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 ~ ■ 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 (-. OJS 1 .^ _C -1.3 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 — "EWUII Y^ 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 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 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 , K O 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