocactus, and Mamillaria. Grusonia Brad- tiana grows on rocky hills in the southwestern part of the State. Muller (1947:39) emphasizes the homogeneous climatic conditions of the area covered by this vegetation-type and points out that edaphic variations are responsible for major diflFerences in vegetation. This vegetation-type occurs east of the Sierra del Carmen-Sierra Madre Oriental Axis around the Serranias del Burro and on the Upper Rio Grande Plain at least as far southeast as the vicinity of Villa Acufia. Some mammals, notably Odocoileus hemionus, from 136 University of Kansas Publs., Mus. Nat. Hist. western Coahuila range into this area, and mammals characteristic of the Tamauhpan Thorn Shrub also occur there in some places. Mammals characteristic of the Chihuahuan Desert Shrub are: Species Dipodomys i\elsoni Notiosorex crawfordi Reithrodontomys megalotis * Pipistrellus hesperus Onychomys torridus * Spermophilus spilosoma * Peromyscus maniculatus * Ammospermophilus interpres Peromyscus melanophrys Thomomys bottae * Neotoma albigtda * Thomomys umbrinus Neotoma goldmani Perognathus nelsoni Vulpes macrotis Perognathus penicillatus Odocoileus hemionus * Dipodomys merriami * Ovis canadensis * * Denotes occurrence also in at least one other vegetation-type. Upper Sonoran Life-zone Less than one-fourth of Coahuila lies within the Upper Sonoran Life-zone ( see fig. 1 ) . This life-zone encompasses the higher parts of the lesser mountains but is more pronounced in the foothills and the southern exposures of upper slopes of the highest mountains. This life-zone occurs in eastern Coahuila at elevations lower than 3000 feet in the Sierra Azul and Serranias del Burro but in western Coahuila occurs usually above 5000 feet. Four of Muller's vege- tation-types, the Piedmont Shrub, the Grassland (and Grassland Transition), the Montane Low Forest, and the Montane Chaparral, pertain to the Upper Sonoran Life-zone. In all of these vegetation- types except the Grassland, oaks {Quercus spp. ) are prominent. Characteristic mammals of the Upper Sonoran Life-Zone are: Sylvilagus floridanus * Peromyscus difficilis * Cynomys mexicanus Peromyscus nasutus * Spermophilus variegatus * Peromyscus pectoralis * Sciurus alleni * Peromyscus truei Thomomys bottae * Sigmodon ochrognathus * Peromyscus boylii * Odocoileus virginianus * * Denotes occurrence also in at least one other life-zone. Piedmont Shrub. — MuUer (1947:43) considers this vegetation- type to be a poorly defined belt of large shrubs and small trees which occur on irregular topography along the east-facing bases of the mountains that bound the western limit of the Tamaulipan Thorn Shrub. Outlying low hills on the Coastal Plain also bear this vegetation. In some places open forests of arborescent palm (Brahea hella) exist on grassy valley floors and lower mountain slopes of the east face of the Sierra del Carmen, Serranias del Burro, and east of Hermanas on the Sierra Azul and other ranges. Species Mammals of Coahuila, Mexico 137 which are most numerous in this vegetation-type include several kinds of oaks {Quercus spp.), black persimmon (Diaspyros texana), woolly buckthorn (Bumelia lanuginosa), and mescalbean (Sophora sectindifiora ) . Montane Low Forest. — This vegetation-type is not a homo- geneous one and consists of open stands of low oaks, piiion and junipers with rounded crowns and gnarled branches, an abundant grass cover, and one or more species of Agave. Such plant com- munities are found principally on eastern slopes of mountains along the eastern escarpment, Muller (1947:50) lists as dominant trees several species of oaks, including Grave's oak (Quercus gravesii), pifion (Pinus cembroides) , alhgator juniper (Juniperus pachy- phloea), madrono (Arbutus xalapensis), and flowering ash (Frax- inus cuspidata. ) . This vegetation-type often is called the oak-pinyon- juniper association. Montane Chaparral. — This shrubby vegetation-type is on moun- tainous slopes, usually where soils are shallow and is especially con- spicuous on upper slopes of the Sierra de los Hecheros, Sierra del Pino, Sierra de la Encantada, Sierra de la Madera, Sierra de San Marcos and other high mountains. Oaks dominate this flora; many of the species are evergreen, and include Quercus intricata, Q. in- vaginata, Q. Pringlei, Q. Laceyi, and Q. hypoxantha. This xeric chaparral is dense and shed oak leaves form thick ground cover. Grasses are sparse; several kinds of grammas (Bouteloua spp.) are present. The three vegetation-types. Piedmont Shrub, Montane Low Forest and Montane Chaparral, offer equal attraction to many mammals of the Upper Sonoran Life-Zone. Acorns and piiion nuts provide seasonal foods for many mammals including such large species as Ursus americanus and Odocoileus virginianus. Some species, Syl- vilagus foridanus, Peromyscus nasutus, P. pectoralis, P. difficilis, prefer the thicker cover, especially the chaparral. Others, including Sciurus alleni, Spermophilus variegatus, Thomomys bottae, Sigmo- don ochrognathus, select more open stands of timber, where grassy cover and rocky prominences are interspersed with larger oaks and pifions. Mammals characteristic of these three vegetation-types are: Kinds Peromyscus difficilis * Sylvilagus floridanus nelsoni * Peromyscus nasutus * Spermophilus variegatus * Peromyscus pectoralis * Sciurus alleni * Peromyscus truei Thomomys bottae * Sigmodon ochrognathus Peromyscus boylii Odocoileus virginianus * * Denotes occurrence also in at least one other vegetation-type. 138 University of Kansas Publs., Mus. Nat. Hist. The fox squirrel (Sciurus niger) occurs locally in oak forests of the Upper Sonoran Life-zone in the Serranias del Burro and where large oaks grow along caiions in the foothills of the mountains near the headwaters of the Rio Sabinas west of the Hacienda La Mari- posa. The mole (Scalopus montanus) is known only in the oak belt in the Sierra del Carmen. Grassland. — According to Muller (1947:46), "True climatic grassland is not extensive in Coahuila. The greater proportion of the grass-covered area never develops much beyond a transition between desert shrub and grassland, further development being ter- minated by failure of elevation or by the development of chaparral. . . . Since grassland requires an area of relatively deep, flat soil and a considerably more mesic climate than that of the desert plains, it is to be sought only at higher elevations on gently sloping hills and plateaus." He adds, "Grassland Transition, by contrast, is very widespread, occurring about the flanks of most of the mountain ranges." The dominant grass is blue grama (Bouteloua gracilis) with side-oats grama (B. ctirtipendula) , silver beardgrass (Andro- pogon saccharoides) , wolftail (Lycurus phleoides), needle grass (Stipa eminens), three-awn {Aristida glaiica), buffalo grass (Buch- loe dactyloides) , and mesa muhly {Muhlenbergia monticola) locally abundant or even prominent. Numerous perennial herbs also are present. Oaks, junipers, sotol {Dasylirion), beargrass (Nolina), and Yucca grow interspersed in grassland, especially in rocky places. In the Grassland Vegetation-type in the highlands of central and northern Coahuila, the most characteristic mammal is Sigmodon ochrognathus. Grasslands, mostly overgrazed, occur in the high intermontane valleys of southeastern Coahuila. Small mammals living in these areas frequently are subspecifically different from related mammals living in the Lower Sonoran Life-zone. Mammals characteristic of this part of southeastern Coahuila are: Kinds Perognathus flaws medius Lepus californicus asellus * Perognathus hispidus zacatecae Cynomys mexicanus Dipodomys ordii durranti * Spermophilus spilosoma pallescens * Onychomys torridus surrnfus Cratogeomys castanops suhnubilus Peromyscus maniculatus blandus * * Denotes occurrence in at least one other vegetation-type. Transition Life-zone Less than one-fifth of Coahuila lies within the Transition Life- zone. The dominant tree is the Arizona pine (Pinus arizonica). Mammals of Coahuila, Mexico 139 It may occur in association with large oaks and Douglas fir (Pseudo- tsnga taxifoUa). Montane Mesic Forest. — Truly mesic forest occurs only near the summits of the highest and most massive mountains of the State. Stands of tall trees with shade-tolerant plants as understory characterize this forest. The surface soil has a well-developed humus overlain by leaf litter. The moisture content of the soil is moderate or high throughout the summer and aestivation of the plants normally does not take place. According to Muller ( 1947:55) the principal trees are Arizona pine (Pinus arizonica), Douglas fir (Pseitdotsuga taxifolia), Arizona cypress (Cupressus arizonica), oaks (Quercus Gravesii, Q. hypoleiicoides, Q. Muehlenbergii) , quaking aspen (Poptdus tremuloides), and southwestern maple (Acer brachypterum) . A fir (Abies coahuilensis) grows on some mountains. The easternmost mountains, such as the Sierra del Carmen and the Sierra Madre Oriental, possess the best developed forests, and logging operations there often are extensive. The lesser ranges and those to the westward, which are more subject to continental influences, bear more depauperate pine forests. Mountains with conspicuous stands of pine are Sierra de la Encantada, Serranias del Burro, Sierra del Pino, Sierra de la Madera, Sierra de San Marcos, Sierra de Guadalupe, Sierra de Parras, and the Sierra Encarnacion. Sparse groves of pine occur as low as 3000 feet in the Serranias del Burro, probably because the eastward posi- tion of this range allows it to receive more moisture from undi- verted air from the Gulf of Mexico. Small stands of pine also grow in protected valleys in many other highlands; such small, isolated areas influence the distribution of mammals only sHghtly. Mammals characteristic of this vegetation-type are: Kind Eutamias bulleri* Sorex milleri* Eutamias dorsalis Mijotis planiceps Reithrodontomys megalotis saturatus Mtjotis volans Peromyscus difficilis* Sylvilagus floridanus nelsoni* Peromyscus nasutus* Sylvilagus floridanus orizabae* Neotoma mexicana* * Denotes occurrence also in at least one other vegetation-type. Canadian Life-zone Plants indicative of the Canadian Life-zone occur near the summits of at least four of the highest mountains of Coahuila. Small groves of quaking aspen (Populus tremuloides) exist on the 140 University of Kansas Publs., Mus. Nat. Hist. Sierra del Pino and the Sierra del Carmen. No mammals character- istic solely of the Canadian Life-zone occur in these mountains. The highest parts of the Sierra Madre Oriental, east of San Antonio de las Alazanas and at the border of Nuevo Leon, possess extensive stands of aspens. In the Sierra Madre Oriental the pine- fir-aspen forests, above 9000 feet in elevation, are mesic and luxuriant. Herbs flourish in the deep, moist humus-soils through- out the growing season. Climatic conditions and the composition of the flora, in these high elevations, resemble closely those de- scribed by Muller (1939:703-704) for the Subalpine Humid Forest Vegetation-type of the Sierra Madre Oriental in Nuevo Leon. Many of the same plants also exist on top of the Sierra de Guada- lupe which is situated westward of the Sierra Madre Oriental. Subalpine Humid Forest. — Mammals characteristic of this vege- tation-type in southeastern Coahuila are: Sorex milleri* Peromyscus melanotis Sorex saussurei Neotoma mexicana* Eutamias bulleri* Microtus mexicanus * Denotes occurrence also in at least one other vegetation-type. Plant-Mammal Relationships in the Sierra de la Madera The Sierra de la Madera, situated in central Coahuila, extends in a west-northwest to east-southeast direction for approximately 30 miles. The highest elevations are at least 9000 feet, and the higher slopes are covered with pine forest. The northeastern foothills give way to the relatively undissected intermontane plain, the Llano de Ocampo, which near the base of the foothills is ap- proximately 3000 feet in elevation. The vegetation-types of the lower elevations extend into the foothills, but the expanse of this growth depends on the amount of exposure to direct rays of the sun and the quality and the moisture-content of the soil. Like- wise, the vegetation-types occurring at the higher elevations extend lower where environmental conditions permit, such as in protected canons. Field parties obtained mammals at 12 localities on the northern approaches and in these mountains in 1952, 1953, and 1954. Parties ascended the mountains on two occasions by way of the Caiion de la Charretera. Base-camps were made at 5300 feet and at 5950 feet, from where excursions were taken to ele- vations of more than 8000 feet. Collections of mammals also were obtained from as low as 3250 feet in elevation on the Llano de Ocampo near the northward-facing base of the foothills of the Sierra de la Madera. Mammals of Coahuila, Mexico 141 Ascending the mountains by way of the Canon de la Charretera, the Chihuahuan Desert Shrub occurs, within the valley, up to approximately 4000 feet in elevation. Above this vegetation-type Grassland Transition flourishes, in which several kinds of daggers (Yucca spp. ), beargrass (Nolina sp.), and sotol (Dasijlirion sp. ) are conspicuous. The grassy cover, ungrazed by domestic live- stock, becomes progressively thicker with increase in elevation. At 4900 feet the first shrubby oaks (Qtiercus spp.) mark the be- ginnings of the Montane Chaparral. Farther up the canon clumps of oaks are larger and grow in association with madroiio (Arbutus xalapensis), junipers (Juniperus spp.) and cherry (Prunus sp. probably capuli). At 5300 feet in elevation grasses, mostly gramas (Bouteloua spp.) are interspersed between clumps of chaparral. Some prickly-pear (Opuntia sp. ) and mesquite (Prosopis glandu- losa) also occur in the floor of the canon. Beginning at 5400 feet the oak trees are larger and grow in more open stands and are -Syfviloflus fforidonus- Peromyscus nosirfus- -Eufomios dorsolis — -Neofoma mtxrcono _Peromyscu» '■ pectoralis ': P«romyscus ertmicus- -Spermophllus spilosomo — - Cratogaomys cosfonops- -Perognothus flavus -P»rognothu$ natsoni -Perognolhus penicillotus- -Dtpodomys marriomi -Oipodomys nelsont -Neofomo otbigula Fig. 2. Diagrammatic profile of the north slope of the Sierra de la Madera, showing altitudes and distributions of vegetation-types and small mammals. indicative of the Montane Low Forest Vegetation-type of MuUer. The first Arizona pine (Pinus arizonica) appears at 5500 feet. A Montane Mesic Forest, principally pine-oak, is reached at approxi- mately 6500 feet in the narrowing canon. At 6500 feet the first 142 Unr'ersity of Kansas Publs., Mus. Nat. Hist. Douglas fir (Psuedotstiga taxifolia) occurs; at 7500 feet the forest is chiefly pine-fir. Above 7500 feet large pine, fir and Arizona cypress (Cupressus arizonica) grow in the canon and on adjacent slopes. Another fir {Abies probably coaJiuilensis) also is found in this area. Seventy-four small mammals, belonging to 15 species, were taken at the 12 collecting stations. The altitudinal distribution of these species is shown graphically on fig. 2. Most kinds of mammals are restricted to one or two vegetation-types. Peromtjs- cus eremicus, common in the Chihuahuan Desert Shrub, occurred as a "straggler" at 6000 feet in the Montane Low Forest. Else- where in Coahuila it is also true that more kinds of small mammals inhabit the less elevated vegetation-types than those more elevated. CHARACTERISTICS OF THE MAMMALIAN FAUNA Composition and Derivation of the Fauna Genera. — Genera of Recent North American mammals, from the standpoint of distribution, may be assigned to one of three major faunal assemblages: the Tropical, the Sonoran, and the Boreal. Recently Hooper (1949:23-24) has emphasized the affinities of the rodents to these basic groupings. Most mammalian genera in each of these faunal assemblages have, in common, somewhat similar distributions and environmental requirements and often similar geographic centers of maximum differentiation (see Hooper, loc. cit.). Coahuila receives mammals from all three of these groupings. As shown in Table 1, 56 genera of native mammals occur in Coahuila. These include 43 non-flying (terrestrial) genera and 13 genera of bats. The Tropical element, having affinities with the fauna of the lowlands of Central America and adjacent parts of South and North America, is represented by 8 genera (19 per cent) of non-flyers and 6 genera (42 per cent) of bats. The So- noran element, with affinities with the fauna of the arid parts of western United States and the Central Plateau, is represented by 18 genera (42 per cent) of non-flyers and one genus (8 per cent) of bats. The Boreal element, with affinities with the fauna of the high latitudes of North America and the higher elevations in the middle and lower latitudes of North America, is represented by 17 genera (39 per cent) of non-flyers and 6 genera (42 per cent) of bats. Most genera (see Table 1) have either Sonoran or Boreal affinities but the majority occur only in the lower elevations; 79 Mammals of Coahuila, Mexico 143 Table 1. — Affinities and Distribution of Genera of Mammals of Coahuila Genus (number of species in parentheses) Didelphis (1) Sorex (2) Notiosorex (1) Scalopus (1) Aello (1) Choeronycteris (1). . . . Leptonvcteris (1) Myotis' (6) Pipistrellus (2) Eptesicus (1) Lasiurus (2) Dasypterus (1) Nycticeius (1) Corynorhinus (1) Antrozous (1) Tadarida (1) Eumops (1) Dasypus (1) Lepus (1) Sylvilagus (2) Cynomys ( 1 ) Spermophilus (3) Ammospermophilus (1) Eutamias (2) Sciurus (2) Thomomys (2) Cratogeomys (1) Perognathus (5) Dipodomys (3) Castor (1) Reithrodontomys (2) . . Perom.yscus (10) Baiomys (1) Onychomys (2) Sigmodon (2) Neotoma (4) Microtus (1) Erethizon (1) Canis (2) Vulpes (1) Urocyon (1) Ursus (1) Bassariscus (1) Procyon (1) Nasua (1) Mustela (1) Taxidea (1) Affinities 03 "E, o Si H X X X xr X X X V? 03 (-1 O c o X X X X? X X X X X X X X X o3 O X X X X X X X X X X X X X X X X X Distribution as to Life-zones O c ^ o QJ X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X c '-3 p H X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X a c o X X 144 University of Kansas Publs., Mus. Nat. Hist, Table 1. — Concluded Genus (number of species in parentheses) Affinities o c a t^ o c o O Distribution as to Life-zones c ^ O S o c 03 a 03 U Spilogale (1) . . Mephitis (2) . . Conepatus (1). FeHs (4) Lynx(l) Pecari (1) Odocoileus (2) . Antilocapra (1) Ovis (1) Totals X X X X X X X X X X X X X X X X X X X X X X X X X X 14 19 23 44 42 21 per cent are in the Lower Sonoran Life-zone. Representatives of 37 per cent of the genera have been recorded from the Transition Life-zone, whereas only 16 per cent hve in the Canadian Life-zone. Species. — The species of mammals that occurred in Coahuila in the Pleistocene are unknown. Probably in Wisconsinan time, some of the presentday species were already living in what is now Coahuila, but others are now extinct or live elsewhere. Cush- ing (1945:182) records Marmota and Synaptomys from Quaternary deposits in Nuevo Leon; these Boreal genera are absent from both Nuevo Leon and Coahuila today. Certainly, many mammals have moved into the State since the glacial epochs. Because no knowledge of earlier mammalian faunae is available, it seems best merely to point out the affinities of Recent species in Coahuila with those which now occur in other areas. Species do not necessarily differentiate within the areas from where genera, to which the species are assigned, are derived. For example, 24 species of non-flying mammals in Coahuila belong to 17 genera of Boreal origin, yet only four species (two of Sorex and one each of Etitamias and Microtus) are restricted, in the State, to the Canadian (and Transition) Life-zone, while others are most common in the Upper and Lower Sonoran life-zones (see Table 1). At least two species (Pcromysctis melanotis and Neo- toma mexicana), which belong to genera of Sonoran derivation, are restricted to the Canadian Life-zone and/or the Transition Life- zone. Mammals of Coahuila, Mexico 145 Some non-flying mammals living at lower elevations in Coahuila have extensive continental distributions, but others with less en- vironmental tolerance have more limited distributions, which reflect their adaptations to restricted habitats. These latter species have afiinities with the present-day faunae, or moved into Coahuila, from: (1) tropical lowlands of eastern Mexico; (2) northern part of the Gulf Coastal Plain; (3) Mesa del Norte, northern part of the Central Plateau; (4) Mesa Central, southern part of the Central Plateau; and (5) Great Plains of the United States. Mammals which have closest affinities with the fauna on the tropical lowlands of eastern Mexico include: Didelphis marsupialis Felis pardalis Dasypus novemcinctus Felis onca Sigmodon hispidus? Felis yagouaroundi Mammals which have closest affinities with the fauna on the northern part of the Gulf Coastal Plain include: Sciurus niger Baiomys taylori Spermophilus mexicanus Neotoma micropus Perognathus merriami Mammals which have closest affinities with the fauna on the Mesa del Norte include: Notiosorex crawfordi Cratogeomys castanops Spermophilus spilosoma Spilogale gracilis Ammospermophilus interpres Vulpes macrotis Perognathus nelsoni Antilocapra americana Perognathus penicillatus Ovis canadensis? Neotoma goldmani? Odocoileus hemionus Mammals which have closest affinities with the fauna on the Mesa Central include: Peromyscus melanophrys Conepatus mesoleucus Mammals which have closest affinities with the fauna on the Great Plains include: Scalopus montanus Omjchomtjs leucogaster Lepus californicus Mephitis mephitis? Cynomys mexicanus Montane mammals, Hving in Coahuila, have affinities with the fauna of either one or another of three different montane areas. Mammals found on the mountains of the northern three-fourths of Coahuila, north of latitude 26° N, are most closely related to mammals living on mountains to the northwest in New Mexico and other parts of the western United States. Montane mammals of the southern one-fourth of the State south of latitude 26° N, are most 146 University of Kansas Publs., Mus, Nat. Hist. closely related either to mammals to the west on the mountains of Durango and Zacatecas or to mammals to the south on the moun- tains of Nuevo Leon, Tamaulipas, Veracruz and Hidalgo. Sorex milleri, a species with northwestern aflBnities, occurs as a relict both north of latitude 26° N, on the Sierra del Carmen, and south of latitude 26° N, on the Sierra Madre Oriental east of San Antonio de las Alazanas and on Cerro Potosi in Nuevo Leon (see Findley, 1955). Montane mammals may have reached Coahuila either by way of: (1) a northern route, along the eastern escarpment of the Rocky Mountains in New Mexico and Texas; (2) a trans-plateau route, through mountains across the Mesa del Norte from Durango to southern Coahuila; or (3) a southern route, along the Sierra Madre Oriental (see fig. 3). Fig. 3. Routes by which montane mammals may have reached Coahuila. (1) Northwestern route. (2) Trans-plateau route. (?) Southern route. Mammals of Coahuila, Mexico 147 These present, discontinuous "routes" at an earlier time may have either been continuous or had suitable habitat. In either case mon- tane mammals would have had opportunity to move into Coahuila. Perhaps these routes did not allow the passage of all possible mon- tane species (see Gushing, 1945:182), because the presentday iso- lated, montane mammalian fauna of the State is an impoverished one. Many mammals, that might have had the opportunity to reach Coahuila by way of a montane passageway did not do so, or at least are not living in Coahuila today. Some of these mammals, unknown in Coahuila today, are: Sorex vagrans, T amiasciurus hud- sonicus, Spermophilus lateralis, Clethrionomijs gapperi, Microtus longicaudus, and Zapus princeps living on mountains to the north- west; Sorex oreopolus, Sciurus aherti, Sciurus nayaritensis, Neotoma ferruginea, and Nelsonia neotomodon Hving on mountains to the west; and Sciurus oculatus and Neotoma torquata living on moun- tains to the south. Montane mammals which have closest aflBnities with the fauna on mountains to the northwest and only known in Coahuila from north of latitude 26° N are: Sylvilagtis floridanus nelsoni Sigmodon ochrognathus Eutamias dorsalis Neotoma mexicana inornata Peromyscus boylii rotvleyi Erethizon dorsatum Peromyscus nasutus Odocoileus virginianus carminis Montane mammals having closest affinities with those on moun- tains to the west and that are known in Coahuila only from south of latitude 26° N are: Sciurus alleni? Peromyscus truei Eutamias bulleri Neotoma mexicana navus Montane mammals having closest aflBnities with those on moun- tains to the south and that are known in Coahuila only from south of latitude 26° N are: Sorex saussurei Peromyscus difficilis Sylvilagtis floridanus orizahae Peromyscus melanotis Reithrodontomys megalotis saturatus Microtus mexicanus Peromyscus boylii levipes Odocoileus virginianus miquihuanensis Topography and Speciation On preceding pages the major habitats of mammals in Coahuila have been described. All mammals, as Hall (1946:64) and others have pointed out, reproduce geometrically and their tendency, as the population grows, is to push outwardly in search of new living spaces. This spread is retarded and ultimately stopped by en- vironmental barriers. Movements of those mammals with narrow 148 University of Kansas Publs., Mus. Nat, Hist. ecological tolerances often are checked more easily by marginal environmental conditions than those mammals with wide ecological tolerances. We have found also that in specific environmental situations in Coahuila certain mammals should be expected. However, filter- barriers, usually of a physical sort, keep some mammals from occupying all suitable habitats available to them but allow others to pass and occupy such areas. Three important filter-barriers — see the concept of the filter bridge in Simpson (1953:21) — to the passage of small, non-flying mammals in Coahuila (see fig. 4) are: ( 1 ) the Sierra del Carmen-Sierra Madre Oriental Axis Filter-Barrier; (2) the Rio Grande Filter-Barrier; and (3) the Southern Coahuila Filter-Barrier. Mammals of the same species that live on either side of these filter-barriers often are isolated from each other and differ in morphology to the extent that they are classified as diflFerent subspecies. Sierra del Car men- Sierra Madre Oriental Axis Filter- Barrier. — It is supposed that at one time, this eastern escarpment of the Cordillera was a continuous highland, separating the Gulf Coastal Plain from the Mesa del Norte, and that subsequent ero- sion and other geological processes produced gaps and low passes in this escarpment which not only restricted montane mammals to isolated highlands but also produced environments allowing some lowland-dwelling plants and animals to pass from east to west and in the opposite direction. Examples of these passes are at the headwaters of the Rio Salado-Sabinas and the Rio Salinas, in Puerto San Lazaro, and through the low pass east of Saltillo. Most of the plants and animals moved from west to east with the result that today the eastern foothills of the Sierra del Carmen, Serranias del Burro and other eastern mountains possess a fauna and flora more characteristic of the Chihuahuan Desert Shrub than of the Tamaulipan Thorn Shrub. Three mammals, Didelphis rtiar- supialis, Spermophilus mexicanus, and Neotoma micropus, moved westward, presumably along the watershed of the Rio Salinas, from the Coastal Plain into the vicinity of Saltillo in southeastern Coahuila. No small mammals have made their way west from the Tamaulipan Thorn Shrub to the deserts of northwestern Coa- huila, but several species from the Coastal Plain, including Spermo- philus mexicanus, Perognathus hispidus, Peromtjsctis leucopus and Onychomys leucogaster, occur on or adjacent to typical Chihuahuan Desert Shrub where this vegetation-type is found east of the montane axis. Mammals of Coahuila, Mexico 149 104 103 102 101 100 1 ^.a^^X^^^;,^ ' ' ~ iu J^^^^^^^o ^^^ >^^^^K C* ^^ ^^^^>% j^Cjm^/ ^£t-rv\, C^ -r?>*^ ^^<'^S(J^^ ^ ^^Si^ ^ 29 /^- 1 29 28 t W -^ 28 \ ^^ S / " \ ^^ 8s ; \ a- .; \ 3 \ ^ ? C ?? — * ^^ 3. ^ — 2/ ^ ^^^^ ^ i \ ^^^ '' \ W-^ /V \ ^^^i>f/^ / ) ^^^ 26 X <<^/^^Si 26 25 25 0 50 IOC ' I 1 1 scale of miles 1 04 103 102 101 100 .. 1 Fig. 4. Major filter-barriers in Coahuila. Small mammals characteristic of the Chihuahuan Desert Shrub, and that show no definable morphologic diflFerence between the west and east side of the montane axis, are: Perognathus nelsoni Peromyscus eremicus Perognathus penicillatus Neotoma albigula Dipodomys merriami 150 University of Kansas Publs., Mus. Nat. Hist. Small mammals characteristic of the Chihuahuan Desert Shrub, which are not found east of the montane axis, are: Spermophilus spilosoma Reithrodontomys megalotis Ammospermophilus interpres Peromyscus maniculatus Perognathus ftavus Neotorna goldmani Dipodomys nelsoni Small mammals, which have representative subspecies on the Mesa del Norte and other subspecies on the Coastal Plain, each subspecies isolated from its corresponding one by the intervening montane axis, are: Sylvilagus audubonii Dipodomys ordii Cratogeomys castanops Peromyscus leucopus Perognathus merriami Lepus calijornicus seems to pass through the montane axis and intergradation between the western subspecies, texianus, and the eastern subspecies, merriami, is discernible in specimens from the intervening mountainous area. Rio Grande Filter-Barrier. — The Rio Grande, although prob- ably a barrier to small mammals where it flows through deep canons along part of the northern boundary of Coahuila, seems not to bar the passage of most mammals where the river's banks are low. Thomomys bottae, Cratogeomys castanops, and Dipodomys ordii are the only mammals which have distinctive subspecies on opposite sides of the Rio Grande, along the Texas-Coahuila border. Probably the Rio Grande, in the lower part of its course, has not always barred the passage of Cratogeomys because that genus reached northeastern Coahuila, northern Nuevo Leon and northern Tamaulipas by crossing the Rio Grande from Texas. At least one mammal, Dipodomys nelsoni, living near the Rio Grande in northwestern Coahuila, is unreported from almost identi- cal habitat in Texas, less than 8 miles to the north. Likewise, Neotoma micropus, unreported from northern Coahuila west of the Sierra del Carmen-Sierra Madre Oriental Axis, is found opposite northwestern Coahuila in Brewster County, Texas. One would suppose that Neotoma micropus is not barred from passage across the Rio Grande along its lower course, because the subspecies, N. m. micropus, occurs in both Texas and northeastern Mexico. Spermophilus spilosoma, unreported in northern Coahuila east of the montane axis, occurs in Texas directly across the Rio Grande from eastern Coahuila. Probably this mammal is able to cross the river, at least near its mouth, because Hall (1951a:38) obtained Mammals of Coahuila, Mexico 151 the South Texan subspecies, Spermophilus spilosoma annectens on a barrier beach in Tamaulipas. Other small mammals occur un- diflPerentiated on both sides of the river. Southern Coahuila Filter-Barrier. — Passage of small mam- mals between broad, intermontane valleys in southeastern Coahuila (within the Upper Sonoran Life-zone) and lower, more arid plains of the vicinity of, and west of, Saltillo (within the Lower Sonoran Life-zone) is for the most part blocked by a high escarpment. In these intermontane valleys there is one endemic species, Cyno- mys mexicantis, and several subspecies of small mammals that occur nowhere else in Coahuila. The subspecies, at least, seem to have reached these valleys by way of the higher plains of Nuevo Leon and San Luis Potosi. On one area, in the vicinity of La Ventura, there is an intermingling of these Upper Sonoran mammals with Lower Sonoran mammals from the northwest. Small mammals of southeastern Coahuila which are not found elsewhere in the State are: Lepus califomicus asellus Perognathtis flavus medius Cijnomys mexicantis Perognathus hispidus zacatecae Cratogeomtjs castanops planifrons Dipodomys merriami atronasus (in intermontane valleys) Onychomys torridus surrufus Cratogeomys castanops stibnubilus Neotoma albigula leucodon (also in (in vicinity of La Ventura and on extreme east-central Coahuila) other lowlands of the southeast) Small mammals that live in the vicinity of La Ventura, and also in other parts of Coahuila, but not on the intermontane valleys, are: Perognathus nelsoni Peromyscus eremicus Perognathus penicillatus Peromyscus melanophrys Dipodomys nelsoni Small mammals that occur on the Mesa del Norte in western Coahuila, and that have reached, undifferentiated, the high inter- montane valleys by way of the north or west, are: Sylvilagus audubonii Reithrodontomys megalotis Spermophilus spilosoma Peromyscus maniculatus Endemic Species Six species, Sorex milleri, Scalopus montanus, Scitirus alleni, Cyno- mys mexicanus, Dipodomys nelsoni, and Neotoma goldmani, either live only in Coahuila or occur there and in adjacent Mexican states. Findley (1955) suggests that Sorex milleri is a relict and that it is most closely related to the Sorex cinereus group. Perhaps Scalopus montanus, a relative of Scalopus aquaticus, also is a relict 152 University of Kansas Publs., Mus. Nat. Hist. species in the Sierra del Carmen. Sciurus alleni lives on oak-covered slopes of the Sierra Madre Oriental in Coahuila, Nuevo Leon and Tamaulipas. Probably this squirrel has been derived from an ancestral stock in western Mexico; alleni seems to be closely re- lated to Sciurus oculatus, found to the south, and to Sciurus natjari- tensis of western Zacatecas. Burt (1949) considers that Cynomys mexicanus is a relict species left when the northward shift of the faunas took place after the retreat of the last ice-sheet. Dipodomys nelsoni is smaller than, but in many ways is closely related to Dipodomys spectabilis that occurs in the western part of the Mesa del Norte and on the Mesa Central. Neotoma goldma7ii, a small woodrat, superficially re- sembles Neotoma lepida, but is more closely related to 'Neotoina alhigula (see Rainey and Baker, 1955). MAMMALOGICAL EXPLORATION IN COAHUILA Probably the first mammal saved for scientific study from Coa- huila was the cotton rat, which Baird examined and named Sig- modon berlandieri [= Sigmodon hispidus berlandieri] in 1855. This animal came from the vicinity of San Pedro de las Colonias along the Rio Nazas. In 1880, Dr. Edward Palmer obtained a small collection of mammals from the vicinities of Monclova, Sal- tillo, Parras, and San Pedro [^ San Pedro de las Colonias]. These specimens went to the Museum of Comparative Zoology and were reported upon by J. A. Allen (1881). From 1892 to 1906, E. W. Nelson and E. A. Goldman collected mammals in Mexico under the auspices of the United States Department of Agriculture. Col- lections in Coahuila were made in eight areas (see Goldman, 1951: 129-135), while visiting the State in December, 1893; July and August, 1896; and April and May, 1902. Their collections form the basis of much of our present knowledge of the kinds and dis- tribution of Coahuilan mammals. At least two other field employees of the Bureau of Biological Survey visited Coahuila. These were Clark P. Streator, who obtained mammals at La Ventura and Jaral, and Jas. H. Gaut, who collected along the Coahuilan side of the Rio Grande in the vicinity of Villa Acuiia in 1903. Goldman (1951:132) revisited Coahuila in 1926, obtaining small mammals at San Pedro [= San Pedro de las Colonias]. Ernest G. Marsh, Jr. (1937) reported on mammals observed while making a biologi- cal survey for the United States National Park Service in the Sierra Mammals of Coahuila, Mexico 153 del Carmen and other areas at least as far south as Cuidad Muz- quiz. Stanley P. Young, Tappan Gregory, and R. S. Stugis visited the Sierra del Carmen in September, 1937 (see Gregory, 1938). Included in the mammals collected and deposited in the United States National Museum were three new rodents of the genera Eutamias, Thomomys, and 'Neotoma, which were described by Goldman (1938a). Mammals obtained in southern Coahuila in 1937 by the Museum of Vertebrate Zoology of the University of California included a previously unknown subspecies of Vulpes macrotis, which was described by Benson (1938). In 1938, Ernest G. Marsh, Jr., of the University of Texas, studied mammals at several localities in the state. From June 12 through August 13, 1938, Gerald Young and I accompanied Marsh, making vertebrate collections in northeastern Coahuila from Villa Acuna to as far south as Ciudad Muzquiz. Mammals obtained are deposited in the Texas Memorial Museum. In 1938, George M. Sutton, Thomas D. Burleigh and John B. Semple took a few mammals for the Carnegie Museum while studying birds in southern Coahuila (see Burleigh and Lowery, 1942). Ted Dealey and Joseph MacDonald Dealey of Dallas, Texas, obtained for the United States National Museum, white-tailed deer at the Carmen Mountain Hunting Club in the Sierra del Carmen in 1939; these were described as Odocoileus virginiamis carminis by Goldman and Kellogg (1940). In April, 1941, Thomas D. Burleigh, George H. Lowery, Jr., Merriam L. Miles and Sam M. Ray collected a few mammals in southern Coa- huila for the Museum of Zoology at Louisiana State University. Frederick W. Miller of the Dallas Museum of Natural History secured mammals in the Sierra del Carmen in October and No- vember, 1940. From this collection, Jackson (1947) named as new the shrew, Sorex milleri. A part of Miller's collection is in the United States National Museimi; the remainder is in the Dallas Museum of Natural History. In the spring of 1945, Walter P. Tay- lor, Walter B. McDougall, Clifford C. Presnall, and Karl P. Schmidt (1945-1946) visited the Sierra del Carmen. Mammals obtained are deposited in the Texas Cooperative Wildlife Collection at A. & M. College of Texas. A. Starker Leopold, Alden H. Miller, and Ward C. Russell of the Museum of Vertebrate Zoology at the University of California spent March and April, 1953, in the Sierra del Carmen where they obtained mammals and other vertebrates. The University of Kansas Museum of Natural History began 154 University of Kansas Publs., Mus. Nat. Hist. field studies of mammals in Coahuila on July 8, 1949. Since that time 30 students and stafiF members in the interests of the Museum visited Coahuila to study vertebrates. These persons are: Albert A. Alcorn Gerd H. Heinrich Joseph Raymond Alcorn Delayne Hudspeth Sydney Anderson William McKee Lynn Rollin Harold Baker Jack M. Mohler James Sheldon Carey George M. Newton Peter Stanley Chrapliwy Roger O. Olmsted W. Kim Clark Robert Lewis Packard Robert William Dickennan Robert Julian Russell John R. Esther WiUiam J. Schaldach, Jr. James Smith Findley Harrison Bruce Tordoff Charles M. Fugler John Wesley Twente, Jr. John Keever Greer South Van Hoose, Jr. Eugene Raymond Hall Olin Lawrence Webb John WiUiam Hardy Robert Gravem Webb Oscar B. Heck J. Spencer Wolf CONSERVATION OF MAMMALS IN COAHUILA In Pre-Spanish times Indians depended for food on big game and other wildlife in Coahuila. These peoples probably lived in small groups near permanent water and took what animals they required for their economy. Gilmore ( 1947 ) identified more than 2000 pieces of mammalian bone from archeological cave-sites near Cuatro Ciene- gas. These bones, probably Pre-Spanish in age, included those of pronghorn, mountain sheep, mule deer, white-tailed deer, wapiti and bison. Bison and wapiti, never numerous in Coahuila, disap- peared probably prior to 1850. Historians write that the Indians went north of the Rio Grande to hunt the bison. Coahuila, under Spanish domination and even in the early years of the Mexican Republic, was not extensively colonized. Webb (1931:86-87) and other writers say that the Spaniards were not attracted to northern Mexico, because of its aridity and because the Plains Indians re- sisted strongly the efforts of Europeans to travel through or oc- cupy the area. The Spaniards did introduce cattle, and ranching proved to be more profitable than farming, because, without irri- gation, crops could not be grown on the arid plains. The settlers confined most in their activities to protected settle- ments until the Indian menace declined in the latter part of the nineteenth century. Following this time Coahuila was developed rapidly into a ranching country, and the game herds began to de- cline as a result of this increased occupation. Since the turn of the twentieth century, hunting has increased, and big game has Mammals of Coahuila, Mexico 155 had to compete with an always increasing number of domestic Hve- stock. Improved roads and landing strips for aircraft have assisted the sportsman to reach favored, out-of-the-way, game-range. The Sierra del Pino, Sierra del Carmen, Sierra de San Marcos, Sierra de Guadalupe and other mountains are well-known to hunters from as far away as Guadalajara, Mexico City and Dallas. Laws prohibiting the taking of game, except in prescribed open seasons, have been enacted, and in October, 1950, a Presidential Decree ordered closed seasons for all game in the State of Coahuila. The desert bighorn, always the prize take of the trophy hunter, has disappeared from all but a few isolated desert ranges. How- ever, competition with domestic goats and sheep probably has been as important as over-himting in causing the decline of the bighorn at many places in the State. The pronghorn also has been reduced to small herds in a few places (see Villa R., 1951:23). In 1952, camp providers for mining operations threatened the few prong- horn remaining in the vicinity of the Sierra de la Encantada. The grizzly bear, probably never common in Coahuila, disappeared before 1900. The black bear, still found in most mountain ranges, is regarded as a menace to livestock and is killed on sight on most ranchos. Javelina is hunted for its meat but is still common on some ranchos in eastern Coahuila. Mule deer, living in the accessible lowlands, is now rare in most places, whereas the mountain-dwelling white-tail deer occurs commonly in many of the more inaccessible montane retreats. Beaver, once numerous along most of the per- manent streams in northeastern Coahuila, is found only along the Rio Grande, at Presa Don Martin at the junction of the Rio Salado and the Rio Sabinas, and at a few places along the Rio de San Diego. The numbers of wolf, coyote, desert fox, gray fox, mountain lion, bobcat, and jaguar (never abundant there) have been reduced on many ranchos, especially those in northern Coahuila, through trap- ping and in the case of the canids by means of the cyanide gun. The endemic Mexican prairie dog is in jeopardy in parts of south- eastern Coahuila where it occurs on agricultural lands. Game laws have been difficult to enforce in Coahuila, especially in rural areas. At present the areas best for game are either on remote mountains or on the larger ranchos where conservation- minded owners have protected the animals from over-hunting. Deer, javelina, wild turkey and other animals are given protection on such ranchos as the Rancho La Mariposa and the Rancho Las Margaritas. Dr. Marcelino Garza-Pena and his family have pro- 156 University of Kansas Publs., Mus. Nat. Hist. tected the desert bighorn which live on their lands in the vicinity of San Lazaro and the javelina on their Rancho Guadalupe. The mule deer and javelina need protection from illegal hunting. The hunting of bighorn, pronghom and beaver should be prohibited by law. A combination of more rigid enforcement of the hunting regulations as well as greater concern by land holders for game on their holdings seems a necessity. Local sportsmen's organizations, such as the progressive one in Sabinas, are needed in every large community to encourage better conservation practices. In the middle 1930's, when the Big Bend National Park of Brewster County, Texas, was first being developed, the formation of the Big Bend International Peace Park to include the Big Bend and the Sierra del Carmen of northern Coahuila was visualized by conser- vationists of both countries. This highly important international project is still in the planning state. Our present, increased knowl- edge of the unusual biological features of the Carmens has made this proposal even more desirable. It is hoped that this Interna- tional Park, to conserve important wildlife species and to promote good-will, will become a reality. METHODS AND ACKNOWLEDGMENTS Mammals in Coahuila were collected with the aim of obtaining maximum information on the kinds present, their distributions, and their geographic varia- tions. As many ecological data as possible were gathered. At each locality visited every effort was made to get an adequate sample of the local mammalian fauna. Most parts of the State were visited by means of automobiles; isolated mountainous areas were reached either on foot or by means of horses and burros. The status of big game and other species was gleaned from reliable local residents whenever possible. In this report the arrangement of genera follows that of Miller and Kellogg (1955). Species and subspecies are arranged alphabetically under the respec- tive genera and species. Under each species account, remarks concerning its natural history are followed by a discussion of subspecies known from the State. Under each subspecies, the citation to the original description is given followed by the citation to the first usage of the currently used name-combina- tion. Measurements, unless otherwise specified, are of adults and are given in millimeters. When measurements of several individuals arc listed, usually only the average and extreme measurements are presented. Weights are in grams. Capitalized color terms are those of Ridgway, Color Standards and Color Nomenclature, Washington, D. C, 1912. Under each of the lists of specimens examined and additional records from the literature, the order of arrangement of the localities is from north to south. Those localities listed in Roman type are represented on the appropriate distri- bution map by blacked-in circles. Each of several circles covers two or more locahties because the localities are close together. In such instances the north- ernmost place is listed in Roman type and the names of the other places in Mammals of Coahuila, Mexico 157 Italic type. Specimens, reported on in the literature, but not seen, are marked on the maps of distribution as blacked-in triangles. Most specimens were taken at localities designated on attached labels as being a certain number of miles in a certain direction from a town or other mapped, geographic place. In the text, for the sake of brevity, some of these localities are referred to as either "near" or "in the vicinity of" a place. To determine the exact place of capture, the reader can consult the list of speci- mens examined that immediately follows the text under a given species or subspecies. Specimens examined are designated as to collections in which catalogued by the following abbreviations: BSC, the Biological Surveys Collection of the United States National Museum; DMNH, the Dallas Museum of Natural His- tory; CNHM, the Chicago Natural History Museiun; MCZ, the Museum of Comparative Zoology at Harvard College; TCWC, the Texas Cooperative Wild- life Collection at the Agricultural and Mechanical College of Texas. Those in charge of these collections are thanked for allowing me to examine specimens. Specimens listed without such abbreviations are in the Museum of Natural His- tory of the University of Kansas. First, I want to thank Ernest G. Marsh, Jr., for inviting me to join his field party in Coahuila in the summer of 1938 and affording me the chance to ac- quaint myself with some of the mammals of that State. Of those who accom- panied me on one or more collecting trips to Coahuila (spring, 1950; sp., 1952; summer, 1952; smn., 1953; sp., 1954; sum., 1955), I am especially indebted to George N. Newton, J. Ray Alcorn and John Keever Greer. Other persons from the University of Kansas who provided valuable cooperation and companionship in the several field undertakings are listed under the heading "Mammalogical Exploration in Coahuila." I owe special thanks to the many friends and hos- pitable residents of Coahuila who furthered our field eflForts in countless ways. Especially helpful was my good friend Roberto Spence of Sabinas who ren- dered many kind services to me and to John Keever Greer, Robert W. Dicker- man and other workers from the University of Kansas. Gracious cooperation and needed counsel were oflFered by Dr. Marcelino Garza-Peiia of Saltillo, David McKellar and Alden McKellar of the Hacienda La Mariposa, Oscar Cardenas of Saltillo, Luis Rannauro of the Hacienda El Tulillo, Alvaro Trevino of the Hacienda La Candalaria, Jesus M. Ramon-Cantu of Villa Acuria, Maxie Machaelis of BoquiUas, Juan Parras of the Hacienda Aqua Verde, Francisco Garcia of Piedras Negras, Professor Rafael Rodriquez-Velasco of Sacramento, Earl C. Johnston of Cuatro Cienegas, Tomas Cantu of Estanque de Norias, Ildefonso Verduzco of the Hacienda San Ildefonso, Juan Jose Zapata of the Hacienda Rincon, Urbano Santos of the Hacienda San Geronimo, Antonio Garza de la Fuente of Cuatro Cienegas and many others. The cooperation of the Direccion General Forestal y de Caza for kindly providing permits for the taking of mammals for scientific purposes in Coahuila is acknowledged. I am grateful to Professor Bernardo Villa-Ramierez of the Instituto de Biologia for his help and many suggestions. The advice and encouragement given me by Professor E. Raymond Hall has been a guiding force seeing this work to completion. I thank also the many others at the University of Kansas for their kind help and would single out Robert J. Russell, Howard J. Stains, J. Knox Jones, Jr., James S. Findley, Richard Van Gelder, Sydney Anderson, Keith R. Kelson, and Ohn L. Webb for their 2—1204 158 University of Kansas Publs., Mus. Nat. Hist. services on matters concerning mammals and Ronald L. McGregor for his de- termination of plants. I thank also my father, Charles Laurence Baker, for his advice concerning the geology of Coahuila. Figures were prepared by Justine Anderson and Victor Hogg. Assistance in financing field work is acknowledged from the Kansas University Endowment Association and the National Science Foundation, GAZETTEER OF LOCALITIES IN COAHUILA The following is a list of place-names in Coahuila used in re- cording the localities from which specimens of mammals have been taken. Each place-name is followed by its location in degrees and minutes of latitude and longitude. Place-names in Coahuila sometimes are vague, because their locations are not exactly at the same geographic position on every map examined. Most lo- calities included in this report occur on the maps of the American Geographical Society of New York, Provisional Edition, scale 1:1,000,000 as follows: No. N.H-13 (Chihuahua), No. N.G-13 (Culiacan), and No. N.G-14 (Monterrey). Part of northeastern Coahuila is mapped on the same scale on sheet No. N.H-13 (Austin) of the Geological Survey, United States Departinent of the In- terior. Also useful are the World Aeronautical Charts of the United States Coast and Geodetic Survey, scale 1:1,000,000, Nos. 469, 470, 521, and 522. The best map of Coaliuila is the Carta General del Estado de Coahuila de Zaragoza prepared by T. S. Abbott, 1905, scale 1:500,000. The reader is referred to Goldman (1951: 129- 135) for detailed descriptions of the localities visited by E. W. Nelson and E. A. Goldman. Acatita.— 26°26', 102°59'. EI Pino.— 28°31', 103° 14'. Acebuches.— 28°17', 102°56'. EI Tulillo.— 25°41', 101° 13'. Allende.— 28°21', 100°51'. Encantada.— 25°17', 101 °04'. Agua Nueva.— 25°12', 101°06'. Encarnacion.— 24°53', 101°03'. Americanos.— 27° 12', 103° 14'. Fortin.— 28°48', 101° 14'. Arteaga.— 25°27', 100°52'. Fuentes.— 28°39', 100°33'. Australia.— 26° 18', 102° 18'. General Cepeda.— 25°22', 101° 14'. Bella Union.— 25°26', 100°51'. Gomez Farias.— 24°58', 101 °02'. Boquillas.— 29°ir, 102°55'. Guadalupe.— 26° 12', 101°21'. Carneros.— 25°07', 101 °06'. Guerrero.— 28° 20', 100°24'. Carricitos.— 28°03', 100°27'. Hechicero.— 28°36', 103°38'. Castillon.— 28°21', 103°33'. Hermanas.- 27°13', 101° 13'. Chorro.— 25°25', 100°12'. Hipulito.— 25°43', 101°25'. Coyote.— 27° 16', 102° 16'. Hisachalo or Huisachalo.— 26°45', Cuatro Cienegas.— 26°58', 102°04'. 101 °04'. Diamante— 25°54', 100°22' La Babia.— 28°33', 102°03'. Don Martin.— 27°32', 100°37'. La Candelaria.— 28°20', 100°20'. EI Hundido.- 26°37', 102° 18'. La Flor.— 25°06', 103° 17'. Mammals of Coahuila, Mexico 159 La Gacha.— 28°09', 101°31'. La Mariposa.— 28° 12', 101 °59'. La Pastora.— 25°39', 101° 39'. La Rosa.— 25°33', 101° 22'. La Ventura.— 24°48', 100°54', Las Delicias.— 26°10', 102°49'. Las Margaritas.— 28°42', 101° 14'. Jaco.— 27°50', 104°00'. Jaral or San Antonio de Jaral. — 25°41', 101°23'. Jimenez.— 29°04', 100°42'. Jimulco.— 25°07', 103° 18'. Monclova.— 26°54', 101 °25'. Mesa de Tablas.— 25°19', 100°24'. Muralla.- 26°19', 101 °22'. Muzquiz.— 27°53', 101°32'. Nadadores.— 27°03', 101°36'. Nava.— 28°25', 100°46'. Nueva Rosita or Rosita. — 27° 58', 101°18'. Nuevo Mundo.— 25°57', 103°09'. Ocampo.— 27°22', 102°26'. Panuco.— 26°43', 101 °00'. Parras.— 25°25', 102° 12'. Piedra Blanca.— 29°02', 102°26'. Piedras Negras.— 28°43', 100°32'. Sabinas.— 27°51', 101 °07'. Saltillo.— 25°26', 101 °00'. San Antonio de las Alazanas. — 25° 16', 100°37'. San Buenaventura.— 27°06', 10r32'. San Geronimo.— 28°30', 101° 12'. San Juan de Sabinas. — 27°55', 101°17'. San Juan Neponuceno. — 24° 42', 100°58'. San Lazaro.— 26°18', 101 °23'. San Lorenzo.— 25° 28', 102° 12'. San Marcos.— 26°41', 102°07'. San Miguel, a hacienda. — 29° 14', 101°22'. San Miguel, a railroad station. — 25°00', 100°59'. San Pedro de las Colonias. — 25°45', 102°58'. Santa Cruz.— 25°47', 101 °06'. Santa Teresa.— 26° 27', 101°21'. Sierra Mojada.— 27°17', 103°42'. Tanque Alvarez.— 27° 56', 102°38'. Torreon.— 25°33', 103°27'. Trevino.— 26°02', 101° 13', Villa Acufia.— 29°19', 100°56'. Mountain ranges have been used as place-names on labels of some specimens. Their latitudes and longitudes, given as follows, are listed using the approximate center of the highland of each range as the point of reference. Pico de Jimulco.— 25°08', 103° 16'. Serranias del Burro. — 29°00', 101 °48'. Sierra del Carmen.— 29°00', 102°30'. Sierra de la Encantada. — 28° 25', 102°30'. Sierra Encamacion. — 24°48', 101°17'. Sierra de Guadalupe. — 25° 13', 101°32'. Sierra de la Paila.— 26°00', 101°45'. Sierra Palo Verde.— 26° 18', 101°23'. Sierra del Pino.— 28° 15', 103°03'. Sierra de la Madera. — 27°03', 102°30'. Sierra de San Marcos. — 26° 15', 101°53'. Sierra Madre Oriental. — 25° 15', 100°25'. 160 University of Kansas Publs., Mus. Nat, Hist. CHECK-LIST OF THE MAMMALS OF COAHUILA The 144 kinds of native mammals of 103 full species found in Coahuila belong to 60 genera of 24 families of 8 orders. In the following list, mammals introduced by man are marked with an * and those thought to be extinct in Coahuila are marked with a f. Class MAMMALIA — mammals Order MARSUPIALIA— marsupials Family Didelphidae — opossums Didelphis marsupialis californica Bennett opossum 166 Didelphis marsupialis texensis J. A. Allen opossum 166 Order INSECTIVORA— insectivores Family Soricedae — shrews Sorex milleri Jackson Miller shrew 167 Sorex saussurei saussurei Merriam Saussure shrew 168 Notiosorex crawfordi crawfordi (Coues) desert shrew 169 Family Talpidae — Moles Scalopus montanus Baker Coahuilan mole 170 Order CHIROPTERA— bats Family PHYLLOSTOMmAE — leaf-nosed bats Aello megalophylla senicula ( Rehn ) lappett-chinned bat 171 Choeronycteris mexicana Tschudi hog-nosed bat 172 Leptonycteris nivalis nivalis ( Saussure ) long-nosed bat 174 Family Vespertilionidae — common bats Myotis californicus calif ornicus (Audubon and Bachman ) California myotis 175 Myotis evotis auriculus Baker and Stains long-eared myotis 176 Myotis planiceps Baker flat-headed myotis 177 Myotis velifer incautus (J. A. Allen) cave myotis 177 Myotis volans interior Miller long-legged myotis 179 Myotis yumanensis yumanensis i H. Allen ) Yuma myotis 179 Pipistrellus hesperus maximus Hatfield western pipistrelle 180 Pipistrellus subfiavus clartis Baker eastern pipistrelle 181 Eptesicus fuscus pallidus Young big brown bat 182 Lasiurus horealis borealis (Miiller) red bat 184 Lasiurus cinereus cinereus ( Beauvois ) hoary bat 185 Dasyptenis ega xanthinus Thomas western yellow bat 185 Nycticeius humeralis mexicanus Davis evening bat 186 Corynurliinus totvnsendii australis Handley lump-nosed bat 187 Antrozous pallidus pallidus (Le Conte) pallid bat 188 Mammals of Coahuila, Mexico 161 Family Molossidae — free-tailed bats Tadarida brasiliensis mexicana ( Saussure ) Eumops perotis californicus ( Merriam ) common free-tailed bat western mastiff bat . . . . PAGE 190 191 Order EDENTATA — sloths, anteaters and armadillos Family Dasypodedae — armadillo Dasypus novemcinctus mexicanus Peters nine-banded armadillo . Order LAGOMORPHA— hares, rabbits and pikas P'amily Leporidae — hares and rabbits Lepus californicus asellus Miller Lepus californicus merriami Meams Lepus californicus texianus Waterhouse Sylvilagus audubonii minor (Meams) Sylvilagus audubonii parvulus (J. A. AUen) Sylvilagus floridanus chapmani (J. A. Allen) Sylvilagus floridanus nelsoni Baker Sylvilagus floridanus orizabae ( Merriam ) 193 black-tailed jackrabbit . . 194 black-tailed jackrabbit . . . 195 black-tailed jackrabbit . . . 196 Audubon cottontail 197 Audubon cottontail 198 eastern cottontail 200 eastern cottontail 201 eastern cottontail 201 Order RODENTIA— rodents Family Sciuridae — squirrels Cynomys mexicanus Merriam Spermophilus mexicanus parvidens Meams Spermophilus spilosoma pallescens (A. H. Howell) Spermophilus variegatus couchii Baird Ammospermophilus interpres ( Merriam ) Eutamias bulleri solivagus A. H. Howell Eutamias dorsalis carminis Goldman Sciurus alleni Nelson Sciurus niger limitis Baird Family Geomyidae — pocket gophers Thomomys bottae analogus Goldman Thomomys bottae angustidens Baker Thomomys bottae humilis Baker Thomomys bottae perditus Merriam Thomomys bottae retractus Baker Thomomys bottae sturgisi Goldman Thomomys bottae villai Baker Thomomys umhrinus goldmani Merriam Cratogeomys castanops bullatus Russell and Baker Cratogeomys castanops consitus Nelson and Goldman Mexican prairie dog 202 Mexican ground squirrel . . 204 spotted ground squirrel . . 205 rock squirrel 207 Texas antelope squirrel . . 209 BuUer chipmunk 210 cliff chipmunk 212 Allen squirrel 213 fox squirrel . . 215 Botta pocket gopher 218 Botta pocket gopher 218 Botta pocket gopher 219 Botta pocket gopher 219 Botta pocket gopher 220 Botta pocket gopher 221 Botta pocket gopher 221 pigmy pocket gopher .... 222 plateau pocket gopher . . . 224 plateau pocket gopher . . . 225 162 University of Kansas Publs., Mus. Nat. Hist. Cratogeomys castanops convexus Nelson and Goldman Cratogeomys castanops excelsus Nelson and Goldman Cratogeomys castanops goldmani Merriam Cratogeomys castanops jucundus Russell and Baker Cratogeomys castanops planifrons Nelson and Goldman Cratogeomys castanops sordidulus Russell and Baker Cratogeomys castanops subnubilus Nelson and Goldman Cratogeomys castanops subsimus Nelson and Goldman Cratogeomys castanops ustulatus Russell and Baker plateau pocket gopher plateau pocket gopher plateau pocket gopher plateau pocket gopher plateau pocket gopher plateau pocket gopher plateau pocket gopher plateau pocket gopher plateau pocket gopher Family Heteromyidae — pocket mice Perognathus flavus medius Baker Perognathus -flavus pallescens Baker Perognathus hispidus hispidus Baird Perognathus hispidus zacatecae Osgood Perognathus merriami gilvus Osgood Perognathus merriami merriami J. A. Allen Perognathus nelsoni canescens Merriam Perognathus nelsoni nelsoni Merriam Perognathus penicillatus eremicus Meams Dipodomys merriami atronasus Merriam Dipodomys merriami merriami Meams Dipodomys nelsoni Merriam Dipodomys ordii durranti Setzer Dipodomys ordii idoneus Setzer Family Castoridae — beavers Castor canadensis mexicanus Bailey and kangaroo rats silky pocket mouse . . . silky pocket mouse . . . hispid pocket mouse . . hispid pocket mouse . . Merriam pocket mouse Merriam pocket mouse Nelson pocket mouse Nelson pocket mouse desert pocket mouse . Merriam kangaroo rat Merriam kangaroo rat Nelson kangaroo rat . Ord kangaroo rat . . . Ord kangaroo rat . . . PAGE 226 226 227 227 228 228 229 230 230 232 232 233 234 236 236 237 238 239 242 242 244 247 248 beaver 248 Family Cricetidae — native rats and mice Reithrodontomys fulvescens canus Benson Reithrodontomys fulvescens intermedius J. A. Allen Reithrodontomys megalotis megalotis (Baird) Reithrodontomys megalotis saturatus J. A. Allen and Chapman Peromyscus boylii levipes Merriam Peromyscus boylii rowleyi J. A. Allen Peromyscus difficilis diffxcilis (J. A. Allen) Peromyscus eremicus eremicus (Baird) fulvous harvest mouse . . 251 fulvous harvest mouse . . 251 western harvest mouse . . 253 western harvest mouse . . 254 brush mouse . 255 brush mouse . 255 big-eared rock mouse . . . 256 cactus mouse . 258 Mammals of Coahuila, Mexico 163 PAGE Peromyscus eremicus phaeurus Osgood cactus mouse 260 Feromtiscus leucopus texanus (Woodhouse) woods mouse 261 Feromijscus leucopus tornillo Mearns woods mouse 262 Peromyscus maniculatus blandus Osgood deer mouse 263 Peromyscus melanophrys coahuiliensis Baker plateau mouse 264 Peromyscus melmiotis J. A. Allen and Chapman black-eared woods mouse . 265 Peromyscus nasutus nasutus (J. A. Allen) rock mouse 267 Peromyscus pectoralis eremicoides Osgood white-ankled mouse 269 Peromyscus pectoralis laceianus Bailey white-ankled mouse 270 Peromyscus truei gentilis Osgood piiion mouse 271 Baiomys taylori taylori (Thomas) Tayloi pigmy mouse 273 Onychomys leucogaster longipes Merriam northern grasshopper mouse 274 Onychomys torridus surrufus HoUister southern grasshopper mouse 275 Onychomys torridus torridus (Coues) southern grasshopper mouse 276 Sigmodon hispidus herlandieri Baird hispid cotton rat 276 Sigmodon ochrngnathus ochrognathus Bailey yellow-nosed cotton rat . . 278 Neotoma albigula alhigula Hartley white-throated wood rat 280 Neotoma albigula leucodon Merriam white-throated wood rat . 281 Neotoma goldmani Merriam pigmy wood rat 282 Neotoma mexicana inornata Goldman Mexican wood rat 284 Neotoma mexicana navus Merriam Mexican wood rat 284 Neotoma micropus micropus Baird gray wood rat 285 Microtus mexicanus subsimus Goldman Mexican vole 287 Family Muridae — introduced rats and mice Rattus norvegicus norvegicus ( Berkenhout ) * Norway rat 289 Raftus rattus alexandrinus (fi. Geoffroy- Saint-Hilaire ) * black-rat 289 Mus musculus brevirostris Waterhouse * house mouse 290 Family Erethizontidae — porcupines Erethizon dorsatum couesi Mearns porcupine 290 Order CARNIVORA— carnivores Family Canidae — wolves, coyotes, and foxes Canis latrans mearnsi Merriam coyote 292 Canis latrans texensis Bailey coyote 293 Canis lupus baileyi Nelson and Goldman gray wolf 294 Vulpes macTotis zinseri Benson kit fox 295 Urocyon cinereoargenteus scottii Mearns gray fox 296 Family Ursidae — bears Ursus americanus eremicus Merriam black bear 298 Ursus sp.f grizzly bear 300 164 Umversity of Kansas Publs., Mus. Nat. Hist. Family Procyoxidae — raccoons and allies page Bassariscus astutus flavus Rhoads ring-tailed cat 301 Procyon lotor fuscipes Meams raccoon 302 Nasua narica tamaulipensis Goldman coati 304 Family MusTELroAE — weasels, skunks and allies Mustela frenata frenata Lichtenstein long-tailed weasel 304 Taxidea taxus berlandieri Baird badger 305 Spilogale gracilis gracilis Merriam spotted skunk 307 Mephitis macroura milleri Mearns hooded skunk 308 Mephitis mephitis varians Gray striped skunk 309 Conepatus mesoleucus mearnsi Merriam hog-nosed skunk 310 Family Felidae — cats Felis concolor stanleyana Goldman mountain lion 311 Felis onca veraecrucis Nelson and Goldman jaguar 313 Felis pardalis albescens Pucheran ocelot 313 Felis yagouaroundi cacomitli Berlandier jaguarundi 314 Lynx rufus baileyi Merriam bobcat 315 Lynx rufus texensis J. A. Allen bobcat 315 Order ARTIODACTYLA— even-toed ungulates Family Tayassuidae — peccaries Pecari tajacu angulatus (Cope) coUared peccary 316 Family Cervidae — deer, wapiti, and allies Cervus merriami Nelson f wapiti 318 Odocoileus hemionus crooki (Meams) mule deer 318 Odocoileus virginianns carminis Goldman and Kellogg white-tailed deer 322 Odocoileus virginianns miquihuanensis Goldman and Kellogg white-tailed deer 323 Odocoileus virginianns texanus ( Meams ) white-tailed deer 323 Family ANTiLOCAPRroAE — pronghorn Antilocapra americana mexicana Merriam pronghom 324 Family Bovidae — cows, bison, and allies Bison bison bison (Lirmaeus)f bison 325 Ovis canadensis mexicana Merriam bighorn sheep 327 Mammals of Coahuela, Mexico 165 ACCOUNTS OF SPECIES AND SUBSPECIES Didelphis marsupialis Opossum The opossum occurs along permanent streams and in some irri- gated areas in eastern and southwestern Coahuila. Normally this species is confined to narrow strips of riparian vegetation and seem- ingly rarely moves into the nearby desert plains or arid foothills. Opossums follow along streams onto the uplands and occasionally are found near the headwaters of the watershed. The presence of the opossum in the Sierra del Carmen (see Marsh, 1937:25) prob- ably is the result of movements of individuals upstream along arroyos from the Rio Grande. The opossum is common in northeastern Coahuila along the Rio de San Diego, the Rio Sa- binas, the Rio Salado and along the Rio Grande downstream from Villa Acuna. At Fuentes, I saw tracks and examined two young opossums in June 1938. At the Hacienda La Candelaria, east of Nava, Seiior Alvaro Tre- vino regards the species as com- mon not only along the Rio Grande but in the live oak brush, which forms extensive thickets on his rancho. Along the Rio Grande, upstream from Villa Acuiia, the opossum becomes progressively less common as streamside vegetation becomes restricted by narrow canons and rimrock. At the Hacienda Aqua Verde, approximately 80 miles (airline) upstream from Villa Acuna, according to Seiior Juan Parras the opossum is present but uncommon. In 1952, no opossum tracks were seen along the Rio Grande bordering this rancho. At Boquillas, local residents con- sider this animal to be rare; Borell and Bryant (1942:6) obtained few records from the north side of the Rio Grande in the Big Bend area of Texas. In 1952, Senor Diego Rubio, who lives ten miles Fig. 5. Distribution of the opossum in Coahuila. 1. Didelphis marsupialis californica. 2. Didelphis marsupi- alis texensis. 166 UNrv'ERSiTY OF Kansas Publs., Mus. Nat. Hist. south of Boquillas, told me that the opossum is found occasionally along the Arroyo de los Alamos. At Presa Don Martin, the opos- sum is well known. Above this Presa, along the Rio Sabinas, the opossum occurs as far upstream as Nacimiento. Senor D. M. Mc- Kellar of the Hacienda La Mariposa reports that this animal is oc- casionally seen on his ranch, which contains arroyos leading into the Rio Sabinas. Above the Presa Don Martin, along the Rio Salado, the opossum occurs upstream at least as far as Cuatro Cienegas, according to Sefior Tomas Cantu of Estanque de Norias. Between Monclova and Saltillo the arid plains and mountains lack permanent streams and probably constitute an effective bar- rier to the opossum. Opossums in southwestern Coahuila are sepa- rated from those in southeastern Coahuila by unsuitable habitat. Don Luis Rannauro of the Hacienda El Tulillo in 1950 informed me that the opossum is unknown at his rancho and adjacent areas; his rancho is along the road from Saltillo to San Lorenzo. The opossum, locally called "tlacuache," is hunted for both its meat and its fur, but since the animal is uncommon in most places, it is unimportant to the economy of the people. Didelphis marsupialis califomica Bennett Didelphis Califomica Bennett, Proc. Zool. Soc. London, p. 40, 1833, type from Sonora. Didelphis marsupialis califomica Hershkovitz, Fieldiana — Zool., Chicago Nat. Hist. Mus., 31 (47):548, July 10, 1951. Distribution. — Southwestern Coahuila, east at least as far as San Lorenzo (see fig. 5). Remarks. — Skulls only (one broken) of two subadults were ob- tained from road-kills in southeastern Coahuila. They have been compared with specimens from Jalisco and Distrito Federal and are referable to califomica because of the shape of the nasal bones, especially the truncated appearance of the posterior ends. Allen (1881:189) records two half-grown specimens taken by Dr. Ed- ward Palmer at Parras as Didelphys. Although not seen I refer them to califomica on geographic grounds. Specimens examined. — Total, 2, from: 1 mi. SW San Pedro de las Colonias 3700 ft., 1; 1 mi. N San Lorenzo, 4200 ft., 1. Additional record (Allen, 1881:189): Parras. Didelphis marsupialis texensis J. A. Allen Didelphis marsupialis texensis J. A. Allen, Bull. Amer. Mus. Nat. Hist., 14: 172, June 15, 1901, type from Brownsville, Cameron County, Texas. Distribution. — Nortliem and eastern Coahuila (see fig. 5). Remarks. — Two subadults examined include one skin with par- Mammals of Coahuila, Mexico 167 tial skull and one partial skull only, both from road-kills. Based on the shape of the nasals ( see Allen, 1901 : 172 ) and comparison with two topotypes of texensis, these two specimens are referable to texensis. The partial skull from Monclova has nasals less pointed posteriorly than those of the specimen from one-half mile south of Sabinas. External measurements of these specimens (the one from Monclova first) are: total length, 730, 732; length of tail, 325, 340; length of hind foot, 57, 62; height of ear from notch, 45, 52. Opos- sums found in the vicinity of Saltillo, although not seen, presumably are referable to texensis, because animals living there probably reached southeastern Coahuila from the east by way of the Rio Salinas. Specimens examined. — Total, 2, from: /2 mi. S Sabinas, 1000 ft., 1; Mon- clova, 2000 ft., 1. Sorex milleri Jackson Miller Shrew Sorex milleri Jackson, Proc. Biol. Soc. Washington, 60:131, October 9, 1947, type from Madera Camp, 8000 ft.. Sierra del Carmen, Coahuila. Distribution. — Montane mesic forests of the Sierra del Carmen in northern Coahuila and the Sierra Madre Oriental in southeastern Coahuila (see fig. 6). Remarks. — Three specimens, obtained by Frederick W. Miller, are known from the Sierra del Carmen. In these mountains this species seems restricted to the Montane Mesic Forest Vegetation- type; no specimens were found by my associates or myself in the Montane Low Forest Vegetation-type in the Carmens at approxi- mately 5000 feet in elevation. Twenty-three shrews were captured at 9350 feet in elevation in an association of Douglas fir, pine, and aspen on July 6-8, 1955, in the Sierra Madre Oriental (13 miles east of San Antonio de las Alazanas ) . Most of these animals were caught between fallen logs in runs on pine needles and other ground litter in moist shaded places on wooded slopes. Peromyscus melanotis was caught also in these runs. Two Peromyscus melanotis and three Sorex milleri were captured in a 24 hour period in two museum special traps that were set within three feet of one another along a runway and baited with chewed rolled oats. The two Peromyscus were taken first and the three Sorex were taken at subsequent inspections of the traps. One of the Sorex was captured sometime between 8 a. m. and 11 a. m. and another sometime between 11 a. m. and 5 p. m. Four Sorex were trapped along with Microtus mexicanus in runways in short grass at the edge of a clearing. In April, 1954, the same area was trapped, but no Sorex was obtained and fewer Peromyscus 168 University of Kansas Publs., Mus. Nat. Hist. melanotis and Microtus mexicanus were taken. Weights of milleri from 13 miles east of San Antonio de las Alazanas were: for 11 males, 4.1 (2.9-4.8); for 8 non-pregnant females, 3.5 (3.3-3.8). One lactating female, taken on July 7, weighed 5.1 grams. Jackson (1947:132) classified this shrew as a small representative of the "vagrans-obscurus" group, but Findley (1955:617) related it more closely to Sorex cinereus Kerr. Findley considers milleri to be a relict population of Sorex cinereus isolated in the mountains of northeastern Mexico; he (loc. cit.) records milleri also on Cerro Potosi in Nuevo Leon. Findley (1953:635) reported also the ocur- rence of Sorex cinereus in Pleistocene deposits from San Josecito Cave in Nuevo Leon. Measurements. — The type specimen (thought to be a female and measured by Jackson, op. cit.ilSl) and three females from 13 miles east of San Antonio de las Alazanas measure: total length, 95, 97, 89, 93; length of tail vertebrae, 44, 42, 37, 41; length of hind foot, 11, 11, 11.5, 12; condylobasal length (of skull), 15.5, 16.1, 15.5, 15.8; palatal length, 6.2, 6.2, 6.2, 6.4; breadth of cranium, 7.4, 7.7, 7.6, 7.9; least interorbital constriction, 3.7, 3.1, 3.1, 3.3; maxillary breadth, 4.4, 4.1, 4.1, 4.3; maxillary tooth-row, 5.5, 5.9, 5.6, 5.8. Specimens examined. — Total, 24, from: Madero Camp, Sierra del Carmen, 1 (DMNH); 13 mi. E San Antonio de las Alazanas. 9350 ft., 23. Sorex saussiirei saussurei Merriam Saussure Shrew Sorex saussurei Merriam, Proc. Biol. Soc. Washington, 7:173, Sep- tember 29, 1892, type from north slope of El Nevada de Colima, about 8000 ft., Jalisco. Distribution. — High mountains of southeastern Coahuila (see fig. 6). Remarks. — Two Saussure shrews were obtained in the Si- erra de Guadalupe by E. W. Nelson and E. A. Goldman. This species probably lives within the coniferous belt above 9000 feet elevation (see Goldman, 1951: 134). The Sierra de Guadalupe is connected, by a ridge dipping to approximately 6800 feet in elevation, with the Sierra Madre Oriental to the eastward; this ridge may have been used as a Fig. 6. Distribution of shrews in Coa- hiula. 1. Sorex milleri. 2. Sorex saus- surei. Mammals of Coahuila, Mexico 169 route by this shrew to reach the Sierra de Guadalupe. Shrews probably occur also in the higher parts of the Sierra Madre Oriental in extreme southeastern Coahuila, where, above 9000 feet elevation, Sorex milleri, Eutamias hulleri and Microtus mexicanus occur. This subspecies is on most of the higher mountains of Central Mex- ico, and northward on the eastern side of the Central Plateau to southeastern Coahuila. Jackson (1928:155-156) writes that there is some "slight local variation" in shrews referred to this subspecies but infers that these variations are not of sufficient magnitude to merit recognition as different subspecies. External measurements of two males from the Sierra de Guadalupe are: total length, 106, 110; length of tail, 37, 42; length of hind foot, 13, 13.5. Specimens examined. — Total, 2, from Sierra de Guadalupe (BSC). Notiosorex crawfordi crawfordi (Coues) Desert Shrew Sorex (Notiosorex) crawfordi Coues, Bull. U. S. Geol. and Geogr. Surv. Terr., 3:651, May 15, 1877, type from near old Fort Bliss, El Paso County, Texas. Notiosorex crawfordi Merriam, N. Amer. Fauna, 10:32, December 31, 1895. Distribution. — Desert plains of Coahuila. Remarks. — The desert shrew probably occurs in suitable places throughout the Chihuahuan Desert Shrub Vegetation-type. Field parties made special effort to obtain this mammal by trapping in various places, by demolishing woodrat houses, and by breaking apart dead stumps of Agave, sotol and other plants, using methods suggested by Hoffmeister and Goodpaster (1954:47) and others. No specimens were obtained by these methods, but parts of two skulls were found in owl pellets taken in a cave three miles north- west of Cuatro Cienegas (see Baker, 1953b: 253). This cave was situated approximately 100 feet up the west side of a narrow caiion cut by a spring-fed branch of the Rio Salado. Riparian growth flourished along the stream, in a few places where grazing was not severe; the area otherwise consisted of arid hills and desert plains. One skull of Notiosorex was in a pellet containing otherwise only remains of the pocket mice, Perognathus flavus and Perogna- thus penicillatus. Since these two mice hve in deserts, possibly the owl caught the shrew in such a place. The two precranial parts of skulls from pellets are assigned to N. c. crawfordi chiefly on geographic grounds, and measure: palatal length, 6.8, 7.2; alveolar length of maxillary tooth-row, 5.9, 5.9; least interorbital constriction, 4.2, 4.1; maxillary breadth, 4.6, 4.9. Specimens examined. — Two from 3 mi. NW Cuatro Cienegas. 170 University of Kansas Publs., Mus. Nat. Hist. Scalopus montanus Baker Coahuilan Mole Scalopus montanus Baker, Univ. Kansas Publ., Mus. Nat. Hist., 5:19, Feb- ruary 28, 1951, type from Club Sierra del Carmen, 2 mi. N and 6 mi. W Piedra Blanca, Coahuila. Distribution. — Sierra del Carmen of northern Coahuila (see fig. 7). Remarks. — This species is known from one specimen taken at 4950 feet elevation in the Montane Low Forest Vegetation-type of the Sierra del Carmen in northern Coahuila. It was caught by a Macabee pocket gopher trap in an underground runway, in which was trapped the next day, a pocket gopher, Thomomys bottae sturgisi. At that time no "raised" tunnels or other sign usually made by members of the genus Scalopus were found in heavily grazed parklike stands of oaks. Moles have been sought in vain in the Sierra de la Encantada, Serranias del Burro, Sierra del Pino, and Sierra de los Hecheros. Pocket gophers of tlie genus Thomomys have been taken in the oak belt in situations comparable to those in the Sierra del Carmen in most of these mountains. Moles have been reported by reliable local residents in the low- lands of northeastern Coahuila, at Sabinas and at the Hacienda La Mariposa. Mr. D. S. Mc- Kellar told me that prior to 1952 a mole, called "topo" by the resi- dents, was brought to the hacienda. Another was seen, ac- cording to McKellar, in Sabinas in 1952; a child had the animal alive and held by a string tied on one leg. Residents in the Sierra del Carmen, when showed the mole taken there, had not previously seen the creature and did not know a Mexican name for it. This mole is closely related to Scalopus aquaticus and the adult male measures: total length, 150; length of tail vertebrae, 27; length of hind foot, 20; greatest length of skull, 32.2; palatilar ^ig. 7. Distribution of the Coahuilan , 1 T r. n .111 11 mole, hcalopus montanus. length, 13.8; mastoidal breadth, 16.2; interorbital breadth, 7.0; alveolar length of maxillary tooth- Mammals of Coahuila, Mexico 171 row, 11.3; greatest depth of skull including auditory bullae, 9.4; weight, 38.6 grams. Specimen examined. — One, from the type locality. Aello megalophylla senicula (Rehn) Lappet-chinned Bat Mormoops megalophylla senicula Rehn, Proc. Acad. Nat. Sci. Philadelphia, 54:169, June 11, 1902, type from Fort Clark, Kinney County, Texas. Aello megalophylla senicula J. A. Allen, Bull. Amer. Mus. Nat. Hist., 22:261, July 25, 1906. Distribution. — Northeastern Coahuila (see fig. 8). Remarks. — In Coahuila, the lappet-chinned bat seems to occur only in the northeastern part and probably reached this area by way of the Gulf Coastal Plain. There are no records of the species in the drier, western and southwestern parts of the State. Likewise, in San Luis Potosi, Dalquest (1953:45) found this bat on the Coastal Plain and not on the arid Central Plateau. In Coahuila, specimens were obtained in two widely separated, limestone caves and also in mist nets stretched over water at two localities in the Serranias del Burro. One colony of approximately 100 bats of this species was found on March 24 in a dry cave, which opens half-way up the side of an arid limestone bluflF bordering the north side of Puerta Santana. The cave has a narrow entrance opening into a large room, the ceiling of which was estimated to be 75 feet above the guano- covered floor. The bats clung to the north wall in a compact group; most of them began to fly when disturbed by our approach. Eleven specimens, including examples of both sexes, were ob- tained; no other kinds of bats were present in the cave. Robert W. Dickerman found a colony of more than 500 Aello in a large limestone cave east of Hermanas on December 7; several were obtained by throwing sticks into clusters of bats clinging to the high ceiling. These bats were much less active than those ob- served at the cave in Puerta Santana in March. Dickerman also recorded Tadarida brasiliensis from the cave east of Hermanas, although this species was found apart from Aello in the cave. Aello was taken in mist nets stretched over water at Fortin and at the Hacienda Las Margaritas, both places in the foothills of the Serranias del Burro, on March 29, May 6, September 25 and 30, and October 9, Perhaps these bats fly less or not at all in late autumn and winter, because none was taken by John Keever Greer in 1953-54, who in these seasons maintained mist nets stretched over the water in an open square, concrete water tank at the Hacienda Las Margaritas. It may be assumed that Aello, 172 Unr'ersity of Kansas Publs., Mus. Nat. Hist. at least in Coahuila, remains inactive in caves or other dark re- treats during the months from November at least through February or early March. Average and extreme weights of 8 adult males from Puerta San- tana and near Hermanas w^ere: 15.0 (13.4-17.0); of 5 non-pregnant females, 14.9 (13.0-16.8). Four of eight females taken on March 24 were pregnant, each contain- ing one embryo. Another preg- nant female, containing one embryo, was secured on May 6. Coahuilan specimens are as- signed to senicula on the basis of the large size of the internal lobe of the second upper pre- molar tooth (see Rehn, 1902:162 and 169). Individuals vary in color of upper parts from cinna- mon-brown to slate-gray. In the cave east of Hermanas, Dicker- man reports that gray bats out- numbered brown ones. Measurements. — Average and ex- treme measurements of 8 adult males from northeastern Coahuila (KU 44- 702, 44703, 44706, 56471, 56578, 56579, 56582, 56583) and 10 adult females (KU 44697-44701, 44704, 44705, 44707, 44708, 58675), respectively, are: total length, 92 (87-100), 92 (88-95); length of tail vertebrae, 25 (24-27), 26 (24-27); length of hind foot, 11 (9-12), 11 (9-12); height of ear from notch, 15 (14-16), 16 (15-17); length of forearm, 54.2 (52.7-56.8), 54.9 (52.3-56.8); condylobasal length of skull, 15.0 (14.8-15.2), 15.0 (14.8-15.3); zygomatic breadth, 9.6 (9.4-9.9), 9.5 (9.4-9.8); least interorbital constriction, 5.2 (5.1-5.3), 5.2 (5.1-5.4); palatal length, 8.9 (8.5-9.1), 8.9 (8.8-9.1); mastoidal breadth, 8.4 (8.3-8.6), 8.4 (8.3-8.6); breadth of braincase, 9.0 (8.9-9.2), 9.0 (8.9-9.1); breadth across M3, 6.8 (6.7-7.0), 6.8 (6.7-7.0); alveolar length of maxillary tooth-row, 7.8 (7.7-8.0), 7.8 (7.7-8.0). Specimens examined. — Total, 37, from: Fortin, 3300 ft., Rancho Las Mar- garitas, 1; Hacienda Las Margaritas, 2800 ft., 6; Puerta Santana, 2900 ft., 6 mi. S and 3 mi. W Hacienda La Mariposa, 11; 6 mi. E Hermanas, 1200 ft., 19. Choeronycteris mexicana Tschudi Hog-nosed Bat Choeronycteris mexicana Tschudi, Untersuchungen uber die Fauna Peruana . . ., p. 72, 1844, type from Mexico. Distribution. — Southern Coahuila (see fig. 9). Fig. 8. Distribution of the lappet- chinned bat, A'ello megalophylla senic- ula, in Coahuila. Mammals of Coahuila, Mexico 173 Remarks. — The hog-nosed bat was taken in southern Coahuila in natural caves at five localities and in one abandoned mine tunnel; none was snared in mist nets or shot over water at dusk. HofiF- meister and Goodpaster (1954:52), in Arizona, took these bats only in dark retreats. One cave, immediately below the summit of the south face of the canon of the Rio Salado near San Buena- ventura, Coahuila, contained approximately 200 bats of the five species, Eptesicus fuscus, Corynorhinus townsendii, Myotis velifer, Tadarida hrasiliensis, and Choeronycteris. According to Olin L. Webb, Albert A. Alcorn and Jack M. Mohler, who obtained ex- amples of these bats on June 29, 1952, each species occupied slightly difiFerent quarters in darkened recesses of this cave. The Choeronycteris were most easily disturbed and flushed from hang- ing places quicker than did the other species. This same cave was revisited on April 8, 1954; only one bat, a Corynorhinus, was present. At a cave at El Chiflon, John W. Twente, Jr., and O. B. Heck found approximately 18 of these bats in one group. When alarmed, many flew out of the cave and hid nearby in small holes in rocky ledges. Tadarida hrasiliensis was taken also in this cave. Single hog-nosed bats were ob- tained at Cueva de Don Jesus and in a small, shallow cave near San Pedro de las Colonias. Speci- mens were obtained on March 23, June 17, June 29, August 23, September 1 and 11. Average and extreme weights of 5 non-pregnant females from several places in southern Coa- huila were 16.8 (12.6-21.0); 2 males weighed 9.7 and 13.0 grams, respectively. A female with one embryo (length, crown to rump, 25 mm.) was obtained on March 23. On June 17, John W. Twente, Jr. found females in a cave at El Chiflon with half- grown young. These young were carried while the females were in flight. Four of these females in lactation were saved as study specimens. Several bats had deciduous teeth; these were studied by Stains and Baker ( 1954:437). Fig. 9. Distribution of the hog-nosed bat, Choeronycteris mexicana, in Coa- huila. 174 University of Kansas Publs., Mus. Nat. Hist. Coahuilan specimens are not distinguishable from examples of Choeronycteris mexicana from Tamaulipas and Jalisco. Measurements. — Average and extreme measurements of 3 adult males from southern Coahuila (KU 34549, 39204, 39207) and 10 adult females from southern Coahuila (KU 34544, 34547, 34552, 34553, 35669, 39205, 39206, 39208, 39209, 48235) are: $ total length, 102 (100-103), 9 92 (81-103); length of tail vertebrae, 7 (6-9), 8 (5-10); length of hind foot, 11 (10-12), 12 (10-13); height of ear from notch, 16 (16-16), 16 (15-18); length of fore- arm, 46.1 (45.1-47.8), 45.5 (43.2-46.6); greatest length of skull, 29.6 (29.2- 29.9), 29.8 (29.5-30.4); condylobasal length, 28.7 (28.2-28.9), 28.9 (28.7- 29.3); least interorbital constriction, 3.8 (3.6-3.9), 4.0 (3.8-4.1); palatal length, 18.1 (17.6-18.4), 18.3 (18.0-18.8); mastoidal breadth, 9.9 (9.9-10.0), 10.0 (9.9-10.5); width of braincase, 9.6 (9.4-9.7), 9.9 (9.6-10.1); depth of brain- case, 7.8 (7.7-7.8), 7.7 (7.3-7.9); alveolar length of upper premolar-molar series, 8.7 (8.6-8.8), 8.8 (8.6-9.3). Specimens examined. — Total, 22, from: 9 mi. W and 4 mi. S San Buena- ventura, 1800 ft., 4; 10 mi. S Santa Teresa, 4000 ft., 5; Cueva de San Jesus, 57 mi. N and 27 mi. W Saltillo, 1; 26 mi. SW San Pedro de las Colonias, 3700 ft., 1; El Chiflon, 21.7 mi. W SaltiUo, 6; 1 mi. S and 4 mi. W Bella Union, 7000 ft., 5. Leptonycteris nivalis nivalis (Saussure) Long-nosed Bat M. [=: Ischnoglossa] nivalis Saussure, Rev. Mag. Zool., Paris, ser. 2, 12:492, November, 1860, type from near snow line on Mount Orizaba, Veracruz. Leptonycteris nivalis Miller, Proc. Biol. Soc. Washington, 13:126, April 6, 1900. Distribution. — Known from a single locality in southeastern Coahuila. Remarks. — Twenty-four Leptonycteris were obtained on July 9 by W. K. Clark from an abandoned mine situated on a mountain slope near Arteaga. The opening was surrounded by piiion and yucca. The bats were concentrated in a crevice in the high ceiling of the mine. Twenty-three specimens were captured; all but three were females. A mummified individual was picked up on the floor of the shaft. No other bats of this species were found or reported from Coahuila. Average and extreme weights of 20 non-pregnant females from near Arteaga (see below) were 21.7 (18.0-31.5); three males weighed 26.2, 22.9 and 21.0 grams, respectively. Females, taken on July 9, were not pregnant. Coahuilan specimens are paler and have a longer third finger and a broader and higher skull than nivalis from Veracruz, Distrito Federal and Oaxaca. Mammals of Coahuila, Mexico 175 Measurements. — Measurements of 2 males and average and extreme meas- urements of 5 females from 12 miles south and 2 miles east of Arteaga are: total length, $ 73, 83, 9 82 (80-86); length of hind foot, 16, 17, 16 (16-17); height of ear from notch, 16, 17, 17 (16-17); greatest length of skull, 27.2, 27.5, 28.5 (27.7-29.1); condylobasal length, 26.5, 26.3, 27.3 (26.4-27.8); least interorbital constriction, 5.0, 4.5, 5.0 (4.8-5.3); palatal length, 14.6, 14.6, 15.1 (14.6-15.3); mastoidal breadtli, 11.7, 11.9, 11.9 (11.6-12.3); depth of braincase, 8.6, 8.3, 8.6 (8.2-8.9); breadth across third upper molars, 7.0, 7.2, 7.1 (6.9-7.2); alveolar length of ma.xillary tooth-row, 9.3, 9.0, 9.5 (9.3-9.8). Specimens examined. — Total, 24, from 12 mi. S and 2 mi. E Arteaga, 7500 ft. Myotis calif omicus calif omicus (Audubon and Bachman) California Myotis Vespertilio califomicus Audubon and Bachman, Jour. Acad. Nat. Sci. Phila- delphia, vol. 8, pt. 2, p. 285, 1842, type from Monterey County, Cali- fornia. Myotis calif ornicus Miller, N. Amer. Fauna, 13:69, October 16, 1897. Distribution. — All of State except northeastern part (see fig. 10). Remarks. — The California myotis was obtained chiefly at lower elevations in desert associations where the bat congregated at ranch tanks and other surface water in the evening. Specimens were taken both with the collecting gun and in mist nets stretched over water. All collecting stations, where califomicus was obtained, were no more than one mile from rocky cliffs, which probably pos- sessed cavities suitable as day- time retreats. Dalquest (1953: 50) describes daytime hiding places of these bats between leaves of tree yuccas in San Luis Potosi. This bat probably oc- curs throughout the State except on the Coastal Plain of north- eastern Coahuila, Average and extreme weights of 4 adult males from localities in Coahuila were 4.2 (3.9-5.0); of 8 adult females, 3.9 (3.5-4.1). One female, taken on June 29, was in lactation. Females that seemed not to be pregnant were Fig. 10. Distribution of the California obtained on March 3, 4, 25, June myotis, Myotis califomicus californi- no j ^i -r^ i n cus, in Coahuila. 29, and another on December 2, 176 University of Kansas Publs., Mus. Nat. Hist. Coahuilan specimens are slightly paler than caUfornicus from Los Angeles County, California, but closely approximate in color a specimen of this subspecies from Socorro County, New Mexico, and are not so pale as topotypes of M. c. stephensi Dalquest from San Diego County, California. The skull of M. c. mexicaniis ( Saus- sure) from Michoacan is slightly larger with a broader rostrum than Coahuilan caUfornicus. Measurements. — Average and extreme measurements of 4 adult males from northern Coahuila (KU 44721, 54360, 54362, 54363) and 6 adult females from northern Coahuila (KU 44720, 44722, 54361, 54364, 57006, 57764), respectively, are: total length, 82 (81-83), 85 (82-88); length of tail verte- brae, 41 (40-42), 40 (35-43); length of hind foot, 7 (6-7), 7 (7-8); height of ear from notch, 13 (13-15), 13 (13-14); length of forearm, 32.5 (31.5-34.4), 33.2 (33.0-33.6); greatest length of skull, 13.5 (13.2-13.8), 14.0 (13.6-14.4); condylobasal length, 12.8 (12.6-13.3), 13.3 (13.1-13.5); zygomatic breadth, 8.3 (8.0-8.4), 8.5 (8.4-8.6); mastoid breadth, 7.0 (6.8-7.2), 7.2 (7.1-7.4); least interorbital constriction, 3.2 (3.2-3.4), 3.3 (3.1-3.5); palatal breadth, 6.9 (6.8-7.4), 7.2 (7.0-7.5); greatest breadth across upper molars, 5.1 (5.1- 5.2), 5.2 (5.0-5.4); length of maxillary tooth-row, 5.2 (5.1-5.4), 5.3 (5.2-5.6). Specimens examined. — Total, 14, from: 10 mi. S and 5 mi. E Boquillas, 1500 ft., 1; 16 mi. S Boquillas, 1600 ft., 1; El Rio Alamos, 3300 ft., Canon Mulato, Rancho Las Margaritas, 1; 6 mi. N and 2 mi. W Castillon, 3750 ft., 4; 4 mi. W Hacienda La Mariposa, 2300 ft., 5; La Gacha, 1600 ft., 1. Additional record (Miller and Allen, 1928:154): Guadalupe [ = Sierra de Guadalupe]. My Otis evotis auriculus Baker and Stains Long-eared Myotis Myotis evotis auriculus Baker and Stains, Univ. Kansas Publ., Mus. Nat. Hist., 9:83, December 10, 1955, type from 10 mi. W and 2 mi. S Piedra. 1200 ft.. Sierra de Tamaulipas, Tamaulipas. Distribution. — Foothills and adjacent mountains in northern and eastern Coahuila. Remarks. — Five long-eared myotis bats were snared in mist nets stretched over a concrete water tank situated in mixed chaparral and mesquite in the arid foothills. These bats, taken on March 25 and 26, were females; average and extreme weights were 6.6 (5.9-7.2). Coahuilan specimens are not so dark as typical auriculus from Tamaulipas but are much darker and richer in color than M. e. evotis (H. Allen) from Colorado and Wyoming. In the degree of rise of the forehead, Coahuilan specimens are somewhat inter- mediate between auriculus and evotis; otherwise, in large size, paleness of the ears and in other features the Coahuilan bats resemble auriculus more than evotis. Mammals of Coahuila, Mexico 177 Measurements. — Measurements of three adult females from 4 miles west of Hacienda La Mariposa are: total length, 97, 86, 96; length of tail vertebrae, 41, 39, 45; length of liind foot, 10, 10, 8; height of ear from notch, 20, 18, 20; length of forearm, 40.2, 37.3, 38.5; greatest length of skull, 16.2, 15.8, 16.1; condylobasal length, 15.6, 15.0, 15.4; zygomatic breadth, 10.1, 9.4, 9.9; least interorbital constriction, 3.9, 3.7, 3.7; breadth of braincase, 7.5, 7.5, 7.6; occipital depth, 5.5, 5.7, 5.6; palatal length, 8.7, 8.7, 8.9; mastoidal breadth, 8.3, 8.2, 8.3; breadth across third upper molars, 6.3, 6.1, 6.1; length of mandible, 11.9, 11.2, 11.7; length of ma.xillary tooth-row, 6.7, 6.6, 6.6; lengtli of mandibular tooth-row, 8.1, 8.1, 8.1. Specimens examined. — Total, 5, from 4 mi. W Hacienda La Mariposa, 2300 ft. Myotis planiceps Baker Flat-headed Myotis Myotis planiceps Baker, Proc. Biol. Soc. Washington, 68:165, December 31, 1955, type from 7 mi. S and 4 mi. E Bella Union, 7200 ft., Coahuila. Distribution. — Known only from the type locality in extreme southeastern Coahuila. Remarks. — The only known specimen of planiceps was shot on the evening of June 24, 1952, by Albert A. Alcorn as it circled over a water-filled earthen tank in an open, intermontane valley near Bella Union. Surrounding mountains supported montane mesic forests; probably the bat lives in mountainous areas. Short forearm, greatly flattened cranium and small teeth dis- tinguish planiceps from all of the other known species of North American Myotis. Measurements. — Measurements of the holotype (an adult male) are: total length, 76; length of tail vertebrae, 25; length of hind foot, 8; height of ear from notch, 10; length of forearm, 26.5; length of thumb, 3.8; length of third metacarpal, 24.3; length of fifth metacarpal, 23.3; condylobasal length of skull, 13.3; zygomatic breadth, 8.1; least interorbital constriction, 3.4; breadth of braincase, 7.1; mastoidal breadth, 7.2; palatal length, 6.7; maxillary breadth across M3, 4.9; length of mandible, 8.8; length of maxillary tooth-row, 4.9; length of mandibular tooth-row, 6.2. Specimen examined. — One, from 7 mi. S and 4 mi. E Bella Union, 7200 ft-, (type locality). Myotis velifer incautus (J. A. Allen) Cave Myotis Vespertilio incautus J. A. Allen, Bull. Amer. Mus. Nat. Hist., 8:239, April 22, 1896, type from San Antonio, Bexar County, Texas. Myotis velifer incautus Miller and G. M. Allen, U. S. Nat. Mus., Bull. 144:92, May 25, 1928. Distribution. — Lower elevations of the State except for northeastern Coa- huila, where the bat is either rare or absent (see fig. 11). Remarks. — The cave myotis was found frequenting watering 178 University of Kansas Publs., Mus. Nat. Hist. places at dusk in desert areas in most parts of Coahiiila except for the Coastal Plain in the northeast. Most ranch tanks and other surface water, where these bats congregated in connection with their foraging activities, usually were close to rocky bluffs that probably contained openings where daytime retreats were available. Bats were taken from limestone caves near San Buenaventura and at Cueva de Don Jesus. Gerd Heinrich received several velifer at La Flor from natives; probably these were gathered from a cave. Bats were taken either by gun or in mist nets over water as early as March 8 and as late as September 23. Average and extreme weights of velifer from Coahuila were: for 21 males, 7.9 (7.0-9.6); for 10 non-pregnant females, 8.5 (7.0-10.0). One lactating female was captured on June 29; females examined on March 8, 17, 28, 30, 31, April 1, and September 23 seemed not to be pregnant. Coahuilan specimens are as- signed to incautus because of their pallor; their skulls are smaller than those of incautus from Comanche County, Kansas, and resemble more closely those of M. V. velifer (J. A. Allen) from Jalisco ( 13 miles south and 15 miles west of Guadalajara ) . Measurements. — Average and ex- treme measurements of 10 adult males from La Flor and 5 adult females from places in Coahuila (KU 44711, 44713, 44719, 54350, 55044), respectively, are: total length, 95 (90-98), 97 (94- 104); length of tail vertebrae, 43 (42- 45), 45 (42-49); length of hind foot, 10 (10-10), 11 (10-11); height of ear from notch, 15 (15-16), 15 (14-16); length of forearm, 42.6 (40.1-43.9), 43.1 (41.7-44.1); greatest length of skull, 16.1 (15.9-16.4), 16.3 (16.1-16.5); condylobasal length, 15.6 (15.3-15.8), 15.7 (15.4-16.0); zygomatic breadth, 10.5 (10.3-10.9), 10.6 (10.2-11.1); least interorbital constriction, 4.0 (3.9-4.2), 4.0 (4.0-4.1); mastoidal breadth, 8.5 (8.3-8.6), 8.5 (8.2-8.7); palatal length, 8.6 (8.4-8.9), 8.7 (8.5-9.0); greatest breadth across upper molars, 6.8 (6.7- 7.0), 6.9 (6.7-7.1); length of maxillary tooth-row, 6.5 (6.4-6.7), 6.8 (6.5-7.1). Specimens examined. — Total, 42, from: S side Rio Grande, 600 ft., 17 mi. S Dryden, Terrell County, Texas in Coahuila, 1; 11 mi. W Hacienda San Miguel, 2200 ft., 1; 1 mi. N Boquillas, 1700 ft., 1; 2 mi. W Jimenez, 2; Fortin, Fig. 11. Distribution of the cave my- otis, Myotic velifer incautus, in Coa- huila. Mammals of Coahuila, Mexico 179 3300 ft., Rancho Las Margaritas, 5; Hacienda Las Margaritas, 2800 ft., 2; 3 mi. S and 8 mi. E Hechicero, 4650 ft.. Chihuahua, in Coahuila, 1; 2 mi. S and 5 mi. E San Geronimo, 1; 6 mi. N and 2 mi. W Castillon, 3750 ft., 2; 8 mi. N and 4 mi. W Muzquiz, 1800 ft., 2; 2 mi. S and 3 mi. E San Juan de Sabinas, 1160 ft., 1; 8 mi. N Hermanas, 1500 ft., 1; 9 mi. W and 4 mi. S San Buenaventura, 2000 ft., 1; San Lazaro, 4800 ft., 62 mi. N and 22 mi. W Saltillo, 1; Cueva de Don Jesus, 5100 ft., 57 mi. N and 27 mi. W Saltillo, 1; La Flor, 4500 ft.. Valley Rio Aguanaval, 19. Myotis volans interior Miller Long-legged Myotis Myotis longicrus interior Miller, Proc. Biol. Soc. Washington, 27:211, Oc- tober 13, 1914, type from 5 mi. S Twining, Taos County, New Mexico. Myotis volans interior Miller and G. M. Allen, U. S. Nat. Mus., Bull. 144:142, May 25, 1928. Distribution. — Known from the Sierra del Carmen, may occur in other mountainous parts of the State. Remarks. — Two long-legged myotis were shot as they foraged over a pool surrounded by oaks and pine in the Sierra del Carmen on April 7. Several others were seen. No other specimens were obtained but this bat may occur in other wooded, montane situa- tions in Coahuila, especially in the central and southeastern parts of the State. One female weighed 6.8 grams; the other, labeled as having no embryos, weighed 5.2 grams. Specimens from Coahuila are assigned to interior and resemble closely bats of this subspecies from Rio Arriba County, New Mexico. Measurements. — One adult female from Club Sierra del Carmen measures: total length, 93; length of tail vertebrae, 46; height of ear from notch, 11; greatest length of skull, 14.6; condylobasal length, 14.3; zygomatic breadth, 8.8; least interorbital constriction, 4.2; mastoidal breadth, 8.1; palatal length, 7.4; greatest breadth across upper molars, 6.0; length of maxillary tooth-row, 5.5. Specimens examined. — Two from Club Sierra del Carmen, 4950 ft., 2 mi. N and 6 mi. W Piedra Blanca. Myotis yumanensis yumanensis (H. Allen) Yuma Myotis Vespertilio yumanensis H. Allen, Smithsonian Misc. Coll., vol. 7, publ. 165, p. 58, June 1864, type from Old Fort Yuma, Imperial County, CaHfomia. Myotis yumanensis Miller, N. Amer. Fauna, 13:66, October 16, 1897. Distribution. — Known from one locality in northern Coahuila, probably found in desert areas in other parts of the State. Remarks. — Two yumanensis were shot at dusk as they foraged along the edge of the Rio Grande. The fact that no others were obtained in desert areas, although collections of bats were made in most parts of the State, suggests that this species may be less common in Coahuila than M. velifer. The two females, taken on 180 University of Kansas Publs., Mus. Nat. Hist. March 18, were seemingly not pregnant and weighed 4.1 and 4.7 grams. In external appearance, Coahuilan specimens compare favorably with yumanensis from Washington County, Utah, and Navajo County, Arizona. Skulls were lost in the field. Measurements. — Two females (see below) measure: total length, 81, 80; length of tail vertebrae, 36, 35; length of hind foot, 8, 10; height of ear from notch, 13, 13; length of forearm, 32.8, 33.3. Specimens examined. — Two from S side Rio Grande, 600 ft., 17 mi. S Dryden, Terrell County, Texas, in Coahuila. Pipistrellus hesperus maximus Hatfield Western Pipistrelle Pipistrellus hesperus maximus Hatfield, Jour. Mammalogy, 17:261, August 14, 1936, type from Dog Spring, Grant County, New Mexico. Distribution. — State-wide except for Coastal Plain (see fig. 12). Remarks. — The western pipistrelle is found on the desert plains and in the mountains of Coahuila but seems not to occur on the Gulf Coastal Plain in the northeastern part of the State, where Pipistrellus subflavus has been taken along wooded streams. Pipis- trellus hesperus reaches the northeastern limit of its range in Coahuila. It is a conspicuous bat at ranch tanks, along open streams and at other surface waters in late evening and early morning. This species is usually the first bat to make its appear- ance in flight in the evening and the last one to go into seclusion in the morning. On cloudy af- ternoons the western pipistrelle forages in narrow mountain carious as early as two hours be- fore sunset. This bat concen- trates most commonly at surface water in caiions or in the vicinity of rocky cliffs, where daytime retreats in crevices are numer- ous. Large numbers of these bats were seen at Boquillas, San La- zaro, and Cuatro Cienegas. Bats Fig. 12. Distribution in Coahuila of the pipistrcllcs. 1. western )iipistrelle, ripistrclltis hesperus maximtis. 2. east- ern pipistrelle, Pipistrellus subflavus clarus. Mammals of Coahuila, Mexico 181 were taken as low as 1700 feet in elevation (vicinity of Boquillas) and as high as 6500 feet (Sierra de Guadalupe). Many specimens were snared in mist nets stretched over their watering places. Average and extreme weights of Hesperus from Coahuila were: for 22 males, 3.4 (2.0-4.2); for 16 non-pregnant females, 4.0 (3.1- 4.8). Lactating females were taken on June 17, 22, and 24. Coahuilan specimens are assigned to maximus although some, but not all, examples from southeastern Coahuila are darker and resemble P. h. potosinus Dalquest from Irturbide, Nuevo Leon (55136 KU). Measurements. — Average and extreme measurements of ten adult bats from Puerto San Lazaro (San Lazaro and 3 miles south and 3 miles east of Muralla) are: total length, 78 (74-83); length of tail vertebrae, 32 (28-37); length of hind foot, 7 (6-8); height of ear from notch, 11 (10-12); length of fore- arm, 31.4 (30.4-32.6); length of tibia, 12.5 (12.1-12.7); greatest length of skull, 12.6 (12.3-13.1); condylobasal length, 12.1 (11.9-12.6); zygomatic breadth, 8.1 (7.8-8.3); least interorbital breadth, 3.4 (3.2-3.5); mastoidal breadth, 6.6 (6.4-6.8); breadth of braincase, 6.3 (6.2-6.5); length of maxillary tooth-row, 4.4 (4.2-4.6). Specimens examined. — Total, 75, from: 1 mi. N Boquillas, 1700 ft., 8 Hacienda Las Margaritas, 2800 ft., 3; 3 mi. S and 8 mi. E Hechicero, 4650 ft. Chihuahua, in Coahuila, 3; 6 mi. N and 2 mi. W Castillon, 3750 ft., 7; 4 mi W Hacienda La Mariposa, 2300 ft., 7; 9 mi. W and 4 mi. S San Buenaventura 1800 ft., 3; 4 mi. NW Cuatro Cienegas, 1; 3 mi. NW Cuatro Cienegas, 2450 ft. 1; 6 mi. E Cuatro Cienegas, 2200 ft., 2; 4 mi. N Acatita, 2; 3 mi. S and 3 mi. E Muralla, 12; San Lazaro, 62 mi. N and 22 mi. W Saltillo, 8; Hacienda El Tuhllo, 5 km. S HipoUto, 5; 17 mi. N and 8 mi. W Saltillo, 1; 11 mi. N and 4 mi. W Saltillo, 4; 3 mi. SE Torreon, 3800 ft., 2; El Chiflon, 21.7 mi. W Saltillo, 2; La Pastora Rancho, 41 mi. W and 15 mi. S Saltillo, 1; W foot Pico de Jimulco, 5000 ft., 2; W foot Sierra de Guadalupe, 6500 ft., 10 mi. S and 5 mi. W General Cepeda, 1. Pipistrellus subflavus clarus Baker Eastern Pipistrelle Pipistrellus subflavus clarus Baker, Univ. Kansas Publ., Mus. Nat. Hist., 7:585, November 15, 1954, type from 2 mi. W Jimenez, 850 ft., Coahuila. Distribution. — Northern Coahuila, restricted to timbered areas along the Rio Grande and its major tributaries (see fig. 12). Remarks. — The eastern pipistrelle reaches the western Hmit of its distribution along the Rio Grande and its tributaries in northern Coahuila. This bat is restricted to parklike stands of cypress, pecan, willow and other trees that border permanent streams, including the Rio Grande, Rio de San Diego, and Rio Sabinas. This bat was not found at stock tanks and stream borders in adjacent areas where no large trees occur. Probably this species roosts in trees or in tree holes (see Findley, 1954:433). The eastern pipistrelle reached Coahuila by way of the watershed of the Rio Grande rather than 182 University of Kansas Publs., Mus. Nat. Hist, directly from the east across the generally unsuitable habitat of the Hill Country of Texas. Except for records along the Rio Grande no eastern pipistrelles have been recorded in southwestern Texas west of Kerr County (see Taylor and Davis, 1947:17). Along the Rio de San Diego not far from Jimenez, eastern pipis- trelles were numerous in June, 1952; a series of 15 was obtained. These bats appeared in early evening at least 30 minutes before dark and foraged among the large trees. They were active also for a short while after daybreak. Few were taken in mist nets stretched across water; most were shot with dust loads. Both the eastern pipistrelle and the western pipistrelle live in Coahuila but have not been found to occur together. The former lives only along the narrow forested streams whereas the western pipistrelle occurs elsewhere throughout the arid desert plains and mountains. Pipistrelhis subflavus clarus is the palest of eastern pipistrelles. It probably intergrades with P. s. subflavus (Cuvier) in Tamauli- pas or extreme southern Texas, but seems to have little relation to P. s. veracrucis (Ward), which occurs to the southeast. Measurements. — Average and extreme measurements of five adult females from the type locality are: total length, 88 (85-92); length of tail vertebrae, 41 (36-45); length of hind foot, 9 (8.5-11); height of ear from notch, 13 (12-13); length of foreann, 33.4 (32.9-33.8); length of tibia, 14.6 (14.5-14.8); greatest length of skull, 13.1 (12.7-13.3); condylobasal length, 12.4 (12.1- 12.6); breadth of braincase, 6.8 (6.7-7.0); zygomatic breadth, 8.0 (7.8-8.2); mastoidal breadth, 6.9 (6.5-7.1); length of maxillary tooth-row, 4.5 (4.4-4.6). Specimens examined. — Total, 18, from: 2 mi. W Jimenez, 850 ft., 15; 2 mi. S and 3 mi. E San Juan de Sabinas, 1160 ft., 3. Eptesicus fuscus pallidus Young Big Brown Bat Eptesicus pallidus Young, Proc. Acad. Nat. Sci. Philadelphia, 60:408, Oc- tober 2, 1908, type from Boulder, Boulder County, Colorado. Eptesicus fuscus pallidus Miller, U. S. Nat. Mus., Bull. 79:62, December 31, 1912. Distribution. — Northern Coahuila, probably less common in the western part of this area (see fig. 13). Remarks. — There are few reports of Eptesicus from the arid, northern part of the Central Plateau ( Mesa de Norte ) . In western Coahuila, only a few specimens were obtained even though con- siderable effort to collect bats was made by each field party visiting the area. In the more humid, northeastern part of the state Eptesi- cus seems more abundant, especially along the Rio Salado where Mammals of Coahuila, Mexico 183 some woody cover grows and suitable daytime retreats in caves in limestone bluflFs occur nearby. Twenty-one big brown bats were collected from one such cave on the south side of the canon of the Rio Salado near San Buenaventura; this cave yielded also examples of Choeronycteris, Tadarida, Corynorhintis, and My Otis velifer. Other big brown bats were obtained over water at dusk. One bat, taken in a mist net stretched over a small stream at San Marcos, was largely eaten by minnows, when that part of the net containing this animal sagged into the water. The series of Eptesicus taken on June 29 from near San Buena- ventura included examples of adults, younger adults that had less worn teeth and less developed cranial ridges and shorter forearms, and Juvenal individuals, many of which probably were still being nursed by females. Most of the young still retained some decidu- ous teeth although permanent replacements had pushed them laterally. Based on examination of six young, each with one or more deciduous teeth, it was found that the deciduous dentition includes three incisors, one canine and three premolars in each jaw, although not all of these teeth are present in any one specimen. The order in which these teeth drop out seems to be: upper premolars, lower premo- lars, canines, lower incisors, and upper incisors. These decidu- ous teeth are slender, hooked spicules, decidedly smaller and grossly different from the per- manent teeth which replace them. One tooth, thought to be an upper, deciduous, third pre- molar, and which is between the in-growing permanent premolar and the first molar, is noticeably different from other deciduous teeth, being premolariform al- though less than one-fourth as large as the permanent premolar. In comparison with E. f. palli- dus from San Diego County, California, and with E. f. fuscus (Beauvois) from Crawford County, Fig. 13. Distribution of the big brown bat, Eptesicus fuscus pallidus, in Coa- huila. 184 University of Kansas Publs., Mus. Nat. Hist. Pennsylvania, Coahuilan specimens are somewhat intermediate in color but closer to pallidus, to which they are assigned. Probably the zone of intergradation between the paler western pallidus and the darker eastern fuscus is a broad one, and assignment based on color of specimens from within this zone to one or the other forms is often a diflScult task. Measurements. — Average and extreme measurements of 7 adult females from 9 miles west and 4 miles south of San Buenaventura are: total length, 123 (116-125); length of tail vertebrae, 52 (49-57); length of hind foot, 12 (12-13); height of ear from notch, 18 (17-19); length of forearm, 50.2 (49.5- 51.1); condylobasal length of skull, 18.7 (18.5-19.1); zygomatic breadth, 13.1 (12.9-13.3); least interorbital constriction, 4.4 (4.2-4.6); mastoidal breadth, 10.2 (10.1-10.5); palatal length, 9.5 (9.3-9.7); breadth across M3, 8.1 (7.8- 8.4); alveolar length of maxillary tooth-row, 7.3 (7.0-7.5). Specimens examined. — Total, 27, from: 1 mi. N Boquillas, 1; 9 mi. W and 4 mi. S San Buenaventura, 1800 ft., 21; 4 mi. NW Cuatro Cienegas, 2; 3 mi. NW Cuatro Cienegas, 1; San Marcos, 1; Acatita, 3600 ft., 1. Lasiurus borealis borealis (Miiller) Red Bat Vespertilio borealis Miiller, Natursyst. SuppL, p. 20, 1776, type from New York. Lasiurus borealis Miller, N. Amer. Fauna 13:105, October 16, 1897. Distribution. — Northern Coahuila. Remarks. — The red bat is known from northeastern Coahuila, probably distributed only where large trees occur along the Rio Grande and its tributaries and in forests of oak and pine of the mountains. Two specimens are reported from the State; both were taken in nets stretched over water. One, a male, was obtained in March at Fortin in the Serranias del Burro where oaks, pine and juniper are present. The other, a lactating female, was taken in June near Jimenez in a parklike stand of cypress and pecan along the Rio de San Diego. The male weighed 8.8 grams. The adult female is dull buffy-brown while the younger male is bright orange-red. The male, taken on a cool evening on March 29 at 3300 feet elevation, may have been a migrant. Measurements. — Measurements of the adult female from 2 miles west of Jimenez are: total length, 118; length of tail vertebrae, 56; length of hind foot, 9; height of ear from notch, 12; length of forearm, 41.5; condylobasal length of skull, 13.3; zygomatic breadth, 9.9; least interorbital constriction, 4.4; mastoidal breadth, 7.9; breadth of braincase, 7.0; length of maxillary tooth-row, 4.9. Specimens examined. — Total, 2, from: 2 mi. W Jimenez, 850 ft., 1; Fortin, 3300 ft., Rancho Las Margaritas, 1. Mammals of Coahuila, Mexico 185 Lasiurus cinereus cinereus ( Beauvois ) Hoary Bat Vespertilio cinereus (misspelled linereus) Beauvois, Catal. Raisonne Mus. Peale, Philadelphia, p. 18, 1796, type from Philadelphia, Pennsylvania. Lasiurus cinereus H. Allen, Smithsonian Misc. Coll., vol, 7, publ. 165, p. 21, June, 1864. Distribution. — Probably occurs throughout Coahuila, especially where trees are present. Records of breeding within the State are lacking. Remarks. — The hoary bat probably occurs in most parts of Coa- huila. It has been taken in desert situations, at Acatita, and in pine and oak in the Serranias del Burro at Fortin. Specimens were ob- tained in March and April; these may have been either migrants or representatives of the resident population. None was recorded in summer. Females taken in March were not pregnant; one weighed 24.6 grams and the other 28.5 grams. These two bats were taken on March 29 sometime between midnight and dayhght in mist nets stretched over water at Fortin. Measurements. — Measurements of an adult female and a subadult female from Fortin, respectively, are: total length, 145, 136; length of tail vertebrae, 65, 61; length of hind foot, 12, 13; height of ear from notch, 19, 20; length of forearm, 56.4, 53.8; condylobasal length of skull, 17.2, 16.5; zygomatic breadth, 13.8, 13.0; least interorbital constriction, 5.2, 5.1; mastoidal breadth, 10.9, 10.4; breadth of braincase, 9.1, 9.0; length of maxillary tooth-row, 6.2, 6.5. Specimens examined. — Total, 4, from: Fortin, 3300 ft., Rancho Las Mar- garitas, 2; 4 mi. N Acatita, 1; Acatita, 3600 ft., 1. Dasypterus ega xanthinus Thomas Western Yellow Bat Dasypterus ega xanthinus Thomas, Ann. and Mag. Nat. Hist., ser. 6, vol. 20, p. 544, December 1897, type from Sierra Laguna, Baja CaUfomia. Distribution. — Known from one place in northern Coahuila but probably state-wide. Remarks. — The western yellow bat probably occurs uncommonly throughout Coahuila. Specimens have been taken in northern Coahuila and within ten miles of the southern border, in Zacatecas (Concepcion del Oro, 57968-71 KU). Little is known of the habits of this bat; it is probably migratory and may not breed in Coahuila. Those from nearby Zacatecas were taken in late February. Coa- huilan specimens, all males, were captured on March 25 and 26 in a mist net stretched over a concrete tank, which was situated among mesquite and other scrub vegetation at the base of low, rocky hills. No large trees were present. The bats were netted more than two hours after dark and were taken in the lower edge of the net where it touched the water in the tank. No other western yellow bats were observed although 23 bats of other species were 186 University of Kansas Publs., Mus. Nat. Hist. netted on these two cool nights; the temperature had dropped to 29° F. two nights before. These three males weighed 16.2, 13.4 and 17.9 grams. Specimens in the University of Kansas Museum of Natural History from Coahuila, Tamaulipas, and Zacatecas together with records from San Luis Potosi (see Dalquest, 1953:62) indicate that this bat occurs throughout northern Mexico, Measurements. — Measurements of three adult male xanthinus (see below), respectively, are: total length, 123, 116, 113; length of tail vertebrae, 51, 52, 52; length of hind foot, 7, 9, 7; height of ear from notch, 12, 12, 13; length of forearm, 45.8, 44.2, 44.8; condylobasal length of skull, 15.9, 15.2, 14.8; zygomatic breadth, 11.5, 10.7, 10.7; least interorbital constriction, 4.9, 5.1, 4.8; mastoidal breadth, 8.9, 8.9, 8.4; breadth of braincase, 8.5, 8.3, 8.0; length of maxillary tooth-row, 5.6, 5.5, 5.4; breadth across upper canines, 6.1, 6.2, 5.9; breadth across upper molars, 7.8, 7.2, 7.3. Specimens examined. — Total, 3, from 4 mi. W Hacienda La Mariposa, 2300 ft. Nycticeius humeralis mexicanus Davis Evening Bat Nycticeius humeralis mexicanus Davis, Jour. Mamm., 25:380, December 12, 1944, type from Rio Ramos, 20 km. NW Montemorelos, Nuevo Leon. Distribution. — Northeastern Coahuila (see fig. 14). Remarks. — The evening bat is common along timbered stream- courses in northeastern Coahuila, especially among the parklike stands of cypress, pecan, willow and other large trees that border the Rio Sabinas, the Rio de San Diego and other streams. The species probably occupies cavi- ties in the larger trees during the day; this habit was observed by Davis (1944:380) in Nuevo Leon. The bat usually appeared at dusk and was shot and also caught in mist nets stretched across water. Most specimens were snared near the bottom edge of the mist net, where it touched the water. On two oc- casions, evening bats were taken at ranch tanks several miles from large timber. The species seemed much less common in these situ- ations ^'^- ^'^^ Distribution of the evening * bat, Nycticeius humeralis mexicanus, Females taken in March in Coahuila. Mammals of Coahuila, Mexico 187 showed no signs of pregnancy. Those taken in mid-June either were not pregnant or seemed to be lactating. Young bats, born earlier in the year, were taken in the period June 18-21. Average and extreme weights of Nycticeius from northeastern Coahuila were: for seven adult females, 8.3 (7.3-10.9); for two adult males, 6.4 and 6.8. Specimens from Coahuila are assigned to mexicanus on the basis of their pale coloring. Measurements. — Average and extreme measurements of 5 adult males and 10 adult females from 2 miles west of Jimenez, respectively, are: total length, 93 (92-95), 98 (94-102); length of tail vertebrae, 39 (37-41), 39 (33-42); length of hind foot, 9 (8-10), 10 (9-10); height of ear from notch, 13 (12-14), 13 (12-14); length of forearm, 34.0 (32.5-36.7), 35.1 (33.6-36.1); greatest length of skull, 14.0 (13.3-14.6), 14.4 (14.0-14.6); zygomatic breadth, 9.6 (9.2-9.9), 10.1 (9.7-10.5); mastoidal breadth, 7.9 (7.5-8.4), 8.2 (7.9-8.7); occipital height, 5.4 (5.1-5.6), 5.4 (5.3-5.5); length of maxillary tooth-row, 5.4 (5.2-5.6), 5.5 (5.3-5.7). Specimens examined. — Total, 75, from: 2 mi. W Jimenez, 850 ft., 40; 2 mi. S and 11 mi. E Nava, 810 ft., 3; Hacienda La Candelaria, 3 mi. S and 13 mi. E Nava, 800 ft., 1; 10 mi. E Hacienda La Mariposa, 3000 ft., 1; 8 mi. N and 4 mi. W Muzquiz, 1800 ft., 7; 2 mi. S and 3 mi. E San Juan de Sabinas, 1160 ft., 23. Corynorhinus townsendii australis Handley Lump-nosed Bat Corynorhinus townsendii australis Handley, Jour. Washington Acad. Sci., 45:147, May, 1955, type from 2 mi. W Jacala, 5500 ft., Hidalgo. Distribution. — State-wide, except for northeastern part (see fig. 15). Remarks. — Bats of this species occur in desert areas of Coahuila but seem to be less common than the western pipistrelle, the pallid bat and the common free-tailed bat. One specimen was snared in a mist net stretched over water near the Hacienda La Mariposa on March 26. Others were obtained in caves. Near San Buenaventura two lump-nosed bats were caught in a cave in company with Choeronycteris, Tadarida, Eptesicus and Myotis velifer on June 29; on April 8, two years later, one Corynorhinus was the only bat found in this large cave. Near Bella Union, Corynorhinus roosted in small limestone caves in company with Choeronycteris in Sep- tember. On November 5, W. K. Clark obtained 19 from mine shafts on a slope a half mile north of the village of Muralla. He found these bats in two of six shafts inspected; all mines contained some guano. Clark collected these bats by hand; most of them were dormant and roosting individually. Of those sexed by him one was a male and ten were females. Average and extreme weights of 4 males from Coahuila were 188 University of Kansas Publs., Mus. Nat. Hist. 7.7 (5.7-10.0); of 10 non-pregnant females from a half mile north of Muralla, 7.5 (6.0-9.2). Non-pregnant females were taken on June 29 and November 5. Coahuilan specimens are paler than australis from 4 kilometers east of La Vigas, Veracruz, and almost as pale as C. t. pallescens Miller from Yavapai and Pima counties, Arizona. Probably these bats from Coahuila are in- tergrades between australis and pallescens but are assigned to australis, following Handley (1955:148). Measurements. — Average and ex- treme measurements of 9 males from several localities in central and south- em Coahuila and 10 females from % mile north of Muralla, respectively, are: total length, 94 (80-100), 101 (98-107); length of tail vertebrae, 47 (41-60), 49 (45-58); length of hind foot, 10 (9-12), 10 (9-10); height of ear from notch, 34 (32-36), 33 (31- 35); length of forearm, 42.6 (40.7- 43.8), 42.7 (39.4-44.3); greatest length of skull, 16.2 (15.7-16.7), 16.3 (15.7-16.6); condylobasal length, 15.0 (14.4-15.3), 15.1 (14.9-15.4); least interorbital constriction, 3.6 (3.5-3.8), 3.6 (3.4-3.9); palatal length, 7.0 (6.5- 7.3), 7.1 (7.0-7.4); mastoidal breadth, 9.2 (9.0-9.4), 9.3 (9.1-9.5); width of braincase, 7.8 (7.7-8.0), 7.9 (7.7-8.1); depth of braincase, 6.0 (5.9-6.1), 6.9 (5.9-6.1); length of maxillary tooth- row, 5.2 (4.9-5.4), 5.2 (5.1-5.4). Specimens examined. — Total, 27, from: 4 mi. W Hacienda La Mariposa, 2300 ft., 1; 9 mi. W and 4 mi. S San Buenaventura, 3; 'A mi. N Muralla, 4500 ft., 19; 1 mi. S and 4 mi. W Bella Um6n, 7000 ft., 3; Sierra Guadalupe, 10 mi. S and 5 mi. W General Cepeda, 1. Antrozous pallidas pallidas (LeConte) PalUd Bat V [espertilio]. pallidus LeConte, Proc. Acad. Nat. Sci. Philadelphia, vol. 7 ( 1854-55), p. 437, 1856, type from EI Paso, El Paso County, Texas. Antrozous pallidus H. Allen, Smithsonian Misc. Coll., vol. 7, publ. 165:68, June 1864. Distribution. — Desert areas in all parts of Coahuila (see fig. 16). Remarks. — The pallid bat is in most parts of Coahuila but seems to prefer desert areas where surface water is adjacent to daytime Fig. 15. Distribution of the lump- nosed bat, Corynorhinus townsendii australis, in Coahuila. Mammals of Coahuila, Mexico 189 retreats in limestone caves and buildings. Most specimens were snared in mist nets stretched over water in tanks or pools in arroyos. Normally, Antrozous appeared at watering places after dark; most individuals were netted in the two hours following the onset of darkness. Twenty-three females were snared in a mist net stretched over a steel cattle-trough on June 29 near Castillon. A well, near Boquillas, which had an opening approximately two and one-half feet in diameter and a water level 20 feet below the opening, was used as a watering place by pallid bats. One was taken in a net stretched over the opening of this well. The pallid bat used, as a night time retreat, an abandoned adobe house near Tanque Alvarez. At 9 p. m. on July 5, Olin Webb and Jack Mohler found 20 of these bats (the eight collected were females) within this structure. These bats had not been present in the adobe earlier that evening. Average and extreme weights of Antrozous from Coahuila were: for 6 adult males, 11.6 (8.7-14.5); for 23 adult females, 16.7 (14.0- 19.0). Of 60 of the palHd bats from Coahuila that were sexed, 50 are females and 10 are males. Females taken on May 10 were pregnant, each containing two embryos; females obtained on July 6 were lactating. Coahuilan specimens have been compared with A. p. palU- dus from northern Chihuahua and with A. p. pacificus Merriam from California (Los Angeles, Riverside and San Bernardino counties ) . In size and color ( see Miller, 1897:45) Coahuilan spec- imens are indistinguishable from pallidiis. Specimens from Jaral, assigned to A. p. pacificus by El- liot (1907:519) and six skulls from owl pellets, identified as A. pallidus by Baker (1953b:- 253), are referred here on geo- graphic grounds to A. p. pal- lidus. Measurements. — Measurements of 2 males from 10 miles south and 5 miles east of Boquillas and from Fortin and _, 1^ T^. , r 1 11 1 average and extreme measurements of riG. 15. Distribution or the pallia ^ i i r tt • j t \i bat, Antrozous pallidus pallidus, in ^ ^^"^^^^^ ^""""^ Hacienda Las Mar- Coahuila. garitas are: total length, 104, 110, 112 a— 1204 190 University of Kansas Publs., Mus. Nat. Hist. (107-119); length of tail vertebrae, 47, 50, 43 (41-46); length of hind foot, 12, 12, 11 (10-12); height of ear from notch, 31, 29, 28 (27-30); length of forearm, 50.3, 53.4, 52.8 (51.8-53.9); greatest length of skull, 18.6, 20.1, 19.6 (19.0-20.1); condylobasal length, 17.5, 18.5, 18.2 (17.9-18.5); zygomatic breadth, 11.7, 13.1, 12.4 (12.2-12.7); least interorbital constriction, 5.1, 5.4, 5.3 (5.0-5.4); palatal length, 8.2, 9.2, 9.2 (9.1-9.4); mastoidal breadth, 9.6, 10.1, 9.8 (9.6-9.9); greatest breadth across molars, 7.7, 8.1, 8.0 (7.8-8.2); alveolar length of maxillary tooth-row, 6.6, 7.1, 6.9 (6.8-7.0). Specimens examined. — Total, 63, from: S side Rio Grande, 600 ft., 17 mi. S Dr>'den, Terrell Co., Texas, in Coahuila, 2; 10 mi. S and 5 mi. E Boquillas, 1; Fortin, 3300 ft., Rancho Las Margaritas, 4; Hacienda Las Margaritas, 2800 ft., 12; 3 mi. S and 8 mi. E Hechicero, 4650 ft.. Chihuahua, in Coahuila, 1; 6 mi. N and 2 mi. W Castillon, 3750 ft., 23; 4 mi. N San Isidro, 3; 18 mi. S and 14 mi. E Tanque Alvarez, 4000 ft., 8; 4 mi. NW Cuatro Cicnegas, 2; 4 mi. N Acatita, 3600 ft., 5; Acatita, 3600 ft., 1; 3 mi. S and 3 mi. E Bella Union, 6750 ft, 1. Additional records: (Baker, 1953b:253): 3 mi. NW Cuatro Cienegas; (Elliot, 1907:518-519): Jaral. Tadarida brasiliensis mexicana (Saussure) Common Free-tailed Bat Molossus mexicanus Saussure, Rev. Mag. Zool., Paris, ser. 2, 12:283, July, 1860, type from Cofre de Perote, Veracruz. Tadarida brasiliensis mexicana Schwartz, Jour. Mammalog\% 36:106, Feb- ruary 28, 1955. Distribution. — State-wide (see fig. 17). Remarks. — The common free-tailed bat was taken in most parts of Coahuila, in caves and other darkened, daytime retreats and over tanks and other watering places at dusk and after dark. At dusk these bats flew swiftly along dry arroyos from mountainous areas to lower elevations, where they foraged and obtained water. Favorite daytime roosting places were in adobe buildings; the bats hid in cracks in the rafters and in the layers of cane supporting the ceilings. Although no large colonies of Tadarida were observed, several caves containing large acculumations of guano, thought to have been deposited by this bat, were examined. The Cueva de Don Jesus on the Rancho Guadalupe, approximately 51 miles north and 27 miles west of Saltillo, has been mined for guano for many years. Another cave, where guano has been mined, is in the Canon de la Alameda west of Nacimiento. In 1952 an extensive guano mining operation was reported in progress in a cave southeast of Torreon. The common free-tailed bat was taken as high as 4950 feet in ele- vation in the Sierra del Carmen, where specimens were shot over water surrounded by oaks and pine. However, the species was most common at lower elevations. In most places these bats were ob- tained in greater numbers, in mist nets stretched over water, than were other species. Mammals of Coahuila, Mexico 191 Average and extreme weights of Tadarida from Coahuila were: for 46 males, 9.4 (6.6-12.4); for 9 non-pregnant females, 10.2 (8.0- 11.3). Five females in lacta- tion were taken on June 28. Specimens from Coahuila are remarkably uniform in color. One female from San Pedro de las Colonias has patches of white fur on the right rump, between the ears and on the right side of the chin. Measurements. — Average and ex- treme measurements of 5 adult males from 8 miles north and 25 miles west of Cuatro Cienegas and 5 adult fe- males from San Pedro de las Col- onias, respectively, are: total length, 101 (94-107), 103 (99-108); length of tail, 36 (33-38), 37 (35-39); length of hind foot, 9 (9-10), 11 (11-12); height of ear from notch, 17 (17-18), 19 (18-20); greatest length of skull, 16.6 (16.3-16.9), 16.7 (16.2-17.1); condylobasal length, 15.8 (15.4-16.0), 15.7 (15.3- 16.2); zygomatic breadth, 9.6 (9.5- 9.9), 9.7 (9.5-9.9); least interorbital constriction, 4.8 (4.7-5.0), 4.9 (4.7-5.0); mastoidal breadth, 9.1 (8.9-9.3), 9.1 (9.0-9.3); maxillary breadth at M3, 7.1 (6.8-7.2), 7.0 (6.8-7.2); alveolar length of ma.xillar>' tooth-row, 6.0 (5.8-6.1); 5.9 (5.8-6.1). Specimens examined. — Total, 105, from: 1 mi. S and 9 mi. W Villa Acufia, 1; Canon del Cochino, 3200 ft., 16 mi. N and 21 mi. E Piedra Blanca, 1; 1 mi. N Boquillas, 700 ft., 1; Club Sierra del Carmen, 4950 ft., 2 mi. N and 6 mi. W Piedra Blanca, 5; Fortin, 3300 ft., Rancho Las Margaritas, 14; Hacienda Las Margaritas, 2800 ft., 13; Hacienda La Candelaria, 3 mi. S and 13 mi. E Nava, 1; 4 mi. W Hacienda La Mariposa, 2300 ft., 4; 8 mi. N and 4 mi. W Muzquiz 1800 ft., 5; 8 mi. N Hermanas, 6; 6 mi. E Hermanas, 1200 ft., 1; 8 mi. N and 25 mi. W Cuatro Cienegas, 4000 ft., 9; 9 mi. W and 4 mi. S San Buenaventura, 2000 ft., 2; 4 mi. N Acatita, 3600 ft., 8; Acatita, 3600 ft., 3; San Lazaro, 62 mi. N and 22 mi. W Saltillo, 1; Hacienda Guadalupe, 52 mi. N and 19 mi. W Sal- tillo, 2; San Pedro [= San Pedro de las Colonias], 9; El Chiflon, 21.7 mi. W Saltillo, 4; W foot Pico de Jimulco, 5000 ft., 1; Jimulco, 4500 ft.. Valley Rio Aguanaval, 11; Valley Rio Aguanaval, La Flor, 4400 ft., 3. Eumops perotis calif omicus (Merriam) Western Mastiff Bat Molossus calif ornicus Merriam, N. Amer. Fauna, 4:31, October 8, 1890, type from Alhambra, Los Angeles County, California. Eumops perotis californicus Sanborn, Jour. Mamm,, 13:351, November 2, 1932. Fig. 17. Distribution of the common free-tailed bat, Tadarida hrasiliensis mexicana, in Coahuila. 192 University of Kansas Publs., Mus. Nat. Hist. Distribution. — Central and western Coahuila, the easternmost sight record obtained was as San Lazaro (see fig. 18). Remarks. — Large, fast-flying bats, thought to be Eumops, were noted at several tanks and other watering places in Coahuila, but first appeared long after dark; J. R. Alcorn shot two at Acatita on April 15. On June 5 Robert W. Dickerman snared another in a mist net hung over water that flowed from a spring at San Marcos (approximately 20 miles south of Cuatro Ciene- gas). This latter specimen was attacked by minnows as it hung in a part of the net which sagged into water; Dickerman, on exam- ining the net the next morning, was able to save only the bones of the tvvo forearms and wings. A female, taken on Af)ril 15, v/as not pregnant. Specimens from Coahuila com- pare favorably with the descrip- tion of californicus in Sanborn ( 1932:351 ) and with examples of this subspecies from Los Angeles and San Bernardino counties, California. The skulls of the Coahuilan Eumops are damaged by dust shot. A single skull, iden- tified by Gilmore (1947:154) from cave deposits from near Cuatro Cienegas, is assigned by him to this subspecies. Measurements. — Measurements of a male and a female, both from Acatita, respectively, are: total length, 188, 177; length of tail vertebrae, 61, 61; lengtli of hind foot, 18, 19; height of ear from notch, 43, 40; length of forearm, 76.3, 72.6; zygomatic breadth, 17.1, — ; lacrimal breadth, 10.0, 9.8; least interorbital constriction, 5.4, — ; mastoidal breadth, 15.0, — ; maxillary breadth at M3, 12.0, 12.8; length of maxillary tooth-row, 12.9, 12.7; lengtli of mandible (an- terior face of incisors to plane of occipital condyles), 24.5, — . Specimens examined. — Total, 3, from: San Marcos, appro.\imately 20 mi. S Cuatro Cienegas, 1 (wing bones only); Acatita, 3600 ft., 2. Other record (Gilmore, 1947:154): 22 mi. W and 4 mi. S Cuatro Cienegas. Fig. 18. Distribution of the western mastiff bat, Eumoi)s perotis californi- cus, in Coahuila. Mammals of Coahuila, Mexico 193 Dasypus novemcinctus mexicanus Peters Nine-banded Armadillo Dasypiis novemcinctus var. mexicanus Peters, Monatsber. k. preuss. Akad. Wissensch. Berlin, p. 180, 1864, type from Matamoras, Tamaulipas, see Hollister, Jour. Mamm., 6:60, February 9, 1925. Distribution. — Found along pemianent streams in northeastern Coahuila. Remarks. — The armadillo is common in live-oak thickets along the Rio Grande, downstream from Piedras Negras. Along the river upstream from tliis city the armadillo is reported to be uncommon at Jimenez; on 22 June 1952 James S. Findley saw armadillo sign two miles west of Jimenez. According to Sefior Juan Parras, there are no armadillos on the Coahuilan side of the Rio Grande south of Dryden, Terrell County, Texas. The armadillo probably ranges no farther upstream along the Rio Grande than Villa Acuiia. On the Texas side, Bailey (1905:52) and Taber (1939:489) report this species upstream as far as tlie mouth of the Pecos River. The armadillo is well known to residents along the Rio Sabinas and occurs upstream from Presa Don Martin at least as far as Naci- miento. Woodlands along this river in the vicinity of Muzquiz pro- vide good habitat, where considerable underbrush is present in the parklike groves of cypress and pecan. Armadillos seem less com- mon on the Rio Salado and occur, upstream from Presa Don Martin, probably no farther than Progreso. Marsh (1937:43) reported armadillos from the Sierra del Carmen but obtained no specimens. The armadillo is of little economic value to local residents, be- cause it is not abundant in the narrow confines of the riparian habitat along the waterways. One male, recorded as Tatu novemcincttim by Elliot (1907:26) and taken by Heller and Barber at Sabinas, is the only specimen known from Coahuila. Lepus californicus Black-tailed Jack Rabbit The black-tailed jack rabbit is found on open lands in all parts of Coahuila — the Gulf Coastal Plain at 810 feet elevation to moun- tain valleys as high as 7500 feet (see specimens examined). This hare is the most conspicuous mammal of the desert plains of the State. During the heat of the day, animals may be found crouching in the shade of shrubs. Surface water seems not to be a factor in 194 University of Kansas Publs., Mus. Nat. Hist. the distribution of this species; jack rabbits hve many miles from per- manent water. The black-tailed jack rabbit is more numerous at higher elevations and in the vicinity of ranch estabhshments than elsewhere. In 1953, jack rabbits were abundant in the vicinity of the Sierra de los Hecheros; 16 specimens were obtained. Jack rabbits seem to favor places where grazing has been severe or where grasses are not abundant and cacti and low shrubs are scattered. Between Tanque Alvarez and the Sierra del Pino, where grassy cover was rank in 1952 and 1953, few jack rabbits were found. Reports were received that jack rabbits damage some fields of corn and wheat. Little local con- trol is attempted. It is consid- ered sporting at some ranches to hunt jack rabbits along the roads. The flesh of this species is not so highly esteemed as that of the cottontails by local residents. A ranch hand who accompanied our field party north of Ocampo in 1952 refused a serving of stewed jack rabbit at an evening meal but readily ate other foods offered him. He expressed sur- prise that we would eat the ani- mal. Gilmore (1947:159) found bone fragments of jack rabbit, Lepus californiciis, common in cave deposits excavated in the vicinity of Cuatro Cienegas. Average and extreme weights of Lepus from Coahuila were: for four males, 2186 (2000-2365), for ten, non-pregnant females, 2617(2300-3100). Pregnant fe- males were taken on April 2 (one with two embryos) and June 25 (one with three embryos). Females in lactation were obtained on January 9, March 17, March 20 and April 17. Lepus califomicus asellus Miller Lepus asellus Miller, Proc. Acad. Nat. Sci., Philadelphia, 51:380, September 29, 1899, type trom San Luis Potosi, San Luis Potosi. Lepus califomicus asellus Nelson, N. Amer. Fauna, 29:150, August 31, 1909. Distribution. — Southeastern Coahuila (see fig. 19). Fig. 19. Distribution in Coahuila of the black-tailed jack rabbit. 1. Lepus califomicus asellus. 2. Lepus califor- nicus nicrriami. 3. Lepus califomicus texianus. Mammals of Coahuila, Mexico 195 Remarks. — Lepus c. aseUus is paler and larger with longer ears than merriami but is not so pale as texianus and is noticeably darker on the head than either of the latter two subspecies. The ears are less buffy than those of merriami but not so gray as those of texianus. This subspecies is especially numerous in the heavily grazed moun- tain valleys south and east of Saltillo. Measurements. — Measurements of two adult males from 7 miles south and 4 miles east of Bella Union and two adult females from 4 miles south and 6 miles east of Saltillo, respectively, are: total length, 573, 575, 560, 600; length of tail vertebrae, 62, 80, 69, 70; length of hind foot, 121, 123, 128, 120; height of ear from notch (taken fresh), 143, 140, 155, 132; basilar length, 75.2, 74.5, 78.0, 73.1; zygomatic breadth, 43.3, 42.2, 42.5, 44.2; postorbital constriction, 11.9, 11.5, 11.0, 11.5; length of nasals, 39.8, 42.5, 43.6, 39.0; parietal breadth, 32.0, 30.6, 32.2, 32.9; greatest width of nasals, 19.4, 17.3, 21.1, 17.9; alveolar length of maxillary tooth-row, 16.3, 16.4, 17.5, 16.1. Specimens examined. — Total, 10, from: 3 mi. S and 3 mi. E Muralla, 3800 ft., 1 (skull only); La Pastora Rancho, 41 mi. W and 15 mi. N Saltillo, 2; 4 mi. S and 6 mi. E Saltillo, 7500 ft., 2 (1 skull only); 7 mi. S and 4 mi. E Bella Union, 7200 ft., 2 (skulls only); N foot Sierra Guadalupe, 6500 ft., 10 mi. S and 5 mi. W General Cepeda, 3 (2 skulls only). Additional records (Nelson, 1909:151): Jaral, Saltillo, Cameros, Encama- cion, La Ventura. Lepus calif ornicus merriami M earns Lepus merriami Meams, Preliminary diagnoses of new mammals from the Mexican border of the United States, p. 2, March 25, 1896 (preprint of Proc. U. S. Nat. Mus., 18:444, May 23, 1896), type from Fort Clark, Kinney County, Texas. Lepus californicus merriami Nelson, N. Amer. Fauna, 29:148, August 31, 1909. Distribution. — Open lands of northeastern Coahuila (see fig. 19). Remarks. — This subspecies occurs at least as far south as Mon- clova and west to the central mountains. Animals from the lower elevations are smaller and have shorter ears than those from the Rancho Las Margaritas in the foothills of the Serranias del Burro (see measurements). Measurements. — Average and extreme measurements of four adult males from 8-9 miles west and one mile south of Villa Acuna and six adult females from within five miles of Hacienda Las Margaritas, respectively, are: total length, 550 (536-588), 579 (560-600); length of tail vertebrae, 76 (69-83), 84 (70-93); length of hind foot, 121 (118-126), 128 (122-132); height of ear from notch (taken fresh), 120 (112-128), 134 (125-145); basilar length, 74.7 (72.5-76.0), 74.3 (69.1-76.7); zygomatic breadth, 42.9 (42.4-43.3), 43.9 (43.0-44.8); postorbital constriction, 11.9 (11.4-12.5), 13.1 (12.3-14.8); length of nasals, 40.6 (38.2-42.4), 39.6 (35.5-43.4); parietal breadth, 31.4 (31.1- 31.7), 31.7 (30.7-32.8); greatest width of nasals, 18.9 (18.0-19.9), 19.5 (17.1- 21.0); alveolar length of maxillary tooth-row, 17.0 (16.6-17.2), 16.9 (16.1- 17.5). 196 University of Kansas Publs., Mus. Nat. Hist. Specimens examined. — Total, 29, from: 9 mi. W and 1 mi. S Villa Acuna, 2 (1 skull only); 8 mi. W and 1 mi. S Villa Acuna, 2 (1 skull only); 5 rnf. W and 1 mi. S Villa Acuna, 1; Canon de Cochino, 3200 ft, 16 mi. N and 21 mi. E Piedra Blanca, 1 (skull only); 11 mi. W Hacienda San Miguel, 2200 tt., 3 (skulls only); 25 mi. S and 8 mi. E Piedra Blanca, 1; 1 mi. W Hacienda Las Margaritas, 2900 ft., 4; Hacienda Las Margaritas, 2900 ft, 3 (2 skulls only); J mi. SW Hacienda Las Margaritas, 1; 2 mi. SW Hacienda Las Margaritas, 2800 ft., 1; 5 mi. SW Hacienda Las Margaritas, 2600 ft., 2; 5 mi. E and 3 mi. S San Geronimo, 1 (skull only); 2 mi. E Nava, 810 ft., 4 (skulls only); 5 mi. SE Guerrero, 1 (skull only); La Gacha, 1600 ft., 1; 10 mi. ESE Sabinas, 1 (skull only). Additional records (Nelson, 1909:151): Sabinas; Monclova. Lepus califomicus texianus Waterhouse Lepus texianus Waterhouse, Nat. Hist. Mamm., 2:136, 1848, type probably from western Texas. Lepus califomicus texianus Nelson, N. Amer. Fauna, 29:142, August 31, 1909. Distribution. — Western Coahuila (see fig. 19). Remarks. — Lepus c. texianus is paler, larger, and longer-eared than merriami. These two subspecies intergrade in mountain val- leys of central Coahuila, but the animals from the lowlands to the east (merriami) and of the mountains to the west (texianus) are distinct. A male from the vicinity of Piedra Blanca, assigned to merriami, has small ears but is intermediate in color. Cranially, texianus averages only slightly larger than merriami. Measurements. — Measurements of two adult males from 4 miles north of Acatita and from 8 miles southeast of San Pedro de las Colonias and average and extreme measurements of six adult females from no more than 9 miles from Castillon, respectively, are: total length, 582, 517, 556 (540-573); length of tail vertebrae, 93, 76, 80 (71-90); length of hind foot, 125, 115, 122 ( 115-124); height of ear from notch (taken fresh), 141, 132, 136 ( 131-145); basilar length, 76.2, 72.4, 73.4 (72.6-74.1); zygomatic breadth, 42.8, 41.2, 42.4 (41.5-43.1); postorbital constriction, 13.1, 12.6, 11.3 (10.0-12.7); length of nasals, 40.5, 36.2, 38.5 (36.6-41.2); parietal breadth, 31.3, 29.7, 31.1 (30.0-31.8); greatest width of nasals, 21.5, 18.4, 19.1 (17.7-20.2); alveolar length of maxillary tooth- row, 17.2, 16.2, 17.1 (16.1-17.7). Specimens examined. — Total, 29, from: 8 mi. NE Castillon, 3400 ft., 1 (skull only); 9 mi. N and 6 mi. W Castillon, 4100 ft., 1 (skull only); 7 mi. N and 3 mi. W Castillon, 3800 ft., 1 (skull only); 6 mi. N and 2 mi. W Castillon, S750 ft., 5 (skulls only); 6 mi. W Castillon, 3400 ft., 1 (skull only); 5 mi. S Ace- buches, 3600 ft., 1 (skull only); 3 mi. S Castillon, 4100 ft., 1 (skull only); 4 mi. S Castillon, 4100 ft., 2 (skulls only); 5 mi. S Castilldn, 4150 ft., 1 (skull only); 44 mi. N and 15 mi. W Ocampo, 4200 ft., 1 (skull only); 12 mi. S and 2 mi. W Castill6n, 4025 ft., 1 (skull onlv); 19 mi. S and 9 mi. W Castillon, 4025 ft., 1; 21 mi. S and 10 mi. \V Castillon. 3850 ft., 1 (skull only); 30 mi. N and 7 mi. W Ocampo, 4100 ft, 1 (skull only); 22 mi. N and 5 mi. W Ocampo, 4125 ft., 1 (skull only); 18 mi. S and 14 mi. E Tanque Alvarez, 4000 ft., 1 (skull onlv); 23 mi. S and 8 mi. E Tanque Alvarez, 3200 ft., 1 (skull only); 3 mi. NE Sierra Mojada, 4100 ft., 1 (skull onlv); 8 mi. N and 25 mi. W Cuatro Cionegas, 1 (skull only); 4 mi. N Acatita, 3600 ft., 2 (skulls only); 8 mi. SE San Pedro de las Colonias, 3700 ft., 1; 10 mi. E Torreon, 3700 ft., 1; 3 mi. SE Torre6n, 3800 ft., 1. Mammals of Coahuila, Mexico 197 Sylvilagus audubonii Audubon Cottontail The Audubon Cottontail lives mostly on the desert plains and occupies arid situations where coarse desert shrubs and sparse grov%i:hs of mesquite along arroyos oflFer some protective cover. Surface water seems not to be always a necessity for this rabbit, and succulent foods, especially cacti and tender growing parts of grasses and forbs, probably sup- ply this need. This cottontail is not so abundant in tlie open des- ert plains as along the larger ar- royos and in scrub vegetation in foothills of the sierras, where ani- mals were taken at elevations as high as 8700 feet. At one place in thick mesquite brush along a wide arroyo near Villa Acuiia, both S. audubonii and S. flori- danus were taken together. Rab- bits were most active when light intensities were low, in the morn- ing and evening, and hunting at those times was most profitable. Specimens were obtained also at night with the aid of headlights. This cottontail — "conejo" of the Mexicans — is a choice food of the local residents. Females containing embryos (two and three) were taken in December, April and July; lactating females were examined in April and June. Average and extreme weights of audubonii from Coa- huila were: for 10 adult males, 688 (525-850); for 10 adult, non- pregnant females, 791 (582-855). Sylvilagus audubonii minor (Meams) Lepus arizonae minor Meams, Proc. U. S. Nat. Mus., 18:557, June 24, 1896, type from El Paso, El Paso County, Texas. S[ylvilagus]. a[uduboni]. minor Nelson, Proc. Biol. Soc. Washington, 20:83, July 22, 1907. Distribution. — Desert plains and foothills of western and southern Coahuila (see fig. 20). Fig. 20. Distribution in Coahuila of the Audubon cottontail. 1. Sylvila- gus audubonii minor. 2. Sylvilagus audubonii parvulus. 198 University of Kansas Publs., Mus. Nat. Hist, RemarJcs. — Coahuilan cottontails of this species, except those of the Coastal Plain of the northeastern part, are assigned to the sub- species S. a. minor. Pale pelage and large auditory bullae distin- guish minor from S. a. parvultis. In minor the length of the auditory bullae is usually more, instead of usually less (as in parvulus), than 26 per cent of the basilar length. Measurements. — Average and extreme measurements of five adult females, all from no more than 9 miles from Castillon, are: total length, 358 (347-365); length of tail vertebrae, 45 (30-53); length of hind foot, 82 (78-85); height of ear from notch (in the flesh), 72 (68-75); basilar length, 49.8 (48.8-50.5); zygomatic breadth, 32.3 (31.4-33.6); postorbital constriction, 11.7 (11.1-12.5); length of nasals, 27.3 (26.7-27.9); length of auditory bullae, 13.9 (13.2-14.5); greatest width of nasals, 13.0 (12.3-13.4); alveolar length of maxillary tooth- row, 12.1 (11.9-12.6). Specimens examined. — Total, 26, from: 10 mi. S and 5 mi. E Boquillas, 1500 ft., 1; 3 mi. S and 7 mi. E Hechicero, 4600 ft., Chihuahua, in Coahuila, 1; 1 mi. SE El Pino, 1; 9 mi. N and 6 mi. W Castillon, 1; 6 mi. N and 2 mi. W CastiUdn, 3760 ft., 3; 1 mi. N and 3 mi. W Castillon, 3800 ft., 1; 3 mi. SSE Castillon, 4050 ft., 1; 6 mi. S Castillon, 4150^ ft., 1; 13 mi. N and 11 mi. W Tanque Alvarez, 4250 ft., 1; 1 mi. S San Lazaro, 1; La Pastora Rancho, 41 mi. W and 15 mi. N Saltillo, 1; 10 mi. E Torreon, 3700 ft., 1; 12 mi. N and 10 mi. E Parras, 3850 ft., 1; Diamante, 7500 ft., 4 mi. S and 6 mi. E Saltillo, 1; 7 mi. S and 4 mi. E Bella Union, 7200 ft., 1; 2 mi. E and 2 mi. N San Antonio de las Alazanas, 8700 ft., 1; N foot Sierra de Guadalupe, 6200 ft., 10 mi. S and 5 mi. W General Cepeda, 5; W foot Pico de Jimulco, 5000 ft., 2; 7 mi. S and 1 mi. E Gomez Farias, 6500 ft., 1. Additional records (Nelson, 1909:237): Saltillo, La Ventura. Sylvilagus audubonii parvulus (J. A. Allen) Lepus (Sylvilagus) parvulus J. A. Allen, Bull. Amer. Mus. Nat. Hist., 20:34, February 29, 1904, type from Apam, Hidalgo. Sylvilagus auduboni parvulus Nelson, N. Amer. Fauna, 29:236, August 31, 1909. Distribution. — Coastal Plain in northeastern Coahuila (see fig. 20). Remarks. — Gilmore (1947:159) records bone fragments from cave deposits from the vicinity of Cuatro Cienegas as Sylvilagus audu- bonii, identifying the material as the species audubonii chiefly on geographic grounds. Audubon cottontails found in the vicinity of Cuatro Cienegas would probably be of the subspecies parvulus. Measurements. — Measurements of 3 adults (2 $ , 1 9 ) from 1 mile south and 9 miles west of Villa Acuiia and 1 adult female from 11 miles west of the Hacienda San Miguel, respectively, are: total length, 372, 328, 336, 354; length of tail vertebrae, 38, 50, 40, 46; length of hind foot, 78, 61, 81, 83; height of ear from notch (in the flesh), 63, 62, 67, 68; basilar length, 51.9, 50.4, 50.2, 49.5; zygomatic breadth, 32.8, 33.1, — , 32.8; postorbital constriction, 12.2, 11.5, 11.4, 11.7; length of nasals, 28.1, 28.9, 27.6, 28.1; length of auditory bullae, 12.8, 13.3, 12.5, 12.4; greatest width of nasals, 14.3, 14.7, 12.6, 13.7; alveolar length of maxillary tooth-row, 11.9, 11.7, 12.2, 12.1. Specimens examined. — Total, 15, from: 1 mi. S and 9 mi. W \'illa Acuiia, Mammals of Coahuila, Mexico 199 J^ 10! 00_ 9 50 , KO scale of milci 55 lij B? 7; 11 mi. W Hacienda San Miguel, 2200 ft., 1; 15 mi. N and 8 mi. W Piedras Negras, 1; 4 mi. N Piedras Negras, 2; La Gacha, 1600 ft., 2; 1 mi. E La Gacha, 1600 ft., 1; 10 mi. ESE Sabinas, 1. Additional records (Nelson, 1909:237): Las Vacas Creek (head of), Sa- binas, Monclova. Sylvilagus floridanus Eastern Cottontail The eastern cottontail occupies the riparian association along major streams and their larger tributaries in northeastern Coahuila and the oak and pine associations of the mountains. The range of the smaller lowland subspecies, S. /. chapmani, does not come in contact with those of the larger mountain subspecies. S. /. nelsoni and S. /. orizahae. In Coahuila the eastern cottontail is wary and rarely ventures as far as 20 feet from brush or other protective cover. On the Coastal Plain the eastern cottontail lives in the narrow strip of mesic growth which borders streams and arroyos. Both S. jiori- danus and S. audubonii occur to- gether in this situation although audubonii occurs also on the ad- jacent desert plains. The east- ern cottontail is found upstream along arroyos to where they en- ter the foothills of the mountains; in the Serranias del Burro, chap- mani of the lowlands lives along the Rio Alamos at an elevation of 3800 feet. This species ranges upstream along the Rio Grande probably no farther than the vi- cinity of Villa Acufia, since de- sirable streamside habitat is pro- gressively reduced as the river becomes more entrenched up- stream. Large cottontails probably are found on most high mountains in Coahuila where there is ex- tensive oak and pine habitat. Specimens are available from the Sierra de la Madera, Sierra Madre Oriental, and the Sierra En- carnacion in southeastern Coahuila. Reports, but no specimens, of large cottontails were obtained in the Sierra del Carmen, Sierra de la Encantada, Sierra de los Hecheros, Sierra del Pino, Sierra de Fig. 21. Distribution in Coahuila of the eastern cottontail, Sylvilagus flori- danus. 1. S. /. chapmani, 2. S. /. nelsoni. 3. S. /. orizahae. 200 University of Kansas Fuels., Mus. Nat, Hist. San Marcos. Marsh (1937:38) records a large cottontail at 8500 feet in the Sierra del Carmen. Nelson (1909:195) also reports a large cottontail in the Sierra de Guadalupe. These mountain cotton- tails seemingly exist in small numbers and remain concealed in brushy cover most of the day. Specimens were taken in late eve- ning or at night with the aid of a headlight. Hunting usually was done where rabbit droppings were observed. Females containing embryos were taken on March 25 (one with three embryos ) , April 4 ( one with two embryos ) , April 9 ( one with three embryos ) , May 22 ( one with four embryos ) , and June 21 ( one with three embryos). Females in lactation were taken on April 6 ( one ) and April 7 ( one ) . An adult, non-pregnant female from near Villa Acuna (chapmani) weighed 745.9 grams; one from 21 miles south and 4 miles west of Ocampo and in the Sierra de la Madera (nelsoni) weighed 1200 grams. A pregnant female from 10 miles east of San Antonio de las Alazanas and in the Sierra Madre Oriental (orizabae) weighed 1300 grams. Sylvilagus floridanus chapmani (J. A. AUen) Lepus -floridanus chapmani J. A. Allen, Bull. Amer. Mus. Nat. Hist., 12:12, March 4, 1899, type from Corpus Christi, Nueces County, Texas. Sylvilagus floridanus chapmani Lyon, Smithsonian Misc. Coll., 45:336, Jxme 15, 1904. Distribution. — Along streams and arroyos on the coastal plain of northeastern Coahuila (see fig. 21). Remarks. — This subspecies is small and brown and strikingly dif- ferent from the much larger and grayer montane subspecies. Exam- ples of chapmani from along arroyos in the foothills ( as high as 3900 feet ) are larger than those from along the Rio Grande but not nearly so large as the subspecies in the mountains. There is no evidence of intergradation between these subspecies. Measurements. — One adult male from 2 miles west of Jimenez and another from 3 miles north and 1 mile east of Allende measure: total length, 382, 365; length of tail vertebrae, 47, 60; length of hind foot, 89, 80; height of ear from notch (in the flesh), 60, 58; basilar length, 50.1, 46.5; zygomatic breadth, 32.0, 31.0; postorbltal constriction, 12.0, 12.8; length of nasals, 27.9, 25.9; length of auditory bullae, 10.6, 10.1; parietal breadth, 25.2, 24.6; greatest width of nasals, 13.6, 10.8; alveolar length of maxillary tooth-row, 12.7, 13.1. Specimens examined. — Total, 14, from: 1 mi. S and 9 mi. W Villa Acufia, 1 (skull only); 2 mi. W Jimenez, 850 ft., 3; El Rio Alamos, in Canon Mulato, Rancho Las Margaritas, 3800 ft., 1 (skull only); 12 mi. S Hacienda Las Mar- garitas, 2300 ft., 1; 20 mi. E Nava, 1; 3 mi. N and 1 mi. E Allende, 1000 ft., 1; 5 mi. SE Guerrero, 2 (1 skull only); 1 mi. W Hacienda La Mariposa, 1; La Gacha, 1600 ft., 1 (skull only); 2 mi. S and 3 mi. E San Juan de Sabinas, 1160 ft., 1; Nadadores, 1 (skull only). Additional records (Nelson, 1909:178): Sabinas, Monclova. Mammals of Coahuila, Mexico 201 Sylvilagus floridanus nelsoni Baker Sylvilagus floridanus nelsoni Baker, Univ. Kansas Publ., Mus. Nat. Hist., 7: 611, April 8, 1955, type from 22 mi. S and 5 mi. W Ocampo, 5925 ft., Coahuila. Distribution. — Oak and pine associations of the Sierra de la Madera and probably other mountains in central and northern Coahuila (see fig. 21). Remarks. — The large, grayish cottontail of the Upper Sonoran and Transition life-zones may occur in most high mountains in the north- ern half of the state. Reports of a large mountain rabbit, probably assignable to nelsoni, have been obtained in the Sierra de San Marcos, Sierra del Pino, Sierra de los Hecheros, Sierra de la Encan- tada and Sierra del Carmen. The subspecies nelsoni is grayer and larger in all respects than chapmani. There is no evidence of intergradation between these two subspecies. Measurements. — Two adult females from 21-22 miles south and 4-5 miles west of Ocampo measure: total length, 421, 405; length of tail vertebrae, 51, 48; length of hind foot, 97, 98; height of ear from notch (taken fresh), 72, 76; basilar length, 57.9, 56.2; zygomatic breadth, 35.9, 34.1; postorbital constriction, 11.0, 11.1; length of nasals, 35.1, 30.3; length of auditory bullae, 12.8, 12.5; parietal breadth, 26.8, 26.7; greatest width of nasals, 16.3, 16.1; alveolar length of maxillary tooth-row, 13.7, 13.4. Specimens examined. — Total, 4, from: 20 mi. S and 4 mi. W Ocampo, 5300 ft., 2; 21 mi. S and 4 mi. W Ocampo, 5700 ft., 1; 22 mi. S and 5 mi. W Ocampo, 5925 ft., 1. Sylvilagus floridanus orizabae ( Merriam ) Lepus orizabae Merriam, Proc. Biol. Soc. Washington, 8:143, December 29, 1893, type from Volcan de Orizaba, Puebla. Sylvilagus floridanus orizabae Nelson, N. Amer. Fauna, 29:183, August 31, 1909. Distribution. — Mountains of southeastern Coahuila (see fig. 21). Remarks. — The subspecies orizabae occurs in the Sierra Madre Oriental southeast of Saltillo, in the Sierra Encarnacion in extreme southeastern Coahuila and probably in the Sierra de Guadalupe, where Nelson (1909:195) reported the presence of a large cotton- tail. This subspecies can be distinguished from nelsoni by its darker, browner color. In size the two cottontails are similar. Measurements. — An adult female from 10 miles east of San Antonio de las Alazanas measures: total length, 441; length of tail vertebrae, 54; length of hind foot, 92; height of ear from notch (in the flesh), 71; basilar length, 56.0; zygomatic breadth, 34.8; postorbital constriction, 12.3; length of nasals, 31.4; length of auditory bullae, 12.3; parietal breadth, 26.5; greatest width of nasals, 15.8; alveolar length of maxillary tooth-row, 13.1. Specimens examined. — Total, 3, from: 10 mi. E San Antonio de las Alazanas, 8700 ft., 1; IS mi. E San Antonio de las Alazanas, 1; Sierra Encarnacion, 9000 ft., 1 (BSC). 202 University of Kansas Publs., Mus. Nat. Hist. Cynomys mexicanus Merriam Mexican Prairie Dog Cynomys mexicanus Merriam, Proc. Biol. Soc. Washington, 7:157, July 27, 1892, type from La Ventura, Coahuila. Distribution. — Extreme southeastern Coahuila (see fig. 22). Remarks. — The Mexican prairie dog was discovered by Dr. Ed- ward Palmer, who (in J. A. Allen, 1881:185) wrote, "Only a single small colony was met with, in a little valley surrounded by moun- tains, not far from Saltillo, confined to an area of some thirty or forty acres." It is Hkely that Palmer found these animals east- southeast of Saltillo in the vicinity of San Antonio de las Alazanas, At La Ventura in March, 1891, Clark P. Streator obtained ten speci- mens which were described by Merriam ( 1892:157-158) as Cynomys mexicanus. Unlike Palmer, Streator found the animals abundant at La Ventura and described (in Merriam, loc. cit.) the colony as the largest one that he had ever seen. Nelson and Goldman visited La Ventura in July and August, 1896, and preserved 27 additional specimens. Goldman (1951:130) described the area around La Ventura as in interior basinlike plain which was known as the "Llano de los Perros" because of the presence of this large colony of prairie dogs. In July and November, 1949, W. Kim Clark ob- tained several prairie dogs in the vicinity of La Ventura for the University of Kansas. His field notes indicate that these animals are still numerous in this area. The Mexican prairie dog, locally called "perro llanero" or "per- rito," is confined to an altitudinal range from 5500 feet to about 7000 feet in broad valleys and intermontane basins in southeastern Coa- huila, adjacent parts of Nuevo Leon, San Luis Potosi and probably in Zacatecas. The species seems to be most numerous in the area from Gomez Farias south to La Ventura. Southeast of Saltillo, in the broad mountain valleys, these animals are less numerous chiefly because less habitat is available. In 1950, one such colony, in this area, extended approximately three miles in an east-west direction but was less than 400 yards wide. The preferred habitat of these animals is deep, rockfree soils. At San Antonio de las Alazanas I was told that the prairie dogs are eaten as food. Farmers consider the animals to be pests and control measures have reduced the colonies, especially where wheat and other crops are planted. Specimens taken in March and April are clothed in heavy pelage, Mammals of Coahuila, Mexico 203 which has considerable underfur especially on the back and rump. According to Hollister (1916:22) a spring pelage closely resembles the winter pelage. Specimens taken in June are in a much worn pel- age and are molting on the head and in spots over the back where shorter, stiffer hairs of the sum- mer coat appear. This molt was found also in animals taken in July and early August. Animals secured in September show the beginnings of a growth of the heavier winter pelage on the up- per parts. In November animals are in typical winter pelage. A female, taken on March 25 at 3 miles north and 4 miles west of San Antonio de las Alazanas, contained 3 embryos averaging 23 mm. in crown to rump length. She weighed 746.2 grams. The Mexican prairie dog is a large black-tailed species resem- bling closely C. ludovicianus (see Merriam, 1892:157-158 and Hol- lister, 1916:21-23). Owing to the isolation of C. mexicanus (ap- proximately 330 miles separates its range from that of C ludovicianus), and the degree of difference between it and ludovicianus, the Mexican prairie dog is considered specifically distinct. Cynomys mexicanus differs more from either subspecies of C. ludovicianus than the subspecies differ from each other. Measurements. — An adult male from 3 miles north of Gomez Farias meas- ures: total length, 440; length of tail, 120; length of hind foot, 63; height of ear (from notch), 15; greatest length of skull, 65.4; zygomatic breadth, 45.3; mastoidal breadth, 31.1; lengtli of nasals, 23.4; postorbital breadth, 14.6; alveo- lar length of upper molariform tooth-row, 15.3. Specimens examined. — Total, 35, from: 7 mi. S and 4 mi. E Bella Union, 7200 ft., 4; 3 mi. N and 4 mi. W San Antonio de las Alazanas, 1; 13 mi. W San Antonio de las Alazanas, 4; 12 mi. W San Antonio de las Alazanas, 13; 8 mi. W San Antonio de las Alazanas, 7000 ft., 4; 7 mi. W San Antonio de las Ala- zanas, 1; 4 mi. E and 2 mi. S Agua Nueva, 6800 ft., 4; 3 mi. N Gomez Farias, 7000 ft., 2; 2 mi. W San Miguel, 5500 ft., 1; 8 mi. N La Ventura, 5500 ft., 1. Additional records (Hollister, 1916:23) :Saltillo [= approximately 15 miles ESE Saltillo]; La Ventura. Fig. 22. Distribution of the Me.xican prairie dog, Cynomys mexicanus, in Coahuila. 204 University of Kansas Publs., Mus. Nat. Hist. Spermophilus mexicanus parvidens Meams Mexican Ground Squirrel Spermophilus mexicanus parvidens Meams, Preliminary diagnoses of new mammals from the Mexican Border of the United States, p. 1, March 25, 1896 (preprint of Proc. U. S. Nat. Mus., 18:443, May 23, 1896), type from Fort Clark, Kinney County, Texas. Distribution. — Arid plains and foothills of eastern Coahuila (see map, fig. 23). Remarks. — The Mexican ground squirrel lives in the lowlands of eastern Coahuila and is numerous in the northeastern part of the State where soils are not rocky. Probably this animal reached south- eastern Coahuila, from Nuevo Leon to the eastward, by way of the watershed of the Rio Salinas rather than from the northward. Speci- mens were taken from as low as 800 feet in elevation (Don Martin) to as high as 6500 feet ( north foot of the Sierra de Guadalupe ) , This species frequently lives on desert flats supporting prickly pear and some grass. Animals were ob- tained by trapping or by luring them to within gunshot range with a squeaking call. Allen (1881:184) records a specimen from Monclova taken by Dr. Ed- ward Palmer, who wrote (in Allen, 1881:184) that the species is "widely distributed at favor- able localities, but not nearly so abundant as the small species [= S. spilosoma]." Marsh (1937: 37 ) found this ground squirrel at Piedra Blanca, which is near the western limit of the range of the species in this part of Coahuila. One pregnant female, contain- ing 5 embryos, was obtained on June 16. Average and extreme weights of three adult males Fig. 23. Distribution of the Mexican ground squirrel, S})cr)7io))hilus mexi- canus parvidens, in Coahuila. from near Villa Acufia, Hacienda San Miguel and Piedras Negras were 186.6 (145.2-210.2); of two adult nonpregnant females from 1 mile soutli and 9 miles west of Villa Acuiia, 126.4 and 135.5. Bone fragments from cave deposits from the vicinity of Cuatro Cienegas, identified as Citellus [= Spermophilus] mexicanus by Gil- Mammals of Coahuila, Mexico 205 more (1947:157), on geographic grounds, here are referred to this subspecies. Measurements. — Of specimens from northeastern Coahuila, average and ex- treme measurements of 4 adult males (KU 35719, 35723, 35725, 48442) and 4 adult females (KU 35720, 35722, 35724, 48444), are: total length, 301 (286- 312), 283 (276-286); length of tail vertebrae, 119 (106-127), 116 (115-120); length of hind foot, 40 (38-42), 39 (37-40); height of ear from notch, 12 (11- 14), 12 (11-13); greatest length of skull, 43.0 (42.5-43.5), 42.0 (41.4-42.8); palatilar length, 20.2 (20.1-20.3), 19.9 (19.7-20.1); zygomatic breadth, 25.4 (24.5-26.1), 23.9 (23.7-24.4); cranial breadth, 18.9 (18.6-19.3), 17.9 (17.8- 18.1); least interorbital breadth, 9.6 (9.2-10.1), 8.8 (8.3-9.0); postorbital breadth, 13.5 (13.1-13.9), 12.6 (12.0-13.3); length of nasals, 14.8 (14.4-15.2), 14.5 (14.1-14.9); length of maxillary tooth-row, 8.5 (8.3-8.6), 8.1 (7.9-8.3). Specimens examined. — Total, 43, from: 2/2 mi. W Villa Acuna, 1; Las Vacas [= Villa Acuiia], 7 mi. E, 1 (BSC); 1 mi. S and 9 mi. W Villa Acuiia, 3; 13 mi. W San Miguel, 2200 ft., 1; 15 mi. N and 8 mi. W Piedras Negras, 1; 2 mi. 5 and 11 mi. E Nava, 810 ft, 3; La Gacha, 2000 ft, 1; Sabinas, 1 (BSC); Don MarU'n, 800 ft, 5; 5 mi. N and 2 mi. W Monclova, 1; Monclova, 1 (BSC); 5 mi. W Trevifio, 3000 ft., 1; 1 mi. N Saltillo, 5000 ft., 1; Saltillo, 8 (BSC); 1% mi. N Parras, 5500 ft., 1; N foot Sierra Guadalupe, 6200-6500 ft., 10 mi. S and 5 mi. W General Cepeda, 13. Additional records (Allen, 1881:184): Monclova; (Gihnore, 1947:157): vicinity of Cuatro Cienegas. Spermophilus spilosoma pallescens (A. H. Howell) Spotted Ground Squirrel Citellus spilosoma pallescens A. H. Howell, Proc. Biol. Soc. Washington, 41:212, December 18, 1928, type from La Ventura, Coahuila. Distribution. — Western and southern Coahuila (see fig. 24). Remarks. — The spotted ground squirrel hves both on the arid des- erts of western Coahuila and in the open, mountain valleys of the southeastern part of the State. Animals have been taken from as low as 2450 feet in elevation to as high as 7000 feet. The species prefers areas of deep soil and seems to avoid rocky terrain. This ground squirrel is uncommon on the desert plain; usually only one or two specimens were obtained at any one place. The animal is wary and when disturbed often eludes the collector by rapidly mov- ing away. In the elevated valleys of southeastern Coahuila, this ground squirrel is more common; it seems to prefer the open places that have been grazed severely by goats and is less wary than in the desert and sits near its burrow alongside the road as vehicles pass. This squirrel eats both grasses and forbs as well as wheat and irri- gated crops. Palmer (in Allen, 1881:185) found this ground squirrel at San Pedro and near Parras and wrote that it "Lives on the open plains and about the edges of fields, where it is a troublesome pest." 206 University of Kansas Publs., Mus. Nat. Hist. Specimens were obtained in January, March, April, May, June, July, August, September and November; probably this species is active throughout the year. In Novem- ber, January, March and April only young were seen. Young are active all year but adults hibernate (see Dalquest, 1953: 93). Pregnant females were ob- tained on June 25 (four females having 4, 4, 4, and 6 embryos), July 1 (one female with 3 em- bryos), July 6 (one female with 4 embryos ) , July 22 ( one female with 7 embryos ) . Young ground squirrels, less than 170 mm. in total length, were trapped on July 18. Locally this ground squirrel is called "ardillon." Specimens from western Coa- huila are assigned to paUescens but some, especially some young individuals, are either darker or lighter than adults from south- eastern Coahuila from the vicinity of tlie type locality. The spot- ting is less conspicuous in some animals than in others. An adult female from the vicinity of Sierra Mojada has a short, broad ros- trum— a resemblance to S. s. canescens (Merriam) of western Chi- huahua. Bone fragments from near Cuatro Cienegas, identified as C. spilosoma by Gilmore (1947:157), are here assigned, on geo- graphic grounds, to S. 5. paUescens. Measurements. — Average and extreme measurements of five adult females and of three adult males from 7 miles south and 4 miles east of Bella Uni6n, respectively, are: total length, 257 (251-265), 250 (248-253); length of tail vertebrae, 81 (79-85), 82 (78-84); length of hind foot, 36 (32-39), 36 (35- 38); height of ear from notch, 12 (11-12), 11 (10-12); greatest length of skull, 42.1 (41.5-43.1), 41.7 (40.0-42.7); basilar length, 33.7 (33.1-34.1), 33.3 (32.2-34.0); palatilar length, 19.0 (18.3-19.3), 19.2 (18.3-19.9); zygomatic breadth, 24.8 (24.1-25.6), 25.3 (23.8-26.1); cranial breadth, 18.9 (18.5-19.6), 18.9 (18.7-19.3); least interorbital constriction, 9.1 (9.0-9.6), 9.3 (8.9-10.0); postorbital constriction, 14.4 (14.1-15.2), 14.8 (14.6-15.1); length of nasals, 14.8 (14.3-15.3), 14.3 (13.3-15.2); alveolar length of maxillary tooth-row, 8.3 (8.1-8.5), 8.1 (7.9-8.3). Fig. 24. Distribution ot the spotted ground squirrel, Spermophihis spilo- soma paUescens, in Coahuila. Mammals of Coahuila, Mexico 207 Specimens examined. — Total, 46, from: 2 mi. S and 3 mi. E Hechicero, 4450 ft., Chihuahua, in Coahuila, 2 (one skin only); 3 mi. SW Carricitos, 3500 ft., 1; 11 mi. E Acebuches, 1; 23 mi. N and 5 mi. W Ocampo, 4125 ft., 1; 18 mi. S and 14 mi. E Tanque Alvarez, 4000 ft., 1; 19 mi. S and 8 mi. E Tanque Alvarez, 1 (skull only); 23 mi. S and 8 mi. E Tanque Alvarez, 3200 ft., 1; 3 mi. NE Sierra Mojada, 4 ( one skull only ) ; 5 mi. N and 19 mi. W Cuatro Ciene- gas, 3250 ft., 2; 3 mi. NW Cuatro Cienegas, 2450 ft., 1; 4 mi. N Acatita, 3600 ft., 6 (four skulls only); Santa Cruz, 25 mi. N and 7 mi. W Saltillo, 1; 12 mi. N and 10 mi. E Parras, 5000 ft., 1; 7 mi. S and 4 mi. E Bella Union, 7200 ft., 12 (one skull only, two skeletons only); % mi. S Encantada, 5500 ft., 5 (one skull only); 8 mi. W San Antonio de las Alazanas, 7000 ft., 2; 7 mi. S and 1 mi. E Gomez Farias, 6500 ft., 2; 8 mi. N La Ventura, 5500 ft., 2. Additional records (Gilmore, 1947:157): vicinity of Cuatro Cienegas; (Howell, 1938:125): Jaral, Torreon, Cameros, La Ventura; (J. A. Allen, 1881: 185): Parras. Spermophilus variegatus couchii Baird Rock Squirrel Spermophilus couchii Baird, Proc. Acad. Nat. Sci. Philadelphia, vol. 7 ( 1854- 55), p. 332, April 1855, type from Santa Catarina, Nuevo Leon. Distribution. — In suitable habitat in western and southern Coahuila (see fig. 25). Remarks. — The rock squirrel, appropriately named because of its preference for rocky situations, occurs in most parts of Coahuila except the Coastal Plain. Specimens were obtained as low as 1220 feet in elevation and as high as 8600 feet. This squirrel Hves in rocky outcrops in mountains and along arroyos but utilizes also stone fences. Canons in mountainous areas are likely places to find rock squirrels especially in the Upper Sonoran Life-zone of the Sierra del Carmen, Sierra de Guadalupe, and the Sierra Madre Oriental. Occasionally variegatus was seen feeding in oak trees, but most individuals taken were on the ground or in rocky outcrops. In the Sierra del Carmen, Marsh (1937:36) saw variegatus eating fruits of cherry and madrono and found acorns and hackberry seeds in caches under rocks. Both the rock squirrel and Sciurus alleni occur together in the Sierra de Guadalupe and the Sierra Madre Oriental, and these two species may compete for acorns and other foods. Because rock squirrels are confined to rocky areas, they do little damage to forage for domestic livestock or to corn or other crops. Stone fences, a favored abode of variegatus, rarely are constructed in Coahuila to enclose crop-lands. In other Mexican states to the southward where stone fences surround many fields, damage to corn by these squirrels is reported (see Dalquest 1953:95). The upper parts are mottled with pale buff or dark brownish 208 University of Kansas Publs., Mus. Nat. Hist. black. Of 30 specimens examined from Coahiiila, one-half are melanistic. Howell (1938:140) writes that "At Saltillo, Coahuila, where 12 specimens were taken, no black ones were seen, but in the canyons of the Sierra Guada- lupe more than half of the ani- mals seen were in the melanistic phase." Residents eat rock squirrels to- day and did so in earlier times, as shown by Gilmore's (1947: 156) finding 145 fragments of bone in caves formerly occupied by Indians near Cuatro Ciene- gas. Average and extreme weights of variegatus from Coahuila for four adult males were 621 (470- 875); for five adult, non-preg- nant females, 546 (450-634). One pregnant female containing five embryos was taken on April 8. Bone fragments from cave de- posits from near Cuatro Ciene- gas, identified by Gilmore ( 1947: 156) as Citellus [= Spermophilus] variegatus, are assigned here on geographic grounds, to couchii. Measurements. — Measurements of two adult males the first from the north foot of the Sierra Guadalupe and tlie second from 23 miles south and 5 miles west of Ocampo, and average and extreme measurements of eight adult females from eastern and southern Coahuila (KU34571, 35726, 35727, 55514, 55515, 55519, 55525, 58011), respectively, are: total length, 420, 456, and 437 (400- 480); length of tail vertebrae, 194, 174 and 174 (150-197); lengtli of hind foot, 59, 61 and 56 (50-61); height of ear from notch, 25, 26 and 26 (22-30); con- dylobasal length, 59.9, 59.7 and 59.2 (56.7-61.5); basilar length, 48.7, 49.1 and 48.6 (46.4-51.0); palatilar length, 28.9, 28.8 and 28.5 (27.3-29.2); zygo- matic breadth, 36.2, 38.2 and 36.5 (35.6-38.2); cranial breadth, 23.6, 25.6 and 24.0 (23.3-25.2); postorbital constriction, 16.4, 18.3 and 16.2 (15.2-18.2); length of nasals, 22.6, 20.6 and 20.6 (19.8-21.3); alveolar length of maxillary tooth-row, 13.1, 12.9 and 12.1 (11.5-12.7). Specimens examined. — Total, 30, from: Club Sierra del Carmen, 4950 ft., 2 mi. N and 6 mi. W Piedra Blanca, 2; 23 mi. S and 5 mi. W Ocampo, 7000 ft., 1; Panuco, 1220 ft., 3; lYi mi. N Parras, 5500 ft., 1; 1 mi. S and 4 mi. W Bella Uni6n, 7000 ft., 5; 3 mi. S and 3 mi. E Bella Union, 6750 ft., 1; Encantada, 5000 ft., 1; Mesa de Tablas, 8600 ft., 1; N foot Sierra Guadalupe, 6700 ft., 10 mi. S and 5 mi. W General Cepeda, 2; N foot Sierra Guadalupe, 6500 ft., 10 Fig. 25. Distribution of the rock squirrel, Spermophilus variegatus couchii in Coahuila. Mammals of Coahuila, Mexico 209 mi. S and 3 mi. W General Cepeda, 5; N foot Sierra Guadalupe, 6400 ft., 11 mi. S and 4 mi. W General Cepeda, 8. Additional records (Gilmore, 1947:156): vicinity of Cuatro Cienegas; (Howell, 1938:141): Saltillo, Sierra Guadalupe, Cameros, Sierra Encamacion. Ammospermophilus interpres (Merriam) Texas Antelope Squirrel Tamias interpres Merriam, N. Amer. Fauna, 4:21, October 8, 1890, type from El Paso, El Paso County, Texas. Ammospermophilus interpres Bailey, N. Amer. Fauna, 25:81, October 25, 1905. Distribution. — Rocky situations, chiefly at lower elevations, in western Coa- huila (see fig. 26). Remarks. — The Texas antelope squirrel occurs in rocky situations usually in the driest parts of Chihuahuan Desert Shrub Vegetation- type. The animal prefers low hills covered witli lechuguilla and topped by massive rimrock. This squirrel was wary and difficult to approach. Before a person was within gunshot range, it would dash out of sight on a rock slide. In a talus slope in the foothills of the Sierra de los Hecheros, one interpres escaped me in this manner. Marsh (1937:37) writes that he saw two of these squirrels in rock slides in the Sierra del Carmen. Taylor et al. (1945:25) saw one in the Carmens at 3000 feet elevation in the Caiion de la Jardin Viejo. Gilmore (1947:157) identified as A. interpres nine bone frag- ments from cave deposits from near Cuatro Cienegas. In the vicinity of Boquillas the Texas antelope squirrel was frequently seen in early March, 1952. Evidently this species is active in winter: Borell and Bryant (1942:21) record individuals in November and on Decem- ber 1 in Brewster County, Texas. In the University of Kansas Mu- seum of Natural History there are two interpres (51990-91 KU) taken on January 10 at 3 miles south and 6 miles east of Presidio, Texas, and a third (58966 KU) obtained on February 17 at a place 3 miles west of Tijeras in Bernalillo Co., New Mexico. Borell and Bryant {loc. cit.) remark that this species may be inactive only in coldest weather. The winter pelage of interpres is dense and soft and underfur is abundant; the over-all color is bright and tawny. The summer pel- age is shorter and harsher, has little underfur and is pale gray. Speci- mens in winter pelage were taken on March 2 and 3; one in summer pelage was taken on May 4. Specimens of interpres in summer pelage were obtained on July 10 and 13 less than ten miles from the Coahuilan border in Durango (3 mi. SW Lerdo and 4 mi. WSW Lerdo ) . Average and extreme weights of four adult, nonpregnant females 210 University of Kansas Publs., Mus. Nat. Hist. from no more than 10 miles from Boquillas were 110.2 (98.7- 121.9). Lactating females were taken on March 2 and 3. Specimens in winter pelage from Coahuila resemble examples of interpres in the same pelage taken in March in Bernarlillo County, New Mexico, and in January in Presidio County, Texas, except that the coloring on the sides, flanks and legs is paler and the upper parts are grayer and paler. Measurements. — Average and ex- treme measurements of four adult fe- males from no more than 10 miles from Boquillas are: total length, 227 (219-235); length of tail vertebrae, 67 (51-75); length of hind foot, 37 (36-39); height of ear from notch, 14 (14-15); greatest length of skull, 40.0 (39.1-40.5); condylobasal length, 39.5 (38.4-40.1); basilar length, 31.9 (30.8- 32.8); palatilar length, 18.2 (17.7- 18.5); zygomatic breadth, 22.9 (22.0- 23.3); cranial breadth, 18.4 (17.8- 18.7); least interorbital constriction, 9.4 (8.9-9.8); postorbital constriction, 13.6 (13.0-14.3); length of nasals, 12.1 (11.2-12.6); alveolar length of maxillary tooth-row, 6.9 (6.7-7.1). Specimens examined. — Total, 5, from: 7 mi. S and 2 mi. E Boquillas, 1800 ft., 1; 10 mi. S and 5 mi. E Boquillas, 1500 ft., 3; 15 mi. S El Hundido, 1. Additional records (Gilmore, 1947:157): 22 mi. W and 4 mi. S Cuatro Cienegas; (Howell, 1938:181): Jaral. Eutamias bulleri solivagus A. H. Howell BuUer Chipmunk Eutamias buUeri solivagus A. H. Howell, jour. Mamm., 3:179, August 4, 1922, type from Sierra Guadalupe, Coahuila. Distribution. — Mesic forests of extreme southeastern Coahuila (see fig. 27). Remarks. — The Buller chipmunk lives in montane mesic forests of the Sierra de Guadalupe and the Sierra Madre Oriental of south- eastern Coahuila. Specimens were obtained in stands of pine, fir and aspen at elevations no lower tlian 9000 feet. Chipmunks, taken in April, were trapped under moss-covered rock ledges along a small mountain stream under a dense canopy of coniferous trees. W, Kim Fig. 26. Distribution of the Texas an- telope squirrel, Ammospermophilis in- terpres, in Coahuila. Mammals of Coahuila, Mexico 211 Clark found chipmunks in the same area much more abundant in late July and early August. According to Howell (1929:105), Nel- son and Goldman regarded Buller chipmunks as ". . . com- mon [in May] in the coniferous forest on top of the Sierra de Guadalupe, above La Concor- dia." Weights of bulleri from east of San Antonio de las Alazanas are: for two adult males, 70.0 and 68.4 grams and for two adult, non-pregnant females, 88.0 and 79.5 grams. Females in lacta- tion were taken on July 28 and August 3. Differences between solivagus and other subspecies, E. b. bul- leri (Allen) and E. b. durangae Allen, are not extreme. Also, E. b. solivagus differs little from E. c. cinereicollis (Allen) from New Mexico. Further study may reveal that bulleri and cinerei- collis should be arranged as one species. Howell (1929:103) of- fers no explanation for their separation as two species except that ". . . there is a considerable gap between the ranges of the sub- species of bulleri and of cinereicollis." White ( 1953:627), however, reports that the baculum of bulleri is distinct from those of all other chipmunks of western North America. Measurements. — Two adult males and two adult females, all from 12 miles east of San Antonio de las Alazanas, respectively, measure: total length, 220, 226, 230, 230; length of tail vertebrae, 90, 95, 86, 95; length of hind foot, 30, 34, 34, 31; height of ear from notch, 19, 20, 20, 18; greatest length of skull, 38.4, 36.5, 37.5, 37.0; condylobasal length, 37.0, 35.5, 36.0, 35.8; basilar length, 29.2, 28.4, 28.7, 28.6; palatilar length, 16.2, 15.8, 15.9, 15.8; zygomatic breadth, 20.3, 19.5, 19.7, 19.5; squamosal breadth, 17.3, 16.7, 16.4, 16.5; least inter- orbital constriction, 8.5, 8.0, 8.7, 8.3; length of nasals, 12.4, 11.5, 12.0, 11.5; alveolar length of maxillary tooth-row, 5.7, 5.7, 5.8, 5.7. Specimens examined. — Total, 13, from: 12 mi. E San Antonio de las Ala- zanas, 9000 ft., 11; 13 mi. E San Antonio de las Alazanas, 9350 ft., 2. Additional records (Howell, 1929:105): Sierra Guadalupe. Fig. 27. Distribution in Coahuila of chipmunks. 1. Eutamias bulleri soliv- agus. 2. Eutamias dorsalis carminis. 212 University of Kansas Publs., Mus. Nat. Hist. Eutamias dorsalis carminis Goldman Cliff Chipmunk Eutamias dorsalis carminis Goldman, Proc. Biol. Soc. Washington, 51:56, March 18, 1938, type from Sierra del Carmen, Coahuila. Distribution. — Oak and pine forests of the Sierra del Carmen and Sierra de la Madera (see fig. 27). Remarks. — The southeasternmost occurrence of the cliff chip- munk is in Coahuila in montane mesic forest north of latitude 26° N in the Sierra del Carmen and the Sierra de la Madera. Possibly, the animal is also in the Sierra del Pino although no evidence, other than questionable reports by local residents, were obtained there in 1952 and 1954. The Coahuilan representative of E. dorsalis is only sub- specifically distinct from the widely-separated population occurring in Arizona, New Mexico, Chihuahua and Durango and is thought to be a relict of a once more widely distributed species. In the Sierra del Carmen, the cliff chipmunk occurs as low as 4900 feet in elevation in protected caiions con'^aining parklike groves of oaks near the upper limit of the Montane Low Forest Vegetation- type. At higher elevations, in the Montane Mesic Forest, the ani- mal is more abundant; the type specimen was taken at 7400 feet. This chipmunk is a wary creature, but traps produced catches when set at the bases of large oaks, at the edge of brush on dense accumu- lations of fallen leaves, and next to rocky outcrops on the sides of narrow canons. Acorns seem favored as food, and caches of these nuts, stored by chipmunks, were found under ledges. Seiior Ed- uardo Lopez, who lived adjacent to the Carmen Mountain Club, told me that the local name for this animal is "chichimuka." In the Sierra de la Madera, the cliff chipmunk was found in the Cafion del Agua and in the Cafion de la Charretera ( south and west of Ocampo) but was less abundant than in the Sierra del Carmen. In July, 1952, and December, 1953, members of field parties saw chipmunks but none was obtained. Finally, in April 1954, four cliff chipmunks were shot in the Canon de la Charretera. In the five days spent in this canon in 1954 by a field party of five persons, only six chipmunks were seen; none was attracted to baited traps, which were set for several hundred trapnights in favorable areas. Most of the chipmunks were located by listening for their high- pitched squeaks, made when one of us approached or when one of us imitated their call. The animals were in parklike stands of oak interspersed with madrono, cherry, juniper, pine and other trees in sheltered canons above 6400 feet in elevation. Most of the chip- munks were taken in oak groves. One young animal was shot at the entrance to its den in a cavity which opened at a knot six feet above Mammals of Coahuila, Mexico 213 ground. The shallow cavity in this oak contained approximately one pint of dry grass. The trees were short, approximately 30 feet high and less than two feet in diameter. One chipmunk was taken in pine-fir forest at 7500 feet elevation. No chipmunks lived in chap- arral that grew in dense thickets on more exposed slopes. One subadult male from Club Sierra del Carmen weighed 63.8 grams; average and extreme weights of four adult, non-pregnant females from Club Sierra del Carmen and from 22-23 miles south and 5 miles west of Ocampo were 76.9 ( 69.8-83.0 ) . Females, taken on April 4 and April 7, were lactating. Although isolated by a vast area of inhospitable desert plains from other populations of dorsalis, carminis is surprisingly like examples of E. d. dorsalis ( Baird ) from New Mexico. Most describable dif- ferences are in coloring, carminis being grayer, having less distinct dorsal stripes, and having the area around the vent and the basal part of the hairs of the underside of the tail slightly darker. Com- pared with carminis from the Sierra del Carmen, specimens from the Sierra de la Madera have even less conspicuous dorsal stripes and are darker around the vent and on the basal part of the hairs of the underside of the tail. Measurements. — Three adult females from Club Sierra del Carmen and one adult female from 23 miles south and 5 miles west of Ocampo, respectively, measure: total length, 234, 239, — , 222; length of tail vertebrae, 98, 113, — , 92; length of hind foot, 35, 36, 35, 34; height of ear from notch, 20, 25, 24, 21; greatest length of skull, 37.1, 37.3, 38.6, 37.0; condylobasal length, 35.7, 35.9, 36.9, 36.1; basilar length, 28.8, 28.8, 29.3, 29.1; palatilar length, 15.9, 16.0, 16.1, 15.6; zygomatic breadth, 19.6, 20.6, 21.0, 20.5; squamosal breadth, 16.7, 17.5, 17.5, 17.5; least interorbital constriction, 7.8, 8.0, 8.7, 8.6; length of nasals, 12.0, 11.8, 12.8, 10.8; alveolar length of maxillary tooth-row, 5.8, 6.0, 6.0, 6.1. Specimens examined. — Total, 15, from: Club Sierra del Carmen, 4950 ft., 2 mi. N and 6 mi. W Piedra Blanca, 11; Sierra de la Madera, 22 mi. S and 5 mi. W Ocampo, 6200 ft., 3; Sierra de la Madera, 23 mi. S and 5 mi. W Ocampo, 7500 ft., 1. Additional records (Goldman, 1938a: 56): Carmen Mountains, 7400 ft. Sciurus alleni Nelson Allen Squirrel Sciurus alleni Nelson, Proc. Biol. Soc. Washington, 12:147, June 3, 1898, type from Monterrey, Nuevo Leon. Distribution. — Mountains of southeastern Coahuila (see fig. 28). Remarks. — The Allen squirrel lives in the mountains of southeast- em Coahuila — in some places numerously; Gerd Heinrich collected eleven in a few hours in oak groves in Canon de Meco in the Sierra de Guadalupe. Where coniferous timber predominates, such as in Diamante Pass and east of San Antonio de las Alazanas, this squirrel 214 University of Kansas Publs., Mus. Nat. Hist. seems to be less numerous. I obtained one at the Pass in a grove of pine and cedar and another was taken by Robert Packard east of San Antonio de las Alazanas in the fir-pine-aspen association. In Tamaulipas this squirrel is numerous in oak woodlands and chap- arral in the foothills of the Sierra Madre Oriental directly west of Ciudad Victoria. No evidence of this squirrel — "ardilla" of the Mexicans — or report of it was obtained from residents in the moun- tains north of Saltillo, although there is chaparral and extensive oak woodland in the Sierra de San Marcos and Sierra de la Madera. In southwestern Coahuila, alleni is unreported from Pico de Jimulco but may occur in the Sierra de Parras. Acorns seem to provide the basic food. Robert W. Dickerman found partly eaten cones, probably of Pinus arizonico, two miles east of Mesa de Tablas, A female taken on March 20 had two embryos averaging 25 mm. long (crown to rump). Two females obtained on April 23 were lactating. Three non-pregnant females and three males, all from Sierra Guadalupe, respectively, weighed: 452, 473, 507 and 486, 490, 491. Specimens from Coahuila closely resemble alleni from the vicinity of Linares, Nuevo Leon. Sciurus alleni bears a super- ficial resemblance to Sciurus carolinensis Gmelin but is more closely related to Sciurus nayari- tensis J. A. Allen and Sciurus oculatus Peters. Cranially, these species show no marked differ- ences. Both Nelson (1899:92) and Dalquest (1953:86) remark on the resemblance of alleni to oculatus. Dalquest writes that alleni is "Strikingly similar to Sciurus oculatus in proportions and color but slightly smaller . . ., under parts white rather than buffy, and postauricular spots absent." In alleni from Coahuila there are pale to dark buffy, postauricular spots con- Fig. 28. Distribution in Coahuila of •^ '- '■ tree squirrels. 1. hcnirus alleni. 2. spicuous on some specimens but Sciurus niger limitis. Mammals of Coahuila, Mexico 215 less so on others. It is suspected that nayaritensis is related to ocu- latus through alleni and that squirrels ancestral to these species may have spread from western Mexico across the Mesa del Norte by way of mountains in western Coahuila, Zacatecas and Durango. Measurements. — Measurements of three adult males from Sierra Guadalupe and one adult female from Diamante Pass, respectively, are: total length, 433, 441, 440, 456; length of tail vertebrae, 200, 219, 209, 216; length of hind foot, 55, 63, 61, 62; height of ear from notch, 30, 31, 31, 34; greatest length of skull, 59.0, 58.8, 58.7, 59.3; basilar length, 44.4, 45.1, — , 45.2; zygomatic breadth, 34.1, 32.2, 33.0, 33.5; length of nasals, 20.4, 19.8, 20.3, 20.2; width across posterior tongues of premaxillae, 16.4, 15.5, 14.8, 16.1; mastoidal breadth, 27.2, 25.9, 26.1, 26.5; postorbital constriction, 17.0, 18.3, 17.3, 18.5; alveolar length of ma.xillary tooth-row^, 10.4, 10.5, 9.5, 10.4, Specimens examined. — Total, 13, from: Diamante Pass, 8200 ft., 4 mi. E and 3 mi. S Saltillo, 1; 13 mi. E San Antonio de las Alazanas, 1; Caiion de Meco, 6400 ft.. Sierra Guadalupe, 10 mi. S General Cepeda, 11. Seiurus niger limitis Baird Fox Squirrel Seiurus limitis Baird, Proc. Acad. Nat. Sci. Philadelphia, vol. 7 (1854-55), p. 331, April, 1855, type from Devil's River, Valverde County, Texas. S[ciurus]. n [iger]. limitis Osgood, Proc. Biol. Soc. Washington, 20:45, April 18, 1907. Distribution. — Northeastern Coahuila, along timbered streams and in oak- forested uplands (see fig. 28). Remarks. — The fox squirrel lives in groves of timber along the Rio Grande and some of its major tributaries and also in isolated oak forests on the Rancho Las Margaritas in the Serranias del Burro. This species occurs along the Rio Grande at least as far upstream as Villa Acufia, Southeast of Piedras Negras the fox squirrel is re- ported by Senior Alvaro Trevino of the Rancho La Candelaria to occur in live oak woodlands near the Rio Grande. Perhaps the squirrel is more abundant in suitable habitat along smaller per- manent streams, such as the Rio de San Diego near Jimenez, where eight specimens were taken in June, 1952. These were obtained in a parklike grove of pecan and cypress bordering the stream. Along the Rio Sabinas fox squirrels are numerous from Presa Don Martin at least as from upstream as Nacimiento. Several squirrels were shot in groves of pecan, cypress and hackberry adjacent to the Rio Sabinas in the vicinity of Ciudad Muzquiz in June, 1938, and in April and June, 1952. Fox squirrels have made their way also along dry arroyos to the foothills of the Sierra de Santana, north of Nacimiento, where oaks grow in the vicinities of springs. The squirrels live in small numbers in oak forests of the Upper Sonoran and Transition Life-zones in the Serranias del Burro. There, acorns 216 University of Kansas Publs., Mus. Nat. Hist, constitute the principal food of squirrels. They reached the Burros either from the north by way of tributaries from the Rio Grande or from the south by way of the watershed of the Rio Sabinas. In color, specimens from the Burros more closely resemble those from the Rio Grande than those from the Rio Sabinas to the south. No fox squirrels have been reported from along the Rio Salado, except near where it flows into Presa Don Martin, Coahuilan residents refer to the fox squirrel as "onza" although the name "ardilla" also is used. Fox squirrels breed twice annually in Coahuila, Females taken in late March and early April were lactating; two obtained in mid- June were pregnant and contained 2 and 4 embryos. Three of eight fox squirrels taken two miles west of Jimenez in mid-June are in molt, Coahuilan fox squirrels resemble closely near topotypes of S. n. limitis, taken in February along the Devil's River ( 18 miles north of Comstock, Valverde County, Texas), Although within the range of color of the near topotypes, specimens from near the Rio Grande (taken in June) and from the Serranias del Burro (taken in March) are darker, especially on the underside of the tail, which is Ochra- ceous Orange, and on the belly, feet and ears, than squirrels from along the Rio Sabinas (taken in April and June), which are much paler (the underside of the tail is Ochraceous-Buff ), In addition to being paler, the underside of the tail in most specimens from along the Rio Sabinas is more suffused with blackish. No significant difference in size was observed between the near topotypes of limitis and Coahuilan fox squirrels, but specimens from along tlie Rio Sabinas average slightly smaller in most cranial measurements. Average and extreme weights of adult, non-pregnant females and adult males from Fortin and from the Rio Sabinas, respectively, were: 5 females, 455,6 (388.2-509.0); 1 male, 433.2 and 4 females, 487.6 (436.0-547.0); 2 males, 460.1, 515.7. Measurements. — Measurements of 3 adult males from 2 miles west of Ji- menez and from Fortin and average and extreme measurements of 5 adult females from Fortin, respectively, are: total length, 452, 485, 467, and 467 (447-494); length of tail, 199, 238, 216 and 215 (200-237); length of hind foot, 65, 65, 63 and 63 ( 62-65 ) ; height of ear from notch, 32, 35, 33 and 32 ( 30-34 ) ; greatest length of skull, 58.0, — , 56.8 and 57.8 (57.2-60.0); basilar length, 44.7, 44.8, 43.9 and 45.0 (43.9-46.8); zygomatic breadth, 31.6, 32.8, 32.0 and 33.3 (32.5- 34.7); length of na.sals, 20.2, 19.5, 20.0 and 20.1 (19.5-20.8); lca.st postorbital constriction, 18.9, 18.8, 17.8 and 18.4 (17.8-19.1); mastoidal breadth, 25.4. 26.3, 26.1 and 26.4 (26.0-27.3); width across posterior projections of pre- maxillaries, 14.6, 15.6, 15.9 and 15.6 (14.8-16.1); alveolar length of maxillary tooth-row, 10.0, 11.0, 10.3 and 10.2 (10.0-10.5). Mammals of Coahuila, Mexico 217 Specimens examined. — Total, 27, from: 2 mi. W. Jimenez, 850 ft., 8; Fortin, 3300 ft., Rancho Las Margaritas, 7; 7 mi. NE Hacienda Las Margaritas, 4000 ft., 4; Rio Sabinas, 8 mi. N and 4 mi. W Muzquiz, 1800 ft., 7; Rio Sabinas, 2 mi. S and 3 mi. E San Juan de Sabinas, 1160 ft., 1. Additional record (Elliot, 1907:125): Sabinas. Thomomys bottae Botta Pocket Gopher The Botta pocket gopher is more abundant in mountains where soils are dark than in arid lowlands, where soils are pale. Isolated groups of these fossorial creatures frequent stands of lechuguilla; the roots and growing parts of this plant provide a basic food for Thomomys. Oftentimes the pres- ence of pocket gophers can be detected by the finding of patches of dead lechuguilla plants, especially those which have been pulled partway under- ground by the gophers. In north- eastern Coahuila bottae occurs also in deep soils along arroyos at least as far eastward as Villa Acuiia. The Rio Grande bars the passage of Thomomys bottae and distinctive subspecies oc- cupy the opposite sides of this river (see Baker, 1953a: 506). Generally, in more mesic situa- tions at higher elevations pocket gophers are darker, while in drier, desert plains, animals are paler. Pocket gophers were difficult to catch, and in many places where they might be expected to live, none was obtained. There was no sign of Thomomys even in such favorable areas as the Sierra de los Hecheros and in the Sierra de la Madera. Locally this species is called "tuza" or "topo." Females with embryos were taken on January 21 ( one with 2 em- Wos), July 30 (one with 4 embryos), and December 7 (one with 4 embryos). Robert Dickerman obtained an adult male and an adult female from the same burrow on January 13. FiG. 29. Distribution in Coahuila of pocket gophers of the species Tho- momys bottae and Thomomys um- brinus. 1. T. b. analogus 5. T. b. rctractus 2. T. b. augustidcns 6. T. b. sturgisi 3. T. b. humilis 7. T. b. villai 4. T. b. perditus 8. T. u. goldmani 218 University of Kansas Publs., Mus. Nat. Hist. Thomomys bottae analogus Goldman Thomomys umbrinus analogus Goldman, Proc. Biol. Soc. Washington, 51:59, March 18, 1938, type from Sierra Guadalupe, Coahuila. Thomomys bottae analogus Baker, Univ. Kansas Publ., Mus. Nat. Hist., 5: 511, June 1, 1953. Distribution. — Mountains and elevated plains of southeastern Coahuila (see fig. 29). Remarks. — This subspecies is characterized by having a large, broad skull. The upper parts are darker than those of T. b. perditus but not so dark as those of T. b. sturgisi. Aside from a few scattered records of analogus from the elevated plains, most specimens were obtained in openings surrounded by oaks and conifers in moun- tainous areas. Specimens assigned to analogus from Nuevo Leon (Cerro Potosi, near La JoUa, Galeana, 48263 CNHM) and from Zacatecas (Con- cepcion del Oro, 58063 KU) indicate that this subspecies occurs in mountainous areas south of Coahuila. Measurements. — Average and extreme measurements of 7 adult males and 4 adult females from the two localities east of San Antonio de las Alazanas ( see below), respectively, are: total length, 213 (203-223), 198 (184-205); length of tail vertebrae, 57 (52-66), 56 (51-60); length of hind foot, 28 (27-30), 27 (27-28); basilar length of skull, 31.8 (30.6-32.9), 30.5 (29.8-31.3); zygomatic breadth, 23.9 (21.7-25.0), 22.3 (21.6-22.8); least interorbital constriction, 6.8 (6.4-7.1), 6.8 (6.5-7.0); squamosal breadth, 18.9 (18.0-19.5), 18.3 (17.7- 18.9); length of nasals, 13.5 (12.5-14.1), 12.4 (12.2-12.6); breadth of rostrum, 6.9 (6.6-7.4), 6.8 (6.6-7.0); alveolar length of maxiUary tooth-row, 7.3 (7.0- 7.5), 6.9 (6.6-7.1). Specimens examined. — Total, 49, from: Jaral, 3860 ft., 1 (BSC); 5 mi. E and 2 mi. S Saltillo, 9000 ft. 1; Diamante Pass, 8500 ft., 3 mi. E and 4 mi. 5 Saltillo, 5; 4 mi. S and 6 mi. E Saltillo, 7500 ft., 1; 1 mi. S and 4 mi. W Bella Union, 7000 ft., 4; 2 mi. SE Bella Union, 5850 ft., 1; 12 mi. E San Antonio de las Alazanas, 9000 ft., 6; 3 mi. S and 13 mi. E San Antonio de las Alazanas, 8700 ft., 9; Sierra de Guadalupe, 7000 ft., 10 mi. S and 8 mi. W General Cepeda, 3; Sierra de Guadalupe, 8200 ft., 10 mi. S and 7 mi. W General Cepeda 2; Sierra de Guadalupe, 11 mi. S and 6 mi. W General Cepeda, 1; Sierra de Guadalupe, 6700 ft., 11 mi. S and 4 mi. W General Cepeda, 9; Sierra de Guadalupe, 2 (BSC); Cameros, 3 (BSC); Sierra Encamacion, 9000 ft., 1 (BSC). Thomomys bottae angustidens Baker Thomomys bottae angustidens Baker, Univ. Kansas Publ., Mus. Nat. Hist., 5:508, Tune 1, 1953, type from Sierra del Pino, 5250 ft., 6 mi. N and 6 mi. W AceDuches, Coahuila. Distribution. — Sierra del Pino of northwestern Coahuila and probably other isolated desert ranges in the vicinity and west of the Sierra de la Encantada (see fig. 29). Remarks. — This subspecies may occur in several desert ranges in northwestern Coahuila. No specimens were obtained on the Sierra de Los Hecheros, but one taken in the Sierra Almagre, Chihuahua ( 19 miles south and 4 miles west of Jaco, 55563 KU ) is assigned to Mammals of Coahuila, Mexico 219 angustidens. This subspecies is small and pale-colored with a short, broad skull and a broad rostrum. In the Sierra del Pino this pocket gopher was trapped from burrows in grassy places surrounded by oaks and pines. Two specimens are in dark color phase. Measurements. — Average and extreme measurements of 5 adult males and 3 adult females from the Sierra del Pino (see below), respectively, are: total length, 192 (174-200), 190 (187-195); length of tail vertebrae, 53 (45-64), 57 (52-64); length of hind foot, 26 (25-27), 25 (24-26); basilar lengtli of skull, 29.5 (28.9-29.9), 27.8 (27.5-28.2); zygomatic breadth, 20.7 (20.4-21.0), 19.6 (18.8-20.3); least interorbital constriction, 6.7 (6.0-7.0), 6.6 (6.2-7.2); squamosal breadth, 17.6 (17.4-17.8), 16.7 (16.4-17.0); length of nasals, 12.4 (11.9-13.0), 11.1 (10.9-11.3); breadth of rostrum, 6.5 (5.9-6.9), 6.9 (6.0-7.3); alveolar length of maxillary tootli-row, 6.9 (6.5-7.0), 6.9 (6.4-7.4). Specimens examined. — Total, 11, from: Sierra del Pino, 5250 ft., 6 mi. N and 6 mi. W Acebuches, 3; Sierra del Pino, 7925 ft., 5 mi. N and 9 mi. W Acebuches, 2; Sierra del Pino, 6200 ft., 5 mi. S and 3 mi. W Acebuches, 6. Thomomys bottae humilis Baker Thomomys bottae humilis Baker, Univ. Kansas Publ., Mus. Nat. Hist., 5:503, June 1, 1953, type from 3 mi. W Hacienda San Miguel, 2200 ft., Coahuila. Distribution. — Desert plains of north-central Coahuila, north of the Ser- ranias del Burro and as far east as the vicinity of Villa Acuna ( see fig. 29 ) . Remarks. — This subspecies occupies lower elevations especially where lechuguilla grows, north and east of the Serranias del Burro. Paler color of upper parts and smaller dimensions of the skull dis- tinguish humilis from T. h. retractus, which occurs in the Serranias del Burro. Specimens of T. b. limitaris Goldman, from the vicinity of Comstock, Texas, across the Rio Grande, resemble humilis in many ways; perhaps the Rio Grande, at least in the vicinity of Com- stock and Villa Acuna, may not completely bar the passage of these pocket gophers. Measurements. — Measurements of two adult males from the type locality and two adult males from 1 mile south and 9 miles west of Villa Acuiia, respec- tively, are: total length, 197, 200, 188, 201; length of tail vertebrae, 55, 55, 52, 63; length of hind foot, 27, 27, 24, 27; basilar length of skull, 30.8, 30.7, 29.5, 29.0; zygomatic breadth, 22.3, 21.0, 21.1, 20.6; least interorbital constriction, 6.7, 6.4, 6.1, 6.7; squamosal breadth, 18.8, 17.2, 17.5, 16.3; length of nasals, 11.7, 12.2, 12.0, 11.2; breadth of rostrum, 6.9, 6.1, 7.2, 6.0; alveolar length of maxillary tooth-row, 7.2, 7.2, 6.7, 7.0. Specimens examined. — Total, 8, from: 1 mi. S and 9 mi. W Villa Acuna, 3; Caiion del Cochino, 3200 ft., 16 mi. N and 21 mi. E Piedra Blanca, 2; 3 mi. W Hacienda San Miguel, 2200 ft., 3. Thomomys bottae perditus Merriam Thomomys perditus Merriam, Proc. Biol. Soc. Washington, 14:108, July 19, 1901, type from Lampazos, Nuevo Leon. Thomomys bottae perditus Baker, Univ. Kansas Publ., Mus. Nat. Hist., 5:510, June 1, 1953. 220 University of Kansas Publs., Mus. Nat, Hist, Distribution. — Mountains and elevated plains of east-central Coahuila (see fig. 29). Remarks. — This pale-colored, medium-sized pocket gopher lives in the northernmost parts of the Sierra Madre Oriental, Populations are isolated on discontinuous hills. Measurements. — Measurements of an adult male from 2 miles north and 18 miles west of Santa Teresa and average and extreme measurements of 5 adult females from 9 miles east of Hermanas, respectively, are: total length, 187, 195 (189-201); length of tail vertebrae, 57, 57 (55-63); length of hind foot, 24, 26 (25-27); basilar lengtli of skull, 30.1, 29.3 (28.3-30.9); zygomatic breadth, 21.4, 20.9 (19.9-21.7); least interorbital constriction, 6.3, 6.9 (6.8- 7.0); squamosal breadth, 17.2, 17.5 (16.9-18.6); length of nasals, 12.4, 11.6 (11.2-12.1); breadth of rostrum, 6.7, 6.7 (6.1-6.8); alveolar length of maxillary tooth-row, 6.6, 7.2 (7.0-7.4). Specimens examined. — Total, 11, from: 9 mi. E Hermanas, 2000 ft., 8; Panuco, 1; 2 mi. N and 18 mi. W Santa Teresa, 7500 ft.. 2, Thomomys bottae retractus Baker Thomomys bottae retractus Baker, Univ. Kansas Publ., Mus. Nat. Hist., 5: 507, June 1, 1953, type from Fortin, 3300 ft., Rancho Las Margaritas, CoahuUa. Distribution. — Serranias del Burro of north-central Coahuila (see fig. 29). Remarks. — This subspecies is distinguished from adjacent sub- species of bottae by its larger size and larger skull. The animal lives in dark soils within the oak-juniper-pine belt of the Serranias del Burro; it has not been obtained below this vegetational level. In the original description (Baker, 1953a: 507) the type locality of retractus at Fortin was hsted as being 33 miles N and 1 mile E of San Geron- imo. This is incorrect; Fortin is a place on the Rancho Las Mar- garitas, approximately 7 miles north of the Hacienda Las Margaritas and 20 miles north and 2 miles east of San Geronimo. Measurements. — Average and extreme measurements of 6 adult males and 5 adult females from the Rancho Las Margaritas (see below), respectively, are: total length, 210 (195-220), 200 (192-205); length of tail vertebrae, 65 (58- 74), 62 (57-71); length of hind foot, 27 (25-28), 25 (24-27); basilar length of skull, 32.1 (30.6-33.6), 29.9 (29.6-30.2); zygomatic breadth, 22.8 (21.5-24.1), 21.5 (21.0-22.4); least interorbital constriction, 6.8 (6.1-7.5), 6.4 (6.2-6.6); squamosal breadth, 18.3 (17.5-18.9), 17.2 (17.0-17.5); length of nasals, 13.7 (13.0-14.0), 11.9 (11.3-12.4); breadth of rostrum, 7.0 (6.5-7.5), 6.6 (6.2-7,2); alveolar length of maxillary tooth-row, 7.3 (7.1-7,6), 7.0 (6.7-7.5), Specimens examined. — Total, 34, from Fortin: 3300 ft., Rancho Las Mar- garitas, 8; 6 mi. NW Hacienda Las Margaritas, 4500 ft., 1; El Rio Alamos, 3300 ft., in Cafion Mulato, Rancho Las Margaritas, 1; 1 mi. N Hacienda Las Margaritas, 2800 ft., 12; I mi. NW Hacienda Las Margaritas, 1; ^ mi, W Hacienda Las Margaritas, 6; Hacienda Las Margaritas, 2900 ft., 5. Mammals of Coahuila, Mexico 221 Thomomys bottae sturgisi Goldman Thomomys sturgisi Goldman, Proc. Biol. Soc. Washington, 51:56, March 18, 1928, type from Carmen Mountains [= Sierra del Carmen], 6000 ft., Coa- huila. Distribution. — Mountains and elevated plains of north-central Coahuila from as far south as Cuatro Cicnegas northward to the northern limits of the Sierra del Carmen at the south side of the Rio Grande (see fig. 29). Remarks. — This subspecies is small and dark with a small skull having a broad roshum. Animals from the Sierra de la Encantada and from the vicinity of Ocampo are larger and paler than those from the Carmens. Like analogus, sturgisi occurs in the darker soils of the sierras and in stands of lechuguilla and yucca on the desert plains. Measurements. — Measurements of 2 adult males and 2 adult females from the Club Sierra del Cannen, respectively, are: total length, — , 204, 194, 199; length of tail vertebrae, — , 68, 65, 64; length of hind foot, 27, 27, 26, 25; basilar length, 29.7, 29.8, 27.2, 28.5; zygomatic breadth, 20.5, 20.5, 18.8, 19.7; least interorbital constriction, 6.6, 6.5, 6.2, 6.5; squamosal breadth, 16.4, 17.1, 16.1, 16.6; length of nasals, 12.7, 12.3, 10.9, 11.5; breadth of rostrum, 6.8, 6.7, 5.9, 6.4; alveolar length of maxillary tooth-row, 7.2, 7.1, 6.6, 7.0. Specimens examined. — Total, 29, from: Club Sierra del Carmen, 4950 ft., 2 mi. N and 6 mi. W Piedra Blanca, 6; Madera Camp, Sierra del Carmen, 3 (DMNH); Juarez Canon, Sierra del Carmen, 3 (DMNH); Sierra de la Encan- tada, 4100 ft., 37 mi. S and 21 mi. E Boquillas, 1; Sierra de la Encantada, 4400 ft., S8 mi. S and 23 mi. E Boquillas, 6; 16 mi. E and 18 mi. N Ocampo, 5; 2 mi. N and 1 mi. W Ocampo, 4050 ft., 1; 5 mi. SE Ocampo, 3; 3 mi. NW Cuatro Cienegas, 2450 ft., 1 (partial cranium only). Thomomys bottae villai Baker Thomomys bottae villai Baker, Univ. Kansas Publ., Mus. Nat. Hist., 5:505, June 1, 1953, type from 7 mi. S and 2 mi. E Boquillas, 1800 ft., Coahuila. Distribution. — Desert lowlands bordering the south side of the Rio Grande and the Arroyo de los Alamos in northwestern Coahuila, eastward no farther than the beginnings of the higher lands of the Sierra del Carmen and related ranges (see fig. 29). Remarks. — This medium-sized pocket gopher is the palest sub- species of bottae known from Coahuila. Specimens taken in late February are pale cinnamon buff above and white to pale buff below. The tympanic bullae are noticeably larger, in relation to the size of the skull, than in adjacent subspecies of bottae. The absence of river bank mantled with soil in the narrow Boquillas Caiion pre- vents villai from coming into contact with hiimilis, which occurs in lowlands downstream from the caiion in Coahuila. These two sub- species are noticeably different. The Rio Grande also is an effective barrier to Thomomys, and villai is distinct from T. b. limitaris that hves in the Big Bend area of Texas (see Baker, 1953a: 505-506). 4—1204 222 University of Kansas Publs., Mus. Nat. Hist. A series of 9 villai was obtained from a lechuguilla-covered bank adjacent to an arroyo. No other sign of Thomomys was found in the vicinity of Boquillas. Measurements. — Average and extreme measurements of 3 adult males and 3 adult females from the type locality, respectively, are: total length, 195 (194- 196), 211 (209-213); length of tail vertebrae, 58 (56-61), 66 (65-68); length of hind foot, 28 (28-29), 28 (27-28); basilar length of skull, 31.7 (31.0-32.5), 30.7 (30.1-31.6); zygomatic breadth, 22.9 (22.2-23.6), 21.5 (21.1-21.7); least interorbital constriction, 6.6 (6.4-6.9), 6.5 (6.4-6.8); squamosal breadth, 18.5 (18.5-18.6), 18.3 (18.0-18.5); length of nasals, 12.4 (12.2-12.6), 11.8 (11.4- 12.2); breadth of rostrum, 7.3 (7.2-7.5), 7.0 (6.5-7.2), alveolar length of maxil- lary tooth-row, 7.1 (6.8-7.3), 7.0 (6.7-7.3). Specimens examined. — Total, 9, from 7 mi. S and 2 mi. E Boquillas, 1800 ft. Thomomys umbrinus goldmani Merriam Pigmy Pocket Gopher Thomomys goldmani Merriam, Proc. Biol. Soc. Washington, 14:108, July 19, 1901, type from Mapimi, Durango. Thomomys umbrinus goldmani Nelson and Goldman, Jour. Mamm., 15:115, May 15, 1934. Distribution. — Desert plains of western and southwestern Coahuila (see fig. 29). Remarks. — Some specimens from Sierra Mojada (taken in May) are duller colored than goldmani from the vicinity of Mapimi ( taken in February). Coahuilan umbrinus have narrower rostra than bottae from there. The pigmy pocket gopher occurs in small numbers in west-central and southwestern Coahuila in desert areas of the Central Plateau. Specimens from Sierra Mojada were trapped in a fallow, terraced field at the edge of the village. Differences between umbrinus and bottae, in Coahuila, are discussed by Baker (1953a: 501-502). The most useful characters to distinguish these species are the curvature of the margin of the anterior base of the maxilla where it meets the frontal (in bottae this margin is convex, in umbrinus it is concave or emarginate) and the extent of attachment of the lachrymal, in dorsal aspect, to tlie maxillary arm of the zygomatic arch (in bottae approximately one-half of the lachrymal is attached and in umbrinus three-fourths or more of the lachrymal is attached to the zygomatic arch). In addition, in bottae the upper incisors are decurved or moderately procumbent while in umbrinus these teeth are strongly procumbent. T. umbrinus, T. bottae, T. baileyi, and T. townsendii have a sphenorbital fissure (vertical slit in the back of tlie orbit) whereas the two other species of the genus, T, talpoides and T. 7nonticola, lack this fissure. A female, taken on January 21, contained two embryos. Mammals of Coahuila, Mexico 223 Measurements. — Measurements of a male from 3 miles northeast of Torreon and a male and two females from Sierra Mojada, and from 3 miles southeast thereof, respectively, are: total length, 193, 195, 201, 192; length of tail ver- tebrae, 61, 56, 61, 58; length of hind foot, 27, 26, 26, 26; basilar length of skull, 30.2, 30.4, 31.2, 30.3; zygomatic breadth, 20.2, 21.6, 21.3, 21.2; least interorbital constriction, 6.5, 6.0, 6.4, 6.4; squamosal breadth, 17.2, 17.5, 17.7, 18.1; length of nasals, 10.9, 11.6, 11.4, 11.7; breadth of rostrum, 6.2, 5.8, 5.7, 5.8; alveolar length of maxillary tooth-row, 7.0, 6.8, 6.9, 7.1. Specimens examined. — Total, 7, from: 3 mi. NE Sierra Mojada, 4100 ft., 2; Sierra Mojada, 3; 3 mi. SE Torreon, 3800 ft., 2. Cratogeomys castanops Plateau Pocket Gopher The plateau pocket gopher lives in open areas in the Lower So- noran Life-zone in Coahuila and also in the Upper Sonoran Life- zone in broad intermontane valleys, at least as high as 8700 feet in elevation. Unlike Thomomys, which occurs in the dark, rocky soils within the oak and pine belts of the mountains as well as in the desert plains, castanops is restricted to open places mantled by suit- able soils, preferably sandy in texture, in which the animals can dig their elaborate underground systems of runways. Thin soils of hard texture and rocky soils provide less favorable places for cas- tanops to live, and the animals usually are uncommon or absent in such situations. As a result, castanops often is partly isolated as small populations where suitable habitat is surrounded on one or more sides by thin, rocky soils of uplands and mountains. The Rio Grande, in many places flowing through steep-walled canons, also is a barrier that this sedentary animal does not cross. Smaller streams within the State seem to be unimportant barriers to the passage of these animals. The food supply of castanops consists of fleshy tuberous roots of lechuguilla and other well-distributed desert shrubs; in elevated intermontane valleys, where desert shrubs are absent, roots and leaves of low-growing forbs are eaten. The food supply seems to be adequate in most situations and is thought not to govern the distribution of this species. Russell and Baker (1955:593-594) have recognized 11 subspecies of Cratogeomys castanops in Coahuila. These Hve in three distinct habitats. These authors write (/oc. aY.: 594) that Coahuila ". . . is crossed by a series of mostly impassable, mountainous ridges [= Sierra del Carmen-Sierra Madre Oriental Axis] beginning at the northwestern boundary at the Caiion de Boquillas on the Rio Grande and extending southeastward to the east-central border. This di- vides Coahuila into a more humid and less elevated northeastern 224 University of Kansas Publs., Mus. Nat. Hist. area which is an inland extension westward of the Coastal Plain and a more arid and higher western and southern area which is a part of the 'Mesa del Norte' of the Mexican Plateau. In the extreme southeast the still higher elevated plains and intermontane valleys within the Sierra Madre Oriental afford a third habitat for popula- tions of this species. The subspecies of these pocket gophers found in any one of these three habitats show greater affinity to each other than they do to any subspecies found in the other habitats." Re- lated subspecies of castanops in northeastern Coahuila (see figure 30) are: convexus, bullatus, us- tulatus, jiictindtis; in the south- east: subnubilis, planifrons; and in the west: convexus, which is smaller, and sordidulus, excelsus, siibsimus, goldmani. Pregnant females contained from one to three embryos (aver- age 1.8). Two distinct breeding periods were noted, in winter (December-March) and in sum- mer ( June-August ) . Pregnant females were taken on December 4, 7, January 8, 11, 25, March 19, June 24, 30, July 6, 9, 18, 22, and August 16. Lactating females were examined on January 11, 13, June 25 and July 9. Non- pregnant, adult females were ob- tained in every month except September and October; no spec- imens with data concerning re- productive condition were available for those two months. Russell (1954:122) reports the taking of a nursing female, an adult male and a young female in the same trap-set in the course of the night of June 24-25, near Muralla. Cratogeomys castanops bullatus Russell and Baker Cratogeomys castanops bullotus Russell and Baker, Univ. Kansas Publ., Mus. Nat. Hist., 7:597, March 15, 1955, type from 2 mi. S and 6J2 mi. E Nava, 810 ft., Coahuila. Distribution. — Desert lowlands of northeastern Coahuila, from the Rio Grande as far southwestward as the Rio Sabinas (see fig. 30). Fic. 30. Distribution in Conhuila of the plateau pocket gopher, Cratogeo- mys castanops. Guide to subspecies 6. C. c. jucundus 1. C. c. bullatus 7. C. c. planifrons 2. C. c. consitus 8. C. c. sordidulus 3. C. c. convexus 9. C. c. subnubilus 4. C. c. excelsus 10. C. c. subsimus 5. C. c. goldmani 11. C. c. ustulatus Mammals of Coahuila, Mexico 225 Remarks. — In size, Cratogeomijs castanops hullatus resembles C. c. tomaulipends Nelson and Goldman of the lower Rio Grande Val- ley in Tamaulipas, but the two differ markedly in cranial features. In color and cranial characters bullatus resembles convexus, but bullattis is smaller. Comparison of this subspecies with C. c. an- gusticeps Nelson and Goldman, which occurs across the Rio Grande but not directly opposite the range of bullatus, indicates that the two are less closely related than bullatus is to tamaulipensis and convexus. C. c. bullatus is common in sandy soils in extreme northeastern Coahuila. In the vicinity of Nava, mounds were found in fallow irrigated fields and other open places between extensive live oak thickets. South and west of the Rio Grande this gopher was less abundant and lived in heavier soils usually as individuals or in small groups. Measurements. — Measurements of one adult male and average and extreme measurements of 5 adult females from the three localities south and east of Nava (see below), respectively, are: total length, 261, 256 (242-263); length of tail vertebrae, 80, 80 (72-85); length of hind foot, 36, 36 (35-37); condy- lobasal length of skull, 49.7, 47.4 (47.0-47.7); zygomatic breadth, 35.3, 30.6 (29.9-31.1); length of palate, 34.4, 32.6 (32.3-32.9); breadth of rostrum, 12.4, 10.7 (10.0-11.6); length of nasals, 17.1, 17.1 (16.5-17.8); squamosal breadth, 29.2, 27.9 (27.5-28.2); alveolar length of maxillary tooth-row, 9.5, 9.5 (9.2-9.8). Specimens examined. — Total, 24, from: 2 mi. S and 6 mi. E Nava, 810 ft., 2; 2 mi. S and 12 mi. E Nava, 800 ft., 1; 3 mi. S and 12 mi. E Nava, 800 ft., 4; 29 mi. N and 6 mi. E Sabinas, 5; 10 mi. E Hacienda La Mariposa, 2000 ft., 1; La Gacha [r= La Concha], 1600 ft., 8; 8 mi. S and 8 mi. E Hacienda La Mariposa, 1900 ft., 1; 9 mi. S and 11 mi. E Sabinas, 1050 ft., 2. Cratogeomys castanops consitus Nelson and Goldman Cratogeomys castanops consitus Nelson and Goldman, Proc. Biol. Soc. Wash- ington, 47:140, June 13, 1934, type from Gallego, 5500 ft.. Chihuahua. Distribution. — Arid high plains of northwestern Coahuila (see fig. 30). Remarks. — Cratogeomys castanops consitus is a small pocket gopher; the largest adult seen is much smaller than the smallest adult of any adjacent subspecies. Specimens from Coahuila as- signed to consitus compare favorably with topotypes although those from the vicinity of Jaco are smaller, paler and have a narrower rostrum and smaller maxillary teeth. This subspecies seems to be rare in northwestern Coahuila and small colonies are widely scat- tered. Cratogeomys castanops clarkii (Baird) may occur along the Rio Grande in extreme northwestern Coahuila but no specimen is ac- tually known from there. 226 University of Kansas Publs., Mus. Nat. Hist. Measurements. — Measurements of an adult male from 3 miles nortli and 9 miles east of El Pino and average and extreme measurements of 4 adult females from 6 miles east of Jaco, respectively, are: total length, 289, 229 (226-232); length of tail vertebrae, 94, 74 (68-78); length of hind foot, 36, 32 (31-32); condylobasal length of skull, 53.8, 43.8 (42.6-45.8); zygomatic breadth, 32.6, 28.1 (27.3-28.8); length of palate, 37.1, 29.6 (29.4-29.9); breadth of rostrum, 12.7, 9.7 (9.4-9.9); length of nasals, 18.8, 16.0 (15.5-16.2); squamosal breadth, 29.5, 26.2 (25.7-26.9); alveolar length of maxillary tooth-row, 9.6, 8.9 (8.1-9.2). Specimens examined. — Total, 8, from: 3 mi. N and 9 mi. E El Pino, 1; 6 mi. E Jaco, Chihuahua, in Coahuila, 6; 3 mi. NE Sierra Mojada, 1. Cratogeomys castanops convexus Nelson and Goldman Cratogeomys castanops convexus Nelson and Goldman, Proc. Biol. Soc. Wash- ington, 47:142, June 13, 1934, type from 7 mi. E Las Vacas [= Villa Acuna], Coahuila. Distribution. — Extreme northern Coahuila, east and north of the Serranias del Burro (see fig. 30). Remarks. — This subspecies is larger than bullatus, which occurs to the southeast. To the west and southwest, mountainous country seems to be a barrier that this animal does not cross. Conspicuous difiFerences between convexus and angusticeps, found across the Rio Grande in Texas, indicate that this river is an eflFective barrier to passage by these fossorial rodents. Measurements. — Measurements of one adult male and two adult females from Villa Acufia, respectively, are: total length, 275, 260, 265; length of tail vertebrae, 89, 86, 83; length of hind foot, 39, 37, 38; condylobasal length of skull, 55.0, 50.6, 49.3; zygomatic breadth, 34.4, 31.7, 31.6; length of palate, 37.0, 33.8, 32.9; breadth of rostrum, 12.6, 11.7, 11.8; length of nasals, 20.0, 16.7, 15.8; squamosal breadth, 30.9, 29.1, 28.9; alveolar length of maxillary tooth-row, 10.4, 9.3, 10.6. Specimens examined. — Total, 14, from: S side Rio Grande. 17 mi. S Dryden, Terrell County, Texas, in Coahuila, 6; Rio Grande, opposite Samuels, Val Verde County, Texas, in Coahuila, 1 (BSC); Villa Acuna, 5; Caiion del Cochino, 21 mi. E and 16 mi. N Piedra Blanca, 1; 11 mi. W Hacienda San Miguel, 1. Cratogeomys castanops excelsus Nelson and Goldman Cratogeomys castanops excelsus Nelson and Goldman, Proc. Biol. Soc. Wash- ington, 47:143, June 13, 1934, type from San Pedro, 10 mi. W Laguna de Mayran, Coahuila. Distribution. — Desert plains of southwestern Coahuila (see fig. 30). Remarks. — Cratogeomys castanops excelsus is characterized by large size and pale color and is the palest subspecies of castanops. Of adjacent subspecies, excelsus most closely resembles subsiinus, which occurs to the east, and least resembles consitus, which occurs to the northwest. This subspecies lives in deep soils of the interior basin of south- western Coahuila and is common in the cultivated areas in, and in the vicinity of, the formerly extensive Laguna de Mayran. Mammals of Coahuila, Mexico 227 Measurements. — Measurements of one adult male from 2 miles east of Tor- reon and average and extreme measurements of 4 adult females from 4 miles north of Acatita, respectively, are: total length, 315, 284 (274-291); length of tail vertebrae, 97, 82 (77-86); length of hind foot, 41, 37 (35-38); condy- lobasal length of skull, 54.7, 51.4 (51.1-51.6); zygomatic breadth, 37.8, 34.1 (33.6-34.9); length of palate, 37.6, 35.4 (34.7-36.1); breadth of rostrum, 12.1, 11.6 (10.4-12.1); length of nasals, 19.5, 18.9 (18.4-20.1); squamosal breadth, 31.4, 31.2 (30.5-31.7); alveolar length of maxillary tooth-row, 9.8, 9.5 (9.2-9.9). Specimens examined. — Total, 36, from: 8 mi. E and 2 mi. S Americanos, 3500 ft., 3; 4 mi. N Acatita, 3600 ft., 9; 20 mi. S El Hundido, 1; Nuevo Mundo, 33 mi. NE Torreon, 3; San Pedro, 2 (BSC); J mi. SW San Pedro de las Colonias, 3700 ft., 4; 10 mi. N and 11 mi. W San Lorenzo, 2; 2 mi. E Torreon, 12. Cratogeomys castanops goldmani Merriam Cratogeomys castanops goldmani Merriam, N. Amer. Fauna, 8:160, January 31, 1895, type from Caiiitas, Zacatecas. Distribution. — Plains of extreme southwestern Coahuila (see fig. 30). Remarks. — Coahuilan specimens assigned to this subspecies com- pare favorably with topotypes of goldmani but are shghtly paler, and in this respect show some relationship to excelsus. The ranges of these two subspecies, however, are partly isolated by mountainous country in southern Coahuila. Both excelsus and subsimus are larger than goldmani. Measurements. — Measurements of one male and one female from the west foot of Pico de Jimulco, respectively, are: total length, 255, 250; length of tail vertebrae, 82, 85; lengtli of hind foot, 36, 35; condylobasal length of skull, 48.9, 46.0; zygomatic breadth, 33.4, 32.6; length of palate, 33.4, 31.4; breadth of rostrum, 11.7, 10.7; length of nasals, 17.7, 16.3; squamosal breadth, 29.6, 27.8; alveolar length of maxillary tooth-row, 9.3, 9.8. Specimens examined. — Total, 6, from: W foot Pico de Jimulco, 4600 ft., 1; Valley Rio Aguanaval, 1 mi. S Jimulco, 4; VA mi. N Parras, 1. Cratogeomys castanops jucundus Russell and Baker Cratogeomys castanops jucundus Russell and Baker, Univ. Kansas Publ., Mus. Nat. Hist., 7:599, March 15, 1955, type from Hermanas, 1205 ft., Coahuila. Distribution. — Arid plains and broad intermontane valleys of east-central Coahuila ( see fig. 30 ) . Remarks. — In external size, jucundus is the largest known cas- tanops but the skull is smaller than in subsimus. Passage to the north and northeast by jucundus is at least partly blocked by in- hospitable mountainous country; the resulting semi-isolation may be one reason for the distinctive characteristics of jucundus com- pared with those of bullatus and ustulatus. Measurements. — Measurements of one adult male and average and extreme measurements of 4 adult females from Hermanas, respectively, are: total length, 311, 296 (294-298); length of tail vertebrae, 80, 85 (83-86); length of hind 228 University of Kansas Publs., Mus. Nat, Hist. foot, 42, 39 (38-39); condylobasal length of skull, 56.9, 50.9 (49.8-51.8); zygo- matic breadth, 38.7, 33.0 (32.1-33.8); length of palate, 40.1, 34.6 (33.8-35.0); breadth of rostrum, 13.3, 11.5 (11.0-11.6); length of nasals, 21.0, 18.0 (17.0- 18.6); squamosal breadth, 32.2, 29.6 (29.0-30.1); alveolar length of maxillary tooth-row, 9.9, 9.4 (9.1-9.6). Specimens examined. — Total, 19, from: Hermanas, 1205 ft., 9; 1 mi. S Her- manas, 2; 1 mi. N and 13 mi. E Cuatro Cienegas, 2; 5 mi. N and 2 mi. W Monclova, 1; 2 mi. N and 1 mi. E Monclova, 1; Monclova, 2 (BSC); Hisachalo [^ Huisachalo], 2. Cratogeomys castanops planifrons Nelson and Goldman Cratogeomys castanops planifrons Nelson and Goldman, Proc. Biol. Soc. Washington, 47:146, June 13, 1934, type from Miquihuana, 5000 ft., Tamaulipas ( Usted, by mistake, as Nuevo Leon ) . Distribution. — Elevated montane valleys of the Sierra Madre Oriental of extreme southeastern Coahuila ( see fig. 30 ) . Remarks. — This subspecies is darker and slightly larger than sub- nubilis but in other ways is most closely related to subnubilis. Cratogeomys c. planifrons is smaller and darker than adjacent sub- species and inhabits both deep and shallow soils of the high, open valleys of the Sierra Madre Oriental; in Coahuila it was taken as low as 7200 feet in elevation and as high as 8700 feet. Measurements. — Measurements of two adult males from 4 miles south and 6 miles east of SaltiUo and average and e.\treme measurements of five adult females from 12 miles west of San Antonio de las Alazanas, respectively, are: total length, 254, 272, 244 (239-247); length of tail vertebrae, 76, 85, 66 (62- 69); length of hind foot, 34, 35, 32 (31-33); condylobasal length of skull, 48.0, 48.8, 43.7 (43.3-44.3); zygomatic breadth, 32.2, 33.2, 28.0 (27.5-28.5); length of palate, 32.6, 34.1, 29.1 (28.7-29.4); breadth of rostrum, 9.8, 10.5, 9.4 (8.9- 9.7); length of nasals, 16.6, 17.5, 14.5 (13.6-15.3); squamosal breadth, 28.0, 29.9, 26.2 (25.3-26.8); alveolar lengtli of maxillary tooth-row, 8.6, 9.5, 8.6 (8.3- 8.9). Specimens examined. — Total, 50, from: 4 mi. S and 6 mi. E SaltiUo, 7500 ft., 7; 7 mi. S and 4 mi. E Bella Uni6n, 7200 ft., 14; 12 7ni. W San Antonio de las Alazanas, 16; 12 mi. S and 2 mi. E Arteaga, 7500 ft., 11; 2 mi. E and 2 mi. N San Antonio de las Alazanas, 8700 ft., 2. Cratogeomys castanops sordidulus Russell and Baker Cratogeomys castanops sordidtdus Russell and Baker, Univ. Kansas Publ., Mus. Nat. Hist., 7:600, March 15, 1955, type from 1.5 mi. NW Ocampo, 3300 ft., Coahuila. Distribution. — Desert plains of north-central Coahuila, surrounded for the most part by higher mountainous country ( see fig. 30 ) . Remarks. — This subspecies, although larger than most adjacent castanops, has a medium-sized, narrow skull and is limited to the Llano de Ocampo, an elevated, desert plain bordered on the west, south and east, by higher mountainous country that seems to bar the Mammals of Coahuila, Mexico 229 passage of this rodent. On the eastern side this barrier extends north to the very banks of the Rio Grande in the Canon de Boquillas. This subspecies, therefore, is in contact with other populations of Crato- geomijs only to the north and northwest. It is not abundant; groups of mounds constructed by one or a few individuals occurred in widely scattered places. Mounds often were small, appeared to be old and, in other ways, were inconspicuous in open desert country dominated by creosote, catclaw and other shrubs. Measurements. — Measurements of one adult male and average and extreme measurements of three adult females from I/2 miles northwest of Ocampo, re- spectively are: total length, 307, 276 (270-288); length of tail vertebrae, 88, 79 (75-85); length of hind foot, 37, 37 (36-39); condylobasal length of skull, 57.5, 50.4 (49.5-51.4); zygomatic breadth, 39.7, 31.7 (30.6-32.4); length of palate, 40.3, 34.6 (33.8-35.2); breadth of rostrum, 13.6, 10.9 (10.3-11.4); length of nasals, 22.1, 18.2 (17.7-18.5); squamosal breadth, 33.1, 30.0 (29.8- 30.1); alveolar length of maxillary tooth-row, 10.3, 9.1 (8.9-9.2). Specimens examined. — Total, 13, from: 50 mi. N and 20 mi. W Ocampo, 4150 ft., 1; 18 mi. S and 14 mi. E Tanque Alvarez, 4000 ft., 4; IY2 mi. NW Ocampo, 3300 ft., 6; Ocampo, 1; 5 mi. N and 19 mi. W Cuatro Cienegas, 3250 ft., 1. Cratogeomys castanops subnubilus Nelson and Goldman Cratogeomys castanops subnubilus Nelson and Goldman, Proc. Biol. Soc. Washington, 47:145, June 13, 1934, type from Cameros, 6800 ft., Coa- huila. Distribution. — Intermontane valleys and high plains of southeastern Coahuila (see fig. 30). Remarks. — According to Russell and Baker (1955:607), sub- nubilus is the smallest subspecies of C. castanops. It is most closely related to planifrons, but subnubilus is smaller and paler. It is dif- ferentiated to a much higher degree from adjacent subspecies than is usual in this species. In the Sierra de Guadalupe, subnubilus was trapped at 6700 feet within two miles of a place where subsimus was taken at 6500 feet. No intergrades between subnubilus and other subspecies have been obtained. Measurements. — Measurements of an adult male and an adult female from 8 miles north of La Ventura, respectively, are: total length, 250, 235; length of tail vertebrae, 79, 76; length of hind foot, 34, 30; condylobasal length of skull, 46.3, 42.2; zygomatic breadth, 31.8, 27.9; length of palate, 31.0, 28.5; breadth of rostrimi, 9.6, 9.0; length of nasals, 16.4, 14.3; squamosal breadth, 28.7, 26.3; alveolar lengdi of maxillary tooth-row, 8.4, 7.8. Specimens examined. — Total, 31, from: 1 mi. N Agua Nueva, 5500 ft., 1; Domingo Canon, Sierra de Guadalupe, 6700 ft., 11 mi. S and 4 mi. W General Cepeda, 1; Cameros, 6800 ft., 6 (BSC); 1 mi. S Cameras, 6000 ft., 4; 2 mi. W San Miguel, 5500 ft., 3; 8 mi. N La Ventura, 6000 ft., 10; La Ventura, 5600 ft., 6 (BSC). 230 University of Kansas Publs., Mus. Nat. Hist. Cratogeomys castanops subsimus Nelson and Goldman Cratogeomys castanops subsimus Nelson and Goldman, Proc. Biol. Soc. Washington, 47:144, June 13, 1934, type from Jaral, Coahuila. Distribution. — Desert plains and lower foothills of mountains of south-central Coahuila (see fig. 30). Remarks. — In cranial dimensions, this subspecies is the largest known of the species but it is exceeded in size of body by C. c. jucundus to the north. Of adjacent subspecies, subsirnus is most closely related to excelsus and shows little or no relationship to the smaller and darker C. c. stibnubilus and C. c. planifrons, found at higher elevations to the south and southeast, respectively. Move- ments by subsimus to the north, east, and south are barred by in- hospitable mountains. Specimens of subsimus from the northeast- ern part of its range are larger and darker than other specimens assigned to this subspecies. Cratogeomys castanops subsimus occurs in scattered colonies in sandy soils principally in the upper drainage of the Rio Salinas. Specimens have been taken also from the foothills of the Sierra Madre Oriental and westward on the elevated desert plains. Measurements. — Measurements of an adult male from the Hacienda El Tul- illo and an adult female from Jaral, respectively, are: total length, 315, 295; length of tail vertebrae, 105, 104; length of hind foot, 40, 40; condylobasal length of skull, 56.4, 53.2; zygomatic breadth, 35.3, 34.1; length of palate, 39.5, 36.9; breadth of rostrum, 12.5, 12.6; length of nasals, 20.8, 18.7; squamosal breadth, 33.8, 29.7; alveolar length of maxillary tooth-row, 10.6, 10.0. Specimens examined. — Total, 22, from: 3 mi. S and 3 mi. E Muralla, 3800 ft., 3; 2 mi. N Santa Cruz, 2; 21 mi. S and 11 mi. E Australia, 4400 ft., 6; Jaral, 3860 ft., 4 (BSC); Hacienda El Tulillo, 5 km. S Hipolito, 2; 17 mi. N and 8 mi. W Saltillo, 5200 ft., 1; 3 mi. N and 5 mi. W La Rosa, 3600 ft., 1; 12 mi. N and 10 mi. E Parras, 5000 ft., 1; N foot Sierra de Guadalupe, 9 mi. S and 5 mi. W General Cepeda, 6200 ft., 1; N foot Sierra de Guadalupe, 10 mi. S and 5 mi. W General Cepeda, 6500 ft., 1. Cratogeomys castanops ustiJatus Russell and Baker Cratogeomys castanops ustulatus Russell and Baker, Univ. Kansas Publ., Mus. Nat. Hist., 7:598, March 15, 1955, type from Don Martin, 800 ft., Coa- huila. Distribution. — Extreme northeastern Coahuila from the vicinity of Presa Don Martin southeastward (see fig. 30). Remarks. — In size of skull, this large pocket gopher is exceeded, among Coahuilan subspecies, only by C. c. subsimus found beyond the mountains in the south. In size, ustulatus differs so markedly from bullatus that the two can be distinguished easily by this feature alone. The skull of C. c. convexus approaches that of ustulatus in size, but is smaller in all respects, save breadth of rostrum. Mammals of Coahuila, Mexico 231 This pocket gopher is common along the Rio Salado and in its watershed. Fallow cotton fields in the vicinity of Anahuac [ = Rodrigues], Nuevo Leon, are preferred living places. Measurements. — Measurements of one adult male and average and extreme measurements of eight adult females from no more than 5 miles from Don Martin, respectively, are: total length, 280, 273 (261-280); length of tail verte- brae, 75, 74 ( 64-83 ) ; length of hind foot, 37, 36 ( 35-38 ) ; condylobasal length of skull, 54.6, 51.4 (50.7-52.1); zygomatic breadth, 37.3, 33.5 (32.6-34.1); length of palate, 38.2, 35.4 (34.8-36.5); breadth of rostrum, 13.7, 11.8 (11.0- 12.5); length of nasals, 20.6, 18.8 (17.8-19.2); squamosal breaddi, 31.8, 30.1 (29.1-30.8); alveolar length of maxillary tooth-rovi^, 10.3, 10.0 (9.3-10.6), Specimens examined. — Total, 10, from: Don Martin, 800 ft., 5; base of Don Martin Dam, 2; 2 mi. SE Don Martin, along Rio Salado, 2; 5 mi. SE Don Martin, 1. Perognathus flavus Silky Pocket Mouse The silky pocket mouse lives on sandy desert soils in western, central and soutliern Coahuila. Rarely was this mouse taken in extreme rocky situations or where vegetation was dense; the animal prefers open places where clumps of plant growth are widely scattered. Mice sel- dom were caught in mouse traps. By using lights near Bella Union we took 20 specimens in two nights by hand. Near Tanque Alvarez, in July, one mouse was spotted at night with a light but eluded the hunter by going into a burrow beneath a dead prickly pear. One of several traps set around the opening of the bur- row caught the animal later that night. The mouse probably walked into one of the traps in- stead of being attracted to the bait of chewed rolled oats. Un- like flavus, other species of Pe- rognathus frequently were taken in traps with their cheek pouches partly filled with bait. As far as could be determined flavus and another small pocket Fig. 31. Distribution in Coahuila of the silky pocket mouse. 1. Perog- nathus flavus meditis. 2. Perognathus flavus pallescens. 232 University of Kansas Publs., Mus. Nat. Hist. mouse, P. merriami, do not occur together in northern Coahuila. In this area flavus is confined to arid flats, adjacent to mountainous topography, where the sparse vegetation is chiefly lechuguilla and creosote bush, whereas, merriami Hves on alluvial soils, usually along arroyos where mesquite is the dominant plant. Three adult males and two adult, non-pregnant females, all from Coahuila, respectively, weighed 5.0, 5.0. 5.5, 5.0, and 5.5 grams. Perognathus flavus medius Baker Perognathus flavus medius Baker, Univ. Kansas Publ., Mus. Nat. Hist., 7: 343, February 15, 1954, type from 1 mi. S and 6 mi. E Rincon de Romos, 6550 ft., Aguascalientes. Distribution. — Intermontane grasslands of southeastern Coahuila (see fig. 31). Remarks. — Specimens from the sparse grasslands in the broad intermontane valleys of southeastern Coahuila are referred to medius on the basis of color and size. Measurements. — Average and extreme measurements of 9 adult males and 11 adult females from 7 miles south and 4 miles east of Bella Union, respec- tively, are: total length, 106 (103-115), 110 (105-118); length of tail, 54 (52- 57), 55 (51-57); length of hind foot, 17 (16-18), 17 (16-18); height of ear from notch, 6 (6-7), 6 (5-7); occipitonasal length, 20.6 (20.0-21.1), 20.2 (19.3- 20.8); frontonasal length, 13.6 (13.2-14.0), 13.4 (13.0-13.8); length of audi- tory bullae, 8.2 (8.1-8.5), 8.2 (7.5-8.6); least interorbital constriction, 4.4 (4.2- 4.6), 4.4 (4.1-4.5); interparietal width, 3.1 (2.6-3.5), 3.0 (2.7-3.3); alveolar length of maxillary tooth-row, 3.1 (2.9-3.1), 3.0 (2.8-3.1). Specimens examined. — Total, 20, from 7 mi. S and 4 mi. E Bella Union, 7200 ft. Perognathus flavus pallescens Baker Perognathus flavus pallescens Baker, Univ. Kansas Publ., Mus. Nat. Hist., 7:345, February 15, 1954, type from 1 mi. SW San Pedro de las Colonias, 3700 ft., Coahuila. Distribution. — Desert areas of western and southern Coahuila (see fig. 31). Remarks. — The subspecies pallescens is characterized by its pale color. Measurements. — Average and extreme measurements of 9 adult males (KU 39226, 40295, 40298, 40306, 40912, 40913, 54555-54557) and 7 adult females (KU 40297, 40300, 40305, 40914, 48547, 54558, 54559), all from southwestern Coahuila, respectively, are: total length, 110 (103-118), 110 (102-119); length of tail vertebrae, 53 (45-62), 53 (48-63); length of hind foot, 16 (15-17), 16 (15-17); height of ear from notch, 6 (5-7), 6 (5-6); occipitonasal length, 20.2 (19.7-20.9), 19.5 (18.8-20.2); frontonasal length, 13.6 (12.9-14.2), 13.0 (12.4- 13.8); length of auditory bullae, 8.0 (7.7-8.1), 7.6 (7.4-8.1); least interorbital constriction, 4.4 (4.1-4.9), 4.2 (3.9-4.9); interparietal width, 3.2 (2.7-3.9), 3.3 (2.9-3.8); alveolar length of maxillary tooth-row, 3.0 (2.9-3.1), 3.0 (2.8- 3.1). Mammals of Coahuila, Mexico 233 Specimens examined. — Total, 37, from: Canon del Cochino, 16 mi. N and 21 mi. E Piedra Blanca, 3200 ft., 1; ^ mi. E Las Margaritas, 2900 ft., 4; 6 mi. N and 2 mi. E La Babia, 3; 11 mi. N and 9 mi. W Tanque Alvarez, 1; 6 mi. NW Tanque Alvarez, 1; 3 mi. NE Sierra Mojada, 4100 ft., 1; 8 mi. N and 25 mi. W Cuatro Cienegas, 4000 ft., 1; 5 mi. N and 19 mi. W Cuatro Cienegas, 3250 ft., 4; 4 mi. N Acatita, 3600 ft., 2; 1 mi. SW San Pedro de las Colonias, 3700 ft., 4; La Pastora Rancho, 41 mi. W and 15 mi. N Saltillo, 2; 3 mi. N and 5 mi. W La Rosa, 1; 12 mi. N and 10 mi. E Parras, 3850 ft., 5; 1 mi. N San Lorenzo, 4200 ft., 4; N foot Sierra Guadalupe, 6400 ft., 10 mi. S and 5 mi. W General Cepeda, 3. Additional record (Baker, 1954:346): 3 mi. NW Cuatro Cienegas. Perognathus hispidus Hispid Pocket Mouse The hispid pocket mouse prefers grasslands to open desert plains and is of more restricted occur- rence in Coahuila than other pocket mice. Most specimens obtained were trapped in grassy situations, either in the vicinity of streams or along fence rows bordering cultivated fields. This pocket mouse is most abundant along the Rio Grande, down- stream from Villa Acuiia, and along the Rio Sabinas. Speci- mens, obtained on sparse grass in the high valleys in extreme southeastern Coahuila reached there from the southwest. Five adult males from north- eastern Coahuila weighed from 31.2 to 43.3 grams. In Coahuila all rodents having external cheek pouches, including pocket mice, pocket gophers and kangaroo rats, commonly are referred to as tuza. Fig. 32. Distribution in Coahuila of the hispid pocket mouse. 1. Perog- nathus hispidus hispidus. 2. Perog- nathus hispidus zacatecae. Perognathus hispidus hispidus Baird Perognathus hispidus Baird, Mammals, in Rep. Expl. Surv. Railr. to Pacific, vol. 8, pt. 1 (1857), p. 421, July 14, 1858, type from Charco Escondido, Tamauhpas. Distribution. — Northeastern Coahuila (see fig. 32). Remarks. — Northeastern Coahuila is within the range of P. h. hispidus as outlined by Glass (1947:176-177). This subspecies reached Coahuila from the Gulf Coastal Plain to the eastward. 234 Univtrsity of Kansas Publs., Mus. Nat. Hist, There seems to be little chance for contact through Coahuila of P. h. hispidus with P. h. paradoxus to the west ( see Glass, loc. cit. ) or with P. h. zacatecae to the southwest because of unsuitable habi- tat. There is no evidence that P, h. paradoxus occurs in western Coahuila, although it is known from Chihuahua. Forty-two skulls from owl pellets from a cave three miles north- west of Cuatro Cienegas, identified by Baker (1953b: 253) as P. hispidus, are here referred, on geographic grounds, to P. h. hispidus. Measurements. — Average and extreme measurements of 4 adult males from 15 miles north and 8 miles west of Piedras Negras and 4 adult females from 2 miles south and 11 miles east of Nava, respectively, are: total length, 203 (195-210), 197 (194-204); length of tail vertebrae, 97 (94-101), 100 (98- 104); length of hind foot, 23 (22-25), 23 (23-24); height of ear from notch, 12 (11-13), 12 (11-13); occipitonasal length of skull, 29.3 (27.7-30.9), 29.0 (28.4-29.5); frontonasal length, 20.0 (18.9-21.2), 20.0 (19.7-20.3); mastoidal breadth, 14.9 (14.2-16.0), 14.5 (14.1-14.7); least interorbital constriction, 7.2 (6.8-7.9), 7.4 (7.3-7.6); interparietal breadth, 7.2 (7.1-7.4), 7.4 (7.1-8.1); alveolar length of upper molariform tooth-row, 4.3 (4.2-4.4), 4.6 (4.5-4.8). Specimens examined. — Total, 29, from: Caiion del Cochino, 3200 ft., 16 mi, N and 21 mi. E Piedra Blanca, 1; 11 mi. W Hacienda San Miguel, 2200 ft., 1; 15 mi. N and 8 mi. W Piedras Negras, 4; 6 mi. N and 2 mi. E La Babia, 3; 1 mi. W Hacienda Las Margaritas, 2; ?i mi. W Hacienda Las Margaritas, 2: 2 mi. S and 11 mi. E Nava, 810 ft., 9; 29 mi. N and 6 mi. E Sabinas, 3; La Gacha, 1600 ft., 2; 2 mi. S and 3 mi. E San Juan de Sabinas, 6; 5 mi. W Nadadores, 3. Additional records: Sabinas (Glass, 1947:177); 3 mi. NW Cuatro Cienegas (Baker, 1953b: 253). Perognathus hispidus zacatecae Osgood Perognathus hispidus zacatecae Osgood, N. Amer. Fauna 18:45, September 20, 1900, type from Valparaiso, Zacatecas. Distribution. — Extreme southeastern Coahuila (see fig. 32). Remarks. — On the basis of dark color and large skull, tvvo speci- mens from an intermontane valley near Bella Union are assigned to zacatecae and extend the known range of zacatecae approxi- mately 230 miles northeast from the nearest point of capture ( 8 miles southeast of Zacatecas, 48666 KU). In Coahuila this subspecies probably occurs only south of the Southern Coahuila Filter-Barrier. Measurements. — One adult female from 7 miles south and 4 miles east of Bella Union measures: total length, 193; length of tail vertebrae, 87; length of hind foot, 22.5; height of ear from notch, 11; occipitonasal length of skull, 31.0; frontonasal length, 21.1; mastoidal breadth, 15.2; least interorbital constriction, 4.6; interparietal breadth, 4.6; alveolar length of upper molariform tooth-row, 7.7. Specimens examined. — Total, 2, from: 3 mi. S and 3 mi. E Bella Union, 6750 ft., 1; 7 mi. S and 4 mi. E Bella Union, 7200 ft., 1. Mammals of Coahuila, Mexico 235 Perognathus merriami Merriam Pocket Mouse This small, yellowish-backed mouse lives on sandy soils on the Gulf Coastal Plain and in northwestern Coahuila. Heavy, rocky soils are barriers to the dispersal of this species, as suggested by Borell and Bryant (1942:24). Denyes (1954:453) has shown ex- perimentally that this mouse is unable to make burrows in hard soil. The species reached northeastern Coahuila from the east and southeast, and reached the northwestern part of the state from the north or northwest. The Sierra del Carmen-Sierra Madre Oriental Axis prevents the eastern and western populations from coming together in Coahuila. Like P. favus, merriami is less attracted than are most other small rodents to mouse traps baited with chewed rolled oats or other foods. This wariness probably accounts for the small number of specimens captured; some animals were caught by hand, by spotting them with a headlight at night. This pocket mouse was taken on deep sand along the Rio Grande (near Guerrero), on and adjacent to fallow, formerly irrigated, fields (vicinity of Monclova), in grass among clumps of prickly pear and mesquite (29 miles north and 6 miles east of Sabinas ) and along mesquite-bordered arroyos (vicinity of Castillon). A pregnant female containing four embryos was trapped on June 12 southeast of the Coahuilan border in Nuevo Leon (5 miles north and 3 miles west of La Gloria). Non-pregnant females were taken in Coahuila in March and April. Average and extreme weights of merriami from northern Coahuila were: for 10 adult males, 7.3 (6.2- 7.9); for 6 adult, non-pregnant females, 6.4 (5.6-7.6). Fig. 33. Distribution in Coahuila of the Merriam pocket mouse. 1. Perog- nathus merriami gilvus. 2. Perogna- thus merriami merriami. 236 University of Kansas Publs., Mus, Nat. Hist. Perognathus merriami gilvus Osgood Perognathus merriami gilvus Osgood, N. Amer. Fauna, 18:22, September 20, 1900, type from Eddy County, New Mexico. Distribution. — Sandy soils in northwestern Coahuila (see fig. 33). Remarks. — Mice from the vicinity of Castillon are referred to gilvus on the basis of pale yellowish upper parts well mixed with black, more slender rostra and larger mastoids. Probably this mouse occurs in alluvial soils bordering most arroyos which lead into the Rio Grande. Specimens of gilvus from Coahuila as listed below were mis-identified earlier as P. flavus pallescens by Baker (1954:345). Measurements. — One adult male and 3 adult females from near Castillon (see below), respectively, measure: total length, 115, 111, 109, 120; length of tail vertebrae, 55, 61, 51, 57; length of hind foot, 15, 16, 15, 17; height of ear from notch, 5, 5, 5, 7; occipitonasal length, 20.0, 20.5, 20.4, 20.7; fronto- nasal length, 13.3, 14.0, 14.1, 14.4; length of auditory bullae, 7.7, 8.0, 7.7, 8.1; least interorbital constriction, 4.8, 4.7, 4.7, 4.9; interparietal width, 3.1, 3.4, 2.8, 3.0; alveolar length of maxillary tooth-row, 3.1, 3.1, 3.3, 3.3. Specimens examined. — Total, 5, from: 6 mi. N and 2 mi. W Castillon, 3750 ft., 1; 2 mi. SSE Castillon, 4050 ft., 4. Perognathus merriami merriami J. A. Allen Perognathus merriami J. A. Allen, Bull. Amer. Mus. Nat. Hist., 4:45, March 25, 1892, type from Brownsville, Cameron County, Texas. Distribution. — Coastal plain of northeastern Coahuila (see fig. 33). Remarks. — Specimens from northeastern Coahuila are assigned to merriami but are generally more yellowish and less dark than specimens from Brownsville, Texas, and Tamauhpas. Coahuilan mice usually have larger bullae, also a tendency toward the char- acters expressed by gilvus. Osgood (1900:22) regards mice from Comstock, Texas, which is across the Rio Grande and upstream ap- proximately 30 miles from Villa Acufia, as merriami but showing intergradation between merriami and gilvus. Measurements. — Average and extreme measurements of 8 adult males (KU 35783, 35784, 35786, 35787, 35791-35793, 35796) and measurements of one adult female (KU 35797) from northeastern Coahuila, respectively, are: total length, 113 (108-122), 119; length of tail vertebrae, 56 (54-60), 60; length of hind foot, 16 (16-17), 16; height of ear from notch, 6 (5-7), 6; occipitonasal length, 20.5 (20.1-21.0), 20.5; frontonasal length, 13.8 (13.5-14.2), 14.2; length of auditory bullae, 7.5 (7.3-7.9), 7.2; least interorbital constriction, 4.5 (4.4-4.7), 4.4; interparietal width, 3.4 (3.0-3.7), 3.0; alveolar length of maxil- lary tooth-row, 3.1 (2.9-3.4), 3.0. Specimens examined. — Total, 23, from: 15 mi. SE Langtry, Val Verde County, Texas, in Coahuila, 2 (BSC); 1 mi. S and 9 mi. W Villa Acuna, 2; 11 mi. W Hacienda San Miguel, 2200 ft., 2; 2 mi. S and 11 mi. E Nava, 810 ft., 2; 5 mi. SE Guerrero, 1; 29 mi. N and 6 mi. E Sabinas, 3; Sabinas, 3 (BSC); Mammals of Coahuila, Mexico 237 10 mi. ESE Sabinas, 1; 10 mi. N Hermanas, 1100 ft., 1; 5 mi. N and 2 mi. W Monclova, 3; 2 mi. N and 1 mi. E Monclova, 1; Monclova, 2 (BSC). Perognathus nelsoni Nelson Pocket Mouse The Nelson pocket mouse is characteristic of the Chihuahuan Des- ert Shrub Vegetation-type. This species is taken rarely on sandy or other fine soils and prefers rocky soils on slopes where cactus, creosote, sotol, and lechuguilla provide scattered cover. A good way to catch nelsoni is to place a baited mouse trap in the midst of, or next to, a clump of lechu- guilla. Normally, this pocket mouse moves no more than a few feet from protective rocks or vegetation (see Dalquest, 1953: 107.) Specimens have been taken in Coahuila as low as 1220 feet in elevation (Panuco) and as high as 6750 feet (3 miles south and 3 miles east of Bella Union ) . Dalquest {op. cit. :108) deter- mined that nelsoni breeds in late summer in San Luis Potosi. Young appear also in spring; pregnant females were examined in Coahuila on March 29 (one female with 3 embryos), April 18 (one with 2 embryos) and April 28 (one with 2 embryos) as well as on June 27 ( one with 5 embryos ) , July 7 ( one with 2 em- bryos) and July 21 (one with 3 embryos). Non-pregnant females were taken in January (2 females), February (2 females), March (4 females), April (6 females), July (14 females), November (1 female ) , December ( 3 females ) . Perognathus nelsoni canescens Merriam Perognathus (Chaetodipus) intermedins canescens Merriam, Proc. Acad. Nat. Sci. Philadelphia, 46:267, September 27, 1894, type from Jaral, Coahuila. Perognathus nelsoni canescens Osgood, N. Amer. Fauna 18:54, September 20, 1900. Distribution. — Desert plains and foothills in northwestern and central Coa- huila, at least as far south as Jaral, the type locality (see fig. 34). Fig. 34. Distribution in Coahuila of the Nelson pocket mouse. 1. Perog- nathus nelsoni canescens. 2. Perogna- thus nelsoni nelsoni. 238 University of Kansas Publs., Mus. Nat. Hist. Remarks. — Perognathus nelsoni canescens is paler and has a smaller rostrum and larger mastoids than P. n. nelsoni of extreme southern Coahuila. Specimens from the vicinities of San Lazaro, La Rosa, Parras, and San Lorenzo are assigned to canescens but are somewhat intermediate between canescens and nelsoni in color and cranial characteristics. One fragmentary skull, recorded as P. nelsoni by Gilmore (1947: 158) from cave deposits from the vicinity of Cuatro Cienegas, is considered here, on geographic grounds, as belonging to the sub- species canescens, as are skulls, identified as P. nelsoni by Baker (1953b:253), from owl pellets taken 3 miles northwest of Cuatro Cienegas. Average and extreme weights of 12 adult males from northwestern Coahuila were 16.8 (13.3-20.4); of 15 adult, non-pregnant females from northwestern Coahuila, 16.2 (13.5-19.0). Measurements. — Average and extreme measurements of 5 adult males from 10 miles south and 5 miles east of Boquillas and 8 adult females from 12 miles north and 10 miles east of Parras, respectively, are: total length, 180 (174- 187), 187 (182-192); length of tail vertebrae, 104 (92-112), 112 (108-121); length of hind foot, 22 (21-23), 21 (20-23); height of ear from notch, 8 (7-9), 10 (9-10); occipitonasal length, 26.0 (25.5-26.7), 25.9 (25.3-26.7); fronto- nasal length, 17.3 (16.9-17.7), 17.4 (16.7-18.1); mastoidal breadth, 13.3 (13.1- 13.5), 13.4 (13.2-13.7); length of auditory bullae, 8.6 (8.2-8.8), 8.1 (7.9-8.3); least interorbital constriction, 6.7 (6.6-6.8), 6.5 (6.1-6.8); interparietal vddth, 7.6 (7.3-7.8), 7.5 (7.3-7.9); alveolar length of upper molariform tooth-row, 3.8 (3.6-3.9), 3.8 (3.5-3.9). Specimens examined. — Total, 70, from: 7 mi. S and 2 mi. E Boquillas, 1800 ft., 2; 10 mi. S and 5 mi. E Boquillas, 1500 ft., 10; 4 mi. W Hacienda La Mari- posa, 2300 ft., 1; 9 mi. NW Don Martin, 2; 2 mi. N and 1 mi. W Ocampo, 4050 ft., 3; 2Ji mi. SE Ocampo, 1; 8 mi. N Hermanas, 1500 ft., 3; 9 mi. E Her- manas, 2000 ft., 4; 3 mi. NE Sierra Mojada, 4100 ft., 2; 8 mi. N and 25 mi. W Cuatro Cienegas, 4000 ft., 2; 5 mi. N and 9 mi. W Cuatro Cienegas, 3250 ft., 3; 9 mi. W and 4 mi. S San Buenaventura, 1800 ft., 1; 3 mi. NW Cuatro Cienegas, 2450 ft., 1; Panuco, 1220 ft., 2; 4 mi. N Acatita, 3600 ft., 5; Acatita, 3600 ft., 1; 2 mi. N San Lazaro, 4800 ft., 64 mi. N and 22 mi. W Saltillo, 1; 3 mi. N and 5 mi. W La Rosa, 3600 ft., 2; 3 mi. SE Torreon, 3800 ft., 3; 12 mi. N and 10 mi. E Parras, 3850 ft., 16; 1 mi. N San Lorenzo, 4200 ft., 7. Additional records (Baker, 1953b: 253): 3 mi. 'NW Cuatro Cienegas; (Gil- more, 1947:158): 25 mi. W and 5 mi. S Cuatro Cienegas; (Osgood, 1900:54): Jaral. Perognathus nelsoni nelsoni Merriam Perognathus (Chaetodipus) nelsoni Merriam, Proc. Acad. Nat. Sci. Phila- delphia, 46:266, September 27, 1894, type from Hacienda La Parada, San Luis Potosi. Distribution. — Desert plains and foothills in extreme southern Coahuila (see fig. 34). Remarks. — Mice, assigned to nelsoni, from the vicinity of Saltillo, the north slope of the Sierra de Guadalupe, and the vicinity of Pico Mammals of Coahuila, Mexico 239 de Jimulco show some relationships to canescens in cranial charac- ters although features resembling neJsoni are more pronounced. One specimen taken at 6750 feet elevation in tlie vicinity of Bella Union was trapped on rocky soil in scrub vegetation in a narrow caiion approximately 50 feet, in elevation, below the beginning of the oak-pifion belt. This record is an indication of the ability of this pocket mouse to live in marginal habitat in the lower mountain valleys and passes. Average and extreme weights of this subspecies from southern Coahuila were: 16 adult males, 17.0 (14.0-20.0); 5 adult, non-preg- nant females, 14.8 (13.0-18.0). Measurements. — Average and extreme measurements of 7 adult males and 2 adult females from the north foot of the Sierra Guadalupe, respectively, are: total length, 186 (183-190) and 174, 187; length of tail vertebrae, 111 (100- 118) and 98, 111; length of hind foot, 24 (23-24) and 23, 23; height of ear from notch, 8 (6-8) and 8, 8; occipitonasal length, 26.1 (25.1-27.1) and 25.9, 25.8; frontonasal length, 17.4 (16.9-17.8) and 17.5, 17.3; mastoidal breadth, 13.6 (12.8-13.9) and 13.6, 12.9; length of auditory bullae, 8.5 (8.3-8.7) and 8.8, 8.3; least interorbital constriction, 6.7 (6.5-6.8) and 6.8, 6.6; interparietal width, 7.9 (7.6-8.3) and 8.3, 7.9; alveolar length of upper molariform tooth-row, 4.0 (3.8-4.1) and 4.0, 3.8. Specimens examined. — Total, 32, from: 17 mi. N and 8 mi. W Saltillo, 5200 ft., 2; 3 mi. S and 3 mi. E Bella Union, 6750 ft., 1; N foot Sierra Guadalupe, 6500 ft., 10 mi. S and 5 mi. W General Cepeda, 20; W foot Pico de Jimulco, 4600 ft., 3; W foot Pico de Jimulco, 5000 ft., 3; 8 mi. N La Ventura, 5500 ft., 2; 2 mi. E and 'A mi. S La Ventura, 5500 ft., 1. Additional records (Osgood, 1900:53): Jimulco, La Ventura, Sierra En- camacion. Perognathus penicillatus eremicus Meams Desert Pocket Mouse Perognathus (Chaetodipus) eremicus Meams, Bull. Amer. Mus. Nat. Hist., 10:300, August 31, 1898, type from Fort Hancock, El Paso County, Texas. Perognathus penicillatus eremicus Osgood, N. Amer. Fauna 18:48, September 20, 1900. Distribution. — Desert plains of Coahuila, except for the northeastern part of the State (see fig. 35). Remarks. — The desert pocket mouse and the Nelson pocket mouse occupy approximately the same parts of Coahuila and often occur together where fine, sandy soils mantle desert areas. Unlike nelsoni, penicillatus rarely lives in coarse, rocky soils. In the vicinity of Boquillas, penicillatus was trapped in deep sand adjacent to the Rio Grande while nelsoni was taken on rocky uplands nearby. Pe- rognathus penicillatus is confined to the Chihuahuan Desert Shrub Vegetation-type; but is less common in this association east of the 240 University of Kansas Publs., Mus. Nat. Hist. Sierra del Carmen-Sierra Madre Oriental Axis than is P. nelsoni. Specimens have been taken at elevations as low as 1300 feet and as high as 5200 feet. Pregnant females were taken on February 9 (one female with 6 embryos ) and on June 30 ( one with 3 embryos ) . Non-pregnant females were trapped in Feb- ruary (9 females), March (8 fe- males), April (5 females), May (1 female), June (3 females), July (4 females), August (2 fe- males), and December (1 fe- male). Average and extreme weights of penicillatus from Coa- huila were: for 18 adult males, 16.3 (13.7-20.0); for 18 adult, non-pregnant females from Coa- huila, 14.6 (11.8-19.0). Specimens from southern Coa- huila are duller colored than topotypes of eremiciis. Speci- mens reported as P. peniciUatiis by Gilmore (1947:158), from cave deposits in the vicinity of Cuatro Cienegas, are here consid- ered, on geographic grounds, as belonging to the subspecies ere- micus. Measurements. — Average and extreme measurements of 8 adult males from 1 mile southv/est of San Pedro de las Colonias and 5 adult females from the west foot of Pico de Jimulco, respectively, are: total length, 181 (177-185), 171 (163-182); length of tail vertebrae, 97 (93-102), 95 (85-104); length of hind foot, 23 (22-23), 23 (21-25); height of ear from notch, 8 (8-8), 7 (7-7); occipitonasal length, 26.1 (25.5-26.8), 25.6 (24.6-26.5); frontonasal length, 18.0 (17.7-18.5), 17.9 (17.3-18.6); mastoidal breadth, 13.3 (13.0-13.5), 12.9 (12.4-13.4); length of auditory bullae, 8.0 (7.7-8.3), 7.9 (7.7-8.2); least inter- orbital constriction, 6.7 (6.5-7.0), 6.4 (6.2-6.6); interparietal breadth, 7.1 (6.7- 8.0), 7.2 (6.9-7.5); alveolar length of upper molariform tooth-row, 3.7 (3.6- 3.8), 3.7 (3.5-4.0). Specimens examined. — Total, 130, from: 1 mi. N Boquillas, 1700 ft., 9; 7 mi. S and 2 mi. E Boquillas, 1800 ft., 5; 10 mi. S and 5 mi. E Boquillas, 1500 ft., 1; 3 mi. S and 8 mi. E Hechicero, 4650 ft.. Chihuahua, in Coahuila, 1; 3 mi. N and 9 mi. E El Pino, 8; 9 mi. N and 6 mi. W Castill6n, 4100 ft., 1; 6 mi. N and 2 mi. W CastilUn, 4; 9 mi. W Castillon, 3640 ft., 1; 2 mi. SSE Castillon, 4050 ft., 2; 50 mi. N and 20 mi. W Ocampo 4150 ft., 1; 7 mi. NE Jaco, 3800 ft., Chihuahua, in Coahuila, 1; 8 mi. E and 2 mi. S Americanos, 1; 4 mi. N San Isidro, 1; 3 mi. NE Sierra Mojada, 4100 ft., 6; 1 mi. S Hermanas, 1300 ft., 3; Fig. 35. Distribution of the desert pocket mouse, Perognathus penicilla- tus eremicus, in Coahuila. Mammals of Coahuila, Mexico 241 5 mi. N and 19 mi. W Cuatro Cienegas, 3250 ft., 1; 6 mi, E Cuatro Cienegas, 2200 ft., 4; 4 mi. N Acatita, 3600 ft., 9; Acatita, 3600 ft., 3; 21 mi. S and 11 mi. E Australia, 1; 1 mi. SW San Pedro de las Colonias, 3700 ft., 26; 1 mi. S and 1 mi. W Coyote, Rio Nazas, 3500 ft., 1; 9 mi. E Torreon, 3600 ft., 10; JO mi. E Torreon, 3700 ft., 1; 17 mi. N and 8 mi. W Saltillo, 5; Hacienda El Tulillo, 5 km. S Hipolito, 3; 12 mi. N and 10 mi. E Parras, 3850 ft., 3; 1 mi. N San Lorenzo, 4200 ft., 6; 4 mi. NNW Saltillo, 5200 ft., 1; W foot Pico de Jimulco, 4600 ft., 11. Additional records (Baker, 1953b:253): 3 mi. NW Cuatro Cienegas; (Gil- more, 1947:158): 25 mi. W and 5 mi. S Cuatro Cienegas, 16 mi. E and 5 mi. S Cuatro Cienegas; (Osgood, 1900:49): Torreon, Jimulco, La Ventura. Dipodomys merriami Merriam Kangaroo Rat The Merriam kangaroo rat is one of the most abundant small mammals of the Lower Sonoran Life-zone, and occurred at most collecting stations in this zone. Specimens examined were caught as low as 810 feet in elevation and as high as 6500 feet. This rat lives in company with Di- podomys ordii on desert flats and in sandy areas but is equally abundant in coarser soils and rocky situations. The ability of the Merriam kangaroo rat to sur- vive in most xeric situations has probably accounted for its wide distribution within the State. The Merriam kangaroo rat digs its burrow usually under protective cover of cactus, creo- sote bush or other shrubby vege- tation. Since burrows are plugged so that openings are usually inconspicuous in the day- time, little evidence of activity of kangaroo rats is visible except for impressions of feet and tail on sandy soils. Merriam kanga- roo rats were trapped on several occasions at the entrances to burrows of Dipodomys nelsoni. Seeds of various grasses, catclaw ( Acacia sp. ) and other plants have been found in cheek pouches. Animals in the process of molt were taken in January, March, July and November. Pregnant females were obtained in January (7 Fig. 36. Distribution in Coahuila of the Merriam kangaroo rat. 1. Dipod- omys merriami atronasus. 2. Dipod- omys merriami merriami. 242 University of Kansas Publs., Mus. Nat. Hist. females), February (3 females), March (5 females), April (2 fe- males), June (4 females), and July (2 females). The usual number of embryos was two; two specimens had one each and one had three. Dipodomys merriami atronasus Merriam Dipodomys merriami atronasus Merriam, Proc. Biol. Soc. Washington, 9:113, June 21, 1894, type from Hacienda La Parada, San Luis Potosi. Distribution. — Desert plains of extreme southeastern Coahuila (see fig. 36). Remarks. — Because of their dark color and large external and cranial dimensions, rats from southeastern Coahuila more closely resemble D. m. atronasus from the vicinity of San Luis Potosi than D. m. merriami from Maricopa County, Arizona. Populations refer- able to atronasus occur at least as far north as 17 miles north of Sal- tillo. Those from tlie north slope of the Sierra de Guadalupe are intermediate, at least in color, between atronasus and merriami and have been referred to D. m. merriami. Specimens from the vicinity of La Rosa, also assigned to D. m. merriami, show no resemblance in coloration to atronasus. Average and extreme weights of atronasus from southeastern Coahuila were: for eleven adult males, 47.8 (43.3-51.9) grams, for five adult, non-pregnant females, 42.0 (38.4- 46.0). Measurements. — Average and extreme measurements of 5 adult males and measurements of 2 adult females from 8 miles north of La Ventura, respectively, are: total length, 255 (248-260) and 242, 248; length of tail vertebrae, 150 (140-158) and 142, 142; length of hind foot, 39 (38-39) and 37, 36; height of ear from notch, 13 ( 13-14) and 12, 11; greatest length of skull, 37.3 (36.7-37.9) and 36.5, 35.1; basilar length, 23.3 (23.1-23.8) and 23.1, 22.0; breadth across maxillary arches, 19.8 (19.5-20.1) and 19.7, 18.4; breadth across auditory bullae, 23.6 (23.3-23.9) and 23.1, 22.3; lengdi of nasals, 13.9 (13.4-14.3) and 13.3, 13.1; least interorbital constriction, 13.1 (12.5-13.5) and 13.1, 12.4; alveolar length of maxillary tooth- row, 4.9 (4.6-5.2) and 4.6, 4.7. Specimens examined. — Total, 31, as follows: 17 mi. N and 8 mi. W Sahillo, 5200 ft., 10; 4 mi. NNW Saltillo, 5200 ft., 9; 10 mi. W Saltillo, 5000 ft., 1; 3 mi. E Arteaga, 5300 ft., 2; 7 mi. S and 1 mi. E Gomez Farias, 6500 ft., 1; 8 mi. N La Ventura, 5500 ft., 8. Dipodomys merriami merriami Meams Dipodomys merriami Meams, Bull. Amer. Mus. Nat. Hist., 2:290, February 21, 1890, type from New River, between Phoenix and Prescott, Maricopa County, Arizona. Distribution. — Desert plains of Coahuila with the exception of the extreme southeastern part (see fig. 36). Remarks. — Geographic variation in Dipodomys merriami needs additional study. Some named subspecies seem well-marked; others have been considered invalid. The subspecies, D. m. merriami, Mammals of Coahuila, Mexico 243 which is currently recognized as occupying an extensive range from Nevada and Cahfornia eastward to Texas and northeastern Mexico, consists of a series of highly variable populations. To this subspe- cies has been assigned most of the Merriam kangaroo rats in Coa- huila, because comparison of them with near topotypes of merriami from Maricopa County, Arizona, reveals no differences. Specimens from the Gulf Coastal Plain of northeastern Coahuila and northern Nuevo Leon have slightly narrower skulls and shorter maxillary tooth-rows than specimens from southwestern Coahuila. Popula- tions in the former area are isolated to a high degree from kangaroo rats in the western and southern part of the State by the Sierra del Carmen-Sierra Madre Oriental Axis. Perhaps the correct relation- ships of these kangaroo rats from northeastern Coahuila and north- em Nuevo Leon will not be known until a revisionary study of D. merriami is completed. Average and extreme weights of 32 adult males from western Coahuila were 45.9 grams (26.8-51.9), of 17 adult, non-pregnant females from western Coahuila, 43.3 (39.1- 57.0). Measurements. — Average and extreme measurements of 11 adult males and 14 adult females from 10 miles east of Torreon, respectively, are: total length, 250 (243-255), 248 (242-255); length of tail vertebrae, 148 (143-155), 146 (137-156); length of hind foot, 38 (37-39), 38 (36-39); height of ear from notch, 14 (14-15), 14 (13-15); greatest length of skull, 36.3 (35.6-37.6), 35.8 (35.1-36.8); basilar length, 23.0 (22.3-24.0); 22.8 (22.2-23.5); breadth across maxillary arches, 20.1 (19.4-21.2), 19.9 (18.8-20.9); breadth across auditory bullae, 23.2 (22.5-24.2), 23.0 (22.1-23.7); length of nasals, 13.4 (12.7-14.2), 13.3 (12.7-14.2); least interorbital constriction, 13.4 (12.3-14.0), 13.1 (12.4- 13.7); alveolar length of maxillary tooth-row, 4.8 (4.6-5.1), 4.8 (4.4-5.1). Specimens examined. — Total, 296, as follows: 11 mi. W Hacienda San Miguel, 2200 ft., 2; 7 mi. S and 2 mi. E Boquillas, 1800 ft., 10; 10 mi. S and 5 mi. E Boquillas, 1500 ft., 15; 15 mi. N and 8 mi. W Piedras Negras, 1; 16 mi. 5 Boquillas, 1600 ft., 2; 17 mi. S and 2 mi. E Boquillas, 1700 ft., 1; 20 mi. S and 5 mi. E Boquillas, 1800 ft., 1; 35 mi. S and 14 mi. E Boquillas, 2350 ft., 2; 9 mi. N and 6 mi. W Castillon, 4100 ft., 1; 3 mi. N and 9 mi. E El Pino, 2; 6 mi. N and 2 mi. W Castillon, 3750 ft., 1; 2 mi. SSE Castillon, 4050 ft., 7; 2 mi. S and 11 mi. E Nava, 810 ft., 1; 50 mi. N and 20 mi. W Ocampo, 4150 ft., 4; 29 mi. N and 6 mi. E Sabinas, 1; La Gacha, 1600 ft., 12; 11 mi. N and 9 mi. W Tanque Alvarez, 4500 ft., 1; 10 mi. ESE Sabinas, 9; 9 mi. NW Don Martin, 1; 8 mi. E and 3 mi. S Americanos, 1; 14 mi. E and 16 mi. N Ocampo, 3; 2 mi. N and 1 mi. W Ocampo, 4050 ft., 2; 5 mi. SE Ocampo, 4700 ft., 2; 3 mi. NE Sierra Mojada, 4100 ft., 2; Sierra Mojada, 4150 ft., 4; 8 mi. N Her- manas, 1500 ft., 7; 8 mi. N and 15 mi. W Cuatro Cienegas, 4000 ft., 1; 5 mi. N and 9 mi. W Cuatro Cienegas, 3250 ft., 9; 3 mi. S and 9 mi. E Cuatro Cienegas, 1; 5 mi. N and 2 mi. W Monclova, 6; Monclova, 1 (BSC); 20 mi. S El Hundido, 2; 4 mi. N Acatita, 3600 ft., 21; Acatita, 3600 ft., 11; 21 mi. S and 11 mi. E Australia, 4400 ft., 5; 3 mi. S and 3 mi. E Muralla, 4000 ft., 1; 1 mi. S San Lazaro, 4000 ft., 3; Trevino, 6 (BSC); 12 mi. N and 10 mi. E Parras, 3850 ft., 9; La Pastora Rancho, 41 mi. W and 15 mi. N Saltillo, 7; Jaral, 1 (BSC); 'A mi. S and 1 mi. E Jaral, 4 mi. S and 2 mi. W Hipolito, 6; Hacienda "El Tulillo", 5 km. S Hipolito, 6; 8 mi. SE San Pedro de las Colonias, 3700 ft., 10; 3 mi. N and 5 mi. W La Rosa, 9; 1 mi. E and 1 mi. S La Rosa, 2; Torreon, 244 University of Kansas Publs., Mus. Nat. Hist. 2 (BSC); 10 mi. E Torreon, 3700 ft., 33; 1 mi. N San Lorenzo, 4000 ft., 27; N foot Sierra Guadalupe, 6400 ft., 10 mi. S and 5 mi. W General Cepeda, 9; W foot Pico de Jimulco, 4600 ft., 6; Jimulco, 7 (BSC). Additional records: 3 mi. NW Cuatro Gienegas (Baker, 1953b:253); ]aral (Elliot, 1907:323). Dipodomys nelsoni Merriam Nelson Kangaroo Rat Dipodomys nelsoni Merriam, Proc. Biol. Soc. Washington, 20:75, July 22, 1907, type from La Ventura, Coahuila. Distribution. — Desert plains of western and southern Coahuila (see fig. 37). Remarks. — The Nelson kangaroo rat occurs in Chihuahuan Desert Shrub Vegetation-type from eastern Chihuahua and northeastern Durango eastward to southeastern Coahuila, southwestern Nuevo Leon, and the northern parts of San Luis Potosi and Zacatecas. In Coahuila, passage to the nortlieast is barred by the Sierra del Carmen-Sierra Madre Oriental Axis but not to the southeast by the Southern Coahuila Filter-Barrier. Like Dipodomys spectahilis, nelsoni constructs low mounds, as large as nine feet in diameter and two feet high. Some mounds have as many as six entrances and are honeycombed with tunnels. One mound examined by me on June 24, 1952, was constructed in sandy soil at a place approxi- mately 32 miles north of Saltillo. Tunnels in the mound were roofed over by thin (less than one inch in some places), crusty coverings which caved-in when stepped on or probed with a shovel. Once this crust of soil caved-in the internal passageways beneath were obscured. I removed loose soil from the burrow system until there was a hole more than four feet across and three feet deep. It was impossible to follow all of the timnels witliin this space but I did find four lateral burrows leading away from the mound and into more substantial soil. No signs of either food, nests or fecal materials were found in the mound. Perhaps such materials were inside cavities or below the level to which I dug. Usually no more than t^vo rats were obtained at any one mound. Unoccupied mounds seemingly cave-in quickly. I purposely stepped on a mound to cave-in one tunnel. The next morning this damage had been repaired by the occupant. Rats cared little for trap baits containing chewed rolled oats. I suspect that most catches resulted because the rats sprung the traps accidentally when they passed over the traps which had been set crosswise in their runways at the entrances to burrows. Single spring steel-traps (size 0) set in burrow openings either with oats sprinkled on the pan or un- baited were as successful as wood-based rat traps in catching the large kangaroo rats. Night hunting by spotting animals with flash Mammals of Coahuila, Mexico 245 lights was a successful means of catching the smaller species of Dipodomys; I was never able to spot nelsoni with a light. The Nelson kangaroo rat was not abundant at any location. Usu- ally in good habitat less than four inhabited mounds were found within normal cruising range of collectors from any one campsite. Only two mounds were found in the vicinity of camps made from Boquillas southward for 30 miles. Oftentimes an inhabited mound would be found surrounded by bare ground for a radius of 50 or 75 feet; evidently these rats have to forage for plant foods over a considerable area under such circumstances. The Nelson kangaroo rat is a large species with a white-tipped tail. Merriam (1907:75) thought that it was related to D. specta- bilis and described nelsoni as being "Much smaller than spectabilis and very much larger than phillipsi group." The two species, nel- soni and spectabilis, do not occur together, but nelsoni was obtained approximately 60 miles from D. spectabilis zygomaticus Goldman (nelsoni from 1 mile north and 3 miles east of Camargo, Chihua- hua, and zygomaticus from 5 miles east of Parral, Chihuahua ) and approximately 90 miles from D. spectabilis spectabilis Mer- riam ( nelsoni from 2 miles north and 6 miles east of Carmago and spectabilis from 4 miles north and 2 miles west of Chihuahua, Chihuahua). Dipodomys nelsoni difiFers from D. s. spectabilis (including subspecies geographically adja- cent, D. s. baileyi Goldman, D. s. cratadon Merriam, D. s. specta- bilis and D. s. zygomaticus) in: size smaller, hind foot shorter, upperparts paler, white tip of tail shorter in relation to total length of tail, skull smaller and narrower, space across maxillary arches narrower in relation to space across auditory bullae, upper incisors narrower, and length of upper molariform tooth-row shorter. Weights of adults (females not pregnant) demonstrate further the diflFerence in size between these two kangaroo rats; average and ex- FiG. 37. Distribution of the Nelson kangaroo rat, Dipodomys nelsoni, in Coahuila. 246 University of Kansas Publs., Mus. Nat. Hist. treme weights of 6 spectabilis from Chihuahua are 124.9 (115-132), of 18 nelsoni from Chihuahua ( Sierra Almagre ) are 86.5 ( 78-102 ) . At the present writing there is no reason to suppose that intergrada- tion between these two kangaroo rats takes place. Dipodomys nelsoni should retain its specific status. Measurements. — Average and extreme measurements of 6 adult males and 4 adult females from La Ventura, respectively, are: total length, 321 (303- 334), 315 (310-330); length of tail vertebrae, 200 (180-211), 193 (186-204); length of hind foot, 49 (48-50), 48 (46-49); greatest length of skull, 42.4 (41.6-43.4), 42.3 (42.0-42.6); basUar length, 28.3 (27.8-28.5), 28.1 (27.6- 28.6); breadth across maxillary arches, 22.8 (22.2-23.3), 22.6 (22.0-23.5); breadth across auditory bullae, 27.0 (26.7-27.3), 26.7 (26.2-27.0); length of nasals, 15.0 (14.6-15.5), 15.0 (14.8-15.2); least interorbital constriction, 14.5 (14.3-14.6), 14.8 (14.3-15.4); alveolar length of maxillary tooth-row, 6.1 (5.8- 6.4), 5.9 (5.6-6.2). Specimens examined. — Total from Coahuila, 65, as follows: 7 mi. S and 2 mi. E Boquillas, 1800 ft., 2; 6 mi. N and 2 mi. W Castillon, 3759 ft., 1; 2 mi. SSE Castillon, 4050 ft., 2; 11 mi. N and 9 mi. W Tanque Alvarez, 4500 ft., 3; 2.5 mi. SE Ocampo, 3300 ft., 1; 2.5 mi. W and 21 mi. S Ocampo, 3500 ft., 3; 4 mi. N Acatita, 3600 ft., 2; Trevifio, 2 (BSC); Jaral, 3 (BSC); La Pastora Rancho, 41 mi. W and 15 mi. N Saltillo, 1; 12 mi. N and 10 mi. E Parras, 5000 ft., 1; 3 mi. N and 5 mi. W La Rosa, 3600 ft., 3; Torreon, 1 (BSC); 4 mi. NNW Saltillo, 5200 ft., 4; 1 mi. N San Lorenzo, 4200 ft., 4; W foot, Pico de Jimulco, 4600 ft., 1; N foot. Sierra Guadalupe, 6500 ft., 10 mi. S and 5 mi. W General Cepeda, 16; 7 mi. S and 1 mi. E Gomez Farias, 6500 ft., 5; La Ventura, 10 (BSC). Total from other Mexican states, 42. Chihuahua: 1 mi. N San Francisco, 4325 ft., 1; Sierra Almagre, 6000 ft., 12 mi. S Jaco, 21; 2 mi. N and 6 mi. E Camargo, 4150 ft., 3; 1 mi. N and 3 mi. E Camargo, 4150 ft., 2; Santa Rosalia [=Camargo], 1 (BSC). Nuevo Leon: Doctor Arroyo, 2 (BSC). Durango: 1 mi. WSW Mapimi, 3800 ft., 3; 2 mi. NW Chocolate, 4500 ft., 3; 2 mi. E Pedricena, 4300 ft., 6. Additional record (Elliot, 1907:325): Jaral. Dipodomys ordii Ord Kangaroo Rat The Ord kangaroo rat is found in desert situations in many parts of Coahuila but is not a common mammal. In most instances both merriami and ordii were taken at tlie same locality. Usually more than twice as many merriami as ordii would be trapped at a given collecting station. One series of 45 ordii was obtained in an elevated intermontane valley in the vicinity of Bella Union. In San Luis Potosi, Dalquest (1953:114) thought that where both species are found together they are equally common but that ordii is less likely to be attracted to traps. In most places ordii avoided rocky soils and preferred sandy soils, often in the vicinity of cultivated fields. In the northern foothills of the Sierra de Guadalupe, Gerd Hcinrich found ordii in open country in the vicinity of cultivated fields; he obtained 12 ordii and 9 merriami in this area. Mammals of Coahuila, Mexico 247 Average and extreme weights of seven adult males from Coahuila were 47.1 grams (40.0-54.0). Two adult, non-pregnant females from Coahuila weighed, 46.0 and 57.7 grams. Specimens were molting in June, July, and August and in November, January, and Feb- ruary. Two females were preg- nant in February; one had two embryos, and the other had four. Dipodomys ordii durranti Setzer Dipodomys ordii fuscus Setzer, Univ. Kansas Publ., Mus. Nat. Hist., 1:555, December 27, 1949, type from Juamave, Ta- maulipas, nee Dipodomys agilis fuscus Boulware, 1943. Dipodomys ordii durranti Setzer, Jour. Washington Acad. Sci., 42: 391, December 17, 1952. Distribution. — Desert plains of northeastern Coahuila and elevated intermontane valleys of extreme south- eastern Coahuila (see fig. 38). Fig. 38. Distribution in Coahuila of the Ord kangaroo rat. 1. Dipodomys ordii durranti. 2. D. ordii idoneus. Remarks. — Two specimens ob- tained in northeastern Coahuila are tentatively referred to dur- ranti, but are paler but no smaller than typical durranti. Animals in this part of Coahuila may have arrived there by way of Nuevo Leon to the southeast; the Sierra del Carmen-Sierra Madre Orien- tal Axis bars their passage from western and southern Coahuila and the Rio Grande seems to bar their passage from the north. In southeastern Coahuila, typical durranti occurs on elevated plains and on broad intermontane valleys. Measurements. — Average and extreme measurements of 9 adult males and 9 adult females from 7 miles south and 4 miles east of Bella Union, respectively, total length, 251 (238-260), 249 (239-264); length of tail vertebrae, are 147 (136-155), 144 (137-157); length of hind foot, 39 (38-40), 39 (37-42); height of ear from notch, 14 (12-16), 14 (13-15); greatest length of skull, 37.5 (36.6-38.3), 37.8 (36.9-38.7); basilar length, 23.4 (22.9-24.2), 23.4 (23.0- 23.8); breadth across maxillary arches, 20.0 (19.5-20.7), 19.8 (19.1-20.2); breadth across auditory bullae, 24.2 (23.5-24.8), 24.3 (23.5-25.4); length of nasals, 13.3 (12.9-14.0), 13.4 (12.7-14.0); least interorbital constriction, 13.5 (12.9-13.9), 13.2 (12.5-13.7); alveolar length of maxillary tooth-row, 4.8 (4.6- 5.0), 4.7 (4.5-5.0). Specimens examined. — Total, 62, from: 11 mi. W Hda. San Miguel, 2200 248 University of Kansas Publs., Mus. Nat. Hist. ft., 1; 1 mi. S Hermanas, 1300 ft., 1; 7 mi. S and 4 mi. E Bella Union, 7200 ft., 47; 12 mi. W San Antonio de las Alazanas, 6; 8 mi. N La Ventura, 5500 ft., 7. Additional records (Setzer, 1949:556): San Juan Neponuceno, 5 mi. N La Ventura; La Ventura. Dipodomys ordii idoneus Setzer Dipodomys ordii idoneus Setzer, Univ. Kansas Publ., Mus. Nat. Hist., 1:546, December 27, 1949, t>Te from San Juan, 12 mi. W Lerdo, 3800 ft., Durango. Distribution. — Desert plains of western and southern Coahuila (see fig. 38). Remarks. — Setzer (1949:546) named idoneus using only three specimens from the type locality but was certainly correct in assum- ing that the probable range of this subspecies {op. cit. -.522, figure 24) extended to the east and north into Coahuila. Ord kangaroo rats obtained for the University of Kansas show that idoneus has an extensive range in southern and western Coahuila and in eastern Chihuahua. Animals from the north slope of the Sierra de Guada- lupe resemble more closely idoneus than the larger and darker durranti of southeastern Coahuila. Specimens from the vicinity of Sierra Mojada are slightly darker than near topotypes of idoneus. Measurements. — Average and extreme measurements of 4 adult males from 9-10 miles east of Torreon and measurements of 2 adult females from 1 mile southwest of San Pedro de las Colonias, respectively, are: total length, 243 (237-251) and 253, — ; length of tail vertebrae, 140 (134-150) and 143, — ; length of hind foot, 37 (37-38) and 38, 36; height of ear from notch, 14 (13- 15) and 14, 14; greatest length of skull, 37.3 (37.0-37.8) and 37.7, 37.3; basilar length, 23.1 (22.8-23.3) and 23.6, 23.0; breadth across maxillary arches, 19.9 (19.6-20.3) and 20.3, 19.5; breadth across auditory bullae, 23.5 (23.2- 23.9) and 24.1, 23.6; length of nasals, 13.7 (13.2-14.2) and 14.3, 13.2; least interorbital constriction, 12.7 (12.4-13.0) and 13.1, 13.0; alveolar length of maxillary tooth-row, 4.8 (4.5-5.0) and 4.8, 4.6. Specimens examined. — Total, 40, as follows: 3 mi. NE Sierra Mojada, 4100 ft., 4; 4 mi. N Acatita, 3600 ft., 2; 1 mi. SW San Pedro de las Colonias, 3700 ft, 5; 8 mi. SE San Pedro de las Colonias, 3700 ft., 1; 9 mi. E Torreon, 3600 ft., 2; 10 mi. E Torreon, 3700 ft., 6; 1 mi. N San Lorenzo, 4200 ft., 8; N foot Sierra Guadalupe, 9 mi. S and 5 mi. W General Cepeda, 6200 ft., 3; N foot Sierra Guadalupe, 10 mi. S and 5 mi. W General Cepeda, 6400 ft., 9. Castor canadensis mexicanus Bailey Beaver Castor canadensis mexicanus Bailey, Proc. Biol. Soc. Washington, 26:191, October 23, 1913, type from Ruidoso Creek, 6 miles below Ruidoso, Lin- coln County, New Mexico. Distribution. — Northern and eastern Coahuila along the Rio Grande and some of its major tributaries (see fig. 39). Remarks. — The beaver occurs along the Rio Grande and its major tributaries in northeastern Coahuila (see Villa-R., 1954). It is most Mammals of Coahulla, Mexico 249 common along the Rio Grande; and evidence of beavers was ob- tained at Boquillas, upstream from Villa Acuna (south of Dryden, Texas), at Piedras Negras, and in the vicinity of Guerrero. One mile north of Boquillas, I saw numerous tracks and cuttings of wil- low in March, 1952. Along the Rio Grande the beaver uses burrows in the bank. Along the Rio de San Diego (two miles west of Ji- menez) old cuttings were seen in June, 1952. Local residents re- ported that a few beaver live upstream on the Rio de San Diego at least as far as a large irrigation dam. Along the watersheds of the Rio Salado and Rio Sabinas beaver still occur in small numbers. According to older residents, beavers were found at one time as far upstream on the Rio Salado (= Rio de los Nadadores) as San Buena- ventura and possibly to Cuatro Cienegas. Mr. D. S. McKellar of the Hacienda La Mariposa told me that beavers formerly lived along the Rio Sabinas upstream as far as Nacimiento and that their dams were observed there in former times. Marsh (1937:38) writes that there were no beaver on the Rio Sabinas near Muzquiz in 1936. Mr. Robert Spence and others of Sabinas reported that at present (1953) the beaver occurs in small numbers at the junction of the Rio Salado and the Rio Sabinas at Presa Don Martin, especially below the irrigation dam. W. Kim Clark (field notes), who camped at Don Martin on August 14-21, 1949, reported seeing sev- eral beaver, and a large beaver dam was seen by me in April, 1954, below the presa. A few animals still occur on the Rio Sabinas between the presa and the town of Sabinas. Down- stream along the Rio Salado, beaver occur in small numbers in Nuevo Leon. Gerd H. Hein- rich (in litt., 12 May 1953) re- ports the presence of beaver ap- proximately 17 miles east of La Gloria in Nuevo Leon in 1953. The beaver is trapped both for its pelt and for its meat which is highly prized as food. The trap- ping of beaver in Coahuila is sporadic but is causing a slow decline Fig. 39. Distribution in Coahuila of Castor canadensis mexicanus. 250 University of Kansas Publs., Mus. Nat. Hist. in the population, especially along the watershed of the Rio Salado and Rio Sabinas. No local efforts to protect the beaver have, to my knowledge, been made. As a result the species has disappeared in many areas and is slowly approaching extinction in the State, ex- cept along the Rio Grande. Measurements of an adult, unsexed skull (partly broken) from Eagle Pass, Maverick County, Texas (directly across the Rio Grande from Piedras Negras), are as follows: occipitonasal length, 137.9; zygomatic breadth, 101.3; least interorbital breadth, 24.1; length of nasals, 49.8; greatest width of nasals, 26.5; alveolar length of upper molariform tooth-row, 32.9. A flat skin in poor condition, obtained five miles south of Don Martin, is extremely pale. Five specimens of beaver from Sabinas, recorded as C. c. frondator by Elliot (1907:190), are considered here, on geographic grounds, as C. c. mexicanus. Specimens examined. — Total, 3, from: Eagle Pass (opposite Piedras Negras), Maverick County, Texas, 1 (skull only); Presa Don Martin, 1 (lower jaw only); 5 mi. SE Don Martin, 1 (skin only). Additional records (Elliot, 1907: 190): Sabinas. Reithrodontomys fulvescens Fulvous Harvest Mouse The fulvous harvest mouse is found from as low as 800 feet in elevation along the Rio Grande to as high as 5300 feet on the western slopes of the Sierra del Carmen, This species prefers a grassy habitat and was found most abundantly on un- grazed areas, especially at the edges of cultivated fields and along streams. At one such place near San Geronimo, 25 fulvous harvest mice in company with Sigmoclon hispidus, Baiomys tay- lori and Peromyscus leucopus were trapped in a single night. In grazed situations, R. ftdves- Fic;. 40. Distribution in Coaluiila of tbe fulvous barvt'st mouse. 1. Reithro- donfomys fulvescens canus. 2. Reith- rodontomys ftdvescens intermedins. cens often lives in the sparse grass that grows within the protection of large clumps of prickly pear. Average and extreme weights of fulvescens from Coahuila were: Mammals of Coahuila, Mexico 251 for 7 non-pregnant females taken in April, 11.2 grams (8.9-13.7); for 18 males taken in April, 11.6 (9.2-13.0). Reithrodontomys fulvescens canus Benson Reithrodontomys fulvescens canus Benson, Proc. Biol. Soc. Washington, 52: 149, October 11, 1939, type from five miles southeast of Chihuahua, Chihuahua. Distribution. — Southern and western Coahuila (see fig. 40). Remarks. — This pale subspecies of R. fulvescens occurs uncom- monly in suitable grassy habitat in western and soutliern Coahuila at lower elevations. Taylor et al. (1945:25) obtained two speci- mens on the western foothills of the Sierra del Carmen in grass near a spring. Mice from the vicinity of Saltillo are classified as canus chiefly because of their pale upper parts. Specimens, assigned tentatively to R. f. griseoflavus Merriam by Hooper (1952:100) from near San Antonio de Jaral, are here considered, on geographic grounds, as belonging to canus. Measurements. — Measurements of 2 adult males and 2 adult females from 1 mile nortli of San Lorenzo, respectively, are: total length, 176, 182 and 166, 162; length of tail vertebrae, 103, 107 and 98, 95; length of hind foot, 21, 20 and 20, 20; height of ear from notch, 15, 15 and 15, 15; greatest length of skull, 21.7, 22.0 and 21.0, 21.4; zygomatic breadth, 10.3, 10.7 and 11.0, 11.0; breadth of braincase, 10.2, 10.2 and 10.6, 10.4; depth of braincase, 8.2, 8.1 and 8.5, 8.1; least interorbital constriction, 3.5, 3.3 and 3.4, 3.6; length of nasals, 8.3, 8.7 and 8.1, 8.2; length of palatine slits, 4.6, 4.6 and 4.3, 4.5; alveolar length of ma.xillary tooth-row, 3.4, 3.6 and 3.5, 3.5. Specimens examined. — Total, 13, from: Campo Centrale, 5300 ft., El Jardin Ranch, 30 mi. S [= 15 mi. SE in Taylor et al, 1945:25] Boquillas, 2 (TCWC); Juarez Canon, 4800 ft.. Sierra del Carmen, 1 (DMNH); 17 mi. N and 8 mi. W Saltillo, 5200 ft., 4; 1 mi. N San Lorenzo, 6. Additional records (Hooper, 1952:98): Sierra del Carmen; (Hooper, op. cit. -.100) : "k mi. S San Antonio de Jaral, 4400 ft. Reithrodontomys fulvescens intermedius J. A. AUen Reithrodontomys mexicanus intermedius J. A. Allen, Bull. Amer. Mus. Nat. Hist., 7:136, May 21, 1895, type from Brownsville, Cameron County, Texas. Reithrodontomys fulvescens intermedius A. H. Howell, N. Amer. Fauna 36: 47, June 5, 1914. Distribution. — Northeastern Coahuila (see fig. 40). Remarks. — Northeast Coahuilan mice are paler and have longer tails but otherwise resemble typical intermedius from Brownsville, Texas. Evidently intermedius occurs as far west and southwest as the Sierra del Carmen-Sierra Madre Oriental Axis. Intergradation between intermedius and canus probably takes place in the vicinity of Puerto San Lazaro, which seems not to bar the passage of this 252 University of Kansas Publs., Mus. Nat. Hist. mouse from one side of the axis to the other. One specimen from within the Puerto (from 2 miles north of San Lazaro) is character- istically intermedius. Four others from 46 miles to the south-south- east (17 miles north and 8 miles west of Saltillo) are assigned to canus but one of these mice is more rufescent than typical canus and approaches intermedius, at least in this respect. Remains of harvest mice, identified as Reithrodontomys fulvescens by Baker (1953b: 253) from owl pellets from three miles northwest of Cuatro Cienegas, are here assigned, on geographic grounds, to intermedius. Measurements. — Average and extreme measurements of 18 adult males and 7 adult females from 6 miles southwest of San Geronimo, respectively, are: total length, 162 (148-177), 161 (153-173); length of tail vertebrae, 93 (79- 104), 92 (86-100); length of hind foot, 20 (18-21), 20 (19-21); height of ear from notch, 15 (14-18), 15 (14-17); greatest length of skull, 21.3 (20.5- 22.3), 21.2 (19.6-22.7); zygomatic breadth, 10.8 (10.1-11.2), 10.7 (10.2- 11.2); breadth of braincase, 10.3 (9.9-10.7), 10.2 (9.9-10.5); depth of brain- case, 8.2 (7.5-8.6), 8.1 (7.8-8.4); least interorbital constriction, 3.3 (3.1-3.5), 3.3 (3.2-3.4); length of nasals, 8.2 (7.6-8.7), 8.3 (7.3-8.9); length of palatine slits, 4.4 (4.2-4.7), 4.4 (3.9-4.9); alveolar length of maxillary tooth-row, 3.5 (3.3-3.7), 3.5 (3.3-3.7). Specimens examined. — Total, 49, from: 9 mi. W and 1 mi. S Villa Acuiia, 2; 2 mi. W Jimenez, 1; 15 mi. N and 8 mi. W Piedras Negras, 4; 6 mi. SW San Geronimo, 25; 29 mi. N and 6 mi. E Sabinas, 5; 3 mi. W Nueva Rosita, 1250 ft., 1; 2 mi. S and 3 mi. E San Juan de Sabinas, 1160 ft., 6; 8 mi. N Hennanas, 1500 ft., 4; 2 mi. N San Lazaro, 64 mi. N and 22 mi. W Saltillo, 1. Additional record (Baker, 1953b:253): 3 mi. NW Cuatro Cienegas. Reithrodontomys megalotis Western Harvest Mouse The western harvest mouse occurs uncommonly both in moun- tainous areas and on desert plains in western and southern Coahuila, but is not known from the Coastal Plain. Like the fulvous harvest mouse, the western harvest mouse prefers grassy situations but the two species were never taken in the same places. One specimen ob- tained south and east of Boquillas lived in association with Sigmo- don hispidus and Peromtjscus eremicus in scattered clumps of grass growing on a dry flat adjacent to an arroyo. This place was sur- rounded by a sparse growth of creosote bush, lechuguilla, mesquite, ocotillo, and other desert plants. Sigmodon and Reithrodontomys, at least, were isolated in this habitat, which may have contained ten acres. Certainly, only a few animals were present; in 220 trap-nights in March, 1952, two Sigmodon and one Reithrodontomys were ob- tained. South and east of Saltillo, mice were trapped in grassy areas on upland valleys, especially along the borders of irrigated fields. Mammals of Coahuila, Mexico 253 At 9000 feet in openings in the boreal forest of the Sierra Madre Oriental, mice were taken in company with Microtus mexicanus. Average and extreme weights of megalotis from Coahuila were: for 14 adult males, 10.7 (8.4- 12.5); for 4 adult females, 11.4 (9.5-13.1). Reithrodontomys megalotis megalotis (Baird) Reithrodon megalotis Baird, in Rep. Expl. Surv. Railr. to Pacific, vol. 8, pt. 1 (1857), p. 451, July 14, 1858, type from between Janos, Chihuahua, and San Luis Springs, Grant Co., New Mex- ico. Reithrodontomys megalotis J. A. Allen, Bull. Amer. Mus. Nat. Hist., 5:79, April 28, 1893. Distribution . — Distributed locally on desert plains and mountains of western and southern Coahuila, except for the higher elevations in the Sierra Madre Oriental (see fig. 41). Remarks. — Coahuilan speci- mens resemble closely megalotis from Hudspeth County, Texas and Pinal County, Arizona. Mice from the foothills of the Sierra Madre Oriental, south and east of Saltillo, are slightly larger and darker, indicating some relationship to R. m. saturatus. Seven specimens, assigned to R. m. megalotis, from the vicinity of San Antonio de las Alazanas are intergrades with saturatus. Most of these mice are browner than typical mega- lotis; three have buffy underparts. Measurements. — Measurements of two adult males and two adult females from 6 miles east and 4 miles south of Saltillo, respectively, are: total length, 151, 151 and 150, 146; length of tail vertebrae, 82, 80 and 79, 75; length of hind foot, 19, 18 and 19, 17; height of ear from notch, 16, 16 and 16, 15; greatest length of skull, 20.8, 20.9 and — , — ; zygomatic breadth, 10.4, 10.9 and 10.5, 10.5; breadth of braincase, 10.2, 10.2 and 10.5, 10.5; depth of braincase, 7.9, 7.9 and 7.9, 7.9; least interorbital constriction, 3.3, 3.1 and 3.3, 3.2; length of nasals, 7.9, 8.3 and — , — ; length of palatine slits, 4.4, 4.5 and 4.2, 4.3; alveolar length of maxillary tooth-row, 3.6, 3.3 and 3.4, 3.4. Specimens examined. — Total, 18, from: 35 mi. S and 14 mi. E Boquillas, 2350 ft., 1; 3 mi. S and 3 mi. E Bella Union, 6750 ft., 2; 6 mi. E and 4 mi. S Saltillo, 7500 ft., 8; 2 mi. N and 2 mi. E San Antonio de las Alazanas, 8700 ft., 7. Additional records (Hooper, 1952:55): Jimulco, Saltillo, Sierra Guadalupe. 5—1204 Fig. 41. Distribution in Coahuila of the western harvest mouse. 1. Reith- rodontomys megalotis inegalotis. 2. Reithrodontomys megalotis saturatus. 254 University of Kansas Publs., Mus. Nat. Hist. Reithrodontomys megalotis saturatus J. A. Allen and Chapman Reithrodontomys saturatus J. A. Allen and Chapman, Bull. Amer. Mus. Nat. Hist., 9:201, June 16, 1897, type from Las Vigas, Veracruz. Reithrodontomus megalotis saturatus A. H. Howell, N. Amer. Fauna, 36:36, June 5, 1914. Distribution. — Known only from the higher parts of the Sierra Madre Orien- tal in extreme southeastern Coahuila (see fig. 41). Remarks. — Three specimens of western harvest mice taken near the crest of the Sierra Madre Oriental, at 9000 feet elevation, are assigned to R. m. saturatus. These resemble topotypes of saturatus in size but color both above and below is slightly paler. Intergrades, assigned to megalotis, and taken approximately 10 miles to the west of the place of capture of saturatus and 300 feet lower in elevation are less dark. The subspecies saturatus occurs in grassy situations bordering coniferous forests. Hooper (1952:58 and 60) records this subspecies near Galeana, Nuevo Leon, at 7000 feet elevation; this place is approximately 30 miles south and east of the locality of cap- ture in Coahuila. Measurements. — Measurements of one adult male from 12 miles east of An- tonio de las Alazanas are: total length, 158; length of tail vertebrae, 83; length of hind foot, 16; height of ear from notch, 14; greatest length of skull, 21.6; zygomatic breadth, 10.4; breadth of braincase, 10.3; depth of braincase, 8.0; least interorbital constriction, 3.0; length of nasals, 8.3; length of palatine slits, 4.6; alveolar length of maxillary tooth-row, 3.3. Specimens examined. — Three from: 12 mi. E San Antonio de las Alazanas, 9000 ft. Peromyscus boylii Brush Mouse The brush mouse is known from the higher parts of the Sierra del Carmen and the Sierra Madre Oriental, southeast of Saltillo, and prefers rocky exposures and scrubby vegetation where yellow pine, Douglas fir and oaks predominate. In the Carmens this mouse was taken in association with Sigmoclon ochrognathus, Neotoma albigtda, and Neotoma mexicana. In the Sierra Madre Oriental the brush mouse is less abundant than Eutamias bulleri, Neotoma mexicana, Peromyscus dijJJciUs, Peromyscus melanotis, and Microtus mexi- canus, which lived in the same general area. Apparently, boylii is more numerous on the eastern slopes of the Sierra Madre Oriental in Nuevo Leon, than on the western exposures of these mountains in Coahuila. In the Sierra del Carmen, Taylor et al. (1945:26), who records their specimens as Peromyscus sp., foimd tlie brush mouse on a rocky hillside in oak cover at Hondido Cafion and in grass and brush adjacent to a spring at Campo Centrale. F. W. Miller of the Mammals of Coahuila, Mexico 255 Dallas Museum of Natural History trapped brush mice as high as 8000 feet elevation, at Campo Madera in the Carmens. An adult male and an adult, non-pregnant female (both with molar teeth slightly worn) from Coahuila, respectively, weighed 26.9 and 28.0 grams. A female taken on October 28 contained four embryos. Peromyscus boylii levipes Merriam Peromtjscus levipes Merriam, Proc. Biol. Soc. Washington, 12:123, April 30, 1898, type from Mount Malinche, Tlaxacala. Peromtjscus hoylei levipes Osgood, N. Amer. Fauna, 28:153, April 17, 1909. Distribution. — Higher elevations of the Sierra Madre Oriental in south- eastern Coahuila ( see fig. 42 ) . Remarks. — ^Two specimens, taken in July, resemble in color, selected specimens of July-taken levipes from two kilometers east of Durango in Hidalgo but are much paler than most specimens examined from that State. Coa- huilan mice are no paler than levipes from the vicinity of Antiguo Morelos, Tamaulipas. This subspecies reached southeastern Coa- huila from the south and seems not to meet the range of rowleyi found in northern Coahuila. Measurements. — One male and one female (see below), both with molar teeth slightly worn, respectively, measure: total length, 196, 197; length of tail vertebrae, 97, 97; length of hind foot, 20, 20; height of ear from notch, 17, 17; greatest length of skull, 26.9, 28.0; basilar length, 20.6, 21.0; zygomatic breadth, 13.0, 12.6; length of nasals, 10.3, 11.3; mastoidal breadth, 11.3, 11.5; length of palatine slits, 5.9, 6.2; alveolar length of maxillary tooth-row, 4.3, 4.5. Specimens examined. — Two from 12 mi. E San Antonio de las Alazanas, 9000 ft. Peromyscus boylii rowleyi (J. A. Allen) Sitomys rowleyi J. A. Allen, Bull. Amer. Mus. Nat. Hist., 5:76, April 28, 1893, type from Noland Ranch, San Juan River, San Juan County, Utah. P[eromyscus]. b[oylii]. rowleyi Mearns, Preliminary diagnoses of new mam- mals from the Mexican border of the United States, p. 3, May 25, 1896 (Preprint of Proc. U. S. Nat. Mus., 19:139, December 21, 1896). Distribution. — Higher elevations in the Sierra del Carmen in northern Coa- huila (see fig. 42). Fig. 42. Distribution in Coahuila of the brush mouse. 1. Peromyscus boylii levipes. 2. Peromyscus boylii rowleyi. 256 University of Kansas Publs., Mus. Nat. Hist. Remarks. — Specimens from the Caimens show no important dif- ferences from rowleyi from New Mexico (Catron and Rio Arriba counties). This subspecies reaches the southeastern Hmit of its range in the Sierra del Carmen and seems not to intergrade with P. b. levipes of extreme southeastern Coahuila. Measurements. — Two males and two females, both with molar teeth slightly worn, and from Hondido Caiion, respectively, measure: total length, 209, 216 and 207, 202; length of tail vertebrae, 111, 121 and 110, 109; length of hind foot, 22.5, 23 and 22, 23; greatest length of skull, 27.4, 28.0 and 27.0, 27.1; basilar length, 20.5, 20.8 and 20.5, 20.2; zygomatic breadth, 13.5, 13.1 and 13.6, 13.1; length of nasals, 10.9, 11.2 and 10.1, 11.0; mastoidal breadth, 11.9, 11.4 and 11.5, 11.6; length of palatine silts, 5.7, 6.0 and 5.8, 5.8; alveolar length of maxillary tooth-row, 4.6, 4.5 and 4.4, 4.3. Specimens examined. — Total, 19, from: Tinaja de Chavez, 6400 ft.. El Jardin Ranch, Sierra del Carmen, 1 (TCWC); Hondido Canon, 6600 ft.. Sierra del Carmen, 5 (TCWC); Campo Centrale, 5300 ft.. El Jardin Ranch, Sierra del Carmen, 5 (TCWC); Botellas Canon, 6300 ft.. Sierra del Carmen, 1 (DMNH); Campo Madera, 8000 ft.. Sierra del Carmen, 6 (DMNH). Peromyscus difficilis difiBcilis (J. A. Allen) Big-eared Rock Mouse Vesperimus difficilis J. A. Allen, Bull. Amer. Mus. Nat. Hist., 3:298, June 30, 1891, type from Sierra de Valparaiso, Zacatecas. [Peromyscus] difficilis Trouessart, Catal. Mamm. viv. foss., fasc. 3, p. 518, 1897. Distribution. — Mountains of southeastern Coahuila (see fig. 43). Remarks. — The big-eared rock mouse occurs in the Montane Mesic Forest Vegetation-type of southeastern Coahuila where it prefers rocky areas. The animal lives less commonly in adjoining, less moist places where chaparral provides the principal vegetative cover. Specimens were obtained as low as 7500 feet in elevation and as high as 9350 feet. Peromyscus difficilis was trapped com- monly in association with Neotoma mexicana, Eiitaniias buUeri, and less commonly in association with P. truei and P. boylii. Average and extreme weights of 12 adult males and 12 adult, non- pregnant females from southeastern Coahuila, respectively, were: 35.0 (28.3-42.8) and 33.3 (28.7-40.0). Pregnant females (each con- taining either 2 or 3 embryos) were trapped July 12, 25, 28, 30, August 2 and 3. A female in lactation was taken on April 25. Coahuilan difficilis are generally grayish, as was noted by Osgood (1909:180), although some specimens are brown. The subspecies, difficilis, has a U-shaped distribution in nortliern Mexico, occurring north in the Sierra Madre Occidental on the western edge of the Central Plateau and north in the Sierra Madre Oriental on the east- ern edge. These two populations are related through mice which Mammals of Coahuila, Mexico 257 live in semi-isolated mountainous areas extending east to west from San Luis Potosi and Hidalgo to Guanajuato and Zacatecas. Measurements. — Average and ex- treme measurements of 5 adult males and 6 adult females from 6 miles east of San Antonio de las Alazanas, re- spectively, are: total length, 230 (223-237), 237 (227-250); length of tail vertebrae, 122 (115-126), 125 (112-138); length of hind foot, 24 (22-25), 24 (23-25); height of ear from notch, 24 (23-28), 24 (23-26); greatest length of skull, 30.1 (29.5- 30.7), 30.5 (29.9-31.3); basilar length, 23.2 (22.8-23.4), 23.4 (22.9-24.4); least interorbital constriction, 4.6 (4.5- 4.8), 4.6 (4.4-4.8); length of nasals, 12.0 (11.5-12.5), 12.3 (11.6-13.1); length of shelf of bony palate, 4.3 (4.2-4.6), 4.4 (4.3-4.6); lengtli of palatine slits, 6.7 (6.4-7.2), 6.8 (6.6- 7.1); length of diastema, 7.8 (7.5- 8.0), 7.9 (7.7-8.0); postpalatal length, 10.9 (10.8-11.0), 11.1 (10.6-11.7); alveolar length of maxillary tooth-row, 4.9 (4.7-5.0), 4.8 (4.7-5.0). Specimens examined. — Total, 81, from: 5 mi. E and 2 mi. S Saltillo, 8000 ft., 1; Diamante Pass, 8500 ft., 3 mi. E and 4 mi. S Saltillo, 9; 6 mi. E San Antonio de las Alazanas, 8000 ft., 4; Mesa de Tablas, 8600 ft., 7; 2 mi. E Mesa de Tablas, 8500 ft., 1; H mi. S Mesa de Tablas, 8600 ft., 8; 2 mi. N and 2 mi. E San Antonio de las Alazanas, 8700 ft., 1; 12 mi. E San Antonio de las Alazanas, 9000 ft., 33; 13 mi. E San Antonio de las Alazanas, 9350 ft., 3; 12 mi. S and 2 mi. E Arteaga, 7500 ft., 5; Sierra Guadalupe, 8000 ft., 11 mi. S and 7 mi. W General Cepeda, 5; Sierra Guada- lupe, 8000 ft., 11 mi. S and 5 mi. W General Cepeda, 4. Additional records (Osgood, 1909:180): Sierra Guadalupe, Carneros, Sierra Encamacion. Peromyscus eremicus Cactus Mouse The cactus mouse occurs at lower elevations in all parts of Coa- huila except for the extreme northeastern part of the State. This species prefers rocky places but occurs both on sandy and clay soils. Where clay soils predominate, eremicus often is the only mouse present. At many localities in northwestern Coahuila eremicus was trapped in rocky places where clumps of lechuguilla, yucca, and cat- claw offered home sites. In the vicinity of Villa Acuiia, Monclova, and Saltillo, eremicus was at home in deeper soils where mesquite and prickly pear provided retreats. The cactus mouse is probably Fig. 43. Distribution of the big-eared rock mouse, Peromyscus difficilis diffi- cilis, in Coahuila. 258 University of Kansas Publs., Mus. Nat. Hist. the most widely distributed, as well as most common, Peromyscus in the State. Average and extreme weights of eremicus from Coahuila were: for 58 adult males (showing wear on their teeth), 21.7 (17.0-26.0); for 19 adult, non-pregnant fe- males, 21.7 (17.0-27.6). Thirty- eight pregnant females contained an average of 2.7 embryos (as few as one embryo in some fe- males and as many as 5 in others ) . Females containing em- bryos were obtained in the fol- lowing months: January (1 fe- male), February (1 female), March (12 females), April (9 females), May (3 females), June (2 females), July (2 females), November (4 females), and De- cember (4 females). In March, 12 of 19 females taken were pregnant. In the months of June, July, November and December the number of young individuals (showing no wear on their teeth) taken exceeded the number of adults captured. Of 301 eremicus trapped in Coahuila, 175 (57 per cent) were males and 132 (43 per cent) were females. Approximately one-half of this entire sample of eremicus was taken in March and April, in which period almost twice as many males as females were collected. Since almost half ( 43 per cent ) of the females obtained in March and April were pregnant, it is possi- ble that relatively fewer females than males were taken because females were less active owing to pregnancy or the presence of young in nests. Peromyscus eremicus eremicus (Baird) Hesperomys eremicus Baird, Mammals, in Rep. Expl. Surv. Railr. to Pacific, vol. 8, pt. 1 (Washington, 1857), p. 479, July 14, 1858, type from Old Fort Yuma, Imperial County, California. Peromyscus eremicus J. A. Allen, Bull. Amer. Mus. Nat. Hist., 7:226, June 1895. Distribution. — Arid lowlands in northern and western Coahuila (see fig. 44). Fig. 44. Distribution in Coahuila of the cactus mouse. 1. Peromyscus eremicus eremicus. 2. Peromyscus eremicus phaeurus. Mammals of Coahuila, Mexico 259 Remarks. — Mice from northern and western Coahuila are classi- fied as P. e. eremicus but are slightly paler and have slightly shorter upper molar tooth-rows than examples of eremicus from Yuma County, Arizona, and from western Texas. In southern Coahuila, the mice are darker, and the upper parts are more heavily mixed with black; this darker color is an approach to P. e. phaeurus Osgood. Three skulls from owl pellets taken from a cave three miles north- east of Cuatro Cienegas, identified by Baker (1953b: 253) as P. eremicus, are here referred, on geographic grounds, to P. e. eremicus. This subspecies is not barred from passage to northeastern Coa- huila by the Sierra del Carmen-Sierra Madre Oriental Axis and oc- curs commonly in parts of northeastern Coahuila in the Chihuahuan Desert Shrub Vegetation-type and adjacent parts of the Tamaulipan Thorn Shrub Vegetation-type. Measurements. — Average and extreme measurements of 9 adult males and 5 adult females from no more than 21 miles from Boquillas, respectively, are: total length, 183 (174-202), 182 (172-189); length of tail vertebrae, 98 (92- 106), 101 (98-104); length of hind foot, 20 (18-21), 20 (19-21); height of ear from notch, 19 (18-21), 20 (18-21); basilar length of skull, 19.0 (18.6-19.3), 19.0 (18.3-19.5); zygomatic breadth, 12.5 (12.3-13.2), 12.5 (12.2-13.1); length of nasals, 8.9 (8.6-9.4), 9.0 (8.6-9.2); least interorbital constriction, 4.0 (3.9- 4.2), 4.1 (3.9-4.3); mastoidal breadth, 11.0 (10.6-11.3), 10.9 (10.7-11.3); length of palatine slits, 5.0 (4.7-5.2), 4.8 (4.4-5.1); alveolar length of maxillary tooth-row, 3.8 (3.7-3.9), 3.7 (3.4-3.9). Specimens examined. — Total, 202, from: 9 mi. W and 1 mi. S Villa Acuiia, 9; Canon del Cochino, 3200 ft., 16 mi. N and 21 mi. E Piedra Blanca, 23; 11 mi. W Hacienda San Miguel, 2200 ft., 12; 1 mi. N Boquillas, 1700 ft., 6; 7 mi. S and 2 mi. E Boquillas, 1800 ft., 8; 10 mi. S and 5 mi. E Boquillas, 5; 17 mi. S and 2 mi. E Boquillas, 3; 20 mi. S and 5 mi. E Boquillas, 1; 15 mi. N and 8 mi. W Piedras Negras, 16; 35 mi. S and 14 mi. E Boquillas, 2350 ft., 3; Sierra de la Encantada, 4100 ft., 37 mi. S and 21 mi. E Boquillas, 1; 6 mi. N and 2 mi. E La Babia, 5; 9 mi. N and 6 mi. W CasUllon, 4100 ft., 2; 2 mi. SSE Castillon, 4050 ft., 1; Sierra del Pino, 6200 ft., 5 mi. S and 3 mi. W Acebuches, 1; La Gacha, 1600 ft., 3; 50 mi. N and 20 mi. W Ocampo, 4150 ft., 3; 18 mi. S and 14 mi. E Tanque Alvarez, 4000 ft., 2; 3 mi. N and 4 mi. E San Francisco, 4850 ft., 2; 14 mi. E and 16 mi. N Ocampo, 1; 3 mi. NE Sierra Mojada, 4100 ft., 1; 2'A mi. SE Ocampo, 1; 2M mi. W and 21 mi. S Ocampo, 3500 ft., 5; 5 mi. W and 22 mi. S Ocampo, 6000 ft., 1; 8 mi. N and 25 mi. W Cuatro Cienegas, 4000 ft., 1; 5 mi. N and 19 mi. W Cuatro Cienegas, 3250 ft., 1; 4 mi. N Acatita, 3600 ft., 12; Acatita, 3600 ft, 1; 20 mi. S El Hundido, 3; 21 mi. S and 11 mi. E Austraha, 4400 ft., 3; 8 mi. SE San Pedro de las Colonias, 3700 ft., 2; La Pastora Rancho, 41 mi. W and 15 mi. N Saltillo, 2; Hacienda El Tulillo, 5 km. S Hipolito, 1; 3 mi. N and 5 mi. W La Rosa, 3600 ft., 11; 1 mi. E and 1 mi. S La Rosa, 3; 12 mi. N and 10 mi. E Parras, 3850 ft., 12; 4 mi. N and 1 mi. W San Lorenzo, 2; 1 mi. N San Lorenzo, 4200 ft., 26; 3 mi. SE Torreon, 3800 ft., 1; VA mi. N Parras, 5500 ft., 1; N foot Sierra Guadalupe, 6200 ft., 9 mi. S and 5 mi. W General Cepeda, 2; iV foot Sierra Guadalupe, 10 mi. S and 5 mi. W General Cepeda, 1; W foot Pico de Jimulco, 4600-5000 ft., 11. Additional records (Osgood, 1909:242): Jaral, Torreon, Jimulco; (Baker, 1953b: 253): 3 mi. NW Cuatro Cienegas. 260 University of Kansas Publs., Mus. Nat. Hist. Peromyscus eremicus phaeurus Osgood Peromyscus eremicus phaeurus Osgood, Proc. Biol. Soc. Washington, 17:75, March 21, 1904, type from Hacienda La Parada, San Luis Potosi. Distribution. — Arid lowlands of southeastern Coahuila (see fig. 44). Remarks. — Specimens from southeastern Coahuila are as dark as phaeurus from San Luis Potosi. Mice from the vicinity of Sabinas and Don Martin are intermediate in color but seem closer to phaeurus, to which they are assigned here. Mice from Monclova and Carneros, referred to P. e. eremicus by Osgood (1909:242), are here assigned, on geographic grounds, to phaeurus. Specimens at hand from the vicinity of Monclova show that mice from that area are nearest to phaeurus. A specimen from Sierra Encarnacion, in- correctly listed by Osgood {loc. cit.) as being from Nuevo Leon and as eremicus, is also referred, on geographic grounds, to phae- urus. Measurements. — Average and extreme measurements of 10 adult males and 8 adult females from 17 miles north and 8 miles west of Saltillo, respectively, are: total length, 183 (170-194), 182 (174-197); length of tail vertebrae, 97 (90-103), 94 (88-101); length of hind foot, 20 (18-21), 20 (19-21); height of ear from notch, 19 (17-19), 19 (17-20); basilar lengtli of skuU, 19.1 (18.8- 19.6), 19.0 (18.5-19.5); zygomatic breadth, 12.7 (12.4-13.0), 12.6 (12.3-13.0); length of nasals, 9.1 (8.8-9.4), 9.1 (8.7-9.4); least interorbital constriction, 4.0 (3.9-4.1), 4.0 (3.8-4.1); mastoidal breadth, 11.0 (10.8-11.3), 11.0 (10.5-11.6); length of palatine slits, 4.8 ( 4.8-5.0 ) , 4.9 ( 4.8-5.2 ) ; alveolar length of ma.\illary tooth-row, 3.8 (3.7-3.9), 3.8 (3.7-3.9). Specimens examined. — Total, 99, from: 10 mi. ESE Sabinas, 1; 9 mi. NW Don Martin, 5; base Don Martin Dam, 3; 8 mi. N Hermanas, 1500 ft., 11; 9 mi. W and 4 mi. S San Buenaventura, 1800 ft., 14; 5 mi. N and 2 mi. W Monclova, 5; Panuco, 8; 2 mi. S San Lazaro, 60 mi. N and 22 mi. W Saltillo, 2; 3 mi. S and 3 mi. E Muralla, 4000 ft., 2; 17 mi. N and 8 mi. W Saltillo, 5200 ft., 27; 4 mi. NNW Saltillo, 5200 ft., 9; 10 mi. W Saltillo, 6000 ft., 4; 1 mi. S Carneros, 6000 ft., 2; 7 mi. S and 1 mi. E Gomez Farias, 6500 ft., 1; 8 mi. N La Ventura. 5500 ft., 5. Additional records (Osgood, 1909:242): Monclova, Cameras, Sierra En- carnacion; (Osgood, 1909:251): Sabinas, Saltillo. Peromyscus leucopus Woods Mouse The woods mouse clearly is common in northeastern Coahuila but is rare in the western and southern part of the State. Although obtained at elevations of more than 3000 feet in the foothills of mountainous areas, the woods mouse prefers lowlands where it is most abundant in grassy or brushy cover along arroyos. Grass pro- tected from grazing within clumps of prickly pear offer sufficient protection to leucopus. One-night catches of 21, 24 and 30 indi- Mammals of Coahuila, Mexico 261 viduals were obtained in such situations, when approximately 200 museum special traps were set. West of the Sierra del Carmen-Sierra Madre Oriental Axis leu- copus is scarce. Evidently it occurs in suitable brushy and grassy areas along arroyos. One specimen was obtained adjacent to the Rio Nazas near San Pedro de las Colonias. No specimens were taken along the Rio Grande at Boquillas, although Borell and Bryant (1942:32) record Zewcopw5 on the Texas side ". . . among tules and cane along the river bottom." Average and extreme weights of leucopus from northeastern Coa- huila were: for 37 adult males, 23.0 (17.8-26.6); for 23 adult, non-pregnant females, 22.1 (15.2- 29.5). Pregnant females con- tained an average of 3.6 em- bryos; females with embryos were taken on March 24 (1 fe- male with 2 embryos), March 31 (1 with 3, 1 with 4), April 6 (1 with 5 ) , April 9 ( 1 with 3 ) , April 10 (1 with 2), June 19 (1 with 4), June 22 (1 with 4), June 23 ( 1 with 3 ) , November 6 ( 1 with 6 ) , December 9 ( 1 with 5 ) . Fe- males in lactation were taken on January 8, 15, April 3, and De- cember 2. Peromyscus leucopus texanus ( Woodhouse ) Woods Mouse Fig. 45. Distribution in Coahuila of the woods mouse. 1. Peromyscus leu- copus texanus. 2. Peromyscus leuco- pus tornillo. Hesperomys texana Woodhouse, Proc. Acad. Nat. Sci. Philadel- phia, vol. 6 (1852-53), p. 242, 1853, type from Mason, Mason County, Texas. Peromyscus leucopus texanus Osgood, N. Amer. Fauna, 28:127, April 17 1909. Distribution. — Coastal plain and adjacent mountainous foothills of north- eastern Coahuila (see fig. 45). Remarks. — Specimens from NE Coahuila resemble closely P. I. texanus from Bexar and Webb counties, Texas. There seems to be little chance, because of inhospitable mountainous country, for leucopus to spread south and west in Coahuila; consequently, no intergradation between P. I. texanus and P. I. tornillo is demon- 262 University of Kansas Publs., Mus. Nat. Hist. strable in the State. The woods mouse does occur along the Rio Grande at least as far upstream as the vicinity of Villa Acuiia. Up- stream from this place steep-walled carious allow for no habitat suit- able for leucopus. Measurements. — Average and extreme measurements of 7 adult males and 7 adult females from 14 miles north and 8 miles west of Piedras Negras, re- spectively, are: total length, 173 (161-186), 168 (160-177); length of tail vertebrae, 80 (73-88), 75 (72-84); length of hind foot, 22 (21-23), 22 (21-23); height of ear from notch, 17 (16-19), 17 (16-18); basilar length of skull, 20.4 (19.7-21.0), 20.4 (20.0-20.7); zygomatic breadth, 13.6 (13.3-14.0), 13.5 (13.2- 13.8); length of nasals, 10.6 (10.0-10.9), 10.7 (10.2-11.2); least interorbital constriction, 4.2 (4.1-4.5), 4.2 (4.0-4.3); mastoidal breadth, 11.3 (10.8-11.7), 11.2 (10.8-11.8); length of palatine slits, 5.1 (4.6-5.5), 5.1 (4.8-5.4); alveolar length of maxillary tooth-row, 3.8 (3.7-4.0), 3.7 (3.6-4.0). Specimens examined. — Total, 196, from: 9 mi. W and 1 mi. S Villa Acuna, 11; Caiion del Cochino, 3200 ft., 16 mi. N and 21 mi. E Piedra Blanca, 30; 11 mi. W San Miguel, 2200 ft., 7; 3M mi. N Jimenez, 850 ft., 1; 2 mi. W Jimenez, 850 ft., 3; 15 mi. N and 8 mi. W Piedras Negras, 24; 8 mi. N and 2 mi. W Piedras Negras, 1; 1 mi. NW Hacienda Las Margaritas, 3000 ft., 5; 1 mi. W Hacienda Las Margaritas, 3000 ft., 5; /i mi. \V Hacienda Las Margaritas, 2900 ft., 2; Hacienda Las Margaritas, 2900 ft., 1; 'A mi. E Hacienda Las Margaritas, 3000 ft., 1; 6 mi. N and 2 mi. E La Babia, 2; 6 mi. SW San Geronimo, 28; 2 mi. S and 11 mi. E Nava, 810 ft., 5; 29 mi. N and 6 mi. E Sabinas, 2; 4 mi. W Hacienda La Mariposa, 2300 ft., 11; 10 mi. E Hacienda La Mariposa, 3000 ft., 4; 4 mi. W and 2 mi. S Hacienda La Mariposa, 2300 ft., 2; La Gacha, 1600 ft., 17; 8 mi. N and 4 mi. W Muzquiz, 1800 ft., 2; 2 mi. S and 3 mi. E San Juan de Sabinas, 1160 ft., 17; 10 mi. ESE Sabinas, 2; 9 mi. NW Don Martin, 1; Don Martin, 800 ft., 1; 5 mi. SE Don Martin, 2; 10 mi. N Hermanas, 1100 ft., 1; 1 mi. S Hermanas, 1300 ft., 6; 5 mi. N and 2 mi. W Monclova, 1; 6 mi. E Cuatro Cienegas, 2200 ft., 1. Additional records (Osgood, 1909:131): Sabinas. Peromyscus leucopus tomillo Meams Peromyscus tomillo Meams, Preliminary diagnosis of new mammals from the Mexican border of the United States, p. 3, March 25, 1896 (Preprint of Proc. U. S. Nat. Museum. 18:445, May 23, 1896), type from Rio Grande, about 6 miles above El Paso, Texas. Peromyscus leucopus tomillo Osgood, N. Amer. Fauna, 28:125, April 17, 1909. Distribution. — In suitable habitats in western Coahuila (see fig. 45). Remarks. — Peromyscus leucopus tomillo is rare in western Coa- huila. One specimen was taken on February 8 near the Rio Nazas in the vicinity of San Pedro de las Colonias. This subspecies is prob- ably restricted to shrubs and grasses occurring along arroyos. The single specimen is referable to tomillo although the skull, which has slightly worn teeth, is smaller than skulls of tomillo, of equal age, from Santa Fe County, New Mexico. Measurements. — The male (teeth slightly worn) measures: total length, 157; length of tail vertebrae, 72; length of hind foot, 21; height of ear from notch, 17; basilar length of skull, 18.4; zygomatic breadth, 12.3; length of nasals, 9.0; least interorbital constriction, 3.9 mastoidal breadth, 10.3; length of palatine slits, 4.7; alveolar length of maxillary tooth-row, 3.9. Mammals of Coahuila, Mexico 263 Specimen examined. — One from 1 mi. SW San Pedro de las Colonias, 3700 ft. Peromyscus maniculatus blandus Osgood Deer Mouse Peromyscus sonoriensis blandus Osgood, Proc. Biol. Soc. Washington, 17:56, March 21, 1904, type from Escalon, Chihuahua. Peromyscus maniculatus blandus Osgood, N. Amer. Fauna, 28:84, April 17, 1909. Distribution. — Western and southern Coahuila (see fig. 46). Remarks. — The deer mouse occurs in Coahuila west of the Sierra del Carmen-Sierra Madre Oriental Axis. This species seems lo- calized in its distribution on both desert plains and foothills of the mountains; specimens were taken as low as 3400 feet in elevation and as high as 8600 feet. Deer mice usually were found along arroyos where deep soils occurred and where mesquite, cacti, and some grassy cover existed. The animals seldom were trapped in thick stands of lechuguilla that often grew at borders of arroyos. In mountain valleys, maniculatus was trapped in low-growing shrubs and in clumps of prickly pear, which often is planted to enclose irrigated fields. Average and extreme weights of 7naniculatus from western Coahuila were: for 14 adult males, 21.6 (17.0-27.0); for 11 adult, non-pregnant females, 22.0 (17.0-25.1). Pregnant females, containing an average of 4.1 em- bryos, were obtained on March 19 (2 females with 3 embryos each), March 24 (1 with 4), April 27 (1 with 3), April 29 (1 with 4), June 29 (1 with 4), and November 11 (1 with 8). Fe- males in lactation were taken on April 29 (2), July 5 (2), and November 11 (2). Specimens from Coahuila are not distinguishable from P. m. blandus from Chihuahua al- FiG. 46. Distribution of the deer mouse, Peromyscus maniculatus blan- dus, in Coahuila. though some, but not all, skins from the foothills of the Sierra Madre Oriental, southeast of Saltillo, are slightly darker on the upper parts than are other examples of blandus. Specimens were obtained in 264 University of Kansas Publs., Mus. Nat. Hist. both gray and buff phases; the pale tufts of hair at the anterodorsal bases of the pinnae are conspicuous in most examples of hlandus. One skull from an owl pellet taken from a cave three miles north- west of Cuatro Cienegas, identified by Baker (1953b: 253) as Pero- myscus maniculatus, is here referred, on geographic grounds, to P. m. blandus. Measurements. — Average and extreme measurements of 10 adult males and 9 adult females from the north foot of the Sierra Guadalupe, respectively, are: total length, 152 (138-167), 153 (142-165); length of tail vertebrae, 68 (62- 78), 69 (65-77); length of hind foot, 21 (20-22), 22 (21-23.5); height of ear from notch, 18 (17-19), 18 (17-19); basilar length of skull, 19.6 (18.7-21.1), 19.5 (18.4-20.4); zygomatic breadth, 12.9 (12.3-13.9), 13.0 (12.5-13.4); length of nasals, 10.2 (9.6-10.9), 10.1 (9.6-10.6); least interorbital constriction, 3.9 (3.6-4.3), 3.9 (3.7-4.1); mastoidal breadth, 10.6 (10.1-11.3), 10.6 (10.4-11.0); length of palatine slits, 5.6 (5.3-5.9), 5.6 (5.3-5.9); alveolar length of ma.xillary tooth-row, 4.0 (3.8-4.3), 4.0 (3.8-4.2). Specimens examined. — Total, 93, from: Campo Centrale, 5300 ft., Sierra del Carmen, 30 mi. S Boquillas, 1 (TCWC); 9 mi. N and 6 mi. W Castillon, 4100 ft., 2; 2 mi. SSE Castillon, 4050 ft., 1; 50 mi. N and 20 mi. W Ocampo, 4150 ft., 1; 11 mi. N and 9 mi. W Tanque Alvarez, 4500 ft., 2; 6 mi. NW Tanque Al- varez, 3400 ft., 1; 3 mi. S San Lazaro, 60 mi. N and 22 mi. W Saltillo, 1; 17 mi. N and 8 mi. W Saltillo, 5200 ft., 6; 12 mi. N and 10 mi. E Parras, 5000 ft., 2; 4 mi. N and 1 mi. W San Lorenzo, 5500 ft., 1; 9 mi. E Torreon, 3600 ft., 1; 1 mi. N San Lorenzo, 4200 ft., 15; 4 mi. NNW Saltillo, 5200 ft., 16; 6 mi. E and 4 mi. S Saltillo, 7500 ft., 5; 3 mi. S and 3 mi. E Bella Union, 6750 ft., 1; 7 mi. 5 and 4 mi. E Bella Union, 7200 ft., 5; Mesa de Tablas, 8600 ft., 1; N foot Sierra Guadalupe, 6200 ft., 9 mi. S and 5 mi. W General Cepeda, 2; N foot Sierra Guadalupe, 6200-6500 ft., 10 mi. S and 5 mi. W General Cepeda, 17; 1 mi. S Cameros, 6000 ft., 1; 7 mi. S and 1 mi. E Gomez Farias, 6500 ft., 8. Additional records (Baker, 1953b: 253): 3 mi. NW Cuatro Cienegas; (Os- good, 1909:85): Saltillo, Jimulco, La Ventura. Peromyscus melanophrys coahuiliensis Baker Plateau Mouse Peromyscus melanophrys coahuiliensis Baker, Univ. Kansas Publ., Mus. Nat. Hist., 5:257, April 10, 1952, type from 7 mi. S and 1 mi. E Gomez Farias, 6500 ft., Coahuila. Distribution. — Desert areas of southeastern Coahuila (see fig. 47). Remarks. — The plateau mouse reaches the northern limit of its distribution in southern Coahuila. It is safe to say that the mouse is uncommon in the State because only nine specimens were trapped in the many hundreds of trapnights in habitats where the species might be expected to live. All individuals were taken in arid situa- tions where yucca, cholla and other cacti and thorny shrubs were abundant. An adult male and three adult, non-pregnant females from south- eastern Coahuila, respectively, weighed 50.0, 51.5, 54.0, and 58.4 grams. A female, taken on March 29. was lactating. Mammals of Coahuila, Mexico 265 This subspecies resembles P. m. consobrinus Osgood, found to the southward, more than any other subspecies. No specimens of this species are known from south- western Coahuila; if the mouse does occur there, specimens may show relationships to, or be re- ferrable to, P. m. xenurus Os- good of Durango. Measurements. — One male (KU 33248) and two females (KU 35018, 35019), with molar teeth moderately worn, from southeastern Coahuila, re- spectively, measure: total length, 247, 256, 273; length of tail vertebrae, 130, 135, 149; length of hind foot, 26, 26, 27; height of ear from notch, 23, 23, 25; basilar length of skull, 24.0, 23.7, 24.8; least interorbital constriction, 4.7, 4.7, 4.9; length of nasals, 10.3, 11.3, 11.8; length of shelf of bony palate, 4.6, 4.5, 5.0; length of palatine slits, 6.5, 6.5, 6.1; length of diastema, 8.1, 8.2, 8.3; alveolar length of maxil- lary tooth-row, 4.5, 4.4, 4.7. Fig. 47. Distribution of the plateau mouse, Peromyscus melanophrys coa- huiliensis, in Coahuila. Specimens examined. — Total, 9, from: 17 mi. N and 8 mi. W Saltillo, 1; 7 mi. S and 1 mi. E Gomez Farias, 4; 8 mi. N La Ventura, 3; 2 mi. E and Yi mi. S La Ventura, 1. Peromyscus melanotis J. A. Allen and Chapman Black-eared Woods Mouse Peromyscus melanotis ]. A. Allen and Chapman, Bull. Amer. Mus. Nat. Hist., 9:203, June 16, 1897, type from Las Vigas, Veracruz. Distribution. — Boreal forests of the Sierra Madre Oriental in southeastern Coahuila (see fig. 48). Remarks. — Peromyscus melanotis lives in the Subalpine Humid Forest Vegetation-type and adjacent areas of the Montane Mesic Forest Vegetation-type in the Sierra Madre Oriental southeast of Saltillo. Specimens were taken no lower than 8500 feet in elevation and were trapped in places as high as 9350 feet. Some mice were found in "fences" of large Agave that bordered small wheat fields in the narrow montane valleys. Mice were most common at wood- land edges but also were caught in traps set on moss and among rocks along a shaded brook and adjacent to fallen logs under a canopy of pine, Douglas fir and aspen. Peromyscus melanotis lives 266 University of Kansas Publs., Mus. Nat. Hist. in association with Sorex milleri, Eutamias bulleri, Peromyscus diffi- cilis, Neotoma mexicana, and Microtiis mexicanus. Average and extreme weights of melanotis from southeastern Coa- huila were: for 7 adult males, 20.4 (17.2-24.7); for 5 adult, non- pregnant females, 24.0 (20.4-30.5), Pregnant females contained an average of 3.8 embryos; females with embryos were taken on Jan- uary 16 (1 female with 3 embryos), April 10 (1 with 4). July 26 (1 with 5), July 27 (2 with 5 each, 2 with 1 each), July 29 (2 with 3 each), July 31 (1 witli 5), August 1 (2 with 5 each), August 2 (1 with 5). Coahuilan specimens resemble closely P. melanotis from the vi- cinity of Las Vigas, Veracruz. Winter-taken animals are paler than those taken at the same places in warmer months. Generally, Coa- huilan mice are sHghtly paler than melanotis taken at the same season in Veracruz, Estado de Mexico, Distrito Federal, and Durango; however, selected skins from Coahuila compare favorably with all but the dark- est individuals from the other states. Perom,yscus melanotis, in northeastern Mexico, occurs at high altitudes in Coahuila, Ta- maulipas (see Osgood, 1909:112) and probably Nuevo Leon. It has not been recorded from San Luis Potosi and owing to absence there of suitable boreal habi- tat in the Sierra Madre Oriental is isolated from populations of melanotis in central Mexico. The two species, P. melanotis and P. maniculatus, were taken to- gether at Mesa de Tablas. However, usually maniculatus is found in the foothills of the Sierra Madre Oriental no higher than tlie be- ginnings of the semi-arid piiiyon-juniper association whereas mel- anotis was not taken below the humid pine-fir association. Measurements. — Average and extreme measurements of 7 adult males and 14 adult females, taken 12 miles east of San Antonio de las Alazanas and 13 Fig. 48. Distribution of the black- cared woods mouse, Peromyscus me- lanotis, in Coahuila. Mammals of Coahuila, Mexico 267 miles east and 3 miles south of San Antonio de las Alazanas, respectively, are: total length, 155 (147-160), 163 (153-170); lengtli of tail vertebrae, 63 (60- 66), 65 (60-70); length of hind foot, 20 (19-22), 20 (19-22); height of ear from notch, 18 (17-19), 18 (16-19.5); basilar length of skull, 20.1 (19.3-20.7), 20.4 (19.5-20.9); zygomatic breadth, 12.9 (12.2-13.6), 13.1 (12.5-13.6); length of nasals, 10.9 (10.5-11.6), 11.0 (10.4-11.5); least interorbital constriction, 4.1 (3.9-4.2), 4.0 (3.6-4.2); mastoidal breadth, 11.0 (10.6-11.3), 11.0 (10.6-11.4); length of palatine shts, 5.8 (5.5-6.2), 5.9 (5.5-6.2); alveolar length of maxillary tooth-row, 3.9 (3.7-4.1), 4.0 (3.8-4.5). Specimens examined. — Total, 62, from: Mesa de Tablas, 8600 ft., 2; 2 mi. E Mesa de Tablas, 8500 ft., 1; ii mi. S Mesa de Tablas, 8600 ft., 1; 12 mi. E San Antonio de las Alazanas, 9000 ft., 38; 13 mi. E San Antonio de las Alazanas, 9S50 ft., 5; 3 mi. S and 13 mi. E San Antonio de las Alazanas, 8900 ft., 15. Peromyscus nasutus nasutus (J. A. Allen) Rock Mouse Vesperimus nasutus J. A. Allen, Bull. Amer. Mus. Nat. Hist., 3:299, June 30, 1891, type from Estes Park, Larimer County, Colorado. Peromyscus nasutus Osgood, N. Amer. Faxma 28:176, April 17, 1909. Distribution. — Montane areas (oak-pinyon association) in central and north- em Coahuila (see fig. 49). Remarks. — The rock mouse occurs as isolated populations in the oak-pinon belt of desert ranges in central and northern Coahuila. Specimens were obtained in the Sierra del Pino, Sierra de la Madera, and the Sierra de San Marcos. On most mountains nasutus is the only Peromyscus present, although in the Sierra de la Madera P. pectoralis occurs with nasutus in some places, especially near the lower edge of the oak-pinon association. Rock mice were trapped near rocky exposures in narrow mountain caiions, in chaparral where the ground was blanketed with a thick layer of oak leaves, and in front of openings in bases of oak trees. A principal food is acorns; caches of these nuts were found beneath rocks in the Sierra de San Marcos in April. It was not discovered whether this mouse either climbed or nested in trees. Average and extreme weights of four adult males and four adult, non-pregnant females, all from 5 miles west and 22 miles south of Ocampo, respectively, were: 28.0 (25.0-31.0) and 32.7 (30.0-35.0). A pregnant female containing three embryos was taken on July 4; one with four embryos was captured on December 15. Three lac- tating females were obtained on December 14. Coahuilan spcimens diflPer from P. n. nasutus from the Capitan Mountains, New Mexico, in being more dusky and more heavily washed with blackish and averaging larger in both external measure- ments and in dimensions of the skull. Size in this species seems to increase in a southerly direction. Actually, specimens from the 268 University of Kansas Publs., Mus. Nat. Hist. Sierra de San Marcos and from 5 miles west and 22 miles south of Ocampo (Sierra de la Madera) approach, in size, P. difficilis of southeastern Coahuila. How- ever, difficilis is larger and has larger auditory bullae and dis- tinctly longer palatine slits. Most Coahuilan nasutus have dark tarsi. Measurements. — Average and ex- treme measurements of four adult males from the Sierra de San Marcos and five adult females from 5 miles west and 22 miles south of Ocampo, respectively, are: total length, 229 (220-233), 236 (226-241); length of tail vertebrae, 129 (121-133), 129 (122-134); length of hind foot, 24 (22-26), 24 (23-24); height of ear from notch, 24 (21-26), 24 (22-25); greatest length of skull, 29.2 (28.8- 29.5), 29.5 (29.4-29.7); basilar length, 22.3 (22.1-22.6), 22.5 (22.2-22.8); least interorbital constriction, 4.5 (4.4- 4.6), 4.5 (4.4-4.6); length of nasals, 11.2 (11.0-11.4), 11.4 (11.0-11.7); length of shelf of bony palate, 4.3 (4.1-4.5), 4.4 (4.1-4.6); length of palatine slits, 6.5 (6.3-6.7), 6.3 (6.1-6.5); length of diastema, 7.7 (7.5-7.8), 7.7 (7.5-8.0); post palatal length, 10.5 (10.3-10.9), 10.7 (10.2-11.3); alveolar length of maxillary tooth-row, 4.7 (4.6-4.8), 4.5 (4.3-4.7). Specimens examined. — Total, 46, from: Sierra del Pino, 7925 ft., 5 mi. N and 9 mi. W Acebuches, 2; Sierra del Pino, 5 mi. W and 3 mi. S Acebuches, 11; 5 mi. W and 22 mi. S Ocampo, 6000 ft., 13; 4 mi. N and 21 mi. W Cuatro Cienegas, 6200 ft., 7; Sierra de San Marcos, 7250 ft., 2 mi. N and 18 mi. W Santa Teresa, 13. Peromyscus pectoralis White-ankled Mouse The white-ankled mouse lives on the brush-covered foothills of the Upper Sonoran Life-zone in chaparral and in piiion-oak associa- tions in the Sierra del Carmen, Sierra del Pino, Sierra Madre Orien- tal and other mountainous areas. The species is usually less abun- dant on exposed slopes where sotol, lechuguilla, yucca and other xeric plants are dominant. Rarely was pectoralis taken either in the more mesic pine-fir association of the higher elevations or in desert situations within the Lower Sonoran Life-zone. At lower elevations this mouse often occupied rocky ledges facing arroyos Fig. 49. Distribution of the rock mouse, Peromyscus nasutus nasutus, in Coahuila. Mammals of Coahuila, Mexico 269 ? T- where vegetation was sparse. Six specimens were taken in such a habitat along the Rio Grande. The observations on occurrence in Coahuila corroborate the findings of Dalquest (1953:145) in San Luis Potosi tliat pectoralis occupies places where most other kinds of Peromyscus are absent, Zonally, this species occurs usually immediately above the areas of occurrence of eremicus, leucopus, and manicnlatus but usually below that of boijlii, difficilis, melanotis, nasutus and truei. Peromyscus pectoralis commonly is associated with eremicus, especially near bases of rocky slopes. In lines of traps set up the sides of such slopes, usually pectoralis was taken in larger numbers in the traps set at the higher elevations while eremicus outnumbered pectoralis in catches at the lower elevations, where the habitat was more closely related to that on the desert plain. Average and extreme weights of pectoralis from Coahuila were: for 35 adult males (show- ing wear on their teeth), 21.1 (18.0-25.0); for 19 adult, non- pregnant females, 22.3 (15.0- 29.5). Twenty pregnant fe- males, containing an average of 2.9 embryos (some females with as few as 2 embryos, others with as many as 5), were obtained in the following months: March (3 females), April (7 females). May (2 females), June (4 females), September ( 2 females ) , October ( 1 female ) , and December ( 1 fe- male). Females in lactation were found in January, March, April, June, September, and December. None of 18 adult females taken near Torreon on January 11 and 12, 1951, was pregnant. Peromyscus pectoralis eremicoides Osgood Peromyscus attwateri eremicoides Osgood, Proc. Biol. Soc. Washington, 17: 60, March 21, 1904, type from Mapimi, Durango. Peromyscus pectoralis eremicoides Lyon and Osgood, U. S. Nat. Mus., Bull. 62:128, January 28, 1909. Distribution. — Rocky exposures at lower elevations in extreme southwestern Coahuila (see fig. 50). Fig. 50. Distribution in Coahuila of the white-ankled mouse. 1. Peromys- cus pectoralis eremicoides. 2. Pero- myscus pectoralis laceianus. 270 University of Kansas Publs., Mus. Nat. Hist. Remarks. — This subspecies is here restricted in its distribution in Coahuila to suitable habitat in the broad basin (Bolson de Mapimi) in the extreme southwestern part of the State. Specimens from the vicinity of Torreon agree closely with near topotypes of eremicoides from Durango and are smaller and grayer than laceianus. Examples of pectoralis from the Sierra de Guadalupe have been referred to laceianus because of larger size both in external and cranial dimen- sions. Evidently there is a rather narrow zone of intergradation between eremicoides and laceianus. Measurements. — Average and extreme measurements of 12 adult males (showing wear on their teeth) and 18 adult females, all from 3 miles southeast of Torreon, respectively, are: total length, 186 (175-203), 188 (177-204); length of tail vertebrae, 106 (97-113), 107 (101-116); length of hind foot, 20 (19-21), 20 (19-21); height of ear from notch, 18 (16-19), 18 (17-19); greatest length of skull, 24.5 (23.5-25.4), 24.7 (23.9-25.8); basilar length, 18.5 (17.5-19.1), 18.6 (17.8-19.4); zygomatic breadth, 11.9 (11.5-12.6), 12.2 (11.6- 12.5), length of nasals, 9.2 (8.6-9.5), 9.3 (8.9-9.8); mastoidal breadth, 10.4 (10.2-11.0), 10.5 (10.2-10.7); length of palatine slits, 5.0 (4.7-5.4), 5.0 (4.7- 5.4); alveolar length of maxillary tooth-row, 3.8 (3.6-4.0), 3.7 (3.6-4.0). Specimens examined. — Total, 33, from 3 mi. SE Torreon, 3800 ft. Additional record (Osgood, 1909:164): Jimulco. Peromyscus pectoralis laceianus Bailey Peromyscus pectoralis laceianus Bailey, Proc. Biol. Soc. Washington, 19:57, May 1, 1906, type from Lacey Ranch, near Kerrville, Kerr County, Texas. Distribution. — Lower elevations on mountainous areas of Coahuila (see fig. 50). Remarks. — Specimens of pectoralis from all parts of Coahuila except the southwestern part are best referred to laceianus. Os- good (1909:165) previously reported pectoralis from several locali- ties in northern Coahuila as laceianus. When compared with laceianus from the Sierra del Carmen, specimens taken from some places along the eastern border — especially in the foothills of the Sierra Madre Oriental — are generally larger, more brownish on the upper parts, and have longer rostra and narrower skulls. These features are well shown in mice from the mountains east of Her- manas and from Panuco, and although these specimens are assign- able to laceianus they show some relationship to P. p. collinus Hooper, which occurs on the eastern slopes of the Sierra Madre Oriental in Nuevo Leon. Mice from southeastern Coahuila (vicini- ties of Saltillo and Bella Union and from Sierra Guadalupe) are assigned to laceianus on the basis of size. On geographic grounds, specimens classified as P. p. eremicoides by Osgood (1909:164) from Mammals of Coahuila, Mexico 271 Jaral, Saltillo, Carneros, and Sierra Encarnacion are here referred to laceianus. Measurements. — Average and extreme measurements of 5 adult males and 3 adult females from the Sierra de la Encantada, 38 miles south and 23 miles east of Boquillas, respectively, are: total length, 195 (178-207), 200 (192- 208); length of tail vertebrae, 106 (98-117), 109 (102-114); length of hind foot, 20 (19-22), 21 (21-21); height of ear from notch, 20 (18-21), 20 (19- 21); greatest length of skull, 26.7 (25.9-27.3), 27.2 (26.5-27.6); basilar length, 20.1 (19.7-20.5), 20.5 (19.8-21.0); zygomatic breadth, 13.2 (12.7-13.8), 13.4 (13.1-13.7); length of nasals, 10.2 (9.6-10.7), 10.2 (9.9-10.5); mastoidal breadth, 11.1 (10.9-11.5), 11.2 (11.0-11.5); length of palatine slits, 5.4 (5.2- 5.5), 5.6 (5.6-5.7); alveolar length of maxillary tooth-row, 4.1 (4.0-4.2), 4.1 (4.1-4.2). Specimens examined. — Total, 259, from: Rio Grande, 17 mi. S Dryden, Terrell County, Texas, in Coahuila, 7; Canon del Cochino, 3200 ft., 16 mi. N and 21 mi. E Piedra Blanca, 1; Tinaja de Chavez, 6400 ft.. El Jardin Ranch, Sierra del Carmen, 1 (TCWC); Campo Centrale, 5300 ft., El Jardin Ranch, Sierra del Carmen, 1 ( TCWC ) ; Club Sierra del Carmen, 5000 ft., 2 mi. N and 6 mi. W Piedra Blanca, 26; Club Shack, 5000 ft.. Sierra del Carmen, 1 (DMNH); Burro Canon, 5700 ft.. Sierra del Carmen, 1 (DMNH); Oso Canon, 6300 ft., Sierra del Carmen, 5 (DMNH); Juarez Canon, 4800 ft.. Sierra del Carmen, 3 (DMNH); Fortin, 3300 ft., Rancho Las Margaritas, 7; El Rio Alamos, in Canon Mulato, 3300 ft., Rancho Las Margaritas, 3; 1 mi. W Hacienda Las Margaritas, 3000 ft., 2; ii mi. W Hacienda Las Margaritas, 2900 ft., 7; Sierra de la Encantada, 4100 ft., 37 mi. S and 21 mi. E Boquillas, 2; Sierra de la Encantada, 4400 ft., 38 mi. S and 23 mi. E Boquillas, 11; 3 mi. S and 8 mi. E Hechiceros, 4650 ft.. Chihuahua, in Coahuila, 2; 6 mi. SW San Geronimo, 9; Sierra del Pino, 5250 ft., 6 mi. N and 6 mi. W Acebuches, 7; Sierra del Pino, 6200 ft., 5 mi. W and 3 mi. S Acebuches, 2; 4 mi. W Hacienda La Mariposa, 2300 ft., 3; 4 mi. W and 2 mi. S Hacienda La Mariposa, 8; 16 mi. E and 18 mi. N Ocampo, 11; 14 mi. E and 16 mi. N Ocampo, 1; 10 mi. N Hermanas, 2; 6 mi. E Hermanas, 6; 9 mi. E Hermanas, 2000 ft., 12; 20 mi. S and 4 mi. W Ocampo, 5300 ft., 1; 22 mi. S and 5 mi. W Ocampo, 5950 ft., 4; 4 mi. N and 21 mi. W Cuatro Cienegas, 6200 ft., 3; 9 mi. W and 4 mi. S San Buenaventura, 1800 ft., 21; Panuco, 3000 ft., 4; 2 mi. N and 11 mi. W Santa Teresa, 5000 ft., 3; 17 mi. N and 8 mi. W Saltillo, 5200 ft., 18; 1 mi. S and 4 mi. W Bella Union, 7000 ft., 16; 3 mi. S and 3 mi. E Bella Union, 6750 ft., 16; N slope Sierra Guadalupe, 7500 ft., 10 mi. S and 5 mi. W General Cepeda, 8; IV slope Sierra Guadalupe, 8000 ft., 11 mi. S and 7 mi. W General Cepeda, 9; N slope Sierra Guadalupe, 8000 ft., 11 mi. S and 5 mi. W General Cepeda, 15; N slope Sierra Guadalupe, 6700 ft., 11 mi. S and 4 mi. W General Cepeda, 1. Additional records (Osgood, 1909:164-165): opposite Langtry, Texas; 15 mi. SW Langtry, Texas; head Las Vacas Creek; 15 mi. E Las Vacas; Jaral; Saltillo; Carneros; Sierra Encarnacion. Peromyscus truei gentilis Osgood Pifion Mouse Peromyscus gratus gentilis Osgood, Proc. Biol. Soc. Washington, 17:61, March 21, 1904, type from Lagos, Jalisco. Peromyscus truei gentilis Osgood, N. Amer. Fauna 28:175, April 17, 1909. Distribution. — Foothills of the Sierra Madre Oriental in southeastern Coa- huila (see fig. 51). Remarks. — This species occupies rocky situations where juniper. 272 University of Kansas Publs., Mus. Nat. Hist. pinon and yucca provide the principal vegetative cover. In March, the piiion mouse was taken in company with Reithrodontomijs megalotis and Peromyscus maniculatiis in traps set along a dry wash bordered by juniper and cactus at 7500 feet elevation in the val- ley east of Diamante Pass. In the vicinity of Carneros, at 6000 feet elevation, truei and Pero- myscus eremicus were taken to- gether in a yucca-mesquite asso- ciation. Near San Antonio de las Alazanas, at 8700 feet elevation, truei was trapped in company with Peromyscus difficilis at the junction between the piiion-juni- per association and yellow pine. Average and extreme weights of truei from southeastern Coa- huilawere: for four adult males, 22.9 (21-24.7); for five adult, non-pregnant females, 27.8 (21- 32.5). Pregnant females were taken March 19 (one with two embryos), June 26 (two, each with two embryos), July 10 (one with 2 embryos), July 15 (one with three embryos), and Septem- ber 4 ( one with five embryos ) . Hoffmeister (1951:47-50) considers Coahuilan truei as belonging to the subspecies gentilis, that is distributed in suitable localities in the northern part of the Mexican Plateau ( Mesa del Norte ) . Like Peromyscus melanopJirys, another species of the Plateau, truei reaches southeastern Coahuila by way of northeastern Zacatecas and northern San Luis Potosi. Measurements. — Average and extreme measurements of eight adult males (KU 33198, 33199, 49158, 49367, 49368, 49372, 58568, 58569) and 8 adult females (KU 33196, 33200, 33202, 36255, 36256, 49371, 49159, 58565), from southeastern Coahuila, respectively, are: total length, 200 (188-222), 198 (183-211); length of tail vertebrae, 108 (97-124), 104 (89-116); length hind foot, 23 (22-24), 22 (20-24); height of ear from notch, 23 (21-24), 22 (20- 24);' greatest length of skull, 26.6 (26.1-27.0), 27.1 (26.3-27.7); basilar length, 20.2 (20.0-20.4), 20.7 (19.9-21.2); greatest breadth of skull, 12.9 (12.7-13.2), 12.8 (12.5-13.1); least interorbital constriction, 4.4 (4.2-4.5), 4.4 (4.2-4.5); length of nasals, 9.9 (9.8-10.2), 10.2 (9.4-10.6); length of shelf of bony palate, Fig. 51. Distribution of the pinon mouse, Peromyscus truei gentilis, in Coahuila. Mammals of Coahuila, Mexico 273 4.0 (3.7-4.4), 4.1 (3.6-4.4); length of palatine slits, 5.6 (5.3-6.0), 5.7 (5.4- 6.0); alveolar length of maxillary tooth-row, 4.2 (4.1-4.4), 4.2 (4.1-4.4). Specimens examined. — Total, 38, from: 3 mi. S and 3 mi. E Bella Union, 6750 ft., 2; 7 mi. S and 4 mi. E Bella Union, 7200 ft., 6; 5 mi. E and 2 mi. S Saltillo, 9000 ft., 1; 6 7ni. E and 4 mi. S Saltillo, 7500 ft., 11; 2 mi. E and 2 mi. N San Antonio de las Alazanas, 8700 ft., 4; 14 mi. W and 1 mi. N San Antonio de las Alazanas, 6500 ft., 6; 12 mi. S and 2 ini. E Arteaga, 7500 ft., 7; 1 mi. S Cameros, 6000 ft., 1. Additional records ( HoflFmeister, 1951:50): Sierra Guadalupe, Sierra En- camacion. Baiomys taylori taylori (Thomas) Taylor Pigmy Mouse Hesperomys (Vesperiinus) tatjlori Thomas, Ann. and Mag. Nat. Hist., ser. 5, 19:66, January 1887, type from San Diego, Duval County, Texas. Baiomys taylori Meams, U. S. Nat. Mus., Bull. 56:381, April 13, 1907. Distribution. — Grassy situations in northeastern Coahuila along tributaries of the Rio Grande and Rio Salado. Remarks. — In Coahuila the Taylor pigmy mouse was found at one locality only. This was in the mountains southwest of the Hacienda San Geronimo where there was tall grass under mesquite and oak trees in the foothills. Livestock did not graze the area heavily, be- cause it was remote from surface water. Small mammals were ex- ceedingly abundant at this place in 1950; our party of four zoologists prepared 110 study specimens of mammals, all taken in traps set on the night of April 9-10. Baiomys, Reithrodontomys fulvescens, Peromyscus leucopus and Sigmodon hispidus were abundant; many were caught in traps set in runways in deep grass. The Taylor pigmy mouse evidently reached this locality by mov- ing upstream from the Rio Sabinas via the Rio de los Alamos and the Arroyo de la Babia. Downstream, grazing by livestock along this watershed is severe. Probably Baiomys is confined only to places in northeastern Coahuila where grassy habitat has been pre- served. Average and extreme weights of five adult males (see below) were 7.9 (7.2-8.5); of four adult, non-pregnant females, 8.9 (7.9- 9.8). Coahuilan specimens resemble closely examples of taylori from Tamaulipas and Texas. I am indebted to Olin L. Webb for critical comparisons of Baiomys from Coahuila with the several subspecies of B. taylori. Measurements. — Average and extreme measurements of five adult males and five adult females (see below), respectively, are: total length, 107 (99-114), 106 (99-115); length of tail vertebrae, 40 (37-45), 43 (39-46); length of hind foot, 13 (12-14), 14 (13-14); height of ear from notch, 11 (10-12), 11 (10- 12); greatest length of skull, 18.3 (18.2-18.4), 18.4 (17.9-18.9); zygomatic 274 University of Kansas Publs., Mus. Nat. Hist. breadth, 9.7 (9.5-9.9), 9.8 (9.5-10.2); breadth of cranium, 8.7 (8.5-8.7), 8.6 (8.4-8.8); depth of cranium, 5.7 (5.4-6.0), 5.9 (5.7-6.0); least interorbital con- striction, 3.6 (3.5-3.8), 3.6 (3.5-3.7); length of nasals, 7.0 (6.8-7.4), 7.0 (6.8- 7.2); length of incisive foramina, 4.1 (3.9-4.2), 4.2 (4.1-4.3); alveolar length of maxillary tooth-row, 3.1 (3.1-3.2), 3.2 (3.0-3.3). Specimens examined. — Thirty-one from 6 mi. SW San Geronimo. Onychomys leucogaster longipes Merriam Northern Grasshopper Mouse Onychomys longipes Merriam, N. Amer. Fauna 2:1, October 20, 1889, type from Concho County, Texas. Onychomys leucogaster longipes Hollister, Proc. Biol. Soc. Washington, 26: 216, December 20, 1913. Distribution. — Northeastern Coahuila (see fig. 52). Remarks. — The northern grasshopper mouse occurs only on the Gulf Coastal Plain of northeastern Coahuila. Mice are most abun- dant where prickly pear, mes- quite and grass grow in places not heavily grazed. One preg- nant female was taken in March and one in April; both contained 5 embryos. Average and ex- treme weights of leucogaster from northeastern Coahuila were: for ten adult males, 39.5 grams (33.0-48.2); for eight, non-pregnant, adult females, 36.3 (32.6-40.7). Specimens assigned to O. I. longipes from Coahuila closely resemble longipes from La Salle and Cameron counties, Texas. This subspecies reaches tlie west- ernmost limit of its range in Coa- huila. Fic. 52. Distribution in Coahuila of the grasshopper mice. 1. Onychomys lcncoa,(istcr longipes. 2. Onychomys torridus surrufus. 3. Onychomys tor- ridus torridus. Measurements. — Two adult males and two adult females from 1 mile south and 9 miles west of Villa Acuiia, respectively, measure: total length, 177, 176 and 168, 170; length of tail vertebrae, 69, 62 and 58, 60; length of hind foot, 25, 24 and 24, 23; height of ear from notch, 20, 18 and 19, 18; greatest length of skull, 29.4, 29.2 and 28.5, 28.5; basilar length, 23.2, 23.2 and 22.7, 22.5; zygomatic breadth, 14.7, 14.8 and 14.4, 14.9; breadth of braincase, 12.1, 12.2 and 12.3, 12.3; least interorbital constriction, 4.9, 4.6 and 4.5, 4.7; length of nasals, 12.3, 12.3 and 11.6, 11.7; Mammals of Coahuila, Mexico 275 length of palatine slits, 6.3, 5.8 and 5.9, 5.6; alveolar length of maxillary tooth- row, 4.6, 4.5 and 4.7, 4.5. Specimens examined. — Total, 33, from: 1 mi. S and 9 mi. W Villa Acufia, 6; 15 mi. N and 8 mi. W Piedras Negras, 2; 6 mi. N and 2 mi. E La Babia, 6; 2 mi. S and 11 mi. E Nava, 810 ft., 1; 29 mi. N and 6 mi. E Sabinas, 4; La Gacha, 1600 ft., 4; 10 mi. N Hermanas, 1100 ft., 3; 9 mi. N Hermanas, 1500 ft., 6; 1 mi. S Hermanas, 1300 ft., 1. Onychomys torridus Southern Grasshopper Mouse This mouse occurs in western Coahuila and in the intermontane valleys of extreme southeastern Coahuila. Populations from these two areas are separated by mountainous barriers and other unsuit- able habitat. The southern grasshopper mouse seemingly reached western Coahuila from Chihuahua to the westward. In contrast to the more abundant population in southeastern Coahuila that lives in higher and less dry surroundings, the species in western Coahuila occurs sparingly on arid plains, where only two specimens were trapped. Another record was obtained from owl pellets (Baker, 1953b: 253). Borell and Bryant (1942:29) report this species as rare in the Big Bend area of Texas. Four adult females taken in June were pregnant, each containing three embryos; one female taken in July contained four embryos. Onychomys torridus surrufus HoUister Onychomys torridus surrufus Hollister, Proc. U. S. Nat. Mus., 47:472, Octo- ber 29, 1914, type from Miquihuana, Tamaulipas. Distribution. — Southeastern Coahuila (see fig. 52). Remarks. — Coahuilan specimens resemble surrufus from 4 miles north of Jaumave, Tamaulipas. Measurements. — Average and extreme measurements of 3 adult males and 4 adult females from 7 miles south and 4 miles east of Bella Union, respectively, are: total length, 150 (146-153), 159 (156-161); length of tail vertebrae, 56 (55-59), 57 (56-58); length of hind foot, 22 (21-22), 22 (21-23); height of ear from notch, 21 (20-22), 19 (18-19); greatest length of skull, 26.9 (26.6- 27.1), 26.9 (26.5-27.3); basilar length, 20.9 (20.7-21.1), 21.0 (20.7-21.4); zygomatic breadth, 12.9 (12.6-13.1), 13.2 (12.6-13.5); breadth of cranium, 11.9 (11.7-12.1), 11.9 (11.7-12.3); least interorbital constriction, 4.7 (4.5-4.8), 4.6 (4.6-4.7); length of nasals, 10.6 (10.5-10.8), 10.9 (10.2-11.8); length of palatine slits, 5.1 (4.8-5.3), 5.1 (4.9-5.5); alveolar length of maxillary tooth- row, 3.8 (3.7-4.1), 4.0 (3.8-4.1). Specimens examined. — Total, 12, from: 7 mi. S and 4 mi. E Bella Union, 7200 ft., 8; 14 mi. W and 1 mi. N San Antonio de las Alazanas, 6500 ft., 2; 8 mi. N La Ventura, 6000 ft., 2. Additional records (Hollister, 1914:473): La Ventura. 276 University of Kansas Publs., Mus, Nat. Hist. Onychomys torridus torridus (Coues) Hesperomys (Onychomys) torridus Coues, Proc. Acad. Nat. Sci. Philadel- phia, 26:183, December 15, 1874, type from Camp Grant, Graham County, Arizona. Onychomys torridus Merriam, N. Amer. Fauna 2:3, October 30, 1889. Distribution. — Western Coahuila (see fig. 52). Remarks. — Two specimens available compare favorably with tor- ridus from Pima and Santa Cruz counties, Arizona, and from Chi- huahua. One skull from an owl pellet, identified as O. torridus by Baker (1953b:253), is assigned, on geographic grounds, to O. t. tor- ridus. Measurements. — One adult female from 6 miles northwest of Tanque Alvarez measures: total length, 152; length of tail vertebrae, 54; length of hind foot, 21; height of ear from notch, 22; greatest length of skull, 26.9; basilar length, 20.9; zygomatic breadth, 13.0; breadth of braincase, 11.6; least interorbital con- striction, 4.6; length of nasals, 10.7; length of palatine shts, 5.2; alveolar length of maxillary tooth-row, 3.8. Specimens examined. — Total, 2, from: 6 mi. NW Tanque Alvarez, 3400 ft., 1; 18 mi. S and 14 mi. E Tanque Alvarez, 4000 ft., 1. Additional record (Baker, 1953b:253): 3 mi. NW Cuatro Cienegas. Sigmodon hlspidus berlandieri Baird Hispid Cotton Rat Sigmodon berlandieri Baird, Proc. Acad. Nat. Sci. Philadelphia, vol. 7 ( 1854- 55), p. 333, 1855, type from Rio Nazas, Coahuila. Sigmodon hispidus berlandieri Bailey, Proc. Biol. Soc. Washington, 15:106, June 2, 1902. Distribution. — Grassy situations at lower elevations tliroughout the State (see fig. 53). Remarks. — The hispid cotton rat occurs in grassy situations at lower elevations, especially in moist places along streams where grass has been protected from heavy grazing by livestock. Ani- mals live also in small numbers in drier situations where isolated clumps of grass are shielded by large growths of prickly pear from grazing by livestock. Runways through grass and short cuttings of grass blades are characteristic sign of Sigmodon. Animals have been taken as low as 800 feet in elevation and as high as 7500 feet. Pregnant females, taken in April and August, contained an aver- age of 5 embryos. The smallest number of embryos found in any one female was 3, and the largest was 9. Average and extreme weights of nine adult males and non-pregnant females, all from Coahuila, were 119.4 (87.0-186.0). Specimens from Coahuila re- garded as adults from a study of cranial features weighed at least 87 grams, although less mature females weighing as little as 57 grams were pregnant. Mammals of Coahuila, Mexico 277 Grassy habitat in the vicinity of the Rio Nazas, from where the type specimen of berlandieri was taken, is now sparse and much of this area is irrigated crop-land. Only one immature topotype was trapped. Specimens obtained from other parts of Coahuila were assigned to berlandieri after comparison with this single specimen and with the discus- sion of berlandieri in Bailey (1902:106-107). Cotton rats from along the Rio Grande (vicinities of Villa Acuna and Pie- dras Negras) have slightly nar- rower skulls than those of speci- mens from the vicinity of Saltillo. Two mandibles from cave de- posits from the vicinity of Cuatro Cienegas, identified by Gilmore (1947:157) as Sigmodon hispi- dus and 25 skulls taken from owl pellets from a cave three miles northwest of Cuatro Cienegas, identified by Baker (1953b: 253) as Sigmodon hispidus, are here referred, on geographic grounds, to S. h. berlandieri. Measurements. — Average and extreme measurements of 4 adult males and measurements of 2 adult females from 15 miles north and 8 miles west of Piedras Negras, respectively, are: total length, 280 (273-283) and 268, 269; length of tail vertebrae, 123 (117-126) and 116, 114; length of hind foot, 32 (31-34) and 33, 34; height of ear from notch, 20 (19-23) and 19, 22; greatest length of skull, 34.7 (34.4-35.1) and 35.1, 34.9; basilar length, 28.6 (28.4- 29.1) and 28.8, 28.5; zygomatic breadth, 20.0 (19.8-20.2) and 19.3, 19.3; least interorbital constriction, 5.2 (5.0-5.4) and 5.3, 5.8; length of nasals, 13.2 (12.8-13.8) and 13.6, 13.8; breadth of braincase, 14.3 (14.1-14.4) and 14.2, 13.9; mastoidal breadth, 14.1 (13.7-14.5) and 14.2, 13.9; alveolar length of maxillary tootli-row, 6.5 (6.4-6.7) and 6.8, 6.5. The greatest length of the skull of an old adult male (34627 KU) from the vicinity of Saltillo measures 38.1 mm. Specimens examined. — Total, 102, as follows: 1 mi. S and 9 mi. W Villa Acuiia, 12; 15 mi. N and 8 mi. W Piedras Negras, 16; 35 mi. S and 14 mi. E Boquillas, 2350 ft., 2; 6 mi. SW San Geronimo, 13; 29 mi. N and 6 mi. E Sabinas, 6; 2 mi. S and 3 mi. E San Juan de Sabinas, 1160 ft., 2; Sabinas, 3 (BSC); 10 mi. ESE Sabinas, 1; Don Martin, 800 ft., 11; 8 mi. N Hermanas, 1500 ft., 1; 5 mi. W Nadadores, 7; 6 mi. E Cuatro Cienegas, 2200 ft., 2; 2 mi. Fig. 53. Distribution of the hispid cotton rat, Sigmodon hispidus berlan- dieri, in Coahuila. 278 University of Kansas Publs., Mus. Nat. Hist. N and 1 mi. E Monclova, 1; 1 mi. SW San Pedro de las Colonias, 3700 ft., 1 1 mi. N San Lorenzo, 4200 ft., 1; 1 mi. N Saltillo, 5000 ft., 6; ^ mi. N SaltiUo 5000 ft., 9; 2 mi. W Saltillo, 5000 ft., 1; 6 mi. E and 4 mi. S Saltillo, 7500 ft., 1 N foot Sierra Guadalupe, 6200 ft., 9 mi. S and 5 mi. W General Cepeda, 3 N foot Sierra Guadalupe, 6200 ft., 10 mi. S and 5 mi. W General Cepeda, 1 Jimulco, 2 (BSC). Additional records (Baker, 1953b:253): 3 mi. NW Cuatro Cienegas; (Gil- more, 1947:157): 25 mi. W and 5 mi. S Cuatro Cienegas; (Bailey, 1902:106): Rio Nazas. Sigmodon ochrognathus ochrognathus Bailey Yellow-nosed Cotton Rat Sigmodon ochrognathus Bailey, Proc. Biol. Soc. Washington, 15:115, June 2, 1902, type from Chisos Mountains, Brewster County, Texas. Distribution. — Grassy uplands on desert mountains in central and northwest- em Coahuila (see fig. 54). Remarks. — The yellow-nosed cotton rat occurs in the Grassland and Grassland Transition Vegetative-types in central and northern Coahuila. On northern foothills of the Sierra de la Madera, this animal was trapped in well-de- fined runs in bunch grass, prickly pear and scrub oak at 5300 feet elevation. In the Sierra de la Encantada ochrognathus was in runs in thick grass on the floor of the Canon del Hillcoat at ap- proximately 7000 feet elevation. These runs led to small burrows mostly under small oaks. Runs, thought to be those of Sigmodon, were found at elevations as high as 7925 feet in the Sierra del Pino. In the Sierra del Carmen, Taylor et al. (1945:26) found cuttings, burrows, and piles of earth in grass at Campo Central and at Tinajas de Chavez. At no place were Sigmodon hispidtis and Sigmodon ochrog- nathus found together although the former species occurs in the oak association in the vicinity of San Ger6nimo at 3300 feet eleva- tion. Sigmodon ochrognathus is easily distinguished from S. his- pidus by smaller size and by the presence, instead of absence, of a w^; V,-, > MO tej- /I \ -^ Fig. 54. Distribution of the vcllow- nosed cotton rat, Sigmodon ochrogna- thus ochrognathus, in Coahuila. Mammals of Coahuila, Mexico 279 yellowish-buflF wash on the nose, face and rump. The yellowish encircles the eyes. Two females, obtained in April, contained two embryos, respec- tively; four females, obtained in October, contained six, seven, eight and nine embryos, respectively. Goldman and Gardner (1947:57) regard specimens from the Sierra del Carmen in Coahuila as typical S. o. ochrognaihus. Ani- mals from the Sierra de la Madera (specimens from 20 miles south and 4 miles west of Ocampo) also seem best referred to this sub- species although specimens available ( young adults taken in April ) are paler and less washed with yellowish-buff than specimens of equal age from the Carmens taken in October. These variations in color may be the result of usual differences in summer and winter pelages. Measurements. — An adult male from El Jardin Ranch and an adult female from Juarez Canon, respectively, measure: total length, — , 242.5; length of tail vertebrae, — , 95; length of hind foot, 30, 28; greatest length of skull, 33.2, 30.1; basilar length, 26.2, 24.6; zygomatic breadth, 18.8, 17.2; least interorbital constriction, 4.8, 4.7; length of nasals, 12.9, 10.1; mastoidal breadth, 13.8, 12.9; alveolar length of maxillary tooth-row, 6.5, 6.0. Specimens examined. — Total, 20, from: Tinaja de Telles, 4800 ft., El Jardin Ranch, Sierra del Carmen, 1 (TCWC); Juarez Canon, 4800 ft., Sierra del Carmen, 16 (DMNH); 20 mi. S and 4 mi. W Ocampo, 5300 ft., 3. Neotoma albigula White-throated Wood Rat Neotoma albigula occurs in all parts of Coahuila except on the Gulf Coastal Plain in the northeast and at the highest elevations in the mountains. This wood rat prefers rocky outcrops, and most specimens were trapped in such places, where accumulations of their debris and droppings were conspicuous. Houses constructed of twigs and other materials were found also within the protective growth of prickly pear and mesquite clumps, but such houses rarely were found more than a few feet from rocky exposures. In a mon- tane valley near Saltillo, wood rats occupied prickly pear planted as a fence around a small corral. Seven wood rats were taken from runways which connected dens in this growth. The cactus was a chief food for these animals. In the Serranias del Burro Neotoma albigula was trapped on rocky cliffs in oaks and yellow pine at the lower edge of the Transition Life-zone. In the Sierra del Carmen, Taylor et al. (1945:26) found this species living in thick growths of cactus and mesquite as low as 1800 feet in elevation and as high as 9000 feet. 280 University of Kansas Publs., Mus. Nat. Hist, This species may breed at any time of the year. Young animals, less than 260 mm. in total length, were taken in January, February, March, April, June, September and December. Females in lacta- tion were trapped in April, June, July and December. Females containing embryos were trapped in March, April, May, and July. Each of 14 pregnant females contained either 1, 2 or 3 em- bryos (an average of 1.5). Aver- age and extreme weights of albi- gula from Coahuila were: for 8 adult, non-pregnant females, 190.6 (162-204); for fifteen adult males, 217.7 (162-265). These wood rats are highly prized as food by local residents. Neotoma albigula albigula Hartley Neotoma albigula Hartley, Proc. California Acad. Sci., ser. 2, 4- 157, May 9, 1894, type from vicinity of Fort Lowell, near Tucson, Pima County, Arizona. Distribution. — Western and central part of the State (see fig. 55). Remarks.— Goldman (1910:32 and 37) concluded correctly that N. a. albigula intergrades with N. a. leucodon in southern Coahuila. The subspecies albigula is paler and smaller than leucodon. The pale color prevails in most Coahuilan specimens save those in the mountains of the southeast, which are darker. In size and cranial characters distinctions are not so clear-cut, and the zone of inter- gradation between these two subspecies is a broad one. In cranial characters specimens from as far northwest as the Sierra Mojada are somewhat intermediate between these two subspecies. Critical study indicates that leucodon occurs only in the mountains of ex- treme eastern and southeastern Coahuila. Bone fragments, recorded as Neotoma albigula by Gilmore (1947: 157) from cave deposits in the vicinity of Cuatro Cienegas, are con- sidered here as belonging to the subspecies albigula. Five skulls from owl pellets taken from a cave three miles northwest of Cuatro Cienegas and incorrectly identified as Neotoma micropus by Baker Fig. 55. Distribution in Coahuila of the white-throated wood rat. 1. Neo- toma albigula albigula. 2. Neotoma albigula leucodon. Mammals of Coahuila, Mexico 281 (1953b: 253) have been re-examined and identified as Neotoma a. albigula. Measurements. — Average and extreme measurements of 7 adult males and 4 adult females from the west foot of Pico de Jimulco, respectively, are: total length, 341 (320-370), 338 (330-348); length of tail vertebrae, 152 (134-170), 155 (150-158); length of hind foot, 35 (33-37), 35 (34-35); height of ear from notch, 31 (30-33), 32 (30-34); greatest length of skull, 44.4 (43.0-46.1), 43.8 (42.8-44.7); condylobasal length, 43.2 (41.7-45.4), 42.2 (41.2-43.2); basilar length, 37.7 (36.0-39.6), 36.7 (35.9-38.2); zygomatic breadth, 23.1 (21.4-24.5), 23.2 (21.8-23.9); length of nasals, 16.0 (15.3-16.8), 15.3 (14.9- 15.8); least interorbital constriction, 6.1 (5.6-6.3), 6.1 (5.9-6.3); length of palatal bridge, 8.2 (7.9-8.4), 7.5 (6.8-8.3); length of palatine slits, 9.6 (9.3- 10.1), 9.8 (9.4-10.2); alveolar length of maxillary tooth-row, 8.8 (8.5-9.2), 8.3 (7.9-8.8). Specimens examined. — Total, 64, from: 7 mi. S and 2 mi. E Boquillas, 1800 ft., 1; 10 mi. S and 5 mi. E Boquillas, 1500 ft., 1; Campo Centrale, 5300 ft.. Sierra del Carmen, 2 (TCWC), Fortin, 3300 ft., Rancho Las Margaritas, 2; 3 mi. N Las Margaritas, 3500 ft., 1; 35 mi. S and 14 mi. E Boquillas, 1; 6 mi. N and 2 mi. E La Babia, 1; Sierra de la Encantada, 4400 ft., 38 mi. S and 23 mi. E Boquillas, 1; 9 mi. N and 6 mi. W Castillon, 4100 ft., 1; 14 mi. E and 16 mi. N Ocampo, 1; 2)2 mi. SE Ocampo, 3000 ft., 1; 3 mi. NE Sierra Mojada, 4100 ft., 2; Sierra Mojada, 4150 ft., 5; 3 mi. S and 9 mi. E Cuatro Cienegas, 2250 ft., 1; 4 mi. N Acatita, 3600 ft., 5; 10 mi. S Santa Teresa, 4000 ft., 1; San Ldzaro, 4800 ft., 64 mi. N and 22 mi. W Saltillo, 3; 2 mi. S San Lazaro, 4700 ft., 60 mi. N and 22 mi. W Saltillo, 1; 3 mi. S and 3 mi. E Muralla, 3; La Pastora Rancho, 41 mi. W and 15 mi. N Saltillo, 6; 1 mi. N San Lorenzo, 4200 ft., 3; 3 mi. SE Torreon, 3800 ft., 2; W foot Pico de Jimulco, 4600 ft., 19. Additional records (Gilmore, 1947:157): vicinity of Cuatro Cienegas; (Baker, 1953b:253): 3 mi. NW Cuatro Cienegas; (Goldman, 1910:33): Mon- clova, Jaral, Jimulco. Neotoma albigula leucodon Merriam Neotoma leucodon Merriam, Proc. Biol. Soc. Washington, 9:120, July 2, 1894, type from San Luis Potosi, San Luis Potosi. Neotoma albigula leucodon Goldm.an, N. Amer. Fauna 31:36, October 19, 1910. Distribution. — Foothills of the Sierra Madre Oriental of eastern and south- eastern Coahuila (see fig. 55). Remarks. — This subspecies occurs at least as high as the be- ginnings of the growth of pine in the mountains of southeastern Coahuila but seemingly does not live in association with Neotoma mexicana, which is confined to the higher and more moist pine-fir forests. Measurements. — Average and extreme measurements of 7 adult males and measurements of 2 adult females from the mountains of southeastern Coahuila (vicinities of Saltillo, Bella Union, and Arteaga), respectively, are: total length, 347 (328-362) and 330, 350; length of tail vertebrae, 152 (130-167) and 143, 167; length of hind foot, 34 (32-36) and 35, 34; height of ear from notch, 31 (28-34) and 31, 30; greatest length of skull, 44.7 (41.3-47.5) and 42.6, 43.5; condylobasal length, 43.3 (40.0-45.9) and 41.2, 42.0; basilar length, 37.0 (34.1- 39.6) and 35.8, 37.0; zygomatic breadth, 23.9 (22.0-24.8) and 23.4, 22.9; 282 University of Kansas Publs., Mus. Nat. Hist. length of nasals, 15.8 (15.1-16.8) and 14.9, 14.9; least interorbital constriction, 6.1 (5.9-6.5) and 6.3, 5.8; length of palatal bridge, 7.7 (7.3-8.6) and 7.7, 7.6; length of palatine slits, 9.7 (9.0-10.8) and 9.1, 9.5; alveolar length of maxillary tooth-row, 8.3 (7.5-8.6) and 8.8, 8.2. Specimens examined. — Total, 58, from: 6 mi. E Hermanas, 1; 9 mi. E Her- manas, 2000 ft., 1; Panuco, 3000 ft., 4; 1 mi. S and 4 mi. W Bella Union, 7000 ft., 3; 3 mi. S and 3 mi. E Bella Union, 6750 ft., 1; 6 mi. E and 4 mi. S Saltillo, 7500 ft., 7; 7 mi. S and 4 mi. E Bella Union, 7200 ft., 4; 14 mi. W and 1 mi. N San Antonio de las Alazanas, 6500 ft., 2; 12 mi. S and 2 mi. E Arteaga, 7500 ft., 5; N foot Sierra Guadalupe, 6500 ft., 10 mi. S and 5 mi. W General Cepeda, 22; N foot Sierra Guadalupe, 6500 ft., 10 mi. S and 4 mi. W General Cepeda, 4; 7 mi. S and 1 mi. E Gomez Farias, 6500 ft., 3; 8 mi. N La Ventura, 5500 ft., 1. Additional records (Goldman, 1910:37): Saltillo; Cameros, 6800 ft.; Sierra Encamacion, 9000 ft.; La Ventura, 5600 ft. Neotoma goldmani Merriam Pigmy Wood Rat Neotoma goldmani Merriam, Proc. Biol. Soc. Washington, 16:48, March 19, 1903, type from Saltillo, Coahuila. Distribution. — Desert plains of southern and southwestern Coahuila (see fig. 56). Remarks. — Neotoma goldmani occupies the desert plains of south- em and southwestern Coahuila. Formerly thought to have a re- stricted distribution on the Mesa del Norte, goldmani is now known from southeastern Chi- huahua to as far south as San Luis Potosi (see Rainey and Baker, 1955:622). This diminu- tive wood rat was trapped in and adjacent to rocky places in the Lower Sonoran Life-zone. Houses or burrows were not ob- served but it was supposed that these were concealed in rocks. Dalquest (1953:163) also found this wood rat in rocky places in San Luis Potosi. J. R. Alcorn took seven specimens in three days at a locality near Torreon. Here goldmani was taken in as- sociation N. albigula. Two pregnant females taken on March 29 each contained one embryo. These females weighed 95.6 and 96.0 grams, respectively. Fig. 56. Distribution of the pigmy wood rat, Neotoma goldmani, in Coa- huila. Mammals of Coahuila, Mexico 283 Neotoma goldmani is most closely related to N. albigula (see Rainey and Baker, 1955). Small size and silky pelage distinguish goldmani from other wood rats in Coahuila. Measurements. — Average and extreme measurements of 4 adult females ( KU 36338, 36339, 40756, 40761 ) and measurements of one adult male (KU 40758), all from southern Coahuila, respectively, are: total length, 273 (265-285) and 275; length of tail vertebrae, 126 (113-136) and 128; length of hind foot, 29 (27-31) and 28; height of ear from notch, 27 (25-29) and 28; greatest length of skull, 37.4 (36.9-38.4) and 37.7; condylobasal length, 36.0 (35.4-37.0) and 35.8; basilar length of skull, 31.0 (30.5-31.9) and 30.4; zygomatic breadth, 18.7 (17.7-19.7) and 18.3; length of nasals, 14.3 (13.8-14.8) and 14.5; least interorbital constriction, 5.5 (5.3-5.7) and 5.7; length of palatal bridge, 6.1 (5.9-6.5) and 6.2; length of palatine slits, 8.9 (8.8-9.0) and 8.7; alveolar length of maxillary tooth-row, 8.2 (7.9-8.3) and 8.2. Specimens examined. — Total, 9, from: 17 mi. N and 8 mi. W SaltiUo, 5200 ft., 2; 3 mi. SE Torreon, 3800 ft., 7. Additional records (Goldman, 1910:82): Jaral, Saltillo. Neotoma mexicana Mexican Wood Rat The Mexican wood rat occurs in Fig. 57. Distribution in Coahuila of the Mexican wood rat. 1. Neotoma mexicana inornata. 2. Neotoma mexi- cana navus. mesic forests of the higher moun- tains of Coahuila. In the Sierra del Carmen specimens have been taken in mixed oaks and conifers from as low as 4800 feet in elevation to as high as 8000 feet. In the Sierra de la Madera two Mexican wood rats were trapped on rocks and deep ground litter consisting of oak leaves in a plant association in- cluding oaks, pine, madroiio, and juniper. In the Sierra Madre Oriental, southeast of Saltillo, mexicana was obtained at eleva- tions of from 8500 feet to 9350 feet in moist situations in a plant association including pine, aspen, and fir. Mexican wood rats were taken where rock ledges or out- crops were prominent, usually in narrow valleys and where a canopy of trees shaded the areas. 284 University of Kansas Publs., Mus. Nat. Hist, No houses were observed; it was assumed that underground dens or dens protected by rock piles were used. In the Sierra de la Madera mexicana seemed to feed principally on acorns; whether or not acorns were stored by the rats was not determined. The Mexi- can wood rat seemed to be ecologically isolated from other species of Neotoma, but probably competes for food, especially for acorns, with Eutamias hulleri, Sciurus alleni, and Peromyscus difficilis in the Sierra Madre Oriental, with Peromyscus nasutus and Eutamias dorsalis in the Sierra de la Madera and with Spermophilus variegatus and Eutamias dorsalis in the Sierra del Carmen. Pregnant females collected east of San Antonio de las Alazanas included two taken on April 11 containing one and two embryos, respectively, and one taken on August 2 containing one embryo. One juvenile was taken on August 2 east of San Antonio de las Ala- zanas; another was taken on October 25 in the Sierra del Carmen. Average and extreme weights of mexicana from Coahuila were: for five adult males, 180.2 (164.0-195.5); for one adult, non-pregnant female, 144.0 grams. Neotoma mexicana inomata Goldman Neotoma mexicana inomata Goldman, Proc. Biol. Soc. Washington, 51:60, March 18, 1938, type from Sierra del Carmen, Coahuila. Distribution. — Oak-yellow pine association of higher mountains of northern and central Coahuila; definitely known to occur in the Sierra del Carmen and the Sierra de la Madera but probably also in suitable habitat in adjacent moun- tains (see fig. 57). Remarks. — Wood rats from the Sierra de la Madera are assigned to inomata although specimens available, an adult male and a younger female (taken on April 5), have more slender rostra and are paler than a subadult male (taken on October 20) from the Sierra del Carmen. Measurements. — An adult male from the Sierra de la Madera measures: total length, 349, length of tail vertebrae, 167; length of hind foot, 34; height of ear from notch, 32; greatest length of skull, 44.2; condylobasal length, 42.6; basilar length, 36.8; zygomatic breadth, 22.9; length of nasals, 18.6; least inter- orbital constriction, 5.4; length of palatal bridge, 7.7; length of palatine slits, 9.5; alveolar length of maxillary tooth-row, 8.3. Specimens examined. — Total, 4, from: Juarez Canon, 4800 ft.. Sierra del Carmen, 1 (DMNH); Campo Madera, 8000 ft.. Sierra del Carmen, 1 (DMNH); Sierra de la Madera, 22 mi. S and 5 mi. W Ocampo, 6200 it., 2. Additional record (Goldman, 1938a: 60): Sierra del Carmen. Neotoma mexicana navus Merriam Neotoma navus Merriam, Proc. Biol. Soc. Washington, 16:47, March 19, 1903, type from Sierra Guadalupe, Coahuila. Distribution. — Mountains of southeastern Coahuila (see fig. 57). Mammals of Coahuila, Mexico 285 Remarks. — Merriam (1903:47) described navus as a full species relating it to mexicana but distinguishing navus by its longer tail and frontals expanded posteriorly. In the eleven specimens of navus available for this study, the tail seems no longer than that of N. m. inornata. The expansion of the frontals posteriorly to form a shelf is conspicuous in some crania and less so in others. This feature serves to distinguish navus from inornata, but is present, to a lesser degree, in at least one other subspecies, N. m. scopulorum Finley. Neotoma navus is here arranged as a subspecies of N. mexi- cana, because characteristics distinguishing navus from subspecies of mexicana are of the magnitude of those which distinguish the several subspecies of mexicana. Measurements. — Average and extreme measurements of four adult males from Diamante Pass and 12-13 miles east of San Antonio de las Alazanas and measurements of one adult female from a place one-half mile south of the Mesa de Tablas, respectively, are: total length, 339 (333-343) and 329; length of tail vertebrae, 156 (154-160) and 155; length of hind foot, 35 (34-36) and 34; height of ear from notch, 30 (26-32) and 30; greatest length of skull, 44.6 (43.8-45.1) and 43.2; condylobasal length, 42.6 (42.1-43.5) and 41.0; basilar length, 36.6 (35.9-37.2) and 35.2; zygomatic breadth, 22.5 (22.2-22.8) and 22.3; length of nasals, 17.5 (16.7-18.2) and 17.0; least interorbital constriction, 5.5 (5.3-5.9) and 5.5; length of palatal bridge, 8.6 (8.2-9.1) and 8.4; length of palatine slits, 9.7 (9.4-10.1) and 9.5; alveolar length of maxillary tooth-row, 8.7 (8.4-9.1) and 8.8. Specimens examined. — Total, 11, from: Diamante Pass, 8500 ft., 3 mi. E and 4 mi. S Saltillo, 1; }i mi. S Mesa de Tablas, 8600 ft., 1; 12 mi. E San An- tonio de las Alazanas, 9000 ft., 6; 13 mi. E San Antonio de las Alazanas, 9350 ft., 2,. Additional records (Goldman 1910:61): Sierra Guadalupe. Neotoma micropus micropus Baird Gray Wood Rat Neotoma micropus Baird, Proc. Acad. Nat. Sci. Philadelphia, vol. 7 (1854- 55), p. 333, April 1855, type from Charco Escondido, Tamaulipas. Distribution. — Lower elevations of northeastern and southeastern Coahuila (see fig. 58). Remarks. — Neotoma micropus occurs on the Gulf Coastal Plain in northeastern Coahuila and along the Rio Salinas and its tribu- taries in southeastern Coahuila. The animal usually avoids rocky areas and thick cover along arroyos, often constructing large houses in clumps of prickly pear. In the vicinity of Nava wood rats oc- curred in live oak thickets. At La Rosa gray wood rats were trapped at houses in low-growing vegetation on desert flats; nests were not seen on adjacent, low rocky hills on which lechuguilla grew. Neo- 6—1204 286 University of Kansas Publs., Mus. Nat. Hist. toma alhigula, which usually prefers rocky situations, and Neotoma micropus, which usually prefers to live on deep soils, were not found living in close association by me. Goldman (1910:28, 29, 33, 37), however, records both species from Monclova, Jaral, and Saltillo. Neotoma micropus probably reached southeastern Coahuila from Nuevo Leon by way of the watershed of the Rio Salinas. The Sierra del Carmen-Sierra Madre Oriental Axis seems to bar the passage of this wood rat in east-central Coahuila. Local residents refer to this wood rat as "rata de campo" and consider its flesh as a delicacy. One male (KU 35055) from 8 miles north of Hermanas has an additional, shallow, re-entrant angle on the anterior face of each first upper molar. Another specimen (KU 35056) from the same locality has a slight indication of this feature on the right, first upper molar. Average and extreme weights of seven adult males were 260 (236-281), of one, non-pregnant female, 243 grams. A pregnant female taken in December contained two embryos. A female in lactation was trapped in March. Young individuals were taken in March and November. A lactating female and two juveniles of approximately equal age, all taken on the same date and from the same locality near Sabinas, have large areas of white fur. The hairs of the underparts are totally white, except for one small patch on the right leg of the adult female where the bases of the hairs are plumbous. The white hairs extend onto the sides of each indi- vidual in an irregular fashion. Goldman (1910:28, 29) had for study 12 Neotoma micropus from Coahuila. He assigned specimens from three localities to N. m. micropus and those from two other localities, situated more or less in between the otlier three, to N. m. canesccns Allen. These two subspecies are separated on the basis of color and size; micropus is darker and larger. Coahuilan specimens are paler than typical micropus from Tamaulipas but not so pale gray as some examples of canescens from New Mexico. Two animals from southeastern Coa- huila (vicinity of La Rosa) are pale and laterally have a dull whitish appearance. In size, relying principally on a series of seven adults from the vicinity of Villa Acuiia, Coahuilan specimens resemble most closely micropus especially in dimensions of the skull. All specimens from Coahuila, including those of Goldman {loc. cit.), are here assigned to the subspecies, micropus. Five skulls from owl-pellets, taken from a cave three miles north- west of Cuatro Cienegas and identified by Baker (1953b: 253) as Mammals of Coahuila, Mexico 287 N. micropus, are here referred, on geographic grounds, to N. in. micropus. Measurements. — Average and ex- treme measurements of six adult males and measurements of one adult fe- male from 1 mile south and 9 miles west Villa Acufia, respectively, are: total length, 343 (330-369) and 331; length of tail vertebrae, 144 (134- 162) and 134; length of hind foot, 36 (36-37) and 36; height of ear from notch, 29 (27-31) and 27; greatest length of skull, 47.6 (46.6-48.2) and 46.4; condylobasal length, 45.3 (44.2- 46.4) and 43.8; basilar length, 39.0 (37.5-40.0) and 37.5; zygomatic breadth, 24.5 (23.7-25.5) and 24.6; length of nasals, 18.7 (17.4-19.4) and 17.8; least interorbital constriction, 6.2 (5.8-6.5) and 6.0; length of palatal bridge, 7.7 (7.4-8.2) and 7.8; length of palatine slits, 10.8 (10.1-11.4) and 10.3; alveolar length of maxillary tooth-row, 8.7 (8.3-9.0) and 8.9. Specimens examined. — Total, 49, from: 1 mi. S and 9 mi. W Villa Acuiia, 8; Caiion del Cochino, 3200 ft., 16 mi. N and 21 mi. E Piedra Blanca, 1; 11 mi. S Hacienda San Miguel, 2200 ft., 1; 15 mi. N and 8 mi. W Piedras Negras, 5; 2 mi. S and 11 mi. E Nava, 810 ft., 4; 29 mi. N and 6 mi. E Sabinas, 10; 10 mi. E Hacienda La Mariposa, 3000 ft., 1; La Gacha [= La Concha], 1600 ft., 1; 2 mi. S and 3 mi. E San Juan de Sabinas, 1160 ft., 1; 10 mi. ESE Sabinas, 2; 9 mi. NW Don Martin, 2; Don Martin, 800 ft., 2; base of Don Martin Dam, 2; 8 mi. N Her- manas, 1500 ft., 2; Hermanas, 1205 ft., 2; 1 mi. S Hermanas, 1200 ft., 1; 5 mi. N and 2 mi. W Monclova, 1; 3 mi. N and 5 mi. W La Rosa, 3600 ft., 3. Additional records (Baker, 1953b: 253): 3 mi. NW Cuatro Cienegas; (Gold- man, 1910:28, 29): 7 mi, E Las Vacas; Sabinas; Monclova; Jaral; Saltillo. Microtus mexicanus subsimus Goldman Mexican Vole Microtus mexicanus subsimus Goldman, Jour. Mamm., 19:494, November 14, 1938, type from Sierra Guadalupe, Coahuila. Distribution. — Subalpine Humid Forest Vegetation-type of the Sierra de Guadalupe and Sierra Madre Oriental of southeastern Coahuila (see fig. 59). Remarks. — The Mexican vole occurs only in the Subalpine Humid Forest Vegetation-type, on moist meadows in the highest parts of the mountains of southeastern Coahuila. None was found at ele- vations lower than 8500 feet. In the narrow valleys meadow mice often live along the edges of small cultivated fields. The animals FxG. 58. Distribution of the gray wood rat, Neotoma micropus micro- pus, in Coahuila. 288 University of Kansas Publs., Mus. Nat. Hist. do not occur in the coniferous forests or in chaparral which often covered the exposed slopes and places where fires and heavy log- ging had removed the stand of timber. Neither heavy grazing of the grass nor the presence of human habitation seems to restrict this rodent. In places where the grass was cropped until it was less than one inch high, the runways of this meadow mouse were most conspicuous, and the animals could easily be seen moving along their runways. W. Kim Clark ob- tained 11 of these mice with the help of his cat and dog; the cat caught 8 and the dog 3 indi- viduals. I found runs within 25 feet of houses; the mice seemed to occupy all open places where some turf existed. Average and extreme weights of mexicanus from Mesa de Tab- las and 12-13 miles east of San Antonio de las Alazanas were: for 12 adult males, 37.2 (28.0- 44.0); for 9 adult, non-pregnant females, 36.6 (30.0-41.6). Fe- males with embryos were taken on January 13 (1 female with 1 embryo), January 15 (1 with 2), July 26 (1 with 1), July 27 (2 females, each with 2 embryos, 2 with 3, 1 with 4), July 28 (3 with 3 each), July 29 (1 with 3), July 30 (1 with 3), August 1 (1 with 3), August 3 (1 with 3). Microtus mexicanus subsimus resembles more closely A/, m. mexi- canus (Saussure) of central Mexico than M. m. guadalupensis Bailey of western Texas. From M. m. mexicanus, subsimus is easily distinguished by its paler color, shorter rostrum and larger auditory bullae. Measurements. — Average and extreme measurements of four adult males and six adult females from 12 miles east of San Antonio de las Alazanas, re- spectively, are: total length, 145 (143-147), 148 (145-150); length of tail vertebrae, 31 (30-34), 30 (29-34); length of hind foot, 19 (18-20), 18 (18- 19); height of ear from notch, 14 (13-15), 14 (13-14); condylobasal length, 26.2 (25.5-26.7), 26.0 (25.4-26.4); length of nasals, 7.8 (7.5-8.1), 7.5 (7.3-7.8); palatilar length, 13.1 (12.6-13.3), 13.3 (12.7-13.6); zygomatic breadth, 15.3 Fig. 59. Distribution of the Mexican vole, Microtus mexicanus subsimus, in Coahuila. Mammals of Coahuila, Mexico 289 (14.8-15.8), 15.2 (14.9-15.7); least interorbital constriction, 3.4 (3.3-3.7), 3.3 (3.1-3.6); mastoidal breadth, 13.1 (12.6-13.5), 12.8 (12.3-13.4); alveolar length of maxillary tooth-row, 6.6 (6.4-6.8), 6.6 (6.3-6.9). Specimens examined. — Total, 58, from: Mesa de Tablas, 8600 ft., 1; 2 mi. E Mesa de Tablas, 8500 ft., 6; 12 mi. E San Antonio de las Alazanas, 9000 ft., 42; 13 mi. E San Antonio de las Alazanas, 9350 ft., 1; 3 mi. S and 13 mi. E San Antonio de las Alazanas, 8900 ft., 8. Additional records (Goldman, 1938b: 495): Sierra Guadalupe. Rattus norvegicus norvegicus (Berkenhout) Norway Rat Mus norvegicus Berkenhout, Outlines of the natural history of Great Britain . . ., vol. 1, p. 5, 1769, type from England. Rattus norvegicus Cabrera, Trab. Mus. Nac. Cien. Nat., Madrid, ser. zool.. No. 57, p. 264, December 30, 1932. Distribution. — In, and in the vicinity of, commuirities and rural dwellings of Coahuila. Remarks. — The Norway rat lives in most communities in Coa- huila. It probably occurs also in the larger haciendas to which foods and other supplies are transported by truck or wagon. No efiFort was made to catch Rattus in or near the vicinity of towns or rural dwellings. An adult taken at Saltillo on July 16, 1949, con- tained six embryos, each 5 mm. long. An adult female from one mile north of Saltillo measures: total length, 425; length of tail vertebrae, 200; length of hind foot, 38; height of ear from notch, 20; length of nasals, 18.2; zygomatic breadth, 23.4; alveolar length of maxillary tooth-row, 7.1. Specimens examined. — Total, 4, from: 1 mi. N Saltillo, 5000 ft., 1; Saltillo,3. Rattus rattus alexandrinus (£. Geoffroy-Saint-Hilaire) Black Rat Mus alexandrinus fi. GeoflFroy-Saint-Hilaire, Catalogue des mammiferes du Museum National d'Histoire Naturelle, Paris, p. 192, 1803, type from Alexandria, Egypt. R[attus]. rattus alexandrinus Hinton, Jour. Bombay Nat. Hist. Soc, vol. 26, no. 1, p. 63, December 20, 1918. Distribution. — Found in, and in the vicinity of, the larger communities in Coahuila, possibly most common in urban areas along the Rio Grande. Remarks. — In Coahuila the black rat usually lives in urban areas or in the near vicinity thereof, especially those along the Rio Grande. The only specimen saved was trapped near the Rio de San Diego more than three miles from the town of Jimenez and at least one mile from the nearest ranch house. The young, gray-bellied male (see below) has the following ex- ternal measurements: total length, 270; length of tail vertebrae, 144; length of hind foot, 31; height of ear from notch, 18. Specimen examined. — One, from 3 mi. W Jimenez, 850 ft., (skull only). 290 University of Kansas Publs., Mus. Nat. Hist. Mus musculus brevirostris Waterhouse House Mouse Mus brevirostris Waterhouse, Proc. Zool. See. London, 1837, pt. 5, p. 19, November 21, 1837, type from Maldonado, Uruguay. Mus musculus brevirostris Schwartz and Schwartz, Jour. Mamm., 24:64, Feb- ruary 20. 1943. Distribution. — Found throughout Coahuila closely associated with the dwell- ings of man. Remarks. — The house mouse is found in and around most tONvns and permanent farm or ranch dwellings in Coahuila. Specimens were taken near dwellings as low as approximately 800 feet in elevation near the Rio Grande, and as high as 7500 feet in the moun- tains southeast of Saltillo. House mice live also several miles from dwellings of man, especially where some permanent water is pres- ent. The mice seem to be as well adapted to living in adobe dwell- ings as to living in those constructed of wood. The house mouse, Norway rat and black rat cause considerable damage to stored food stuffs each year. There is no concerted effort to control these species. Two adult males from no more than four miles from Saltillo measure: total length, 190, 177; length of tail vertebrae, 94, 95; length of hind foot, 19, 18 height of ear from notch, 15, 16; occipitonasal length (of skull), 22.8, 22.4 basilar length, 18.6, 17.9; palatilar length, 10.6, 10.0; length of nasals, 8.2, 8.4 zygomatic breadth, 11.7, 11.5; mastoidal breadth, 9.8, 9.5; alveolar length of maxillary tooth-row, 3.8, 3.7. Specimens examined. — Total, 29, from: 2 mi. W Jimenez, 850 ft., 2; 15 mi. N and 8 mi. W Piedras Negras, 1; 4 mi. N Sabinas, 1; Don Martin, 800 ft., 1; Ocampo, 1; 8 mi. N and 25 mi. W Cuatro Cienegas, 4000 ft., 1; 9 mi. W and 4 mi. S San Buenaventura, 1800 ft., 1; 1 mi. SW San Pedro de las Colonias, 3700 ft., 2; 1 mi. S and 1 mi. W Coyote, Rio Nazas, 3500 ft., 4; 2 mi. E Torreon. 3700 ft., 2; 4 mi. NNW Saltillo, 5200 ft., 1; 1 mi. N Saltillo, 5000 ft., 2; 'A mi. N Saltillo, 5; Diamante, 7500 ft., 1; 3 mi. S and 3 mi. E Bella Union, 6750 ft., 3; Jimulco, 1 (BSC). Erethizon dorsatum couesi Meams Porcupine Erethizon epixanthus couesi Meams, Proc. U. S. Nat. Mus., 19:723, July 30, 1897, type from Fort Whipple, Yavapai County, ArizoTia. Erethizon dorsatum couesi Hall, Mammals of Nevada, p. 649, July 1, 1946. Distribution. — Northern Coahuila. Reported as being rare. Remarks. — The porcupine is well known to the residents of north- ern Coahuila. No specimens were obtained but reports of an oc- casional porcupine in most of the mountainous areas were received. Reports which I judged to be authentic were obtained in the Sierra del Pino, Sierra de los Hecheros, Sierra del Carmen, and the Ser- Mammals of Coahuila, Mexico 291 ranias del Burro. The species is seemingly nowhere common. Marsh (1937:38) writes that Senor Victoriano Villareal saw three porcupines in the Sierra del Carmen in the seven years that he lived in those mountains. Harvey Pauley told me that a porcupine was killed at La Babia in 1933. In the Serranias del Burro, Roberto Spence informed me in 1953 that he had seen "one or two" porcu- pines in the past ten years. Gilmore (1947:158), records as Ere- thizon dorsatum epixanthum, 42 bone fragments from cave deposits containing archeological materials from the vicinity of Cuatro Ciene- gas. No late records from this area were obtained. The porcupine in Coahuila is assigned to couesi on geographic grounds following Davis and Robertson (1944:271), who use this name for animals from western Texas (Culberson County). Canis latrans Coyote The howl of the coyote can be heard in all parts of Coahuila. The animal has a preference for the desert and occurs in many areas that are remote from water. The coyote is at home in mountainous areas and has been taken as high as 8900 feet. Marsh (1937:31) recorded this animal at 7000 feet in the Sierra del Carmen. Coyotes are perhaps most numerous in the vicinity of Presa Don Martin and other bodies of water on the Gulf Coastal Plain of northeastern Coa- huila and in tlie southern part of the State in the area between Saltillo and Torreon. In most places the local residents tolerate the coyote even though the animal is condemned for taking some goats, sheep and poultry. The losses of sheep and goats are probably kept low because of the vigilance of the ever-present herder. Usually the herder, often a boy, escorts his small flock to forage in the day- time and returns with his flock before evening to the dwelling. Some ranchers in northeastern Coahuila have made considerable effort to reduce coyote populations. Poison baits and steel traps have been used and more recently cyanide guns. Ranch -workers bring ears of victims to the hacienda for credit and usually hang the bodies of the coyotes along the roads, presumably so that the owner will see them. In April, 1950, I saw more than a score of coyote carcasses hung on acacia bushes along the road from Villa Acuna to San Miguel. Most of our coyotes were obtained from the ranchers; some were road-kills; others were shot by J. R. Alcorn, who lured indi- viduals within gunshot range by using his successful calling tech- nique. A female taken on April 9 contained five embryos, sexed 292 University of Kansas Publs., Mus. Nat. Hist. as four males and one female, averaging 125 mm. long (crown to rump). This pregnant female from 6 miles north and 2 miles east of La Babia weighed 25 pounds (11,340 grams); two non-preg- nant females from 10 miles south of Hacienda La Mariposa and 11 miles west of Hacienda San Miguel, respectively, weighed, 20.5 pounds (9,299 grams) and 30.5 pounds ( 13,835 grams ) . An adult male from 6 miles north and 2 miles east of La Babia weighed 32 pounds (14,515 grams ) . Canis latrans mearnsi Merriam Canis mearnsi Merriam, Proc. Biol. Soc. Washington, 11:30, March 11, 1897, type from Quitoba- quito, Pima County, Arizona. Canis latrans mearnsi Nelson, Proc. Biol. Soc. Washington, 45:224 November 26, 1932. Distribution. — Southern Coahuila (see fig. 60). Fig. 60. Distribution in Coahuila of the coyote. 1. Canis latrans mearnsi. 2. Canis latrans texensis. Remarks. — Compared to tex- ensis, specimens of mearnsi from southern Coahuila are bright gray instead of dull gray in over-all appearance; upper parts and legs deeper cinnamon instead of paler cinnamon; comparisons were made of pelts taken in late autumn and early winter. These pelts are more richly colored than those of mearnsi from California and than is indicated as normal coloring for mearnsi by Jackson (in Young and Jackson, 1951:298-299) and, thereby, show some resemblance to the more reddish southern subspecies, C. I. cagottis H. Smith and C. I. impavidiis J. A. Allen. One skin from the vicinity of San Antonio de las Alazanas is especially richly colored. A specimen from the vicinity of Monclova, assigned to texensis, is intermediate in color between mearnsi and texensis. Measurements of these Coahuilan coyotes compare most favorably with tliose for mearnsi listed by Jackson ( loc. cit. ) . Measurements. — Two adult males from 8-9 miles southeast of San Pedro de las Colonias measure: total lengtli, 1116, 1211; length of tail vertebrae, 322, 366; length of hind foot, 182, 191; height of ear from notch, 109, 114; condy- lobasal length, 176.3, 181.8; palatal length, 90.1, 97.3; squamosal breadth, 54.2, Mammals of Coahuila, Mexico 293 58.6; zygomatic breadth, 96.6, 96.4; least interorbital constriction, 30.7, 31.5; length of maxillary tooth-row (crown-length), 85.3, 87.5; crown-length of upper camassial, 20.1, 20.2; crown-length of first upper molar, 13.0, 13.0; crown- breadth of first upper molar, 16.8, 15.8; crown-length of lower camassial, 21.6, 22.4. Specimens examined. — Total, 7, from: 3 mi. S and 1 mi. W Hacienda Guadalupe, 1 (skull only); 8 mi. SE San Pedro de las Colonias, 3700 ft., 1; 9 mi. SE San Pedro de las Colonias, S700 ft., 1; 12 mi. N anl 15 mi. E Parras, 5000 ft., 1 (skull only); 2 mi. SSE Torreon, 3700 ft., 1; 3 mi. SE Torreon, 3800 ft., 1; 3 mi. S and 12^2 mi. E San Antonio de las Alazanas, 8900 ft., 1 (skin only). Canis latrans texensis Bailey Canis nehracensis texensis Bailey, N. Amer. Fauna 25:175, October 24, 1905, type from forty-five miles southwest of Corpus Christi, at Santa Gertrudis, in Kleberg County, Texas. Canis latrans texensis Bailey, N. Amer. Fauna, 53:312, March 11, 1932. Distribution. — Northern Coahuila (see fig. 60). Remarks. — Coyotes of the desert plains and adjacent mountains of northern Coahuila resemble in color examples of C. I. texensis from Duval County, Texas. The cranial measurements of most of these specimens from northern Coahuila are less than the average dimensions, but more than the minimum dimensions, given for texensis by Jackson (in Young and Jackson, 1951:276-277). Bone fragments of the coyote, identified as Canis latrans by Gilmore (1947:156) from cave deposits from near Cuatro Cienegas, are as- signed, on geographic grounds, to texensis. Measurements. — An adult male and an adult female from 6 miles north and 2 miles east of La Babia, respectively, measure: total length, 1199, 1102; length of tail vertebrae, 372, 314; length of hind foot, 194, 167; height of ear from notch, 109, 102; condylobasal length, 186.7, 178.4; palatal length, 97,7, 91.6; squamosal breadth, 62.5, 59.4; zygomatic breadth, 98.1, 94.1; least inter- orbital constriction, 32.4; 30.4; length of maxillary tooth-row (crown-length), 88.5, 83.3; crown-length of upper camassial, 20.5, 18.6; crown-length of first upper molar, 13.5, 12.2; crown-breadth of first upper molar, 16.1, 15.6; crown- length of lower camassial, 22.9, 20.7. Specimens examined. — Total, 24, from: 9 mi. W and 1 mi. S Villa Acufia, 1 (skull only); 11 mi. W Hacienda San Miguel, 1; 10 mi. E Hacienda San Miguel, 1 (skull only); 4 mi. W Hacienda Las Margaritas, 2900 ft., 1; Pino Solo, 4100 ft., Rancho Las Margaritas, 1 (skull only); 2 mi. W Hacienda Las Margaritas, 2800 ft., 3; i mi. W Hacienda Las Margaritas, 3000 ft., 3 ( 2 skins only, 1 skull only); 6 mi. N and 2 mi. E La Babia, 2; 9 mi. W Castillon, 3650 ft. 2 (skulls only); 2 mi. S and 6 mi. E Nava, 810 ft, 1 (skull only); Hacienda La Mariposa, 1940 ft., 1 (skull only); 10 mi. E Hacienda La Mariposa, 3000 ft., 1 (skuU only); La Gacha, 1600 ft., 1; 10 mi. SE Sabinas, 1 (skull only); 11 mi. S and 2 mi. W Sabinas, 1 (skull only); Don Martin, 800 ft., 1 (cranium only); 5 mi. SW Hermanas, 1 (skull only); 8 mi. N and 1 mi. E Monclova, 3000 ft., 1. Additional records (Jackson, in Young and Jackson, 1951:278): near Ha- cienda de La Palma, 25 mi. NW Muzquiz; 110 mi. NW Muzquiz; Gilmore (1947:156): 16 mi. E and 3 mi. S Cuatro Cienegas, 22 mi. W and 4 mi. S Cuatro Cienegas, and 16 mi. W and 26 mi. S Cuatro Cienegas. 294 University of Kansas Publs., Mus. Nat. Hist. Canis lupus baileyi Nelson and Goldman Gray Wolf Canis lupus baileyi Nelson and Goldman, Jour. Mammalogy, 10:165, May 19, 1929, type from Colonia Garcia (about 60 miles southwest of Casas Grandes), Chihuahua. Distribution. — Western Coahuila. Remarks. — Although formerly found throughout the State the wolf now seems to be restricted to a few places in western Coa- huila. It is reported to live in the Sierra de las Cruces, east of Jaco, where the wolf is well known to the people, who call it "lobo" or "lobo griz." In other places along the border of Coahuila and Chihuahua, wolves reported there may be stragglers from the west. In the Sierra de los Hecheros wolves are rare although in June, 1953, Senor Juan Jose Zapata of the Hacienda Rincon told me that one had been seen earlier that year. To the eastsvard, records of wolves are fewer. East of the Sierra del Pino, Senor Armando Verduzco of the Hacienda San Ildefonso told me that one lobo was seen in 1951 on the road south in the direction of Ocampo. Mr. Earl C. Johns- ton of Cuatro Cienegas reported no wolves in the vicinity of Ocampo but in 1954 said that a few may be present west of the Sierra del Pino. Dr. Walter P. Taylor et al (1945:25 and 1946:21) write that wolves are uncommon in the Sierra del Carmen and tliat one was reported near Campo Central (at Agua Chile) in April, 1945. In 1936, Marsh (1937:32) reported that wolves were increasing in the Sierra del Carmen. He wrote that 200 cattle were killed in 1934 and 1935 at Santa Domingo by wolves. He obtained records of wolves also from the desert plains southwest of Muzquiz. There have been no recent reports of such severe depradation on livestock. Traps and cyanide guns are used as control measures at present on many ranches in northern Coahuila. Seiior Roberto Spence says that in the past 30 years a few wolves have been taken at Piedra Rlanca, which is on the southeastern side of the Sierra del Carmen. Spence told me in 1953 that only one record ( ti-acks only ) of a wolf has been reported in the past 15 years in the Serranias del Rurro; these tracks were reported on the Enfante Ranch. He also doubts if wolves have been numerous since the 1880's, when flocks of sheep being pastured and driven through northeastern Coahuila were preyed on by wolves. Gilmore (1947:156) tentatively identified three bone fragments from cave deposits from the vicinity of Cuatro Cienegas as Canis lupus. Goldman (in Young and Goldman, 1944:466-471, fig. 14) records Mammals of Coahuila, Mexico 295 Canis lupus baileyi from Trans-Pecos Texas and Chihuahua but places Coahuila, although no specimens were available from there for examination, within the range of C. I. inonstrabilis Goldman. Taylor et al. (1945:25) use tliis name for the wolf in the Sierra del Carmen. Since wolves now occur only in the western part of Coa- huila and may be replenishing their depleted populations there through the spread of wolves eastward from Chihuahua, it seems likely that wolves in western Coahuila may be more representa- tive of baileyi than inonstrabilis. Perhaps in earher times, wolves found in eastern Coahuila were referable to monstrabilis. Probably Goldman (op. cit. :468) was correct in saying that wolves have never been very numerous in northeastern Mexico, Vulpes macrotis zinseri Benson Kit Fox Vulpes macrotis zinseri Benson, Proc. Biol. Soc. Washington, 51:21, February 18, 1938, type from San Antonio de Jaral, southeastern Coahuila. Distribution. — Western and southern Coahuila (see fig. 61). Remarks. — The kit fox is characteristic of the Chihuahuan Desert Shrub Vegetation-type in west- ern and southwestern Coahuila. It is rare in northeastern Coa- huila, east of the Sierra del Carmen-Sierra Madre Oriental Axis, although Mr. David Mc- Kellar of the Hacienda La Mari- posa reports that the kit fox is present in small numbers on Rancho La Mariposa, which is northwest of Muzquiz. No other records of kit fox east of the central mountainous area of the State were obtained. Bailey (1905:179) gives the "mouth of the Pecos" as the eastern limit of the range of the kit fox in Texas. This small fox is well known to the ranchers at Jaco, in the vi- cinity of the Sierra de los Hech- eros and at Boquillas, all in northwestern Coahuila. None was caught in this part of Coahuila in steel traps, although several Fig. 61. Distribution of the kit fox, Vulpes macrotis zinseri, in Coahuila. 296 University of Kansas Publs., Mus. Nat. Hist. gray foxes ( Urocyon ) were taken. Gray foxes probably outnumber kit foxes in most situations, but the kit fox is more numerous in areas where surface water is absent or at a minimum. Two specimens from the vicinity of Torreon and two from adja- cent places in Durango (one from 8 miles south and 11 miles west of Lerdo and one from 2 miles south-southeast of Chocolate) com- pare favorably with the published description of zinseri and are referred to this subspecies. Gilmore (1947:156) records two bone fragments of Vulpes macrotis from deposits in two caves in the vicinity of Cuatro Cienegas. These are considered, on geographic grounds alone, to belong to zinseri. It is likely that kit foxes of central and northern Coahuila, may prove to be closer to V. m. neomexicana Merriam. Measurements. — The two kit foxes examined from Coahuila (the first, KU 40775, is an adult male; the second, KU 40774, is an adult female) and two from Durango (the first one, KU 40776, from 8 miles south and 11 miles west of Lerdo is an adult male; the second one, KU 40777, from 2 miles south-south- east of Chocolate is an adult female), respectively, measure: total length, 751, 751 and 787, 806; length of tail vertebrae, 296, 275 and 297, 315; length of hind foot, 112, 115 and 125, 115; height of ear from notch, 78, 86 and 91, 79; condylobasal length (of skull), 108.5, 108.3 and 112.7, 110.6; zygomatic breadth, 63.3, 61.0 and 63.5, 61.3; greatest width of cranium, 42.7, 42.1 and 43.2, 41.4; least width of rostrum (measured behind and lateral to the roots of the canines), 17.0, 16.6 and 17.2, 16.5; least interorbital breadth, 21.8, 21.3 and 23.3, 22.6; alveolar length of maxillary tooth-row, 50.3, 49.7 and 52.8, 51.0; length of upper fourtli premolar (crown lengdi of outer side), 9.5, 9.7 and 10.3, 9.8; width of upper fourth premolar (anterior end), 5.4, 4.9 and 5.0, 4.8. Specimens examined. — Total, 2, from: 10 mi. E Torreon, 3700 ft. Additional records (Gilmore, 1947:156): 16 mi. E and 3 mi. S Cuatro Cienegas; and 22 mi. W and 4 mi. S Cuatro Cienegas; (Benson, 1938:21): San Antonio de Jaral [= Jaral]. Urocyon cinereoargenteus scottii Meams Gray Fox Urocyon virginianus scottii Meams, Bull. Amer. Mus. Nat. Hist., 3:236, June 5, 1891, type from Pinal County (probably near Oracle), Arizona. Urocyon cinereo-argenteus scottii J. A, Allen, Bull. Amer. Jklus. Nat. Hist., 7:253, June 29, 1895. Distribution. — State-wide (see fig. 62). Remarks. — The gray fox is the "zorro" of residents and is the most common canid in the State. Found everywhere (specimens taken at elevations of 850 feet to 8900 feet ) , this animal is most abundant on the desert plains and in the foothills of the mountains; its sign is most frequently seen in the vicinity of water and along brushy arroyos where rocky outcroppings are prominent. Most specimens were trapped in sets, along trails or roads, baited with canned fish Mammals of Coahuila, Mexico 297 or other meats. None was taken in summer although traps were set in Hkely places on several occasions in that season. Two half- grown gray foxes were shot with the aid of headlights at night near Jimenez. These pups were found near a rocky ledge; we did not find their den. Weights of Urocyon from Coahuila were: for two adult males, 3856 and 3037 grams; for two adult, non- pregnant females, 3742 and 3402 grams. Gilmore (1947:156) re- cords bone fragments from cave deposits taken in the vicinity of Cviatro Cienegas. Coahuilan gray foxes are re- ferred to scottii after being com- pared with scottii from Pinal County, Arizona, and with U. c. coUmensis Goldman from Nay- arit. Specimens from southwest- ern Coahuila are slightly smaller, especially in cranial dimensions, than those from the vicinity of the Rio Grande. This smallness, noted particularly in the breadth of the cranium and rostrum, shows relationship to coUmensis and indicates that intergradation between these two subspecies occiurs over a large part of the Central Plateau. Dalquest ( 1953:177) refers gray foxes from San Luis Potosi to coUmensis but judging from the measurements that he lists, his specimens are no smaller than Coa- huilan specimens here assigned to scottii. Gray foxes from Coa- huila are variable in color. A darker one from the vicinity of Tor- reon approaches coUmensis but others fully as dark are known from the northeastern part of the State, Measurements. — From northeastern Coahuila, two adult males (KU 45038, 45042) and two adult females (KU 45040, 45041), respectively, measure: total length, 973, 980 and 992, 995; length of tail vertebrae, 414, 418 and 421, 441; length of hind foot, 141, 130 and 136, 156; height of ear from notch, 83, 81 and 78, 79; condylobasal length, 122.1, 117.1 and 122.4, 120.5; palatal length, 61.1, 58.1 and 60.6, 58.7; least squamosal constriction, 46.9, 41.7 and 43.9, 43,1; zygomatic breadth, 67.8, 62.3 and 66.1, 65.4; least interorbital constriction, 25.9, 23.8 and 24.4, 25.0; cranial breadth, 47.1, 43.0 and 45.6, 44.0; crown- length of maxillary tooth-row, 54.2, 49.8 and 54.2, 53.5; crown-length of upper Fig. 62. Distribution of the gray fox, Urocyon cinereoargenteus scottii, in Coahuila, 298 University of Kansas Publs., Mus. Nat. Hist. camassial (from inner side), 11.4, 10.6 and 12.1, 10.8; crown-length of first upper molar, 8.2, 7.6 and 8.5, 7.2; crown-breadth of first upper molar, 10.0, 9.3 and 11.0, 9.7; crown-length of lower carnassial, 12.2, 11.2 and 12.7, 11.8. Specimens examined. — Total, 18, from: S side Rio Grande, 17 mi. S Dr\'den, Terrell County, Texas, in Coahuila, 1; 2 mi. W Jimenez, 850 ft., 2; 1 mi. N Boquillas, 1700 ft., 1 (skull only); Boquillas, 1 (skull only); 10 mi. S and 5 mi. E Boquillas, 1; 11 mi. S Boquillas, 1 (skin only); Fortin, 3300 ft., Rancho Las Margaritas, 1; Rio Alamos, 3300 ft.. Canon Mulato, Rancho Las Margaritas, 1 (skull only); Hacienda Las Margaritas, 2900 ft., 1 (skull only); 4 mi. W Hacienda La Mariposa, 2300 ft., 1; La Gacha, 1600 ft., 2; 8 mi. N and 20 mi. W Santa Teresa, 1; 21 mi. S and 11 mi. E Australia, 4400 ft., 1 (skull only); 3 mi. SE Torreon, 3800 ft., 1; Mesa de Tablas, 1 (skin only); 3 mi. S and 13 mi. E San Antonio de las Alazanas, 8900 ft., 1 (skin only); W foot Pico de Jimulco, 5000 ft., 1. Ursus americanus eremicus Merriam Black Bear Ursus americanus eremicus Merriam, Proc. Biol. Soc. Washington, 17:154, October 6, 1905, type from Sierra Guadalupe, Coahuila. Distribution. — Mountainous parts of Coahuila (see fig. 63). Remarks. — Black bears live in mountainous areas in most parts of Coahuila but are fairly common only in more remote places, such as the Sierra de la Madera, Sierra del Carmen, Sierra del Pino and Sierra de San Marcos, where fewer people live and hunting pres- sures are not great. Hunting black bear is a popular sport for both Mexicans and hunters from the United States. The Carmen Moun- tain Hunting Club, located near Piedra Blanca in the Sierra del Carmen, formerly operated with a membership chiefly from the United States and was attractive because of bear and other game present. The Sierra del Pino, the Sierra de San Marcos and the Sierra de Guadalupe are locales for hunts by persons from Mexico City, Guadalajara, Saltillo, Monterrey, and other cities. The bear population is being slowly reduced because the animal is shot at every opportunity. Ranch riders are often equipped with rifles for this purpose. On June 21, 1938, two bears, a female and her cub, were killed in a mountain canon near La Babia by a rider and left where they fell. We retrieved the specimens being able to save only the skeletons. It was reported to me by Senor Juan Jose Zapata of the Hacienda Rincon in 1953, that bear are entirely gone in the Sierra de los Hecheros and that the last one had been killed three years before. These mountains contain abundant montane chapar- ral, attractive to black bear, but intensive hunting seems to have eliminated the population. The mountains in the vicinity of Saltillo have few bear present because of the great number of hunters op- erating in the area. Black bear are rarely trapped or hunted by using dogs, but are occasionally roped by ranchmen; the hide of one Mammals of Coahuila, Mexico 299 captured in this manner was obtained near Ocampo; Greer reported that at least one bear was roped on the Rancho Las Margaritas in the Serranias del Burro in 1954. Acorns are the principal food of the black bear in autumn, winter and spring; fecal deposits found in late winter and early spring con- tained this food. In late spring, summer and early autumn fruits, berries and animal foods are utilized. In this period of the year bears often move from protective montane mesic forest and mon- tane chaparral and follow down arroyos to the desert plain, being attracted to places when berries and fruits of desert plants ripen. In late summer, bears are attracted to the fruit of the prickly pear. The sotol (Dasylirion) is consumed in spring and perhaps at other times of year. I have seen these plants cropped almost to the ground; the bear seems to prefer the tender bases of the shoots. Taylor et ah (1946:20) describes the bear's habit of feeding on sotol as well as on the succulent leaves of the giant dagger (Yucca car- nerosana ) . The bear also feeds on the blooms of this yucca, climb- ing the plant to reach the edible parts. On December 6 in the Sierra Azul, east of Hermanas, Robert Dickerman saw a half grown bear in the top of a palm ( Bau- hinia hella). Probably the bear was feeding on the tender shoots of this arborescent palm. Ac- cording to Ranchman Roberto Spence of Sabinas, bears prey on goats, sheep and cattle at all times of the year, but damage is heaviest when natural foods are scarce. The presence of at least one herder with each flock of goats or sheep probably keeps predation by bears at a mini- mum. The bear of Coahuila is almost always black. Roberto Spence told me that brown ones are rare but that some blacks have con- spicuous brown markings on the face. Skins examined have pale browm on the muzzles. The cubs are seen beginning in May; Spence Fig. 63. Distribution of the black bear, Ursus americanus eremicus, in Coahuila. 300 Unr^rsity of Kansas Publs., Mus. Nat. Hist, doubts if the adult bears hibernate but thinks that they are less active in winter than at other times of the year. I am indebted to J. Knox Jones, Jr., for comparing specimens from northern Coahuila with the type of U. a. eremicus and with repre- sentative specimens of U. a. a7nbltjceps Baird from the Jemez Moun- tains and the Black Range of New Mexico. His opinion {in lift., 30 March 1955) is that black bears from northern Coahuila are intermediate in character between amblyceps and eremicus but closer to eremicus, especially in forehead not rising so abruptly, larger molars and longer upper molariform tooth-row. Remains of black bear, identified as Euarctos americanus by Gil- more (1947:155) from cave deposits near Cuatro Cienegas, are assigned here, on geographic grounds, to U. a. eremicus. Measurements. — Cranial measurements of the type (female), four adult fe- males (USNM 262695, KU 57818, 56518, 57088) and three adult males (KU 57819, 57089, 56519) from northern Coahuila, respectively, are: condylobasal length, 246.0 and 244.0, 256.0, 257.0, 264.0 and 293.0, 280.0, 293.0; basilar length, 223.0 and 222.0, 235.0, 239.0, 245.0 and 274.0, 261.0, 271.0; least inter- orbital constriction, 53.7 and 57.8, 61.7, 63.4, 59.1 and 71.6, 61.7, 70.8; mastoidal breadth, 108.3 and 109.8, 120.8, 118.6, 118.7 and 141.0, 121.9, 141.0; zygo- matic breadth, 148.5 and 151.2, 158.0, 157.0 162.0 and 190.0, 160.0, 184.0; length of nasals, 62.3 and 60.9, 73.1, 74.0, 74.7 and 80.6, 72.4, 84.2; palatal length, 121.5 and 120.9, 139.0, 134.0, 139.0 and 153.0, 142.0, 152.0; alveolar length of upper molariform tooth-row, 49.9 and 50.6, 58.0, 55.1, 44.5 and 53.6, 57.3, 57.2. Specimens examined. — Total, 17, from: Club Sierra del Carmen, 4950 ft., 2 mi. N and 6 mi. W Piedra Blanca, 2 (skulls only); 7 mi. NW Hacienda Las Margaritas, 4000 ft, 1 (skull only); 1 mi. N Pino Solo, 3500 ft., Rancho Las Margaritas, 1 (baculum only); 1 mi. E Pino Solo, 2800 ft., Rancho Las Mar- garitas, 1; 2 mi. E Pino Solo, 4000 ft., Rancho Las Margaritas, 1 (skull only); Canon Mulato, 3000 ft., Rancho Las Margaritas 2 (skulls only); 2 mi. NW Hacienda Las Margaritas, 3000 ft., 1 (skull only); 4 mi. W Hacienda Las Margaritas, 2200 ft., 1 ( skull only ) ; 2 mi. W Hacienda Las Margaritas, 2800 ft., 1; 6 mi. SW San Geronimo, 1 (skin only); 6 mi. NW Hacienda La Mariposa, 2400 ft., 1 (.skull only); 3'A mi. NE Hacienda La Mariposa, 2300 ft., 1; 5 mi. SW Hacienda La Mariposa, 2300 ft., 1 (skull only); 2 mi. N Ocampo, 1 (skin only); 18 km. W San Lazaro, 62 mi. N and 40 mi. W Saltillo, 1 (skull only). Additional records (Taylor et al. 1945:24): Campo Centrale, 5400 ft.. Sierra del Carmen; (Gilmore, 1947:155): 16 mi. E and 3 mi. S Cuatro Ciene- gas; (U. S. Natl. Mus.): Carmen Mountains; Santa Rosa Mountains, 160 mi. NW Muzquiz; 25 mi. NW Muzquiz, near Hacienda de la Palma; Sierra En- camacion; (Merriam, 1904:154): Sierra Guadalupe. Ursus sp. Grizzly Bear The grizzly bear is extinct in Coahuila today but lived in most mountainous parts of the State in former times. Residents have oc- casionally referred to an "oso grande" that once occurred in the Mammals of Coahuila, Mexico 301 Sierra del Carmen and in other mountains as distinct from the existing "oso." Gilmore (1947:155) records one "doubtful frag- ment of a fibula" of a grizzly bear of the planiceps group from cave deposits obtained in the vicinity of Cuatro Cienegas. He suspects that these remains are at least pre-Spanish in age. On geographic grounds alone, the Coahuilan grizzly probably is related more closely to Ursus horribilis texensis Merriam than to any other kind of grizzly. This subspecies, texensis, once Uved in western Texas to the north- ward. Bassariscus astutus flavus Rhoads Ring-tailed Cat Bassariscus astutus jlavus Rhoads, Proc. Acad. Nat. Sci. Philadelphia, 1893, p. 417, January 30, 1894, type from Texas, exact locality unknown. Distribution. — Found in suitable habitat throughout Coahuila (see fig. 64), Remarks. — The ring-tailed cat is widely distributed in Coahuila, and favors cliffs and other rocky situations. This species is more common in the lower elevations than in the pine-oak belts of the higher mountains although Greg- ory (1938:118) and Taylor et al (1945:24 and 1946:21) record it from the higher parts of the Sierra del Carmen, and Marsh (1937:28) found it as high as 8000 feet there. The ring-tailed cat is common along the Rio Grande; one was trapped in dense cane at the water's edge near Boquillas, and another was shot at night in woodlands near Jimenez. This animal is well- known to the natives. They sel- dom call it by the usual Mexican name, cacomixtle. Usually it is called "cbmadreja" or sometimes "onza." Comadreja is used to refer to the weasel (Mustela frenata) and onza to the tree squirrel (Sciiirus niger). The term "ringtail" is also used occa- sionally and seems to be recognized ever>^where in the State as re- ferring to this species. Fig. 64. Distribution of the ring- tailed cat, Bassariscus astutus flavus, in Coahuila. 302 University of Kansas Publs., Mus. Nat. Hist. The animal is trapped for its pelt. I saw one pelt for sale at San Lazaro in 1950. Marsh (op. cit. •.29) reports that the Kickapoo Indians sell a number of skins each year in Muzquiz. Coahuilan ring-tailed cats seem to be typical flavus although those from near Torreon in southeastern Coahuila resemble B. a. consitus Nelson and Goldman from Jalisco more than astutus in having a narrower braincase in relation to greatest length of skull and a broader rostrum. Bone fragments from cave deposits in the vicinity of Cuatro Cienegas, identified as B. astutus by Gilmore (1947:155), are as- signed here, on geographic grounds, to B. s. fkivus. Measurements. — One adult male from 1 mile north of Boquillas and two adult males from 3 miles southeast of Torreon, respectively, measure: total length, 780, and 736, 780; length of tail vertebrae, 411, 378, 394; length of hind foot, 73, 67, 71; length of ear from notch, 54, 50, 52; greatest length of skull, 82.8, 78.2, 80.2; basilar length (of Hensel), 73.1, 70.7, 73.7; zygomatic breadth, 47.4, 48.7, 51.3; least interorbital constriction, 15.8, 15.9, 17.8; alveolar length of maxillary tooth-rovv^, 31.5, 30.0, 31.5; crown-length of upper fourth premolar (outerside), 7.3, 7.4, 7.6; crown-width of upper fourth premolar, 4.4, 4.9, 4.8. Specimens examined. — Total, 12, from: S side Rio Grande, 17 mi. S Dryden, Terrell Co., Texas, in Coahuila, 1; 15 mi. SE Langtry, Val Verde Co., Texas, in Coahuila, 1 (BSC); 1 mi. N Boquillas, 700 ft., 1; 7 mi. S and 2 mi. E Boquillas, 800 ft., 1; 2 mi. W Jimenez, 850 ft., 1; near Hda. La Palma, 25 mi. NW Muzquiz, 1 (BSC); 3 mi. SE Torreon, 3800 ft., 2; Saltillo, 1 (BSC); Sierra Guadalupe, 1 (BSC); Sierra Encamacion, 2 (BSC). Additional records (Gilmore, 1947:155): 16 mi. E and 3 mi. S Cuatro Ciene- gas, 22 mi. W and 4 mi. S Cuatro Cienegas. Procyon lotor fuscipes Meams Raccoon Proctjon lotor fuscipes Meams, Proc. Biol. Soc. Washington, 27:63, March 20, 1914, type from Las Moras Creek, Fort Clark, Kinney County, Texas. Distribution. — Found in suitable habitat throughout Coahuila (see fig. 65). Remarks. — Raccoons are common along the major streams of Coahuila but occur also in other places where some permanent water is present. Along the Rio Grande, downstream from Cafion de Boquillas, raccoons are numerous. Tracks were seen along the Mexican side of the Rio Grande 17 miles south of Dryden, Texas; at nine miles west and one mile south of Villa Acuiia; at Jimenez, near the mouth of the Rio de San Diego; at Nava and Guerrero. Upstream from Caiion de Boquillas raccoons are rare; I saw no tracks and obtained only a few reports from residents at Boquillas in 1952. Across the river in Brewster County, Texas, Borell and Bryant (1942:13) also obtained few records of raccoons in 1936 Mammals of Coahuila, Mexico 303 and 1937. Marsh (1937:27-28) reported racoons as uncommon in the foothills of the Sierra del Carmen in 1936. Along the water- shed of the Rio Salado-Rio Sa- binas, woodlands and thickets of cane provide good habitat for the animals. Marsh (loc. cit.) saw a caged animal at Muzquiz in 1935. Raccoons are reported also along the larger arroyos, such as the Arroyo de la Babia, and in the irrigated areas in the vicinity of Saltillo and Torreon. Locally the animal is called either "tejon" or "mapache." Raccoons have little economic value in the State; some are trapped at Presa Don Martin, probably by persons primarily in quest of beaver. At Muzquiz, Marsh {loc. cit.) received re- ports from fur buyers that a few ^ ^^ T^ .1 ■ r .L pelts of raccoon are brought in Fig. 65. Distribution of the raccoon, , , t ^ » i i Procyon iotor fuscipes, in Coahuila. by Indians each year. An adult male from 2 miles south and 12 miles east of Nava weighed 13.0 pounds. Specimens examined answer to the description of fuscipes ( Gold- man, 1950:50). Four fragments of skulls recorded as Procyon Iotor by Gilmore (1947:155) from cave deposits in the vicinity of Cuatro Cienegas are considered here as belonging to the subspecies fuscipes. Elliot (1907:427) used the name Procyon Iotor hernandezi for spec- imens from Sabinas, which place is within the range now ascribed to P. I. fuscipes. Measurements. — The adult male from 2 miles south and 12 miles east of Nava measures: total length, 819; length of tail vertebrae, 274; length of hind foot, 128; height of ear from notch, 68; condylobasal length of skull, 115.2; basilar length, 105.5; zygomatic breadth, 83.4; least interorbital constriction, 24.2; alveolar length of maxillary tooth-row^, 42.5; length of upper camassial tooth, 7.9; width of upper camassial tooth, 7.7. Specimens examined. — Total, 11, from: opposite Langtry, [Val Verde County], Texas, 3 (BSC); 2 mi. S and 12 mi. E Nava, 1; Mariposa Ranch, 36 mi. NW Muzquiz, 1 (BSC); La Gacha, 1600 ft, 1; Sabinas, 3 (BSC); base Don Martin Dam, 1; 34 mi. W Chorro, 3 mi. S and 11 mi. E Saltillo, 1. Additional records (Elliot, 1907:427): Sabinas; (Gilmore, 1947:155): 16 mi. E and 3 mi. S Cuatro Cienegas. 304 University of Kansas Publs., Mus. Nat. Hist. Nasua narica tamaulipensis Goldman Coati Nasua narica tamaulipensis Goldman, Proc. Biol. Soc. Washington, 55:80, June 25, 1942, type from Cerro de la Silk, near Monterrey, Nuevo Leon. Distribution. — Eastern Coahuila. Remarks. — The coati occurs in small numbers along forested streams and in the foothills of tlie higher desert ranges of eastern Coahuila. Goldman (1942:80) writes that in Coahuila the coati is found on warmer, low mountain slopes and other forested parts and that it is rare along the Rio Grande as far upstream as the mouth of the Pecos River. I have received no definite records of the coati along the Rio Grande in Coahuila. The species is well known to residents of the mountains of southeastern Coahuila, especially in the area east and south of Saltillo. In a northwestward direction from Saltillo the coati seems to be progressively less common. A few were reported in the Sierra de San Marcos by Seiior Tomas Cantu of Estanque de Norias in 1952. Mr. David McKellar of the Hacienda La Mariposa told me coati occur in the mountains north- west of Muzquiz; Marsh (1937:27) quotes McKellar as saying that one was taken ". . . near the mouth of Santa Anna Canyon [=: Puerta Santana] in 1928." Marsh also reports that the coati lives in the Sierra de la Encantada and along the Arroyo de la Babia. Goldman (1942:81) says that Stanley Young found coati in the foot- hills of the Sierra del Carmen. Some records of the coati in Coa- huila result from the finding of escaped pets. Mr. Roberto Spence told me that he liberated a pair of coati at Fortin, which is in the pine-oak association of the Serranias del Burro. Goldman (1942:80) hsts Coahuila as within the range of N. n. tamaulipensis. The only actual record of coati from the State is that of Gilmore (1947:155), who identified as Nasua narica, bone frag- ments from cave deposits from the vicinity of Cuatro Cienegas. Probably Nasua reached this area by moving upstream along the watershed of the Rio Salado. Hershkovitz (1951:56) considers tamaulipensis as a synonym of N. narica molaris Merriam. Mustela frenata frenata Lichtenstein Long-tailed Weasel Mustela frenata Lichtenstein, Darstcllung neuer oder wenig bekannter Siingc- thier . . ., 1832, pi. 42, and corresponding, unpaged text, type from Mexico City, Mexico. Distribution. — Eastern Coahuila, perhaps no farther north than the northern end of the Sierra Madre Oriental, east of Monclova. Mammals of Coahuila, Mexico 305 Remarks.— The long-tailed weasel is presumed to be a rare mam- mal in Coahuila. None was obtained by University of Kansas field parties. Reports from competent local observers indicate that the animal is unknown in most parts of the State. Diego Rubio of Boquillas, David McKellar of La Mariposa, Roberto Spence of Sabinas, Juan Jose Zapata of Hacienda Rincon all have told me that this weasel does not occur in their areas. In southeastern Coahuila one specimen is known and several residents in the mountains south- east of Saltillo are acquainted with the animal, calling it "onza." At the Canon de la Carolina in the Sierra Madre Oriental, 13 miles east of San Antonio de las Alazanas, one resident reported to me that a weasel entered his chicken house one night. Aside from the one individual this life-long resident of the mountains could recall having seen no other weasels. One long-tailed weasel was taken by Dr. Edward Palmer on August 11, 1880. Allen (1881:183) records the animal as Putorius brasiliensis jrenatus, remarking that Dr. Palmer obtained it in the mountains near Saltillo and that the species is uncommon. I suspect that the mountains referred to are those south and east of the city. Through the courtesy of Dr. Charles P. Lyman of the Museum of Comparative Zoology, I have been able to examine the skull of this male animal which bears MCZ number 6375; the skin, number 6230, is not extant. The skull is that of a young animal (as defined by Hall, 1951b: 25) and measures: basilar length, 47.2; length of upper tooth-rows. 18.0; breadth of rostrum, 13.9; least interorbital constric- tion, 11.4; orbitonasal length, 16.6; mastoidal breadth, 26.1. This specimen is referred to M. f. frenata principally on geographic grounds. Hall (op. cit. :221) regards western Coahuila as within the range of M. /. neomexicana (Barber and Cockerell). Specimens examined. — One from mountains near Saltillo (MCZ). Taxidea taxus berlandieri Baird Badger Taxidea berlandieri Baird, Mammals, in Rep. Expl. Surv. Railr. to Pacific, vol. 8, pt. 1 (Washington, 1857), July 14, 1858, type from Llano Esta- cado, Texas, near border of New Mexico. Taxidea taxus berlandieri J. A. Allen, Bull. Amer. Mus. Nat. Hist., 7:256, June 29, 1895. Distribution. — Found in suitable localities throughout Coahuila, (see fig. 66). Remarks. — The badger occurs in most parts of the State with the exception of the higher mountains. It is nowhere common but is well-knovm to local residents. Most specimens obtained were either 306 UNrvTERsiTY OF Kansas Publs., Mus. Nat. Hist. received from native hunters or were road-kills. Burrows and other sign of badger often were observed where pocket gophers (Crato- geomys castanops) or ground squirrels {Spermophilus) occurred in abundance. I saw one badger run across the highway approxi- mately 20 miles south of San Lazaro in April. This badger moved rapidly keeping a definite course across the road rather than at- tempting to dodge the car or to turn back. Schantz (1949:301) referred one Coahuilan specimen from La Ventura to littoralis. This subspecies is described as differing from berlandieri in color and size of skull. Two skins from northeastern Coahuila, taken approximately 47 miles apart, differ markedly in color. One, an unsexed, January-taken skin of either subadult or adult age from the vicinity of Sabinas, is pale, less washed with black above and has a short, rela- tively inconspicuous white me- dian-dorsal line down the back; and, by these characters, resem- bles berlandieri. The other, an August-taken skin of an adult female from Hermanas, is much darker above and has a conspic- uous white median-dorsal stripe down the back as far as the rump and seems closest to the descrip- tion of littoralis. The one avail- able adult skull, that of a fe- male from Hermanas, is slightly smaller but in other ways com- pares favorably with a female of equal age of berlandieri from McKinley County, New Mexico. Owing to the variability of the few specimens available, I am tentatively referring badgers from eastern Coahuila to ber- landieri. Also assigned to this subspecies are bone fragments iden- tified as Taxidea taxus by Gilmore (1947:156) from cave deposits obtained from the vicinity of Cuatro Cienegas. Measurements. — An adult female from Hermanas measures: total length, 662; length of tail vertebrae, 130; length of hind foot, 100; height of ear from crown, 46; condylobasal length of skull, 117.2; basilar length, 106.5; zygomatic Fic:. 66. Distribution of the badger, Taxidea taxus berlandieri, in Coahuila. Mammals of Coahuila, Mexico 307 breadth, 71.0; mastoidal breadth, 71.7; least interorbital constriction, 24.3; least postorbital constriction, 23.5 length of upper tooth-rows, 48.4; outside length of first upper molar, 13.0; outside length of fourth upper premolar, 11.9; breadth of fourth upper premolar, 9.8. Specimens examined. — Total, 3, from: Boquillas, 1700 ft., 1; 4 mi. N Sabinas, 1; Hermanas, 1300 ft., 1. Additional records: 22 mi. W and 4 mi. S Cuatro Cienegas and from 16 mi. E and 3 mi. S Cuatro Cienegas (Gilmore, 1947:156); Saltillo (Allen, 1881: 183). La Ventura (Schantz, 1949:302). Spilogale gracilis gracilis Merriam Spotted Slcunk Spilogale gracilis Merriam, N. Amer. Fauna 3:83, September 11, 1890, type from Grand Canyon of the Colorado River, north of San Francisco Moun- tain, 3500 ft., Coconino County, Arizona. Distribution. — Lower elevations of western and southern Coahuila (see fig. 67). Remarks. — The spotted skunk is less common in Coahuila than either the striped or hooded skunks. Marsh (1937:29) reports that night hunting with dogs at the Rancho La Mariposa yielded 30 skunks, of which only three were Spilogale. Called "zorrilla" or "zorrilla chica" by the natives, this species is well known in the western and southern parts of the State. Probably Spilogale indianola Merriam occurs on the Gulf Coastal Plain, but no rec- ords are known. Spotted skunks prefer rocky areas and foothills, where en- trance may be gained to cavities in outcroppings. One animal was trapped at the base of the rimrock of a low hill near Bo- quilas. A ringtail cat was ob- tained in another steel trap set the same night within 40 feet of the one taking the spotted skunk. The only evidence that this spe- cies occupies the higher areas of mountains was obtained by Marsh ( loc. cit. ) who observed a dead animal at Jardin del Sur in the Sierra del Carmen. He writes that three coyotes were fighting FiG. 67. Distribution of the spotted skunk, Spilogale gracilis gracilis, in Coahuila. 308 University of Kansas Publs., Mus. Nat. Hist. over the carcass of this animal as he approached it. One adult male from near Boquillas (see below) weighed 399.2 grams. Eight bone fragments from cave deposits from the vicinity of Cuatro Cienegas, identified as Spilogale leucoparia by Gilmore (1947:156), are referred here, on geographic grounds, to S. g. gracilis. Measurements. — An adult male from 7 miles south and 2 miles east of Boquillas and another from the west foot of Pico de Jimulco, respectively, meas- ure: total length, 400, 380; length of tail vertebrae, 140, 130; length of hind foot, 47, 45; height of ear from notch, 30, 23; condylobasal length (of skull), 56.3, 52.6; basilar length, 49.9, 46.3; least interorbital breadth, 15.9, 16.0; orbitonasal length, 19.1, 18.5; mastoidal breadth, 32.1, 30.2; zygomatic breadth, 35.3, 33.8; length of upper tooth-rows, 21.1, 20.4. Specimens examined. — Total, 6, from: 7 mi. S and 2 mi. E Boquillas, 1800 ft., 1; Saltillo, 1 (BSC); W foot Pico de Jimulco, 4700 ft., 1; Sierra Encamacion, 3 (BSC). Additional records (Gilmore, 1947:156): 22 mi. W and 4 mi. S Cuatro Cienegas; 16 mi. E and 3 mi. S Cuatro Cienegas. Mephitis macroura milleri Meams Hooded Skunk Mephitis milleri Meams, Prelimin- ary diagnoses of new mammals of the genera Mephitis, Dorcela- phus, and Dicotyles, from the Mexican Boundary of the United States, p. 1 February 11, 1897. Preprint of Proc. U. S. Nat. Mus., 20:467, December 24, 1897, type from Fort Lowell, Pima County, Arizona. Mephitis macroura milleri Allen, Bull. Amer. Mus. Nat. Hist., 14: 334, November 12, 1901. Distribution. — Western and south- em Coahuila (see fig. 68). Remarks. — The hooded skunk, known to residents as "zorillo," is found in western and southern Coahuila. The ranges of M. macroura and M. mephitis over- lap in northern Coahuila, but both species were not taken at the same places. Like M. mephitis, the hooded skunk is seemingly more abundant in streamside vegetation and near water in forested or shrubby uplands than in arid situations on the desert plain. Measurements. — A subadult male from Juarez Canon measures: total length, Fig. 68. Distribution of the hooded skunk, Mephitis macroura milleri, in Coahuila. Mammals of Coahuela, Mexico 309 r=po 617; length of tail vertebrae, 290; length of hind foot, 65; basilar length, 52.0; zygomatic breadth, 39.1; mastoidal breadth, 32.9; length of upper tooth-rows, 24.4; outside length of first upper molar, 6.5; outside length of fourth upper premolar, 6.2; breadth of fourth upper premolar, 5.9. Specimens examined. — Total, 5, from: Juarez Cafion, Sierra del Carmen, 1 (DMNH); Sierra Guadalupe, 1 (BSC); Jimulco, 4050 ft, 2 (BSC); La Ven- tura, 5600 ft., 1 (BSC). Mephitis mephitis varians Gray Striped Skunk Mephitis varians Gray, Charlesworth's Mag. Nat. Hist., new ser., 1:581, 1837, type from Texas. Mephitis mephitis varians Hall, Carnegie Inst. Washington, Publ. no. 473: 66, November 20, 1936. Distribution. — Northern and northeastern Coahuila (see fig. 69). Remarks. — The .striped skunk occurs in most of northern and east- ern Coahuila but is most com- mon on the Gulf Coastal Plain. This animal is well known to the residents, who call it "zorillo." Specimens have been taken at localities along the watersheds of the Rio Grande, Rio Sabinas, and Rio Salado, The species seems more abundant in stream- side vegetation at lower eleva- tions. Animals were taken also near water in chaparral and other dense shrubs on foothills, often adjacent to rocky ledges. Marsh (1937:30) observed skunks both in the Sierra del Carmen and in the Sierra Hermosa de Santa Rosa; he saw one animal at an elevation of approximately 7000 feet in the Carmens. He writes also that "In the spring of 1935 I saw and captiu-ed at least thirty specimens of [striped] skunk on the plains to the south of Piedra Blanca and east of Muzquiz." Taylor et al. (1945:25) saw no specimens but reports diggings probably of this species at Tinajas de Chavez in the Sierra del Carmen. An adult male, taken in April 6 miles southwest of San Geronimo, weighed 1144.2 grams. Bone-fragments from cave deposits from the vicinity of Cuatro 100 _ I Fig. 69. Distribution of the striped skunk. Mephitis mephitis varians, in Coahuila. 310 University of Kansas Publs., Mus. Nat. Hist, Cienegas, identified by Gilmore (1947:156) as Mephitis mephitis, are here referred, on geographic grounds, to M. m. varians. Measurements. — Two adult males from 1 mile north of Boquillas and 6 miles southwest of San Geronimo and one adult female from 2 miles west of Jimenez, respectively, measure: total length, 666, 680 and 650; length of tail vertebrae, 320, 310 and 325; length of hind foot, 74, 72 and 64; height of ear from notch, 30, 32 and 30; basilar length, 61.1, 61.2 and 57.5; zygomatic breadth, 43.6, 43.1 and 41.5; mastoidal breadth, 38.1, 35.2 and 36.8; length of upper tooth-rows, — , 27.4 and 27.9; outside length of first upper molar, 7.4, 7.0 and 7.0; outside length of fourth upper premolar, 7.4, 6.8 and 6.5; breadth of fourth upper premolar, 7.7, 7.1 and 7.3. Specimens examined. — Total, 13, from: opposite Langtry, [Val Verde County], Texas, 1 (BSC); Las Vacas [= Villa Acufia], 1 (BSC); 1 mi. N Boquillas, 1700 ft., 1; 2 mi. W Jimenez, 850 ft., 2; 1 mi. E Las Margaritas, 3100 ft., 1; 2 mi. S and 5 mi. E San Geronimo, 1; 6 mi. SW San Geronimo, 2; 4 mi. W Hacienda La Mariposa, 2300 ft., 3; Sabinas, 1 (BSC). Additional record (Gilmore, 1947:156): 16 mi. E and 3 mi. S Cuatro Ciene- gas. Conepatus mesoleucus meamsi Merriam Hog-nosed Skunk Conepatus mesoleucus meamsi Merriam, Proc. Biol. Soc. Washington, 15: 163, August 6, 1902, type from Mason, Mason County, Texas. Conepatus pediculus Merriam, Proc. Biol. Soc. Washington, 15:164, August 6, 1902, type from Sierra Guadalupe, Coahuila. Conepatus mesoleucus pediculus Hall and Kelson, Univ. Kansas Publ., Mus. Nat. Hist., 5:335- December 5. 1952. Distribution. — Suitable habitat throughout Coahuila (see fig. 70). Remarks. — The hog-nosed skunk occurs in most parts of Coahuila but prefers mesic areas where shrubby vegetation grows either along arroyos and streams at low elevations or in oak forests on uplands. Specimens were obtained from elevations from as low as 800 feet at Presa Don Martin to as high as 7000 feet in the Sierra de Guada- lupe. Most specimens were taken in steel traps or shot at night with the aid of a headlight; one was taken with the help of a dog from a burrow in a cane thicket at the bank of a stream. Coahuilan hog-nosed skunks are assignable to C. mesoleucus although one male from Don Martin is large and in some respects approaches C. leuconotus texensis Merriam but not sufficiently so to be regarded as an intergrade between these two alleged species (see Hall and Kelson, 1952:336-337). Merriam (1902:164) de- scribed C. pediculus as the smallest hog-nosed skunk. E. Raymond Hall, who has examined the type, informs me that the holotype is approximately two-thirds grown. Recently Hall and Kelson (op. cit.:SS5) arranged this species as C. mesoleucus pediculus, stating that they tentatively recognize the subspecies as being different from C. m. meamsi. I am here placing the name pediculus as a Mammals of Coahuila, Mexico 311 synonym of meartisi. The type of pediculus is small, but a second specimen from the Sierra de Guadalupe, taken in 1953 by a Univer- sity of Kansas field party, is an adult female the measurements of which are within the size- range of females of mearnsi, and has larger cranial measurements tlian those listed by Hall and Kelson {op. cit.:SS7) for an adult female from 8 miles south of Langtry, Val Verde County, Texas. Bone fragments from cave de- posits from the vicinity of Cua- tro Cienegas, identified as Cone- patiis mesoleucus by Gilmore (1947:156), are here referred to mearnsi. Measurements. — Measurements of an adult male from 2 miles west of Jimenez and an adult female from Sierra Guadalupe, 10 miles south and 5 miles west of General Cepeda, re- spectively, are: total length, 728, 600; length of tail vertebrae, 287, 255; length of hind foot, 85, 70; height of ear from notch, 28, 28; condylobasal length, 76.1, 66.3; zygomatic breadth, 51.7, 44.2; mastoidal breadth, 44.1, 37.6; length of upper tooth-rows, 28.9, 27.0; outside length of fourth upper pre- molar, 7.0, 6.4; outside length of first upper molar, 7.7, 7.8; breadth of fourth upper premolar, 7.7, 7.6. Specimens examined. — Total, 8, from: 2 mi. W Jimenez, 850 ft., 2; Don Martin, 800 ft, 1; 3 mi. SE Torreon, 3800 ft., 1; Encantada, 5000 ft., 1; 3 mi. S and 13 mi. E San Antonio de las Alazanas, 8900 ft., 1; Sierra Guadalupe, 7000 ft., 10 mi. S and 5 mi. W General Cepeda, 1; Sierra Guadalupe, 1 (BSC); Cameros, 6500 ft., 1 (BSC). Additional records (Gilmore, 1947:156): 16 mi. E and 4 mi. S Cuatro Cienegas. Felis concolor stanleyana Goldman Mountain Lion Felis concolor youngi Goldman, Proc. Biol. Soc. Washington, 49:137, August 22, 1936, type from Bruni Ranch, near Bruni, southeastern Webb County, Texas. Felis concolor stanleyana Goldman, Proc. Biol. Soc. Washington, 51:63, March 18, 1938. Substitute name for Felis concolor youngi, preoccupied by Felis youngi Pei (Paleo. Sinica, ser. C, vol. 8, fasc. 1, p. 133, May 1934). Distribution. — State-wide. Most common in mountains of central and north- em Coahuila (see fig. 71). Fig. 70. Distribution of the hog-nosed skunk, Conepatus mesoleucus mearnsi, in Coahuila. 312 Unr^ersity of Kansas Publs., Mus. Nat. Hist. Remarks. — The mountain lion or puma occurs throughout Coa- huila, but is most common in the mountains of northern and cen- tral Coahuila. This large cat is regarded everywhere as a menace to livestock, especially to goats and horses, and is shot or trapped whenever possible. At least one rancher, Senor Urbano Santo at San Geronimo, defends the animal and regards it as an efiEective predator on the coyote. Sefior Santos told me in 1950 that where the Hon occurs on his ranches the coyote is absent. The use of coyote as food by the puma is recorded also by Young (in Young and Goldman, 1946:136- 137). A fascinating account of the puma in the Sierra del Carmen was written by Gregory ( 1938 ) , who photographed animals by night flash. He and his party were successful in getting pic- tures of several pumas. Taylor et at. (1945:25 and 1946:21) re- ported tracks in the Carmens ( at Tinajas de Chavez and between Campo Central and the Tinajas de Chavez) in April, 1945. Marsh (1937:33-34) also records the puma in the Sierra del Carmen. He adds that the lion is known to feed on deer and colts. Lion are reported to be numerous in the Sierra del Pino and Sierra de los Hecheros in northwestern Coahuila and as far southeast as Sierra de San Marcos. Six fragments of bone from cave deposits from near Cuatro Ciene- gas, identified as Felis concolor by Gilmore (1947:156), are as- signed here, on geographic grounds, to F. c. stanleyana. Measurements. — Cranial measurements of a young adult male from 5 miles northwest of the Hacienda Las Margaritas are: greatest length of skull, 192; condylobasal length, 173; least interorbital constriction, 39.1; width of nasals (at anterior tips of frontals), 13.9; width of palate (across interpterygoid fossa), 28.0; mastoidal breadth, 79.3; length of maxillary tooth-row (from front of canine to back of camassial), 60.0; lengtli of fourth upper premolar, 28.5; length of first lower molar, 17.5. Fig. 71. Distribution of the puma, Felis concolor stanleijana, in Coahuila. Mammals of Coahuila, Mexico 313 Specimens examined. — Total, 3, from: Carmen Mountains (southern end), 1 (BSC); 5 mi. NW Hacienda Las Margaritas, 3000 ft, 1; 6 mi. N San Ger6nimo, 1 ( skin only ) . Additional record (Gilmore, 1947:156): 16 mi. E and 3 mi. S Cuatro Cienegas. Felis onca veraecrucis Nelson and Goldman Jaguar Felis onca veraecrucis Nelson and Goldman, Jour. Mammalogy, 14:236, August 17, 1933, type from San Andres Tuxtla, Veracruz. Distribution. — An infrequent straggler into Coahuila from the southeast. Remarks. — The jaguar is now not a permanent resident in Coa- huila, and the occasional reports received probably are of stragglers that range into the State from the southeast. In colonial days and earher this cat may have been resident in eastern Coahuila. The watersheds of the Rio Grande and the Rio Salado-Rio Sabinas pro- vide Hkely avenues for the movement of wandering jaguars into Coahuila. An early record along the Rio Grande is that of Baird (1859:8) who writes of seeing the skin of a large jaguar ". . . killed near the mouth of the little stream Las Moras, above Eagle Pass." This place is in Texas directly across the Rio Grande from Coahuila. Records of jaguar in Coahuila provide widely circulated stories. Reports of a single animal may be heard from several sources, each version being slightly different. Such was the case concerning a jaguar roped on the Rancho La Mariposa in 1908. Mr. Roberto Spence told me that another jaguar was taken at El Oso which is northeast of the Mariposa. Marsh ( 1937:34-35) writes that a jaguar was seen by Susano Treviiio at El Jardin in the Sierra del Carmen in the summer of 1932. Three phalanges, found in cave-deposits from the vicinity of Cuatro Cienegas, thought to be Felis onca by Gilmore (1947:156), provide the only part of any specimen saved for study from the State. Felis pardalis albescens Pucheran Ocelot Felis albescens Pucheran, in J. GeoflFroy-Saint-Hilaire, Voy. Venus, Zool. Mamm., p. 149, Atlas, pi. 8, 1855, type from Arkansas. Felis pardalis albescens J. A. Allen, Bull. Amer. Mus. Nat. Hist., 22:219, July 25, 1906. Distribution. — Eastern Coahuila. Remarks. — The ocelot is found infrequently in shrub vegetation along waterways and in the foothills of the mountains of eastern Coahuila. None was obtained, but specimens from along the Rio Grande, on the Texas side opposite Coahuila have been taken at Eagle Pass and in Brewster County (Goldman, 1943:380). The 314 University of Kansas Publs., Mus. Nat. Hist. ocelot is well known as "tigrillo" to residents in eastern Coahuila. Reliable persons have told me that this cat lives along the Rio Sabinas, at least as far upstream as Ciudad Muzquiz. Senor Tomas Cantu of Estanque de Norias said that there was an occasional re- port of ocelot in the vicinity of the Sierra de San Marcos. It occm^s also in the mountains south and east of Saltillo. The margay, Felis wiedii cooperi Goldman, may occur in Coa- huila, because one was obtained directly across the Rio Grande from Coahuila by Col. S. Cooper of the U. S. Army at Eagle Pass, Mav- erick County, Texas, more than 100 years ago (see Goldman, 1943: 384-385). Although no other margays are known from this part of North America, Goldman considers the locality for this specimen to be vahd remarking that this ". . . isolated specimen tends to emphasize the comparative rarity of the species." Certainly it would be advisable to be watchful for margays in eastern Coahuila. Felis yagouaroundi cacomltli Berlandier Jaguarundi Felis cacomitli Berlandier, in Baird, Mammals of the boundary, in Emory, Rep. United States and Mexican boundary survey, . . ., vol. 2, pt. 2, p. 12, January, 1859, type from Matamoros, Tamaulipas. Felis yagouaroundi cacomitli Miller and Kellogg, U. S. Nat. Mus., Bull. 205: 776, March 3, 1955. Distribution. — Brushy areas along major streams in northeastern Coahuila. Remarks. — Like the ocelot, the jaguarundi occurs uncommonly within the boundaries of Coahuila. Brush lands affording suitable habitat for this species are found along the Rio Grande in the vi- cinity of Nava and Guerrero and along the Rio Salado in the vi- cinity of Presa Don Martin. Reliable local residents know the ani- mal, but no specimens were obtained. Lynx rufus Bobcat Bobcats live in most parts of Coahuila but are confined chiefly to thick brush along arroyos in the lowlands and to rocky and forested areas in the mountains. Most specimens examined are from lower elevations, although one was obtained at 8900 feet eleva- tion near San Antonio de las Alazanas. Gregory (1938) made ex- cellent flashlight pictures of bobcats in the Sierra del Carmen; Marsh (1937:35) and Taylor et al. (1945:25) also report the occur- rence of this animal in the Carmens. I shot a large male on March 30 while hunting at night with a headlight in dense brush along the Rio Salado less than one mile below the Presa Don Martin. As I Mammals of Coahuila, Mexico 315 approached a beaver pond, from which I could hear sounds of the activity of beaver, I spotted a bobcat ahead of me; the animal also seemed to be stalking the beaver since it was slinking along look- ing in the direction from which the sounds of the beaver were coming. The cat sensed my presence and turned to look into my light, moved away several steps, turned to look again as I shot him with a no. 6 load from my 16 gauge shotgun. The bobcat, called "gato mon- tez" by residents, has a bad reputation, because of its occa- sional predations on goats and poultry. However, few ranchers trap bobcats. One subadult fe- male from La Gacha weighed 6360 grams. Lynx rufus baileyi Merriam Fig. 72. Distribution in Coahuila of the bobcat. 1. Lynx rufus baileyi. 2. Lynx rufus texensis. Lynx baileyi Merriam, N. Amer. Fauna, 3:79, September 11, 1890, type from Moccasin Spring, north of Colorado River, Coconimo County, Arizona. Lynx rufus baileyi V. Bailey, N. Amer. Fauna 53:291, March 1, 1932. Distribution. — Western part of Coahuila (see fig. 72). Remarks. — Skulls of two bobcats from northwestern Coahuila are slightly smaller than those of specimens from the eastern part of the State and are presumed to be baileyi, which occurs throughout the Mesa del Norte. Measurements. — Two skulls of subadult bobcats (see below), se.x unde- termined, measure: basilar length, 104.2, 100.6; least interorbital constriction, 25.2, — ; orbitonasal length (after Hall, 1951b:417), 54.7, 54.7; mastoidal breadth, 54.1, 54.3; zygomatic breadth, 88.4, 85.3; length of tooth-rows, 43.9, — ; length of first lower molar, — , 10.2; length (lateral) of fourth upper pre- molar, — , 14.5; length (medial) of fourth upper premolar, — , 14.7. Speciinens examined. — Total, 2, from: 2 mi. SW Carricitos, 3500 ft., 1; 9 mi. N and 6 mi. W Castillon, 4100 ft., 1. Lynx rufus texensis J. A. Allen Lynx texensis J. A. Allen, Bull. Amer. Mus. Nat. Hist., 7:188, June 20, 1895 (based on Lynx rufus var. maculatus Audubon and Bachman, The vivi- parous quadrupeds of North America, 2:293, 1851.), type from vicinity of Castroville, on headwaters of Medina River, Medina County, Texas. 316 University of Kansas Publs., Mus. Nat. Hist. Lynx rufus texensis Mearns, Preliminary diagnoses of new mammals of the genera Lynx, Urocyon, Spilogale, and Mephitis, from the Mexican bound- ary line, p. 2, January 12, 1897 (preprint of Proc. U. S. Nat. Mus., 20: 458, December 24, 1897). Distribution. — Eastern part of Coahviila (see fig. 72). Remarks. — Specimens from the Coastal Plain and southeastern Coahuila are assigned to texensis because of their large size, rich coloring and extensive blackish markings. Evidently this subspe- cies occurs at least into the central mountains and intergrades with L. r. baileyi in the western part of the State. Gilmore (1947:156) records bone fragments of Lynx rufus from cave deposits from the vicinity of Cuatro Cienegas; these are probably representative of texensis. Measurements. — An adult male from Don Martin and a subadult female from La Gacha, respectively, measure: total length, 940, 805; length of tail vertebrae, 170, 155; length of hind foot, 191, 162; height of ear from notch, 83, 78; basilar length, 107.3, 97.9; least interorbital constriction, 27.0, 24.0; orbitonasal length (after Hall, 1951b:417), 58.1, 51.7; mastoidal breadth, 57.2, 53.0; zygomatic breadth, 92.6, 82.4; length of tooth-rows, 43.5, 41.9; length of first lower molar, 10.8, 11.0; length (lateral) of fourth upper premolar, 12.4, 14.8; length (medial) of fourth upper premolar, 12.5, 14.6. Specimens examined. — Total, 5, from: 4 mi. E Hacienda Las Margaritas, 3160 ft., 1; La Gacha, 1600 ft, 2; Don Martin, 1; 3 mi. S and 13 mi. E San Antonio de las Alazanas, 8900 ft., 1. Additional records (Gilmore, 1947:156): 16 mi. E and 3 mi. S Cuatro Cienegas, 22 mi. W and 4 mi. S Cuatro Cienegas. Pecari tajacu angulatus (Cope) Collared Peccary D[icotyles] angulatus Cope, Amer. Nat., 23:147, February 1889, type from Guadalupe River, Texas. Pecari tajacu angulatus Hershkovitz, Fieldiana — Zool., Chicago Nat. Hist. Mus. Publ. 670, 31:566, July 10, 1951. Distribution. — Desert plains of Coahuila (see fig. 73). Remarks. — The collared peccary or javelina is a resident of the desert plains of Coahuila (see Leopold, 1947:443, fig. 5 and Villa R., 1951:8, map 1). The animal lives in small bands and seems to con- centrate its activities in the vicinity of surface water; at least tracks of the javelina are most numerous at watering places. The species is most common in the eastern part of the State, especially where ranchers have protected these animals from excessive hunting. Even so, javelina meat is often a major food ration in cattle camps. The javelina is easily obtained with the aid of dogs, who after short chases bring the animals to bay in thickets or in retreats in caves or beneath rocky ledges. It is considered highly sporting to hunt these animals on horseback using as weapons either spears or pistols. Mammals of Coahuila, Mexico 317 Doctor Marcelino Garza-Pena of Saltillo told of several such hunts taking place on the Rancho Guadalupe. The collared peccary is perhaps most numerous in dense brush on ranches north and east of Ciudad Muzquiz, where prickly pear and other favored foods are abundant and ranch tanks and other watering places, built by the ranchers, are well distributed (see Marsh, 1937:39-40). Near the Rio Grande east of Nava, Seiior Alvaro Trevifio told me that the javelina is scarce because of overhunting. West of the Sierra del Carmen, in the vicinity of Boquillas, javelinas are rare ac- cording to Seiior Diego Rubio, a goatherder living near Boquillas. Also on the plains surrounding the Sierra del Pino, peccaries are uncommon; Sefior Armando Verduzco of the Hacienda San Ildefonso told me that one ani- mal was killed on his ranch in 1949. To the west of the Pinos, near the Chihuahuan border and south of the Sierra de los Hech- eros, collared peccaries are re- ported to be more numerous. The animal is not in imme- diate danger of extinction in Coahuila, but it is becoming less numerous. Local protection by interested land operators has been of primary importance in preserving these animals. Specimens from Jaral, recorded as Tagassu angulatum hnmerale (Merriam) by Elliot (1907:35), are referred to angulatus on geo- graphic grounds. Measurements. — External measurements of a young male and an adult female from 2 miles north of Hacienda La Mariposa, respectively, are: total length, 810, 940; length of tail vertebrae, 45, 55; length of hind foot, 180, 180; height of ear from notch, 90, 100. Cranial measurements of tw^o adult males from 2 miles north of Hacienda La Mariposa and of La Gacha and two adult females from 2 miles north of Hacienda La Mariposa and 2 miles northwest La Gacha, respectively, are: condylobasal length, 207, 205 and — , 213; basilar length, 190, 186 and — , 195; zygomatic breadth, 113.9, 114.7 and 105.3, — ; palatal length, 143, 147 and 142, 150; least interorbital constriction, 60.0, 55.0 and 58.8, — ; breadth of braincase across postorbital processes, 81.8, 82.3 and 7—1204 Fig. 73. Distribution of the collared peccary, Pecari tajacu angulatus, in Coahuila. 318 University of Kansas Publs., Mus. Nat. Hist, 80.4, — ; alveolar length of upper molarifomi tooth-row, 65.1, 60.1 and 60.1, 65.1; lengtli of post-canine diastema, 19.7, 21.5 and 20.3, 23.1. Specimens examined. — Total, 29, from Loma Gorda, 1700 ft., 6 mi. NE Hacienda La Mariposa, 12 (10 skulls only, 2 skins only); 2 mi. N Hacienda La Mariposa, 2300 ft., 10 (6 skulls only, 1 skin onl>); 3 mi. NE Hacienda La Mariposa, 2000 ft., 1; 7 mi. N La Gacha, 1600 ft., 2; 2 mi. NW La Gacha, 1600 ft., 1; La Gacha, 1600 ft., 3 ( 1 skin only). Additional record (Elliot, 1907:35): Jaral. Cen'us merriami Nelson Wapiti Cervus merriami Nelson, Bull. Amer. Mus. Nat. Hist., 16:7, January 16, 1902, type from head of Black River, White Mountains, Apache County, Arizona. Distribution. — Extinct in Coahuila but once occurred in mountainous parts of northern and central Coahuila. Remarks. — The wapiti, or American elk, once occurred in the mountainous parts of northern and central Coahuila. At least two lots of wapiti have been introduced into the state. A. Starker Leo- pold and local ranchers informed me that in March, 1941, 18 Cervtis canadensis from the Wichita National Wildlife Refuge, Oklahoma, were released on the Carmen Mountain Club in the Sierra del Carmen. These animals disappeared by 1943. In February, 1952, Jaime F. Garza of Monterrey brought 25 Cervus canadensis nelsoni (5 bulls, 15 cows and 10 calves) from Yellowstone National Park, Wyoming, via Piedras Negras, to his Rancho, Santa Hermosa, ap- proximately 50 kilometers west of Ciudad Muzquiz, and on the west side of the Sierra Hermosa de Santa Rosa. Nineteen animals, including 8 calves born at the rancho, remained in March, 1956. Gilmore (1947:160) records, as C. canadensis, 4 bone fragments from archaeologic cave-sites from the vicinity of Cuatro Cienegas. Probably this material represents the nominal species C. merriami. Odocoileus hemionus crooki (Meams) Mule Deer Dorcelaphus crooki Mearns, Preliminary diagnoses of new mammals of the genera Mephitis, Dorcelaphus, and Dicotyles, from the Mexican border of the United States, p. 2, February 11, 1897 (Preprint of Proc. U. S. Nat. Mus., 20:468, December 24, 1897), type from summit of the Dog Mountains, Grant County, New Mexico. Odocoileus hemionus crooki Goldman and Kellogg, Jour. Mamm., 20:507, November 14, 1939. Distribution. — Western Coahuila (see fig. 74). Remarks. — The mule deer is known as "venado burro" or simply "burro" to residents of Coahuila. It occurs on the desert plains and the rugged terrain of the arid foothills in western and southern Mammals of Coahuila, Mexico 319 Coahuila (see Leopold, 1947:442, fig. 4 and Villa-R., 1950:23, map 2 ) . This species does not occur on the Gulf Coastal Plain of north- eastern Coahuila being found no farther east than the western slopes of the Serranias del Burro and La Babia, according to ranch- ers Alden McKellar and Robert Spence. It is safe to say that the mule deer's range, at least to the eastward, extends no further than the northeastern limits of the lechuguilla. Marsh (1937:42) reports that the mule deer may occur to as high as 6000 feet elevation. For a large mammal the mule deer is notably successful in re- maining inconspicuous in its habitat. The "venado burro" seems nowhere abundant in Coahuila. Unlike the white-tailed deer, which is somewhat isolated in higher parts of the sierras, the mule deer lives in closer proximity to man and more people have the opportunity to hunt the mule deer for food. In the vicinity of Boquillas local residents shoot an occasional mule deer, usually by surprising an animal where it is bedded down at midday or by still-hunting animals along trails or at watering places. Dogs are sometimes used to track the animals. The mule deer is in danger of being exterminated, because the population is so reduced in numbers. Most individuals that remain are found only in the most re- mote areas in Coahuila. Where the human population is more dense in the southern part of the State, few mule deer remain; such is also the case in San Luis Potosi where Dalquest (1953:- 212) regards this species as nearly extinct. At the present time more local protection to the mule deer by interested land- owners is needed. Aside from a few mounted heads in homes in Saltillo, I found little evidence of the mule deer in Coahuila, except for tracks at watering holes. The antlers of one specimen were obtained in the vicinity of Boc- FiG. 74. Distribution of the mule quillas; another head was ob- deer, Odocoileus hemionus crooki, in . . , /- ., „. , , Coahuila. tamed from the Sierra de la 320 University of Kansas Publs., Mus. Nat. Hist. Paila. Taylor et al. (1945:27) record skulls only from Agua Chili on the Rancho El Jardin in the Sierra del Carmen. Both Leopold ( loc. cit. ) and Villa-R. ( loc. cit. ) cite f om- records of occurrence of the mule deer in Coahuila without giving exact locahties. Goldman and Kellogg (1939:507) refer the mule deer of Coa- huila to crooki. Bone fragments, identified as Odocoileiis hemionus by Gilmore (1947:160) from cave deposits from near Cuatro Ciene- gas, are assigned, on geographic grounds, to crooki. Measurements. — Cranial measurements of a young adult male from Sierra de los Hecheros are: condylobasal length, 266; greatest breadth, 122.5; least interorbital constriction, 77.5; length of nasals, 83.7; alveolar length of maxil- lary tooth-row, 83.3. Specimens examined. — Total, 3, from: 10 mi. S and 5 mi. E Bocquillas, 1 (partial skull and antlers only); Sierra de los Hecheros, 1 (cranium only); Sierra de la Paila, 1 (cranium only). Additional record (Taylor et ah, 1945:27): Agua Chili, Rancho El Jardin in Sierra del Carmen. Odocoileus virginianus White-tailed Deer White-tailed deer occur in two distinct habitats in Coahuila. One is the Gulf Coastal Plain of the northeastern part of the State; the other is the higher elevations of the mountains where oaks or conifers grow. The only overlap in the ranges of these two popu- lations seems to be in the eastern and southern foothills of the Serranas del Burro. In the lowlands of the northeast white-tailed deer prefer the more dense woody vegetation along waterways and occur also in open mesquite and huisache brush and in the live oak thickets east of Nava, where it was reported that acorns and green brier are favored foods. Crops in the vicinity of the Rio Grande and elsewhere are occasionally attractive to deer; Senor Alvaro Trevifio of the Rancho La Candelaria reports that these animals damage oats in winter. Marsh (1937:41) found the deer of the lowlands no higher in the mountains than 3500 feet elevation. He recorded animals north of Muzquiz as far as Santa Domingo and suggested that the animal occurs in grassy places in the lower valleys of the Sierra del Carmen. Along the Rio Sabinas between Sabinas and Nacimiento white-tailed deer occur in brush and parklike stands of pecan, cypress, and other trees. Deer receive little protection from hunting along this river. Often the animals are hunted on dark nights with a headlight and rifle. Dogs are seldom used in this area. In the mountains the deer are more abundant because hunting pressure is less severe, especially in Mammals of Coahuila, Mexico 321 isolated areas. These mountain white-tails occur as low as the lowest growth of oaks, where the ranges of the mule deer and the white-tailed deer overlap. Frequently small groups of white-tails will be seen feeding, usually on bunch grasses, on the side of moun- tain slopes, across the valley from the observer. In such situations tlie deer are easy targets for the marksman with a powerful rifle. These deer seem not to need surface water regularly. In the Sierra del Pino, Sierra de los Hecheros, Sierra de la Madera, and Sierra de San Marcos, permanent surface water is often lacking or in widely spaced situations, yet deer and their sign are abundantly observed in places several miles from the nearest water. Where permanent surface water is absent, domestic animals are not grazed, and deer have no serious competition for favored foods. Principal foods are acorns, oak browse, bunch grasses and sotol. It is reason- able to think that mountain populations of the white-tailed deer are holding their own in most places in Coahuila while the low- land populations of northeastern Coahuila, being exposed to more hunters, are fewer and in greater danger of extermination. The white-tailed deer, called "venado con cola blanca" or simply "ve- nado" is the principal big-game species in Coahuila and its im- portance as sport and as food for residents cannot be over emphasized. Venison is fre- quently the main dish at meal- time in livestock camps and for fencing crews. In the latter op- erations usually one man is assigned the task of keeping a supply of fresh meat in camp. Miners working out flourite de- posits in the Sierra de la Encan- tada in 1952 were depending to a large degree on jerked venison (and beans); it was reported to me that the white-tailed deer population was considerably re- duced in the vicinity of these mining operations. Utilization of white-tailed deer by pre- Conquest Indians is illustrated Fig. 75. Distribution in Coahuila of ^^ Giimore's (1947:160) report- the white-tailed deer, Odocoileus vir- ing 67 bone fragments of this ginianus. 1. O. u. carminis. 2. O. v. . . i •.. c miquihuanensis. 3. O. v. texanus. species m cave deposits from 322 University of Kansas Publs., Mus. Nat. Hist. near Cuatro Cienegas. Poaching and overhunting of deer on many ranches are being discouraged. Hunting clubs, such as the one at Sabinas, are urging landowners not to allow excessive hunting of their game populations. A spotted fawn was taken on August 13. A buck in molt, espe- cially noticeable on the flanks and rump, was obtained on Septem- ber 21. Two males, which had dropped their antlers a short time previously, were shot on April 5 in the Sierra de San Marcos. Odocoileus virginianus carminis Goldman and Kellogg Odocoileus virginianus carminis Goldman and Kellogg, Proc. Biol. Soc. Wash- ington, 53:81, June 28, 1940, type from Botellas Caiion, 6500 ft., Sierra del Carmen, Coahuila. Distribution. — Mountains of northern and central Coahuila at least as far south as Jaral (see fig. 75). Remarks. — According to Goldman and Kellogg (1940:81-82), carminis is smaller than texanus and has antlers with shorter tines. Some specimens from the Serranias del Burro (Rancho Las Mar- garitas), especially those from elevations as low as 2900 feet, measure larger than the type and a topotype of carminis; these animals are referable to carminis but show some resemblance to texanus. Probably carminis and texanus intergrade in the foothills of the Burros, especially on the southern and eastern sides. The antlers of a partial cranium only from the Sierra de la Encantada, directly south of the Sierra del Carmen, have longer tines than do any other examples of carminis at hand. To the southward, carminis occurs at least to Jaral. Skins of two males (taken in April) in the Sierra de San Marcos resemble those of carminis from the Burros; one of the two has the upper side of the tail dark although not so dark as in specimens of O. v. miquihuanensis from Tamaulipas. White-tailed deer, not seen by me, from Cuatro Cienegas and Jaral, classified as Odontocoelus battyi Allen by Elliot (1907:48), are here referred, on geographic grounds, to carminis. Measurements. — According to Goldman and Kellogg {loc. cit.), the type (male) and a female topotype, respectively, measure: total length, 1512, 1386; length of tail, 214, 182; length of hind foot, 403, 365; condylobasal length (of skull), 242.3, 234; length of maxilhiry tooth-row, 69.7, 70; width across orbit at frontojugal suture, 107.5, 95. Average and extreme measure- ments of the skulls of three adult males (with worn teeth) and four adult females (with worn teeth) from the Rancho Las Margaritas, respectively, are: condylobasal length, 253 (239-260), 239 (233-246); width across orbit at frontojugal suture, 106.6 (104.5-110.6), 97.8 (96.1-99.5); least interorbital constriction, 62.7 (59.7-66.7), 54.8 (53.7-55.7); length of nasals. 83.8 ^81.2- Mammals of Coahuila, Mexico 323 85.5), 80.4 (77.5-84.0); alveolar length of maxillary tooth-row, 72.3 (69.4- 74.5), 67.9 (64.5-72.6). Specimens examined. — Total, 33, from: Club Sierra del Carmen, 2 mi. N and 6 mi. W Piedra Blanca, 1 ( partial skull and antlers ) ; 10 mi. NW Hacienda Las Margaritas, 3800 ft., 1; 1 mi. N Pino Solo, 3500 ft., Rancho Las Margaritas, 1 (skull only); 2 mi. E Pino Solo, 3900 ft., Rancho Las Margaritas, 7 (6 skulls only); 1 mi. N Hacienda Las Margaritas, 3800 ft., 1 (skull only); 4 mi. W Hacienda Las Margaritas, 2900 ft., 1; 1 mi. W Hacienda Las Margaritas, 1 (skull only); Hacienda Las Margaritas, 2900 ft., 3 (skulls only); 4 mi. E Hacienda Las Margaritas, 4000 ft., 4 (3 skulls only); 3 mi. S and 8 mi. E Hechicero, Chihuahua, in Coahuila, 1 (partial skull and antlers); Sierra de la Encantada, 4400 ft., 38 mi. S and 23 mi. E Boquillas, 1 (partial skull and antlers); Sierra del Pino, 7925 ft., 5 mi. N and 9 mi. W Acebuches, 2 (antlers only); 1/2 mi. NW Ocampo, 1 (cranium and antlers); 4 mi. N and 21 mi. W Cuatro Cienegas, 6500 ft., 2 (partial skulls only); Sierra de San Marcos, 2 mi. N and 18 mi. W Santa Teresa, 1; Sierra de San Marcos, 26 mi. W Santa Teresa, 1; 15 mi. S Las Delicias, 4000 ft., 1 (partial cranium only). Additional records (Taylor et al., 1945:27): Campo Central, El Jardin Ranch, Sierra del Carmen; (Goldman and Kellogg, 1940:81): Botellas Canon, Sierra del Carmen; (Elliot, 1907:48): Cuatro Cienegas and Jaral. Odocoileus virginianus miquihuanensis Goldman and Kellogg Odocoileus virginianus miquihuanensis Goldman and Kellogg, Proc. Biol. Soc. Washington, 53:84, June 28, 1940, type from Sierra Madre Oriental, near Miquihuana, Tamaulipas. Distribution. — Mountains of southern Coahuila (see fig. 75). Remarks. — This subspecies occurs in extreme southern Coahuila. Goldman and Kellogg (1940:84) refer a specimen from the Sierra Guadalupe to miquihuanensis. A female from Sierra Encarnacion, in the U. S. National Museum (Biological Surveys Collection), also belongs to this subspecies. Measurements. — An adult male from the Sierra Guadalupe, according to Goldman and Kellogg (loc. cit.), measures: total length, 1530; length of tail, 270; length of hind foot, 420; condylobasal length (of skull), 247.5; length of maxillary tooth-row, 71.4; width across orbit at frontojugal suture, 108.5. Specimen examined. — One from Mesa de Tablas, 8500 ft. (skin only). Additional records. (Goldman and Kellogg, 1940:84): Sierra Guadalupe; ( BSC ) : Sierra Encarnacion. Odocoileus virginianus texanus (Meams) Dorcelaphus texanus Meams, Proc. Biol. Soc. Washington, 12:23, January 27, 1898, type from Fort Clark, Kinney County, Texas. Odocoileus virginianus texanus Lydekker, Catalogue of the ungulate mam- mals in the ... British Museum, 4:158, 163, 1915. Distribution. — Plains and adjacent foothills of northeastern Coahuila (see fig. 75). Remarks. — The large white-tailed deer of the Gulf Coastal Plain occurs chiefly along arroyos and in open brush country. According to Miller and Kellogg (1955:803), texanus occurs as far south as Monclova and along the Rio Grande upstream as far as Ojinaga, 324 University of Kansas Publs., Mus. Nat. Hist. Chihuahua. Reports recently obtained indicate that texanus does not occur further upstream on the Rio Grande than Villa Acuna. A partial cranium including antlers from the vicinity of Sabinas is large with long tines and resembles a male specimen of texanus from Colorado County, Texas. Specimen examined. — One partial cranium from 16 mi. N and 4 mi. E Sabinas. Antilocapra americana mexicana Merriam Pronghom Antilocapra americana mexicana Merriam, Proc. Biol. Soc. Washington, 14:31, April 5, 1901, type from Sierra en Media, Chihuahua. Distribution. — Formerly found in suitable locahties in most parts of Coa- huila, but now present at only a few places in northern and central Coahuila. Remarks. — The pronghom, locally called "berrendo," formerly occurred in most parts of Coahuila (see Leopold, 1947:438, fig. 1 and Villa R., 1951:22, map 2). In Coahuila, it probably was most numerous on the open grassland plains of the northeast and the broad, intermontane grasslands of the nortliern and central part of the State. Bands of pronghorns remained for many years rela- tively undisturbed by man on these intermontane grasslands. Nel- son (1925:62) estimated that 600 head of pronghom were in Coa- huila in 1922-1924; most of these were presumed to have been in this intermontane range. The animal usually avoids thick, brush lands, preferring the more open plains, where grasses dominate. Such habitat was undoubtedly much more extensive in former times before the introduction of domestic Hvestock. Today, a few animals occur on the intermontane grasslands and on ranches in the vicinity of Presa Don Martin along the border between Coahuila and Nuevo Leon according to Roberto Spence of Sabinas. The latter area is open grassland, seemingly ideal for pronghom. The animals are being given protection by landowners. Nelson (loo. cit.) described "bands of 50 to 100 antelope" pres- ent in 1922-1924 in the valley of the Encantada (tlie broad inter- montane valley surrounding the Hacienda La Encantada). Marsh (1937:42) writes that small herds were still present in 1936, and that numbers had not decreased or increased appreciably in the past ten years. In July, 1938, a few pronghorns were still foimd in this area; probably less than 50 head were present. At that time prong- horns were being given protection, altliough a few were shot, chiefly by guests at the ranch, who approached the animals by automobiles. When chased in this manner, the pronghorns would oftentimes Mammals of Coahuila, Mexico 325 escape by moving into thick stands of yucca which grew at the periphery of the grasslands. In 1952, mining operations for flourite in the Sierra de la Encantada brought many workmen into the gen- eral area; these people lived in scattered camps where surface de- posits of flourite were being worked. In March, 1952, much big game in the area, especially white-tailed deer, had been shot for use as food at these mining camps. Senor Francisco Gracia of Piedras Negras told me that one pronghorn had been seen on the Encantada in early 1952 but that the animals were doomed because of the presence of the mining operations. Pronghorn may occur along the Coahuila-Chihuahua border in the vicinity of Castillon, south of the Sierra de los Hecheros. Seiior Juan Jose Zapata of the Hacienda Rincon and other ranchmen in the vicinity of Castillon told me in July, 1953, that pronghorns were found west of Castillon, only a few miles beyond the border in Chihuahua on ranches where landowners protected the animals. Possibly, these animals also range into Coahuila or may eventually do so if given adequate protection. Pronghorn occur in central and south-central Coahuila, accord- ing to Sefior Guillermo Aral Espinosa, Dr. Gilberto Martinez Car- denas and Seiior Joaquin Belloc (in Villa R., 1951:20-23). Accord- ing to these persons, localities where pronghorns occurred in 1950 are Valle de la Paila, 40 kilometers north-northeast of Parras; Llano de Ocampo, 9 kilometers north-northeast of Ocampo; Llano del Guaje, 108 kilometers north of Ocampo ( Sierra del Pino ) ; La Playa, 80 kilometers north of Ocampo; the Picacho de Santa Tecla; Sierra del Rey and Sierra de la Fragua. In most of these localities the pronghorn is receiving at least some protection from hunting. Bone fragments, identified as A. americanus by Gilmore (1947:160), were obtained from cave deposits near Cuatro Cienegas. The subspecific status of the pronghorn in Coahuila has never been established. The name mexicana is used purely on geo- graphic grounds. Specimen examined. — None. Elliot (1907:54) records specimens taken at Jaral by Heller and Barber. Bison bison bison (Linneaus) Bison [Bos] bison Linnaeus, Systema naturae, ed. 10, 1:72, 1758, type from north- eastern Mexico (see below). [B]ison. bison Jordan, Manual of the vertebrae animals of the United States, ed., 5, p. 337, 1888. Distribution. — Extinct in Coahuila but thought to have occurred, within historic times, in the northeastern part of the State. 326 University of Kansas Publs., Mus. Nat. Hist. Remarks. — The bison does not occur in Coahuila today. Whether this species actually lived in Coahuila or any other part of Mexico, within historic times, has been debated. Recently, Reed (1952) presented evidence, from early writings, which he considers as proof that the bison never was found south of the Rio Grande (Rio Bravo del Norte), at least since the time of the Conquest. He shows reason to doubt that a bison was present in the menagerie of Montezuma at the time of the coming of Cortez to the Aztec capital, although Hornaday (1889:373) thought that this animal existed and might have come from what is now the state of Coa- huila. Accordingly, Reed suggests that the type locality of Bison bison bison (Linnaeus) should be reassigned from "Mexico" to the "southern or southeastern United States." However, there is evidence that bison occurred, even as late as the beginning of the Nineteenth Century, on the Gulf Coastal Plain of northeastern Coahuila, as far south as Monclova. Actually, the bison may have lingered in Coahuila to as late as the middle of the Nineteenth Century, owing to the slow settlement of the southern Great Plains and the northern part of the Central Plateau. Evi- dence of the presence of bison in Coahuila is based on the identi- fication, as Bison bison by Gilmore (1947:161) of eight bone frag- ments from cave deposits containing an accumulation of archeologic materials from the vicinity of Cuatro Cienegas. Further evidence is that of Dobie (1953:150-151) who cites a document, dated Jan- uary 24, 1806, at Monclova, recording the presence of bison in Coa- huila. This document states, in translation, that "Buffalo hunting expeditions in the settlements of this province are the cause of the neglect of families." Of course, there is a possibility that this refer- ence was to bison in northern Coahuila, which at that time, in- cluded what is now Texas. J. A. Allen (1881:184) records field notes of Dr. Edward Palmer concerning the bison in Coahuila as follows, "According to the testimony of old people . . . the Bison was very abundant about Monclova and Parras when the first settler reached these points, probably half a century after the con- quest. For some years they killed large numbers for food, but soon the bison ceased to appear. There seems to be no reason why, so far as the nature of the country is concerned, the Bison may not have ranged also to Saltillo." From this statement, it might seem that bison moved into Coahuila from the north periodically. Dobie (loc. cit.) also calls attention to the arroyo in northern Coahuila called Arroyo de los Cibolos, which means, in English, Buffalo Creek. Mammals of Coahuila, Mexico 327 Thomas (1911:154) regards Mexico as the type locaHty of the American bison because of the reference to "Taurus mexicanus" of Hernandez in the original "description" by Linnaeus. Judging from our present knowledge of the former range of this species in Mexico, I would restrict this type locahty to "northeastern Mexico" (see Seton, 1929:647, map 7). Ovis canadensis mexicana Merriam Bighorn Sheep Ovis mexicanus Merriam, Proc. Biol. Soc. Washington, 14:30, April 5, 1901, type from Lago de Santa Maria, Chihuahua. Ovis canadensis mexicanus Lydekker, The great and small game of Europe, western and northern Asia and America, p. 11, 1901. Distribution. — Formerly found in arid mountainous areas in the northwest- em half of Coahuila, now found in a few localities in the central and northwest- em part of the State ( see fig. 76 ) . Remarks. — Bighorn sheep probably occurred formerly in most desert ranges of western and northern Coahuila, at least as far south as the Sierra de la Paila. This is a considerably larger range than that shown by Leo- pold (1947:439, fig. 2). From reports received from older resi- dents, bighorns are more at home in the arid rimrock of the foothills and less elevated moun- tains than in associations of oaks or conifers at the higher eleva- tions. The gradual reduction in number in Coahuila has been the result of both hunting and competition with livestock, chiefly goats. Most of the big- horns living in Coahuila now are in areas either where few people live or where land operators are giving the animals protection from hunting. The bighorn sheep is known with certainty, to occur in only a few places in the State. One is the mountainous area to the north- westward of San Lazaro. This place is situated in a gap which Fig. 76. Distribution of the bighorn sheep, Ovis canadensis mexicana, in Coahuila. Fomier distribution in par- allel lines. Present distribution in cross-hatched lines. 328 University of Kansas Publs., Mus. Nat. Hist. cuts through an east-west range of rugged, rimrock mountains. The mountains are of massive hmestone and estimated to be less than 7000 feet in maximum elevation; some oak, juniper and piiion grow at higher elevations. To the northwestsvard of San Lazaro these mountains are known locally as Sierra Palo Verde (Sierra de Santa Rosaha on Abbott's map, 1905). According to Dr. Mar- celino Garza-Peiia of Saltillo, bighorns occur in these mountains and as far northwest as the Sierra de San Marcos. The foreman of the Rancho San Lazaro told me that several bighorns have been seen in recent years; he saw a large ram in August, 1952, on the rimrock directly west of the Puerto San Lazaro. He thinks that bighorns are increasing, because the species is getting maximum protection from ranchers. The mounted head of a ram, killed in 1934 on Picacho de San Antonio, approximately 5 miles east of San Lazaro, by Sefior Santiago Gonzales-Guajardo, is in the home of Sefior Marcelino Garza in Saltillo. Another head, taken in the same area in 1915, is reported to be in the possession of Sefior Fermin Soto in Monclova. Another place where bighorns have been reported in recent years is in north-central Goahuila. Ernest Marsh (1937:42-43) cites records of bighorn from the Sierra del Carmen. Harvey Pauley reported (in Marsh, 1937:42) that one bighorn was killed in a cafion west of the Hacienda La Encantada in 1932. Further reason to think that bighorns did occur in this general area at least until 20 years ago, is that Borell and Bryant (1942:44) report a set of horns belonging to J. O. Langford of Hot Springs, Texas, which ". . . were picked up in the Music Mountains of Mexico, about 40 miles south of Hot Springs, in the fall of 1935." I know of no such locality in northern Goahuila but suppose that this name refers to some low desert range to the west of the Sierra de la En- cantada. Information obtained since 1949 indicates that no big- horn now live in the Sierra del Garmen and associated mountains. A third area where bighorn sheep may occur is in the Sierra de los Hecheros (Sierra de los Hechiceros on Abbott's map, 1905) in extreme northwestern Goahuila. In 1953, Sefior Juan Jose Zapata of the Hacienda Rincon reported that a band of bighorns lives in that part of the Hecheros which are in Ghihuahua. If bighorns do live there these animals may range also into Goahuila. Bighorn sheep may still Hve in mountainous country in the vi- cinity of Sierra Mojada. Mr. Golin V. McLaren of Nueva Rosita told me that in 1929 he was a member of a party which hunted mountain sheep in the Sierra de Aparajo, a low desert range south Mammals of Coahuila, Mexico 329 of the Sierra del Pino. One sheep was killed by the party and a band containing several sheep was seen. Both Mr. McLaren and Mr. Earl C. Johnston of Cuatro Cienegas believe that bighorn still occur in the desert ranges south of the Sierra del Pino, although Seiior Mario Villeareal of Americanos told Robert W. Dickerman in May, 1954, that it was doubtful that bighorn were still present. Villeareal said that bighorn occurred formerly, and possibly at present, in the Sierra del Rey, south and east of Sierra Mojada. Mr. Johnston told me that he had heard that one ram was killed south of the Sierra del Pino in 1952. He said that these mountains are well-kno^\^l as bighorn country to hunters, who have come from distant places hoping to obtain trophies. Reliable reports from several sources at Sierra Mojada and at Jaco and Camargo, in Chihuahua, tell of a sizeable band of sheep on the Rancho La Ventura, in the Sierra de Diablo, approximately 25 miles west of Sierra Mojada in Chihuahua. Bighorn sheep might range into Coa- huila from this area. No specimens from Coahuila are known to be preserved in mu- seums. A mounted head of a ram taken in 1934 approximately 5 miles east of San Lazaro and in the possession of Sefior Marcelino Garza of Saltillo has been examined. Bone fragments, identified by Gilmore (1947:160) as O. canadensis, were found in cave deposits near Cuatro Cienegas. LITERATURE CITED Allen, J. A. 1881. List of mammals collected by Dr. Edward Palmer in northeastern Mexico, with field-notes by tlie collector. Bull. Mus. Comp. Zool., 8:183-189, March. 1901. 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Amer, Fauna, 29:1-314, 13 pis., 19 figs., August 31, 1925. Status of the pronghomed antelope, 1922-1924. U. S. Dept. Agric, Dept. Bull. 1346:1-64, 6 pis., 21 figs., August. Osgood, W. H. 1900. Revision of the pocket mice of the genus Perognathus. N. Amer. Fauna, 18:1-72, 4 pis., 15 figs., September 20, 1909. Revision of the mice of the American genus Peromyscus. N. Amer. Fauna, 28:1-285, 8 pis., 12 figs., April 17. Rainey, D. G. and Baker, R. H, 1955. Distribution and systematic position of the pigmy wood rat, Neo- toma goldmani. Univ. Kansas Publ,, Mus. Nat. Hist., 7:619-624. 2 figs, in text, June 10. Reed, E. K. 1952. The myth of Montezuma's bison and the type locahty of the species. Jour. Mamm., 33:390-392, August 19, Rehn, J. A, G, 1902. A revision of the genus Mormoops. Proc. Acad. Nat, Sci. Phila- delphia, 54:160-172, June 11, Russell, R. J. 1954. A multiple catch of Cratogeomys. Jour. Mamm., 35:121-122, Feb- ruary 10. Russell, R. J. and Baker, R. H. 1955. Geographic variation in the pocket gopher, Cratogeomys castanops, in Coahuila, Mexico. Univ. Kansas Publ., Mus. Nat. Hist., 7:591- 608, 1 fig., March 15. 334 University of Kansas Publs., Mus. Nat. Hist. Sanborn, C. C. 1932. The bats of the genus Eumops. Jour. Mamm., 13:347-357, No- vember 2. SCHANTZ, V. S. 1949. Three new races of badgers (Taxidea) from southwestern United States. Jour. Mamm., 30:301-305, August 17. Seton, E. T. 1929. Lives of game animals. Doubleday, Doran and Co., Inc., New York City, vol. 3, pt. 2:413-780, pis. 61-90. Setzer, H. W. 1949. Subspeciation in the kangaroo rat Dipodomys ordii. Univ. Kansas Publ., Mus. Nat. Hist., 1:473-573, 27 figs, in text, December 27. Simpson, G. G. 1953. Evolution and geography. An essay on historical biography with special reference to mammals. Condon Lectures, Oregon State System Higher Ed., Eugene, 64 pp., 30 figs., December. Stains, H. J. and Baker, R. H. 1954. Deciduous teeth in the hognose bat, Choeronycteris mexicana. Jour. Mamm., 35:437-438, August 20. Taber, F. W. 1939. Extension of the range of the armadillo. Jour. Mamm., 20:489- 493, 2 figs., November 14. Taylor, W. P. and Davis, W. B. 1947. The mammals of Texas. Texas Game, Fish and Oyster Commis- sion, Bull. 27:1-79, illus., August. Taylor, W. P., McDougall, W. B., Presnall, C. C, and Schmidt, K. P. 1945. Preliminary ecological survey of the northern Sierra del Carmen, Coahuila, Mexico. April 1-10, 1945. (mimeo.), 48 pp., 16 figs., 1 map. 1946. The Sierra del Carmen in northern Coahuila. Texas Geogr. Mag., pp. 11-22, 5 figs., 1 map, Spring. Thomas, O. 1911. The mammals of the Tenth Edition of Linnaeus; an attempt to fix the types of the genera and exact bases and localities of the species. Proc. Zool. Soc. London, pp. 120-158, March 22. Villa R., B. 1950. Los venados en Mexico. Dir. Gen. Forestal y de Gaza, Dept. Gaza, Bol. 1:1-32, 3 unnumbered figs., 2 maps. 1951. Jabahes y berrendos. Dir. Gen. Forestal y de Gaza, Dept. Gaza, Bol. 2:1-30, 3 figs., 2 maps. 1954. Distributi6n actual de los castores en Mexico. Anales Inst. Biol., 25:443-450, 2 pis., 1 map. Webb, W. P. 1931. The Great Plains. Ginn and Company, Boston, xv -f- 525 pp., illustrated with unnumbered pis. and maps. White, J. A. 1953. The baculum of the chipmunks of western North America. Univ. Kansas Publ., Mus. Nat. Hist., 5:611-631, 19 figs, in text, Decem- ber 1. Mammals of Coahuila, Mexico 335 Young, S. P. and Goldman, E. A. 1944. The wolves of North America. Amer. Wld. Inst., Washington, D. C, XX + 636 pp., 131 pis., 15 figs., May 29. 1946. The puma. Mysterious American cat. Amer. Wld. Inst., Washing- ton, D. C, xiv + 358 pp., 93 pis., 6 figs., November 16. Young, S. P. and Jackson, H. H. T. 1951. The clever coyote. Wildhfe Mgt. Inst., Washington, D. C, xv + 411 pp., 81 pis., 28 figs., November 29. Transmitted September 21, 1955. D 26-1204 (Continued from inside of front cover) 17. The subspecies of the Mexican red-bellied squirrel, Sciurus aureogaster. By Keith R. Kelson. Pp. 243-250, 1 figure in text. April 10, 1952. 18. Geographic range of Peromyscus melanophrys, with description of new sub- species. By Rollin H. BaVer. Pp. 251-258, 1 figure in text. May 10, 1952. 19. A new chipmunk (Genus Eutamias) from the Black Hills. By John A. White. Pp. 259-262. April 10. 1952. 20. A new pinon mouse (Peromyscus truei) from Diuango, Mexico. By Robert B. Finley, Jr. Pp. 263-267. May 23, 1952. 21. An annotated checklist of Nebraskan bats. By Olin L. Webb and J. Knox Jones, Jr. Pp. 269-279. May 31. 1952. 22. Geographic variation in red-backed mice (Genus Clethrionomys ) of the south- ern Rocky Mountain region. By E. Lendell Cockrum and Kenneth L. Fitch. Pp. 281-292, 1 figure in text. November 15, 1952. 23. Comments on the taxonomy and geographic distribution of North American microtines. By E. Raymond Hall and E. Lendell Cockrum. Pp. 293-312. November 17, 1952. 24. The subspecific status of two Central American sloths. By E. Raymond HaU and Keith R. Kelson. Pp. 313-317. November 21, 1952. 25. Comments on the taxonomy and geographic distribution of some North Ameri- can marsupials, inseclivores, and carnivores. By E. Raymond Hall and Keith R. Kelson. Pp. 319-341. December 5, 1952. 26. Comments on the taxonomy and geographic distribution of some North Ameri- can rodents. By E. Raymond Hall and Keith R. Kelson. Pp. 343-371. December 15. 1952. 27. A synopsis of the North American microtine rodents. By E. Raymond Hall and E. Lendell Cockrum. Pp. 373-498, 149 figures in text. January 15, 1953. 28. The pocket gophers (Genus Thomomys) of Coahu:la, Mexico. By Rollin H. Baker. Pp. 499-514, 1 figure m text. June 1, 1953. 29. Geographic distribution of the pocket mouse, Perognathus fasciatus. By J. Knox Jones, Jr. Pp. 515-526, 7 figures in text. August 1, 1953. 30. A new subspecies of wood rat ( Neotoma mexicana ) from Colorado. By Robert B. Finley, Jr. Pp. 527-534, 2 figures in text. August 15. 1953. 31. Four new pocket gophers of the genus Cratogeomys from Jalisco, Mexico. By Robert J. RusseU. Pp. 535-542. October 15, 1953. 32. Genera and subgenera of chipmunks. By John A. ^^Vhite. Pp. 543-561, 12 figures in text. December 1, 1953. '- '> ^ 33. Taxonomy of the chipmunks, Eutamias quadrivittatus ' and Eutamias um- brinus. By John A. White. Pp. 563-582, 6 figures in text. December 1, 1953. 34. Geographic distribution and taxonomy of the chipmunks of Wyoming. By John A. WTiite. Pp. 584-610, 3 figmres in text. December 1, 1953. 35. The baculum of the chipmunks of western North America. By John A. White. Pp. 611-631, 19 figures in text. December 1, 1953. 36. Pleistocene Soricidae from San Josecito Cave, Nuevo Leon, Mexico. By James S. Findley. Pp. 633-639. December 1, 1953. 37. Seventeen species of bats recorded from Barro Colorado Island, Panama Canal Zone. By E. Raymond Hall and William B. Jackson. Pp. 641-646. De- cember 1. 1953. Index. Pp. 647-678. 'Vol. 6. (Complete) Mammals of Utah, taxonomt/ and distribution. By Stephen D. Dur- rant. Pp. 1-549, 91 figures in text, 30 tables. August 10, 1952. Vol. 7. *1. Mammals of Kansas. By E. Lendell Cockrum. Pp. 1-303, 73 figures in text, 37 tables. August 25, 1952. 2. Ecology of the opossum on a natural area in northeastern Kansas. By Henry S. Fitch and Lewis L. Sandidge. Pp. 305-338, 5 figures in text. August 24. 1953. 3. The silky pocket mice (Perognathus flavus) of Mexico. By Rollin H. Baker. Pp. 339-347, 1 figure in text. February 15, 1954. 4. North American jumping mice (Genus Zapus). By Philip H. Krutzsch. Pp. 349-472, 47 figures in text, 4 tables. April 21, 1954. 5. Mammals from Southeastern Alaska. By Rollin H. Baker and James S. Find- ley. Pp. 473-477. April 21, 1954. 6. Distribution of some Nebraskan Mammals. By J. Knox Jones, Jr. Pp. 479- 487. April 21, 1954. 7. Subspeciation in the montane meadow mouse, Microtus montanus, in Wyoming and Colorado. By Sydney Anderson. Pp. 489-506, 2 figures in te.xt. July 23. 1954. (Continued on outside of back cover) (Continued from inside of back cover) 8. A new subspeciej of bat (Myotis velifer) from southeastern California and Arizona. By Terry A. Vaughn. Pp. 507-512. July 23, 1954. 9. Mammals of the San Gabriel Mountains of CaUfomia. By Terry A. Vaughn. Pp. 513-582, 1 figure in text, 12 tables. November 15, 1954. 10. A new bat (Genus Pipistrellus ) from Northeastern Mexico. By Rollin H Baker. Pp. 583-586. November 15, 1934. 11. A new subspecies of pocket mouse from Kansas. By E. Raymond Hall. Pp. 587-590. November 15, 1954. 12. Geographic variation in the pocket gopher, Cratogeomvs castanops, in Coa- huila, Mexico. By Robert J. Russell and Rollin H. Baker. Pp. 591-608. March 15, 1955. 13. A new cottontail (Sylvilagus floridanus) from northeastern Mexico. By Rollin H. Baker. Pp. 609-612. April 8, 1955. 14. Taxonomy and distribution of some American shrews. By James S. Findley Pp. 613-618. June 10, 1955. 15. The pigmy woodrat, Neotoma goldmani, its distribution and systematic posi- tion. By Dennis G. Rainey and Rollin H. Baker. Pp. 619-624, 2 figs, in text. June 10, 1955. Index. Pp. 625-651. Vol. 8. 1. Life history and ecology of the five-lined slunk, Eumeces fasciatus. By Henry S. Fitch. Pp. 1-156, 26 figures in text. September 1, 1954. 2. Myology and serology of the avian family Fringillidae, a taxonomic study. By William B. Stallcup. Pp. 157-211, 23 figures in text, 4 tables. November 15, 1954. 3. An ecological study of the collared lizard (Crotaphytus collaris). By Henry S. Fitch. Pp. 213-274, 10 figures in text. February 10, 1956. 4. A field study of the Kansas ant-eating frog, Gastrophryne obvacea. By Henry S. Fitch. Pp. 275-306, 9 figures in text. February 10, 1956. 5. Check-list of the birds of Kansas. By Harrison B. TordofiF. Pp. 307-359, 1 figure in text. March 10, 1956. 6. A population study of the prairie vole (Microtus ochrogaster) in Northeastern Kansas. By Edwin P. Martin. Pp. 361-416, 19 figures in text. April 2, 1956. 7. Temperature responses in free-living amphibians and reptiles of northeastern Kansas. By Henry S. Fitch. Pp. 417-476, 10 figures in text, 6 tables. June 1, 1956. 8. Food of the crow, Corvus brachyrhynchos Brehm, in south-central Kansas. By Dwight Piatt. Pp. 477-498, 4 tables. June 8, 1956. More numbers will appear in volume 8. Vol. 9. 1. Speciation of the wandering shrew. By James S. Findley. Pp. 1-68, 18 figures in text. December 10, 1955. 2. Additional records and extensions of ranges of mammals from Utah. By Stephen D. Durrant, M. Raymond Lee, and Richard M. Hansen. Pp. 69-80. December 10, 1955. 3. A new long-eared myotis (Myotis evotis) from northeastern Mexico. By Rollin H. Baker and Howard J. Stains. Pp. 81-84. December 10, 1955. 4. Subspeciation in the meadow mouse, Microtus pennsvlvanicus, in Wyoming. By Sydney Anderson. Pp. 85-104, 2 figures in text. 'May 10, 1956. 5. The condylarth genus Ellipsodon. By Robert W. Wilson. Pp. 105-116, 6 figures in text. May 19, 1956. ^ 6. 1 Additional remains of the multituberculate genus Eucosmodon. By Robert W. Wilson. Pp. 117-123, lO figures in text. May 19, 1956. 7. Mammals of Coahuila, Mexico. By Rollin H. Baker. Pp. 125-335, 75 figures in text. June 15, 1956. More numbers wUl appear in volume 9. 'A- ujj^^'^^^^^'ji LIBRAitV OCT 1 1955 _J1AMJ?0 wmw, University of Kansas Publications Museum of Natural History Volume 9, No. 8, pp. 337-346, 1 fig. in text, 1 table August 15, 1956 Comments on the Taxonomic Status of Apodemus peninsulae, with Description of a New Subspecies from North China BY J. KNOX JONES, JR. University of Kansas Lawrence 1956 University of Kansas Publications, Museum of Natural History Editors: E. Raymond Hall, Chairman, A. Byron Leonard, Robert W. Wilson Volume 9, No. 8, pp. 337-346, 1 fig. in text, 1 table Published August 15, 1956 University of Kansas Lawrence, Kansas PRINTED BY FERO VOILAND. JR.. STATE PRINTER TOPEKA. KANSAS 1956 26-3854 niw«. uumr. LliUL I I LIBRARY OCTl 1956 HARVARD Comments on the Taxonomic --StatiAreJ — Apodemus peninsulae, with Description of a New Subspecies from North China BY J, KNOX JONES, JR. In the past several years the United States National Museum has received a large number of mammals from central and southern Korea through the auspices of the Commission on Hemorrhagic Fever of the Armed Forces Epidemiological Board. Among these Korean collections are more than a hundred specimens of a murine rodent originally described as "Micromys speciosus peninsulae" by Oldfield Thomas but currently placed in the genus Apodemus. In attempting to ascertain the specific relationships of this mouse I have examined, through the generosity of Dr. David H. Johnson, Acting Curator of Mammals, most of the other Oriental specimens of the subgenus Sijlvaemus in the U. S. National Museum and it is on this combined material that the following comments and de- scription are based. Three general groups of the genus Apodemus are presently known to occur on the mainland of northeast Asia. One is the distinctive Apodemus agrarius, lone representative of the subgenus Apodemus. The others, both in the subgenus Sylvaemus and closely resembling each other, are represented by a small animal that is currently re- garded as conspecific with Apodemus sylvaticus and a larger animal of which the Korean mouse, peninsulae, is representative. The oldest trivial name applied to the large Sylvaemus is major of Radde, 1862, in the combination [Mus sylvaticus] vrt. major. This is, however, twice preoccupied (see Ellerman and Morrison-Scott, 1951:566). The next available name is peninsulae of Thomas, 1907, which was applied to mice from central and southern Korea (type from Mun'gyong, 110 mi. SE Seoul, Korea), and was originally pro- posed as a subspecies of the insular Japanese species, Apodemus speciosus. G. M. Allen (1940:949), who recognized peninsulae as a monotypic species, was the first investigator to make the important distinction that it was not conspecific with the Japanese speciosus, although HolHster (1913:1-2) and Miller (1914:89) had previously used the combination Apodemus peninsulae, evidently with the same thought in mind. (339) 340 University of Kansas Publs., Mus. Nat. Hist. More recently, Ellerman (1949:32) and Ellerman and Morrison- Scott (1951:566) have arranged peninsulae as a subspecies of Apodemiis flavicollis under the assumption that all the members of the subgenus Sylvaemns on the eastern Asiatic mainland are sub- species of one or another of the species of western Europe, A. -ftavi- collis or A. sylvaticus. Ellerman (in Ellerman and Morrison-Scott, 1951:564) states: "The majority of the forms I distribute in a some- what arbitrary manner between sylvaticus, average smaller skull, and flavicollis, average larger skull; occurring together nearly throughout the Palaearctic. I feel fairly sure that there are some errors of judgment in my arrangement, and equally sure that there is no other way to define species in this very large and difficult group." I have compared the specimens of peninsulae available to me from central and southern Korea with specimens of A. /. flavi- collis from Denmark, Germany and Sweden and find, although the Fig. 1. Ventral views of skulls and left maxillary tootli-rows of two species of the genus Apodemus. a. Apodemus -flavicollis flavicollis (Melcliior), Lolland, Denmark, adult $, No. 141691 USNM, x 2. b. Apodemus flavicollis flavicollis (Melchior), Mauseklippe, Germany, young $, No. 112895 USNM, x 10. c. Apodemus peninsulae peninsulae (Thomas), Central Nat'l Forest, near Pup'yong-ni, 200 m., Korea, subadult 9 , No. 300650 USNM, X 10. ^ d. Apodemus peninsulae peninsulae (Thomas), 6 mi. S Yongdongp'o, Korea, adult $ , No. 299554 USNM, x 2. In comparing the ventral views of skulls note especially the size and loca- tion of incisive foramina and posterior palatine foramina as well as the breadth of mesopterygoid fossae. In comparing the left maxillary tooth-rows note es- pecially the size of M3 and the reduced posterointernal cusp on Ml in A. peninsulae. Taxonomic Status of Apodemus peninsulae 341 two are similar in many ways, that peninsulae differs from flavicollis in several important characters: Mammae 1-2 == 6 in flavicollis, and 2-2 := 8 in peninsulae; incisive foramina reaching level of alveoli of Ml, or nearly so, in flavicollis, but ending conspicuously short of that level in peiiinsulae; posterior palatine foramina large in flavi- collis and opposite a point where Ml and M2 meet, but small in peninsulae and situated farther back on the palate, opposite M2. Moreover, peninsulae lacks the characteristic buffy throat patch of flavicollis, has a much reduced posterointernal cusp on the Ml, a relatively (frequently actually) larger M3 and, on the average, a broader mesopterygoid fossa. In view of these differences, all of which appear to be constant, I consider peninsulae specifically dis- tinct from flavicollis. Throughout its known geographic range ( see below) peninsulae is evidently confined to wooded terrain, either scrub or brush types or forested areas, and the vernacular name wood mouse, therefore, seems appropriate for this species. The type specimens of Apodemus praetor Miller (type from Sungari River, 60 mi. SW Kirin, Manchuria) and Apodemus nigri- talus Hollister ( type from Tapucha, Altai Mountains, Siberia ) agree with peninsulae as concerns the above characters and differ from it only in minor external and cranial features. They are, therefore, here considered as subspecies of the latter. Ellerman (1949:32) and Ellerman and Morrison-Scott (1951: 567) regarded nigritalus, like peninsulae, as a subspecies of flavi- collis. The subspecies praetor, on the other hand, has generally been regarded as a synonym of peninsulae by recent authors. Howell (1929:58) noted that the holotype was, "... a phe- nomenally large specimen such as is encountered occasionally in almost all groups of rodents." He ascribed the color differences noted by Miller to "seasonal" variation. The holotype of praetor is undeniably larger than the other adult specimens listed in the original description. These paratypes and other specimens of praetor available to me are approximately the same size externally and average only slightly larger cranially than specimens of penin- sulae from central and southern Korea. However, the dorsal colora- tion of praetor is somewhat darker and duller than that of penin- sulae, especially in summer pelage when praetor lacks the conspic- uous bright ochraceous tinge of the Korean specimens. In addition, praetor has broader zygomatic plates with correspondingly deeper zygomatic notches and the color on the face of the upper incisors averages much more orange than in peninsulae. 342 University of Kansas Publs., Mus. Nat. Hist. In the north then, wood mice range from Korea and Manchuria westward at least as far as the Altai Mountains. For mice from the intervening Siberian areas Russian workers have used the name major which, as noted above, is unavailable. The exact relation- ships of the mice of these areas to previously named subspecies is unknown to me and I have not seen specimens of "Mus (Alsomys) major niftdus" of Dukelsky, 1928, the type locality of which is 75 versts (approximately 50 miles) SE Vladivostok, Siberia. It ap- pears to be of the same species as peninsitlae and judging from the original description it closely resembles praetor. Neither have I seen specimens of the Sakhalin Island mouse, giliacus, which Eller- man (1949:32) regards as a subspecies of Apodemus sylvaticus. I feel reasonably sure, however, that it will prove to be a subspecies of peninsulae. In the original description giliacus was referred to as, "Most closely allied to the Korean subspecies . . ." (Thomas, 1907:411). In China the extent of the distribution of Apodemus peninsulae is also uncertain. Allen (1940:949-50) reported its occurrence from Jehol and Hopeh in the northeast, southwestward through Shansi, Shensi and eastern Kansu to Szechuan and northwestern Yunnan. Throughout most of this region it occurs with another mouse, cur- rently regarded as conspecific with Apodemus sylvaticus, and tlie two kinds have been confused by some previous authors. Howell (1929:58), for instance, reported twelve specimens of peninsulae from 65-75 mi. NE Peking but my examination of these mice indi- cates that only four are peninsulae while the others are referrable to what is currently regarded as Apodemus sylvaticus draco. Another subspecies of sylvaticus, A. s. orestes, occurs in Szechuan and Yunnan and it is certain that some records of distribution as- cribed to peninsulae from those provinces actually represent orestes (see Allen, 1940:949-50). A. sylvaticus is distinguishable from peninsulae by darker ears, blackish preauricular patches, dark eye rings, a noticeably smaller skull, incisive foramina that reach the level of Ml (or nearly so), much larger auditory bullae, and a more fully developed posterointernal cusp on Ml. Too, sylvaticus typi- cally has 1-2 = 6 mammae although Allen reports finding a 2-2 = 8 formula in some specimens. Apodemus latronum, regarded as a full species by Osgood (1932:318) and G. M. Allen (1940:950) but as a subspecies of flavicollis by Ellerman (1949:32) and Eller- man and Morrison-Scott (1951:567), also occurs in Szechuan and Yunnan. Its relatively dark color, large feet and large ears, flavi- Taxonomic Status of Apodemus peninsxilae 343 coUis-like skull and large molar teeth immediately separate it from peninsulae although the two possibly have been confused in the earlier literature. Until a complete revisionary study of the Asiatic members of the subgenus Sylvaemus can be undertaken the pres- ence of peninsulae in southwestern China must remain in question. The western limits of the geographic range of Apodemus penin- sulae are unknown. Apodemus gurkha Thomas, 1924, from Nepal is said to have 2-2 — 8 mammae but the description is not otherwise suggestive of close relationship to peninsulae. Farther to the west, Apodemus jlavicollis nisiges Miller, 1913, from Kashmir seems to have been properly assigned as a subspecies of flavicollis ( cotypes and large series in USNM). Wood mice almost certainly do not occur in the Gobi Desert. They are known as far west as the Altai Mountains to the north of the Gobi and at least as far west as Kansu (see below) to the south of it. Whether the geographic range of the species skirts the western edge of the arid regions of northern China is at present unknown; perhaps it does not. At any rate, mice available to me from the North Chinese provinces of Jehol, Shansi, Shensi and Kansu are notably different in certain external and cranial features from other known races of Apodemus peninsulae and are here given subspecific recognition. All measurements are in millimeters. Cap- italized color terms are from Ridgway (1912). Apodemus peninsulae sowerbyi, new subspecies Type. — Adult female molting from winter to summer pelage, skin and skull, U. S. National Museum no. 175523, from 30 miles west of Kuei-hua-cheng, 7000 ft., northern Shansi, China; obtained on 23 May 1912 by Arthur de Carle Sowerby, original no. 456. Distribution. — Known presently from eastern Kansu eastward through Shensi, Shansi and Hopeh to southern Jehol, probably also in northeastern Szechuan, exact limits of range unknown. Diagnosis. — Size small for species (see measurements). Color: Upper parts (fresh summer pelage) averaging near (15'a) Ochraceous-BufiF, suffused with blackish (especially mid-dorsally ) ; winter pelage much paler; underparts grayish- white, individual hairs plumbeous at base, tipped with white; ears pale brownish; feet whitish above, darker below; tail bicolor, pale brownish above, whitish below. Skull: Small (see measurements); rostrum somewhat shortened and conspicuously down-curved; zygomatic notches relatively shallow; zygo- matic plates narrow; braincase proportionally more inflated tlian in other sub- species of the species; auditory bullae moderately inflated; upper incisors slen- der, their faces averaging bright yellowish-orange. Measurements. — External measurements of the holotype, followed by those of an adult male and female from the type locality, are, respectively: Length 344 University of Kansas Publs., Mus. Nat. Hist. of head and body, 101, 102, 100; length of tail, 93, 102; length of hind foot (su), 21, 21.5, 23; length of ear from notch, 14, 16, 15.5. Corresponding measurements for an adult female from 20 mi. E Taiyuan, Shansi, are: 91, 99, 23, 16. For cranial measurements see Table 1. Comparisons. — From Apodetnus peninsiilae peninsiilae (speci- mens from various localities in central Korea), A. p. sowerbiji dif- fers in: External size smaller throughout, especially hind foot; upper parts, especially in summer pelage, and dorsal aspect of tail paler; Table 1. — Cranial Measurements of Adults of Several Subspecies of ApODEMUS PENDSrSULAE ^ ;ti TS ^ « bH "--c Q ai -1.2 > o CO E ■♦J p "^ ^ ^ Q Apodermis peninsidae peninsiilae, various localities in central Korea Average 10 (46^,6 9)-. Minimum Maximum 29.2 14.2 11.8 4.7 20.1 11.4 10.2 28.3 13.8 11.5 4.6 19.2 10.8 9.9 29.8 14.6 12.2 5.1 20.7 12.0 10.5 4.3 4.1 4.4 Apodemus peninsulae nigritalus, Tapucha, Altai Mts., Siberia USNM 175164, d^ (type) USNM 175171, 9 28.8 28.2 14.8 13.7 12.4 11.8 4.5 4.5 20.8 19.8 11.7 11.2 11.0 10.3 4.4 4.5 Apodemus peninsiilae praetor, Sungari River, GO mi. SW Kirin, Manchuria USNM 197792, cf(tvpe) USNM 197798, 9 30.5 30.2 14.4 12.5 11.8 4,7 4.6 21.5 21.6 12.5 12.7 10.3 10.6 4.6 4.6 Mukden, Manchuria USNM 197782,cf | 29.5 | 14.8 1 12.4 | 4.8 20.6 I 12.2 I 10.5 I 4.2 19.6 11.4 9.9 4.0 18.9 11.4 9.7 4.1 19.4 11.3 9.8 4.2 Apodemus peninsulae soiverbyi, Kuei-hau-cheng, Shansi USNM 175523, 9 (type) 27.9 13.3 11.7 4.5 USNM 175521, c^ 27.6 11.5 4.6 USNM 175522, 9 27.9 11.8 4.6 20 mi. E Taiyuan, Shansi USNM 172558,9 1 27.4 | 13.8 | 11.5 | 4.6 | 19.4 | 11.6 | 10.1 | 4.4 12 mi. S Yenan, Shensi USNM 155072, cf . USNM 155073, 9 . USNM 155075, d^ . 27.8 27.7 27.9 14.1 13.3 13.5 11.5 11.4 4.4 4.5 4.5 19.5 19.4 19.2 11.0 11.0 io.o 11.0 10.0 4.3 4.2 4.3 Hsin-lung-shan, 65 mi. NE Peking, Jehol USNM 219229,0^ ] 27.7 | 13.8 ] 11.4 | 4.5| 19.0 1 10.9 | 10.4 | 4.4 15 mi. S Lanchow, Kansu USNM 155171, d' I 27.7 I 13.6 I 11.7 I 4.6 | 19.0 | 11.3 | 9.9 | 4.5 Taxonomic Status of Apodemus peninsulae 345 skull smaller and less massive; braincase proportionally more in- flated; rostrum shorter and noticeably down-curved. From Apo- demus peninsulae praetor of Manchuria (holotype and paratypes), A. p. sowerbiji differs in most of the same ways in which it does from peninsulae as well as in having more shallow zygomatic notches, narrower zygomatic plates and smaller, more slender, upper incisors. From Apodemus peninsulae nigritalus of the Altai Moun- tains of Siberia (holotype and paratypes), A. p. sowerbiji differs in: Smaller size, both external and cranial; paler dorsal coloration; less convex cranial outline in lateral view; smaller auditory bullae. Remarks. — Apodemus peninsulae sowerbyi is named in honor of the late Artliur de Carle Sowerby whose collections of mammals from North China and Manchuria have added so much to our meager knowledge of that part of the world. Four specimens from Hsin-lung-shan, 65 mi. NE Peking, here assigned to sowerbyi, are darker dorsally than mice from farther to the west and in this respect may show approach to A. p. praetor. In all other features, however, they closely resemble the new sub- species. All of the specimens of sowerbyi available to me are from alti- tudes of 3000 feet or higher. At lower elevations in North China, destruction of wooded habitats owing to intense land-use practices has probably restricted the distribution of sowerbyi primarily to hilly and mountainous areas where brushy, scrub and forest habitats still prevail. Specimens examined. — Thirty-three, all from North China, as follows: JEHOL: Hsin-lung-shan, 65 mi. NE Peking, 3000 ft., 4. KANSU: 15 mi. S Lanchow, 7400 ft., 1. SHANSI: Chiao-cheng-shan, 90 mi. W Taiyuan, 7000- 8000 ft., 4; 30 mi. W Kuei-hau-cheng, 7000 ft., 5; Lung-wang-shan, 20 mi. E Taiyuan, 4000 ft., 10; 18 mi. W Taiyuan, 5000 ft., 1; 50 mi. NW Taiyuan, 5500 ft., 4. SHENSI: 12 mi. S Yenan, 4000 ft., 4. Apodemus peninsulae, then, is known or suspected to occur over much of southeastern Siberia, Manchuria, Korea and North China. The western limits of its geographic range are unknown. Over this vast area only four subspecies, one newly named, can be as- cribed with certainty to peninsulae whereas only two other kinds, giliacus of Thomas from Sakhalin and rufidus of Dukelsky from ex- treme southeastern Siberia are probably conspecific with it, the latter possibly a synonym of praetor. These considerations under- score the preliminary nature of the present paper. The mammalian fauna of northeastern Asia is scarcely better known today tlian was 346 University of Kansas Publs., Mus. Nat. Hist. that of North America in 1885 when Dr. C. Hart Merriam organized what was later to become the U. S. Biological Survey. It seems to me that the correct names of four kinds of wood mice discussed above are as follows: Apodemus peninsulae peninsulae (Thomas, 1907) Apodemus peninsulae nigritalus Hollister, 1913 Apodemus peninsulae praetor Miller, 1914 Apodemus peninsulae sowerbyi Jones, 1956 LITERATURE CITED Allen, G. M. 1940. The mammals of China and Mongolia. Amer. Mus. Nat. Hist., New York, 2:XXVI + 621-1350, September 3. Ellerman, J. R. 1949. The families and genera of living rodents. British Mus., London, 3:V + 1-210, March. Ellerman, J. R., and T. C. S. Morrison-Scott. 1951. Checkhst of Palaearctic and Indian mammals, 1758 to 1946. British Mus., London, 810 p., November 19. Hollister, N. 1913. Two new mammals from the Siberian Altai. Smith. Misc. Coll., 60:1-3, March 13. Howell, A. B. 1929. Mammals from China in the collections of the United States Na- tional Museum. Proc. U. S. Nat. Mus., 75:1-82, June 7. Miller, G. S., Jr. 1914. Two new murine rodents from eastern Asia. Proc. Biol. Soc. Wash- ington, 27:89-92, May 11. Osgood, W. H. 1932. Mammals of the Kelley-Roosevelts and Delacour Asiatic expedi- tions. Field Columb. Mus. Nat. Hist., Zool. Series, 18:193-339, August 19. RiDGWAY, R. 1912. Color standards and color nomenclature. Washington, D. C, pub- lished by the author. Thomas, O. 1907. The Duke of Bedford's zoological explorations in eastern Asia. — IV. A list of small mammals from the islands of Saghalien and Hokkaido. Proc. Zool. Soc. London, 1907:404-414, August 1. Transmitted May 12, 1956. D 26-3854 fV • '/ University of Kansas Publications Museum of Natural History Volume 9, No. 9, pp. 347-351 August 15, 1956 Extensions of Known Ranges of Mexican Bats BY SYDNEY ANDERSON m%. COMP. ZOOL LIBRARY SEP 2 7I955 KARVARO UNI'fERSiTY University of Kansas Lavi^rence 1956 University of Kansas Publications, Museum of Natural History Editors: E. Raymond Hall, Chairman, A. Byron Leonard, Robert W. Wilson Volume 9, No. 9, pp. 347-351 Published August 15, 1956 University of Kansas Lawrence, Kansas PRINTED BY FERD VOILAND. JR.. STATE PRINTER TOPEKA. KANSAS 1956 Extensions of Known Ranges iMiis. mm, zfloi I mm SEP 2 7 1955 HARVARD Innnvnnu IJNIVERSITY I BY SYDNEY ANDERSON Incidental to studies of speciation of North American mammals, made possible by assistance from the National Science Foundation and the Kansas University Endowment Association, a number of bats have been taken beyond the limits of their previously known geographic ranges. Pending the completion of more detailed faunal accounts, these notes are published so that the distributional records will be available to interested students of Mexican mammals. Many of these bats are essentially tropical and the new records here reported, extend the known geographic ranges to the north- ward on either the east or the west coast of Mexico. Continued collecting, especially by the intensive application of a variety of methods including the use of mist nets, in the northern parts of the zone of tropical vegetation can be expected to yield other species of tropical bats beyond the limits of the ranges now known. Catalogue numbers cited in parentheses are those of the Museum of Natural History. Chilonycteris psilotis Dobson. — Six specimens (36426-36431) taken 7 mi. W and /2 mi. S Santiago, at sea level, Colima, by J. R. Alcorn, on March 17, 1950, extend the known range of this species 330 miles westward from the most northwestern recorded occur- rence at Alpuyeca, Morelos (Davis and Russell, 1952:234). Use of the name psilotis is explained by de la Torre (1955:697). Chilonycteris parnellii mexicana Miller. — One specimen (54934) from 10 mi. W, 2 mi. S Piedra, 1200 ft., Tamauhpas, taken by Gerd H. Heinrich, on June 13, 1953, extends the known range of this spe- cies 76 miles east-northeast (Goodwin, 1954:4), previously the most northern recorded occurrence in northeastern Mexico. Thirty other specimens have been taken from four additional localities between El Pachon and the place 10 mi. W, 2 mi. S Piedra. Pteronotus davyi fulvus (Thomas). — Ten specimens (57525- 57534) from Rancho Santa Rosa, 25 km. N, 13 km. W Cd. Victoria, 260 meters, Tamaulipas, taken by W. Schaldach, V. Grissino, and R. Grimsley, from December 26, 1953, to January 5, 1954, extend the known range of this species 360 miles northward from Mirador, Veracruz (Davis and Russell, 1952:235). Another specimen from (349) 350 Unwersity of Kansas Publs., Mus. Nat. Hist, Tamaulipas in our collection is from Rancho Pano Ayuctle, 8 mi. N Gomez Farias, 300 feet. Glossophaga soricina leachii (Gray). — Two specimens (54942- 54943) from 16 mi. W, 3 mi. S Piedra, Tamaulipas, taken by Gerd H. Heinrich, on June 15, 1953, extend tlie known range of this spe- cies on the Gulf Coast of Mexico northward 60 miles from 5 mi. NE Antiguo Morelos, near El Pachon, Tamaulipas (de la Torre, 1954: 114). Anoura geoffroyi lasiopyga (Peters). — Three specimens (36574- 36576) from 2 mi. SE Jalcocotan, 3000 ft., Nayarit, the first records for the state, taken by J. R. Alcorn, on February 15, 1950, extend the known range of the species 50 miles northward from San Sebastian, Jalisco (Sanborn, 1933:27). Choeronycteris mexicana Tschudi. — Three specimens (60176- 60178) from 4 km. N Joya Verde, near Huisachal, 4000 ft., Tam- auHpas, taken by W. Schaldach, on July 17, 1954, extend tlie kno^vn range of this species eastward into the state of Tamaulipas. East- ern marginal records in northeastern Mexico were 1 mi. S and 4 mi. W Bella Union, 7000 ft., Coahuila (Baker, 1956:174), and Hacienda Capulin, San Luis Potosi (Dalquest, 1953:27). Centurio senex Gray. — One specimen (54958) from 14 mi. W, 3 mi. S Piedra, Tamaulipas, extends the known range of the species 58 miles northward from Pano Ayuctle, near Gomez Farias, Tam- aulipas (de la Torre, 1954:114). Natalus mexicanus saturatus Dalquest and Hall. — Two specimens (54999-55000) from the Sierra de Tamaulipas, 14 mi. W, 3 mi. S Piedra, Tamaulipas, taken by Gerd H. Heinrich, on June 12, 1953, extend the known range of this species 61 miles northward from El Pachon, Tamaulipas (Goodwin, 1954:5). Forty-one other speci- mens have been taken from three localities between El Pachon and the new record station mentioned above. Rhogeessa parvtila tumida H. Allen. — One specimen (55192) from 4 mi. N La Pesca, Tamaulipas, taken by Gerd H. Heinrich, on May 24, 1953, extends the known range of the species on the east coast of Mexico northward 115 miles from 10 kms. WSW Ebano, Veracruz (Dalquest, 1953:58), and also northeastward 115 miles from Santa Maria, Tamaulipas (Goodwin, 1954:6). Another new marginal locality for this species, represented by 39 specimens in our collection, is 10 mi. W, 2 mi. S Piedra, 1200 ft., Tamaulipas. Baeodon alleni (Thomas). — A specimen (68773) from 6 mi. W and 2 mi. N Nejapa, Oaxaca, taken by A. A. Alcorn, on August 6, Extension of Ranges of Mexican Bats 351 1955, extends the known range of this species southeastward 100 miles from Cuicatlan, 590 meters, Oaxaca (Hall, 1955:2). Until recently this species was known from the type locality only. Lasiurus borealis ornatus Hall. — Sixteen specimens (55325-55340) from near Piedra, 1200 to 1400 ft., Tamaulipas ( 1 from 10 mi. W, 2 mi. S; 1 from 14 mi. W, 3 mi. S; and 14 from 16 mi. W, 3 mi. S Piedra) tend to fill a gap in the known distribution of this species. Previously it was known from Matamoros, Tamauhpas (Miller, 1897:108), to the north and from Bledos, San Luis Potosi (Dal- quest, 1953:61), to the south. Assignment to the subspecies L. b. ornatus is tentative and is based primarily on the scanty cover of hair toward the margin of the interfemoral membrane and scanty cover of hair on the ventral surface of the membrane along the forearm. Adequate comparative material of L. b. ornatus from southern Mexico is not available. LITERATURE CITED Baker, R. H. 1956. The Mammals of Coahuila. Univ. Kansas Pub!., Mus. Nat. Hist., 9:125-335, 75 figs, in text, June 15. Dalquest, W. W. 1953. Mammals of the Mexican State of San Luis Potosi. Louisiana State Univ. Studies, Biol. Sci. Ser., No. 1:1-233, December 28. Davis, W. B., and R. J. Russell 1952. Bats of the Mexican state of Morelos. Jour. Mamm., 33:234-239, May 16. DE LA Torre, L. 1954. Bats from southern Tamaulipas, Mexico. Jour. Mamm., 35:113- 116, May 26. 1955. Bats from Guerrero, Jalisco and Oaxaca, Mexico. Fieldiana: Zool- ogy, 37:695-703, 2 plates, June 19. Goodwin, G. G. 1954. Mammals from Mexico collected by Marian Martin for the Ameri- can Museum of Natural History. American Mus. Novit., 1689:1-16, November 12. Hall, E. R. 1955. Nuevos murcielagos para la fauna Mexicana. Acta Zool. Mexicana, l(No. 3): 1-2, September 10. Miller, G. S., Jr. 1897. Revision of the North American bats of the family Vespertilionidae. N. Amer. Fauna, 13:1-135, October 16. Sanborn, C. C. 1933. Bats of the genera Anoura and Lonchoglossa. Zool. Ser., Field Mus. Nat. Hist, XX: 23-28, December 11. Transmitted May 12, 1956. n 26-4058 University of Kansas Publications Museum of Natural History Volume 9, No. 10, pp. 353-356 January 21, 1957 WW. coMP. mi UBRARy JUN 13 19571 HAJtVARO i/NieslTY A New Bat (Genus Leptonycteris) From Coahuila BY HOWARD J. STAINS University of Kansas Lawrence 1957 University of Kansas Publications, Museum of Natural History Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Harrison B. Tordoff Volume 9, No. 10, pp. 353-356 Published January 21, 1957 University of Kansas Lawrence, Kansas PRINTED BY FERD VOILAND. JR.. STATE PRINTER TOPEKA. KANSAS I9S7 26-5800 A New Bat (Genus Leptonycteris From Coahuila BY HOWARD J. STAINS Assistant Professor, Department of Zoology, Southern Illinois University In a collection of mammals obtained in Coahuila, Mexico, there is a series of 24 long-nosed bats, Leptonycteris nivalis. These bats have a larger skull and a longer third finger than other bats of this species found to the south of Coahuila. On the basis of these dis- tinctive characters, it seems appropriate to recognize these long- nosed bats from Coahuila as belonging to a new subspecies, named and described as follows: Leptonycteris nivalis longala new subspecies Type. — Female, adult, skin and skull, No. 33087, Univ. Kansas Mus. Nat. Hist; 12 mi. S and 2 mi. E Arteaga, 7500 ft., Coahuila; 11 July 1949; obtained by W. K. Clark, original number 787. Range. — Southern Coahuila north to the Big Bend (Brewster County) of Texas. Diagnosis. — Size large (see measurements); third finger long; color pale, upperparts Hair Brown (capitalized color terms are after Ridgway, Color Standards and Color Nomenclature, Washington, D. C, 1912), underparts Smoke Gray; skull large and broad. Comparisons. — From Leptonycteris nivalis nivalis (specimens from Vera- cruz, Oa.xaca, Distrito Federal, Hidalgo, Jalisco, and Sonora), L. n. longala differs as follows: color paler, more whitish and less brownish; third finger longer (longala from Coahuila averaging 111.3 mm.; nivalis from Sonora averag- ing 91.0, from Jalisco 96.4, from Hidalgo 98.0, from Veracruz 100.0, from Distrito Federal 100.2, and from Oaxaca 98.6); skull larger, breadth of cranium greater (longala from Coahuila averaging 10.7mm.; nivalis from Sonora 9.8, from Jalisco 9.8, from Hidalgo 9.6, from Veracruz 9.9, from Distrito Federal 9.9, and from Oaxaca 9.8); mastoidal breadth greater (longala from Coahuila averaging 11.6 mm.; nivalis from Sonora 10.5, from Jalisco 10.8, from Hidalgo 10.6, from Veracruz 10.9, from Distrito Federal 10.8, and from Oaxaca 10.7); skull higher (longala from Coahuila averaging 10.0mm.; nivalis from Sonora 9.3, from Jalisco 9.2, from Hidalgo 9.2, from Veracruz 9.3, from Distrito Federal 9.3, and from Oa.xaca 9.1). The average of each dimension of longala listed above exceeds the maximum of the corresponding dimension in nivalis. Remarks. — Leptonycteris nivalis longala inhabits the northeastern end of the Mexican Plateau. Bats from Brewster County, Texas, referred to longala, average slightly larger in all measurements taken than do specimens from southern Coahuila. Specimens from Cerro Potosi, Municipio de Galeana, Nuevo Leon, also are referred to longala on the basis of the length of their third fingers. (355) 356 University of Kansas Publs., Mus. Nat. Hist. Like the specimens from Texas, these bats possess longer forearms, on the average, than do bats from Coahuila. Topotypes of L. n. nivalis from Mount Orizaba were not available, but 111 specimens referable to the subspecies nivalis were examined. These specimens were from the following places: Sonora: Y* mi. W Aduana, 1600 ft., 4 speci- mens. Jalisco: Hda. San Martin, 5000 ft., 18 mi. W Chapala, 3; 11 mi. W Chapala, 5000 ft., 1; 5 mi. W Chapala, 5000 ft., 58; 8 mi. NE Ocotlan, 5100 ft., 1. Hidalgo: 6 km. NW Tasquillo, 500 ft., 1. Veracruz: 3 km. W Boca del Rio, 25 ft., 22. Distrito Federal: Chicomostoc, Cerro Teutli, 2!-,^ mi. NNW Milpa Alta, 2620 ft., 18. Oaxaca: Cuicatlan, 600 ft., 1; 3 km. WNW Dominguillo, 730 ft., 2. All these specimens are in the Museum of Natural History at the University of Kansas. Little discernible geographic variation was found in these specimens of L. n. nivalis. No specimens could, with cer- tainty, be classed as intergrades between longala and nivalis, but it is thought that intergrades will be found in western San Luis Potosi or in Zacatecas or in both states. Dalquest (Mammals of the Mexican State of San Luis Potosi, Louisiana State Univ. Studies, Biol. Sci. Ser. No. 1, pp. 27-28, 1953) refers five specimens taken from Hda. Capulin, southeastern San Luis Potosi, to L. n. nivalis. Measurements by Dalquest are in accordance with other measurements of L. n. nivalis listed above. The name L. n. yerhahuenae Martinez and Villa, was based on specimens from Yerbabuena in the state of Guerrero. The specimens, including the holo- type, on which this name was based have been destroyed. Luis de la Torre (Fieldiana, 37:698, 1955) examined a topotype of yerhahuenae and was un- able to distinguish this specimen from a topotype of nivalis from Orizaba. Davis and Russell (Jour. Mamm., 33:236, 1952) identified as L. n. nivalis one bat from Morelos, taken approximately 32 miles NE of the type locality of yerhahuenae, noting that its third finger was much shorter than in specimens from the Big Bend of Texas. I judge L. n. yerhahuenae to be a synonym of nivalis as does de la Torre. Acknowledgment is made to Dr. W. B. Davis of the Agricultural and Me- chanical College of Texas and Mr. Colin C. Sanborn of the Chicago Natural History Museum for loan of comparative material. I am grateful also to the Kan- sas University Endowment Association and National Science Foundation for support of field work, and to Dr. RoUin H. Baker for guidance in my study. Measurements. — The following measurements in milHmeters include those of the type, and the average and extreme measurements of the type and 22 adult topotypes: total length, 79, 79 (73-86); length of hind foot, 16, 16 (14- 17); length of ear, 16, 16.5 (15-17); length of forearm, 50.0, 50.6 (47.3-55.0); greatest length of skull, 28.1, 27.5 (26.1-29.0); zygomatic breadth, 9.2, 9.6 (8.6-11.2); interorbital constriction, 4.6, 4.8 (4.1-5.4); mastoidal breadth, 11.7, 11.6 (11.0-12.1); breadth of braincase, 10.5, 10.7 (10.1-11.2); greatest height of skull, 9.8, 10.0 (9.6-10.5); alveolar length of maxillary tooth-row, 8.9, 9.1 (8.5-9.6); and length of third finger, 110.8, 111.3 (106.9-116.1). Specimens of L. n. longala examined. — Total number, 109, as follows: Texas: cave W side Emory Peak, Chisos Mts., Brewster Co., 7500 ft., 5 speci- mens (A and M College of Texas). Coahuila: 12 mi. S and 2 mi. E Arteaga, 7.500 ft., 24 (KU). Nuevo Leon: Cerro Potosi, Municipio de Galeana, 11,500 ft., 80 (Chicago Nat. Hist. Mus.). Transmitted December 5, 1955. D O -/Vf-f ^Tt ^m^ m^, 200L MAR 311958 Geographic Variation in the Pocket Gopher, Thomomys bottae, in Colorado BY PHILLIP M. YOUNGMAN University of Kansas Law^rence 1958 University of Kansas Publications, Museum of Natural History Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Robert W. Wilson Volume 9, No. 12, pp. 363-384, 7 figs, in text, 1 table Published February 21, 1958 w. ^^"p, rni Ubhhuj 3 1 1958 UHlVERSiiY University of Kansas Lawrence, Kansas PRINTED IN THE STATE PRINTING PLANT TOPEKA. KANSAS 1958 27-1765 Geographic Variation in the Pocket Gopher, Thomomys bottae, in Golorado BY PHILLIP M. YOUNGMAN INTRODUCTION Two species of pocket gophers of the genus Thomomys ( Family Geomyidae) occur in Colorado, Thomomys bottae (see fig. 1) in the low valleys in the south-central and southwestern parts of tlie state and Thomomys talpoides mainly in the mountains and high valleys. Thomomys bottae occurs primarily in the Pinon-juniper, Pon- derosa Pine, and Short Grass zones of Daubenmire (1943) but in some localities is found in the Douglas Fir Zone. Thomomys talpoides occupies primarily the Douglas Fir Zone and Engelmann Spruce-Subalpine Spruce Zone but is found also in the Pinon-juniper and Short Grass zones in some localities. The ranges of the two species do not overlap in the strict sense but interdigitate in a parapatric type of distribution. Two other pocket gophers, Geomys bursariiis and Cratogeomys castanops, also occur in Colorado — in the Upper Sonoran Life-Zone. Geomys bursarius occupies much of the Great Plains, whereas Cratogeomys castanops is found only on the plains in the south- eastern part of the state. The objectives of the study, reported on here, were to learn the geographic distribution of Thomomys bottae in Colorado, to find means for recognizing the different subspecies, and to describe indi- vidual and geographic variation. I am indebted to Mr. Sydney Anderson and Professor E. Raymond Hall for many helpful suggestions and for their critical reading of the manuscript, to Dr. Richard S. Miller, who made the collection of many of the specimens possible, and to Dr. Richard M. Hansen for numerous suggestions. I wish to express my appreciation also to the following for the loan of specimens in their care: Alfred M. Bailey and A. A. Rogers, Colorado Museum of Natural History, Denver, Colorado; David H. Johnson, United States National Mu- seum, Washington, D. C; Robert W. Lechleitner, Colorado State University, Fort Collins, Colorado; and Robert Z. Brown, Colorado College, Colorado Springs, Colorado. (365) 366 UNivERSiTi' OF Kansas Publs., Mus. Nat. Hist. METHODS Adults of approximately equal age were compared in the study of geographic variation. Three criteria of adulthood are: (a) suture obliterated between supraoccipital and exoccipital, (b) suture at least partly obliterated between basisphenoid and basioccipital, (c) supraorbital crests not widely separated and almost parallel. In males the crests encroach on the lateral borders of the interparietal; in females the crests approach the lateral borders of the inter- parietal but are more widely separated than in males. 105 Fig. 1. Geographic distribution of Thomomys bottae in southwestern Colorado. 1. T. b. howelli 4. T. b. cultellus 2. T. b. aureus 5. T. b. internatus 3. T. b. pervagus 6. T. b. rubidus In studying geographic variation, greater emphasis was placed on females than on males. As noted by Grinnell ( 1931:4), males vary more than females, especially in length of rostrum and associated nasal measurements. Color terms are those of Munsell (1954). Color measurements were stand- ardized by the use of a single 100 watt General Electric blue daylight bulb in a 12 inch white reflector suspended 24 inches above the specimen. All other light was excluded. The individual hairs of Thomomys bottae are either bi- colored or tricolorcd. The darkness of a specimen often may be attributed to the presence of dark-tipped hairs. The color given in tlie description is the basic reddish or yellowish color of the hairs. The presence of a grizzled eflFect or a dark dorsal stripe, or any other pattern resulting from dark hairs, is noted in the remarks. Pocket Gopher in Colorado 367 Specimens examined are listed by counties in the following order: Mesa Montezuma Chaffee Custer Montrose La Plata Fremont Huerfano San Miguel Archuleta El Paso Alamosa Dolores Conejos Pueblo Las Animas Localities are listed from north to south within a county. If two localities lie on the same line of latitude, the western precedes the eastern. Localities omitted on the map in order to prevent overlapping of symbols are in Itahcs. Unless otherwise indicated, specimens are in the University of Kansas, Museum of Natural History. The following initials are used to designate specimens in other collections: CSU — Colorado State University, Fort Collins, Colorado. CMNH — Colorado Museum of Natural History, Denver, Colorado. ERW — E. R. Warren Collection, Colorado College, Colorado Springs, Colo- rado. USNM — United States National Museum, Washington, D. C. The following measurements of the skull are listed in the tables: Condtjlobasal length. — The shortest distance between the anterionnost pro- jections of the premaxillaries and a line touching the posterior surfaces of the exoccipital condyles. Length of nasals. — The distance from the most anterior projection of the nasal bones to the most posterior projection of a nasal bone. Zygomatic breadth. — The greatest distance across the zygomatic arches, at right angles to the long axis of the skull. Squamosal breadth. — The greatest distance between the mastoidal processes of the squamosal. Length of rostrum. — The shortest distance from the shallow notch that lies lateral to the hamulus of the lacrymal bone, to the tip of the nasal on the same side of the skull. Breadth of rostrum. — The greatest width of the rostrum, anterior to the maxillae, transverse to the long axis of the skull. Alveohr length of upper maxillary tooth-row. — Distance between the an- terior margin of the alveolus of the first cheek-tooth and the posterior margin of the alveolus of the last upper cheek-tooth, on one side of the skull. Lea.st interorbital breadth. — The least distance across the frontal bones at the interorbital constriction as seen in dorsal view. PHYSIOGRAPHY Thomomys bottae occurs in the Colorado Plateau Province (ter- minology of Fenneman, 1931), the Southern Rocky Mountain Prov- ince and a small part of the Great Plains Province. The Colorado Plateau Province, in the southwestern part of the state, is mostly above 5000 feet and is characterized by the great number of canyons cut by rivers and streams in the nearly horizon- tal strata. Prominent features of the landscape are cuestas, such as Mesa Verde, and laccoliths, such as Ute Peak. The Southern Rocky Mountain Province consists mainly of high granitic mountains running north and south, many of which extend 368 University of Kansas Publs., Mus. Nat. Hist. to more than 14,000 feet above sea level. Included in this region are several large basins, such as North Park and South Park and the San Luis Valley. The San Juan Mountains, which separate the Colorado Plateau Province from the San Luis Valley, and the Sangre De Cristo and Wet mountains, which intervene between the San Luis Valley and the Great Plains, importantly influence the distri- bution of Thomomijs bottae. The Great Plains Province is a broad highland that slopes grad- ually eastward from tlie Rocky Mountains. Of importance to the present study are two subdivisions of the Great Plains, the Colorado Piedmont and the Raton Section. The Colorado Piedmont is a much dissected fluviatile plain, roughly extending from the vicinity of the Arkansas River to the northern boundary of the state. In general the topography of the Colorado Piedmont is broadly rolling with greater relief than the high plains to the east; however, buttes and steep bluffs occur lo- cally. The Raton Section imperceptibly blends into the southern bound- ary of the Colorado Piedmont and extends south into New Mexico and Texas. A trenched peniplane of greater relief and altitude than the Colorado Piedmont, it is characterized by high mesas, ex- tensive dissected lava-capped plateaus, deep canyons, and moun- tains of volcanic origin. GEOGRAPHIC VARIATION Six subspecies of Thomomijs bottae occur in Colorado. T. b. aureus and T. b. howelli occupy the Colorado Plateau Province (see fig. 1 ) and are characterized by a yellowish color; nasals posteriorly truncate or rounded; posterior extensions of premaxillae long; basi- occipital wide; and interpterygoid space U-shaped with a median spicule. T. b. internatus, T. b. cultellus, and a new subspecies from the vicinity of Carion City described on page 376, inhabit the Sangre De Cristo and Wet mountains in the Southern Rocky Mountain Prov- ince and adjacent parts of the Colorado Piedmont and Raton Sec- tion of the Great Plains Province ( see fig. 1 ) . This group of closely related subspecies is characterized by reddish color; posterior mar- gins of nasals forming a V; posterior extensions of premaxillae short; basioccipital narrow; and interpterygoid space V-shaped, lacking a median spicule. T. b. pervagus occupies part of the San Luis Valley to the west of the Rio Grande (see fig. 1). In Colorado T. b. pervagus is iso- Pocket Gopher in Colorado 369 lated from T. h. internatus and T. h. cultellus by the Sangre De Cristo and Culebra ranges and is separated from T. b. aureus by the San Juan Mountains. T. h. pervogus occupies an area geographi- cally intermediate between T. b. aureus to the west and T. b. inter- naius and T. b. cultellus to the east and has some characters in com- mon with tliese subspecies. T. b. pervagus resembles T. b. aureus in having long posterior extensions of the premaxillae and in some- times having rounded posterior margins of the nasals. T. b. per- vagus resembles T. b. internatus and T. b. cultellus in color, the presence of a V-shaped interpterygoid space, and a narrow basi- occipital. Kelson ( 1951:69) has pointed out that in New Mexico the separation of the ranges of T. b. pervagus and T. b. cultellus is probably complete, but probably incomplete between T. b. pervagus and r. b. aureus. Nevertheless, the similarities between T. b. per- vagus and T. b. cultellus and T. b. internatus suggest that T. b. pervagus was originally derived from the more eastern stock. T. b. aureus is a variable subspecies which, according to Dm-rant (1952:211), intergrades with T. b. howelli in Utah. Specimens of T. b. aureus showing the greatest amount of geographic variation cranially are from the ecotone between the Pinon-juniper and Doug- las Fir zones at the edge of the range of the subspecies. T. b. howelli is a markedly distinct subspecies that shows certain similarities to T. b. aureus, but the degree of cranial difference from T. b. aureus suggests an isolation of long duration, or a rapid evolu- tion from the parent stock. T. b. internatus and T. b. cultellus probably intergrade east of the Sangre De Cristo Range in the vicinity of the Colorado-New Mexico boundary. The amount of intergradation is obscured by the great amount of geographic variation occurring in T. bottae at the edge of the plains and by the lack of specimens from this area. T. b. internatus is a widespread subspecies showing its greatest variation at the edge of the plains. This area is an ecotone between the coniferous forest and the grassland and is by nature an area of change owing to the alternation of wet and dry periods such as the pluvial, interpluvial, and postpluvial periods. This seems to sup- port Durrant's observation (1952:496) that "the greatest range of morphological variation is in animals from the least stable environ- ments." Specimens from a small area north of the Arkansas River in the vicinity of Caiion City ( see fig. 1 ) differ sufficiently from T. b. inter- natus to be given nominal recognition. High mountains and the Arkansas River isolate the new subspecies found at Canon City from 370 University of Kansas Publs., Mus. Nat. Hist. populations of T. b. internatiis to the west and south; however there are no apparent geographic barriers between the newly named sub- species and populations of T. b. internatiis twelve miles to the north or from the vicinity of Pueblo to the east. This new sub- species is the most extreme of the variants occurring in the unstable environment at the edge of the plains. Thomomys bottae aureus Allen Thomomtjs aureus Allen, Bull. Amer. Mus. Nat. Hist., 5:49, April, 1893; Warren, Colorado College Publ., 19:252, January, 1906; Warren, Colo- rado College Publ., 33:77, January, 1908; Warren, Mammals of Colorado, p. 79, 1910; Cary, N. Amer. Fauna, 33:136, August 17, 1911. Thomomys bottae aureus, Goldman, Proc. Biol. Soc. Washington, 48:156, October 31, 1935; Warren, Mammals of Colorado, p. 158, 1942. Thomomys apache Bailey, Proc. Biol. Soc. Washington, 23:79, May 4, 1910. Holotype from Lake La Jara, 7500 feet, Rio Arriba County, New Mexico. Thomomys perpallidus aureus, Bailey, N. Amer. Fauna, 39:74, November 15, 1915. Thomomys perpallidus apache, Bailey, N. Amer. Fauna, 39:75, November 15, 1915. Thomomys bottae apache, Goldman, Proc. Biol. Soc. Washington, 48:157, October 31, 1935; Warren, Mammals of Colorado, p. 160, 1942. Thomomys bottae optabilis Goldman, Jour. Washington Acad. Sci., 26:116, March 15, 1936. Holotype from Coventry, 6500 feet, Montrose County, Colorado; Warren, Mammals of Colorado, p. 159, 1942, part. Holotype. — Adult female, skin and skull number f fit, American Museum of Natural History, obtained at Bluff City, San Juan County, Utah, May 12, 1892, by Charles P. Rowley. Distribution. — Colorado Plateau Province of southwestern Colorado (see fig. 1), northwestern New Mexico, soutlieastern Utah, and northeastern Arizona. Distinctive characters. — Size large ( see measurements ) ; usually pale in west- ern part of range, dark in eastern part; posterior extensions of premaxillae long, wide, and deeply serrated; posterior margins of nasals truncate or slightly rounded (see fig. 2); interpterygoid space U-shaped, with median spicule; basi- occipital wide; bullae well inflated, rounded ventrally. Comparisons. — For comparisons witli T. b. howelli and T. b. pervagus, see accovmts of those subspecies. Remarks. — T. b. aureus is a variable subspecies, which differs con- siderably from T. b. internatus, T. b. culteUus, and T. b. riibidus and includes several microgeographic races distinguishable to a taxonomist specializing in the group. These slightly varying popu- lations are here not considered sufficiently distinct for nominal recognition. Characters such as color of the pelage and conformation of the bullae and zygomatic arches vary with the locality, and to some extent vary among specimens from a single locality. The name Thomomys bottae optabilis, given to specimens from Pocket Gopher in Colorado 371 Covenby by Goldman (1936:116), is here placed in synonymy under T. b. aureus Allen. The characters originally used to describe T. b. optobilis are of the type that vary between populations only a few miles apart, or often vary within a population. The skulls of specimens from Coventry are not lighter in structure than those of T. b. aureus. The premaxillae are not narrower, nor is the frontal region narrower or more constricted than in T. b. aureus. The name Thomoinys bottae apache, given to specimens from Lake La Jara, New Mexico, by Bailey (1910:79), and later applied to specimens from Colorado by Bailey (1915:75), is here also placed in synon)Tny under T. b. aureus. Specimens from Lake La Jara, New Mexico, and nearby localities in Colorado may be separated from topotypes of T. b. aureus on the basis of color only. The topo- types of r. b. aureus are mostly pale; some, however, are dark. The number of pale specimens in any given series decreases gradually in a clinal pattern from west to east. Since there is no noticeable step in the cline and since all specimens show close cranial simi- larity, it is felt that nominal recognition of the darker specimens does not present a realistic picture of the relationships of the rela- tively unisolated populations in the Colorado Plateau Province. Since Thomomys bottae in the Colorado Plateau Province is espe- cially plastic, varying from locality to locality, emphasis is here placed on similarities that unite specimens from different localities. The individual and microgeographic variations are outlined below. Specimens from Bedrock have zygomatic arches that are heavy anteriorly. Specimens from Coventry are dorsally almost uniformly Strong Brown (7.5YR 5/6) and lack a strong dorsal stripe. The venters are Reddish Yellow (7.5YR 8/6). Specimens from 15 miles west of Cortez are the palest specimens of T. b. aureus from Colo- rado, and closely resemble topotypes. The basic color varies from Reddish Yellow' (7.5YR 7/6 and 6/6) to Strong Brown (7.5YR 5/6). Specimens are marked with a narrow dark dorsal stripe. The venters are white. Specimens from Ute Peak and Cortez have Reddish Yellow (7.5YR 6/6) flanks and are slightly darker dorsally. Many specimens from Mesa Verde are indistinguishable from speci- mens from Coventry and from Cortez. Others have dark diffuse dorsal stripes. The venters are Pink (7.5YR 7/4) or Pinkish White (7.5YR 8/2). Some specimens from the Mancos River have wide dorsal stripes. Specimens from three miles west of Durango have especially wide-spreading zygomatic arches posteriorly and have wide black dorsal stripes. The venters are Pink (7.5YR 7/4). One specimen from Florida is dark and grizzled and has a dark dorsal 372 University of Kansas Publs., Mus. Nat. Hist. stripe. Another specimen is pale and has only a small dorsal stripe. Specimens from 12 miles west of Pagosa Springs have thin rostra and diffuse dorsal stripes. Specimens from Bondad have a V-shaped interpterygoid space and in it a small median spicule. One speci- men is uniformly grizzled and lacks a dorsal stripe, giving an over- all effect of Dark Yellowish Brown ( lOYR 3/3 ) . Another specimen has Stiong Brown (7.5YR 5/6) flanks and is only slightly darker dorsally. Specimens examined. — Total 114. Colorado: Montrose Co.: West Paradox Valley, 5 (CMNH); Bedrock, 5150 ft., 5 (ERW); Coventry, 6800 ft., 14 (12 ERW, 2 USNM). San Miguel Co.: 19 mi. N Dove Creek, 6100 ft., 1. Montezuma Co.: Ashbaugh's Ranch (T.36N, R.ISW) 5S50 ft., 5 (4 ERW, 1 USNM); 15 mi. W Cortez (Sec. 2, T.35N, R.19W), 5400 ft., 8; Major Ranch, Cortez, 7 (CSU); 3 mi. SSW Cortez, 6400 ft., 1; Ute Peak, 2 (CMNH); Four Corners, 1 (CMNH). Mesa Verde National Park: Upper Well, Prater Canyon, 7575 ft., 1; % mi. S, 1% mi. W Park Point, 8000 ft., 3; « mi. N Middle Well, 7500 ft., 1; Sec. 27, Head of E Fork, Navaho Canyon, 7900 ft., 2; I'A mi. S, 1% mi. W Park Point, 8000 ft., 1; Middle Well, Prater Canyon, 7500 ft, 9; 3 mi. N Rock Springs, 8200 ft., 4; PA mi. S, 2 mi. W Park Point, 8075 ft., 1; 2'A mi. N, ii mi. W Rock Springs, 8100 ft., 3; 2 mi. N, 'A mi. W Rock Springs, 7900 ft., 2; ?i mi. N Far View Ruins, 7825 ft., 1; Far View Ruins, 7700 ft., 1; 1 mi. NNW Rock Springs, 7500 ft., 1; Rock Springs, 7400 ft., 1; Mancos River, 6200 ft., 9; Mesa Verde, 1 (USNM). La Plata Co.: 1 mi. N La Plata, 1; 3 mi. W Durango, 5; Florida, 6800 ft., 5; Bayfield, 1 (USNM); Bondad, 6 (CMNH); Archuleta Co.: 12 mi. W Pagosa Springs, 6700 ft., 2; Arboles, 1 (USNM). New Mexico: Rio Arriba Co.: La Jara Lake, 7500 ft., 2 (USNM). Thomomys bottae howelli Goldman Thomomys bottae howelli Goldman, Jour. Washington Acad. Sci., 26:116, March 15, 1936; Warren, Mammals of Colorado, p. 161, 1942. Thomomys aureus, Cary, N. Amer. Fauna, 33:136, August 17, 1911, part. Thomomys jjcrpallidtis aureus, Bailey, N. Amer. Fauna, 39:74, November 15, 1915, part. Holottjpe. — Adult female, skin and skull, number 75684, United States Na- tional Museum, obtained by Arthur H. Howell at Grand Junction, 4600 feet, Mesa County, Colorado, November 7, 1895. Distribution. — Colorado Plateau Province of west-central Colorado and cast- central Utah, in the Colorado River Valley cast of the Green River (see fig. 1). Distinctive characters. — Pale (Pinkish White 7.5YR 8/2); cranium flattened; nasals short and wide; posterior tongues of premaxillae long, thin, and at- tenuate (see fig. 3). Comparisons. — Compared with T. b. aureus, T. b. hoivelli differs as follows: paler; nasals shorter and wider; cranium more flattened; posterior extensions of premaxillae longer, thinner, and more acuminate. Remarks. — T. h. howelli most closely resembles T. h. aureus; how- ever, since only one adult specimen of T. b. howelli is known, it is impossible to appraise adequately its characters. Durrant (1952: 211 ) records intergradation between T. b. howelli and T. b. osgoodi, and between T. b. howelli and T. b. aureus in Utah, Pocket Gopher in Colorado 373 An attempt to collect specimens of T. h. howelli, in March, 1957, by Richard S. Miller and the writer was unsuccessful. Speci7nens examined. — Total 2. Mesa Co.: Grand Junction, 4600 ft., 1 (USNM); Sieber Ranch, Little Doloris River, 1 (ERW). Thomomys bottae pervagus Merriam Thomomys aureus pervagus Merriam, Proc. Biol. Soc. Washington, 14:110, July 19, 1901; Gary, Proc. Biol. Soc. Washington, 20:26, March 27, 1907; Warren, Golorado Gollege Publ., 33:77, January, 1908; Warren, Mammals of Golorado, p. 79, 1910, part; Gary, N. Amer. Fauna, 33:137, August 17, 1911, part. Thomomys bottae pervagus, Goldman, Proc. Biol. Soc. Washington, 48:157, October 31, 1935. Thomomys fulvus pervagus, Bailey, N. Amer. Fauna, 39:82, November 15, 1915. Holotype. — Adult male, skin and skull, number 58293, United States Na- tional Museum, Espanola, Rio Arriba Gounty, New Mexico, obtained by J. Alden Loring, January 4, 1894. Distribution. — Upper Rio Grande and San Luis valleys of the Southern Rocky Mountains, in northern New Mexico and southern Golorado ( see fig. 1 ) . Distinctive characters. — Yellowish Red (5YR 4/6); size large (see measure- ments); posterior tongues of premaxillae long, thin, and acuminate; nasals long, tliin, posterior margins usually forming a wide V (see fig. 4); bullae rounded ventrally; interpterygoid space V-shaped, lacking median spicule. Comparisons. — From T. b. aureus, T. b. pervagus differs as follows: reddish, never yellowish or blackish; posterior tongues of premaxillae thin and not deeply serrated; posterior margins of nasals forming a shallow V; interpterygoid space V-shaped, lacking a median spicule; basioccipital narrow. For compari- sons with T. b. internatus, T. b. cultellus, and T. b. rubidus, see accounts of those subspecies. Remarks. — T. b. pervagus is a well-defined subspecies. There is little variation between the topotypes and specimens from Colorado. Specimens examined. — Total 20. Colorado: Gonejos Go.: Antonito, 5 (USNM); 7 mi. E Antonito, 2 (USNM); 12 mi. E Antonito, 1 (USNM); Gonejos River, 6 mi. W Antonito, 8300 ft., 2 (USNM). New Mexico: Rio Arriba Go.: Espanola, 10 (USNM). Thomomys bottae intematus Goldman Thomomys bottae intematus Goldman, Jour. Washington Acad. Sci., 26:115, March 15, 1936; Warren, Mammals of Golorado, p. 160, 1942; Kelson, Univ. Kansas Publ., Mus. Nat. Hist., 5:63, October 1, 1951. Thomomys aureus pervagus, Warren, Mammals of Golorado, p. 80, 1910, part; Gary, N. Amer. Fauna, 33:137, August 17, 1911, part. Thomomys fulvus pervagus, Bailey, N. Amer. Fauna, 39:82, November 15, 1915, part. Holotype. — Adult male, skin and skull, number 150997, United States Na- tional Museum, obtained at Sahda, 7000 feet, Ghaffee Gounty, Golorado, by Merritt Gary, November 10, 1907. 374 University of Kansas Publs., Mus. Nat. Hist. Distribution. — Southern Rocky Mountain Province; southwestern part of the Colorado Piedmont, and Raton Section of the Great Plains, to the east of the Sangre De Cristo Range (see fig. 1). Distinctive characters. — Yellowish Red (SYR 5/6.5); size medium (see measurements); posterior tongues of premaxillae short; posterior margins of nasals forming a V (see fig. 6); bullae pointed ventrally; interpterygoid space V-shaped, lacking a median spicule; basioccipital narrow. Comparisons. — From T. b. pervagus, topotypes of T. b. internatus differ as follows: imifomily paler, not so reddish; smaller; skull smaller; posterior tongues of premaxillae shorter; bullae smaller, less inflated, and more pointed ventrally; zygomata less angular. For comparisons with T. b. cultellus and T. b. rubidus, see accoimts of those subspecies. Remarks. — The dividing line between T. b. internatus and T. b. cultellus is drawn arbitrarily since only one specimen has been col- lected between La Veta Pass and the border of New Mexico. When Goldman (1936:115) named T. b. internatus he included specimens from Union and Colfax counties, New Mexico, and speci- mens from Gardner, Colorado (not Garfield as stated by Kelson, 1951:66). The specimens from New Mexico and a specimen from Fishers Peak, Colorado, were subsequently assigned to T. b. cul- tellus by Kelson ( loc. cit. ) . The specimen from Fishers Peak shows some characters that might be interpreted as intermediate between internatus and cul- tellus, but shows also some unique characters that can be understood only by further collecting in the regions north and northeast of the type locality of T. b. cultellus. Variation is slight in the large series of topotypes of T. b. inter- natus. Specimens from other localities in the western part of the range diflFer little from the topotypes. Specimens from one mile west of Coaldale have slightly more inflated bullae that are more flattened ventrally. Specimens from five miles south of Cotopaxi also have the bullae more flattened ventrally. Specimens from localities bordering the plains differ from the topotypes and near topotypes, and in general show greater varia- tion from locality to locality. Specimens from 12 miles north of Cafion City are dark, resembling T. b. rubidus, but cranially agree with specimens from near Colorado Springs in being indistinguish- able from specimens from Salida. Specimens from St. Charles Mesa and Bear Creek near Walsenburg differ from the topotypes in having wider rostra. The specimens from St. Charles Mesa have more inflated bullae. Pocket Gopher in Colorado 375 Specimens examined. — Total 93. ChafFee Co.: 2 mi. NNW Salida, 7100 ft., 3; Salida, 28 (20 ERW, 8 USNM). Fremont Co.: 12 mi. N Canon City, 5; 1 mi. W Coaldale, 8; Cotopaxi, 1 (CSU); 5 mi. S Cotopaxi, 12. El Paso Co.: 1/4 mi. S Colorado Springs, 2; 9 mi. SSW Colorado Springs, 2; 17 mi. S Colo- rado Springs, 1. Custer Co.: 2/2 mi. S Wetmore, 3; Santa Fe Drive and 20th Lane, Blende, 1; St. Charles Mesa, 5600 ft., 2 (CSU); Fork of Huerfano and Cucharas rivers, 2 (CMNH). Huerfano Co.: 11 mi. WNW Gardner, 7000 ft., 3; Gardner, 7000 ft., 2 (USNM); lYi mi. S Redwing, 3; Bear Creek, near Walsenburg, 2 (CSU); 1 mi. E La Veta, 8; 5 mi. SE La Veta, 2. Thomomys bottae cultellus Kelson Thomomys bottae cultellus Kelson, Univ. Kansas Publ., Mus. Nat. Hist., 5:64, October 1, 1951. Thomomys fulvtis, Gary, Proc. Biol. Soc. Washington, 20:26, March 27, 1907; Warren, Colorado College Publ., 33:76, January, 1908; Warren, Mammals of Colorado, p. 80, 1910. Thomomys fidvus fulvus, Bailey, N. Amer. Fauna, 39:80, November 15, 1915. Holotype. — Adult male, skin and skull, number 70919, United States Na- tional Museum, Halls Peak, Mora County, New Mexico; January 13, 1895, ob- tained by C. Barber. Distribution. — Raton Section of the Great Plains in northern New Mexico and extreme southern Colorado ( see fig. 1 ) . Distinctive characters. — Dark (topotypes); size medium (see measure- ments); posterior tongues of premaxillae short; posterior margins of nasals fomning a V (see fig. 5). Comparisons. — From T. b. pervagus, topotypes of T. b. cultellus differ as follows: darker, not so reddish; smaller; skull smaller; zygomatic arches rela- tively longer; bullae proportionately smaller and less inflated; basioccipital pro- portionately wider; posterior tongues of premaxillae shorter. Topotypes of T. b. cultellus most closely resemble tliose of T. b. internatus but differ as follows: darker; zygomatic arches more widely spreading, not so nearly parallel; nasals not so wide; bullae slightly more inflated. For a comparison with T. b. rubidus see the account of that subspecies. Remarks. — Kelson (1951:64) named T. b. cultellus on the basis of six dark specimens (Dark Reddish Brown 5YR 3/4 and 2/2). Nowhere else within the range of this subspecies, as defined by Kelson, do any specimens resemble tlie topotypes in color. After comparing topotypes of T. b. cultellus with topotypes of T. b. internatus of approximately equal age, I disagree with Kelson {loc. cit.) on some of tlie characters which he used to separate cultellus from internatus. My findings indicate that T. b. cultellus is not smaller, that its skull is not smaller and not less angular, and that the tympanic bullae are not less pointed ventrally. Further collecting is needed better to limit and diagnose this subspecies. Specimens examined. — Total 13. Colorado: Las Animas Co.: Fishers Peak, about 8000 ft., 1 (USNM), New Mexico: Union Co.: Near Folsom, 4 376 University of Kansas Publs., Mus. Nat. Hist. (CMNH); Colfax Co.: Philmont Ranch, Cimarroncito, 8100 ft., 2. Mora Co, Halls Peak, 6 (USNM). 5 6 7 Figs. 2-7. Dorsal views of skulls of Thomomys bottae. X 1. Fig. 2. Thomomt)s b. aureus, 3 mi. W Durango, La Plata Co., Colorado. No. 72967, $. Fig. 3. Thomomys h. howelli, holotype, Grand Junction, 4600 ft., Mesa Co., Colorado. No. 75684 USNM, $ . Fig. 4. Thomomys b. pervagus, Espanola, 5000 ft., Rio Arriba Co., New Mexico. No. 133614 USNM, 9 . Fig. 5. Thomomys h. cultellus. Fishers Peak, 8000 ft.. Las Animas Co., Colorado. No. 129285 USNM, 9 . Fig. 6. Thomomys b. internatus, Salida, 7050 ft., Chaffee Co., Colorado. No. 2757 ERW, 9 • Fig. 7. Thomomys b. rubidus, holotype, 29-10 mi. E Canon City, Fremont Co., Colorado. No. 72954, 9 . Thomomys bottae rubidus new subspecies Holotype. — Adult female, skin and skull, number 72954, Museum of Natural History, University of Kansas, trapped by Richard S. Miller and Phillip M. Youngman, original number 253 (PMY), 2%o miles east of Caiion City, 5344 feet, Fremont County, Colorado, March 17, 1957. Pocket Gopher in Colorado 377 Distribution. — Known only from Garden Park in Canon City and from the type locality ( see fig. 1 ) . Distinctive characters. — Dark (Reddish Brown SYR 3/3); size large (see measurements); skull large; rostrum wide; zygomatic arches rounded and broadly spreading (see fig. 7); alveolar length of upper maxillary tooth-row small. Comparisons. — From topotypes of T. b. internatus, T. b. rubidus differs as follows: uniformly darker; skull averages larger in all measurements, except alveolar length of upper maxillary tooth-row, which is smaller; rostrum pro- portionately wider and tapered anteriorly; zygomatic arches more rounded; bullae more rounded in lateral view. Specimens of T. b. rubidus differ from topotypes of T. b. pervagus in darker color; rostrum wider posteriorly; posterior extensions of premaxillae shorter; bullae smaller, proportionately more inflated posteriorly; zygomatic arches more rounded; wider across squamosals; alveolar length of upper maxillary tooth- row greater. From topotypes of T. b. cultellus, T. b. rubidus differs as follows: paler; larger in all measurements taken; rostnim proportionately wider; zygomatic arches more rounded, less angular; angle formed by zygomatic arch and rostrum greater; bullae proportionately smaller, not so pointed anteriorly; alveolar length of upper maxillary tooth-row shorter. Remarks. — The range of T. b. rubidus is surrounded by the range of T. b. internatus; nevertheless, intergradation has not been found. For a discussion of the geographic relation of T. b. rubidus to T. b. internatus see page 374. Specimens examined. — Total 7. Fremont Co.: Garden Park, Caiion City, 5344 ft., 1; 2^Ao mi. E Caiion City, 5344 ft., 6. SUMMARY A study of 249 specimens of Thomoniys bottae from Colorado re- veals six subspecies in the state. T. b. aureus and T. b. howelli occupy the Colorado Plateau Region in the western and southwest- em parts of the state. T. b. internatus, T. b. cultellus, T. b. per- vagus, and the newly named T. b. rubidus occupy part of the South- ern Rocky Mountain Region and a narrow strip of the Great Plains. The greatest amount of geographic variation, in Thomomys bottae in Colorado, occurs in the ecotone between the grassland and co- niferous forest at the edge of the Great Plains, and in the ecotone between the Piiion, juniper, and sage of the Colorado Plateau and the Coniferous forest of the southern Rocky mountains. 378 University of Kansas Publs., Mus. Nat. Hist. < O n o 05 en 1-1 >< ^ O O U ^ o s o H M-l O H CO H Z in 3 o en O .1-* O c t{jpi?Djq CC QCO (N 000=0 t^OiTj" IR^iq-ioaa^ui ^ CC t^ t^ (Xil>0 CO TO CO 'jSBa'^j MOJ-l^c^oo:^ b- C<>03 o CM ICO iCCOO •x\3ra jaddn jo t^ doo d 00 00 00 OOiOO qqSuai jnp9A|Y mnj'^soj lO O— 1 1— ( oocM >o CM CO CM JO q-^pBajg CO 00 00 o i^odi>- c lO »o -f 1-H ^H .— 1 r^ooi^ ^H i-H T-H q^ptiajq o 'o o CC^ 00 CO CO CO l^t^CM {-Bsomi^nbg d to" do (N CM >> ddc5 CMCM —1 C-J COCM CM CM CM mpB9jq -« l^ e CC* '^ a; > o T}< XO CO 05CO oi'^'Buio2i^2 C5 5 00 (N iC lO i-O CM CM CM 05 .— ■ t^ o o O lO 60 £ gl 1— 1 S5 ooo Tt< -H 00 -* O O rj< q^jSaa^ {tjsbx g r— 1 o coco iC CMC!" 'C COTf< ? 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CD s '^•- rt 03. = (M C^ IN T-H 5 ci.Z O "OiC i-H 1-H loSS IN O h 't ) O t >'b o ^ ) 'b 380 University of Kansas Publs., Mus. Nat. Hist. I C c U PQ < q^pxjojq iO?OtJ < Tt> (N cs lo Oi lu-^iqjOJS^HU «D«D«: arte cc cc CO 'js'Baq Moj-q'^oc; CO iC-- ' 00(N r> CO o •XBiu joddn JO oocdco oJoo « ) GO 00 q;Sua{ JtjpaA^y tuna^soj ^ '^ Tj < cot- t 00 iC JO q'^p^aaa OOOOOO O500 N X t^ luna'^soj OOOiC t>" ■^ '^ = lO lO h- t^ CD • rt «• = OQ O: 05 co-o oo (^ (N CO c3 CO'<<' t^r^ t^ t^ -H CO xog o t 'b o ■b o Pocket Gopher in Colorado 381 ^ 00 y-> o 0606 (N O 00 00 CO I (O ^ o 00<£) CD 00 00 06 cc 06 OCO (N 005 >-H ;0 05 C5 i:d OOlM -H coco CO T3 02 to s o lO 05 CO CC CC CD oc* o CO rt CO t^ o6i> -< -* (N 05 CO Tt< (M 00OI> CO-^ CO o 00 "* CO-* CO CO CO CO O ^ C5 0006 1^ C^ 00 C5 00 00 t^ •^(N 00 C -- 03 iC CO Tfi 00 00»> O (N 00 05 t^ 10 050 00 O -H o r-^ C-^ --1 (N (M (N I -^ 00 10 CO '-0 (N (M (M i-H CO 05 CO CO lO CO Tt^ CD OI>(N C^ CO(N t3^ Tp Tf* ^ Tt^ Oi (N CO (M CO CO(M CO CO CO ^^ Tf^ ■^ C2 (M CO t-^ oor^ (M (N --I (M 01 (M 6 o O S I>00 coco 06 06 t^ 00 O 03 •* CO OJCT) CO CO C^ CO 00 cof 00 00 CO CO 00 I> 00 t>- o oc. ro(N O U CD CD I> CO 10 CO CO 00 CO o CO o a o o O 02 c CO CD 1— t 06 1> 0 10 1 — 1 06 r— 1 d 00 CO 10 00 CO C<1 CD oi CO 0 C5 CD ^ t^ 0 CD C5 1— ( T-H CO coco (35 coco (N coco 00 t— ( y~^ 10 00 'X) rt i c Co .^ o CO^^ r< 'b o o (N O o o (N LO CO (N O 'h 382 University of Kansas Publs., Mus. Nat. Hist. IB^Tqjoja^ui Avoj-q^oo'; •xBui jaddn jo 3 o s o O l-l JO q;pT39Ja mnj^soj JO q-jSuaT q'}pB8jq jiJSOuiTjnbg q'^p'B9jq ij-^Suaj iT;st3]\j q^giisy I'BS'BqOIvtpUOQ -JOOJ pUTIJ n^x q^Suai ii?:jox en O -a o3 o Oi CO -^ 00 00 00 «5 ^ 00 iCrf C^ t^ coo coco (NIM ■^ CO 00 t^ 00 r^ CO CO O5 00 (M(N (N — I (N(N u O s -4-3 CO e (N C5 OS 00 CO CO CO CO lO O CO 00 o 3 Oh a 00 > 03 "J 73 >-. O O b-cd lO. oo 05(N COiM COO C3 00 COCO O Oi CO(N CO X) CO CO 03 CO cc Co' CO CO LO CO (N 00 CO 03 o 05 00 oi CO ■* C3 — —1 (N 00 CO CO CO O paSBjaAt? S[T3nj)IAipUI JO J0C(UJI1U JO joqiunu 2o\v.%v'j X9g (N CO OS OS (N(N O CO t^oo OS OS coco oo o CO 00 o CO 00 Pocket Gopher in Colorado 383 t^ 00 iC-"** lO 05 05 05000 o «o coco CO COCO cot^co o o CO 00 TjH t^ t^ 00 CO l-HTfOO t^ t^ 00 00 t^ 00 05 OOO f— ( 1—* a 00 CD COOS CO t^CD COOOiO ■* lO t-b» J: eo~ -k9 - ^ "o ^ lO tf f-H o3 1-5 lOOO C3 M — CO 00— lO ^ c^ i30 M t>^t^ •0 lO'* t^ OOJ^ (M (N H (N(N v> S (N(N CO CO CO . 'a ■ s 1 S o T— ( ,-(iC o (N(N t^OiO CO CO l.-^CO ^ eo e ■* Tft iCiO>0 f-H T-H 1-H 1— 1 S o i-H ^ i-H i-H 1-H f-H 1-H 05 o ■<* CO 1 &H (N S COCO t^ — N t^ o> (NCI I> s oo -*l-OTt< CO CO Tt; c; >c CO coc^ tOt^ lO (N (N (N(M (N coco coco CO ■^ ^ >h * yr c>^ c OlTf 00 c -^ iC ■^ (M lO »^ > '>' 1 c; ("^ c O-c 05 0- S c:.= O: o c C^ c? CM c; '•-' — — b- b- t^J> 1— 1 a 1 t^b- CO'^'^ o h o h t )'t ) O o K> ► \ ) 3 2S I B W5 ■*-• .2 2.3 0.>HH 'in m -^ iirrt 2 '^ ^ S c C .t; Oh O CO "^ . . . . S ^' > ,;^ May 12, 1958 L..,^!(J^^^|^ A New Bog Lemming (Genus Synaptomys) From Nebraska BY J. KNGX JONES, JR. University of Kansas Laurence 1958 UNTVERSiry OF Kansas Publications, Museum of Nattjbal History Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Robert W. Wilson Volume 9, No. 13, pp. 385-388 Published May 12, 1958 University of Kansas Lawrence, Kansas PRINTED IN THE STATE PRINTING PLANT TOPEKA. KANSAS I 958 27-3033 A New Bog Lemming (Genus Synaptomys) From Nebraska BY J. KNOX JONES, JR. In the autumn of 1952, I obtained a southern bog lemming, Synaptomys cooperi, at Rock Creek State Fish Hatchery, Dundy County, in extreme southwestern Nebraska. This locality of record is the westernmost for the species in North America. Subsequently, 1 reported this specimen in the literature ( Univ. Kansas Publ., Mus. Nat. Hist., 7:486, 1954), provisionally assigning it to Synaptomys cooperi gossii, the subspecies occurring in eastern Nebraska. In late November of 1956, J. R. Alcorn collected three additional bog lemmings at the Rock Creek Hatchery. These specimens from Dundy County represent a relict popula- tion that differs in several characteristics from S. c. gossii, and that differs also from all other subspecies of the species. This relict popu- lation is, therefore, here given subspecific recognition. Synaptomys cooperi relictus, new subspecies Type. — Adult female, skin and skull, University of Kansas Museum of Natural History no. 51617, from Rock Creek State Fish Hatchery, 5 mi. N, 2 mi. W Parks, Dundy County, Nebraska; obtained November 1, 1952, by J. Knox Jones, Jr., original no. 995. Distribution. — Known only from the type locality. Diagnosis. — Size large for the species, both externally and cranially; dorsal pelage dark; nasals broadly flared anteriorly (especially broad in relation to nasal length); auditory bullae small; molariform tooth-rows and incisive foramina long. Measurements (in millimeters). — External measurements of the type speci- men, followed by those of another adult female (KU 72603), are: Total length, 141, 145; length of tail-vertebrae, 24, 21; length of hind foot, 20, 20; length of ear from notch, 11, 12. The type specimen weighed 46.3 grams. Cranial measurements were taken in the manner described by Wetzel (Jour. Mamm., 36:2-3, 1955) except that he did not record the occipitonasal length. These cranial measurements of the type and KU 72603 are: Occipitonasal length, 30.2, 29.8; condylobasilar length, 27.2, 27.1; zygomatic breadth, 18.1, 17.9; lambdoidal breadth, 14.2, 13.8; length of nasals, 8.2, 8.3; breadth of nasals, 4.2, 4.0; length of rostrum, 6.6, 6.6; breadth of rostrum, 6.1, 5.9; breadth of upper incisors, 4.6, 4.2; length of maxillary tooth-row, 8.5, 8.4; length of incisive foramen, 5.8, 5.5; interorbital breadth, 3.1, 3.5, Comparisons. — From Synaptomys cooperi gossii (specimens from eastern Nebraska and eastern Kansas, including one topotype), S. c. relictus differs in: (387) 388 University of Kansas Publs., Mus, Nat. Hist. Dorsal coloration, in comparable pelages, darker, venter lacking buffy tinge; skull averaging larger in all cranial dimensions ( except rostral length, which is approximately the same), especially breadth of upper incisors and length of molariform tooth-rows; nasals broader anteriorly; auditory bullae nearly equal in size, thus relatively smaller. From Synaptomijs cooperi paludis (holotype and paratypes) of Meade County, Kansas, S. c. relictus differs in: Skull aver- aging smaller in all cranial measurements except rostral length and breadth of upper incisors, which are approximately the same, and breadth of nasals, length of incisive foramina and length of molariform tooth-rows, which measure more; nasals relatively (48 per cent of length of nasals) as well as actually broader anteriorly; anterior border of zygomatic plate more concave; auditory bullae smaller; infraorbital foramina larger when viewed anterolaterally. S. c. relictus closely resembles S. c. paludis in color and external proportions. Remarks. — The total population of Synaptomijs cooperi relictus may be small because the only suitable habitat known to me for these mice is the dense, grassy area, approximately 100 yards wide and one mile long, around some of the rearing ponds and along the creek at Rock Creek Hatchery. It has been taken there in associa- tion with Cryptotis parva parva, Blarina brevicaiida caroUnensis, Reithrodontomys megalotis dychei, Peromysctis maniculattis nehras- censis, Microtus ochrogaster haydenii, and another relic, Microtus pennsylvanicus finitis. All specimens of the newly named bog lem- ming are from the border zone between the wet-substrate habitat of M. p. finitis and the drier habitat occupied by M. o. haydenii. Approximately 3000 trap nights produced the four known specimens. S. c. relictus, like S. c. paludis, represents a relict population of the more southwesterly distribution of the subgenus Synaptomys during Wisconsin and post- Wisconsin times. Additional relict pop- ulations likely will be found in the eastern Great Plains. The new subspecies is intermediate in some features between paludis and gossii. The type locality is separated from that of paludis (14 mi. SW Meade, Meade Count)', Kansas) by a dis- tance of approximately 220 miles over habitats largely unsuit- able for bog lemmings. The nearest locality of record for S. c. gossii to the east of the type locality of relictus is at Hunter, Mitch- ell County, Kansas (see Cockrum, Univ. Kansas Publ., Mus. Nat. Hist., 7:196, 1952), approximately 200 miles distant. The locality of record of gossii in Nebraska nearest to the type locality of relictus is even farther eastward — 1 mi. N Pleasant Dale, Seward County (KU 50188). Specimens examined. — Four, from the type locahty (KU 51617, 72601-03). Transmitted March 11, 1958, n 27-3033 -/vA-LLA>v^«v^ceJ University of Kansas Publications Museum of Natural History Volume 9, No. 14, pp. 389-396 December 19, 1958 FEB 2 1959 i. lO MliyEKSIIY mff Pleistocene Bats from San Josecito Cave, Nuevo Leon, Mexico BY J. KNOX JONES, JR. University of Kansas Lawrence ^ 1958 UNrV'EBSITY OF KaNSAS PUBLICATIONS, MuSEUM OF NATURAL HiSTORY Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Robert W. Wilson Volume 9, No. 14, pp. 389-396 Published December 19, 1958 University of Kansas Lawrence, Kansas PRINTED IN THE STATE PRINTING PLANT TOPEKA. KANSAS I 958 27-5516 Pleistocene Bats from San Josecito Ca\b, Nuevo Leon, Mexico BY J, KNOX JONES, JR. Some of the Pleistocene mammals from San Josecito Cave, near Aramberri, Nuevo Leon, Mexico, collected by field parties of the California Institute of Technology under the direction of the late Professor Chester Stock, have been reported previously (see Fur- long, 1943; Cushing, 1945; Stock, 1950; Hooper, 1952; Findley, 1953; Stock, 1953; Handley, 1955; Jackway, 1958). In 1950, Pro- fessor Stock loaned a portion of the San Josecito material to the University of Kansas for identification. Included therein were 89 crania and rami of bats, representing three families ( Phyllostomidae, Desmodontidae and Vespertilionidae ) and five genera, each repre- sented by a single species. One of the species is here described as new. Three of the kinds are known only from the Pleistocene and two are Recent species. The only previous mention of fossil bats from Mexico known to me concerns material from San Josecito Cave. Cushing (1945:182) mentioned a "vampire bat" from the cave (see also Maldonado- Koerdell, 1948:17), and Handley (1955:48) based his description of Corynorhinus tetralophodon on a specimen from San Josecito. Brief descriptions of the cave have been published by Miller (1943) and Stock (1943). The precise age of the deposits is un- known; stratification data did not accompany the material sent on loan to the University of Kansas. However, most of the micro- fauna is thought to have come from the higher levels in the cave and is probably late Pleistocene. The San Josecito Cave collections are currently the property of the Los Angeles County Museum. I am grateful to Dr. E. Raymond Hall for permission to study the bats from San Josecito Cave, to Dr. Robert W. Wilson for criticism of the manuscript, and to Mr. Philip Hershkovitz for permission to use comparative material at the Chicago Natural History Museum. Lucy Rempel made the drawings from photographs by John M. Legler. (391) FEB 2 1959 OHiyERSilY 392 University of Kansas Publs., Mus. Nat. Hist. Leptonycteris nivalis (Saussure) Referred material— Seventy crania, LACM (CIT) 2951-54, 2956-64, 3114-22, 3124-25, 3127, 3131-35, 3137-41, 3143-55, 3942, 21 unnumbered, of which 35 are nearly complete, lacking zygomatic arches, auditory bullae and some teeth; three rami, one right, LACM (CIT) 3126, and two left, un- numbered. Remarks. — The long-nosed bats from San Josecito Cave do not differ appreciably from Leptonycteris nivalis longala Stains, the largest Recent subspecies of the species, and the subspecies that occurs in the same geographic area today. Average and extremes of three cranial measurements of 22 specimens from San Josecito Cave, followed in parentheses by the average and extreme measure- ments of 23 adult L. n. longala from the type locality, 12 mi. S and 2 mi. E Arteaga, 7500 ft., Coahuila (after Stains, 1957: 356), are: Greatest length of skull, 28.2, 27.2-28.9 (27.5, 26.1-29.0); least inter- orbital constriction, 5.0, 4.8-5.4 (4.8, 4.1-5.4); breadth of braincase, 11.1, 10.6-11.6 (10.7, 10.1-11.2). The San Josecito specimens aver- age larger than the series of Recent specimens in all of these meas- urements, especially breadth of braincase, but there is considerable overlap in each case and the extremes of greatest length of skull and of least interorbital constriction do not exceed the extremes in the Recent series. Desmodus stocki, new species Holottjpe. — Cranium, lacking post-incisor dentition on the left side, zygo- matic arches and auditory bullae; Los Angeles County Museum (CIT) No. 3129; from Pleistocene deposits of San Josecito Cave, near Aramberri, Nuevo Leon, Mexico. Referred material. — Twelve additional partial crania, LACM (CIT) 2946-50, 3127-30, 3940-41, 2 unnumbered. Diagnosis. — Resembling the Recent Desmodus rottindus but differing from it as follows: Skull larger (see measurements and Figs. 1-2), heavier and more massive; rostrum and braincase relatively as well as actually broader, interorbital region relatively more constricted; braincase more rounded (less elongate) as viewed from above; nasals less concave in lateral view; narial vacuity broader in relation to greatest length of skull, more nearly heart-shaped; palate broad, less concave medially; mesopterygoid fossa relatively and actually broader anteriorly, the sides nearly parallel; zygomatic arches (judging from No. 2950, the only specimen with a complete arch, the left) less rounded in outline, appearing broader owing to the more constricted interorbital region. Dentition larger and heavier than that in rotundas, but otherwise differing only slightly from it; upper incisor less concave on cutting surface ( see Figs. 3-4); premolar and molar slightly less bladelike, with heavier roots. The peculiar shape of the incisor of stocki is shared to some extent with Diaemus youngi, a Recent South American desmodontid. However, stocki Pleistocene Bats from San Josecito Cave 393 does not otherwise resemble D. youngi, differing from it as follows: Skull larger and heavier; interorbital constriction much narrower; zygomatic arches less strongly bowed; skull less compact, more elongate; braincase and rostrum relatively much narrower in relation to greatest length of skull. Furthermore, specimens of stocki show no trace of the minute M2 attributed to youngi by de la Torre (Proc. Biol. Soc. Washington, 69: 191, 1956). For cranial measure- ments of youngi see Sanborn (Jour. Mamm., 30: 283, 1949). Figs. 1-4. Fig. 1. Dorsal view of holotype of Desmodus stocki, X 1/2. Fig. 2. Dorsal view of Desmodus rotundus mur- inus, $ , KU 54969, La Mula, 13 mi. N Jaumave, Tamaulipas, X 1/2. Fig. 3. Lateral view of left upper incisor of D. stocki, LACM (CIT) 2950, X 2M. Fig. 4. Lateral view of left upper incisor of D. r. murinus, 5 , KU 54967, La Mula, 13 mi. N Jau- mave, Tamaulipas, X 2/2. Remarks. — The essential differences between D. stocki and D. rotundus are in size and proportion. I do not doubt that the two species are closely related; possibly stocki is ancestral to rotundus. The species is named in honor of the late Professor Chester Stock, under whose direction the fossil materials from San Josecito Cave were obtained, and who, at the time of his death, was studying the mammalian fauna from the cave. Eptesicus cf. grandis (Brown) Referred material. — One rostrum, with P4-M3 on the right side and P4 only on the left, LACM (CIT) 2990. Remarks. — This specimen is referred provisionally to E. grandis. The dentition is larger and heavier, and the ridges and depressions on the dorsal surface of the rostrum are more pronounced than in Recent E. fuscus. The P4-M3 length is 6.1 (approximately 6.1 in the holotype of grandis, less in fuscus); least interorbital con- striction, 4.2 ( 4.3 in the holotype of grandis, more in fuscus ) ; breadth of rostrum between infraorbital canals, 6.4; breadth across P4, 7.3. 394 University of Kansas Publs., Mus. Nat. Hist. Table 1. — Cranial measurements of two species of Desmodus. 03 -c s c o3 « '-■s <» - Catalogue number 5 -- u£ to ^ 03 t; cj 6 S or number of -*^ 7j -QJ3 -t-i--3 O o - i c specimens averaged 03 o §0^ 2-^ C to •- C t o >, ti a> t. O O CS3 « H-5 M Desmodus rotundus murinus, La Mula, 13 mi. N Jaumave, Tamaulipas 10 (3cf, 79) Ave.. Max. Min. 24.3 21.4 12.0 12.1 5.5 24.9 22.0 12.5 12.5 5.6 23.9 21.0 11.7 11.9 5.3 5.3 5.1 Desmodus stocki, San Josecito Cave, Nuevo Le6n 2946 27.3 24.5 14.2 13.6 13.9 13.9 13.5 13.5 13.5 13.7 13.8 13.7 6.1 "■6;2"' 6.1 "e^o" 6.2 5.9 6.2 6.0 5 8 2947 5.7 2948 24.3 24.7 "u.i" 5 3 2949 5.5 2950 5 7 3127 5.7 3128 26.5 28.2 27.4 5.5 3129 (type) 24.5 24.4 24.6 "l4'6" 5.7 3940 3941 5.6 Brown (1908:174) originally named grandis as a subspecies of fuscus. Gidley and Gazin (1938:11) considered it a distinct spe- cies. Whether grandis is only a subspecies of E. fuscus or a separate species, grandis is closely related to fuscus, and probably is ances- tral to it. Lasiurus cinereus (Palisot de Beauvois) Referred material. — One cranium, lacking basioccipital, tympanic and mas- toid regions, and most of the dentition, having only M3 on the right side and M2-M3 on the left, LACM (CIT) 3160. Remarks. — The cranium of No, 3160 is inseparable from those of 10 spring-taken specimens of L. c. cinereus from the San Gabriel Mts., Los Angeles Co., California (KU 49727, 49729-37). Measure- ments of No. 3160, followed by the average and extremes ( in paren- theses) of the Californian series, are: Condylobasal length, 16.1, 16.5 (15.9-17.2); zygomatic breadth, 12.3, 12.4 (12.0-12.7); least interorbital constriction, 5.2, 5.4 (5.2-5.6); breadth of braincase, 8.7, 9.0 (8.5-9.3); length of palate not including terminal spine, 5.1, 5.3 (4.8-5.9). The teeth of the San Josecito specimen are com- paratively unworn. A label with the skull bears the notation "talus" Pleistocene Bats from San Josecito Cave 395 in parentheses, which, in so far as I am able to determine, indicates surface talus inside the cave. Therefore, the specimen in question may be of Recent origin. It is perhaps worthy of note that Lasiurus cinereus is primarily a tree-dwelling bat, although a few Recent specimens have been re- ported from caves (see Beer, 1954:116). Corynorhinus tetralophodon Handley A single cranium of a Coryriorhinus LACM (CIT) 2989 was in- cluded in the original materials sent to Kansas by Professor Stock. Subsequently, this specimen was loaned to Charles O. Handley, Jr., who described it as a new species, C. tetralophodon. The latter is said to differ from all other plecotine bats by the retention of a well-developed fourth commissure (ridge extending posteroexter- nally from metacone) on the M3 (Handley, 1955:48). LITERATURE CITED Beer, J. R. 1954. A record of the hoary bat from a cave. Jour. Mamm., 35:116, February 10. Brown, B. 1908. The Conard Fissure, a Pleistocene bone deposit in northern Ar- kansas: with description of two new genera and twenty new species and subspecies of mammals. Mem. Amer. Mus. Nat., 9:155-208, pis. 14-25. Gushing, J. E., Jr. 1945. Quaternary rodents and lagomorphs of San Josecito Cave, Nuevo Leon, Mexico. Jour. Mamm., 26:182-185, July 19. FiNDLEY, J. S. 1953. Pleistocene Soricidae from San Josecito Cave, Nuevo Leon, Mexico. Univ. Kansas Publ., Mus. Nat. Hist., 5:633-639, December 1. Furlong, E. L. 1943. The Pleistocene antelope, Stockoceros conkUngi, from San Jose- cito Cave, Mexico. Carnegie Inst. Washington Publ., 551:1-8, 5 pis., February 3. GiDLEY, J. W., and Gazin, C. L. 1938. The Pleistocene vertebrate fauna from Cumberland Cave, Mary- land. BuU. U. S. Nat. Mus., 171 :vi + 99, 50 figs., 10 pis. Handley, G. O., Jr. 1955. A new Pleistocene bat (Corynorhinus) from Mexico. Jour. Wash- ington Acad. Sci., 45:48-49, March 14. Hooper, E. T. 1952. A systematic review of the harvest mice (genus Reithrodontomys) of Latin America. Misc. Publ. Mus. Zool., Univ. Michigan, 77: 1-255, 9 pis., 24 figs., 12 maps, January 16. 396 University of Kansas Publs., Mus. Nat. Hist. Maldonado-Koerdell, M. 1948. Los vertebrados fosiles del Cuaternario en Mexico. Revista Soc. Mexicana Hist. Nat., 9:1-35, June. Jackway, G. E. 1958. Pleistocene Lagomorpha and Rodentia from the San Josecito Cave, Nuevo Leon, Mexico. Trans. Kansas Acad. Sci., 61 : in press. Miller, L. 1943. The Pleistocene birds of San Josecito Cavern, Mexico. Univ. California Publ. Zool., 47:143-168, April 20. Stains, H. J. 1957. A new bat (genus Leptonycteris ) from Coahuila. Univ. Kansas Publ, Mus. Nat. Hist, 9:353-356, January 21. Stock, C. 1943. The cave of San Josecito, Mexico. New discoveries of vertebrate life of the ice age. Eng. Sci. Monthly, California Inst. Tech., Balch Grad. School Geol. Sci. Contrib., 361:1-5, September. 1950. Bears from the Pleistocene cave of San Josecito, Nuevo Leon, Mexico. Jour. Washington Acad. Sci., 40:317-321, 1 fig., Octo- ber 23. 1953. El caballo pleistocenico {Eqiius conversidens leoni, subsp. nov.) de la cueva de San Josecito, Aramberra, Nuevo Leon. Mem. Congr. Cient. Mex., 3:170-171. Transmitted August 18, 1958. n 27-5516 /V Pf-L L^^^^'^CC J FEB 2 1959 iS< I.. . k - w.;J University of Kansas Publications Museum of Natural History Volume 9, No. 15, pp. 397-404 December 19, 1958 New Subspecies of the Rodent Baiomys From Central America BY ROBERT L. PACKARD University of Kansas Law^rence ^ 1958 University of Kansas Publications, Museum of Natural History Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Robert W. Wilson Volume 9, No. 15, pp. 397-404 Published December 19, 1958 University of Kansas Lawrence, Kansas PRINTED IN THE STATE PRINTING PLANT TOPEKA. KANSAS 1956 27-5660 New Subspecies of the Rodent Baioitiys " ' - 1 FEB 2 1959 MM From Central America BY ROBERT L, PACKARD The southern pygmy mouse, Baiomijs musculus, is known as far north as the Mexican states of Jalisco, Michoacan, south of the Mesa Central, east to central Veracruz (see Hooper, 1952a: 90), and south to western Nicaragua (see Goodwin, 1942:161). Previously, two subspecies have been recognized from the southern part of tlie known range of this species: B. m. nigrescens, blackish mice from Chiapas, Mexico, and Guatemala, and B. m. grisescens, gray- ish-brown mice from Honduras and western Nicaragua. Study of recently acquired specimens from Guatemala, El Salvador, and Nicaragua reveals two additional subspecies. For the loan of comparative material, I am grateful to the United States National Museum (USNM) and the American Museum of Natural History (AMNH). Unless otherwise indicated, specimens are in the University of Kansas Museum of Natural History. Meas- urements are as taken by Hooper (1952b: 10). Postpalatal length is the distance from the posterior margin of the hard palate to the anterior margin of the foramen magnum. Unless otherwise noted, statistical significance as used in this paper is at the 95 per cent confidence limit or higher. The two heretofore undescribed subspecies are characterized below and may be known as: Baiomys musculus handleyi, new subspecies Type. — Adult female, USNM No. 275604 (Biological Surveys Collection), skin and skull; from Sacapulas, El Quiche, Guatemala; obtained on April 24, 1947, by Charles O. Handley, Jr., original number 991. Distribution. — Known only from the type locality; probably inhabits parts ot the east-west drainage of the Rio Negro, Diagnosis. — General ground color of upper parts between Wood Brown and Buffy Brown (all capitalized color terms are those of Ridgway, Color Standards and Color Nomenclature, Washington, D. C, 1912), dorsal parts of fore- and hind-feet, and ankles white; in region of median venter, throat, and chin, hairs white to base; in lateral regions hairs Neutral Gray at base; dorsal hairs below tips Avellaneous, Neutral Gray at base; guard hairs black- tipped; tail white below, brownish above; nasals truncate anteriorly; frontal- parietal suture forming an obtuse angle with median-parietal suture; alveolar- length of upper molar tooth-row and tail long. (399) 400 University of Kansas Publs., Mus. Nat. Hist, Comparisons. — From Baiomys musculus nigrescens (paratypes, from the Valley of Comiti'm, Chiapas, Mexico), found to the north, B. m. handleyi differs in: color paler dorsally and ventrally; fore- and hind-feet whitish instead of dusky to sooty; hairs in region of facial vibrissae white instead of brown; tail bicolored instead of unicolored; anterior tips of nasals square, not rounded; frontoparietal suture forming obtuse angle with median parietal suture instead of a right angle; tail and alveolar length of upper molar tooth- row significantly larger ( see table 1 ) ; zygomatic breadth, breadth of braincase, occipitonasal length, least interorbital constriction, and length of rostnun all averaging larger (see table 2). From Baiomys musculus grisescens (paratypes, from Comayabuela, Hon- duras) to the south, B. m. handleyi differs in: buff-colored hairs in dorsal and ventral region lacking; fore- and hind-feet white, not flesh-colored with gray overtones; tail bicolored, not unicolored; face paler, lacking buff-brown coloration; anterior tips of nasals squared rather than flaring; tail and upper molar tooth-row significantly longer (see Table 1); hind foot, ear from notch, and rostrum longer; braincase averaging broader (see Table 2). Remarks. — The occurrence of these pale mice in the Rio Negro Valley was first noted by Goodwin (1934:39, 40) when he referred specimens from Sacapulas and Chanquejelve, Guatemala, to B. m. musculus. Hooper (op. df.;92-94) correctly assigned specimens from the southern coast and eastern part of Chiapas to B. m. ni- grescens. The continued assignment of specimens from Sacapulas, Guatemala, to the subspecies musculus produces a hiatus both in the range of B. m. nigrescens and B. ni. musculus. Twenty-four specimens, 14 from 1 mi. S Rabinal, and 10 from /2 mi. N, 1 mi. E Sa- lama, Guatemala, are intergrades between handleyi and grisescens, but show more resemblance to the latter and, therefore, are referred to that subspecies. To the north, handleyi intergrades with ni- grescens. The specimen from Chanquejelve is an intergrade between the two subspecies just mentioned. Osgood suggested ( 1909:259) that the degree of relative humidity might in some way control color of pelage in this species. Relative humidity and its subsequent effect on other related environmental factors indeed may account for the superficial resemblance of B. m. musculus to B. m. handleyi (although handleyi averages paler throughout than the paratypical series of musculus). Both sub- species inhabit relatively arid country. According to Goodwin (op. ciY.:39 and Plate 5, Fig. 1), and Handley {in verbis), the Rio Negro Valley in the vicinity of Sacapulas is extremely hot, dry, and rather isolated. Extremes of climate there may exceed those in the arid habitat occupied by B. m. musculus. The resemblance between these two subspecies may result from nearly parallel selective forces New Subspecies of the Rodent Baiomys 401 that have given rise to two distinct subspecies. B. vi. handleyi may have developed in situ. Specimens examined. — Total 49, from the type locality, including the type (12, USNM; 37, AMNH). Baiomys musculus pullus, new subspecies Type. — Adult female, skin and skull, University of Kansas Museum of Natural History, No. 71605, from 8 mi. S Condega, Esteli, Nicaragua; ob- tained on July 15, 1956, by A. A. Alcorn, original No. 4218. Distribution. — West-central Nicaragua, from Matagalpa northwest into the valley of the Rio Esteli, east as far as Jinotega. Diagnosis. — Dorsum Fuscus-Black (see remarks), individual dorsal hairs being black-tipped with a subterminal Ochraceous-BuflF band. Neutral Gray at base; some hairs on dorsum all black to Neutral Gray at base; hair on sides Neutral Gray tinged with blackish; facial region blackish becoming more buffy ventrally; vibrissae black; tail unicolored Chaetura Black; fore- and hind-feet whitish to dusky-white; mid-ventral region of belly white to as far anteriorly as region of throat, hairs being white to base; in region of anus and throat, hairs white-tipped. Neutral Gray at base; tail long; upper molar tooth-row short as in B. r?i. nigrescens; zygoma bowed as in B. m. grisescens. Comparisons. — From B. m. grisescens (paratypes from Comayaguela, Hon- duras), B. m. pullus differs in: dorsal ground-color and tail darker; sides and distal region of belly grayish instead of buffy-brown, thus making white stripe in region of belly distinct; average length of body and tail significantly longer, thus, total length greater; length of hind foot averaging longer (68 per cent confidence limits); alveolar length of upper molar tooth-row significantly shorter; occipitonasal and rostral length averaging longer; zygomatic spread and interorbital region narrower; length of incisive foramina, depth of cranium, postpalatal length, and breadth of braincase all averaging larger (see table 2). From B. m. nigrescens (paratypes from Valley of Comitan), B. m. pullus differs in: dorsal ground-color slightly darker; facial region grayish, not sooty; mid-ventral white stripe present on belly and becoming grayish laterally; tail darker and less hairy, average length significantly longer; body, occipitonasal length of skull, incisive foramina, and postpalatal length averaging smaller; hind foot shorter; zygomatic spread, interorbital region and braincase broader (see table of measurements); cranium deeper. Remarks. — B. m. pullus is the darkest dorsally of any subspecies of this species. Dalquest (1953:156) pointed out that preserved specimens of one of the subspecies of the northern pygmy mouse, Baiomys toylori taylori, tended to fade considerably over a period of four years. Post-mortem changes in color also are apparent in the southern species musculus. For example, the series of specimens from 8 mi. S of Condega, and 9 mi. NNW Esteli, Nicaragua, have faded from near Chaetura Black to the present Fuscous-Black in a period of two years. The most notable change in color came after the first six months of preservation. Allowing for this fading, the 402 University of Kansas Publs., Mus. Nat. Hist. Table 1. — Analysis of Variation in Adults of Four Subspecies of Baiomys musculus (measurements in millimeters) Number of adults averaged Total length Length of body Length of tail Length of hind foot Upper molar length (alveolar) 9 Av 121.44 128.00 115.00 3.60 Baiomys musculus handleyi Sacapulas, El Quiche, Guatemala 70.77 50.67 15.33 77.00 54.00 16.00 66.00 49.00 15.00 3.22 1.26 .44 3.48 Max 3.60 Min 3.40 2xStand. error .05 17 Av 117.29 121.00 111.00 1.27 Baiom 8 mi. S 70.42 74.00 66.00 1.51 ys miisculus Dondega, N 47.18 50.00 44.00 .75 pullus caragua 15.47 17.00 14.00 .35 3.13 Max 3.20 Min 3.00 2xStand. error .03 7 Av 103.71 118.00 97.00 5.50 Baiomyi Coma .59.00 68.00 51.00 4.16 J musculus g yaguela, Ho 44.71 50.00 42.00 2.40 risescens nduras 14.57 15.00 13.00 .78 3.31 Max 3.40 Min 3.20 2xStand. error .06 11 Av 115.00 120.00 108.00 2.12 Baiomyt Va 72.09 77.00 69.00 1.59 • jjiuscidus n ley of Comi 42.91 45.00 39.00 1.0 igrescens tdn 15.31 16.00 14.50 .23 3.15 Max 3.40 Min 2.90 2xStand. error .10 several color differences between pullus, nigrescens and grisescens are, nevertheless, distinctive. Geographically, pullus is partly isolated by the Cerros De Villa- guaire and the Cerros El Zapotillo to the west and the Cerros De Azaculapa to the north. Certain individuals of a series of speci- mens, referable to B. m. nigrescens, from 1 mi. NW San Salva- dor and 1 mi. S Los Planes, El Salvador, are intermediate in col- oration between that subspecies and pullus. Three of 28 specimens from El Salvador possess the mid-ventral white stripe. New Subspecies of the Rodent Baiomys 403 Table 2. — Cranxal Measurements (in millimeters) of Adults of Four Subspecies of Baiomys musculus Number of specimens Type 275604 9 USNM Average Maximum Minimum Number of specimens . . . Type 71605 9 KU Average Maximum Minimum 03 .2" S o o rt -(J ^ (h c3 a; oJ3 n^ 2-a gth of incisive Foramen m2 1"^ tpalat length o S C3^ bD CO cS G d CJ >> O 0) ^ S. Fitch and Lewis L. Sandidge. Pp. 305-338, 5 figures in text. August' 24, 1953. 3. The silky pocket mice (Perognathus flavus) of Mexico. By Rollin H. Baker. Pp. 339-347, 1 figure in text. February 15, 1954. 4. North American jimiping mice (Genus Zapus). By Philip H. Krutzsch. Pp. 349-472, 47 figures in text, 4 tables. April 21, 1954. 5. Mammals from Southeastern Alaska. By RoLUn H. Baker and James S. Findley. Pp. 473-477. April 21, 1954. 6. Distribution of Some Nebraskan Mammals. By J. Knox Jones, Jr. Pp. 479- 487. April 21, 1954. 7. Subspeciation in the montane meadow mouse. Microtus montanus, in Wyo- ming and Colorado. By Sydney Anderson. Pp. 489-506, 2 figures in toxt. July 23, 1954. 8. A new subspecies of bat (Mvotis velifer) from southeastern California and Arizona. By Terry A. Vaughan. Pp. 507-512. July 23, 1954. 9. Mammals of the San Gabriel mountains of California. By Terry A. Vaughan. Pp. 513-582, 1 figure in text, 12 tables. November 15, 1954. 10. A new bat (Genus Pipistrellus ) from northeastern Mexico. By RoUin H. Baker. Pp. 583-586. November 15, 1954. 11. A new subspecies of pocket mouse from Kansas. By E. Raymond Hall. Pp. 587-590. November 15, 1954. 12. Geographic variation in the pocket gopher, Cratogeomys castanops, in Coa- huila, Mexico. By Robert J. Russell and Rollin H. Baker. Pp. 591-608. March 15, 1955. 13. A new cottontail (Sylvilagus floridanus) from northeastern Mexico. By Rollin H. Baker. Pp. 609-612. April 8, 1955. 14. Taxonomy and distribution of some American shrews. By James S. Findley, Pp. 613-618. June 10, 1955. 15. The pigmy woodrat, Neotoma goldmnni, its distribution and systematic posi- tion. By Dennis G. Rainey and Rollin H. Baker. Pp. 619-624, 2 figiu-es in text. June 10, 1955. Index. Pp. 625-651. (Continued on inside of back cover) University of Kansas Publications Museum of Natural History Volume 9, No. 17, pp. 415-511, 12 figs, in text, 2 tables August 1, 1959 Distribution, Variation, and Relationships of the Montane Vole, Micro tus montanus BY SYDNEY ANDERSON University of Kansas Lavv^rence 1959 University of Kansas Publications, Museum of Natx^al History Editors: E, Raymond Hall, Chairman, Henry S. Fitch, Robert W, Wilson Volume 9, No. 17, pp. 415-511, 12 figs, in text, 2 tables Published August 1, 1959 University of Kansas Lawrence, Kansas filUS. COMP. ZflOll LIIRAftY SEP -3 1959 HARVARD UNiYERSin PRINTED IN THE STATE PRINTING PLANT TOPEKA, KANSAS I 959 Distribution, Variation, and Relationships of the Montane Vole, Microtus montanus BY SYDNEY ANDERSON CONTENTS PAGE Introduction, materials and acknowledgments 418 Distribution 419 Method of study 419 Influence of habitat 421 Influence of other species 424 Variation 432 Non-geographic sources 432 Geographic variation 437 Accounts of subspecies 447 M. m. canescens 447 M. m. pratincola 451 M. m. canicaudus 452 M. m. nanus 456 M. m. codiensis 462 M. m. zygomaticus 464 M. m. montanus 465 M. m. dutcheri 472 M. m. undosus 473 M. m. micropus 474 M. m. amosus 476 M. m. rivularis 478 M. m. nevadensis 480 M. m. fucosus 480 M. m. fusus 481 M. m. arizonensis 483 Relationships of Microtus montanus 48 Summary 497 Literature Cited 508 (417) INTRODUCTION, MATERIALS, AND ACKNOWLEDGMENTS The distribution and the variation of animals have fascinated naturalists from antiquity, but more especially from the time of Charles Darwin, to the present day. The search for an explana- tion of the distribution and variation of animals gave rise to the theory of organic evolution, a theory of pervasive import in biology and in other fields. The details of the distribution and the variation of one species of mammal, the montane vole, Microtus montanus, are the sub- ject of this dissertation. The relationship of this species to other species of the same genus is considered, and various characteristics, especially the structure of the baculum, are considered in other species. Specimens numbering 4722, mostly skins with skulls, in the col- lections listed below were studied. Most of these collections were visited. Selected specimens of the collections visited, and of collections not visited were borrowed for study and comparison. Every state in which Microtus montarms occurs was visited, and approximately six months were spent in the field from 1950 through 1956. This experience enabled me to study the animal in certain areas, and to gain a general acquaintance with the habitat occupied throughout the range of the species. Even overnight stops at times yielded valuable distributional records; for example the specimens from Gilliam and Morrow counties in Oregon obtained on two different nights bridged a gap of more than 100 miles in the known geographic range. Specimens examined are listed in the accounts of the subspecies concerned. The number of specimens in each museum from each locahty is noted using the abbreviations for museums hsted below. I am indebted to the persons named who made specimens readily available for study. (AM) American Museum of Natural History, G. G. Goodwin and the late G. H. H. Tate. (BC) University of British Columbia, Department of Zoology, I. M. T. Cowan. ( BY ) Brigham Young University, C. L. Hayward. (CN) Canadian National Museum, A. W. Cameron. ( CM ) Chicago Natural History Museum, C. C. Sanborn. ( DM ) Denver Museum of Natural History, A. C. Rogers. (EB) Ernest S. Booth, College Place, Washington, personal collection. (KU) University of Kansas, Museum of Natural History; including the G. C. Rinker collection and the Ralph Ellis collection. (418) Anderson — Microtus montanus 419 ( LA ) Los Angeles County Museum, K. Stager. (MV) Museum of Vertebrate Zoology, S. B. Benson and O. P. Pearson. (MZ) Museum of Zoology, University of Michigan, W. H. Burt and E. T. Hooper. (NM) University of New Mexico, J. S. Findley. ( OS ) Oregon State College, Franklin Sturges. (PM) Provincial Museum of Natvural History and Anthropology, British Columbia, C. J. Guiguet. ( PS ) College of Puget Sound, M. L. Johnson. ( RO ) Royal Ontario Museum of Zoology, R. L. Peterson. (SD) San Diego Society of Natural History, L. M. Huey. (UA) University of Arizona, E. L. Cockrum. (UC) University of Colorado Museum, H. G. Rodeck. ( UM ) University of Montana, Dept. of Zoology, P. L. Wright. (US) U. S. National Museum, including the Biological Surveys Collec- tion, D. H. Johnson, Viola S. Schantz. ( UU ) University of Utah, S. D. Durrant. (WC) E. R. Warren collection, Colorado Springs, R. M. Stabler. (WS) Washington State College, C. R. Conner Museum, G. M. Hudson. (WW) Walla Walla College, E. S. Booth. I am grateful also to the many naturalists vv'hose labors through the years built the above collections, and to colleagues at the University of Kansas with whom I have discussed problems arising in this study and whose thoughts have contributed to those here written as mine. Professors E. Raymond Hall and Rolhn H. Baker have aided me through counsel since the inception of the study. My wife, Justine Anderson, has assisted in the field, given helpful encouragement, and aided in the final mechanics of preparing manuscript. DISTRIBUTION Method of Study Most small mammals are secretive in habit, and can be recog- nized only with difficulty even when examined at close range. Therefore specimens must be preserved for accurate identification. The specimens with accompanying data are the evidence on which an accurate account of distribution must be based. The listing and the permanent preservation of the specimens make possible the essential scientific process of verification. The locality of capture, as recorded on labels of specimens, was noted for each specimen examined by me. These localities were plotted on a large map (scale 1:2,500,000). This map, reduced one half with a pantograph, was the basis for the maps shown in Figs. 8 through 11; in making the pubUshed maps the figures were reduced another one half or to one fourth the original size. The dots, on the map showing localities, have a radius equivalent to approximately six miles. To prevent overlapping of dots, other localities within 12 miles of a locality shown were omitted. Those omitted are indicated by italic type in the lists of specimens examined. There are 479 locahties mapped. They provide a reasonably accurate pictiure of the dis- 420 University of Kansas Publs., Mus. Nat. Hist. tribution of the species. Lines showing the probable ranges of the subspecies and of the species as a whole are estimates taking into account the habitat requirements of the animals and our acquaintance with the habitat. The re- quirements differ in diflFerent parts of the range as will be pointed out below, and our knowledge of different areas is unequal. Therefore, the accuracy of the estimated range varies. Although intensity of the field work has varied in 117 107 ^ ■■•.. • 1 * '"■■■ ^ \i ' ' 50 \ 11 \\ i ! \ ■•. •:--^*''T\*) H j \ 50 o\^--i%^. ^ i --- (j 7^1 : u 7\-.if 1 \ \ '•■••' /f IS \ A ^ i ••■" -'l 'f V ^S )• ■ ■"■ j^ ' "^^ i ..•■\./_ .. ( X, }■ P ' V/ 1 / 45 \Ji P ''''^^'^ /^\_^i\ ^i^7\ <4v "" / 45 / i } ) ^S^^klV^ ■ s 1'^ "■■!'' Ej_ '.!. ' $(^k'r" 40 -I \ \ivi i*w t: 40 35 35 K ^ ''■<■■■.. ... 1 0 100 800 \ *" ,'^. ' ' SCAL I OF MILES \ L.^ ^^ --^ S 117 107 Fig. 1. The geographic range of Microtus 7nontanus (solid lines) and the distribution of boreal life-zones (dotted lines). That part of the range of M. montanus within the boreal zones is stippled. Note that in the Great Basin and in the northern part of the range of M. montanus areas are occupied that are zonally lower than in the southern part of the range. Each of the question marks is the center of an area at least 70 miles in diameter from which no specimens have been taken. Anderson — Microtus montanus 421 different areas, sufficient work has been done in all parts of the range to sug- gest that any large hiatus remaining is not accidental. To focus attention on remaining gaps, question marks are placed on Figure 1 in each area 70 miles or more in diameter from which no specimen has been preserved. Influence of Habitat In a general way at least the influence of habitat on the occur- rence of most species is obvious. The details of the interrelation- ship of organism and environment are not obvious; they are, in fact, incredibly complex. Numerous approaches have been used to study these interrelationships. The approaches include carefully controlled experiments in the laboratory, intensive study of fluctu- ations in local populations, and analysis of data gathered through- out the range of a species. The last approach is used here. One classification of habitats that has been widely used in the western United States is that of life-zones, originally outlined by C. H. Merriam (1890). These zones are based, in theory, on the temperature in the reproductive season, but in practice, on the occurrence of various species of both plants and animals that serve as indicators. The theoretical basis has been debated, but the es- sential fact of altitudinal and latitudinal zonation remains. The lower boundary of the boreal region and the range of the montane vole are mapped in Figure 1. The boundary drawn is an approxi- mation based on the maps of various authors. That part of the range of the vole within the boreal region is indicated. It is ap- parent that except in the Great Basin the species occurs not only altitudinally but zonally higher in the southern part of its range than in the northern part of its range. Voles of the genus Microtus are commonly associated with grass, and their characteristics and habits fit them for a grassy habitat. Persistently growing, prismatic teeth are adapted to a harsh, often- times abrasive, bulky diet of grass. The habit of clearing and maintaining open runways beneath the grass is characteristic. The voles move, when necessary, with rapidity along these paths. Grass provides both food and shelter. With some modifications, grass is the essential factor that characterizes the habitat of Mi- crotus montanus throughout its range. The relative elevation zon- ally, as well as altitudinally, mentioned above, is explainable in terms of this association with grass. Grass depends on moisture, more moisture than is found in most areas at lower elevations in the southern part of the range of the species. The elevated moun- tainous areas in the essentially arid, subtropical, hot Southwest 422 University of Kansas Publs., Mus. Nat. Hist, provide a local climate that supports pine trees of species of more northern afiBnities. These trees are prominent and easily observ- able organisms and are commonly used as indicators of life-zones. As noted earlier (Findley and Anderson, 1956), with the progres- sive dehydration that followed the latest Pleistocene glaciation boreal species retreated to higher elevations and more northern climes. The sometimes disjunct distribution of many of these boreal species, including Microtus montanus, in the southern parts of their ranges attest to this retreat. The montane climate of the Southwest provides moisture not only for trees but for grass. Lower areas are desert except for scattered drainages, springs, or irrigated areas that provide water to support a permanent growth of grass luxuriant enough to satisfy the needs of Microtus. In the northern states conditions are dif- ferent. The lowlands there have a lower mean annual temperature and more humidity; consequently, they are temperate prairie rather than desert. The montane forest of the Canadian Life-zone is al- titudinally lower and is more uniform than farther south. At higher altitudes meadows, perhaps, are fewer than farther south. The trees that indicate the boreal region, the chief areas of grass- land, and the areas in which Microtus montanus occur, are at lower altitudes than in the south. The reduction from south to north in the altitude at which the grass and the voles occur is greater than the reduction in the altitude at which the trees occur. This difiFerence helps to explain the pattern shown in Figure 1. A de- scription of the habitat of M. montanus in several states will illus- trate the above relationships. In Washington and British Columbia M . montanus occupies the relatively arid valleys of the Columbia, Snake, and Okanogan rivers, the Palouse Prairie, and the Blue Mountains of the southeastern part of Washington, and is absent from the Cascade Mountains and the mountains to the northeast in these two states and in Idaho. Specimens, not previously reported, from the Grand Coulee area and from Adams County and vicinity have been generously made available to me by Drs. Murray L. Johnson and George M. Hudson. These specimens from the Columbia Basin demon- strate that M. montanus occurs throughout the eastern part of the state of Washington, except perhaps in the extreme northeastern corner. Specimens from eastern Washington described earlier as Microtus montanus kincaidi by Dalquest (1941:145) proved to be Microtus pennsylvanicus. In Oregon, M. montanus occupies the Anderson — Microtus montanus 423 valleys of the Deschutes, Columbia, and Snake rivers and their tributaries east of the Cascade Mountains and by way of the Columbia Valley has reached the Willamette Valley west of the Cascades. In the southeastern part of the state where conditions are most arid the species occupies not only suitable local areas at lower elevations but occurs in and near the Steens Mountains. In south-central Oregon voles of this species occupy marshes and other grassy areas at various elevations. Farther to the south, in northern California, the voles occur at higher elevations, and are not present at lower elevations on the west side of the Sierra Nevada. In the southern Sierra Nevada they occur only in the boreal environment at higher elevations. To the east of the Sierra Nevada, in Nevada, they do occur at the lowest available eleva- tions in more or less separated areas where springs or other waters provide suflScient grass. The same trend is apparent in the eastern part of the range of the species; in Montana the Bitterroot Valley is occupied, but not the surrounding highlands, and farther south, in Colorado and New Mexico, the mountains but in general not the lowlands are occupied. Probably less than ten per cent of even the more favorable parts of the region where M. montanus occurs is suitable for occupancy. The population density fluctuates not only seasonally, but ir- regularly from year to year. Although less noticeable, and less regular than the widely reported fluctuations in microtine popula- tions in the arctic, these fluctuations in temperate regions at times reach similar extremes. The total population of the species thus is discontinuously distributed, and is present in local populations that vary in size and in degree of isolation from other local popu- lations from time to time. Fluctuations are not always concurrent in different parts of the range of the species. For example, in August of 1954 Dr. Philip L. Wright of Missoula, Montana, in- formed me that in 1953 and 1954 he had not seen a Microtus mon- tanus although he had trapped in areas that in other years yielded numerous specimens. In Missoula County, on the night of August 24, 1954, 1 set 56 museum special mouse traps in a dry, grassy fence- row (2/2 mi. S, 3 mi. E Frenchtown) but captured no Microtus. Only 160 miles distant, 2^2 miles south-southeast of Pullman, Whit- man County, Washington, two nights later, 55 museum special mouse traps yielded 14 Microtus montanus. Animals distributed, in the above manner, over a large region 424 University of Kansas Publs., Mus. Nat. Hist. and subject to differences in environment may be expected to evolve so that local populations come to differ from each other. And, as there are degrees of isolation, population size, and fluctuations, the amount of difference between local populations would be expected to differ. This is the fact in Microtus montanus. Influence of Other Species Explanation of the distribution of M. montanus in terms of the distribution of a certain type of habitat does not account for both the occurrence of M. montanus at lower elevations in the Great Basin, and the absence of M. montanus at lower elevations west of the Sierra Nevada and east of the Rocky Mountains. Competition with closely related species having habits resembling those of Microtus montanus may affect distribution. This compe- tition has been discussed for the species M. montanus, M. pennsijl- vanicus, and M. ochro gaster hy Findley (1954:419) who concluded that each of these species, where occurring by itself, occupies a greater variety of habitats than where occurring with one of the other species. The segregation of two species, each in a different local habitat, where they occur in the same area was noted. Such ecological displacement supports the thesis that two species having identical ecological requirements cannot persist together in the same area (Matthew, 1930:272; Cause, 1934:113). Close observa- tion of two related species in an area where their ranges overlap, may reveal more clearly the difference in their requirements than will comparison of tlieir habits and habitat in areas that each oc- cupies alone. Closely related species are likely to have more nearly the same requirements than more distantly related species. Therefore, com- petition may be relatively direct between closely related species, and their ranges may tend to be mutually exclusive. A corollary of this relationship and of the theory that species evolving from a common ancestor ordinarily do not arise in the same area (Mayr, 1942:185), is Jordan's Rule that the nearest rela- tive of a species is found not in the same district, nor in a distant one, but in an adjacent one separated from the first by some barrier. There are nine species of the genus Microtus that occur at some place with Microtus montanus. Conversely, from the above con- siderations, it may be hypothesized that those species the ranges of which border on but do not broadly overlap the range of M. montanus are more nearly related to M. montanus and are more Anderson — Microtus montanus 425 nearly similar in their requirements than are those species the ranges of which broadly overlap the range of M. montanus. This hypothesis is considered below in the discussion of the ecological and distributional relationships between these nine species. Fig. 2. The geographic range of Microtus montanus (solid lines) and of six largely allopatric related species of Microtus (dashed lines) showing the part of the range of montanus also occupied by one or more of these other species (stippled areas). The six allopatric species are: M. townsendii, M. oregoni, M. califomicus, M. mexicanus, M. pennsylvanicus, and M. ochrogaster. Only two other species of Microtus, M. richardsoni and M. longicaudus, occur within the range of Microtus montanus but their ranges are not shown on this map. The seven species that are mapped tend to have mutually exclusive ranges. 426 University of Kansas Publs., Mus. Nat. Hist. The ranges of seven species of Microtus are shown in Figure 2. That part of the range of M. montaniis also occupied by one or more of the other species is indicated. These species are M. tawn- sendii, M. oregoni, M. californicus, M. pennsyhanicus, M. ochro- gaster, and M. mexicantis. Only two other species of Microtus, M. richardsoni and M. longicaudus, occur within the range of Microtus montanus but their ranges in contrast to those of the six named above, overlap the range of M. montanus, and are not shown on the map. Microtus oregoni, the creeping vole, occupies the humid region from within the Cascade Mountains westward to the coast and from southern British Columbia south through Washington and Oregon into northern California. The species is the smallest and the most fossorial of the voles within the geographic range of M. montanus. The creeping vole in British Columbia was said by Cowan and Arsenault (1954:198) to be characteristic of the forest edge. Dalquest (1948:358) states that in Washington the species occupies almost every habitat conceivable for a mouse, including wet marshes, damp ravines, dry forest, damp, mossy forest, meadows, alpine meadows and fields of short grass. The vole was said to be rare except in fields of short grass. In Oregon, Bailey (1936:212) reports that it occupies dense forests as well as brushy and open grassy areas of the uplands and dry slopes, and that it seems to avoid the marshes and wet bottoms. These ac- counts indicate that a wide range of habitats is occupied, that the optimum habitat is short dry grass, and that the vole is scarce in moist lowland habitats. It is significant that moist lowland is the optimum habitat of M. townsendii, the range of which almost com- pletely overlaps that of M. oregoni. The geographic range of the creeping vole, in Washington and Oregon, extends down the east slope of the Cascade Mountains to within 25 miles of the known range of Microtus montanus in at least three places. Also, in south- western Oregon and northern California M. montanus and M. ore- goni have been taken within 25 miles of each other. The Willamette Valley of Oregon and Clark County, Washington, on the north side of the Columbia River are the only areas west of the Cascade Mountains occupied by M. montanus. It is my im- pression that in the Willamette Valley M. montanus occupies eco- logically an area between the optimum habitats of M. townsendii and M. oregoni. A detailed ecological study in this valley would be of interest in order to determine if the presence of M. montanus Anderson — Microtus montanus 427 has restricted the creeping vole to the drier, higher, marginal grass- land and small grassy areas in forests and prevented M. oregoni from occupying also areas of more luxuriant grass in the valley. Such a finding would support my theory that where M. montanus and M. oregoni meet in Washington and Oregon the habitat, and per- haps to a lesser extent competition with M. oregoni, limits M. mon- tanus to the west. Owing to the greater variety of habitats occu- pied by M. oregoni than by M. montanus, I suspect that the presence of M. montanus east of the Cascade Mountains is more important than the habitat in limiting M. oregoni. Microtus towiuendii is distributed in the same coastal area occu- pied by Microtus oregoni. The ranges of these two species diflFer in that M. townsendii occurs on certain islands, including Vancouver Island and the San Juan Islands, where M. oregoni is absent. In the Cascade Mountains of both Washington and Oregon and in some parts of northwestern California where M. oregoni occurs, M. townsendii is absent. M. townsendii occupies less varied habitat than M. oregoni, preferring, as Bailey (1936:206) stated, wet ground densely covered by rank grass or tules in the low country. M. town- sendii occurs with M. oregoni and M. montanus in the Willamette Valley. In southwestern Oregon Townsend's vole and the montane vole occur within 25 miles of each other, but have not been re- ported from the same place, nor have their ranges been shown to overlap. The ecological relationships of these species in the Wil- lamette Valley have been mentioned above. On Vancouver Island, where M. townsendii is the only vole, it occurs not only in moist fields and sedge meadows of alluvial areas but to timber-line and above in alpine meadows (Cowan and Guiguet, 1956:221). Microtus calif ornicus, the California vole, occurs west of the Cas- cade-Sierra Nevada mountain chain from southwestern Oregon, through California, into northern Baja California. In Oregon, M. californicus lives mainly in dry upland meadows and on grassy slopes of the open valleys of the Upper Sonoran Life-zone ( Bailey, 1936:207). In California this vole inhabits environments diverse in both humidity and temperature (Kellogg, 1918:1). The sub- species M. c. paludicola, and in places M. c. aestuarinus are almost aquatic, and in other areas other subspecies inhabit wet or moist ground or seasonally dry hillsides in the Upper Sonoran and Transi- tion life-zones. In southern California the species was said to range without interruption from the tidal marshes of the seacoast to the boreal zone of adjacent mountain tops. Along the west slope 428 University of Kansas Publs., Mus. Nat. Hist. of the Sierra Nevada, M. californicus at lower elevations and M. montanus at higher elevations occur within 25 miles of each other at numerous places but the two species actually occur closely to- gether only in the Klamath and Shasta valleys in northern California and southern Oregon, in eastern Shasta County, California, and in the Owens River valley in Mono County, California. The occur- rence of M. californicus in boreal habitat in southern California where M. montanus is absent, and the occurrence of M. montanus in Sonoran habitat east of the Sierra Nevada where M. californicus is absent support the theory that competition between these two species limits their distribution where they meet. Microtus pennsylvanicus, the meadow vole, has the largest range of any species of Microtus in North America. Most of its range is north and east of that of M. montanus. Of the largely allopatric species mapped with M. montanus in Figure 2, M. pennsylvanicus overlaps more of the range of M. montanus than does any other species. Nevertheless, the extent of the overlap is less than is sug- gested by the rather inclusive range shown by Hall and Cockrum (1953:407). Areas where the meadow vole and the montane vole occur together are in British Columbia, in northeastern Washington, in the Bitterroot Valley and south-central Montana, in eastern Idaho, in Utah along the west face of the Wasatch Mountains, in northwestern Wyoming, on the east slope of the Big Horn Moun- tains, and along a narrow front in Colorado. In British Columbia the meadow vole occurs in a variety of habitats that provide an abundance of cover, usually near water, and in this same area the montane vole occupies arid grasslands of the sagebrush community and the grassland community altitudinally higher (Cowan and Guiguet, 1956:217, 220). In each of the other areas where these two voles occur together, M. pennsylvanicus is found most com- monly in denser cover, nearer water, and at lower elevations than M. montanus. Microtus ochrogaster, the prairie vole, meets the montane vole where the prairie meets the mountains in Colorado and Wyoming. In this region M. ochrogaster occurs altitudinally lower, and in drier grasslands. In the northern part of its range M. ochrogaster encounters M. pennsylvanicus and there yields the lower, wetter, more grassy areas to Af . pennsylvanicus. Microtus mexicanus, the Mexican vole, occurs in the mountains of central Mexico and northward through the Sierra Madre Occi- dental into the plateaus and mountains of Arizona and New Mexico Anderson — Microtus montanus 429 where it occupies much of the Transition Life-zone. Its northern hmits are in south-central Utah and southwestern Colorado. To my knowledge M. mexicanus and M. montanus meet only in Mesa Verde National Park, Colorado, and in the White Mountains of Arizona. The montane vole, in this part of its range, occupies a more restricted area than does the Mexican vole. The latter is often found in relatively arid, open, sparse grassland, sometimes in open parks in Yellow Pine forest, or in rocky areas, whereas M. montanus is likely to be found in areas aflFording more ample cover of grasses and sedges. The Mexican vole, nevertheless, is not averse to wet and densely vegetated habitat. At Apache Creek, New Mexico, east of the White Mountains of Arizona, in 1956, 1 captured M. mexi- canus, and no other species of vole, in the wet, dense, matted mix- ture of grasses, sedges, and cattails of several artesian bogs. At Upper Nutrias, in the Zuni Indian Reservation, farther north in New Mexico, I captured M. mexicanus, and no other species of vole, in dense, green grasses, sedges, and cattails in a moist, low area below the dam of the reservoir that retains spring waters for irrigation. In Mesa Verde National Park, Colorado, the Mexican vole has been taken at five places on the top of the mesa and in varying amounts of grass. I captured the montane vole in the park only on the floor of the upper part of one of the higher canyons in grass and sedges more dense than those at any of the five places where the Mexican vole was taken. It seems that the thesis advanced by Findley to explain the re- lationships ecologically of M. montanus, M. ochrogaster, and M. pennstjlvanicus is applicable also to the relationships of M. town- sendii, M. oregoni, M. californicus, and M. mexicanus to each other and to the above three species. In all cases except two, that is to say between M. townsendii and M. oregoni, and between M. penn- sylvanicus and M. ochrogaster, the above-named species are mutu- ally exclusive to the extent that 90 per cent or more of the range of each species is not occupied by any of the other species. Each species occupies a greater variety of habitats where it occurs alone than where it occurs with one of the other species discussed. And finally, where two species occur in the same area they usually ex- hibit ecological displacement. M. richardsoni and M. longicaudus, in contrast to the above species, occur with M. montanus but are not mutually exclusive in a geographic sense. M. richardsoni, the water vole, clearly dif- fers ecologically from M. montanus. In British Columbia, in the 430 University of Kansas Publs., Mus. Nat. Hist. Cascade Mountains and in the northern Rocky Mountains of Idaho and Montana the water vole occurs near streams in the mountains and the montane vole occurs in the valleys with the result that their ranges do not overlap. In the southern part of their ranges M. montanus becomes more montane and the ranges of the two species do overlap in northwestern Wyoming and south into Utah. The ecological separation persists however. For example at Green River Lake, Wyoming, in the Wind River Mountains, in 1951, we captured the water vole in pine forests, having little grass, along cold running streams, and we captured M. montanus at a slightly lower elevation, one half mile distant, in open grassy meadows and along streams below the lake. There, as well as elsewhere, no ap- preciable ecological displacement results. The two voles normally occur in different habitats. Microtiis longicaudus, the long-tailed vole, occurs throughout the range of the montane vole except in a few places. In many places they live in the same local areas. The most obvious differ- ence in the ranges of the two species is that M. longicaudus is more widespread, extending beyond the limits of the range of M. mon- tanus on all sides, on the north reaching into Alaska, on the west reaching to the Pacific Coast from Washington to California, on the east reaching to the Black Hills of Wyoming and the Sangre de Cristo Mountains of Colorado and New Mexico, and on the south reaching to the San Bernardino Mountains of California, the plateau area of north central Arizona, and numerous scattered mountain ranges in southeastern Utah, and in New Mexico. The larger range may result from the occupation of an equally varied but more wide spread habitat than that required by M. montanus. The long-tailed vole seems to occupy a greater variety of habitats than does the montane vole; my own experience, as well as the reports of many naturalists, indicates that it does so. The tvvo most notable places in which A/, montanus, but not M. longicaudus, occurs, are the Willamette Valley of Oregon and the vicinity of some local springs in the desert of southern Nevada. In north- western Oregon M. longicaudus is not abundant and frequents the mountains rather than the valleys. In British Columbia Cowan and Guiguet (1956:224) state that the habitat of M. longicaudus is "extremely variable" but usually is the edges of forests where the long-tailed vole seldom makes runways and seldom lives in large colonies. In Washington, Dalquest (1948:352) reports that this vole has been taken in many habitats, along a small stream through Anderson — Microtus montanus 431 sagebrush in the Upper Sonoran Life-zone, in marshes, near water, in forest-free, grassy places in Canadian and Hudsonian hfe-zones, near talus slides, and from sea level up to 6000 feet. The species is said to be rather rare, and not ordinarily to make trails, resulting in specimens being taken unexpectedly. I have not always found this last observation to be true elsewhere. In Oregon, according to Bailey (1936:208), the long-tailed vole was partial to creek banks and cold water in the mountains, but remained in grassy places that thoroughly dried out in summer. The habit of seasonally ex- tending its range in winter under cover of snow was noted. This habit has been described also by Linsdale (1938:193) in Nevada. In Idaho (Davis, 1939:319) the species has been trapped in sage- brush in the boreal zone, near water in thickets, in boggy areas, in a rock slide at 8500 feet in association with pikas, at the bases of willow clumps, and at the edges of streams. In Nevada Hall (1946:550) reported that M. longicaudus prefers herbacious vege- tation as much as grass and does not make well defined runways, in contrast to M. montanus that ordinarily lives in grass and makes well defined runways. In addition the long-tailed vole was said to be less restricted to the vicinity of water and to occur altitudinally higher than the montane vole, Jenkins (1948:66) studied M. mon- tanus and M. longicaudus in three meadows in the Sierra Nevada of California, in three summers. M. longicaudus was less abundant, less restricted to meadows, less variable in numbers, and individuals occupied larger ranges than M. montanus. The occasional flooding of meadows was thought to contribute to the fluctuation in num- bers of M. montanus. These reports and others depict a Microtus that is more varied in habitat requirements, more widely ranging geographically and zonally, less closely associated with grass, and less likely to construct runways than any of the other species dis- cussed above. This greater vagility and tolerance accounts for the ability of M. longicaudus to exist sympatrically with these other species, among themselves largely allopatric. That this ecological distinctness may be correlated with taxonomic distinctness will be considered later. The reports of naturalists and knowledge of distribution of species of Microtus based on specimens provide data that support the hypothesis that competition is an important limiting factor. The interactions involved in this "competition" are not usually apparent, however. Competition may be direct; for example when the amount of available shelter becomes critical in a local area actual 2—7800 432 University of Kansas Publs., Mus. Nat. Hist. strife may ensue. Competition may be less direct; for example the depletion of a supply of food by one species may limit another species dependent on this same food, although individuals of the two species may seldom meet directly. Or, behavioral responses to environmental factors or other species may preclude the more direct or extreme forms of competition. In any case, knowledge of distribution makes it clear where later field studies may profitably be carried out, and knowledge of general ecological relations indi- cates behavioral and physiological factors that need more detailed analysis in the laboratory and in the field. Form and function are not unrelated; nevertheless, the adaptive or functional values of characters used taxonomically are often not clear. In other cases relationships may be easily postulated; for example the morpholog- ical fact that one species of Microtus has on the average four teats and another species has six may be directly related to a difference in the number of young produced in the average litter, and this in turn to their relative vulnerability to predation. VARIATION Non-geographic Sources Sex, age, season of capture, and methods of sampling are the most obvious sources of variation within a species other than "indi- vidual" variation and geographic variation. Individual variation is simply the variation that remains when the easily ascertainable sources have been minimized. When geographic variation is to be analysed, as in many taxonomic studies, these other sources must be understood, and not allowed to influence the assessment of geographic diflPerences. In many animals sexual dimorphism in size, color, or other char- acters that may be of taxonomic importance is evident. This is not true of Microtus montanus. To determine if more subtle dif- ferences existed in characters being considered for taxonomic pur- poses, males and females were compared in two series from Carbon County, Wyoming. The numbers of adult males were 29 and 23, and the numbers of adult females were 14 and 25 in these two series. Three external measurements and 23 cranial measurements were taken. The coefficient of variability was greater in males than in females in 27 of 46 comparisons of cranial measurements and in 5 of 6 comparisons of external measurements, and the males averaged larger in most measurements, but not significantly larger. Anderson — Microtus montanus 433 That males in microtines may be slightly more variable than females and/or on the average slightly larger has been noted by several authors, including Hall (1946:544), Coin (1943:223), Sny- der (1954:208), and Howell (1924:998). This difference in size between the sexes is less than the difference owing to any of the other sources discussed here, and therefore it seemed advantageous for statistical reasons to use the larger samples gained by including both sexes for taxonomic comparisons. In the study of geographic variation, the changes that occur with increasing age in Microtus montanus are perhaps the most troublesome. Young are produced throughout a large part of each year, with the result that most samples include voles of various ages. Age-classes such as are readily recognizable in species that have a single, short breeding season are not distinct in Microtus montanus. The attainment of sexual maturity is not a useful criter- ion, inasmuch as breeding commences in some individuals when they have not attained one half of their adult weight. In some species of mammals the eruption of a certain tooth, the appearance of wear on certain cusps of the teeth, or the fusion of certain sutui'es in the skull provide convenient landmarks for segregating age- classes. In M. montanus the teeth are persistently growing, lack enamel crowned cusps after early life, and the skulls continue to grow throughout adult life, although the rate of growth then de- creases. In order to take into account this continuous change with growth two methods were devised. The "method of pairs" (previously described by Anderson, 1956: 90 ) was designed for using cranial characteristics that are not easily measureable, but the method can be used for other characters. Comparisons in this study were of the structures hsted when the method was described, unless other structures are mentioned. Com- paring individuals of carefully matched pairs of two series rather than comparing averages of the two series minimizes bias from unrecognized differences in age. The second method was use of the "index of maturity," Different measurements do not change at the same rate, as I have illustrated with selected measurements in Figure 3. Proportions change with increasing age, and these changes continue throughout the range of sizes that may be considered as "adults" for taxonomic pur- poses. By selecting measurements that remain relatively constant and by expressing these as percentages of measurements that in- crease much, a proportion is derived that can be used as a measure 434 University of Kansas Publs., Mus. Nat. Hist, of maturity. The index of maturity (see Figure 4) is the sum of the lambdoidal breadth and the zygomatic breadth divided by the sum of the prelambdoidal breadth and of two times the interorbital breadth. Both the interorbital breadth and the prelambdoidal mm 18 20 22 CONDYLOBASILAR 24 LENGTH 26 mm. Fig. 3. Measurements of the skull and the length of the hind foot (all in milHmeters) in a population of Microtus montanus tiantis. Each point represents the average for ten individuals. In any given group of ten individuals the condylobasilar length varies no more than 1 mm. if the average is less than 20 mm., and no more than Vi mm. if the average is more than 20 millimeters. The rates of increase in the measure- ments shown differ even in adulthood. It is evident that averages and ratios between measurements may differ in samples from the same adult population. Such differences must be considered in the study of geographic variation. The adults all are from Carbon County, Wyoming; younger animals also are mostly from there, but a few individuals are from nearby counties in Wyoming and Colorado. Anderson — Microtus montanus 435 breadth ( see Figure 3 ) change relatively httle. These are measure- ments of parts of the skull associated with the brain; it attains nearly maximum size early in life. The rostral part of the skull and those parts associated with developing musculature, such as the 2ygo- matic arches and the lambdoidal, paraoccipital and other crests con- tinue their development and attain their greatest size late in hfe. The index of maturity increases as the voles become more mature. Change in the index of maturity with increasing size in three sub- species is shown in Figure 4. In that figure, average values for successive age-classes do not differ significantly in any case, owing to the amount of variation in individuals, but the trends are mean- ingful. The index of maturity was calculated for the samples indicated in Table 2. Assuming, from inspection of Figure 4, that, on the average, an increase in the index of maturity of 0.1 unit is equivalent to an increase in condylobasilar length of 1.3 mm., the size at which 25 26 27 28 29 CONDYLOBASILAR LENGTH OF SKULL IN MILLIMETERS Fig. 4. The increase in the Index of Maturity (explained in text) with increasing size in three subspecies of Microtus montanus. Dots represent the subspecies undosus (District 24 ) ; circles represent amosus ( District 66 ) ; and circles enclos- ing smaller dots represent nanus (District 46). Symbols rep- resent averages of groups of specimens. The condylobasilar length in each group has a miximum variation of one milli- meter. The subspecies that average larger in size reach a given degree of maturity at larger sizes. The indices substantiate the diflFerences observed in average size. The nvmiber in each size-class of each subspecies is shown by the numerals below the symbols. 436 University of Kansas Publs., Mus. Nat. Hist. a certain average maturity is reached can be calculated. The calcu- lated size at the 1.75 index of maturity was used in comparisons cited in accounts of various subspecies. DiflFerences in size between voles from diflFerent places need to be evaluated, both in subspecific comparisons and in the testing of ecological rules such as Bergmann's Rule. After considerable study I concluded that for use in comparisons involving averages, the simplest and least objectionable basis on which to separate adults from less mature individuals was a single measure of the length of the skull, the condylobasilar length. Individuals less than 24.0 mm. in condylobasilar length, or if this measurement was not available those of less than 140 mm. in total length, were not included. The range above 24.0 mm. is designated as "adults" in Figure 3. Indi- viduals of larger subspecies in the lower part of this size-range may be relatively less mature than corresponding individuals of smaller subspecies. The greater differences in average size will be apparent despite this bias and the evaluation of lesser differences will not be subject to the bias that may be introduced unknowingly if more subjective criteria are used to select series for comparison. I considered this latter bias more hkely to lead to a serious error than the former. Also the index of maturity provides a means use- ful in detecting the former bias. The composition of a population, discussed in connection with sampling below, or individual animals themselves may vary with the season. Seasonal differences, except in pelage, have rarely been considered important in taxonomic comparisons, and the date of capture is not traditionally included with lists of specimens examined. The length, thickness, and color of pelage vary with season. This variation is greater in arctic than in temperate regions and is greater in temperate than in tropical regions. Important differences between the skulls of Microtus miurus in Alaska that grew up at different times of the year have been described by Bee and Hall (1956:147). Individuals born in winter grew more slowly than, and never so large as, individuals bom in spring; also, in grow- ing, those bom in the two mentioned seasons developed different proportions in the skull. This difference was attributed to more favorable conditions of food, light, and climate in spring and sum- mer. The extremes observed in the arctic do not occur in temperate regions. Nevertheless, within the range of M. montanus, local dif- ferences in food and climate may cause some morphological varia- tion. As mentioned above in the discussion of the index of maturity, Anderson — Microtus montanus 437 individuals of the same size in any population vary as to morpho- logical maturity. It is fortunate, for purposes of minimizing seasonal differences, that most specimens of the montane vole have been taken in summer. Most are taken then because collecting is easier at that time and because the schedules of many collectors allow them to do field work only in that season. Methods of sampling may cause diflFerences between samples drawn from the same population at the same time or at diflFerent times. The composition of a population varies in the relative abundance of different age-classes. A sample taken in spring may contain mostly old adults, and a sample taken in autumn may con- tain mostly young adults and immature individuals. If the time of year influences the course of development, as noted in M. miurus above, then the time of year a sample was taken may need to be considered. If climate and abundance of food in a given year in- fluence growth, many individuals will be stunted in a poor year, and the year of capture and the local conditions in that year may need to be considered. The responses of individuals of different sex, age, experience, or place in the social system may vary with types of traps and bait used, and time of setting and checking traps. If larger, more dominant males are more active than smaller, less dominant males, or vice versa, the sample of males may be biased. Another source of bias is the collector. Suppose that he traps twice as many specimens in a given day as he can prepare for preservation. If he discards all the smaller animals, the sample is biased toward the larger sizes. This bias can be evaluated if well kept notes of the collector are available, but oftentimes no such notes are available. All of these possible sources of bias obviously cannot be eHminated. The presence of such influences may ac- count for some of the variation observed. Some of these variables, as has been noted, can be eliminated, and the importance of the others in the over-all geographic picture of variation is diminished because many of the differences observed betwen populations oc- cur between numerous pairs of samples taken at different places, in different years, by different collectors, and under different climatic conditions. Geographic Variation The impression may be gained by viewing the numerous locali- ties recorded in figures 8 through 11 that ample material is avail- able for the study of geographic variation. This favorable impres- sion is somewhat modified by noting in the lists of specimens that 438 University of Kansas Publs., Mus. Nat. Hist. 117 107 50 45 40 35 . ^MH 1 \ \ N 50 rr \ 36 o42 I (i^-24 ^\ 062^64 §!?5_^Pv 58^ ^ -p;; r I 7'a 72 ^1 \r-- .__.J A- 45 40 35 L 117 107 Fig. 5. Map showing districts into which adult Microtus montanus were grouped for convenience of study. See Table 1; see Figure 6; and see discus- sion on pages 442, 449, 452, and 453. The boundary of any given district was drawn insofar as possible to include an adequate sample of adults of only one subspecies without crossing any major physiographic barrier, or any major hiatus in the knowTi range. numerous localities are represented by fewer than five specimens, and the further realization that the numbers listed include on the average at least 40 per cent that are immature, damaged or lack complete data and that therefore cannot be used in comparisons. A single correctly identified specimen, of any age, provides proof of the occurrence of a species, but assessment of geographic varia- Anderson — Microtus montanus 439 tion in most instances requires a series of specimens of comparable maturity. For example, in two series containing 20 and 34 speci- mens the careful matching process for a comparison of pairs of skulls might yield seven pairs. The more subtle the differences studied, the larger the series required. Geographic differences in size have been used in the descrip- tions of most subspecies of M. montanus. The correlation of size and also the lengths of extremities with latitude, temperature, and altitude has been the subject of various ecological rules. Berg- mann's Rule, for example, states that homoiothermal animals from colder, more northern climates tend to be larger than their relatives from warmer climates. Largeness results in less surface in pro- portion to weight and is thought to be a mechanism for heat con- servation. In order to obtain adequate samples from all parts of the range it was necessary to group specimens from more than one locality. Localities yielding usable specimens were accordingly arranged in the 72 districts shown in Figure 5. These districts were so delimited that: usually at least 10 adults were included; a single district did not cross a major physiographic or ecological barrier to the species; and districts did not cross boundaries of subspecies as understood to date. Averages, extremes, and other data for ex- ternal measurements derived for each of these districts are included in Table 1, and these data form the basis for Figure 6. The number of adults varies from 3 to 101 (beyond which point additional data were not included even though available) and averages 36 per district. The size of sample and, for selected districts, the standard deviations are included so that the reader can, if he wishes, evaluate differences for himself. To test the applicability of Bergmann's Rule to M. montanus the average length of the head and body, derived by subtracting the average length of the tail vertebrae from the average total length, for each district listed in Table 1 was plotted against two values in Figure 7. These two values are the average annual temperature and the "equivalent latitude," that is to say, the latitude corrected for altitude. The average annual temperature for each district was estimated from the maps and data in the Yearbook of Agri- culture, 1941, entitled Climate and Man (Hambidge, et al., 1941: 663-1210). If most specimens were from one part of a district, this was taken into account in making the estimate. Average tem- peratures for January were estimated and also plotted against size, but no better correlation was observed than with average annual temperatures. The microclimate where the voles live almost cer- 440 University of Kansas Publs., Mus. Nat. Hist. tainly differs from the crude estimate plotted, but the estimate is based on the best data now available. In order to test the validity of the observed relationship of size to temperature, the estimated average latitude for specimens in the sample representing a given Fig. 6. Map showing the average length of head and body (collector's total length less length of tail), and the average length of the hind foot and of the tail. The length of the hind foot and the length of the tail are expressed as percentages of length of head and body, for each of the districts shown in Figure 5. The larger size of mice from the southern part of the range of Microtus montanus than from the northern part, the relatively larger feet of mice from the southwestern part of the range than from other parts, and the random pattern throughout tne range in relative length of tail are apparent. Other trends and local differences are discussed in the text. Anderson — Microtus montanus 441 district was plotted against size. The latitude was "corrected" by assuming that each 5000 feet in altitude was equivalent to a north- ward displacement in latitude of 16.3 degrees. The correlation of size with temperature and latitude is not close, but in general larger voles occur in areas of higher temperatures and lower latitudes. But also some of the smallest voles (see Figure 7) occur in warmer areas of lower latitudes. The variation in size as shown with these data and as mapped in Figure 6 demonstrates that in M. montanus the trend is toward a reversal of Bergmann's Rule. Allen's Rule states that the extremities of warm-blooded animals in colder climates tend to be shorter than the extremities of their UJ 1 1 1 ' 1 1 o: 3 1- < • cr ^60 - • - S UJ t- • _j .• • • <50 3 — • •• •••.•••»•:•• • • ^. - Z • • Z < • •• • •• • J^40 _ •« ••* : • • ^ < V* •• cr • • • UJ > < - • \ • • / UJ__ \ o50 — \ • • • • • / — Z3 • ^ \t • • / • 1- < -J ,_60 X • • • •• •^ • • z \. 1^-^* • • UJ -J ^ >-rrr • • < • •••• 1^0 — •• • — UJ 1 • • 1 » no LENGTH OF 115 HEAD 120 AND BODY 125 130 IN MILLIMETERS Fig. 7. The size (average length of head and body) of Microtus montanus representing districts shown in Figure 5 related to the average annual temperature and to the "equiv- alent latitude" (the actual latitude corrected for altitude, as discussed in text ) . The curve was drawn by eye, not computed. 442 University of Kansas Publs., Mus. Nat. Hist, relatives in warmer climates. To test this hypothesis in M. mon- tanus an index was derived by expressing the sum of the average length of the tail and of the hind feet as a percentage of the length of the head and body for the voles in each district. This index was plotted against mean annual temperature and equivalent latitude as was done with size in Figure 7. No significant correlation or trend was apparent. Size of the montane vole in different parts of its range and rela- tive lengths of its tail and hind feet are represented graphically in Figure 6. It may be noted that the smaller voles occur in the north- ern part of the range and the larger voles occur in the southern part of the range, especially in parts of southern Nevada and Utah. As has been discussed, the area where the voles are larger is, in general, the warmer, more southern, and altitudinally lower part of the range, and also the part of the range where there is little competition from closely related species. In the relative length of the hind feet the populations in the western part of the range on the average exceed those in the eastern part. In contrast to length of hind feet and size, there seems to be no major geographic trend in length of tail. In addition to clinal geographic variation Figure 6 illustrates similarities and differences in local populations. For example the two samples from north- western Oregon, the two samples from British Columbia and central Washington, the two samples from west-central Nevada, and the two samples from eastern Arizona are in each case alike in all three characteristics. In other cases adjacent populations differ notably in one or more characters, for example the tsvo southern- most samples in Nevada and the two samples in central-northern Wyoming. In most of the geographic range the differences ap- parent in the three external measurements illustrated are neither so uniform, nor so different as in the examples mentioned above. The varying degrees of distinctness between populations seen in these characters are seen also in most other characters studied. A comparison of the ranges of subspecies shown in Figures 8 through 11 with the examples noted above in Figure 6 will reveal that the uniform, adjacent populations are included in one subspecies, and that distinct populations are included in different subspecies. Nevertheless, the majority of populations cannot be so clearly delineated and boundaries of subspecies and differences shown in external measurements agree less precisely. There are two reasons for this. Firstly, characters of the skull and of the color Anderson — Microtus montanus 443 of the pelage are used in addition to external measurements in distinguishing subspecies; and not all characters could be con- veniently included in the figure. Secondly, many degrees of dif- ferences do exist between local populations within an entity re- garded by me as a subspecies. The differences of lesser degree or of lesser geographical uniformity are disregarded in comparing subspecies. These are the differences said not to be taxonomically significant. Other than in size and accompanying characters I observed no geographic differences in the skull that extend throughout the range of the species. The lesser differences noted are included in the accounts of subspecies. The weights of 1058 adult Microtus montanus are available. Of these, 964 are from 30 districts from each of which ten or more adults of known weight are available. Considering the variability in weight, at least ten in each sample were judged to be necessary for averages to be compared meaningfully. The data on these 30 districts have been summarized in the accounts of the seven subspecies concerned. Twenty-three of the 72 districts were repre- sented by no adults of known weight, and 19 districts by fewer than 10 (average 4.7). Unfortunately, data concerning weight are inadequate for most of the subspecies, and for more than half of the adults examined. Certain variations in color occur in more than one subspecies; for example, three adjacent subspecies are blacker than surrounding subspecies. Differences in color are described in the accounts of subspecies. Microtus is not noted for its brilliant colors nor does it have streaks, spots, or other embellishments seen in some mammals. To the casual observer, the coloration of Microtus is dull and relatively uniform. The most noticeable variations in color are degree of pigmentation and admixture of certain hues, primarily reddish or yellowish. Brown is a dull yellow or orange and is thus included in the hues mentioned. The first of these variables, paleness (slight pigmentation), also has been referred to by the perceptual term dullness and the physical terms luminosity or reflectance. Two of the major contributors to color nomenclature have used "value" (Munsell, 1916:18) and "tone" (Ridgway, 1912:17) for this same variable. The second of the variables important in Microtus involves hue and saturation. A comparison of a yellowish and a reddish color is an evaluation of hue, and a comparison of 444 University of Kansas Publs., Mus. Nat. Hist. a "pure" yellow with a less brilliant yellow (yellow mixed with other hues), is an evaluation of saturation. As Bowers (1957:159) has pointed out in discussing methods of color determination and their application to a study of birds, wren-tits, paleness is the variable that is perhaps most easily judged by eye. In an earlier study of variation in the color of Microtus pennsyl- vanicus (Anderson, 1956:88) a method of comparison was devel- oped primarily for use in evaluating paleness. This method had the advantage of comparing specimens with specimens, rather than with some printed, painted, or otherwise concocted standard. The method also attached numerical values that could be treated sta- tistically and mapped for convenience in visualizing the pattern of geographic variation. The method is in essence like that ar- rived at by Bowers (1957:158) but differs in being less refined. This difference in refinement ( I differentiated 5 stages, as compared to Bowers' 9) is, in part at least, owing to the kind of animals being studied. The range of variation in color in the species Mi- crotus montanus is comparable to the range in the Microtus penn- sylvanicus mentioned above. It was therefore possible to use the same procedure, including even the same standard specimens used in the study of M. pennsylvanicus. The method was used when needed but not throughout the study of M. montanus. Color of the montane vole varies as follows. Blackishness is most pronounced in the southwestern part of the range of the species, and especially in southern Nevada. The blackish color becomes more diluted with paleness and/or various hues of color to the north and east. This blackishness is most prominent, although not uni- form, in the ranges of the subspecies M. m. nevadensis, M. m. fu- cosus, M. m. undosus, M. m. micropus, and M. m. montanus. The more northern of these subspecies, the three listed last, are paler; consequently grayish rather than blackish is perhaps an appropriate term; however they are not so gray as more northern subspecies and lack, in general, the reddish or yellowish hues that characterize populations in other parts of the range of the species. The black- ishness of the southwestern populations is often more noticeable in young than in adults, and this is not simply the more blackish color to be found generally in young, or animals with worn pelage. The palest populations are northern, in the subspecies M. m. canescens, M. m. pratincole, M. m. codiensis, M. m. zygomaticus, and to a lesser extent M. m. nanus. The last-named subspecies is the most nearly central, the most widespread, and the most varied in color as well as the most varied in other characters. Anderson — Microtus montanus 445 An admixture of yellow with the basic dull blackish color has produced a brownish color in one peripheral subspecies, M. m. dutcheri, of the southwest, and in another peripheral subspecies, M. m. canicaudus, of the western part of the range of the montane vole. Southeastern populations of the species show varying degrees of reddishness. These populations belong to the subspecies M. m. rivularis, M. m. amosus, M. m. fusus, and M. m. arizonensis. The populations of the species that are more abruptly set apart from adjacent populations, that differ most from adjacent populations, and that at the same time are most nearly uniform in color locally, are peripheral. The nature of subspecies has been discussed at length by many authors. In the periodical "Systematic Zoology" the matter has been discussed recently by Smith and White (1956:183), and it seems unlikely that the last word has been said. The use of tri- nomens in this study warrants some comment, although a thorough review is not here called for. The basic diflBculties in the formula- tion of a clear concept of a single taxonomic category beneath the rank of a species, in the present instance a subspecies, arise be- cause the name is static and the animals are dynamic. Confronted with this problem authors have either tacitly or in some cases with adequate elucidation proceeded along different paths. These paths have varied from that leading to advocacy of a complete abandon- ment of the trinomen to that leading to the use of the trinomen for minor and sometimes even abnormal variants. A subspecies of one author is not equivalent, therefore, in the degree of its mor- phological distinctness, in its uniformity, or in the certainty with which individual animals can be correctly recognized and assigned, with a subspecies of another author. This inconsistency results from differences in the points of view of authors, but more im- portant it results from differences in the animals concerned. Some species are more varied than are others. Pocket gophers of the genus Thomomys are more varied than are hares of the genus Lepus. An author studying pocket gophers may incorrectly gain a reputation for having a different point of view from another author studying hares, when actually their criteria for recognition of subspecies may be the same. Also one species, Microtus mon- tanus as an example, may exhibit varying degrees of distinctness in different parts of its range. It is therefore impossible even for a single investigator, using previously established criteria, to des- ignate subspecies that are equivalent. If the impossible is not demanded, or expected, the concept of a subspecies can be seen 446 University of Kansas Publs., Mus. Nat. Hist. more easily for what it is, simply a nomenclatorially useful device for designating geographic variation in animals. The subspecies is not the only useful device in such studies, and when not useful the concept need not be used. In my opinion both the concept and the terminology have been useful in the study of Microtus montanus. A subspecies, as here used, is considered to occupy one continuous geographic range, to intergrade with adjacent subspecies, and to have morphological characters by means of which upwards of 75 per cent of adult in- dividuals from most parts of the range can be correctly assigned on morphological grounds. The conservative approach to more doubtful cases has been to retain the previously used classifica- tion unless I felt that I had more conclusive evidence than did previous authors. A subspecies, accordingly, has both morphological and geo- graphical identity. If the geographic area occupied by a popula- tion is smaller, greater morphological distinctness from other named populations should be required before the population is named as a subspecies. If a population being considered occupies a greater geographic area the degree of morphological difference need not be so great if uniformity is apparent. Uniformity here refers not to a single character, but to the complex of characters distinguishing the subspecies. These considerations result from the pragmatic need for subspecies that can be recognized, and from the fact that, statistically speaking, larger samples and smaller standard deviations make differences more demonstrable and make smaller differences more significant. It is easier to make rules than to apply them, and the numerous variables that need to be evaluated in studying geographic variation preclude simple statistical analysis. It is, for example, easy to con- clude that the difference in the average length of the hind foot between two populations is, or it not, significant statistically, but it is not so easy to conclude that the two populations are, or are not, significantly different taxonomically, that is to say whether they belong to different subspecies or not. The degree of morphological difference characterizing geographic units of M. montanus varies from one unit to another. It follows that the number of subspecies recognized by a person depends on the degree of morphological difference selected as a minimum. By choosing successively higher degrees as the acceptable minimum, M. m. caryi can be synonymized with M. m. nanus as Anderson (1954:493) did; M. m. nexus can be synonymized with nanus, and Anderson — Microtus montanus 447 yosemite with montanus, as is done beyond. If one continued, fusus would be synonymized with nanus; amosus with rivularis; pratincola with nanus; codiensis with nanus; dutcheri with mon- tanus; zygomaticus with nanus; fucosus with nevadensis; and arizonensis with rivularis. At this stage there would remain eight subspecies, micropus, canescens, canicaudus, nanus, montanus, un- dosus, nevadensis, and rivularis. To continue, one might then com- bine montanus, micropus and nanus ( under the oldest name, M. m. montanus); canescens with m,ontanus; undosus with montanus; canicaudus with montanus; nevadensis with rivularis, and finally rivularis with montanus. The exact order in which the units should be combined could be debated, but not the existence of some order. The relatively greater distinctness of peripheral subspecies and the relatively smaller ranges of these peripheral subspecies in com- parison to subspecies in the central part of the geographic range of the species are apparent at each level in the above process of combining units. The high degree of morphological difference and small geographic ranges of these peripheral units are of evo- lutionary import and not accidental results of the taxonomic history of the species. Accounts of Subspecies In the synonomy the original description is cited first. The second entry cites the first use of the name combination here used, and other references follow. In the list of specimens examined, localities are arranged from north to south first by countries, then by states, by counties, and by geographic position. Localities that are not mapped in figures 8 through 11, because undesired crowding of the symbols would re- sult, are in italic type. Accounts of the subspecies are arranged in general from north to south and from west to east so that accounts of adjacent, and therefore usually closely related, subspecies are together. Microtus montanus canescens Bailey Microtus nanus canescens Bailey, Proc. Biol. Soc. Washington, 12:87, April 30, 1898; EUiot, Field Col. Mus., 115:291, February 9, 1907; Taylor and Shaw, Occ. Papers Chas. R. Conner Mus., 2:24, December, 1929, in part; R. M. Anderson, Nat. Mus. Canada, Bull. 102:157, January 24, 1947. Microtus montanus canescens. Hall, Proc. Biol. Soc. Washington, 51:133, August 28, 1938; Hall and Cockrum, Univ. Kansas Publ., Mus. Nat. Hist., 5:413, January 15, 1953; Cowan and Guiguet, British Coliunbia Prov. Mus., Handbook No. 11:220, July 15, 1956. Microtus canicaudus, Bailey, N. Amer. Fauna, 17:32, June 6, 1900, one specimen ( 89357 US ) from North Yakima, Washington. 3—7800 448 University of Kansas Publs., Mus. Nat. Hist. Fig. 8. The distribution of Microtus montanus in British Columbia, Washing- ton, Oregon, and western Montana and Idaho. Dots indicate localities from which specimens have been examined. The place of this region in North America is shown on the inset map in Figure 9. The heavy hnes outline the probable ranges of the following subspecies: A. M. m. canescens D. M. m. nanus B. M. m. pratincola G. M. m. montanus C. M. m. canicaudus J. M. m. micropus Anderson — Microtus montanus 449 Type. — Adult male; No. 90577, U, S. Nat. Museum, Biological Surveys Collection, from Conconully, Okanogan County, Washington; obtained by J. Alden Loring, original number 4654, September 12, 1897. Range. — British Columbia and Washington; lower altitudes in the Yakima, Columbia, Okanogan, and Frazer river-valleys (see Figure 8). Distinctive characters. — Size small; color pale, grizzled grayish; hind feet relatively large; braincase broad relative to its length; bullae large relative to size of skull; prelambdoidal fenestrae broad relative to length. See Table 1 for external measurements and Table 2 for cranial measurements. An index of maturity of 1.75 is reached at 24.7 mm. condylobasilar length in District 1, and at 25.6 mm. in District 2. Comparison. — In small size canescens differs from all other subspecies ex- cept nanus, fusus, pratincola and canicaudus. In color canescens differs from all other subspecies, being approached most closely by nanus of which both scattered individuals and some local populations are pale grayish and little suffused with the reddish, blackish, or yellowish so common in southern subspecies. Six pairs of skulls representing canescens ( District 1 ) and nanus ( from within a radius of 6 miles of PocateUo, Idaho) were compared; in 5 of 6 pairs canescens has broader, less constricted, prelambdoidal fenestrae and larger bullae; in each of the 6 pairs canescens has a broader braincase. Twelve pairs of skulls representing canescens (District 1) and nanus (from within 4 miles of Pullman, Washington) were compared; in 9 of 11 pairs (one skull has badly broken bullae) canescens has larger bullae; in 10 of 12 the braincase is broader in canescens. Nine pairs of skulls representing canescens (District 1) and canicaudus (Districts 7 and 8) were compared; in 7 of 9 pairs canescens has broader jugal plates and smaller auditory meatuses; in 8 of 9 the incisive foramina are more constricted especially in the maxillary part in canescens. Six pairs of skulls representing two popula- tions of canescens were compared; in 5 of 6 pairs the specimens from the Okano- gan River Valley have more open incisive foramina and smaller bullae than specimens from the Yakima River Valley. In comparison with those of nanus (from within 6 miles of Pocatello, Idaho, and within 4 miles of Pullman, Washington) skulls of canescens (all series) in relation to length are broader across the zygomata (see measurements). Differences, other than the above, which have been cited by Bailey (1898:88 and 1900:31), were not apparent to me in the specimens examined. Specimens from the Yakima River Valley are less gray than specimens from farther north. In this respect they are intergrades between canescens and nanus, but more nearly like canescens. Specimens obtained by Dr. Murray Johnson from the Grand Coulee are plainly referable to canescens. Two from one mile east of Bluestem, Washington, are intergrades. In color one resembles canescens, but is less grayish than the specimens from the Grand Coulee. The other is darker, but not so dark as nanus (to which they are referred) from Blaine County, Idaho. Some specimens from the range of canescens resemble other subspecies; for example tlie brownish specimen from North Yakima referred by Bailey (1900:32) to canicaudus, has been commented on by Dalquest (1948:349) and Hall and Kelson (1951:78). 450 University of Kansas Publs., Mus. Nat. Hist. Remarks. — Taylor and Shaw (1929:24) record specimens of Mi- crotus montanus from Bumping Lake, Yakima County, Washington, which have been cited by Hall and Cockrum (1953:413). I have I 12 107 T 0 100 '■■' ■■ ■ ■ SCALE OF MILES 50 /\ ^ "1 't^T'I -U--\ ' -7 Sr) X '•• •i|— .jVZ eA— ■> -u-4:k 1 .J \ \ \ ■ " 1 — - — ■ ^ J J MJ-1 } LI. \ . Mr7i_Ar-.A^*^^c^^-' 112 107 Fig. 9. The distribution of Microtus montanus in central Montana, eastern Idaho, and in Wyoming. Dots indicate localities from which specimens have been examined. The inset shows the range of the species in black and the regions shown in Figures 8 and 9. The heavy lines indicate probable ranges of the following subspecies: D. M. m. nanus E. M. m. codiensis F. M. m. zygomaticus Anderson — Microtus montanus 451 not examined specimens from this locality, do not know the where- abouts of specimens from there, and therefore have omitted the locality from my distribution map. The northernmost specimen was reported as "Cariboo," British Columbia, by Elliott, 1907:291, and has been overlooked by later authors. The original label bears the notation "Frazer R. 40 miles above mouth of Chilcotin R." The specimen (8201 CM) is an immature specimen and has unusually small bullae for M. m. canescens, but in other regards is clearly canescens. A specimen (2247 PS) from 18 miles east of Yakima is unusually pale and reddish. Specimens examined. — Total number, 176. British Columbia: Cariboo, Frazer R. 40 mi. above mouth of Chilcotin R., 1 CM; Westwick Lake, 12 mi. W of Williams Lake, 1 BC; Lac du Bois, 11 mi. NW Kamloops, 3 BC; Kam- loops, 1 MV; GHmpse Lake, 20 mi. NE Nicola, 1 BC; Ducks, 2 AM; Vernon (3 localities within 3 miles), 4 MV, 11 US; Okanagan Landing (=:Okanagan), 3 KU, 21 US, 10 PM, 1 CM; % mi. SE Okanagan Landing, 1500 ft., 11 MV; Cold- stream, 4 KU; Trout Creek (1 mi. up), Summerland, Okanagan Lake, 1 MV; Oliver, 1 CN; Osoyoos, 2 CN; Osoyoos-Bridesville Summit, 5 CN; Anarchist Mountain, 3 PM, 2 BC; Midway, Kettle River Valley, 4 CN. Washington: Okanogan Co.: Molson, 3800 ft., 1 US; 30 mi. Lake, Loomis (E base Windy Mts.), 1 SD; ConconuUy, 2280 ft., 4 MV, 2 US; Okanogan, 3 UM; 5 mi. SW Okanogan, 4 US. Chelan Co.: Chelan, 3 US; Wenatchee, 3 US. Douglas Co.: Waterville (8 mi. SW Badger Mts.), 3000 ft., 4 US; Orondo, 1 PS; Badger Mtn., 15 mi. NE Wenatchee, 4 PS; Exp. Station, E. Wenatchee, 2 PS; 4 mi. S Wenatchee, 4 PS; 4 mi. S East Wenatchee, 5 PS. Kittitas Co.: Blewett Pass, 1 US; Ellensburg, 1500 ft., 7 US, 2 WS; 18 mi. E EUensburg, 3 PS. Grant Co.: Steamboat Rock, Grand Coulee, 2 PS; I mi. S Devil's Punch Bowl, Grand Coulee, 2 PS; Dave Lewis Ranch, 10 mi. N Coulee City, 2 PS. Yakima Co.: Selah, 15 KU; North Yakima, 6 US; Wiley City, 10 mi. W Yakima, 2000 ft., 10 US; 18 mi. E Yakima, 1 PS; 1 mi. W Moxee, 1 MV; 3 mi. NW Sawyer, 4 WS; Zillah, 2 WS; Sunnyside, 1 US. Benton Co.: Benton City (Oberg Ranch), 1 US. Microtus montanus pratincola Hall and Kelson Microtus montanus pratincolus Hall and Kelson, Univ. Kansas Publ., Mus. Nat. Hist., 5:75, October 1, 1951. Microtus montanus pratincola. Hall and Cockrum, Univ. Kansas Publ., Mus. Nat. Hist., 5:417, January 15, 1953. Microtus nanus canescens, Bailey, N. Amer. Fauna, 17:31, June 6, 1900, specimens from Montana only. Type. — Subadult female. No. 34004, skin and skull. University of Kansas, Museum of Natural History, from 6 mi. E Hamilton, 3700 ft., Ravalli County, Montana; obtained by John A. White, original number 577, August 14, 1949. Range. — Flathead and Bitterroot valleys of western Montana, at lower altitudes (see Figure 9). Distinctive clxaracters. — Size small; color grayish; upper molariform tootli- row relatively short; bullae relatively large. Comparison. — The only adjacent subspecies (see Figure 8) is nanus, from which pratincola diflFers in smaller size, relatively smaller upper molariform teeth, and larger auditory bullae. The average diflFerence in total length of 6.5 between 148.1 and 154.6 millimeters of a series of 20 adults of pratincola 452 University of Kansas Publs., Mus. Nat. Hist. from various parts of its range and a series of 62 adults of nanus from Dis- tricts 29, 33, and 34 (Idaho only) was found to be significant at the 95 per cent confidence limit, but not at 99 per cent. Excepting size of bullae and molarifomi teeth, cranial characters cited by Hall and Kelson in the original description are duplicated in specimens of nanus of the same size. Only 2 specimens of nanus of 65 (from Districts 4, 33, 34, and 35) compared in detail with the two topotypes, equalled the topotypes of pratincola in the size of the bullae, and these two nanus were larger animals than the topotypes. Remarks. — A single specimen, skin only, No. 78205 MV, referred to by Davis (1937:26) and by Hall and Kelson (1951:76), from Canyon Creek, "a few miles west of Hamilton," Montana, is a young Microtiis longicaudus mordax, as evidenced by its grayish color and the length of the tail, which is more than 50 per cent of the length of the head and body (total length 143, length of tail 50 millimeters). Available records indicate a gap in the distribu- tion of Microtiis montaniis of at least 100 miles between pratincola and nanus (see Figure 8). This gap includes in large part moun- tainous habitat different from the more grassy valley floors and low- lands occupied by pratincola and by nanus to the west in Wash- ington and to a lesser degree to the south in Idaho. Trapping at a number of localities in this gap has yielded numerous records of Microtiis longicaudus, as well as Microtus pennsylvanicus and Microtus richardsoni (see Davis, 1939:314-324; Russell and An- derson, 1956:3). These facts suggest that this gap is at least a partial barrier to interbreeding between populations of Microtus montaniis on either side. Specimens examined. — Total number, 45. Montana: Sanders Co.: Hot Springs Creek, Flathead Reservation, 5 US; 15 mi. S Hot Springs, 1 US; Bison Range, 5 UM. Lake Co.: West arm of Flathead Lake, 5 US; Ravalli, 8 US. Missoula Co.: Missoula, 8 UM; Lolo, 1 UM, 1 WS. Ravalli Co.: Florence, 2 US; 8 mi. NE Stevensville, 2 US; Corvallis, 4 US; Hamilton, 1 MZ; 6 mi. E Hamilton, 2 KU. Microtus montanus canicaudus Miller Microtus canicaudus Miller, Proc. Biol. Soc. Washington, 11:67, April 21, 1897; Elliot, Field Col. Mus., 115:291, February 9, 1907. Microtus montanus canicaudus. Hall and Kelson, Univ. Kansas Publ., Mus. Nat. Hist, 5:77, October 1, 1951. Type. — Adult male, No. 75841, U. S. Nat. Museum, Biological Surveys Collection, from McCoy, Polk County, Oregon; obtained by B. J. Brctherton, December 1, 1895, original number 2119. Range. — The Willamette Valley, Oregon, and adjacent areas along the Co- lumbia River, at lower altitudes, in Oregon and Washington. Distinctive characters. — Size small; color brownish, and/or yellowish (the latter especially on sides); tail, in some specimens, more or less grayish above, in others bicolor; braincase relative to its length broad; incisive foramina little Anderson — Microtus montanus 453 constricted posteriorly; interpterygoid space comparatively acuminate anteriorly in a majority of specimens; bullae actually and relatively large in most speci- mens, auditory meatuses large; lateral pits at posterior edge of bony palate relatively shallow; rostral portion of skull relatively less well developed, size of nasals and upper incisors smaller on the average than in adjacent subspecies. An index of maturity of 1.75 is reached at 25.3 mm. condylobasilar length in canicaudus (Districts 7 and 8 combined). Comparison. — In small size canicaudus differs from M. montanus m,ontanus whose distribution lies to the south of canicaudus, but separated from it by a gap of 85 miles from which no specimens of the species are known. In the relatively rich brownish color of the upper parts and the yellowish suffu- sion of color especially noticeable on the sides, canicaudus differs from the three adjacent subspecies, canescens, nanus, and montanus. A series of cani- caudus was compared with three different series of nanus (from Districts 4, 34, and 35). From nanus from near Walla Walla, Washington (District 4) canicaudus differs as follows: in 9 of 12 pairs the interpterygoid space is more acuminate anteriorly and the distance between the first upper molars is less; in 10 of 12 pairs the auditory meatuses are larger and the incisive foramina are wider; and in 11 of 12 pairs the braincase is broader relative to its length and the distance between the incisive alveolus and the incisive foramen is less. In addition to the 21 characters used in the "method of pairs" the depth of the lateral pits at the posterior edge of bony palate, a character cited by Miller (1897:68) and Hall and Kelson (1951:78), was examined; in 9 of 12 pairs the palatal pits are shallower in canicaudus. None of the skulls of canicaudus used in these comparisons is included in either the series cited by Miller or by Hall and Kelson. Therefore, the observation of many of the same dis- tinctive features, is given added significance. In comparison with nanus (District 34, near Pocatello, Idaho) canicaudus differs as follows: in 10 of 13 pairs of skulls the zygomatic arches are more bowed; in 11 of 13 pairs the nasals are broader and the braincase relative to its length is broader; and in 13 of 13 pairs the incisive foramina are wider. Also, in 10 of 13 pairs the palatal pits mentioned above are shallower. In comparison with nanus from Teton County, Wyoming (District 35), canicaudus differs as follows: in 8 of 10 pairs of skulls the foramen magnum is larger; in 9 of 12 pairs the braincase is broader relative to its length; in 10 of 12 pairs the bullae are larger; and in 12 of 12 pairs the incisive foramina are less constricted. Also, in 9 of 12 pairs the upper incisors are less massive, and in 10 of 12 pairs the palatal pits are shallower. Comparison with montanus is reported on in the account of montanus. Remarks. — In the western half of Oregon, thirteen species of five genera of microtines occur. This is a more diverse fauna of mi- crotines than can be found elsewhere in North America in an area of the same size. The distributions of these species, as a result of the complex pattern of the topography, climate and vegetation, interdigitate or to a greater or lesser degree overlap. The inter- actions of some of these species, especially the species of Microtus (M, calif ornicus, M. montanus, M. longicaudus, M. oregoni, M. 454 University of Kansas Publs., Mus, Nat. Hist. townsendii, and M. richardsoni) , in local areas where two or more occur together would be a productive subject for ecological study. Amount and nature of interspecific competition, differences in re- production, different responses to variations in microclimate and vegetation, and seasonal and multi-annual fluctuations in popula- tions are some aspects of their natural history meriting study. Bailey (1936:206) reported the occurrence of M. m. canicaudus from Eugene, Lane County, Oregon. I have not seen specimens from there and have not mapped Eugene in Figure 8. A specimen (276122 US) was taken in 1927 by A. W. Moore on Lady Island "in Columbia River off of Washington shore at Camas, Washington" (A. W. Moore in litt.). The first specimens from the north side of the Columbia River and west of the Cascade Moun- tains were captured in 1957 by Dr. Murray L. Johnson. The Columbia River is a geographic barrier to free movement of voles. A comparison by pairs of seven skulls from Clark County, Washing- ton, with seven from the Willamette Valley in Oregon reveals the following differences: in 6 of 7 pairs the voles from Clark County have more pronounced zygomatic notches, more holes in the post- palatal shelf, and less nearly parallel tooth-rows, and in 7 of 7 pairs the voles from Clark County have a less acuminate interptery- goid space. In other cranial characteristics and in color of pelage the voles from Clark County resemble M. m. canicaudus rather than M. m. canescens, or M. m. nanus and are best regarded, in my opinion, as a variant population of canicaudus. Evidence for intergradation of M. m. canicaudus and M. m. nanus is scanty. Hall and Kelson (1951:78) referred specimens from Hood River and Wapinitia to M. m. canicaudus and were of the opinion that these specimens showed intergradation with M. in. nanus. These specimens and others from six miles east of Cascade Locks and "East of the Dalles" have been compared with M. m. canicaudus from the Willamette Valley and M. vi. nanus from Idaho and southeastern Washington. The specimens from six miles east of Cascade Locks bear some resemblance to canicaudus in relatively broad incisive foramina, in large auditory meatuses, and in broad braincase; in other cranial characters and in the darker, less reddish, or yellowish color of pelage these specimens resemble nanus, to which they are here referred. Specimens from Hood River resemble canicaudus only in breadth of incisive foramina and in breadth of braincase; in other cranial characters and in color they resemble Anderson — Microtus montanus 455 nanus. The specimen from East of the Dalles in Oregon is inter- mediate between canicaudus and nanus in cranial characters dif- ferentiating these two subspecies, but is nearer nanus; the slightly brownish tinge of pelage is nearer that of the average canicaudus but there is less yellowish on the sides. The skulls of the four specimens from Wapinitia vary considerably. One skull (79987) is damaged. Another (79986) resembles canicaudus in broad in- cisive foramina, broad braincase, and large auditory meatuses, but resembles nanus in heavier rostrum, smaller bullae, less acuminate interpterygoid opening, deeper palatal pits, and less fenestrated palatal plates. The other two intact skulls resemble canicaudus less than does 79986. Number 79988 differs from canicaudus, and resembles nanus, in possessing a heavier rostrum, narrower brain- case, more constricted incisive foramina, deeper palatal pits, smaller bullae, and smaller auditory meatuses. Skull number 79985 has been broken in two and glued together out of place so that the breadth of cranium cannot be ascertained; in size of rostrum, con- stricted incisive foramina, deep palatal pits, and small auditory meatuses the specimen resembles nanus. The reddish-brown pelage of the specimens from Wapinitia resembles that of the average canicaudus, but is less yellowish and more grizzled. In my judg- ment, the specimens from each of these localities resemble nanus in the aggregate of their characteristics more than they do cani- caudus and I have referred the specimens to nanus. There is a distinct morphological and geographic hiatus between nanus east of the Cascade Mountains and canicaudus west of these mountains. No specimen mentioned in this paragraph is certainly an intergrade. But, because of the closer relationship (in morphology) of cani- caudus with the other subspecies of Microtus montanus than with any other species of Microtus, and because of the precedent set by Hall and Kelson {loc. cit.) and followed by Hall and Cockrum (1953:413), and Miller and Kellogg (1955:590), canicaudus is here regarded as a subspecies of Microtus montanus. Specimens examined. — Total number, 143. Washington: Clark Co.: S. Ridgfield, 1 PS; 7 mi. NE Vancouver, 12 PS. Oregon: Columbia Co.: Old Fort Warren, 1 SD; Scapoose Airport, 1 PS. Washington Co.: Banks, 1 US; Hilhhoro, 2 US; Forest Grove, 1 MZ; Beaverton, 1 SD. Multnomah Co.: Portland, 22 AM, 4 KU, 4 LA, 1 PS, 4 SD; Gresham, 1 US. Yamhill Co.: Dayton, 13 PS; McMinnville, 1 PS; Sheridan, 2 US. Clackamas Co.: Canby, 4 US; Molalla, 4 PS; 7 mi. SE Molalla, 1 PS. Polk Co.: McCoy, 11 US, 7 CM. Marion Co.: Woodbum, 11 PS; Salem, 2 MV, 5 SD, 2 US, 2 MZ. Benton Co. : Corvallis ( 14 localities within 12 miles of Corvallis ) , 3 US, 5 KU, 12 OS, 3 PS. Linn Co.: Albany, 1 US; 5 mi. SE Albany, 1 OS. 456 University of Kansas Publs., Mus. Nat. Hist. Microtus montanus nanus (Merriam) Arvicola (Mynomes) nanus Merriam, N. Amer. Fauna, 5:63, July 30, 1891. Microtus montanus nanus. Hall, Proc. Biol. Soc. Washington, 51:133, Au- gust 23, 1938; Davis, The Recent Mammals of Idaho, The Caxton Printers, Caldwell, Idaho, p. 316, April 5, 1939, in part; Hall and Kelson, Univ. Kansas Publ, Mus. Nat. Hist, 5:77, October 1, 1951, in part; Durrant, Univ. Kansas Publ., Mus. Nat. Hist., 6:365, August 10, 1952; Hall and Cockrum, Univ. Kansas Publ., Mus. Nat. Hist., 5:416, January 15, 1953. Arvicola montana, Baird, U. S. Pac. R. R. Expl. and Surv., 8:530, July 14, 1857, in part (specimens from localities in "Nebraska," actually Ne- braska Territory, wow Wyoming, not examined by me ) . Microtus montanus, Bailey, N. Amer. Fauna, 17:27, June 6, 1900, speci- mens from Oregon ( in part ) and from Utah. Microtus nanus, Bailey, op. cit., p. 30, in part; Elliot, Field Col. Mus., 115:290, February 9, 1907, in part; Warren, The Mammals of Colorado, Putnam's Sons, p. 98, 1910, in part; Cary, N. Amer. Fauna, 33:123, August 17, 1911, in part. Microtus nanus canescens, Taylor and Shaw, Occ. Papers Chas. R. Conner Mus., 2:24, December, 1929, in part; Bailey, op. cit., p. 31, specimens from Washington ( in part ) and from Oregon. Microtus montanus cariji Bailey, Proc. Biol. Soc. Wasliington, 30:29, Febru- ary 21, 1917, type from Milford, Fremont Co., Wyoming; Hall and Cockrum, op. cit., p. 414, in part. Microtus montanus montanus, Bailey, N. Amer. Fauna, 55:202, in part; Hall and Cockrum, op. cit., p. 416, January 15, 1953, in part (204691 US, and 31599 MV). Microtus nanus nanus. Whitlow and Hall, Univ. California Publ. Zool., 40:266, September 30, 1933; Bailey, op. cit., p. 204. Microtus canicaudits, Bailey, op. cit., p. 205, in part. Microtus montanus fusus Hall, Proc. Biol. Soc. Washington, 51:131, August 23, 1938, specimens from Kinney Ranch, Wyoming; Warren, The Mam- mals of Colorado, Univ. Oklahoma Press, p. 229, 1942, in part. Microtus montanus nexus Hall and Hayward, The Great Basin Naturalist, 2:106, July 20, 1941, type from West Canyon, Oquirrh Range, Utah Co., Utah; Durrant, op. cit., p. 365. Microtus montanus canicaudus. Hall and Kelson, op. cit., p. 78, in part. Microtus montanus amosus, Durrant, op. cit., p. 368, in part. Microtus montanus rivularis, Durrant, op. cit., p. 369, in part. Type.— Subadult male. No. 23853/31253, U. S. National Museum, Bio- logical Surveys Collection, from Pahsimeroi Mountains, head of Pahsimeroi River, 9350 ft., Custer Co., Idaho; obtained by C. H. Merriam and V. Bailey, original number 1809, September 16, 1890. Range. — Southeastern Washington in the Palouse Prairie and the Blue Mountains; Oregon from the Deschutes Valley to the eastward in the drainages of the Great Basin and Snake River; Idaho on tlie Snake River Plains and adjacent highlands in the western and southeastern parts of the state; south- central Montana; Wyoming west and south of the Absaroka Range and the Shoshone Basin and as far east as Cheyenne; Utah north, east and south of the Great Salt Lake, at lower elevations and in the Wasatch and Uinta moun- tains at higher elevations, southward to southwestern Utah in mountainous part of state; Colorado in the mountains of the north-central part of the state as far south as the upper Arkansas River. Anderson — Microtus montanus 457 Distinctive characters. — Small; color usually grayish, sometimes pallid, but rarely grizzled with untinted silvery-gray (as in canescens), sometimes dark- ened with blackish, or reddish, or tinged with yellowish; length of tail, length of hind foot, and cranial characters average for the species, but all of these characters more variable from population to population than in any other subspecies. In nineteen districts (Table 2) representing nanus an index of maturity of 1.75 was reached at sizes ranging from 25.0 to 26.5 mm. condylobasilar length, and in ten of the districts at sizes from 25.5 to 25.8 mm. The greatest average sizes at the above level of maturity are in central southern Wyoming (26.0, 26.5, and 26.0 in Districts 40, 41, and 47) and in Utah (26.2 and 26.3 in Districts 52 and 57). Comparison. — In the preceding accounts nanus has been compared with canescens, canicatidtis, and pratincola the ranges of which he to the west and north of that of nanus. Subspecies that adjoin nanus on the south, listed from west to east, are montanus, micropus, amosus, and fusus. In north-central Wyoming the subspecies codiensis and zygomaticus adjoin nanus. In small size nanus differs, on the average, and in most samples significantly, from typical montanus, micropus (to a lesser extent), amosus, codiensis, and zy- gomaticus. In moderate grayish color most individuals of nanus differ from those of fusus, many of wliich show a paler wash of reddish and/or yellowish. Certain other characters are cited in the following accounts of these sub- species. The comparison, aheady discussed, of a series of canicaudus with three different series of nanus showed that in several characters, but not in all, canicaudus differed in the same manner from all three series of nanus. Results of several comparisons between series of nanus are listed below. The following series (and number of pairs) were compared: Teton County, Wyoming, and within 6 miles of Pocatello, Idaho ( 12 pairs ) ; Encampment and vicinity, Carbon County, Wyoming, and northwestern Colorado (8 pairs); Uinta County, Wyoming, and Encampment, Carbon County, Wyoming ( 14 pairs); Uinta County, Wyoming, and within 6 miles of Pocatello, Idaho (11 pairs); within 4 miles of Pullman, Washington, and Walla Walla County, Washington (8 pairs); within 4 miles of Pullman, Washington, and within 6 miles of Pocatello, Idaho (9 pairs). No significant differences were de- tected between any of these series, wdth one exception. In 8 of 9 pairs of skulls the specimens from Pocatello, in comparison with specimens from Pullman, have prelambdoidal fenestrae that are more elongate relative to their breadth. As is illustrated in Figure 6 and in Table 1 and Table 2 certain measurements and proportions differ from sample to sample within the range of nanus. Some of these differences are statistically significant. For example, the length of the tail expressed as a percentage of the length of the head and body differs in District 48 (Carbon County, Wyoming; 107 adults in series; 34.1 per cent) from the corresponding percentage in District 49 (Larimer County, Colorado; 12 adults; 37.7 per cent). This difference is significant. Averages for this proportion in districts assigned to nanus vary from approximately 33 to 38 per cent (the median and mode are within /2 per cent of 35 per cent). The average size (as indicated by the length of the head and body in millimeters) in districts in tlie range of nanus varies from approximately 110 to 118. The total length of the largest specimen of nanus recorded, other than in southeastern Oregon, is 185 milHmeters. The 458 University of Kansas Publs., Mus. Nat. Hist. largest individual from southeastern Oregon is 189 millimeters. The average length of the hind foot in districts assigned to nanus varies from 18.3 to 19.8 except in southeastern Oregon where the hind foot averages 20.1 millimeters. The average size of the hind feet expressed as a percentage of the length of the head and body varies in districts assigned to nanus from approximately 16.0 to 17.5. In this subspecies as in the species as a whole the larger percentages are in most, but not in all, cases in the western part of the range. Remarks. — The large, more or less central part of the range of the species occupied by nanus, the large size of this region in com- parison to the ranges of other subspecies, the more or less average position morphologically among the subspecies, and the locally more variable populations included are interesting aspects of the distribution and variation. The significance of this pattern in rela- tion to the evolution of subspecies is discussed elsewhere. The subspecies that are least diflFerentiated from nanus (listed in ap- proximate order of the amount of differentiation, the least differ- entiated form first) are as follows: fusus, pratincola, codiensis, zygomaticus, micropus, montanus, and canescens. Intergradation, in the sense of the existence of populations both geographically and morphologically intermediate between two different subspecies, is evident between nanus and each of the following subspecies: canescens, micropus, amosus, codiensis, zygomaticus, and fusus. Although less evident, intergradation occurs also between nanus and montanus. Individual specimens of nanus, not always in inter- grading populations, may be selected which resemble typical rep- resentatives of various of these other subspecies, in one or more characters. This overlapping of variability and the small size of many samples, especially from marginal localities, makes the sub- specific assignment of some specimens difficult or even arbitrary. This is, of course, to be expected; and is, in fact, one line of evidence for the validity of the concept of subspecies. There is no theoreti- cal or practical reason for assuming that all subspecies are equally differentiated, or for assuming that the areas in which they inter- grade must be of a certain width. Certain specimens, which have been assigned by various authors to the subspecies canescens, coni- caudus, and montanus in Washington and Oregon, are here referred to nanus as I have indicated in Figure 8 and the list of specimens examined to follow. A specimen from 2 mi. NE Weippe, Clear- water County, Idaho, referred by Davis (1939:318) to nanus and hsted by Hall and Cockrum (1953:416) as a marginal record, is an immature Microtus longicaudus (No. 54790 MV). A specimen from Mt. Harrison, 10 mi. S Albion (No. 72360 MV) also referred by Davis to nanus, is a Microtus longicaudus. Anderson — Microtus montanus 459 Microtus montanus occupying the eastern drainage of the Great Basin in Utah was described by Hall and Hayward (1941:106) as a subspecies, Microtus montanus nexus, differing from nanus in being blacker and less grayish, and slightly larger in size. Other distinctive characteristics cited were used to distinguish nexus from other subspecies than nanus. Essentially the same characters were noted by Durrant ( 1952:364). When samples of nanus other than those used by Hall and Hay- ward are used, neither external measurements or measurements of skull (see Tables 1 and 2) show significant differences. The aver- age condylobasilar length of skull in millimeters is 25.42 in nexus (23 adults. District 52) and 25.29 in nanus (32 adults, Uinta Co., Wyoming ) ; this difference is not significant. The index of maturity calculated for one millimeter size classes in both series indicates however that nanus reaches a corresponding degree of maturity on the average at a smaller size than does nexus ( nanus at 25.6 mm. in condylobasilar length is equivalent in maturity to nexus at 26.2 mm., on the average). Approximately 65 per cent of individuals can be identified correctly using this character. Specimens from certain localities in the ranges that have been assigned to both nanus and nexus are paler than the average for either subspecies, and conversely some localities are represented by specimens that are darker than the average for either subspecies. Series of nexus are on the average slightly darker than series of nanus. Individual variation in larger series of either nexus or nanus duplicates each hue of color and degree of darkness found in the other. On the basis of color I judge that approximately 60 per cent of specimens could be correctly assigned. The only significant cranial difference detected by the method of pairs (12 pairs of nexus. District 52, and nanus, within 6 miles of Pocatello, Idaho; and 15 pairs of nexus. District 52, and nanus, Uinta Co., Wyoming) is in size of the auditory bullae. The bullae are larger in 10 of 13 pairs (two skulls were damaged) in nexus than in nanus from Uinta County, Wyoming. In color, in size, and in cranial features nexus differs slightly from nanus — enough so that approximately 75 per cent of adult individuals can be correctly assigned. In these three characters nexus is intermediate between nanus and the three subspecies to the south and west, namely, amosus, rivularis, and micropus, but more closely resembles nanus. The difference noted is less than the difference between the other subspecies here recognized as vahd of the species Microtus montanus. 460 Unr^rsity of Kansas Publs., Mus. Nat. Hist. For these reasons the population heretofore known as nexus is here considered as nanus showing intergradation with more south- ern subspecies. Kuns and Rausch (1950:185) studied the helminths of four species of Microtus in the area of Jackson Hole, Wyoming. Micro- tus montanus was the most wide ranging of the four. Seven spe- cies of parasites were recorded from Microtus montanus; none was limited to M. montanus. Host specificity did not seem to be an im- portant factor in restricting the distribution of the parasites. More kinds and greater numbers of parasites were in habitats where the most voles were. Species of voles may interact through popula- tions of their parasites as well as in more direct ways. The total length, length of tail, length of hind foot, and size of sample of adults from each of 28 districts in the range of nanus are summarized in Table 1. The weights of ten or more adults are available from 13 of the districts in the range of nanus. The district number, the size of sample, and the average and the minimum and maximum weights in grams for each of these districts are as follows: 9, 13, 41.9(22.7-57.6); 12, 15, 41.8(31.0-53.5); 34, 10, 45.2 (36.0-62.9); 35, 60, 40.4(21.0-60.0); 37, 15, 37.8(28.0-48.5); 39, 26, 45.7 (33.4-56.6); 40, 18, 42.8(31.5-65.0); 41, 34, 44.9(29.7-61.2); 42, 23, 39.3 (30.0-53.0); 47, 22, 41.3(30.6-60.0); 48, 122, 42.4(20.1-56.8); 49, 43, 38.6 (21.0-56.2); 65, 12, 54.7(31.5-79.0). Specimens examined. — Total number, 1970. Washington: Stevens Co.: Gifford, 2 US. Lincoln Co.: 1 mi. E Bluestem, 2 PS. Spokane Co.: Liberty Lake, 2 US; Cheney, 1 US; Turnbull Slough, 1 WS. Grant Co.: RuflF, 2 WS. Adams Co.: 2 mi. W Ritzville, 2 WS; 2 mi. SW Ritzville, 1 PS; Roxboro, 1 WS. Whitman Co.: Oakesdale, 2 US; 1 mi. N Ewan, 2 CM; Colfax, 4 US; 1 mi. N Pampa, 11 WS; Union Flat Creek, W of Pullman, 3 MV; Pullman (Several localiries within 4 mi. of), 15 KU, 7 MZ, 5 CM, 134 WS, 3 MV; Steptoe Butte, 1 WS; Almota, 1 WS; Union Flats, 1 WS; Wawawai, 1 US, 1 WS; Wawawai (5-6 mi. NE of), 3 US; Putnam P., Wawowai, 1 MZ. KUcki- tat Co.: Smithville-Centerville Road, about 1500 ft. above Columbia River, 6 PS; 8 mi. E Bingen, 7 PS. Frankhn Co.: Connell, 1 WS. Walla Walla Co.: Prescott, 3 MZ, 3 MV; }2 mi. N Waitsburg, 1 KU, College Place (viathin S mi. of ), 3 WW, 3 EB, 1 KU, 1 MV; Fort Walla Walla, 1 US; Walla Walla, 2 KU, 1 WW; Walla Walla County Game Reserve, 1 WW; Blue Creek Ridge, 12 mi. E Walla Walla, 12 EB. Columbia Co.: Starbuck, 2 US; 5 mi. W Dayton, 1 OS; Dayton, 2 AM. Garfield Co.: Pomeroy, 1 US. Asotin Co.: Bly, 7 US; Grand Ronde River, 6 mi. S Anatone, 1 US; Rogersburg, 5 US. Idaho: Lewis Co.: Nez Perce, 3 US; 4 mi. NW Pollack, 6 KU, 9 MZ; Seven Devils Mountains, 3550 ft., 4 US. Adams Co.: New Meadows, 4 US; Tama- rack, 3 US; Goodrich, 2 US. Valley Co.: Van Wyck, 2 US; Smiths Ferrv, W side Payette River, 2 PS. Washington Co.: Midvale, 4 US; Crane Creek, 15 mi. E Midvale, 13 KU; Weiser, 9 US. Payette Co.: Payette, 4 US. Gem Co.: Sweet, 1 US. Custer Co.: Mill Creek, Challis Nat. Forest, 1 US; Challis, 7 US; Sawtooth Lake, 2 US; Pahsimeroi Mountains, head Pahsimeroi River, 1 MV, 13 US; Lost River Mts., 1 US. Butte Co.: 26 mi. SW Arco, 12 MZ. Fremont Co.: N fork Snake River, 10 mi. SW Island Park, 6200 ft., 2 AM; Black Springs Cr., 4 mi. W Ashton, 5200 ft., 1 AM; 5 mi. W St. Anthony, 5000 ft., 1 AM. Canyon Co.: Nampa, 10 US, 1 MV; Bowmont, 1 US. Ada Co.: Boise, 1 US. Camas Co.: Camas Prairie, Corral, 5100 ft., 2 US. Blaine Co.: Alturas Lake. 7000 ft., 3 BS, 1 MV; Ketchum, 5 PS; 19 mi. NE Gary, Lava Lake, 5200 ft., 8 MZ; Craters of the moon, Laidlow Park, 2 MZ; Ticura, 10 Anderson — Microtus montanus 461 mi. S Picabo, 1 US. Bingham Co.: Shelley, 6 US. Bonneville Co.: S side South Fork, 3 mi. W Swan Valley, 3 MV; 10 mi. SE Irwin, 4 US; Gray's Lake, 2 WS. Owyhee Co.: Three Creeks, 3 US. Gooding Co.: 2 mi. E Hagerman, 1 MV. Minidoka Co.: 2 mi. E Acequia, 3 MV; Heyhurn, 2 US. Twin Falls Co.: 2 mi. S Hagerman, 1 MV; 9 mi. S, 11 mi. W Twin Falls, Castleford Fenced Plot, 1 KU; 6 mi. S, 8 mi. W Twin Falls, Berger Fenced Plot, 3 MV. Cassia Co.: 2 mi. S, 5 mi. W Burley, 6 MV; 2 mi. S, 2 mi. W Burley, 5 KU; Elba, 5 MV. Power Co.: Bannock Creek, 10 mi. NW Poca- tello, 3 MV; Indian Springs, 4 mi. S American Falls, 3 MV. Bannock Co.: Pocatello (within 6 mi. of ), 18 MV, 28 KU, 4 US; Swan Lake, 1 US. FrankUn Co.: Strawberry Creek, 20 mi. NE Preston, 6700 ft., 5 MV. Bear Lake Co.: 15 mi. E Montpelier, 1 PS; MontpeUer Creek, 6700 ft., 3 US; N rim Copen- hagen Basin, Wasatch Mts., 8400 ft., 1 MV. Montana: Fergus Co.: 22 mi. S, 12 mi. E Lewistown, on Fergus Co. hne, 2 KU; Big Snowy Mts., 10 US. Beaverhead Co.: East Fork of Blacktail Creek, 1 MV, Gallatin Co.: West Fork of West Fork, Gallatin River, 1 US. Park Co.: Lamar River, 1 US; Gardiner, 1 US. Sweet Grass Co.: Cremer's Ranch, 13 mi. ENE Melville, 3 MZ; Cayuse Hills, 11 mi. ENE Melville, 1 MZ; "near" head Big Timber Creek, Crazy Mts., 1 US; Big Timber Creek, Crazy Mts., 1 US; McLeod, 1 US; 14 mi. S Big Timber, 1 US; West Boulder Creek, 18 mi. SE Livingston, 2 US. Oregon: Hood River Co.: Hood River, 9 SD, 15 US; 6 mi. E Cascade Locks, 6 PS. Wasco Co.: The Dalles (East of), 1 PS; Wapinitia, 7 mi. W Des Chutes River, 4 US. Gilliam Co.: VA mi. S Heppner Jet., 3 KU. Morrow Co.: 3 mi. S, 2 mi. E Hardman, 3 KU. Umatilla Co.: Milton, 1 US; 7 mi. W Langdon Lake, 1 WW; 4 mi. W Tailgate, 1 EB; Langdon Lake, 1 EB; Pendleton, 1 SD; Blue Mts., Meacham Lake, 3800 ft, 4 MV. Union Co.: Elgin, 2 US; Starkey Range (30-36 mi. SSE Pendleton), 8 US; Telocaset, 1 US. Wallowa Co.: 3 mi. N Troy, 1 PS; 15 mi. N Paradise, at Horse Creek, 2000 ft., 1 US; Doug Bar, Snake River, 2 SD; Wallowa R. Minam-Wallowa, 1 PS; Wallowa, 1 SD; Joseph, 1 PS; Wallowa Lake, 6 US; College Creek R. S. (Amnaha R., Wallowa N.F.), 1 US. JeflFerson Co.: Haycreek, 1 SD, 17 US; Foley Creek, 12 mi. E Haycreek, 2 US; Trout Creek, Ochoco Natl. Forest, 1 US; Camp Sherman, 1 PS. Wheeler Co.: 11 mi. W, 7 mi. S Mitchell, 4850 ft., 11 MV; Derr Rgr. Sta., Ochoco Natl. Forest, 2 SD. Grant Co.: Summit Creek, 7 mi. E Austin, 7400 ft., 3 MV; Cold Spring, 8 mi. E Austin, 4900 ft., 28 MV; Prairie City, 1 US; Schauffler Ranch, 5 mi. W Mt. Vernon, 1 MV; Strawberry Mts., 1 US; 5000 ft. N Fk. Malheur River, 21 mi. SE Prairie City, 2 MV. Baker Co.: Homestead, 1800 ft., 4 US; Home, 1850 ft., 5 US; 18 mi. SW North Powder, Beaver Meadows, 5300 ft., 1 US; Keating, 1 US; Baker, 1 EB; Jenning's Landing, 1 SD. Deschutes Co.: Sisters, 7 US, 2 OS, 1 SD; Upper Des Chute, 2 US; Bend, E bank of Deschutes River, 9 US; Little Lava Lake, 6 PS; Lapine, 1 US. Crook Co.: Marks Creek, Ochoco Natl. Forest, 2 US; Prineville, 1 US; Mouth of Davis Creek, 4 OS; Crooked River at mouth of Bear Creek, 3400 ft., 1 MV; Crooked River, 3 mi. E mouth of Bear Creek, 3600 ft., 1 MV; 2 mi. SW Barnes, 10 MV. Lake Co.: 25 mi. SE Lapine (Deschutes Co., in Lake Co., S. 17, T. 24S, R. 14E), 1 OS. Malheur Co.: Ironsides, 16 AM; Vale, 5 AM; George Harper Ranch, 1 US; Watson, 2 US; Sheaville, 2 US; 16 mi. W Jordan Valley, Arcuenaga Ranch, 1 US; /i mi. W Jordan Valley, 2 PS; Jordan Valley, 4200 ft, 5 US; Bowden Ranch, 1 US; Crooked Creek (head of), Owyhee Desert, 4000 ft., 6 US. Nevada: Elko Co.: Goose Creek, 2 mi. W Utah Boundary, 5000 ft., 19 MV. Utah: Boxelder Co.: Standrod, Raft River Mts., 5500 ft., 4 UU; George Creek, 7^ mi. SE Yost, Raft River Mts., 6500 ft., 3 UU; Lynn Canyon, 1 UU; Kelton, 4225 ft., 2 MV. Cache Co.: Logan, 1 UU, 4 MV; Anderson Ranch, Blacksmith Fork, 5500 ft., 1 UU. Weber Co.: Upper Beaver Creek at Junction with road, 1 UU; South Fk. of Ogden River, Beaver Creek, 1 UU; "Willows" Picnic Grounds, South Fk. of Ogden River, 1 UU; Ogden, 4293 ft., 18 US, 3 MV; 1 mi. NW Uintah, S of New Highways, 1 UU, Fremont Island, Great Salt Lake, 4300 ft., 1 UU; Fremont Island at South End, 1 UU; 2 mi. W of Roy, 2 UU. Davis Co.: Bountiful, 1 UU; West Bountiful, 4342 ft., 1 UU. Morgan Co.: 1 mi. East Morgan, 2 UU. Tooele Co.: 10 mi. S Tooele, 2 UU, 4 US; St. Johns, 4300 ft., 3 UU; Little Valley, 462 University of Kansas Publs., Mus. Nat. Hist. Sheeprock Mountains, 5500 ft., 1 UU. Salt Lake Co.: Lakeshore, 17 mi. W Salt Lake City, 4320 ft., 1 UU; 9.5 mi. W of Salt Lake City (2 mi. N Utah), 1 UU; 3 mi. W Salt Lake Airport, 1 UU; H mi. W Salt Lake Airport, U. S. Highway 40, 4200 ft., 6 UU; Salt Lake City, 4320 ft., 1 UU; 3 US; Utah Capper Gun Club, 4250 ft., 2 UU; E. shore Great Salt Lake at Saltair High- way, 4231 ft., 1 UU; Salamander Lake, Lamb's Canyon, 9000 ft., 3 UU; Butterfield Canyon, VA mi. above Butterfield Tunnel, 6300 ft., 1 UU; Taylors- ville, 2 mi. W Murray, Jordan River, 4500 ft., 1 UU; 1 mi. above Alta, 2 UU; Draper (and environs vi^ithin 2 mi.), 4500 ft., 10 UU. Summit Co.: 14 mi. S, 2 mi. E Robertson [Wyoming], 9300 ft., 1 KU; Henrys Park, Henrys Fork, 9000 ft., 1 UU; 4 mi. NE Snyderville, 5600 ft., 2 UU. Daggett Co.: Un- vi'ood, 1 MZ. Utah Co.: West Canyon, Oquirrh Mountains, 1 UU, 6 BY, 1 MV; Provo, 4 BY, 1 US, 1 MV; Fairfield, 10 US; 7 mi. W Spanish Fork, 4563 ft., 2 UU; Spanish Fork, 2 US; 1 mi. W Payson, 4600 ft., 1 UU; 1 mi. east of Payson Lake, 8300 ft., Nebo Mts., 1 UU. Wasatch Co.: Midway Fish Hatchery, 5450 ft., 3 UU. Duchesne Co.: Uinta Mts., head of Smith Fork, 1 US; Uinta Mts., 1 US. Uintah Co.: La Point, 1 UU. Juab Co.: Goshun Canyon, ^A mi. S Utah-Juab Co. line, 2 UU; Pines Picnic Area, SE Mt. Nebo, 7500 ft., 1 UU. Sanpete Co.: Manti, 20 US. Sevier Co.: 8 mi. E Glenwood, 1 US; Richfield, 2 US, 2UU; 10 mi. N Fishlake, Fishlake Mts., 10,000 ft., 3 MV; 9 mi. N Fishlake, 1 MV; Fish Lake, 1 BY. Piute Co.: Marysvale, 3 US. Iron Co.: Buckskin Valley, 1 US; ii mi. SE Sunset Point, Cedar Breaks National Monument, 1 MV; Brian Head, Parawan Mts., 5 US. Garfield Co.: Panguich Lake, 4 BY. Washington Co.: Blue Springs, Zion National Park, 3 MV. Wyoming, 831 specimens (the locality listed below from Carbon County is the only locaHty in Wyoming shown in Figure 9 that was not represented on the map previously published, Anderson, 1954:498; the 800 specimens previously examined also have been listed, Anderson, op. cit. -.496, 497), the additional 31 specimens being as follows: Sublette Co.: Togwotee Pass, 2 KU; Brooks Peak, 5 KU. Teton Co. (in addition to specimens from the same locaUties previously listed): Two Ocean Lake, 2 KU; Moran, 4 KU. Sweetwater Co.: Kinney Ranch, 23 mi. SW Bittercreek, 10 MV. Carbon Co.: 25 mi. E, 10 mi. S Saratoga, 9800 ft., 8 MV. Colorado: MoflFat Co.: Lay, 6160 ft., 2 WC, 1 AM. Routt Co.: Steamboat Springs, 6750 ft., 3 WC; Wright's Ranch, Yampa, 7700 ft., 4 WC, 2 AM; Gore Range, 8 mi. E Toponas, 8000 ft., 2 US. Jackson Co.: Boettcher Ranch, 1 DM; Homestead Ranch, 1 DM (locahty not known); 3 mi. below Cameron Pass, 9450 ft., 2 WC; Larimer Co.: 11 mi. W, 1 mi. S Rustic, 2 KU; 12'A mi. W, I'A mi. S Rustic, 2 KU; Cache La Poudre River, 1 CM; 16 mi. W Loveland, 6840 ft., 1 KU; 6 mi, W, M mi. S Loveland, 5200 ft., 1 KU; 19'A mi. W, 2% mi. S Loveland, 7280 ft., 6 KU; Estes Park, 2 AM, 1 US; 3 mi. S, 3 mi. W Estes Park, 5 KU; Bear Lake, Estes Park, 1 DM. Rio Blanco Co.: Meeker, 1 US; 9'A mi. SW Pagoda Peak, 7700 ft., 3 KU; 5 mi. S Pagoda Peak, 9100 ft., 2 KU; 2 mi. above Buford, 2 UC. Garfield Co.: Deep Lake, 16 mi. N Glenwood Springs, 1 KU. Eagle Co.: 20 mi. SW Toponas, 1 DM; Eagle, 1 US; Camp Hale (Pando), Yoder Gulch, 10,000 ft., 2 UC; Pando, 2 US. Grand Co.: Mt. Whiteley, 2 US; Arapahoe Pass, Rabbit Ear Mountains, 2 US; Grand Lake, 8300 ft., 1 WC; 9;» mi. N Kremmling, Antelope Creek, 7800 ft., 1 UC; Coulter (near Granby), 5 US; Arrowhead (near Dale), 1 US; Camp near Sheephom Pass, 8200 ft., 1 WC. Summit Co.: Mt. McClellan, 11,000 ft., 2 US. Boulder Co.: % mi. N, 2 mi. W Allenspark, 8400 ft., 4 KU; 3 mi. S Ward, 9000 ft., 3 KU; Nederland, 17 CM. Gilpin Co.: Tolland, 3 UC; Lump Gulch, 5 UC; Moon Gulch (2'A mi. SW RoUinsville ) , 3 UC. Clear Creek Co.: Berthoud Pass, 4 KU; Loveland Pass, 11,800 ft., 7 LA; Gray's Peak, 1 KU. Park Co.: Mt. Bross (4 mi. NW Alma), 1 DM; Buckskin Creek, 1 mi. above Alma, 10,500 ft., 2 WC; Trout Creek Ranch, 2 mi. N Garo, 9200 ft., 1 US. Microtus montanus codiensis Anderson Microtus montanus codiensis Anderson, Univ. Kansas Publ., Mus. Nat. Hist., 7:497, July 23, 1954. Microtus nanus, Bailey, N. Amer. Fauna, 17:30, June 6, 1900, specimens from Beartooth Mountains, Montana. Anderson — Microtus montanus 463 Microtus nanus nanus, Owen and Shakelford, Jour. Mamm., 23:307, August 13, 1942 (169926 and 169927 US). Microtus montanus nanus, Owen and Shakelford, op. cit., p. 311, also referring to specimens 169926 and 169927; Hall and Kelson, Univ. Kansas Publ., Mus. Nat. Hist., 5:77, October 1, 1951, specimens from Carbon County, Montana. Microtus montanus cartji. Hall and Cockrum, Univ. Kansas Publ., Mus. Nat. Hist., 5:414, January 15, 1953, in part (169250 US, and 37801 KU). Type. — Adult female, skin and skull; No. 27578, Museum of Natural History, University of Kansas, from 3% mi. E, % mi. S Cody, 5020 ft.. Park County, Wyoming; obtained by James W. Bee, original nvunber 18-8-11-48, August 11, 1948. Range. — In northwestern Wyoming eastward from the Absaroka and Wind River ranges into the western part of the Big Horn Basin, and in the Bear Tooth Mountains in Montana. Distinctive characters. — Size moderate for the species; tail actually and relatively long; hind foot actually but not relatively long; skull large; zygo- matic expanse actually and relatively large; alveolobasilar length relatively great; upper molar tooth-row relatively long; color pale, not reddish. An index of maturity of 1.75 is reached at 26.1 mm. condylobasilar length (Dis- trict 31). Comparison. — The only adjacent subspecies with which codiensis inter- grades is nanus. From it (material from Idaho), codiensis differs as foUows: larger; paler; tail and hind feet longer; upper incisors slightly more pro- cumbent; zygomatic arches more robust; upper molariform tooth-rows larger and tooth-rows more vddely separated. A series of six pairs of skulls of codiensis (topotypes), and nanus (from near Pocatello, Idaho) were com- pared. In 5 of 6 pairs codiensis has more procumbent incisors, wider jugal bone dorsoventrally, and greater width between alveoli of first upper molariform teeth. The geographically intermediate specimens from Jackson Hole, Teton County, Wyoming, seem to be intergrades between nanus and codiensis. The series of 18 pairs of skulls representing codiensis, and nanus from Teton County, Wyoming, differ significantly only in that the distance between the alveoli of the first upper molariform teeth is greater in codiensis in 15 of 18 pairs. A comparison of 12 pairs of skulls of nanus from Pocatello (within 6 miles of), Idaho, and from Teton County, Wyoming, revealed no significant differences. In three proportions, of the skull, that differentiate nanus (Poca- tello, Idaho) from codiensis, the intergrades from Teton County, Wyoming, more closely resemble codiensis. Average measurements of 17 specimens from Teton County, and of 27 topotypes of codiensis are, respectively, as follows: condylobasilar length, 25.6, 25.6; alveolobasilar length, 14.93, 14.90; length of upper molariform tooth-row, 6.58, 6.57; and zygomatic expanse, 15.49, 15.6. None of the differences are significant. The average length of the tail, expressed as a percentage of the length of the head and body, in these three series of specimens is as follows: Pocatello, 33.0; Teton Co., Wyoming, 34.2; and topotypes of codiensis, 38.3. In this feature the intermediates from Teton County are closer to nanus (from Pocatello). The numbers in the above three series are 34, 94, and 30. The difference between the length of the tail (expressed as a percentage of the length of head and body) in the sample from Teton County and in codiensis is significant. The difference in length of the tail between the sample 4—7800 464 University of Kansas Publs., Mus. Nat. Hist. from Teton County and nanus (from Pocatello) is not significant. The few specimens of codiensis available from places other than the type locahty are intergrades. From the subspecies zygomaticus, in the Big Horn Mountains to the east, codiensis difi^ers in relatively longer tail, relatively longer hind feet, longer upper molar tooth-row, larger and less flattened bullae, and other char- acters listed below. Twelve pairs of skulls of zygomaticus and codiensis (topotypes of each) reveal that the latter differs as follows: in 9 of 12 pairs auditory meatuses larger; in 10 of 12 pairs less bowed zygomatic arches, larger maxillary septa, and larger bullae; in 11 of 12 pairs wider jugal plates dorso- ventrally, and greater distance between alveoh of first upper molariform teeth; and in 12 of 12 pairs wider first upper molariform teeth. External measure- ments of topotypes (District 31) and of other specimens assigned to codiensis (District 30) are included in Table 1. The average weight of 29 adult topo- types is 46.4 (26.0-68.0) grams. Specimens examined. — Total nimiber, 50. Montana: Carbon Co.: "Bear- tooth Mountains," 2 US; "Beartooth Lake," 1 US. Wyoming: Park Co.: 13 mi. N, 1 mi. E Cody, 6300 ft., 1 US; Black Mountain, head of Pat O'Hara Creek, 3 US; SW slope Whirlwind Peak, 9000 ft., 1 KU; type locality, 31 KU; Ishawooa Creek, 6300 ft., 2 US; Valley, 1 US; Needle Mountain, 10.500 ft., 4 US. Hot Springs Co.: 3 mi. N, 10 mi. W Thermopofis, 4950 ft., 6 KU. Microtus montanus zygomaticus Anderson Microtus montanus zygomaticus Anderson, Univ. Kansas Publ., Mus. Nat. Hist, 7:500, July 23, 1954. Microtus nanus, Bailey, N. Amer. Fauna, 17:30, June 6, 1900, specimens from Big Horn Mountains, head of Powder River, Wyoming. Microtus montanus caryi. Hall and Cocknim, Univ. Kansas Publ., Mus. Nat. Hist, 5:414, January 15, 1953, in part (32755 and 27606 KU). Type. — Adult male; 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, July 8, 1949. Range. — The Big Horn Mountains of north-central Wyoming, at higher altitudes. Distinctive characters. — Size moderate for the species; tail relatively short; molar series short; zygomatic arches broad and well rounded in lateral outline when viewed from above; bullae small and flattened; basioccipito-basisphcnoid suture raised. An index of matiurity of 1.75 is reached at 25.7 mm. and 26.4 mm. in districts 32 and 36. Comparison. — The differences between zygomaticus and the subspecies codiensis the range of which lies to the west of, but separated by the Big Horn Basin from, the range of zygomaticus have been discussed previously in the account of codiensis. 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. Remarks. — Intergradation with nanus has been noted previously (Anderson, 1954:501). External measurements of two series of Anderson — Microtus montanus 465 zygomaticus are included in Table 1. Weights of more than 9 adults are available from each of two districts. The number of each district, the size of sample, and the average and extreme weights, in grams, are as follows: 32, 19, 47.6(32-67); 36, 14, 43.5 (30-56.9). Specimens examined. — Total number, 55. Wyoming; Big Horn Co.: type locality, 30 KU; W slope, head of Trappers Creek, 9500 ft., 2 US. Washakie Co.: 9 mi. E, 5 mi. N Tensleep, 7400 ft, 1 KU. Johnson Co.: IVz mi. W, 1 mi. S Buffalo, 6500 ft., 3 KU; Big Horn Mountains, head of Powder River, 3 US. Natrona Co.: BufiEalo Creek, 27 mi. N, 1 mi. E Powder River, 6075 ft., 16 KU. Microtus montanus montanus (Peale) Arvicola montana Peale, Mammalia and ornithology. U. S. Expl. Exped. (Wilkes) 8:44, 1848; Baird, U. S. Pac. R. R. Expl. and Surv., 8:528, 741, July 14, 1857, in part. Microtus montanus montanus. Miller, Bull. U. S. Nat. Mus., 79:216, De- cember 31, 1912; Kellogg, Univ. California Publ. ZooL, 21:259, April 18, 1922; Grinnell, Univ. California Publ. ZooL, 40:185, September 26, 1933; Hatfield, Jour. Mamm., 16:265, August 12, 1935; Hall and Cock- rmn, Univ. Kansas Publ., Mus. Nat. Hist., 5:416, January 15, 1953. Arvicola longirostris Baird, op. cit., p. 530, type from Upper Pit River, California. Arvicola (Myonomes) riparius, Coues, Monogr. N. Amer. Rodentia, Rapt. U. S. Geol. Survey . . ., XI (No. I): 156, 1877, in part. Ft. Crook, California (3867 US), and Lost River, Cahfornia (2223/1271 US). Microtus longirostris, Trouessart, Catal. mammalium . . . fossiUum, p. 564, 1897. Microtus montanus, Merriam, N. Amer. Fauna, 16:95, October 28, 1899; Bailey, N. Amer. Fauna, 17:27, June 6, 1900, in part; Quay, Misc. Publ., Mus. Zool., Univ. Michigan, 82:22, March 17, 1954; Hoffman, Ecol. Monographs, 28:80, January, 1958. Microtus dutcheri Bailey, op. cit., p. 36, in part (specimens from Mammoth, Pine City, and Head of San Joaquin River). Microtus montanus yosemite Grinnell, Proc. Biol. Soc. Washington, 27:207, October 31, 1914, type from Yosemite Valley, 4000 ft., Mariposa Co., California; Kellogg, Univ. California Publ. Zool., 21:264, April 18, 1922; Howell, Jour. Agric. Res., 28:977, June 7, 1924; Grinnell, Univ. Cali- fornia Publ. Zool., 40:186, September 26, 1933; Hall and Cockrum, op. cit., p. 416, Microtus montanus micropus. Hall, Mammals of Nevada, Univ. California Press, p. 544, July 1, 1946, in part. Type. — Sex unknown; no. unknown, U. S. Nat. Mus.; headwaters of Sacra- mento River, near Mount Shasta (probably close to Mount Shasta Village, formerly Sisson, 3600 ft., Siskiyou County), Cahfornia; October 4, 1841; ob- tained by Titian R. Peale (type specimens cannot now be found). Range. — Oregon, California, and Nevada, south of the drainage systems of the Deschutes and John Day rivers in Oregon, in western Nevada, and in the Sierra Nevada of Cahfornia south to the Kings River; at lower altitudes in northern part of its range than in the southern; see figures 8 and 10. Distinctive characters. — Size medium; frequently blackish (especially in northern part of range) and/ or reddish; tail relatively long; relatively large 466 University of Kansas Publs., Mus. Nat. Hist. hind feet. In no single character of the skull does montanus differ in hke manner from all adjacent subspecies, namely canescens, nanus, micropus, undosus and dutcheri; therefore cranial characters are described in the com- parisons below. An index of maturity of 1.75 is reached at 26.6, 26.6, 27.0, 26.4, 25.8, 26.5, 26.1, and 26.0 mm. in condylobasilar length in the eight districts listed in Table 2. Comparison. — M. m. montanus differs in larger size from canescens and nanus, in smaller size from undosus, and in darker, more blackish color from canescens, nanus, and dutcheri. Three series, here assigned to montanus, were compared by the method of pairs: topotypes of montanus, specimens from the Steens Mountains in south- east Oregon, and specimens from Yosemite National Park and vicinity (District 26, heretofore referred to Microtus montanus yosemite Grinnell). Thirteen pairs of skulls representing montanus (topotypes) and the series from Oregon (Steens Mts. ) revealed no significant difference by this method. Ten pairs of skulls (Steens Mts. and Yosemite Park) revealed one significant difference; in 8 of 10 pairs the bullae are larger in the series from Oregon. An additional difference w^as noted in that in 9 of 10 pairs the supraoccipital region at the midpoint of the lambdoidal crest is more rounded in specimens from Yosemite Park, and the supraoccipital is more tilted forward from a dorsoventral plane. Fifteen pairs of skulls (topot>'pes of montanus, and Yosemite Park) revealed only one significant difference; in 11 of 14 pairs (one skull is damaged so the character could not be observed) the distance between the incisive alveolus and incisive foramen is greater in the series from Yosemite Park. To test the signifi- cance of this difference between the two series the distance mentioned above was measured, expressed as a percentage of the condylobasilar length of the skull, and for each series the average was calculated (11.12 from Yosemite Park and 11.99 from montanus) . This difference is of borderline significance at the 95 per cent level of confidence. Specimens, on the average, from the northern part of the range of montanus are darker than from the southern part. Speci- mens from the north (39 from District 14) average significantly darker (3.46 on the color scale explained on page 444 ) than specimens from the south ( 27 from District 26 that average 2.56). The range of variation in these two series, however, overlaps completely (1 to 5 on the color scale in both series). In short, if one attempts to separate individuals in these series on the basis of color, one out of three would be mistakenly assigned. The following differ- ences cited by Kellogg (1922:260) were studied: The skull in mojUanus was said to be slightly smaller than in yosemite. The condylobasilar lengths of 51 yosemite and 28 montanus average 26.14 and 26.63 mm.; the opposite of the difference cited. Compared with montanus the zygomatic arches of yosemite were said to be more widely spreading. The average zygomatic breadths for the series mentioned above are 15.89 (yosemite) and 15.98 (montanus). Expressed as a percentage of the condylobasilar length of the skull the two breadths are, respectively, 60.7 and 59.8. Fifteen skulls compared by the method of pairs for the most widely bowed zygomatic arches revealed no significant difference. The differences in the averages noted above are not significant in any event. The interorbital region in yosemite was said to be less depressed than in montanus. With this feature in mind, 15 pairs of skulls were compared; in 4 cases yosemite is more depressed, in 1 case montanus is, Anderson — Microtus montanus 467 and in 10 cases the skulls are not distinguishably different. The occiput of yosemite was said to be more abruptly truncate, permitting the condyles to be visible when the skull is viewed from above. Of 15 pairs of skulls compared,, 8 show the condyles more concealed in yosemite, 6 in montanus, and 1 not different. This is not a significant difference. The rostrum in yosemite was: said to be usually deeper proximally. A comparison visually of 15 pairs of skulls shows the rostrum deeper in 10 cases in montanus, 2 cases in yosemite,. and 3 cases not different. This difference is not significant. Average depth of the rostrum is not significantly different in the two series, when measured. Differences in the paraoccipital processes and in the jugal bones were cited. Examination by the method of pairs revealed no significant differences. Only two differences between the two series were detected; the color, and the mea- surement from the incisive alveolus to the incisive foramen. The difference in color holds for other samples from the northern and southern parts of the ranges. The difference in the average distance from incisive alveolus to in- cisive foramen does not vary geographically. Other minor differences are de- tectable between samples within the range of the subspecies montanus (here considered to include the subspecies yosemite). Two series of montanus (topotypes and Steens Mts., Oregon) were com- pared with micropus (from Ruby Mountains, Nevada). In 8 of 10 pairs of skulls the specimens from the Steens Mountains have more bowed zygomatic arches and less elongate prelambdoidal fenestrae than m,icropus. In 6 of 7 pairs of skulls montanus (topotypes) have less elongate prelambdoidal fen- estrae, greater distance between the incisive alveolus and incisive foramen, and smaller bullae. In each of the same 7 pairs the palatine opening is more acute in micropus. In 7 of 9 pairs of skulls of montanus (Yosemite Park) and of dutcheri, montanus has smaller auditory meatuses, more open incisive foramina, larger bullae, larger pterygoid foramina, and in addition a less inflated braincase. In 6 of 7 pairs of skulls of montanus (Yosemite Park) and undosus (Dis- trict 24) montanus has less inflated braincase and lesser relative distance between the occipital condyles and the paraoccipital processes than does undosus. Also, in larger size and grayer, less reddish or blackish color of pelage undosus differs from montanus. Topotypes of montanus were com- pared with undosus. In 6 of 7 pairs of skulls montanus has relatively more elongate and posteriorly attenuate nasals, less inflated braincase, lesser rela- tive distance between the occipital condyle and the paraoccipital process, less acuminate palatine opening, relatively smaller bullae, and more raised basioccipito-basisphenoidal suture between bullae. In comparing, by the method of pairs, a series of nanus from within 6 miles of Pullman, Washing- ton, with a series of m,ontanus from the Steens Mountains of Oregon, the latter differs as follows: in each of 5 pairs distance from occipital condyle to paraoccipital process less; in 6 of 7 pairs relatively broader prelambdoidal fenestrae, and greater distance between incisive alveolus and incisive fora- men; in 5 of 6 pairs smaller bullae. Corresponding comparison of a series of nanus from Pocatello, Idaho, with the montanus from the Steens Moun- tains reveals in the latter, in 6 of 8 pairs, more bowed zygomatic arches, less elongate braincase, less depressed nasals, wider incisive foramen, smaller maxillary septum, and wider jugal plate. 468 University of Kansas Publs., Mus. Nat. Hist. In comparing canicaudus and montanus, 21 features of the skull were in- vestigated. In 12 of the 21 features significant differences were found. M. vi. montanus in 5 of 6 pairs has less elevated basioccipito-basisphenoidal suture between bullae, and relatively broader nasals, in 6 of 7 pairs less bowed zygomatic arches, relatively more elongate braincase, wider jugal plates, less elongate prelambdoidal fenestra, comparatively larger foramen magnum, greater distance between the occipital condyle and paraoccipital process, larger maxillary septa, less acuminate palatine opening, and wider first upper molars, and in 7 of 7 pairs more procumbent incisors. Differences in color and in size have been noted in the account of canicaudus. Remarks. — The differences between populations of the subspecies montanus (including M. m. yosemite Grinnell) are less in degree and number than between any one of the adjacent subspecies and montanus. Surrounding subspecies in order of increasing degree of difference from montanus, are micropus, dutcheri, nanus, undosus, and canicaudus. M. m. montanus intergrades with undosus and micropus in western Nevada, and with nanus in eastern Oregon. The total length, length of tail, length of hind foot, and size of sample of adults from each of 11 districts in the geographic range of montanus are in- cluded in Table 1. Weights of ten or more adults are available from each of seven of the districts in the range of montanus. The district number, the size of the sample, and the average and the extreme weights, in grams, for these districts are as follows: 17, 28, 52.1(35.7-66.2); 18, 18, 44.5(25.6- 56.7); 20, 27, 41.3(27.1-60.0); 22, 21, 51.3(26.0-72.0); 25, 25, 38.9(25.5- 62.5); 26, 87, 45.9(22.8-74.8); 27, 41, 45.0(29.8-74.7). Specimens examined. — Total number, 1189, as follows: Oregon: Douglas Co.: Diamond Lake, 2 SD. Jackson Co.: Brownsboro, 7 US. Klamath Co.: Yamsay Mts., 5 US; Fort Klamath, 1 AM, 8 US, 1 PS, 16 OS; Upper Klamath Marsh, 4 US; Swan Lake Valley, 5 US; Dairy, 1 SD; Klamath Falls, 13 US; Klamath Basin {Lost River), 7 US; Tule Lake, 1 US. Lake Co.: West Silver Creek, 10 mi. SW Silver Lake, 8 US; Sycan Marsh, 4 US; Gearhart Mtn., 14 PS; HaH Mtn., 5 PS; Plush (west side Warner Lake), 9 US; 6 rni. W Lake- view, 6 MV; Lakeview, 1 US; 15 mi. E Lakeview, 1 SD; Adel, 1 SD; /2 mi. S West Side School, 1 MV; Goose Lake Mountains, 1 US; Goose Lake Valley, 2 PS. Harney Co.: 28 mi. NW Burns, 1 OS; 5 mi. NNE Burns, 1 MV; Burns (Campbell Ranch), 28 US; 4 mi. E Bums, 5 MV; Squaw Butte, 5 US; 2 mi. E Crane, 1 US; Malheur Lake, 1 US; Malheur National Wildlife Refuge (Sec. S4, T. 28S, R. 31E), 1 US; Narrows (Refuge Hdqts.), 1 SD; Narrows, 22 US, 2 LA; Voltage, 12 US; Diamond, 19 US; Summit NE of Steens Mountains, 2 US; N end of Steens Mountain, 5 CM; Frenchglen, 1 PS; "Steens Moun- tains" (various localities in the following townships: T. 31S, R. 31E; T. 31S, R. 32;^E; T. 32S, R. 3232E; T. 32S, R. 32?iE; T. 32S, R. 33E; T. 33S, R. 33E; including localities known as Fish Lake, Blitzen Canyon, Big Fish Creek, Keiger Gorge, Wild Horse Gulch, Sod House Ranch, and Blitzen Valley), 63 OS, 2 CM, 2 KU, 7 US, 34 SD, 1 PS; Blitzen, 1 US; Shirk P. O., 2 US. California: Siskiyou Co.: Stud Horse Canyon, Siskiyou Mts., 6500 ft., 1 US; Tule Lake, 3 SD; Brownell, 5 US; Medicine Lake, 8 MV; Big Spring, 1^ mi. S Mayten, Shasta Valley, 2 MV; 5 mi. N Weed, 1 MV; Parks Cr., Ui mi. SW Edgewood, 2900 ft., 3 MV; Sisson (=Mt. Shasta, city of), 2 SD, 25 MV, 56 US; Wiley Ranch, 8 mi. NW Day, 3750 ft., 2 MV. Modoc Co.: Tule Lake, NE corner of, Carr's Ranch, 5 US; Tule Lake, 2 SD; Lost River, 1 US; Steele Meadow, 4700 ft., 5 MV; Petroglyph Cliffs, Lava Bed Nat'l Monument, 3 MV; Anderson — Microtus montanus 469 Goose Lake, 5150 ft., 7 US, 3 MV, 1 SD; Sugar Hill, 2 MV; Davis Creek, Goose Lake, 1 US; Joseph Creek, 4800 ft., 10 MV; Happy Camp, 1 US; Kelly Hot Springs, 2 mi. E Canbv, 4500 ft., 3 MV; Likely, 3 LA; A.din, 2 US. Shasta Co.: Dana, 2 US; Fort Crook, 8 mi. WSW Dana, 3000 ft., 2 MV, 25 US; Fall Lake, Fall River Valley, 2 US; 12 mi. NE Burney, 4300 ft., 1 US; Cassel, 3000 ft., 1 US; Lassen Peak, 1 US. Lassen Co.: Bieber, 4 US; Hayden Hill, 2 US; Red Rock P. O., 5300 ft., 7 MV; 7 mi. E Ravendale, 5000 ft., 3 MV; 3 mi. W Observation Peak, 5400 ft., 6 MV; Pine Creek, S from Poison Lake, 2 US; Eagle Lake, Spaldings, 2 MV; Eagle Lake Resort, 2 MV; Eagle Lake, 3 mi. N Merrillville, 2 US; Petes Valley, 4500 ft., 2 MV; 10 mi. N Amadee, 1 MV; Susanville, 2 US; Honey Lake Refuge, 4000 ft., 3 mi. W Wendel, 4 MV; Amedee, 9 US; Milford, Honey Lake, 4 US; Willow Ranch, Long Valley, 4225 ft., 2 US. Tehama Co.: Battle Creek Meadows, 4800 ft., 4 MV; Lyons- ville P. O., 3500 ft., 3 MV. Plumas Co.: Hot Spring Valley, 5200 ft., 5 MV; Kelley's, Warner Creek, 5200 ft., 2 KU; 8 7ni. NW Greenville, 1 SD, 3 US; Prattville, 4300 ft., 2 US; U mi. WNW McKessick Peak, 6000 ft., 2 MV; Quincy, 3400 ft., 3 US; Spring Garden Ranch, Grizzly Mts., 4100 ft., 4 US; Buck's Ranch, 5100 ft., 1 US; Beckwith, Sierra Valley, 5000 ft., 17 US, 1 MV; Mohawk, 4600 ft., 3 US. Butte Co.: Jonesville, 1 MV. Sierra Co.: Weber Lake, 7300 ft., 4 US. Nevada Co.: Independence Lake, 7000 ft., 2 MV; Sagehen Creek, 6500 ft., 3 mi. N Hobart Mills, 15 MV. Placer Co.: Cisco, 7 MV; Summit, 1 US; Lake Mary, /2 mi. S Donner Summit, 2 MV; Donner, 10 US. Alpine Co.: Tallac, 4 US. Tuolumne Co.: Sonora Pass, 9600 ft, 5 US; Tuolumne Meadows, Yosemite Park, 8600 ft., 19 MV, 38 US; Head of Lyell Canyon, 9700 ft., 1 MV. Mono Co.: Swager Cr., Sweetwater Range, 7600 ft., 1 MV; Tioga Pass, 9800 ft., 2 KU; Mono Lake P. O., 6500 ft., 22 MV; Salmon Ranch, Mono Lake, 6500 ft., 6 MV; Farrington's Ranch, Mono Lake, 6800 ft., 10 MV, 16 US; Warren Fork of Leevining Creek, 9200 ft., 4 MV; Mono Pass, 7 US; Mt. Unicorn, 10 US; Mt. Lyell, North base of, 9 US; Mt. Dana, 4 US; J mi. S of Williams Butte, 6900 ft., 2 MV; Walker Lake, 8 MV; Dexter Creek Meadows, 23 SD; Mammoth (head of Owens River, Sierra Nevada Mts.), 1 AM, 2 MZ, 1 KU, 19 US, {including localities known as Pine City and above Lone Pine Camp); Convict Creek, 2 SD, 14 MV, 4 KU, 7 AM; Cottonwood Creek, 9500 ft.. White Mts., 2 MV. Mariposa Co.: Lake Tenaya, 2 US; Vogelsang Lake, 1 MV; Fletcher Creek near Vogelsang Lake, 10,100 ft., 10 MV; Yosemite, 4000 ft., 20 MV, 2 US; Porcupine Flat, Yosemite Park, 8100 ft., 3 MV; Sentinel Bridge, 4000 ft., Yosemite Valley, 2 MV; Yosemite Valley, 4000 ft., 8 MV; near Mono Meadmvs, Yosemite Park, 7300 ft., 6 MV; Gentry's Big Oak Flat Road, 5800 ft., 2 MV. Madera Co.: Banner Peak, 9500 ft., 3 US; Head of San Joaquin River, Alt. 4100 ft. above Lone Pine Camp, 8000 ft., 4 US. Fresno Co.: Cascade Valley, 2 SD; Shaver Ranger Station, 5300 ft., 13 MV. Inyo Co.: Robert's Ranch, 8200 ft., Wyman Creek, White Mts., 1 MV; Deep Spring Lake, 5000 ft., 12 MV; Big Pine Creek, 2 LA. Nevada: Washoe Co.: Barrel Spring (9M mi. E, 3M mi. N Ft. Bidwell, Cahfomia), 5700 ft., 4 MV; 4 mi. SW Diessner, 5800 ft., 3 MV; Bound Hole, 3900 ft., 3 MV; 4 mi. S Flanigan, 4200 ft., 1 MV; Cottonwood Cr., Virginia Mts., 4400 ft., 2 MV; Pyramid Lake, 2 mi. W Sutcliffe, 5 MV; S}i mi. E Reno, S. side Truckee R., 4500 ft., 12 MV; 7 mi. S Reno, 3 MV, 7 KU; % mi. S Mt. Rose, 9500 ft., 1 MV, 2 KU; 3 mi. S Mt. Rose, 8600 ft., 5 MV; Washoe, 4700 ft., 1 US; Incline Creek, 7100 ft., 1 MV; Gal-Neva Lake Beach, 6250 ft., Lake Tahoe, 1 MV. Storey Co.: Gilpin Spillway, Truckee Canal, 3 mi. WSW Wadsworth, 1 MV. Ormsby Co.: Carson, 1 KU, 18 US; Clear Creek, 3 mi. W Stewart, 1 KU. Lyon Co.: 2 mi. E Fernley, 3 MV; 2ii mi. W Hazen, Truckee Canal, 1 MV; 2 mi. N Yerington, 4350 ft., 3 MV; West Walker River, 6 mi. S Yerington, 4500 ft., 1 MV; % mi. S Wellington, 6 KU. Douglas Co.: Minden, 3 US; 5 mi. S Gardnerville, 2 KU; Carson River, 5 mi. SE Minden, 4900 ft., 1 MV. Mineral Co.: N fork of Cat Creek, Mt. Grant, 8800 ft., 5 MV; Lapon Canyon, Mt. Grant, 8800-8900 ft., 24 MV; Schurz Indian dam. Walker River, 4300 ft., 2 MV. Esmeralda Co.: Indian Creek, 7400 ft.. White Mts., 2 MV; Fish Lake, 4800 ft., 37 MV. 470 University of Kansas Publs., Mus. Nat. Hist. 122 122 117 Fig. 10. The distxibution of Microtus montanus in California and Nevada. Dots indicate localities from which specimens have been examined. The inset shows the range of the species in black and the regions shown in Figures 10 and 11. The heavy lines indicate probable hmits of the ranges of the following subspecies: Guide to subspecies I. M. m. undosiis D. M. m. nanus J. M. m. micropus G. M. m. montanus M. M. m. nevadensis H. M. m. dutcheri N. 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O 00 o o 00 1 o 1 t- 1 o 1 -*( 1 fN 1 cc 1 r^ 1 loco oo 00 "* 00^ t>iC OCO oo or~-i 00 00 O5 00 &5 00 00 00 00 00 00 ooo ooo 00 00 3 ^ >o^^ s 2s? 00 ^-v 1-1 1^ t^ o I> »o 1-Ht^ 00 417 CO :§ ^7 CO CO CO ■«7 CO (M 1 CO 1 o i CO i o i ^ 1 CO i o 1 rfHlM lOCO rp,-! Tto CO lO lO •CO CO M^* iM ^^7 (^^ (N (M M 7 (M TP 1 ^ 1 ^ 1 CO 1 o 1 ^ 1 .o 1 r. 1 1>C0 Olio O'* 1-H 1-H Tji o 00 (N oo o o T)o^ '^ CO C5(M (N(M (MCM (MiM (N (M (N (M (N (M (N (M """ 00 t> o T-H 00 1-H CO o 1-H 1— I T-H 1-H 1— 1 (N iC a> 1-H (M CO ■* 1-H ^H CO CO Anderson — Microtus montanus 503 C<1 C! t^ CD CO T-^ t>» O 00 ^O O C<) 05 r- (N lO ^h nco CO cOjir^ CO t>- cOj|t>» cojjco co^t>- t^ •^ \ ojl Tt< \ 1^1 oo' pol -*! »cl os' col co' co' -^ ' iO(N CO--" (MOO CCt>- coo -"ti ^ ICCO tJ< ^ (M 05 IM CO CC 03 CO IM t^ lO coco coco coio coio coco coco coco coco coio co^c coic coco coco coec •* ec co ■* (n ^ os (m lO oo c^ o co im co i^ n'od 00 od^oo 00 00 r^^oo od^oo oo^oo i> fo' lo' oo' Oil ■^1 ^hI o' co' col r-' (mI co' oi' ooeo t^ »-i oom i^iM ooco coci coco ano omo coo ooco oo^o -"fiM t>.t>. t^t^ r>.t^ t^t^ t^t^ i>i^ t>i^ i>b- t^t^ t^t^ t^ir^ t^t^ t>.r^ coo eO t>- •* IC t^ ■* "5 OOO lO iCO OJ 0000 ,~^ •^ ■ ■ . -^ . . -^ . . ^ . -OS- o • CO (N Cai o-^ i>C5 ano <-! o ooim ooi-i ooo oco oo i o C>1>— I 1— li— I 1— lO <— lO I— Oi OijOi 05 OS OSjjOJ 05ji05 05jj05 Oi CO' ■^' oo' co' io' Oil o' co' co' CO' t^-' o' o' OOIM OeO coo 05C0 Ot> t^lM ooo ooco 0"3 l^-OO OOIM o-* --iiO 0000 0500 0000 odoo ooo 0000 ooo 0000 ooo odi^ 0000 ooo ooo y-iOi t-' t^ •-I 00 O 00 !>• i-H>. 00 ^00 00 --H t^ 00 ^CO CO CO^CO CO CO CO^CO CO^CO CO^CO CO i-h' oo' c^' lo' i-h' ■^i ■^' 00' o' oo' «o' o' co' lOCO -^IM »«C0 lOCO coco lOCO lOCO -^IM CO-* -"i^fM »0 Tf lO iM -^ O coco coco coco coco coco coco coco coco COCO coco COCO coco coco oO'TjH o CO oco ■* 00 «o tam ^ oco ■* imo co ■^. . (o ■ ■ • ■ iti ■ . (o ■ • in • - «o CO CO -co o »o ic -co r>- -co co -co ic In.' m ' co' o' ic' o' ■*' <— i' ■*' co' o' ic' co' C^ ■* 000 CO ■* ^ IC •<** •Tt< CO COO O CO -^IM CO O O iM 00 (M ■* CO t^ l^ C<1 O CO- • • la ■ -CO. -co- -t>.- • t>. CO CO -00 CO »0 lO -co CO -co CO -t^ CO ^ a (M (N^IM (N IM IM^IM (M^IM (M^IM (N •-*' ■*' t^l •>*' t^l co' o' o' oo' co' im' im' o' CO-<** t>.0 OIM r-iO --HO oo ooo iM ■* OOO OO OO ■* (M O "-1 O •* ■* •^ iC ■* 1/5 •* lO •* •* ■* •>*•»*< lO Tj< •'f •* lO ■* ■* ■*! IC •* •* ■* IM C<) IM IM (MIM IM (M.-H^r-4rH-H CO'HOOlMIM m t>. o o ^H CO CO CO -"ti ■># IM ^ CO t^ 00 o (N t^ -V Tj^ TfH ■* >* ■* ifi iC 504 University of Kansas Publs., Mus. Nat. Hist. 3 C «.> C o H CQ < rt o o "o-c S O 03 03 O T3 03 03- o3 a> 03 .■Sja O 03 (D t< ■■5 .a o a> CS3 c3 3^3 O bC c o O s s 4J .2 00 ^co 00 S cT CO^v «D ^T ^"f CO CD t^ 41^7 coco ^ 1 COOl 0 1 CO 1 0 1 t-»co 01 ■* oo ?oco CO 3, ;OCO coco coco coco o 00^ (NCO coco f-M P CO CO — (NO 00 ^^«' 41 «> 00 00 00 ■H °P CO 1 00 CO ^ 1 OJCO CO 1 coco 0 1 (NOi *^ t- •^w •^w 00 t>^ 00 t^ 00 r^ 1— ( N.00 CO ■ 4)7 00 S" CO • -H 7 0 1-H en CO CO 1-H -H^ •CO 4<7 (N 1 »oco CO 1 .-H(N 0 1 1^0 T3 00 00 1 10 03 f-H 1— t 1-H ^H bO a; 1-H ^H 1-H 1-H IN -H 1-H ^H 9 CO »o -H =f 00^ 4)'«> bo 3 OS OS coo CO • •0 ■H 7 caudus 9.38 (9.1- 00 1 00^ "B 00 00 1 CO 1 coco 00 GO 00 50 CJ C S c ,-(CO oioo 6» 3 c c £ © 1 O5 00 10 1 O5Q0 "^ So lOCO t^ 00 S n S ■H7 CO C CO CO ■H7 :§■ 00 1 >oco ^ 1 coco :^ c^ 1 UOCO MCC COCO coco coco COCO coco coco 3 C C CO I— 1 t^CD •CO -H V •CO ■H 7 1— ( 10 CO 1-H 1-H CO • •1^ -H 7 ^ 1 COIN COI> CO'* 0 1 0 05 £ CO 1 COt^ CO 1 coco c^ 1 iCTt* lO-* UO^ M • •CO CO CO (N 00 h-CD • 00 41^ 05 1 oo ^ 1 r^ 1 C5 1 0)03 ^ 1 O3 00 CO 1 COiM (N03, lO't' kCco (NC3, co»o CO 10 CO 1-H 10 05 CO 0 T-H 00 =a t--00 CO CO CO CO CO 1-H 10 T-H 1-H Anderson — Microtus montanus 505 Ifi^ 05^ a>.~. IN.-^ IM CA CO C^ l-H (M lO lO ^(M C0 05 00 CO t^ ^^ t^ t^ M^ r^ ■H*r 41 »7 CO I^ CD 1 o 1 o 1 t- 1 Oi 1 00 1 o ' ^ 1 r^ 1 CO 1 00 r^ I>CO 00 CO t^tC OKM 00 IC t^. •* cot^ I^ (MiO C0 03 (N »o 00 ^^« 00 00 ^«> 00 ■H°° •ti °? 00 00 Soo CO 1 ^ 1 -H 1 o 1 ^ 1 -H 1 o 1 o 1 o 1 05C0 oeo ot^ OSiM COOi oco cot^ 00 CD O'f 00 t^ t^t- t^t^ 00 t^ l^t^ 00 t^ OOt^ 00 t^ I^t^ I>t~- 00 CO • CO CO CO^ CO 00 o' CO 00 o CO (N •CO CO CO •CO CO •CO •CO oi CO T-H ^^ 7 l-H T-H ■H V 1-H ■H T ■H T 1— t 1-H CD 1 c^ 1 00 1 CO 1 r^ 1 CD 1 Z i Ji 1 iO 1 ^ 1 CO 00 C<1 00 . CI •* C^-H CO 00 Ol-H 00 00 00 CO C^~^ .^~^ 00 -H CO-— . ^— s^ ^— ^ d (M 00 d l-H CO coco CO ^•o ■H 7 C^05 CO t^ •H =f Oi ^ =f o> ■H ? 05 OS M 1 ^ 1 cn 1 ^ 1 r-, 1 CO 1 ^ 1 ^ 1 .o 1 ^ 1 c^»o c^oo l-H 00 -H t^ OiM iMt^ coo lOi-H 65 . • 05 OiO (MiO doo dob dob dod doo doo ^ O500 S °^S- ^ dod ooo -c o o .£J -^ 3 « -ti 3 ^ ,.— ^ t^^ ^— s, y—^ »o^. ^-^ . co.^ . t^.— N ,„— ^ ^-^ 00 ^t^ OS 00 l-H 05 CO g 1-H 00 s -^.o^ «o 00 CO ^ 1 jj CO 1 1 CO o. 1 CO CO 1 CO ^co ai7 CO ^ 1 CO o 1 lOCO iO(M lO-0(M COCO coco lOlM coco coco coco coco coco coco coco coco coco coco N.—' 00 CO--* os" o CO 00 1> • Oi 00 CO lO • 00^ 00 • c^ 1-H d ■1^ d b^ •t^ t>- •(•^ •00 d t>^ r— 1 41 V 1— t l-H ■H 7 i-H -H V ■H T 1-H 1-H ^ 1 CO 1 r^ 1 ^ 1 o 1 00 1 I- 1 ci 1 cr^ 1 CO 1 coco ■^iCl lOCO t^»o OliC lOt^ ot^ 00 'f ooo T}<^ lOco l-H l-H i-H -H 1-H T-H COiC f-H 1-H CO-^ 1-H 1-H CDiO 1-H 1-H TfCO 1-H r-H f-H 1-H c^ 00 Cl' t^ cT CO 00 S" 05CD 05 • t^ ^ l^ •00 00 00 • t^ 00 •1> •Oi d 00 ^ 00 t^ 00 t^ 00 t^ 5-^ 00 J>^ ■H 7' CO CO CO • •CO 41 7 0 (N I-H CO 00 t^ • •CO 41 7 ^ 1 (NOi ^ 1 00 -H ci 1 coco 1 (N CO-"** IN y-i (N^ (N -H ? o> O) 41 ? C5 4i ^ ^ 1 CO 1 CO 1 &5 C0 05 ^ 1 CiCT) 1 CO 1 om O5Q0 O5 00 g aJoo 1 ^ o CJ 3 |ocoo ^00 00 . 00^ g r-l 1-H ^•00^ B CO^ 417 lOlM CO ^ 1 >< ICCO S 1 m coco COCO coco coco coco CO coco iCOO CO ■ •CO ■H 7' 00 00 .— 1 r-H cJ ^00 ■H 7 06 mco 05 • •b- 41 7 ^ 1 cot^ CO 1 oco coco ^ 1 ■*o 1 CiO >-H 1— I CO'** t^CO 1-H 1— * com m 0 1— ( 1-^ .-hS" o 1— ( moo ^00 -HOO 4i^ 00 CO .-H05 05 (N rlOO 417 CD 1 0-* w 1 CO 1 mo ^ 1 1 -H i CO-* (NCJ, cO-t^ •^i^ *^t- 00 t>^ CO (M 1—1 iCCD 1— ( O5 00 ■H T" ^ 1 00 (N C5 1 '-<0 O) 1 coo ^1 1 OiCO ^ 1 oco o 1 1—1 i-H 1—1 1—1 (N 1-1 1—1 ^H 1—1 t— 1 1—1 1—1 1—4 1— ( 1— t 1— ( 1— H 1— t 1—1 1— ( cq c^ CO CO (MCO o f? ot^ O) OS •H ^^ 05 OS -H "^ CO 1 t^co o 1 coco 0, 00 1 05 1 I— 1 o 00 00 00 00 00 00 00 00 1 8 00 oo CiOO 00 t^ o.^ t^ ^ CO oco CO CO ^^ ^ CO s CO M ^ ^ 1 loco CO 1 ■* CO ^ 1 ^ '^(M >0 lO ecfo coco coco coco coco coco I— 1 CO CO 1—1 41 -r 00 1—1 CO CO 1—1 oo? t- ■ -H T" o 1 i-iO lOOO CD 1 ooo ^ 1 (N CO o 1 CO lo ^ 1 o^ 1—1 1— 1 1—1 -H 1—1 1— t 1— < 1-H coic 1—1 1—1 CO (M CO t^o' CO • ■CO 41 ^ 1—1 CO CD (MIC 41 ^ CO 1 coco 00 1 coco ^ 1 C50 ^ 1 ^ 1 Tt