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Mammalogy Papers: University of Nebraska State Museum Museum, University of Nebraska State

August 1993

"Taxonomy" from Biology of the Heteroeromyidae

Daniel F. Williams California State University - Stanislaus

Hugh H. Genoways University of Nebraska-Lincoln, [email protected]

Janet K. Braun University of Oklahoma

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Williams, Daniel F.; Genoways, Hugh H.; and Braun, Janet K., ""Taxonomy" from Biology of the Heteroeromyidae" (1993). Mammalogy Papers: University of Nebraska State Museum. 112. https://digitalcommons.unl.edu/museummammalogy/112

This Article is brought to you for free and open access by the Museum, University of Nebraska State at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Mammalogy Papers: University of Nebraska State Museum by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. TAXONOMY

DANIEL F. WILLIAMS, HUGH H. GENOWAYS, AND JANET K. BRAUN

Introduction

erein the taxonomy of the family is a single term for consistency except in cases H reviewed and diagnoses of Recent where we were unsure how an author mea­ species and accounts of all currently rec­ sured a trait. Ifno measurements were given ognized Recent species and subspecies are for holotypes (more than halfofthe species provided. The objective is to present a cur­ and subspecies) or where the measurements rent taxonomy for the family rather than a in the original description differed from our systematic review. Unfortunately, there own, either the published measurements in have been no comprehensive reviews for a subsequent revision, with appropriate ci­ most genera for 80 years or more and the tation, or our measurements were used. For current taxonomy of many species clearly a few taxa, measurements were made by is unsatisfactory, as is the understanding of others at our request. These are noted in the the relationships between most of the de­ acknowledgments. Museums where holo­ scribed fossil and Recent taxa. types are stored are identified by acronyms Arrangement ofsubfamilies, genera, spe­ preceding the catalog numbers; acronyms cies, and subspecies is alphabetical rather are those listed by Yates et al. (1987) for than phylogenetic. Recognition of species North American collections or by conven­ and subspecies, and taxonomic treatment tion for those few outside North America. oftaxa at the generic level and above, were Type localities are as given in the original generally based on the most recent system­ description except all numbers ~ 1,000 have atic reviews and published analyses of ar­ commas inserted for clarity, accent marks rangements. For cases where there was dis­ are used in spelling locality names where agreement in the literature on taxonomy, appropriate, and ft and m have no period. opinion derived from studies ofdistribution Other necessary amendments to the type and variation was favored over unsubstan­ localities are enclosed in brackets. tiated belief. Measurements ofholotypes (except weight in g) are given in mm. Measurements are Taxonomic History 0/the as listed in the original description, except Family Heteromyidae where conversion to mm was necessary. Where different terms were used in the lit­ The name Saccomyna Gray, 1843 was the erature for the same measurement, we used first applied to members ofthe family, based

38 TAXONOMY 39 on the genus Saccomys Fr. Cuvier, 1823, Alston (1876) first used the name Hetero­ with type species S. anthophilus. The origin myinae to contain the species included in of the type specimen was uncertain and its the family Saccomyidae of Gill (1872). Al­ type locality, given as North America, was ston (1876) discarded the family and sub­ thought to be one ofthe West Indian islands family names of earlier authors because (Baird, 1858). Because Cuvier's name could Saccomys was a synonym ofHeteromys and not be applied to any ofthe known species, he believed it necessary to derive the name it was later considered a nomen dubium and from the latter. Alston (1876) combined the Saccomys was relegated to the synonymy of Geomyinae and Heteromyinae in the fam­ Heteromys Desmarest, 1817 (Peters, 1874, ily Geomyidae. Coues (1877) reviewed the as quoted in translation in Coues, 1877:487). prior taxonomic history ofthe heteromyids Coues (1877), based on remarks by Peters and included other family or subfamily (as quoted in Coues, 1877:487), reached this names that had been used for various het­ conclusion but believed it unnecessary to eromyids. Allen and Chapman (1893) first discard the synonym Saccomys as the basis used the name Heteromyidae to constitute ofthe family name. Waterhouse (1848) used the family as currently defined. Weber (1904) the name Saccomyina to contain all of the first used the superfamily name Geomy­ North American rodents with external cheek oidea for the Geomyidae and Heteromyi­ pouches, encompassing the pocket gophers dae. Wood (1936) described a new subfam­ and heteromyids. The family name, Dipod­ ily ofHeteromyidae, Florentiamyinae, based omyna Gervais, 1853, which applied only on Florentiamys loomisi; Rensberger (1973) to the currently defined heteromyids, was transferred Florentiamyinae to the Geo­ proposed next. Baird (1858) amended Gray's myidae, stating that it shared lineage with name to Saccomyidae, and used the sub­ Entoptychinae and Pleurolicinae. family names Geomyinae for the pocket go­ Recognition ofone or two families for the phers and Saccomyinae for the hetero­ geomyids and heteromyids and their group­ myids. Later, Gray (1868) used the name ing as a superfamily have generated persis­ Saccomyinae for the known species of het­ tent disagreements for nearly 150 years. eromyids and the subdivisions (tribes), Di­ Most recently, Shotwell (1967) and Lindsay podomyina (Dipodomys) and Heteromyina (1972) classified geomyids and heteromyids (Abromys, Cricetodipus, Dasynotus, Hetero­ in a single family, while others (e.g., Hafner, mys, Perognathus, and Saccomys-Abro­ 1982; Hafner and Hafner, 1983; Wahlert, mys and Cricetodipus are synonyms of Pe­ 1985) classified them as separate families in rognathus, and Dasynotus and Saccomys are the superfamily Geomyoidea. Although the synonyms of Heteromys). Gray (1868:199) two groups are recognizably distinct bio­ used Baird's (1858) subfamily spelling but chemically and structurally, recognition of wrote only of"the family" ofpouched mice, one or two families is arbitrary; thus, there from which he excluded the pocket gophers. probably will continue to be disagreements Gray (1868) treated Saccomys anthophilus about their classification at the family level. as a valid species, not mentioning its pos­ Herein, the taxonomic arrangement ofHet­ sible synonymy with Heteromys (Hafner and eromyidae above the family level follows Hafner, 1983:4, incorrectly implied that the classification presented by Carleton Gray recognized that Saccomys was a syn­ (1984). onym ofHeteromys; see Peters above). Gill (1872) first formally used the name Sacco­ Classification and Species Accounts myidae as equivalent to the subfamily Sac­ comyinae ofBaird (1858) and Gray (1868), Class Mammalia Linnaeus, 1758 and combined the Saccomyidae and Geo­ Subclass Theria Parker and Haswell, 1897 myidae in the superfamily Saccomyoidea. Infraclass Eutheria Gill, 1872 40 WILLIAMS ET AL.

Order Rodentia Bowdich, 1821 zygomatic arches; nasals extend far beyond Suborder Sciurognathi Tullberg, 1899 incisors; zygomatic arches slender, with Infraorder Myomorpha Brandt, 1855 greatly reduced jugal (malar) and nearly Superfamily Geomyoidea Weber, 1904 contacting the tympanic; frontal trapezoid and broad; parietal broad and quadrate to Family Heteromyidae Allen and Chapman, pentagonal or triangular; interparietal broad 1893 to exceedingly tiny or obsolete; squamosal Diagnosis. -Size small to medium-small mostly or entirely confined to orbit; infra­ for order, total length varying from about orbital canal long and opening at a large 100 to 370 mm and mass ranging from about perforation anteriorly on rostrum and pro­ 5 to 170 g; tail shorter to much longer than tected from muscle pressure by counter­ length of head and body; body forms qua­ sinking in vacuity; mastoid foramen minute drupedal, subricochetal (some fossil forms), or absent; sphenopterygoid foramen occu­ and ricochetal; locomotion scansorial to ric­ pied only by a large vein or by a large vein ochetal; hind limbs considerably larger than and part of the internal pterygoid muscle; forelimbs; hind feet longer than typical vascular canal in basisphenoid vestigial; murids and cricetids; manus with four, jugular (posterior lacerate) foramen situated clawed digits; pes with four or five clawed between tympanic bulla and basioccipital digits; claws ofmanus elongate and slender; and large and slit-like; coronoid process fore- and hind feet slender; soles of hind small, sloping, and below level of mandib­ feet naked to clothed with dense covering ular condyle; mandibles small and widely of fur; invagination of skin on face forms diverging posteriorly; baculum with bul­ fur-lined cheek pouch opening anteriorly bous base tapering to a thin, upturned tip near the mouth; hairs oftwo orthree general or, exceptionally, an ornate, trifid tip; male types, long overhairs with or without a flat­ reproductive tract typically with the full tened cross-section and concave trough on complement of accessory glands common the dorsal surface, and underfur of shorter, for muroid rodents, or without ventrome­ often slightly curved hairs; hairs with im­ dial prostate or preputial or both glands bricate, flattened cuticular scales; scales of (Heteromys in so far as is known is atypical hairs in single layer; medulla cells of hairs in having only three of the seven glands); variable, but always compound; dental for­ origin of temporal muscle restricted to lat­ mula i 1/1, c 0/0; p 1/1, m 3/3, total 20 eral part of roof of skull; M. platysma cer­ teeth; anterior face ofupper incisor smooth icale inserts into pouch retractor muscle at or with weak or strong groove; cheek teeth fusion zone; M. cervico-auricularis divided brachydont to hypsodont and definitive to into three slips; M. buccinatorius pars in­ evergrowing; usually six cusps per molar, termaxillaris anterior on palate; M. adduc­ three on each loph; enamel ofmolars rarely tor brevis double; M. semitendinosus with divided into two plates, never reduced to one head; and pedal flexor muscles reduced one; skull light and thin; width across skull (Arata, 1964; Burt, 1936; Ellerman, 1940; at mastoid bullae often greater and never Hafner and Hafner, 1983; Hill, 1935; Ho­ much less than width across maxillary arch­ man and Genoways, 1978; Merriam, 1889; es; mastoids inflated and bullous and form­ Ryan, 1986, 1989; Wahlert, 1985; Wood, ing part of lateral and (in all but Hetero­ 1935). myinae) dorsal cranial surface; occipital Remarks. - The diagnosis is based mostly small and limited in area, but extending onto on Recent species. The Heteromyidae is di­ dorsal surface of cranium; interorbital re­ vided into three subfamilies: Dipodomyi­ gion wider than rostrum; tympanic inflated nae, Heteromyinae, and Perognathinae. The or highly inflated, and vesicular; palate earliest known heteromyid dates from the nearly horizontal, little ifany below level of Middle Oligocene genus Heliscomys Cope, TAXONOMY 41

1873, which was not assigned to any ofthe Subfamily Dipodomyinae Coues, 1875 recognized subfamilies (Wood, 1935), but was later included in the subfamily Perog­ Diagnosis. - Size varying from small to nathinae (Lindsay, 1972). Rensberger large for family; total length from 130 to (1973), however, depicted Heliscomys as the 370 mm; weights from 10 to about 170 g; common ancestor ofgeomyids and hetero­ body form ricochetal in Recent species, sub­ myids. For a comprehensive review offossil ricochetal in some fossil forms; forelimbs forms see the chapter, Fossil Record, in this slender; hind limbs enormously developed volume by J. H. Wahlert. compared to forelimbs; length of tail from slightly greater than head and body to 1.65 Key to the Subfamilies times head and body length; soles of hind feet with dense covering of hair; hind feet 1. Body form ricochetal; soles of hind feet long; manus with four, clawed digits; pes densely covered with short hairs; mastoid with four or five, clawed digits; hair long, bullae enormously enlarged, forming half smooth, slightly curved, and oval in cross or more ofthe dorsal surface ofthe skull section; no obvious distinction between posterior to the orbits; mastoid bulla with overhair and underfur; upper incisors hollow interior...... Dipodomyinae strongly grooved; molars hypsodont in Re­ I'. Body form scansorial; soles of hind feet cent forms, rooted, and tuberculate; cheek naked or partly covered with sparse, short teeth nearly evergrowing or evergrowing in hairs; mastoid bullae moderately to advanced forms; cusps short relative to strongly inflated but not forming more elongated base of tooth ("alveolar part" of than about half of the dorsal surface of the skull posterior to the orbits; bullae Wood, 1935:110); enamel pattern ofcheek with spongy network of bony trabeculae teeth not persisting with age; lophs of P4 in interior 2 unite first at or near center of tooth; pro­ 2. Adult pelage coarse with abundant stiff, toloph ofP4 usually single-cusped; lophs of flattened spine-like hairs on dorsal and upper molars unite first at the lingual mar­ lateral surfaces of body; juvenile pelage gin and progress to buccal margins; lophs and adult pelage ofa few taxa offine hairs ofp4 unite at center oftooth, presenting an and soft, flattened and anteriorly-grooved X-pattern; lophs oflower molars unite first bristles; mastoid bullae moderately in­ at buccal margin and next at the lingual flated; mastoid bullae never projecting to margin, always forming an H-pattern; fo­ postero-dorsal plane ofocciput (dorsal to the external auditory meatus the lateral ramen and pit between m3 and the base of wall of the skull is composed entirely of the coronoid process present or absent; pal­ the squamosal); ventral plane oftympan­ ate broad, extending to or behind level of ic bulla higher than occlusal plane ofcheek M3; center of palate between cheek teeth teeth Heteromyinae with or without ridges; squamosal perfo­ 2'. Pelage silky or with a sparse to moderate rated by auditory bulla; incisive foramen mix of coarse, spine-like hairs and silky small; auditory bullae enormously expand­ hairs on dorsal and lateral surfaces of ed; mastoid bullae appearing on dorsal sur­ body; mastoid bullae strongly inflated; face ofskull; inflated interior ofbullae most­ mastoid bullae projecting to or beyond ly hollow, without spongy network of posterior plane of occiput; except for a trabeculae; middle ear with stapedius mus­ thin spine of the squamosal extending over the external auditory meatus, lateral cle; auditory bullae extend posteriorly well wall of the skull dorsal to the meatus is beyond plane ofocciput; median ventral fo­ composed ofmastoid bulla; ventral plane ramina in the "central" ofcaudal vertebrae of tympanic bulla lower than plane of present or absent; scapula prolonged pos­ occlusal surface ofcheek teeth . teriorly; obturator foramen triangular; ar­ ...... Perognathinae ticulation of trapezium with scapholunar; 42 WILLIAMS ET AL. articulation ofpes bones variable: calcaneo­ reduced to vestige nor absent; auditory navicular, calcaneo-cuneiform, or astraga­ bullae enormously inflated, tympanic ex­ lus-cuboid; dorsal surface ofectocuneiform tending below level ofocclusal surface of hour-glass-shaped; baculum of moderate upper cheek teeth and anteriorly to be­ length relative to body size, slender, with a yond the mandibular fossa ... Microdipodops swollen base, and an upturned tip; tip of I'. Size relatively large, total length from about 205 to 370 mm; tail relatively long, baculum simple; phallus moderate in length, averaging from about 1.05 to 1.6 or more slender, and with external spines and ure­ of head and body length, not wider in thral lappets; M. bucco-naso-labialis with middle than at base, and with a promi­ anterior origin from dorsal margin of the nent distal crest and terminal tuft oflong bulge created by the incisor root; M. clei­ hairs; digit 1 ofhind foot reduced to ves­ domastoideus absent; extensores breves tige located about half-way up the foot muscles absent; M. adductor minimum su­ or absent externally; auditory bullae perficial at origin; M. abductor hallucis ab­ greatly inflated, but tympanic rarely ex­ sent; lumbricales muscles reduced or absent tending below level ofocclusal surface of (Burt, 1936; Hafner, 1978; Hafner and Haf­ cheek teeth and never extending anteri­ ner, 1983; Hall, 1941; Nader, 1966; Ryan, orly beyond mandibular fossa ... Dipodomys 1989; Webster and Webster, 1975; Wood, 1935). Remarks. - The above diagnosis is based Genus Dipodomys largely on Recent species. The Dipodomyi­ nae includes the kangaroo mice, Microdi­ 1841. Dipodomys Gray, Ann. Mag. Nat. Hist., ser. 1,7:521, August. podops, which share many structural fea­ tures with the Perognathinae. Hafner and Type species. - Dipodomys phillipsii Gray, Hafner (1983) considered sharing to be co­ 1841, Ann. Mag. Nat. Hist., ser. 1, 7:521, incidental retention of ancestral traits. The August. union ofDipodomys and Microdipodops in Diagnosis. - Body form ricochetal, with the subfamily Dipodomyinae, first advo­ greatly enlarged hind limbs, short fore­ cated by Reeder (1956) and sustained by limbs, short neck with compressed and Hafner and Hafner (1983), Lindsay (1972), partly fused cervical vertebrae, and with tail Ryan (1989), Wahlert (1985), and Webster usually longer than length ofhead and body; and Webster (1975), is based on structural digit 1 of hind feet vestigial or absent ex­ similarities of muscular, skeletal, phallic, ternally; underparts, foreleg (with individ­ and auditory systems, and patterns of pro­ ual exceptions), forefoot, dorsal surface of tein electrophoresis. Development of rico­ hind foot, upper lip, spot above eye, and chetallocomotion in the two living genera spot behind ear white; white stripe across ofthe subfamily may be the result ofparallel flank, extending to base of tail; base of tail evolution (Hafner and Hafner, 1983; Wood, white all around; white stripes extending 1935). along each side oftail from base to near tip; dorsal side of tail dark (not white); ventral side of tail usually dark (except most indi­ Key to the Genera viduals ofD. deserti); hairs long and narrow for family; hairs smooth and soft or silky; 1. Size relatively small, total length from no distinct underfur layer to pelage; hairs about 130 to 180 mm; tail relatively short, averaging only slightly longer than length oval in cross section; base of hairs often of head and body, wider in middle than wide compared to shafts; tail with crest of at base, and without a crest oflong hairs longer hairs distally, terminating in a tuft; along its distal one-third or a prominent upper parts some shade of buff or brown; tuft at its tip; digit 1 of hind foot not large, dermal sebaceous gland on back be- TAXONOMY 43 tween shoulders; cheek teeth hypsodont, Dipodomys and provided the most recent crown height in adults generally exceeding review of the genus. length ofroot; molars evergrowing or nearly evergrowing, single-rooted in advanced spe­ cies, the root tubular and continuous with Key to the Species the crown; molars ofprimitive species with more than one root, but roots generally 1. Four, clawed toes on hind foot (minute fused; p4 two-rooted and P4 three-rooted vestigial, fifth toe [digit 1orhallux] rare- in primitive species; in advanced species, ly present, but without a claw) ...... 2 upper and lower premolars single-rooted; I'. Five, clawed toes on hind foot (the hal- lux vestigial and located about halfway anterocone of P4 lost early by wear; in ad­ up the inner side of the foot) 10 vanced species, cheek teeth with enamel 2. Tip of tail with conspicuous, white tuft limited to anterior and posterior plates; p4 ofhairs; tuft usually measures about 20 never with more than 5 cusps, the fifth, when mm or greater in length, but ranges from present, appearing in center ofmetalophid; 6 to about 40 mm or more ...... 5 third upper and lower molars small, with 2'. Tip oftail without a white tuft of hairs .. 3 oval enamel pattern; auditory bullae huge; 3. Distal one-third of tail, including tuft, ventral surface of tympanic bullae rarely black; found only in central Mexico .. reaching level of occlusal surface of cheek ...... Dipodomys phillipsii teeth, never extending noticeably below that 3'. Distal one-third of tail not all black, level; frontal without ethmoid foramen; zy­ rather a mix oflight- and dark-colored hairs; distributed widely in western gomatic process ofmaxilla expanded in an­ North America, including central Mexi- teroposterior plane; center of palate be­ co 4 tween premolars ridged; pterygoid fossae 4. Length ofhind foot usually less than 36 double; masticatory and buccinator foram­ mm; length of nasals usually less than ina usually united; median ventral foramina 12.4 mm; occurs only in central Cali­ in caudal vertebrae; notch in transverse pro­ fornia north of the Tehachapi Moun­ cesses of caudal vertebrae gently curved; tains and west of the Sierra Nevada .. calcaneo-navicular or calcaneo-cuneiform ...... Dipodomys nitratoides articulation offoot; phallus with spines and 4'. Length of hind foot usually 36 mm or urethral lappets, and without dorsal groove; greater; length of nasals greater than tip of phallus strongly upturned; baculum 12.6, averaging greater than 13.0 mm; not occurring in central California, north moderate in length relative to body size and of the Tehachapi Mountains and west with strongly upturned tip and swollen base; of the Sierra Nevada . preputial gland absent (Burt, 1936; Hall, ...... Dipodomys merriami 1981; Hafner and Hafner, 1983; Homan and 5. Width across auditory bullae less than Genoways, 1978; Wahlert, 1985; Wood, 25 mm; length ofhind foot usually less 1935; Zakrzewski, 1981). than 46 mm; greatest length of skull Remarks. - The genus Dipodomys in­ usually less than 41 mm 8 cludes as synonyms: Perodipus Fitzinger, 5'. Width across bullae greater than 25.5 1867 (type species D. agilis Gambel, 1848), mm; length ofhind foot usually greater a name that was subsequently applied to the than 46 mm; greatest length of skull species of Dipodomys with five toes on the usually greater than 41 mm 6 6. White tail tuft not bordered proximally hind feet; Dipodops Merriam, 1890; and by band of blackish hairs; underside of Macrocolus Wagner, 1846 (Coues, 1875; tail usually not darker than sides oftail; Hall, 1981; Jones and Genoways, 1975). interparietal usually not visible in dor- Some species ofDipodomys have also been sal view ...... Dipodomys deserti listed as species ofCricetodipus Peale, 1848. 6'. White tail tuft bordered proximally by Grinnell (1919) synonymized Perodipus with band ofblackish hairs; underside oftail 44 WILLIAMS ET AL.

dark, contrasting with light sides oftail; 14'. Ratio ofbreadth across maxillary arches interparietal conspicuous from dorsal to greatest length ofskull less than 0.548 view 7 ·...... 15 7. White tail tuft small, length from about 15. Numerous white hairs measuring 2.0 to 6 to 20 mm; width across maxillary 2.5 mm on inner surface of ear pinna; arches less than 23.5 mm . crown length ofear generally more than · Dipodomys nelsoni 16 mm 16 7'. White tail tuft large, length from about 15'. No noticeable long, white hairs on inner 25 to 40 mm; width across maxillary surface ofear pinna; crown length ofear arches greater than 25 mm . generally less than 16 mm 17 · Dipodomys spectabilis 16. Crown length of ear generally greater 8. Length of hind foot usually between than 16.7 mm .. Dipodomys elephantinus about 42 and 49 mm; not occurring in 16'. Crown length ofear generally less than central Mexico...... 9 16.7 mm Dipodomys venustus 8'. Length of hind foot usually between 17. Length ofhind foot greater than 43 mm; about 34 and 42 mm; found only in greatest length ofskull greater than 39.4 central Mexico Dipodomys phillipsii mm Dipodomys agilis 9. Found only in north-central California 17'. Length of hind foot less than 42 mm; and south-central Oregon . greatest length of skull less than 39.3 · Dipodomys californicus mm Dipodomys simulans 9'. Occurs only in north-central Texas and 18. Ratio oflengths oftail to head and body southwestern Oklahoma . normally greater than 1.42; not in Baja · Dipodomys elator California 19 10. Lower incisors with flattened anterior 18'. Ratio oflengths oftail to head and body faces (chisel-like) .... Dipodomys microps normally less than 1.42; occurs only in 10'. Lower incisors with rounded anterior Baja California ..... Dipodomys gravipes faces (awl-like) ...... II 19. Occurs only in western Nevada and II. Length of head and body normally southeastern California east of the greater than 130 mm and ratio oflengths coastal mountain ranges . oftail to head and body less than 1.35; · Dipodomys panamintinus width ofskull across bullae greater than 19'. Not in Nevada and southeastern Cali­ 27 mm; breadth across maxillary arches fornia east of the coastal mountain greater than 26 mm .. Dipodomys ingens ranges 20 11'. Length of head and body less than 130 20. Found only in central California north mm; width of skull across bullae less of the Transverse ranges . than 26.2 mm; breadth across maxillary · Dipodomys heermanni arches less than 25.5 mm 12 20'. Found only in southern California south 12. Ratio oflengths oftail to head and body and west of the San Bernardino, San less than 1.30; not occurring in western Jacinto, and Santa Rosa ranges . California or Baja California 13 · Dipodomys stephensi 12'. Ratio oflengths oftail to head and body more than 1.30; mayor may not occur in western California or Baja California 14 Species Accounts 13. Width of skull across bullae less than 22.2 mm; ratio oflengths oftail to head Dipodomys agilis and body less than 1.08 ...... Dipodomys compactus 13'. Width ofskull across bullae greater than Diagnosis. -A medium-sized kangaroo 22.2, ratio oflengths oftail to head and rat with large ears, five toes on the hind feet, body greater than 1.13 ... Dipodomys ordii and a skull narrow across the maxillary 14. Ratio ofbreadth across maxillary arches arches; breadth of skull across maxillary to greatest length of skull greater than arches less than 54.8% of greatest skull 0.548 18 length; head and body length generally av- TAXONOMY 45 erages greater than 119 mm in males and northern, 62-chromosome forms (agilis, 116 in females; greatest length of skull av­ perplexus, andfuscus) were consistently dis­ erages greater than 39.5 mm in females; and tinguishable from the southern group with breadth ofmaxillary arches averages greater 60 chromosomes (simulans and others). Best than 21 mm in females (male are larger). et al. (1986) provided analyses ofgenic dif­ Comparisons. - D. agilis is most similar ferences between the northern and southern to D. simulans, D. venustus, D. elephanti­ karyotypic forms; and Sullivan and Best (in nus, and D. heermanni. From D. heerman­ press) determined current taxonomic ar­ ni, D. agilis can be distinguished by the nar­ rangements from bacular, morphometric, rower breadth across the maxillary arches, and previous genic and karyotypic studies. smaller hind feet, and larger ears. From D. elephantinus and D. venustus, D. agilis can be distinguished by its smaller size, shorter Dipodomys agilis agilis ears, and narrower maxillary arch (breadth Gambel, 1848 at midpoint less than 5.2 rather than greater than 5.2 mm). From D. simulans, agilis dif­ 1848. Dipodomys agilis Gambe1, Proc. Acad. fers in being larger with wider maxillary Nat. Sci. Philadelphia, 4:77. arches and longer ears; length of hind foot 1853. D[ipodomys]. wagneri Le Conte, Proc. Acad. Nat. Sci. Philadelphia, 6:224, January. averages greater than 43 in agilis and less than 42 mm in simulans; greatest length of Holotype. - Holotype lost, sex unknown, skull averages greater than 39.4 in agilis and from Los Angeles, Los Angeles Co., Cali­ less than 39.3 mm in simulans; size differ­ fornia; obtained by William Gambel (Grin­ ences are greater when comparisons are nell, 1922). made between animals ofthe same sex (Best, Measurements ofholotype. - Total length, 1978, 1983a; Grinnell, 1922; Hall, 1981; 267; length of tail, 165. Sullivan and Best, in press). Distribution. -Coastal plains and Pacific Distribution. - Dipodomys agilis occurs slope of mountains from Cuyama Valley, primarily in open woodland and chaparral Santa Barbara County, southward to Or­ communities from the Temblor Mountains ange County, California. In the Los Angeles and Transverse ranges (Tehachapi Moun­ Basin, generally occupies chaparral com­ tains on the east and Santa Ynez Mountains munities above 500 m in elevation. on the west) in west-central California, Remarks. - Taxonomy follows the ar­ southward in the San Gabriel and San Ber­ rangement suggested by Best et al. (1986) nardino ranges to the lower hills ofthe Los and Sullivan and Best (in press). Angeles Basin (Sullivan and Best, in press; unpubl. data). Remarks. -Grinnell (1922) and subse­ Dipodomys agilis perplexus quent authorities have remarked that D. (Merriam, 1907) venustus is similar to D. agilis, and may prove not to warrant specific recognition; if 1907. Perodipus perplexus Merriam, Proc. BioI. this is true, D. elephantinus may also prove Soc. Washington, 20:79, 22 July. 1943. Dipodomys agilisfuscus Boulware, Univ. to be conspecific with D. agilis. Stock (1974), California Publ. Zool., 46:393, 16 September. however, found identical karyotypes for D. simulans, D. venustus, and D. elephantinus, Holotype. -Adult male, skin and skull, which differed from D. agilis perplexus by USNM 29261/41328, from Walker Basin, having 2n = 60 versus 62 and FN = 116 3,400 ft, Kern Co., California; obtained on versus 110. Best et al. (1986) reported on 15 July 1891 by Vernon Bailey. karyologic, morphologic, and genic inves­ Measurements ofholotype. - Total length, tigations ofthe agilis group, finding that the 320; length oftail, 195; length ofhind foot, 46 WILLIAMS ET AL.

46; greatest length of skull, 41.75; width Overall, D. californicus is perhaps most across bullae, 25.55; breadth across maxil­ similar in size, structure, and color pattern lary arches, 22.55; nasal length, 15.55; in­ to D. elator, a species ofnorth central Texas terorbital breadth, 12.10. and adjacent areas in Oklahoma. The in­ Distribution. -Found in semiarid com­ terorbital region ofD. elator is broader, the munities in the mountains bordering the bullae larger, and the rostrum shorter than southern end ofthe San Joaquin Valley and in D. californicus. the western edge ofthe Mojave Desert; from Distribution. -California kangaroo rats the Temblor Range, Kern and San Luis primarily occur in chaparral and other shrub Obispo counties and southern Sierra Ne­ communities from central California, north vada and Tehachapi Mountains on the of the San Pablo-Suisun bays/Sacramento­ northeast to the desert slopes of the San San Joaquin rivers estuary system, north­ Bernardino Mountains on the southeast, ward in the coastal mountains and Sacra­ westward to the vicinity ofMt. Pinos, Ven­ mento Valley to south-central Oregon in the tura County, and coastal chaparral com­ Klamath and Rogue river watersheds, and munities, Santa Barbara Co., California. eastward on the Modoc Plateau ofCalifor­ Generally occupies chaparral communities nia to the Surprise Valley on the California­ in areas above 500 m in elevation. Nevada border, and thence southward on Remarks. -Grinnell (1921) first treated the eastern slopes of the Sierra Nevada to perplexus as a subspecies of D. agilis. Best about the northern border ofSierra County, (1983b) remarked that he could find no dis­ California. tinct separation of specimens of perplexus Remarks. -Grinnell (1922) treated D. and fuscus, based upon multivariate anal­ californicus as a subspecies ofD. heermanni, yses using 5 skeletal and 14 cranial char­ primarily because he found a few individ­ acters. uals with rudimentary fifth toes on the hind feet, and because ofwhat he termed "round­ about intergradation- from berkeleyensis Dipodomys californicus through tularensis, heermanni, and exi­ mius, to californicus. " He apparently failed Diagnosis. -A medium-sized, four-toed to consider, however, D. heermanni dixoni, (with exceptions) kangaroo rat with a broad the population geographically intermediate face, dark color, and with a tail with broad, to heermanni and tularensis. He remarked dark dorsal and ventral stripes and a white that the latter two taxa were alike in external tuft at its tip; length of head and body av­ appearance (but failed to note the distinc­ erages from about 112 to 117 mm; skull tive white tip of the tail of eximius), but with widely spreading maxillary arches, noted great differences in the skulls (see small bullae, thick incisors, relatively long above diagnosis). Stock (1974) found widely and broad (heavy) rostrum, and broad su­ divergent karyotypes for heermanni and praoccipital and interparietals, the length californicus (also Fashing, 1973), and noted and breadth ofthe latter about 3.5 and 3.0, the great similarity of the karyotypes of D. respectively; average and extreme measure­ merriami and D. nitratoides to that of D. ments for 10 adult and subadult specimens californicus. Patton et al. (1976) compared were given by Grinnell (1922). allele frequencies at 18 loci among popu­ Comparisons. - Dipodomys californicus lations ofD. californicus, D. heermanni, D. is readily distinguished from all other spe­ panamintinus, D. merriami, and D. nitra­ cies ofkangaroo rats occupying the same or toides, and concluded that the differences adjacent geographic areas by the combina­ between the four- and five-toed populations tion offour toes on the hind foot, medium of D. heermanni were greater than either size, dark color, and small auditory bullae. population was to D. panamintinus, and that TAXONOMY 47 the electrophoretic data clearly supported Holotype.-Adult male, skin and skull, karyotypic evidence that D. califamicus was MVZ 18347, from Marysville Buttes, 300 not conspecific with D. heermanni. Hall ft, 3 mi NW Sutter, Sutter Co., California; (1981) did not follow Patton et al. (1976) obtained by F. H. Holden on 5 April 1912. in recognizing D. califamicus as a species Measurements ofholotype. - Total length, separate from D. heermanni. 303; length oftail, 192; length ofhind foot, 42; length ofear (crown), 13; greatest length of skull, 36.7; width across bullae, 22.0; Dipadamys califamicus califamicus length of nasals, 13.9; breadth across max­ Merriam, 1890 illary arches, 21.9; width of maxillary arch at middle, 5.3; width of rostrum near end, 1890. Dipodomys califomicus Merriam, N. 4.3. Amer. Fauna, 4:49, 8 October. Distributian. - Found only in the vicinity 1899. Dipodomys califomicus pallidulus Bangs, ofMarysville Buttes, Sutter Co., California. Proc. New England Zooi. Club, 1:65,31 July. Remarks. - This population shares with 1916. Dipodomys califomicus trinitatis L. Kel­ D. c. saxitilis the characteristic ofbeing sub­ logg, Univ. California Pubi. ZooI., 12:366, 27 stantially smaller in all dimensions than in­ January. dividuals of D. c. califomicus; eximius dif­ 1925. Dipodomys heermanni gabrielsoni Gold­ fers from saxitilis primarily by slightly lighter man, Proc. BioI. Soc. Washington, 38:33, 12 March. color, differences that perhaps do not war­ rant recognition as separate subspecies. Holotype. -Adult male, skin and skull, USNM 16618/23544, from Ukiah, Men­ docino Co., California; obtained on 4 May Dipodomys califomicus saxitilis 1889 by Theodore S. Palmer. Grinnell and Linsdale, 1929 Measurements ofholatype. - Total length, 302; length oftail, 183; length ofhind foot, 1929. Dipodomys heermanni saxitilis Grinnell 43; greatest length of skull, 39.60; width and Linsdale, Univ. California Pubi. ZooI., 30: across bullae, 24.75; breadth across maxil­ 453, 15 June. lary arches, 22.70; nasal length, 14.55; in­ Holatype. - Adult female, skin and skull, terorbital breadth, 12.90. MVZ 34963, from Mesa near Dale's, on N Distribution. -Occurs from north of the side Paine's Creek, 700 ft, Tehama Co., Cal­ Sacramento-San Joaquin estuary system in ifornia; obtained on 27 December 1924 by the coastal mountains and on the western Joseph Grinnell. edge of the Sacramento Valley, northward Measurements ofholotype. - Total length, to southern Oregon, and eastward on the 300; length oftail, 183; length ofhind foot, Modoc Plateau to the border between Cal­ 41; length of ear (crown), 12; weight, 57.5 ifornia and Nevada. g; greatest length ofskull, 37.3; width across Remarks. -Grinnell and Linsdale (1929) bullae, 22.85; breadth across maxillary regarded gabrielsoni as inseparable from arches, 21.4; nasal length, 14.9; interorbital califomicus. breadth, 12.0. Distribution. - Foothills on the eastern edge ofthe Sacramento Valley, from about Dipodamys califamicus eximius the South Fork of the American River, Grinnell, 1919 northward to the south bank ofBattle Creek, Tehama County, California. 1919. Dipodomys califomicus eximius Grinnell, Remarks. -Measurement of 14 adult Proc. BioI. Soc. Washington, 32:205, 31 De­ males from the type locality were listed by cember. Grinnell and Linsdale (1929). 48 WILLIAMS ET AL.

Dipodomys compactus ditional analyses demonstrating specific separation ofD. compactus and D. ordii and Diagnosis. -A medium-sized, five-toed their sympatric occurrence in portions of kangaroo rat with a remarkably short tail; the geographic range ofcompactus. Schmid­ the tail has a relatively small crest; pelage ly and Hendricks (1976) listed average mea­ is relatively short and coarse; color ofdorsal surements for 12 males and 13 females. parts has an orangeish hue; skull with rel­ Setzer (1949) gave means and ranges of atively broad interorbital region, small bul­ measurements for each sex ofeach subspe­ lae, and rounded or rectangular-shaped in­ cies. terparietal. The ratio of lengths of tail to head and body averages from about 1.02 to 1.08, values from lower than other species Dipodomys compactus compactus ofDipodomys. The skull is narrower across True, 1889 the bullae than all other species, averaging 21.7 mm in males and 22.1 in females 1889. Dipodomys compactus True, Proc. U.S. (Schmidly and Hendricks, 1976). Natl. Mus., 11:160,5 January. 1942. Dipodomys ordii parvabullatus Hall, Univ. Comparisons. - Dipodomys compactus is Kansas Publ., Mus. Nat. Hist., 5:38, 1 Octo­ most likely to be confused only with D. or­ ber. dii, the only other medium-sized, five-toed 1951. Dipodomys ordii [argus Hall, Univ. Kan­ kangaroo rat in the southeastern portion of sas Publ., Mus. Nat. Hist., 5:40, 1 October. the geographic range ofDipodomys. D. com­ pactus compactus has a shorter tail which is Holotype. - None designated, but Poole less bushy and with a shorter dorsal crest; and Schantz (1942) assumed that it was an the ventral pencil of the tail is lighter col­ adult female, skin only (skull lost), USNM ored and does not extend to the tip. The 19665/35227, from Padre Island, Cameron pelage ofD. compactus is shorter and coars­ Co., Texas; obtained on 3 April 1888 by C. er than that ofD. ordii. The auditory bullae K. Worthen. ofD. ordii are more inflated, giving the skull Measurements ofholotype. - Total length, a triangular outline. In D. ordii, the supraoc­ 209.5; length of tail, 114.5; length of hind cipital is narrower and the interparietal is foot, 31; length of ear (crown), 6.5. narrower, more pointed posteriorly, and Distribution. -Occurs on Mustang and more triangular in outline than the rectan­ Padre Islands ofsoutheastern Texas and on gular to roundish shape ofthe interparietal the barrier islands ofTamaulipas, Mexico. of D. compactus. Remarks. - Baumgardner and Schmidly Distribution. -Southeastern Texas, in­ (1981), using both univariate and multi­ cluding Padre and Mustang islands, and the variate statistical routines, found no distinct mainland eastward from Zapata Co. and separation of the Mustang Island popula­ southward from Bexar and Gonzales coun­ tion from that ofPadre Island or the island ties to the barrier islands of Tamaulipas, populations from Tamaulipas, and there­ Mexico. fore treated parvabullatus and largus as ju­ Remarks. - The characteristics listed by nior synonyms of compactus. Schmidly and Hendricks (1976) as indicat­ ing specific separation ofD. compactus and D. ordii were considered by Hall (1981) to Dipodomys compactus sennetti indicate "hardly more than subspecific (J. A. Allen, 1891) grade;" therefore, Hall provisionally re­ tained compactus, largus, parvabullatus, and 1891. Dipodops sennetti J. A. Allen, Bull. Amer. sennetti as subspecies of D. ordii. Baum­ Mus. Nat. Hist., 3:226, 29 April. gardner and Schmidly (1981) provided ad- Holotype. - Adult male, skin and skull, TAXONOMY 49

AMNH 3478/2733, from near Brownsville, ofkangaroo rats by its large size, absence of Cameron Co., Texas (amended to "Santa a vestigial fifth toe on the hind feet, and the Rosa, near Sarita, Kenedy Co., Texas" by usual absence ofa darker ventral stripe on Baumgardner [1989: 103]); obtained on 9 the tail. Overall, D. deserti is most similar March 1888 by J. M. Priour. in size and proportions to Dipodomys spec­ Measurements of holotype. -[Approxi­ tabitis, but lacks a dark-colored band ofhairs mated by Allen from the dried skin] Total bordering the white tail tuft, generally has length, 210; length of tail, 110; length of larger hind feet, and a more inflated skull hind foot, 35; length ofear (crown), 9; great­ with a narrower interorbital region and nar­ est length ofskull, 38.1; width across bullae, rower breadth across the maxillary arches 23.7; length of nasals, 14.0; breadth across (Hall, 1981; Nader, 1978). maxillary arches, 19.4; interorbital breadth, Distribution. - Desert kangaroo rats are 13.1; length ofrostrum, 20.1; length ofmax­ found almost exclusively on loose sandy soils illary toothrow, 5.0. in association with Larrea orProsopis in the Distribution. - Found in the eastern two­ low, hot deserts ofNevada, extreme south­ thirds of the mainland of southern Texas, western Utah, western Arizona, southeast­ east of Zapata and Willacy counties and ern California, northwestern Sonora, and south of Atascosa, Bexar, and Gonzales northeastern Baja California (Nader, 1978). counties (Baumgardner, 1989). Remarks. -Nader (1978) reviewed indi­ Remarks. -Davis (1942) treated sennetti vidual, secondary sexual, and geographic as a subspecies of D. ordii; Schmidly and variation in external and skull dimensions Hendricks (1976) assigned it to D. compac­ and provided the most recent systematic re­ tus. According to Baumgardner and view of the species. Schmidly (1981), topotypes of sennetti are the most atypical of the mainland samples of D. compactus, showing more similarity Dipodomys deserti aquilus to the samples of D. c. compactus of the Nader, 1965 barrier islands than do other samples from the mainland. 1965. Dipodomys deserti aquilus Nader, Proc. BioI. Soc. Washington, 78:52, 21 July. Holotype. -Adult male, skin and skull, Dipodomys deserti MVZ 126411, from 1.5 mi NW High Rock Ranch, about 12 mi SE Wendel, 4,080 ft, Diagnosis. - A large kangaroo rat with Lassen Co., California; obtained 21 July relatively large, four-toed hind feet and a 1960 by Charles S. Thaeler. tail with ventral coloration the same as or Measurements ofholotype. - Total length, only slightly darker than the lateral light 321; length oftail, 191; length ofhind foot, stripes, and with no dark band proximal to 54; length ofear, 18; weight, 108.9 g; width the white, distal tuft; length of head and across bullae, 30.00; breadth across maxil­ body averaging from about 134 to 155 mm lary arches, 23.50; nasal length, 11.85; in­ (Hoffmeister, 1986; Nader, 1978); skull with terorbital breadth, 15.40; length of maxil­ greatly inflated auditory bullae; interpari­ larytoothrow, 5.40; depth ofcranium, 14.40. etal usually absent in dorsal view in adult Distribution. -Sandy deserts ofeast-cen­ specimens; supraoccipital so compressed as tral Lassen Co., California and adjacent to be barely visible dorsally (1 mm or less). northwestern Nevada in Humboldt and External and skull measurements were giv­ Washoe counties (Nader, 1978). en by Hoffmeister (1986) and Nader (1978). Remarks. - D. d. aquilus is characterized Comparisons. - Dipodomys deserti is by its small size in comparison to other sub­ readily distinguished from all other species species. 50 WILLIAMS ET AL.

Dipodomys deserti arizonae Distribution. - Found on loose, sandy soils Huey, 1955 in west-central and southern Nevada, ex­ treme southeastern Utah, western Arizona, 1955. Dipodomys deserti arizonae Huey, Trans. northwestern Sonora, northeastern Baja San Diego Soc. Nat. Hist., 12:99, 10 February. California, and east-central and southeast­ Holotype. -Adult male, skin and skull, ern California. SDSNH 12532, from 3 mi SE Picacho, Pi­ Remarks. - Measurements for 8 males nal Co., Arizona; obtained on 14 May 1937 and 6 females were given by Nader (1978), by Laurence M. Huey. and Hall (1946) gave measurements for 10 Measurements ojholotype. - Total length, males and 6 females. 345; length oftail, 193; length ofhind foot, 52; length ofear, 14; greatest length ofskull, 45.3; width across bullae, 30.8; breadth Dipodomys deserti sonoriensis across maxillary arches, 23.7; length ofna­ Goldman, 1923 sals, 16.2; width of maxillary arch at mid­ die, 6.5. 1923. Dipodomys deserti sonoriensis Goldman, Distribution. -Generallylimited to sandy Proc. BioI. Soc. Washington, 36:139, 1 May. soils in south central Arizona in Maricopa Holotype. - Adult male, skin and skull, and Pinal counties (Nader, 1978). USNM 242306, from La Libertad Ranch, Remarks.-According to Nader (1978), 30 mi E Sierra Seri, Sonora, Mexico; ob­ W. Goodpaster collected desert kangaroo tained on 3 January 1922 by Charles Shel­ rats living in gravelly soil, the only recorded don. observations ofthis species occupying sub­ Measurements ojholotype. - [No external strates other than sand. Measurements for measurements recorded] Greatest length of 13 males and 10 females were given by Na­ skull, 46.6; width across bullae, 31.5; length der (1978). of nasals, 16.9; breadth across maxillary arches, 24.7; interorbital breadth, 14.6; length ofrostrum, 22.2; length ofmaxillary Dipodomys deserti deserti toothrow, 6.0; depth ofcranium, 13.3. Stephens, 1887 Distribution. - Found on the coastal plain of west central Sonora. 1887. Dipodomys deserti Stephens, Amer. Nat., Remarks. - D. d. sonoriensis is the largest 21:42, January. subspecies and the darkest colored, being 1903. Dipodomys deserti helleri Elliot, Field Co­ "nearly blackish" (Nader, 1978). Measure­ lumbian Mus., PubI. 87, ZooI. Ser., 3:249, 7 ments for seven males and eight females January. were presented by Nader (1978). Holotype. -Juvenile female, skin and skull, USNM 15629/22522, from Mojave River [3 or 4 miles from, and opposite Hes­ Dipodomys elator peria], San Bernardino Co., California; ob­ tained on 29 June 1886 by Frank Stephens. Diagnosis. -A medium-large sized kan­ Measurements ojholotype. - Total length, garoo rat with four toes on the hind foot 328; length oftail, 196; length ofhind foot, and a prominent white tuft at the tip ofthe 48.3; greatest length of skull, 44.2; width tail; skull with relatively widely positioned across bullae, 29.5; length of nasals, 16.0; auditory bullae, large interparietals and wide breadth across maxillary arches, 21; inter­ supraoccipital; rostrum wide, interorbital orbital breadth, 13.1; length of rostrum, region narrow, and orbits large; incisors rel­ 22.2; length of maxillary toothrow, 6.0; atively thick and stout (Carter et al., 1985; depth ofcranium, 13.3. Merriam, 1894a). TAXONOMY 51

Comparisons. - Dipodomys elator is su­ relatively long tail, dark coloration, and a perficially similar to D. spectabilis, but is narrow face; auditory bullae large, supraoc­ smaller. It resembles D. phillipsii ornatus, cipital and interparietal very narrow, and except for the white, ratherthan black-tipped incisors relatively robust; the length of the tail (Dalquest and Collier, 1964). It is also ear pinna exceeds that of all other species similar in size, proportions, and coloration ofDipodomys (Grinnell, 1922). to D. californicus (see the account of the Comparisons. - Dipodomys elephantinus latter species for distinguishing characters). can be distinguished from all other kanga­ Distribution. - The Texas kangaroo rat is roo rats except D. venustus by a combina­ known from a small area in north-central tion offive toes on the hind feet, ear (crown Texas and one locality in adjacent Okla­ length) in excess of 16.7 mm, a relatively homa in association with mesquite (Pro­ long, heavily crested tail averaging 155% of sopis) and short grasses (Carter et aI., 1985). length of head and body, large body size, Remarks. - Dipodomys elator is consid­ and relatively narrowly spread maxillary ered to be related to D. phillipsii, although arches; from typical individuals ofD. venus­ its specific relationships are not clear (Jan­ tus, D. elephantinus differs in being lighter nett, 1976). in color and having larger ears. Character­ istics ofcoloration and size listed by Grin­ nell (1922) as distinguishing elephantinus Dipodomys elator Merriam, 1894 and venustus are not diagnostic for popu­ lations of the venustus group in the Diablo 1894. Dipodomys elator Merriam, Proc. BioI. Range ofwestern Stanislaus and southeast­ Soc. Washington, 9: 109, 21 June. ern San Benito counties (D. F. Williams, Holotype. - Adult male, skin and skull, unpubi. data). USNM 64802, from Henrietta, Clay Co., Distribution. - The big-eared kangaroo rat Texas; obtained on 13 April 1894 by J. Al­ is known from chaparral communities in den Loring. the southern Gabilan Range in San Benito Measurements ofholotype. - Total length, and Monterey counties, California (Best, 292; length oftail, 173; length ofhind foot, 1986). 47; length ofear (dry, from notch), 14; great­ Remarks. -Dipodomys elephantinus is est length ofskull, 40.0; width across bullae, closely similar in structure, genetics, and 24.3; length of nasals, 14.5; breadth across ecology to D. venustus; Best (1986) noted maxillary arches, 23.2; interorbital breadth, that they possibly were conspecific. 12.7; length ofrostrum, 21.2; length ofmax­ illary toothrow, 5.4; depth ofcranium, 12.9. Remarks. - Ranges of external measure­ ments and means of cranial measurements Dipodomys elephantinus for 15 adults were given by Carter et ai. (Grinnell, 1919) (1985). Best (1987) provided summary sta­ tistics of 19 characters by sex for 206 adults 1919. Perodipus elephantinus Grinnell, Univ. California Pub1. Zoo1., 21 :43, 29 March. in three geographic samples, finding signif­ icant sexual dimorphism and geographic Holotype. -Adult male, skin and skull, variation. MVZ2851l,from 1 miNCookP.O., 1,300 ft, Bear Valley, San Benito Co., California; obtained on 9 July 1918 by Halsted G. Dipodomys elephantinus White. Measurements ofholotype. - Total length, Diagnosis. - A moderately large-sized, 333; length oftail, 200; length ofhind foot, large-eared, five-toed kangaroo rat with a 48; length of ear (crown), 18; weight, 90.7 52 WILLIAMS ET AL. g; greatest length ofskull, 43.2; width across Dipodomys gravipes Huey, 1925 bullae, 26.2; breadth across maxillary arch­ es, 22.95; nasal length, 15.5; interorbital 1925. Dipodomys gravipes Huey, Proc. BioI. Soc. breadth, 12.15. Washington, 38:83, 26 May. Remarks. -Grinnell (1922) gave mea­ Holotype. - Adult male, skin and skull, surements for 10 adults. SDSNH 4703, from 2 mi W Santo Domingo Mission, Baja California Sur, Mexico, lat. 30 0 4YN, long. 115°58'West; obtained on 28 Dipodomys gravipes February 1925 by Laurence M. Huey. Measurements ofholotype.-Total length, Diagnosis. - The San Quintin kangaroo 310; length oftail, 180; length ofhind foot, rat has five toes on the hind feet, is medium­ 44; length ofear, 10; greatest length ofskull, large in size with a small ear pinna, and a 41.8; width across bullae, 26.5; length of thick tail ofmedium length; the skull is rel­ nasals, 14.7; breadth across maxillary arch­ atively wide, the width across the maxillary es' 24.0; width ofmaxillary arch at middle, arches averaging more than 54.8% of the 5.9. greatest length of skull; the maxillary root Remarks. -Best (1983a) presented exter­ of the zygomatic arch has a sharp postero­ nal and cranial measurements for D. gra­ external angle (Best and Lackey, 1985). vipes. Comparisons. -Dipodomys gravipes is larger than the sympatric species, D. mer­ riami and D. simulans. It can be distin­ Dipodomys heermanni guished from the former by having five toes on the hind feet; in comparison to D. simu­ Diagnosis. -A medium large-sized kan­ lans, D. gravipes has shorter ears, longer hind garoo rat with five toes on the moderately­ feet, larger body size, and a tail that is thick­ sized hind feet, a relatively broad face, and er, paler, and less sharply bicolored (Best moderate-sized ear pinnae and auditory and Lackey, 1985); the head is wider, which bullae; the width across the maxillary arches is apparent externally as well as being re­ averages more than 54.8% of the greatest flected in measures of skull width. D. gra­ length ofskull and the maxillary root ofthe vipes is most similar to the allopatric spe­ zygomatic arch has a sharp postero-external cies, D. stephensi, from which it differs in angle. having larger hind feet (length greater than Comparisons. -Dipodomys heermanni is 44 versus less than 44 mm), larger body size similar in size and general appearance to (weights averaged about 79 to 85 versus 67 contiguous populations of D. agilis and D. g), relatively smaller auditory bullae that are venustus, and the allopatric species D. ste­ less globular in shape, and less constriction phensi and D. gravipes. From D. agilis, ve­ ofthe interbullar region dorsally (Best and nustus, and elephantinus, D. heermanni dif­ Lackey, 1985; Bleich, 1977; Grinnell, 1922). fers in having smaller ear pinnae, lighter Distribution. -Dipodomys gravipes is coloration, a less heavily crested tail, and a known from an area in Baja California bor­ broader face that is apparent externally and dering the Pacific Ocean, from near San Tel­ in greater width across the maxillary arches. mo on the north to just south ofEl Rosario From D. gravipes, heermanni differs in hav­ on the south. Its range extends from the ing shorter hind feet. The auditory bullae seashore to approximately 20 km inland of D. stephensi are more globular in shape (Best and Lackey, 1985). (i.e., shorter in length and deeper and wider) Remarks. - Best and Lackey (1985) re­ than those of heermanni, and the length of viewed the relationships of D. gravipes to tail ofheermanniaverages greater than 147% other species ofDipodomys. ofthe length ofhead and body versus 145% TAXONOMY 53 in D. stephensi (Grinnell, 1922). There is no 290; length oftail, 171; length ofhind foot, known physical characteristic that will dis­ 40; length ofear, 16; greatest length ofskull, tinguish all D. heermanni from D. pana­ 39.2; width across bullae, 23.9; breadth mintinus, although they are not known to across maxillary arches, 22.3; nasal length, be sympatric. From the sympatric species, 15.2; interorbital breadth, 13.05. D. nitratoides, heermanni differs in being Distribution. -Open sandy soils along the much larger (average weights about 70 to Pacific slope of the coastal ranges in San 80 versus 35 to 40 g) and having five rather Louis Obispo and Santa Barbara counties, than four toes on the hind feet. From the California, from about Oceano on the north sympatric D. ingens, heermanni differs in to the Santa Inez River on the south (Boul­ being smaller (ingens averages greater than ware, 1943). 110 g in weight), having shorter hind feet Remarks. - Boulware (1943) gave exter­ (45 or less versus greater than 45 mm), and nal and skull measurements for five males having a relatively longer tail (the length of and seven females. tail of ingens averages about 128% ofhead and body length). Dipodomys heermanni berkeleyensis Distribution. - Found in west central Cal­ Grinnell, 1919 ifornia south of the Sacramento Valley; known from south of the American River 1919. Dipodomys berkeleyensis Grinnell, Proc. on the east and the Suisun Bay on the west, Bio!. Soc. Washington, 32:204, 31 December. southward through the San Joaquin Valley below about 3,000 ft; and extending through Holotype. -Adult male, skin and skull, the interior valleys and mountains to the MVZ 28729, from head of Dwight Way, coast at Morro Bay and from Point Sal to Berkeley, Alameda Co., California; ob­ Point Conception in Santa Barbara County tained on 6 October 1918 by Joseph Grin­ (Grinnell, 1922; Hall, 1981). Within this nell and D. D. McLean. area, Heermann kangaroo rats are found on Measurements ofholotype. - Total length, a variety ofsubstrates in a diversity ofplant 301; length oftail, 180; length ofhind foot, communities. 41; length ofear (crown), 12; greatest length Remarks. -Grinnell (1922) and Hall of skull, 39.5; width across bullae, 24.1; (1981) included the four-toed populations length of nasals, 14.5; breadth across max­ of D. californicus in D. heermanni (see ac­ illary arches, 23.0; width of maxillary arch count ofD. californicus for details). External at middle, 5.0; greatest width of rostrum and skull measurements and details ofcol­ near end, 4.5; weight, 77 g. oration were given by Grinnell (1922). Distribution. - Known from the hills and valleys east of San Francisco Bay, in Ala­ meda and Contra Costa counties, California Dipodomys heermanni arenae (Grinnell, 1922). Boulware, 1943 Remarks. - Heermann kangaroo rats from adjacent areas in Santa Clara and Stanislaus 1943. Dipodomys heermanni arenae Boulware, counties, in the Diablo Range, are probably Univ. California Pub!. Zoo!., 46:392, 16 Sep­ of this subspecies. Grinnell (1922) listed tember. measurements for two specimens. Holotype. -Adult male, skin and skull, MVZ 84840, from C. A. Davis Ranch, 2 mi Dipodomys heermanni dixoni NNW Lompoc, 400 ft, Santa Barbara Co., (Grinnell, 1919) California; obtained on 16 February by Richard M. Bond. 1919. Perodipus dixoni Grinnell, Univ. Cali­ Measurements ofholotype. - Total length, fornia Pub!. Zoo!., 21:45, 29 March. 54 WILLIAMS ET AL.

Holotype. -Adult male, skin and skull, Dipodomys heermanni heermanni MVZ 26805, from Delhi, near Merced Riv­ Le Conte, 1853 er, Merced Co., California; obtained on 23 March 1917 by Joseph Dixon. 1853. D[ipodomys]. heermanni Le Conte, Proc. Measurements ofholotype. - Total length, Acad. Nat. Sci. Philadelphia, 6:224. 280; length oftail, 165; length ofhind foot, Holotype. -Juvenile [sex unknown], par­ 40; length of ear (crown), 14; weight, 72.5 tial skin and skull, ANSP 229, from Sierra g; greatest length ofskull, 37.5; width across Nevada, California [probably along the Ca­ bullae, 23.5; breadth across maxillary arch­ laveras River, Upper Sonoran life zone, Ca­ es, 21.65; nasal length, 13.55; interorbital laveras Co.; Grinnell, 1922]. breadth, 12.45. Measurements of holotype. - Length of Distribution. - Known from grassland and hind foot, 37.5; length of ear (crown), 10; savanna communities on the eastern mar­ greatest length of skull, 32.3; width across gin of the San Joaquin Valley, California, bullae, 20.0; breadth across maxillary arch­ in Merced and Stanislaus counties (Grin­ es, 17 (measured on intact side and dou­ nell, 1922). bled); nasal length, 10.2; interorbital Remarks. -Grinnell (1922) gave mea­ breadth, 10.8 (external measurements from mounted skin by Grinnell, 1922). surements for 10 adult and subadult spec­ Distribution. - Found in grassland and imens. This is the smallest sized subspecies savanna communities between about 500 of D. heermanni (Grinnell, 1922). and 3,200 feet in the northeastern San Joa­ quin Valley, California, from Amador Co. on the north to Mariposa Co. on the south. Dipodomys heermanni goldmani Remarks. -Grinnell (1922) augmented (Merriam, 1904) the scanty diagnosis ofLe Conte (1853) and discussed features ofthe type specimen and 1904. Perodipus goldmani Merriam, Proc. Biol. the probable type locality. Soc. Washington, 17: 143, 14 July. Holotype. - Adult male, skin and skull, USNM 118924, from Salinas, mouth ofSa­ Dipodomys heermanni jolonensis linas Valley, Monterey Co., California; ob­ Grinnell, 191 9 tained on 4 September 1902 by Luther J. Goldman. 1919. Dipodomysjolonensis Grinnell, Proc. Biol. Soc. Washington, 32:203, 31 December. Measurements ofholotype. - Total length, 312; length oftail, 185; length ofhind foot, Holotype. - Adult male, skin and skull, 45.4; greatest length of skull, 40.45; width MVZ 29087, from valley floor, 1 mi SW across bullae, 25.35; breadth across maxil­ Jolon, Monterey Co., California; obtained lary arches, 20.80; nasal length, 14.55; in­ on 18 October 1918, by Joseph Dixon. terorbital breadth, 12.55. Measurements ofholotype. - Total length, Distribution. - Ranges from near sea level 310; length oftail, 185; length ofhind foot, to about 1,300 ft, from the sea coast at Mon­ 44; length of ear (crown), 15; weight, 82.9 terey Bay northeastward to San Jose, Santa g; greatest length ofskull, 43.0; width across Clara Co., and southward in the Salinas Val­ bullae, 27.0; length ofnasals, 16.3; breadth ley to the vicinity ofSoledad, Monterey Co. across maxillary arches, 22.9; width ofmax­ (Grinnell, 1922). illary arch at middle, 5.7; greatest width of Remarks. - External and cranial mea­ rostrum near end, 4.6. surements for 10 adult males and females Distribution. -Chiefly occupies Lower were given by Grinnell (1922). Sonoran associations in west central Cali- TAXONOMY 55 fornia in the Salinas Valley, from near 45; length ofear (crown), 11; greatest length Peachtree and San Lucas, Monterey Co., of skull, 42.35; width across bullae, 25.95; southward at least to Creston, San Luis breadth across maxillary arches, 23.55; na­ Obispo Co. sallength, 15.00; interorbital breadth, 13.45. Remarks. -External and cranial mea­ Distribution. -Occurs in Bromus, Atri­ surements for 10 adults were given by Grin­ plex, and Ephedra associations along the nell (1922). southwestern margin of the San Joaquin Valley, Kern Co., westward to the Carrizo Plain, San Louis Obispo Co., and the upper Dipodomys heermanni morroensis Cuyama Valley, Santa Barbara Co., Cali­ (Merriam, 1907) fornia. Remarks. - Measurements of six adults 1907. Perodipus morroensis Merriam, Proc. Biol. were listed by Grinnell (1922). Soc. Washington, 20:78, 22 July.

Holotype. -Adult female, skin and skull, Dipodomys heermanni tularensis USNM 31626/43499, from Morro [south (Merriam, 1904) side Morro Bay, about 4 mi S oftown; Grin­ nell, 1922], San Luis Obispo Co., Califor­ 1904. Perodipus agilis tularensis Merriam, Proc. nia; obtained on 11 November 1891, by Biol. Soc. Washington, 17:143, 14 July. Edward W. Nelson. Measurements ofholotype. - Total length, Holotype. - Adult female, skin and skull, 300; length oftail, 182; length ofhind foot, USNM 127158, from Alila [Earlimart], Tu­ 45; greatest length of skull, 40.00; width lare Co., California; obtained on 23 June across bullae, 23.60; breadth across maxil­ 1903 by Luther 1. Goldman. lary arches, 21.45; nasal length, 14.80; in­ Measurements ofholotype.-Total length, terorbital breadth, 13.10. 308; length oftail, 182; length ofhind foot, Distribution. - Limited to coastal plant 41; length ofskull, 40.10; width across bul­ communities on sandy soils along the shore lae, 24.80; breadth across maxillary arches, of Morro Bay, San Louis Obispo Co., Cal­ 20.65; nasal length, 14.75; interorbital ifornia. breadth, 10.70. Remarks. - Boulware (1943) first treated Distribution. -Grassland and savanna morroensis as a subspecies ofD. heermanni. associations on the floor ofthe San Joaquin Grinnell (1922) gave external and cranial Valley, California, from the vicinity ofTra­ measurements for 10 adults. cy, San Joaquin Co., on the north to the foothills of the Tehachapi and Temblor ranges, Kern Co., on the south and south­ Dipodomys heermanni swarthi west, respectively. (Grinnell, 1919) Remarks. -Specimens from the Temblor Mountains, 3,000 ft, 12 mi W McKittrick, 1919. Perodipus swarthi Grinnell, Univ. Cali­ were assigned to tularensis by Grinnell fornia Publ. Zool., 21 :44, 29 March. (1922), but are surrounded geographically by specimens Grinnell (1922) assigned to Holotype. -Adult male, skin and skull, D. h. swarthi. MVZ 14440, from 7 mi SE Simmler, Carri­ zo Plain, San Luis Obispo Co., California; obtained on 26 May 1911, by H. S. Swarth Dipodomys ingens and W. L. Chandler. Measurements ofholotype. - Total length, Diagnosis. -Dipodomys ingens is the 313; length oftail, 187; length ofhind foot, largest kangaroo rat with five toes on the 56 WILLIAMS ET AL. hind feet, and has a relatively short tail and Remarks. - Measurements for eight sub­ short ears; other diagnostic traits are the adult and adult specimens were given by wide maxillary arches (width at mid-length Grinnell (1922). averaging 5.9 mm). Comparisons. -Giant kangaroo rats can be readily distinguished from the other large­ Dipodomys ingens (Merriam, 1904) sized kangaroo rats, D. deserti, D. nelsoni, and D. spectabilis, by having five, rather 1904. Perodipus ingens Merriam, Proc. BioI. Soc. than four, toes on each hind foot and by its Washington, 17:141, 14 July. greater average weight and relatively shorter Holotype. - Adult male, skin and skull, tail (about 128% of head and body length, USNM 128805, from Painted Rock, 20 r= range about 112 to 135%, D. F. Williams, 25.5] mi SE Simmler, Carrizo Plain, San unpubl. data). From the sympatric kanga­ Luis Obispo Co., California; obtained on 6 roo rats, D. heermanni and D. nitratoides, August 1903 by Luther J. Goldman. D. ingens can be distinguished by having a Measurements ofholotype. - Total length, hind foot longer than 47 mm (rarely to 46 360; length oftail, 191; length ofhind foot, mm in immature individuals), whereas the 54; greatest length ofskull, 48; width across length ofthe hind foot in D. heermanni does bullae, 30.5; length of nasals, 18; breadth not exceed 46 mm, and is usually less than across maxillary arches, 23.5. 45 mm. The huge hind feet will distinguish Remarks. - Dipodomys ingens appears to evenjuvenile giant kangaroo rats from sym­ be most closely related to the sympatric spe­ patrie species. The hind foot of D. nitra­ cies, D. heermanni. toides has only four toes and its length does not exceed 39 mm (Grinnell, 1922). The skull of D. ingens is large and broad, aver­ Dipodomys merriami aging 27 mm or more across the maxillary arches, a width much larger than that ofall Diagnosis. -A small-sized kangaroo rat other kangaroo rats except D. spectabilis, (head and body length averages less than which averages from 25.8 to 26.9 mm about 105 mm and greatest length of skull (Grinnell, 1922; Nader, 1978). The average averages less than about 37 mm in all pop­ greatest length ofthe skull ofD. ingens, 45.9 ulations; Lidicker, 1960) with four toes on (Grinnell, 1932), is equaled only by D. de­ the relatively slender hind feet, and with a serti and D. spectabilis (Nader, 1978). tail terminating in a crest and tuft ofdusky Distribution. -Dipodomys ingens occu­ or blackish-brown hairs; mastoid bullae pies a narrow band ofgently sloping ground usually relatively more inflated and rostrum along the western edge of the San Joaquin narrower than in most other species; ros­ Valley, California. Records of occurrence trum not decidedly shortened and face not extend from the base of the Tehachapi noticeably narrower than most other spe­ Mountains on the south to a point about 16 CIes. km S Los Banos, Merced Co. Inhabited ar­ Comparisons.-Dipodomys merriami is eas west ofthe San Joaquin Valley include most similar in size and appearance to the the Carrizo and Elkhorn plains west of the other small-sized, four-toed species, D. ni­ Temblor Mountains, and the upper Cuya­ tratoides; all other four-toed species are ma Valley, adjacent to and nearly contigu­ considerably larger and most have a prom­ ous with the Carrizo Plain, and scattered inent tuft ofwhite hairs at the tip ofthe tail. colonies in the Ciervo, Kettleman, Panoche, D. merriami differs from D. nitratoides in and Tumey hills, and the Panoche Valley generally being larger in size, and in having (Grinnell, 1922; Hall, 1981; Williams, in a longer and wider rostrum (nasal length press; Williams and Kilburn, 1991). usually averages greater than 13.1 in D. TAXONOMY 57 merriami and less than 12.3 mm in D. ni­ hua, northeastern Durango, southern Coa­ tratoides; Hoffmann, 1975; Lidicker, 1960). huila and central Nuevo Leon (Lidicker, Distribution. -Dipodomys merriami is 1960). widely distributed in desert and arid grass­ Remarks. - Lidicker (1960) gave mea­ land associations in southwestern North surements and comparisons for D. m. am­ America, extending from northwestern Ne­ biguus; Anderson (1972) listed means and vada southward through southeastern Cal­ ranges ofmeasurements for a sample of 37 ifornia, the length of the Baja California from Chihuahua. Peninsula, and along the mainland ofMexi­ co through Sonora to northern Sinaloa; and ranging eastward across southern Nevada Dipodomys merriami annulus to extreme southeastern Utah, western and Huey, 1951 southern Arizona, central and southern New Mexico to western Texas; and thence south­ 1951. Dipodomys merriami annulus Huey, ward through the plateau regions ofMexico Trans. San Diego Soc. Nat. Hist., 11 :224, 30 to Aguascalientes and Zacatecas (Hall, April. 1981). Remarks. - Hall (1981) treated Dipodo­ Holotype. -Adult female, skin and skull, mys margaritae as a species distinct from SDSNH 15522, from El Barril, GulfofCal­ D. merriami, although Lidicker (1960) con­ ifornia, lat. 28°20'N, long. 112°50'W, Baja sidered it to be a subspecies ofD. merriami; California, Mexico; obtained on 24 March we concur with Lidicker. Lidicker (1960) 1928 by Laurence M. Huey. provided measurements for each ofthe sub­ Measurements ofholotype. - Total length, species ofD. merriami. 251; length oftail, 150; length ofhind foot, 35; length ofear, 10; greatest length ofskull, 37.9; width across bullae, 23.8; length of nasals, 14.1; breadth across maxillary arch­ Dipodomys merriami ambiguus es, 21.0; width ofmaxillary arch at middle, Merriam, 1890 5.5. Distribution. - Found along the coastal 1890. Dipodomys ambiguus Merriam, N. Amer. plain of the Gulf of California from near Fauna, 4:42, 8 October. Bahia Los Angeles on the north to Bahia Holotype. -Adult male, skin and skull, Santa Teresa on the south (Huey, 1951). USNM 18147125045, from El Paso, El Paso Remarks. -Huey (1951) and Lidicker Co., Texas; obtained by Vernon Bailey on (1960) gave measurements for samples of 13 December 1889. D. m. annulus. Measurements ofholotype. - Total length, 233; length oftail, 133; length ofhind foot, 37; length ofear (crown), 7, (notch, dry), 12; Dipodomys merriami arenivagus greatest length of skull, 36.7; width across Elliot, 1904 bullae, 23.0; length ofnasals, 13.6; breadth across maxillary arches, 20.8; interorbital 1904. Dipodomys m[erriami]. arenivagus Elliot, breadth, 13.5; length ofnasals, 13.6; length Field Columbian Mus., Publ. 87, Zool. Ser., ofrostrum, 19.7; length ofmaxillary tooth­ 3:249, 7 January. row, 4.3; depth ofcranium, 10.6. Distribution. -Occurs in the northern Holotype. -Adult skin and skull, FMNH parts of the central Mexican deserts, from 10733, from San Felipe, Baja California, western New Mexico and western Texas Mexico; obtained on 1 April 1902 by Ed­ southward to southern and eastern Chihua- mund Heller. 58 WILLIAMS ET AL.

Measurements ofholotype. - Total length, Dipodomys merriami brunensis 225; length oftail, 134; length ofhind foot, Huey, 1951 36; length ofear (notch), 15; greatest length ofskull, 34; width across bullae, 22; length 1951. Dipodomys merriami brUliensis Ruey, ofnasals, 12; length ofmaxillary toothrow, Trans. San Diego Soc. Nat. Rist., 11 :225, 30 3; greatest width ofrostrum, 5; palatal length, April. 11. Holotype. -Adult male, skin and skull, Distribution. -Arid communities east of SDSNH 6904, from Llano de San Bruno, the Sierra San Pedro Martir and Sierra Jua­ Baja California, Mexico; obtained by Lau­ rez, from just south ofthe U.S. border and rence M. Huey on 24 March 1928. west of the Colorado River Delta, south­ Measurements ofholotype. - Total length, ward to San Felipe, all in Baja California, 252; length oftail, 152; length ofhind foot, Mexico (Huey, 1951). 37; length ofear (crown), 10; greatest length Remarks. -Huey (1951) provided means ofskull, 37.6; length ofnasals, 13.8; breadth and ranges of measurements for five males across maxillary arches, 19.8; width ofmax­ and five females. Elliot's paper was listed as illary arch at middle, 4.8. being published in December, 1903, but was Distribution. -Costal areas along the Gulf actually published on 7 January 1904. of California from near El Valle de Yaqui, about 12 km NW Santa Rosalia, southward to the southern end of Bahia Concepcion, Baja California (Huey, 1951). Dipodomys merriami atronasus Remarks. - Huey (1951) gave means and Merriam, 1894 ranges of measurements for five males and three females; Lidicker (1960) listed average 1894. Dipodomys merriami atronasus Merri­ measurements for an unspecified sample am, Proc. BioI. Soc. Washington, 9:113, 21 size. June. Holotype. - Adult male, skin and skull, USNM 50276, from Hacienda La Parada Dipodomys merriami collinus [about 25 mi NW San Luis Potosi], San Luis Lidicker, 1960 Potosi, Mexico; obtained by Edward W. Nelson on 20 August 1892. 1960. Dipodomys merriami collinus Lidicker, Measurements ofholotype. - Total length, Univ. California Pubi. ZooI., 67: 194, 4 Au­ gust. 250; length oftail, 162; length ofhind foot, 40; length of nasals, 13.6; breadth across Holotype. -Adult female, skin and skull, maxillary arches, 21.6; length of maxillary MVZ 123455, from 3.25 mi S, 2.25 mi E toothrow, 4.8. Scissors Crossing, Earthquake Valley, San Distribution. -Occupies the southern Diego Co., California; obtained on 10 Sep­ portions ofthe central Mexican desert, from tember 1958 by William Z. Lidicker, Jr. central Durango and southeastern Coahui­ Measurements ofholotype. - Total length, la, southward to southeastern Aguascalien­ 252; length oftail, 148; length ofhind foot, tes, southwestern San Luis Potosi, and ex­ 39; length ofear, 14; weight, 39.3 g; greatest treme southwestern Tamaulipas (Lidicker, length of skull, 35.8; width across bullae, 1960). 22.55; breadth across maxillary arches, Remarks.-Dalquest (1953) gave means 18.65; nasal length, 12.9; interorbital for measurements oftwo samples from San breadth, 13.0. Luis Potosi; Lidicker (1960) listed average Distribution. - Occupies areas in San Fe­ measurements for specimens ofD. m. atro­ lipe, Earthquake, and Mason (La Puerta) nasus. valleys in eastern San Diego Co., and TAXONOMY 59

Aguanga Valley in southern Riverside Co., Holotype. -Adult female, skin and skull, California (Lidicker, 1960). USNM 79053, from San Jose Island, Gulf Remarks. - Lidicker (1960) listed aver­ of California, Baja California Sur, Mexico; age measurements for an unspecified num­ obtained on 6 August 1895 by J. E. Mc­ ber of individuals of this subspecies. Lellan. Measurements ofholotype. - Total length, 243; length oftail, 143; length ofhind foot, Dipodomys merriami frenatus 39; greatest length ofskull, 36.0; width across Bole, 1936 bullae, 22.7; length ofnasals, 13.2; breadth across maxillary arches, 21.3; interorbital 1936. Dipodomys merriami frenatus Bole, Sci. breadth, 13.0; length ofrostrum, 20.1; length PubIs., Cleveland Mus. Nat. Rist., 5: 1, 17 Jan­ of maxillary toothrow, 4.4; depth of crani­ uary. um, 10.5. Holotype. -Adult female, skin and skull, Remarks. -Lidicker (1960) included UMMZ 121258 from Toquerville, 3,200 ft, samples of insularis in his analysis of geo­ Washington Co., Utah; obtained on 14 May graphic variation in D. merriami because of 1934 by F. 1. Tobin. the possibility that it might prove to be a Measurements ofholotype. - Total length, subspecies ofthe latter species. He retained 207; length oftail, 113; length ofhind foot, specific rank for D. insularis, however, be­ 35; occipitonasallength ofskull, 33.5; width cause of its greater structural divergence across bullae, 21.9; length of nasals, 12.7; from the norm among samples of all sub­ breadth across maxillary arches, 19.5; width species of D. merriami. Best and Janecek of maxillary arch at middle, 5.2; width of (1992) found that insularis was significantly nasals in front of incisors, 2.7; length of different in several morphological traits from maxillary toothrow, 4.1. samples ofD. merriami from the mainland, Distribution. -Occupies arid communi­ but they classified it as a subspecies of D. ties from the Virgin River Valley in Wash­ merriami based on allozymic similarities. ington Co., Utah, westward to the edge of Lidicker (1960) found considerable sexual the Beaver Dam Mountains, north as far as dimorphism, with males being larger than Veyo, east to Springdale, and southward to females; he provided external and cranial the Kanab Plateau in northwestern Arizona measurements for samples ofmales and fe­ (Lidicker, 1960). males. Remarks. - Durrant and Setzer (1945) considered frenatus to be a synonym of D. m. vulcani, but Lidicker (1960) found that Dipodomys merriami margaritae the two subspecies were recognizably dis­ Merriam, 1907 tinct. Hoffmeister (1986) considered it to be most like, and possibly a synonym of, D. 1907. Dipodomys margaritae Merriam, Proc. m. merriami. Hoffmeister (1986) listed sta­ BioI. Soc. Washington, 20:76, 22 July. tistics for measurements of12 males and 11 Holotype. - Young adult male, skin and females from Arizona. The holotype was skull, USNM 146058, from [Santa] Mar­ originally in the Cleveland Museum ofNat­ garita Island, Baja California Sur, Mexico; ural History (CMNH 6771). obtained on 1 December 1905 by Edward W. Nelson and Edward A. Goldman. Dipodomys merriami insularis Measurements ofholotype. - Total length, Merriam, 1907 234; length oftail, 144; length ofhind foot, 38; greatest length ofskull, 34.6; width across 1907. Dipodomys insularis Merriam, Proc. BioI. bullae, 21.7; length ofnasals, 12.2; breadth Soc. Washington, 20:77, 22 July. across maxillary arches, 19.5; length ofros- 60 WILLIAMS ET AL. trum, 19.0; length of maxillary toothrow, Dipodomys merriami melanurus 4.7; depth ofcranium, 11.1. Merriam, 1893 Distribution. -Occurs only on Santa Margarita Island off the Pacific Coast, in 1893. Dipodomys merriami melanurus Merri­ Baja California Sur. am, Proc. California Acad. Sci., Ser. 2, 3:345, Remarks. -Lidicker (1960), in his anal­ 5 June. ysis ofgeographic variation in D. merriami, 1951. Dipodomys merriami llanoensis Huey, concluded that margaritae did not warrant Trans. San Diego Soc. Nat. Hist., 11 :226, 30 April. specific status and assigned it as a subspecies ofD. merriami. Huey (1964) concluded that Holotype. -Adult male, CAS 539, from "specific instead of subspecific rank seems San Jose del Cabo, Baja California Sur, preferable." Hall (1981) listed margaritae Mexico; obtained by Walter E. Bryant on as a species without reference to Huey (1964) 19 March 1892. or Lidicker (1960). We believe Lidicker's Measurements ofholotype. - Total length, (1960) conclusions are based on the most 239; length oftail, 144; length ofhind foot, comprehensive investigations, and concur 35. with his assignment of margaritae as a sub­ Distribution. - From near San Jorge on the species of D. merriami. Huey (1951) and north end of the Magdalena Plain, south­ Lidicker (1960) listed measurements for ward through the Cape region ofBaja Cal­ samples of this subspecies. ifornia Sur (Lidicker, 1960). Remarks. - The holotype was destroyed in the San Francisco earthquake and fire of Dipodomys merriami mayensis 1906. After the fire, a new catalog was start­ Goldman, 1928 ed; thus, CAS 539 now represents a different specimen. Lidicker (1960) synonymized 1928. Dipodomys merriami mayensis, Proc. BioI. llanoensis with melanurus, because the for­ Soc. Washington, 41:141, 15 October. mer exhibited intermediate characteristics Holotype. - Adult male, skin and skull, between typical melanurus to the south and USNM 96437, from Alamos, Sonora, Me­ platycephalus to the north. Huey (1964) re­ xico; obtained on 19 December 1898 by tained llanoensis "as a means ofcataloging Edward A. Goldman. the geographic variation shown by the spec­ Measurements ofholotype. - Total length, imens." Hall (1981) cited Huey (1964) as 240; length oftail, 138; length ofhind foot, his reason for recognizing llanoensis. We 37; greatest length of skull, 35.8; occipito­ concur with Lidicker (1960). nasal length ofskull, 34.3; width across bul­ lae, 22.8; length of nasals, 13.1; breadth across maxillary arches, 20.5; interorbital Dipodomys merriami merriami breadth, 12.4; length ofrostrum, 19.5; length Mearns, 1890 of maxillary toothrow, 4.1; depth of crani­ um, 10.9; least width ofsupraoccipital, 1.6. 1890. Dipodomys merriamiMearns, Bull. Amer. Distribution. -Occupies open shrub1ands Mus. Nat. Hist., 2:290, 21 February. of the coastal plains from the Rio Yaqui, 1894. Dipodomys simiolus Rhoads, Proc. Acad. Nat. Sci. Philadelphia, 45:410, 27 January. southern Sonora, southward to extreme 1894. Dipodomys similis Rhoads, Proc. Acad. northern Sinaloa to the south side of the Nat. Sci. Philadelphia, 45:411, 27 January. Rio del Fuerte (Lidicker, 1960). 1894. Dipodomys merriami nevadensis Merri­ Remarks. - Lidicker (1960) gave average am, Proc. BioI. Soc. Washington, 9:111, 21 measurements for an unspecified number of June. specimens. 1894. Dipodomys merriami nitratus Merriam, TAXONOMY 61

Proc. BioI. Soc. Washington, 9:112, 21 June. Holotype. - Adult female, skin and skull, 1904. Dipodomys merriami mortivallis Elliot, USNM 63188, from Tiburon Island, Gulf Field Columbian Mus., PubI. 87, ZooI. Ser., ofCalifornia, Sonora, Mexico; obtained by 3:250, 7 January. J. W. Mitchell on 23 December 1895. 1907. Dipodomys merriami kernensis Merriam, Measurements ofholotype. - No external Proc. BioI. Soc. Washington, 20:77, 22 July. 1937. Dipodomys merriami regillus Goldman, measurements recorded; greatest length of Proc. BioI. Soc. Washington, 50:75, 22 June. skull, 36.1; width across bullae, 22.4; length of nasals, 12.9; breadth across maxillary Holotype. -Adult male, skin and skull, arches, 19.8; interorbital breadth, 12.3; AMNH 2394, from New River, between length ofrostrum, 19.0; length ofmaxillary Phoenix and Prescott, Maricopa Co., Ari­ toothrow, 4.8; depth ofcranium, 10.2. zona; obtained by Edgar A. Mearns on 16 Distribution. -Occurs only on Tiburon May 1885. Island, Gulf of California, Sonora (Hall, Measurements ofholotype. - Total length, 1981). 259; length oftail, 149; length ofhind foot, Remarks. - Lidicker (1960) listed aver­ 36; length ofear (crown), 10, (meatus), 13. age measurements for a sample ofunspeci­ Distribution. - Extends from northwest­ fied size. ern Nevada, southward through the deserts of southeastern California, extreme south­ eastern Utah, western and southern Arizona Dipodomys merriami olivaceus as far east as the Pe10ncillo Mountains, and Swarth, 1929 to the Rio Yaqui in southern Sonora (Li­ dicker, 1960). 1929. Dipodomys merriami olivaceus Swarth, Remarks. - Durrant and Setzer (1945) Proc. California Acad. Sci., ser. 4, 18:356,26 gave measurements for 6 males and 2 fe­ April. males from Utah; Hall (1946) listed mea­ Holotype. -Adult male, skin and skull, surements for 10 males and 10 females from CAS 6235, from Fairbank, Cochise Co., Ar­ Nevada; and Hoffmeister (1986) gave sta­ izona; obtained by Sam Davidson on 28 tistics for measurements oftwo samples (18 October 1928. males, 13 females; 6 males, 10 females) from Measurements ofholotype. - Total length, Arizona. Mearns (1890) referred to the skull 243; length oftail, 141; length ofhind foot, in his description ofthe type specimen; but 37; length ofear, 12; greatest length ofskull, provided no measurements. Goodwin 36.45; width across bullae, 22.80; length of (1953) stated that the skull was not cata­ nasals, 14.40; breadth across maxillary logued into the collections at the American arches, 18.90; length ofmaxillary toothrow, Museum of Natural History, and was pre­ 4.65; depth of cranium, 12.15; width of sumed to be lost. Goodwin (1953) listed the maxillary arch at middle, 5.0; greatest width total length ofthe ho10type as 259 mm but of rostrum near end, 3.2. Mearns (1890) listed it as 281 mm, the dif­ Distribution. - Desert associations from ference apparently being that Mearns mea­ southeastern Arizona in Cochise and Santa sured to the end of the hairs on the tail. Cruz counties and western New Mexico east of the Peloncillo Mountains, eastward to eastern Luna and western Dona Ana coun­ Dipodomys merriami mitchelli ties and southward around the northern end Mearns, 1897 of the Sierra Madre Occidental into north­ eastern Sonora on the west and central Chi­ 1897. Dipodomys mitchelli Mearns, Proc. U.S. huahua on the east (Hall, 1981). Natl. Mus., 19:719, 30 July. Remarks. -Anderson (1972) listed mea- 62 WILLIAMS ET AL. surements for a sample of 19 from Chihua­ by Edward W. Nelson and Edward A. Gold­ hua; Hoffmeister (1986) listed statistics for man. measurements of 15 males and 11 females Measurements ofholotype. - Total length, from Arizona; and Swarth (1929) listed 238; length oftail, 145; length ofhind foot, means and ranges ofmeasurements for two 38; greatest length ofskull, 35.6; occipitona­ samples of olivaceus from Arizona. sal length, 34; width across bullae, 23.6; length of nasals, 12.5; breadth across max­ illary arches, 20.9; interorbital breadth, 13.7; Dipodomys merriami parvus length ofrostrum, 18.7; length ofmaxillary Rhoads, 1894 toothrow, 4.5; depth of cranium, 11.0. Distribution. - Found along the Pacific 1894. Dipodomys parvus Rhoads, Amer. Nat., slope from the southern end of the Sierra 28:70, January. San Pedro Martir and San Fernando, south­ Holotype. -Adult female, skin and skull, ward to about lat. 26°l5'N, including the ANSP 8213, from San Bernardino [Reche entire Viscaino Desert, but not extending to Canyon, 4 mi SE Colton], San Bernardino the Gulf coastal regions (Lidicker, 1960). Co., California; obtained on 12 June 1892 Remarks. - Huey (1951) listed means and by R. B. Herron. ranges for males and females; Lidicker Measurements ofholotype. - Total length, (1960) listed ranges ofmeans for unspecified 248; length oftail, 154; length ofhind foot, samples. Lidicker (1960) synonymized 35; length ofear (dry, from crown), 10; bas­ semipallidus with platycephalus because he ilar length ofskull, 21; width across bullae, found them to be intergrades between quin­ 22.5; length of nasals, 13; interorbital tinensis and platycephalus. Huey (1951, breadth, 13; crown length of mandibular 1964) distinguished semipallidus entirely on toothrow, 3.6. the basis ofslightly darker coloration, which Distribution. - Found in the San Bernar­ Lidicker (1960) did not consider worthy of dino and San Jacinto valleys, in San Ber­ subspecific recognition. Hall (1981) re­ nardino and Riverside counties, California. tained semipallidus for reasons not stated. Remarks. -Grinnell (1922) amended the We concur with Lidicker (1960) that the type locality and provided measurements of slight and inconsistent differences in color 10 adult and subadult specimens. Lidicker attributed to semipallidus by Huey (1951, (1960) noted that parvus differed from typ­ 1964) do not merit recognition as a separate ical samples of merriami in being much subspecies. smaller, and remarked that it was one ofthe most highly differentiated subspecies of D. merriami. He believed that it had nearly Dipodomys merriami quintinensis achieved species rank. Huey, 1951

1951. Dipodomys merriami quintinensis Huey, Dipodomys merriami platycephalus Trans. San Diego Soc. Nat. Hist., 11 :222, 30 Merriam, 1907 April. Holotype. - Adult male, skin and skull, 1907. Dipodomys platycephalus Merriam, Proc. SDSNH 4205, from 5 mi E San Quintin, BioI. Soc. Washington, 20:76, 22 July. Baja California Mexico; obtained on 9 April 1927. Dipodomys merriami semipallidus Huey, 1923 by Laurence M. Huey. Trans. San Diego Soc. Nat. Hist., 5:66,6 July. Measurements ofholotype. - Total length, Holotype. -Adult male, skin and skull, 240; length oftail, 148; length ofhind foot, USNM 139882, from Calmalli, Baja Cali­ 39; length ofear (crown), 10; greatest length fornia, Mexico; obtained on 1October 1905 of skull, 36.4; width across bullae, 22.6; TAXONOMY 63 length of nasals, 13.0; breadth across max­ es of measurements for samples of males illary arches, 20.1; width of maxillary arch and females from Baja California. at middle, 5.1. Distribution. - Occurs on the San Quintin Plain along the Pacific coast, from near San­ Dipodomys merriami vulcani to Domingo, southward to the region of El Benson, 1934 Rosario, Baja California. Remarks.-Huey (1951) gave means and 1934. Dipodomys merriami vulcani Benson, ranges of measurements for five males and Proc. BioI. Soc. Washington, 47:181, 2 Octo­ five females. ber. Holotype. -Adult male, skin and skull, MVZ 56002, from lower end Toroweap Dipodomys merriami trinidadensis Valley, about 0.5 mi E Vulcan's Throne, Huey, 1951 Mohave Co., Arizona; obtained by Annie M. Alexander on 11 November 1932. 1951. Dipodomys merriami trinidadensis Huey, Measurements ofholotype. - Total length, Trans. San Diego Soc. Nat. Hist., 11:220, 30 241; length oftail, 138; length ofhind foot, April. 39; length of ear (crown), 10; weight, 39.5 Holotype. -Adult male, skin and skull, g; greatest length ofskull, 35.3; width across SDSNH 11531, from Aguajito Spring, El bullae, 23.45; breadth across maxillary Valle de la Trinidad, Baja California, Mexi­ arches, 19.6; nasal length, 12.85; interor­ co; obtained by Laurence M. Huey on 13 bital breadth, 12.9. March 1936. Distribution. -Known only from Toro­ Measurements ofholotype. - Total length, weap Valley, Mohave Co., Arizona. 243; length oftail, 144; length ofhind foot, Remarks. - Durrant and Setzer (1945) 38; length ofear (crown), 10; greatest length listed measurements for six males and three of skull, 35.5; width across bullae, 22.1; females; Hoffmeister (1986) listed statistics length of nasals, 13.0; breadth across max­ for measurements of five males and six fe­ illary arches, 19.5; width of maxillary arch males. at middle, 5.0. Distribution. - Lidicker (1960) outlined a discontinuous range for trinidadensis: the Dipodomys microps Jacumba Valley and Mountain Springs re­ gion ofsouthern San Diego Co., California, Diagnosis. -A medium-sized kangaroo and nearby Baja California; and from El rat (head and body length averages from Valle de la San Rafael and El Valle de la about 108 to 120 mm) with a relatively short Trinidad along the western border of the tail (averaging about 135% ofhead and body Sierra Juarez, Baja California, Mexico. length), with five toes on the hind feet, a Remarks. - Lidicker (1960) discussed the narrow face, reflected in the breadth across possibility that trinidadensis was polyphy­ the maxillary arches, and with lower inci­ letic, and that the diagnostic characters ex­ sors that are flattened on their anterior faces hibited by the disjunct populations were ex­ and chisel-like. amples of convergence. A more likely Comparisons. -Other species ofDipodo­ hypothesis, according to Lidicker (1960), is mys that are most similar in size and general that the populations are relicts ofa formerly appearance to D. microps, and which oc­ more widely-distributed population. The cupy the same geographic areas, are D. ordii northern populations of this taxon were and D. merriami. Other, larger-sized, sym­ classified as D. m. simiolus by Grinnell patric species are D. panamintinus and D. (1922). Huey (1951) listed means and rang- deserti. D. agilis occupies a contiguous area 64 WILLIAMS ET AL. along the southwestern margin of the geo­ 4,300 £1, Great Salt Lake, Box Elder Co., graphic range of D. microps, and may be Utah; obtained on 1 June 1937 by Alfred marginally sympatric. From D. merriami M. Bailey and Robert 1. Niedrach. and D. deserti, D. microps differs in having Measurements ofholotype. - Total length, five, rather than four toes on the hind feet. 286; length oftail, 170; length ofhind foot, From D. agilis and D. panamintinus, D. mi­ 44; greatest length ofskull, 38.5; width across crops differs in being decidedly smaller and bullae, 25.0; length ofnasals, 12.8; breadth having chisel-like rather than awl-like lower across maxillary arches, 20.6; interorbital incisors. From D. ordii, D. microps differs breadth, 12.5; length ofrostrum, 19.2; length in having chisel-like, rather than awl-like of maxillary toothrow, 4.3; depth of crani­ lower incisors, and a narrower face, reflect­ um, 11.5. ed in a shorter breadth across the maxillary Distribution. -Confined to Gunnison Is­ arches in sympatric populations (Grinnell, land, Great Salt Lake, Box Elder Co., Utah 1922; Hall and Dale, 1939; Setzer, 1949). (Hall, 1981). Distribution. -Occurs throughout most Remarks. - Hall and Dale (1939) listed of the Great Basin desert ofwestern North measurements for three males and three fe­ America, from southeastern Oregon and ad­ males and noted that members ofthis sub­ jacent Owyhee Co., Idaho, southward species were the largest of the species. through Nevada and the Mojave Desert re­ gion ofsoutheastern California, eastward in Dipodomys microps aquilonius northwestern Utah to the Great Salt Lake. Disjunct populations occur in extreme Willett, 1935 southwestern Utah and adjacent north­ 1935. Dipodomys microps aquilonius Willett, J. western Arizona, and in the vicinity ofJosh­ Mamm., 16:63, 14 February. ua Tree National Monument in Riverside and San Bernardino counties, California Holotype. -Adult female, skin and skull, (Csuti, 1979; Hall and Dale, 1939). LACM 3935, from 3 mi E Eagleville, Mo­ Remarks. -Hall and Dale (1939) provid­ doc Co., California; obtained by George ed the only systematic review ofDipodomys Willett on 4 June 1934. microps; they listed measurements for each Measurements ofholotype. - Total length, ofthe subspecies then known (two were de­ 272; length oftail, 156; length ofhind foot, scribed subsequent to their revision). Dur­ 41; greatest length ofskull, 36.1; width across rant (1952) and Hall (1946) gave measure­ bullae, 23.9; width ofmaxillary arch at mid­ ments for subspecies found in Utah and die, 3.2. Nevada, respectively. Csuti (1979) re­ Distribution. - Lower portions of the viewed patterns of variation in structure, Great Basin Desert in northeastern Califor­ plasma and tissue proteins, physiology, and nia and adjacent Nevada, from the Surprise ecology, but provided no taxonomic review Valley on the northwest to the south end of of the subspecies. Pyramid Lake on the southeast. Remarks.-Hall and Dale (1939) gave measurements for four males and two fe­ Dipodomys microps alfredi males. Goldman, 1937 Dipodomys microps bonnevillei 1937. Dipodomys microps alfredi Goldman, Proc. BioI. Soc. Washington, 50:221, 28 De­ Goldman, 1937 cember. 1937. Dipodomys microps bonnevillei Gold­ Holotype. - Adult female, skin and skull, man, Proc. BioI. Soc. Washington, 50:222,28 USNM 262846, from Gunnison Island, December. TAXONOMY 65

Holotype. - Adult female, skin and skull, Remarks.-Hall and Dale (1939) listed USNM 31894/43755, from Kelton, about the type specimen as USNM 243093, but 4,300 £1, Box Elder Co., Utah; obtained on the number of the designated type in the 7 November 1891 by Vernon Bailey. U.S. National Museum is number 243101; Measurements ofholotype. - Total length, measurements are for the latter numbered 260; length oftail, 156; length ofhind foot, specimen. Stock (1970) regarded woodburyi 41; greatest length ofskull, 36.0; width across as indistinguishable from celsus. bullae, 23.7; length ofnasals, 11.9; breadth across maxillary arches, 18.9; interorbital breadth, 11.2; length ofrostrum, 17.8; length Dipodomys microps centralis of maxillary toothrow, 4.6; depth of crani­ Hall and Dale, 1939 um, 11.1. Distribution. - Great Basin Desert of 1939. Dipodomys microps centralis Hall and northeastern Nevada and northwestern Dale, Occas. Papers Mus. ZooI., Louisiana State Utah, corresponding closely with the former Univ., 4:52, 10 November. outline of the Pleistocene Lake Bonneville. Holotype. -Adult male, skin and skull, Remarks. -Hall and Dale (1939) listed MVZ 70817, from 4 mi SE Romano, Dia­ means and ranges of measurements for 8 mond Valley, Eureka Co., Nevada; ob­ males and 10 females; Durrant and Setzer tained by William B. Richardson on 3 June (1945) gave measurements for 17 males and 1936. 15 females. Measurements ofholotype. - Total length, 282; length oftail, 164; length ofhind foot, 43; length ofear, 13; weight, 72.8 g; greatest Dipodomys microps celsus length of skull, 37.65; width across bullae, Goldman, 1924 23.55; breadth across maxillary arches, 19.65; nasal length, 12.6; interorbital 1924. Dipodomys microps celsus Goldman, J. Washington Acad. Sci., 14:372, 19 September. breadth, 12.2. 1942. Dipodomys microps woodburyi Hardy, Distribution. - Great Basin Desert ofcen­ Proc. BioI. Soc. Washington, 55:89, 25 June. tral Nevada, from the Humboldt River Val­ ley southward to Pahute Mesa; from north­ Holotype. -Adult male, skin and skull, eastern Pershing Co. on the northwest, to USNM 243101, from 6 mi N Wolf Hole, Spring Valley on the northeast. 3,500 £1, Mohave County, Arizona; ob­ Remarks. - Measurements for three males tained on 16 October 1922 by Edward A. and three females were given by Hall and Goldman. Dale (1939). Measurements ofholotype. - Total length, 284; length oftail, 170; length ofhind foot, 44; greatest length ofskull, 37.9; width across bullae, 25.3; length ofnasals, 13.0; breadth Dipodomys microps idahoensis across maxillary arches, 19.9; interorbital Hall and Dale, 1939 breadth, 11.4; length ofrostrum, 18.5; length of maxillary toothrow, 5.1; depth of crani­ 1939. Dipodomys microps idahoensis Hall and um, 11.5. Dale, Occas. Papers Mus. ZooI., Louisiana State Univ., 4:53, 10 November. Distribution. - Occupies an apparently disjunct range in extreme southwestern Utah Holotype. -Adult male, skin and skull, in the Virgin River Valley and in adjacent MVZ 67568, from 5 mi SE Murphy, Owy­ northwestern Arizona north of the Colora­ hee Co., Idaho; obtained on 26 May 1935 do River and east to Kanab Creek (Hall and by Howard Twining. Dale, 1939). Measurements ofholotype. - Total length, 66 WILLIAMS ET AL.

201; length oftail, 155; length ofhind foot, Holotype. -Adult male, skin and skull, 43; length ofear, 13; greatest length ofskull, USNM 27176/34575, from Perognathus 37.15; width across bullae, 24.05; breadth Flat, Emigrant Gap, 5,200 ft, Panamint across maxillary arches, 20.75; nasal length, Mountains, Inyo Co., California; obtained 12.85; interorbital breadth, 12.55. by Vernon Bailey on 16 April 1891. Distribution. - Known only from south of Measurements ofholotype. - Total length, the Snake River in the Snake River Valley 288; length oftail, 156; length ofhind foot, of northwestern Owyhee Co., Idaho. 43; greatest length of skull, 38.20; width Remarks. - Measurements for three males across bullae, 24.25; breadth across maxil­ and one female were listed by Hall and Dale lary arches, 20.55; nasal length, 13.50; in­ (1939). terorbital breadth, 12.30. Distribution. - Limited to an area in Inyo Dipodomys microps leucotis Co., in the Panamint Valley, California from Goldman, 1931 the Panamint Mountains westward to the vicinity of Darwin. 1931. Dipodomys microps leucotis Goldman, Remarks. - Grinnell (1922) listed means Proc. BioI. Soc. Washington, 44: 135, 17 Oc­ and ranges of measurements for five indi­ tober. viduals of each sex. Hall and Dale (1939) Holotype. -Adult male, skin and skull, restricted the application of the name levi­ USNM 250036, from 6 mi W Colorado pes to the population in the Panamint Val­ River Bridge, Houserock Valley, N [= W] ley; the name had been applied widely in side Marble Canyon, Colorado River, 3,700 previous publications. ft, Coconino Co., Arizona; obtained on 8 June 1931 by Edward A. Goldman. Measurements ofholotype. - Total length, Dipodomys microps microps 281; length oftail, 170; length ofhind foot, (Merriam, 1904) 43.5; greatest length of skull, 35.5; occipi­ tonasal length, 34.6; width across bullae, 1904. Perodipus microps Merriam, Proc. BioI. 23.7; length of nasals, 12.0; breadth across Soc. Washington, 17:145, 14 July. maxillary arches, 19.6; interorbital breadth, 11.7; length ofrostrum, 18.4; length ofmax­ Holotype. -Adult male, skin and skull, illary toothrow, 4.9; depth ofcranium, 11.1. USNM 25288/32701, from Lone Pine, Ow­ Distribution. - Known only from a nar­ ens Valley, Inyo Co., California; obtained row strip between the Vermilion Cliffs and by Edward W. Nelson on 22 December the brink ofMarble Canyon ofthe Colorado 1890. River, in northwestern Arizona (Goldman, Measurements ofholotype. - Total length, 1931). 282; length oftail, 165; length ofhind foot, Remarks.-Hall and Dale (1939) listed 41; greatest length of skull, 36.05; width measurements for five male and five female across bullae, 22.50; breadth across maxil­ topotypes, and Hoffmeister (1986) gave lary arches, 18.35; nasal length, 12.10; in­ measurements for one male and three fe­ terorbital breadth, 11.75. males. Distribution. - Limited to the western Mojave Desert, California, in the Owens River watershed, from near the Nevada Dipodomys microps levipes border N of Benton Station, southward to (Merriam, 1904) near Victorville. Remarks. - Measurements were given by 1904. Perodipus microps levipes Merriam, Proc. Csuti (1979), Grinnell (1922), and Hall and BioI. Soc. Washington, 17:145, 14 July. Dale (1939). TAXONOMY 67

Dipodomys microps occidentalis across maxillary arches, 19.2; length of Hall and Dale, 1939 maxillary toothrow, 4.2; least width of su­ praoccipital, 2.5. 1939. Dipodomys microps occidentalis Hall and Distribution. -Great Basin Desert of Dale, Occas. Papers Mus. Zoo1., Louisiana State southeastern Oregon, and northwestern Ne­ Univ., 4:56, 10 November. vada; westernmost locality is Summer Lake, Holotype. - Adult female, skin and skull, Lake Co. Oregon, and the easternmost is MVZ 64119, from 3 mi S Schurz, 4,100 ft, near the Idaho-Nevada-Oregon border in Mineral Co., Nevada; obtained by E. Ray­ the Owyhee River Valley, Humboldt Co., mond Hall on 8 July 1934. Nevada. Measurements ofholotype. - Total length, Remarks.-Hall and Dale (1939) gave 273; length oftail, 160; length ofhind foot, measurements for three males and one fe­ 41; length ofear, 13; weight, 55.8 g; greatest male. length of skull, 35.1; width across bullae, 23.25; breadth across maxillary arches, 18.3; nasal length, 12.0; interorbital breadth, Dipodomys microps russeolus 11.95. Goldman, 1939 Distribution. -Great Basin Desert in western and southern Nevada, and eastern 1939. Dipodomys microps russeolus Goldman, California in Mono and Inyo counties; south J. Mamm., 20:353, 14 August. from Humboldt Co., Nevada to Death Val­ ley, Inyo Co., California and Las Vegas, Holotype. -Adult male, skin and skull, Clark Co., Nevada; the eastern limits in Ne­ USNM 263895, from Dolphin Island, Great vada are Smiths Creek Valley, Lander Co., Salt Lake, 4,250 ft, Box Elder Co., Utah; Mud Lake, Nye Co., and west ofPanaca in obtained by William H. Marshall on 5 June Desert Valley, Lincoln Co. (Hall and Dale, 1938. 1939). A disjunct population occurs in the Measurements ofholotype. - Total length, vicinity of Joshua Tree National Monu­ 248; length oftail, 136; length ofhind foot, ment in Riverside and San Bernardino 41; greatest length ofskull, 37.5; width across counties, California (Csuti, 1979). bullae, 24.0; length ofnasals, 12.1; breadth Remarks. - Hall (1946) and Grinnell across maxillary arches, 19.2; interorbital (1922) gave means and ranges of measure­ breadth, 11.4; length ofrostrum, 18.5; length ments for samples from Nevada and Cali­ of maxillary toothrow, 5.0; depth of crani­ fornia, respectively. um, 11.0. Distribution. - Limited to Dolphin Is­ land, Great Salt Lake, Box Elder Co., Utah. Dipodomys microps preblei Remarks. -According to Durrant (1952), (Goldman, 1921) russeolus is known only from the type spec­ imen. 1921. Perodipus microps preblei Goldman, J. Mamm., 2:233, 29 November. Holotype. - Adult female, skin and skull Dipodomys microps subtenuis USNM 79340, from Narrows, Malheur Goldman, 1939 Lake, Harney Co., Oregon; obtained on 23 July 1896 by E. A. Preble. 1939. Dipodomys microps subtenuis Goldman, J. Mamm., 20:354, 14 August. Measurements ofholotype. - Total length, 262; length oftail, 154; length ofhind foot, Holotype. - Adult female, skin and skull, 41; greatest length ofskull, 33.5; width across USNM 263917, from Carrington Island, bullae, 22.4; length ofnasals, 11.6; breadth Great Salt Lake, 4,250 ft, Tooele Co., Utah; 68 WILLIAMS ET AL. obtained by William H. Marshall on 30 June a conspicuous black border to the terminal 1938. white tuft of hairs on the tail, and a dark Measurements ofholotype. - Total length, underside to the tail (see account ofD. cal­ 267; length oftail, 157; length ofhind foot, ifornicus, D. deserti, and D. elator for other 40; length ofear (crown), 10; greatest length diagnostic information). of skull, 37.6; width across bullae, 23.7; Distribution. -Open desert communities length of nasals, 12.6; breadth across max­ ofnorth-central Mexico, from southeastern illary arches, 19.7; length ofrostrum, 19.4; Chihuahua and western Coahuila south­ length of maxillary toothrow, 4.7; depth of ward through northeastern Durango and cranium, 11.5. southern Coahuila to northern Zacatecas, Distribution. - Known from Badger, Car­ northern San Luis Potosi, and southern rington, and Stansbury islands, Great Salt Nuevo Leon (Nader, 1978). Lake, Tooele Co., and south on the main­ Remarks. -Controversy about the spe­ land through Rush Valley to Cedar Valley cific status ofD. nelsoni is not satisfactorily west ofUtah Lake, Utah Co., Utah (Durrant resolved. Nader (1978), in reviewing the and Setzer, 1945). distribution, variation, and systematics of Remarks. - Durrant and Setzer (1945) re­ D. deserti, D. spectabilis, and D. nelsoni, marked that no specimens of D. microps found evidence of intergradation between had ever been taken on the eastern or south­ D. s. zygomaticus and D. nelsoni in a narrow ern shore ofGreat Salt Lake, adjacent to the zone ofcontact in south central Chihuahua islands supporting populations of this spe­ and north central Durango. He character­ cies-ofhundreds ofspecimens trapped on ized the intergradation as secondary, the the shore ofthe mainland, all were D. ordii. zone narrow, and the character gradients between taxa as steep. Nader (1978) con­ cluded that nelsoni was best treated as a Dipodomys nelsoni subspecies of D. spectabilis. Anderson (1972), working with less material found no Diagnosis. -A relatively large, four-toed overlap in a combination of characters, kangaroo rat with a white tuft ofhair at the maxillary breadth plotted against total length tip of the tail; length of head and body of of skull, between the two taxa. Hall (1981) adults ranges from about 118 to 134 mm stated that specimens unavailable to An­ and length of hind foot ranges from about derson (1972) were "less easily identified to 46 to 52 mm; white tail tuft relatively small, taxon, by means ofthe graph," but even so, measuring from about 6 to 20 mm in length; concluded that they were not intergrades, face of moderate width, not narrow nor and did not concur with Nader (1978). Mat­ broad; rostrum relatively short andauditory son (1980) interpreted Nader's (1978) anal­ bullae relatively large. yses as providing "no direct evidence ofin­ Comparisons. -Dipodomys nelsoni is tergradation between D. nelsoni and D. most similar in size and appearance to D. spectabilis zygomaticus." We believe, how­ spectabilis, from which it can be distin­ ever, that the indirect evidence is strong. guished by its average smaller size, shorter Matson (1980) analyzed structural relation­ white tail tuft (the tuft of spectabilis is typ­ ships between samples of the spectabilis ically 25 mm or longer), and lesser breadth group from areas near the boundaries ofthe across the maxillary arches (breadth in D. ranges of nelsoni and D. s. cratodon, which nelsoni is less than about 24.5 and in D. ranges to the south ofnelsoni. Discriminant spectabilis is greater than 25 mm in all but function analyses clearly separated refer­ D. s. intermedius). It can be distinguished ence samples of the two taxa and unequiv­ from all other four-toed kangaroo rats by ocally placed "unknowns" from areas be­ either its larger size or darker coloration with tween the ranges of the two taxa with one TAXONOMY 69 or the other ofthe reference groups. He in­ about 88 to 100 mm; length of hind foot terpreted these analyses as demonstrating generally less than 36 mm; skull with rela­ the D. nelsoni and D. spectabilis cratodon tively short and narrow rostrum with rela­ were separate species. Questions that re­ tively straight sides; nasal length averaging main to be resolved concern the relation­ less than 12.3 mm, and rostral breadth av­ ship of cratodon to D. spectabilis (see re­ eraging 3.1 mm or less. marks in the account ofD. spectabilis), and Comparisons. - Dipodomys nitratoides is the significance of the narrow zone of in­ most similar in size and appearance to the tergradation in structural characteristics be­ other small-sized, four-toed species, D. tween D. s. zygomaticus and D. nelsoni. merriami and D. insularis; all other four­ Considering all of the evidence available, toed species are considerably larger and most we believe that it points to specific status have a prominent tuft ofwhite hairs at the for D. nelsoni. Additional samples from tip ofthe tail. See the account ofD. insularis possible zones ofcontact and genic analyses for characters distinguishing that species. D. are needed to help resolve remaining issues, merriami differs from D. nitratoides in gen­ however. erally being larger in size, and in having a longer and wider rostrum (nasal length usu­ ally averages greater than 13.1 in D. mer­ Dipodomys nelsoni Merriam, 1907 riami; Hoffmann, 1975; Lidicker, 1960). Diagnosis. -Dipodomys nitratoides oc­ 1907. Dipodomys nelsoni Merriam, Proc. Bioi. cupies arid, alkaline grassland and desert Soc. Washington, 20:75, 22 July. shrub associations in the San Joaquin Val­ Holotype. -Adult male, skin and skull, ley and contiguous areas to the west, in west­ USNM 79439, from La Ventura, Coahuila, central California. Its range extends from Mexico; obtained on 2 July 1896 by Edward the valley floor in Merced Co., south ofthe W. Nelson and Edward A. Goldman. Merced and San Joaquin rivers to the south­ Measurements ofholotype. - Total length, ern edge of the valley; it also occurs in the 330; length oftail, 204; length ofhind foot, Panoche Valley of eastern San Benito Co., 50; greatest length ofskull, 42.7; width across the Carrizo Plain, San Luis Obispo Co., and bullae, 27.0; length ofnasals, 14.3; breadth the upper Cuyama Valley in San Luis Obis­ across maxillary arches, 22.8; interorbital po and Santa Barbara counties. breadth, 14.1; length ofrostrum, 21.9; length Remarks. -Lidicker (1960) did not in­ of maxillary toothrow, 5.8; depth of crani­ clude taxa assigned to D. nitratoides in his um, 12.5. systematic review ofD. merriami, although Remarks. - Means and ranges of mea­ he acknowledged the two taxa were closely surements for 24 specimens from Chihua­ related and possibly conspecific. Hoffmann hua were given by Anderson (1972). (1975) found that diagnostic characteristics Dalquest (1953) gave cranial measurements for D. nitratoides clearly separated speci­ of 3 specimens, and Nader listed measure­ mens ofthat species from D. merriami, ex­ ments for 11 males and 3 females. cept for a sample from the Cuyama Valley, California, geographically and ecologically contiguous to populations of D. n. brevi­ Dipodomys nitratoides nasus. These specimens exhibited larger size and longer, more robust rostra, character­ Diagnosis. -A small-sized, four-toed istics attributed to D. merriami. Popula­ kangaroo rat with a tail terminating in a tions ofsimilar sized D. merriami, however, small tuft ofbuffy and blackish-brown hairs, such as D. m. parvus, are not geographically and with a face that is not noticeably nar­ adjacent to populations of D. nitratoides. row; length ofhead and body averaging from Measures ofgenetic similarity (Johnson and 70 WILLIAMS ET AL.

Selander, 1971; Patton et aI., 1976), and Dipodomys nitratoides exilis chromosomal data (Hoffmann, 1975; Stock, Merriam, 1894 1974) provide strong evidence that D. ni­ tratoides is related to D. merriami, but is 1894. Dipodomys merriami exilis Merriam, specifically distinct. Proc. BioI. Soc. Washington, 9:113, 21 June. Holotype. - Young adult male, skin and skull, USNM 34843/43823, from Fresno, Dipodomys nitratoides brevinasus San Joaquin Valley, Fresno Co., California; Grinnell, 1920 obtained on 23 September 1891 by Vernon Bailey. 1920. Dipodomys merriami brevinasus Grin­ Measurements ofholotype. - Total length, nell, J. Mamm., 1:179,24 August. 241; length oftail, 143; length ofhind foot, 33; greatest length ofskull, 32.5; width across Holotype. -Adult male, skin and skull, bullae, 20.9; length ofnasals, 10.8; breadth MVZ 28634, from Hayes Station, near across maxillary arches, 17.8; interorbital bench mark 503, 19 mi SW Mendota, Fres­ breadth, 11.9; length ofrostrum, 16.8; length no Co., California; obtained on 30 June 1918 of maxillary toothrow, 3.8; depth of crani­ by Richard Hunt. urn, 10.3. Measurements ofholotype. - Total length, Distribution. -Occupies alkaline grass­ 252; length oftail, 145; length ofhind foot, land and shrub communities on the floor of 36; length of ear (crown), 11; weight, 43.9 the San Joaquin Valley, from approximate­ g; greatest length ofskull, 35.0; width across ly the Merced River, Merced Co., on the bullae, 23.2; length ofnasals, 12.4; breadth north, to Kings River on the south and Fres­ across maxillary arches, 19.5; width ofmax­ no Slough on the west; the area in southern illary arch at middle, 4.7; greatest width of Fresno and northern Kings counties, north rostrum near end, 3.1; width ofrostrum close of historic Tulare Lake, was probably an to base, 5.2. area ofintergradation between typical pop­ Distribution. -Annual grassland and des­ ulations of exilis and nitratoides. ert shrub associations (principally Atriplex Remarks. - Dipodomys nitratoides exilis spp.) along the western margin of the San is the smallest ofthe kangaroo rats. Its cur­ Joaquin Valley, California, from Merced Co. rent range is restricted by irrigated culti­ on the north to the mouth of San Emigdio vation ofits habitat to a few small, isolated Creek, Kern Co., on the south; then ex­ areas ofno more than a few hundred acres. tending eastward in a narrow band above Hoffmann (1975) gave measurements for the valley floor to the east edge ofthe valley samples of exi/is. and northward, east of Bakersfield to ap­ proximately Poso Creek; also occurs in the Panoche Valley in eastern San Benito Co., on the Carrizo Plain, San Luis Obispo Co., Dipodomys nitratoides nitratoides and in the upper Cuyama Valley in San Luis Merriam, 1894 Obispo and Santa Barbara counties. Remarks. -Specimens from iodine bush 1894. Dipodomys merriami nitratoides Merri­ (Allenrolfea) shrublands on the floor of the am, Proc. BioI. Soc. Washington, 9:112, 21 June. San Joaquin Valley, in Merced Co., exhibit characteristics intermediate to brevinasus H olotype. - Adult male, skin and skull, and exilis, but overall, seem closest to the USNM 54674, from Tipton, San Joaquin latter subspecies (unpubI. data). Hoffmann Co., Tulare Co., California; obtained by (1975) provided means and ranges of mea­ Clark P. Streator on 25 June 1893. surements for samples of brevinasus. Measurements ofholotype. - Total length, TAXONOMY 71

246; length oftail, 148; length ofhind foot, other allopatric, five-toed species, except D. 36; greatest length of skull, 34.5; length of ingens, have significantly greater ratios of nasals, 11.8; breadth across maxillary arch­ lengths of tail to head and body; the hind es, 18.6; interorbital breadth, 11.6; length foot of ingens measures 48 mm or more, ofrostrum, 17.6; length ofmaxillary tooth­ whereas the hind foot of ordii measures 43 row, 4.4; depth of cranium, 10.6. mm or less. Distribution. -Occupies the San Joaquin Distribution. - Dipodomys ordii occupies Valley floor, in alkaline shrub and annual an extensive range in the interior grasslands grassland communities, from approximate­ and deserts of western North America; ex­ ly the northern boundaries ofKings County, tending from southeastern Washington east ofthe Kettleman and Lost Hills, south­ southward through the Great Basin region, ward to the edge of the Tehachapi Moun­ and from southwestern Saskatchewan and tains, in Kern County. Limits on the south­ southeastern Alberta southward through the east are approximately Bakersfield, and on western Great Plains region to western Tex­ the southwest, the old shore line of Lake as and Coahuila, southern Texas nearly to Buena Vista. the Gulf coast, and Tamaulipas, Nueva Remarks. - Extant populations are small Leon, and Coahuila; its range also includes and isolated, being scattered in a few sites the intermountain basins and plateaus of throughout the former range ofthe subspe­ Arizona, Colorado, New Mexico, and Utah, cies. Hoffmann (1975) listed measurements and extensive areas ofthe tablelands ofcen­ for samples of nitratoides. tral Mexico; the southernmost portion ofits range is in eastern Zacatecas, Aguascalien­ tes, northeastern Jalisco, San Luis Potosi, Dipodomys ordii Hidalgo, and Guanajuato (Baumgardner and Schmidly, 1981; Hall, 1981; Setzer, 1949). Diagnosis. -A small- to medium-sized Remarks. -Setzer (1949) reviewed the kangaroo rat with five toes on the hind feet species and provided documentation ofdis­ and relatively short tail and ears; the max­ tributions and means and ranges of mea­ illary arch is relatively slender; and the ros­ surements for subspecies. Schmidly and trum is comparatively short. The ratio of Hendricks (1976) and Baumgardner and lengths of tail to head and body normally Schmidly (1981) reported on distribution, averages between about 1.16 and 1.26 and structure, and systematics ofpopulations of the width ofthe skull across the bullae rang­ D. ordii and D. compactus in the south­ es between about 22.3 and 26.3 (Grinnell, eastern portion ofthe range ofD. ordii. Hall 1922; Setzer, 1949). (1981) treated compactus as conspecific with Comparisons. - D. ordii exhibits great D. ordii (see account of compactus). Ken­ variation in size over its extensive geo­ nedy and Schnell (1978) reported on mul­ graphic range, but it is sympatric with few tivariate analyses of geographic variation other five-toed kangaroo rats. D. ordii can and sexual dimorphism of samples of D. be distinguished from D. compactus by its ordii from throughout its geographic range. longer tail and wider skull (see account of the latter species); from D. microps, ordii differs in having a shorter tail and awl-like Dipodomys ordii celeripes lower incisors; from panamintinus, ordiidif­ Durrant and Hall, 1939 fers in being smaller with a relatively shorter 1939. Dipodomys ordii celeripes Durrant and tail and, within their area of sympatry, a Hall, Mammalia, 3:10, March. shorter hind foot (length less than 44 mm; Hall, 1981). All other species sympatric with Holotype. - Adult male, skin and skull, D. ordii have four toes on the hind feet. All UU 1956, from Trout Creek, 4,600 ft, Juab 72 WILLIAMS ET AL.

Co., Utah; obtained by Stephen D. Durrant Dipodomys ordii cinderensis on 5 May 1937. Hardy, 1944 Measurements ofholotype. - Total length, 225; length oftail, 126; length ofhind foot, 1944. Dipodomys ordii cinderensis Hardy, Proc. 41; length ofear, 13; greatest length ofskull, Biol. Soc. Washington, 57:53, 31 October. 36.5; width across bullae, 22.95; breadth Holotype. -Adult male, skin and skull, across maxillary arches, 19.70; nasal length, UU 4611, from immediately N of the 13.30; interorbital breadth, 11.75. northern of two large cinder cones, about Distribution. -Great Basin Desert of 4,000 ft, Diamond Valley, 10 mi N Saint northeastern Nevada, and northwestern George, Washington Co., Utah; obtained by Utah. Ross Hardy on 13 February 1944. Remarks. -Setzer(1949) gave means and Measurements ofholotype. - Total length, ranges ofmeasurements for four males and 232; length oftail, 124; length ofhind foot, two females; Hall (1946) gave measure­ 38; length of ear (notch), 14; condylobasal ments for six males and nine females. length, 29.9; occipitonasal length, 34.4; width across bullae, 23.2; length of nasals, 13.9; breadth across maxillary arches, 19.5; Dipodomys ordii chapmani interorbital breadth, 12.1; length of maxil­ Mearns, 1890 lary toothrow, 5.0; greatest length of bulla as measured parallel with auditory opening, 1890. Dipodomys chapmani Mearns, Bull. Amer. 15.0; ventral symphysis of bulla to ventral Mus. Nat. Hist., 2:291, 21 February. edge of auditory opening, 13.0; length of Syntypes. - Adult male and female, skins diastema, 7.9; length ofleft auditory open­ and skulls, AMNH 2400/10560 and 2398/ ing, 3.7; width of supraoccipital at dorsal 10561, respectively, from Fort Verde, Ya­ crest between bullae, 2.6. vapai Co., Arizona; obtained on 26 January Distribution. -Occupies sandy soils in 1887 and 1 October 1885, respectively, by sagebrush communities in southwestern Edgar A. Mearns. Utah in northern Washington Co. and most Measurements ofholotype.-[2400, 2398] of Iron County, Utah. Total length, 256, 280; length of tail, 137, Remarks. - Hardy (1944) listed measure­ 148; length of hind foot, 36, 38; length of ments for seven males and two females; ear (crown), 11, 11, (notch), 13, 13; greatest Setzer (1949) gave measurements for two length ofskull, 36.7, 37.1; width across bul­ males and two females. lae, 23.2, 22.7; length ofnasals, 12.3, 12.7; breadth across maxillary arches, 19.7, 18.7; interorbital breadth, 12.4, 12.7; length of Dipodomys ordii cineraceus rostrum, 18.9, 19.2; length of maxillary Goldman, 1939 toothrow, 4.6, 4.7. Distribution. - Found in west-central Ar­ 1939. Dipodomys ordii cineraceus Goldman, J. Mamm., 20:352, 14 August. izona from near Kingman and Seligman south to the Bradshaw Mountains and Holotype. -Adult male, skin and skull, southeastward to the Verde Valley (Hoff­ USNM 263890, from Dolphin Island, Great meister, 1986). Salt Lake, 4,250 ft, Box Elder Co., Utah; Remarks. -Setzer (1949) noted that pop­ obtained on 4 June 1938 by William H. ulations of this taxon were relatively iso­ Marshall. lated from those ofother subspecies. Hoff­ Measurements ofholotype. - Total length, meister (1986) gave statistics for 16 males 231; length oftail, 130; length ofhind foot, and 16 females. Setzer (1949) listed means 37; occipitonasallength, 35.7; width across and ranges for six males and four females. bullae, 23.8; length ofnasals, 13.8; breadth TAXONOMY 73 across maxillary arches, 20.1; length of southern boundary of Kaibab Indian Res­ maxillary toothrow, 5.1; width ofnasals (in ervation, Mohave Co., Arizona; obtained front of incisors), 3.5; least width of su­ by Edward A. Goldman on 12 October 1922. praoccipital, 2.9; width of cutting edge of Measurements ofholotype. - Total length, upper incisor, 1.8. 257; length oftail, 150; length ofhind foot, Distribution. - Limited to Dolphin Is­ 41; greatest length ofskull, 38.3; occipitona­ land, Great Salt Lake, Box Elder Co., Utah. sal length, 36.4; width across bullae, 24.9; Remarks. -Goldman (1939) gave mea­ length of nasals, 13.3; breath across max­ surements for three male and one female illary arches, 21.1; interorbital breadth, 11.9; topotypes. length ofrostrum, 19.1; length ofmaxillary toothrow, 4.8; depth ofcranium, 11.6; least width of supraoccipital, 2.5. Dipodomys ordii columbianus Distribution. - Known from an area north (Merriam, 1894) of the Grand Canyon of the Colorado in northwestern Arizona and southwestern 1894. Perodipus ordii columbianus Merriam, Utah. Proc. BioI. Soc. Washington, 9: 115, 21 June. Remarks. - Measurements were given by Holotype. -Adult female, skin and skull, Hoffmeister (1986) for 10 males and 6 fe­ USNM 24181/31594, from Umatilla, at males from Arizona. mouth ofUmatilla River, plains ofColum­ bia, Umatilla Co., Oregon; obtained by Clark P. Streator on 18 October 1890. Dipodomys ordii durranti Measurements ofholotype. - Total length, Setzer, 1952 254; length oftail, 148; length ofhind foot, 40; length of ear (anterior base, dry), 13; 1949. Dipodomys ordii juscus Setzer, Univ. greatest length ofskull, 37.30; width across Kansas PubI. Mus. Nat. Hist., 1:555,27 De­ bullae, 23.30; breadth across maxillary cember. arches, 20.30; nasal length, 13.65; interor­ 1952. Dipodomys ordii durranti Setzer, J. Wash­ bital breadth, 12.60. ington Acad. Sci., 42:391,17 December. Re­ Distribution. - Found on the Columbia naming ofjuscus Setzer, 1949. Plateau and parts of the Great Basin from Holotype. -Adult male, skin and skull, southeastern Washington, eastern Oregon, USNM 93886, from Juamave [Jaumave], and southwestern Idaho southward to Tamaulipas, Mexico; obtained on 3 June northeastern California and parts of north­ 1898 by Edward W. Nelson and Edward A. ern Nevada (Hall, 1981). Goldman. Remarks. - Hall (1946) listed means and Measurements ofholotype. - Total length, ranges of measurements for six males and 241; length oftail, 146; length ofhind foot, six females from Nevada. Setzer (1949) gave 38; greatest length ofskull, 37.5; width across measurements for three male and two fe­ bullae, 23.8; length ofnasals, 12.3; breadth male topotypes. across maxillary arches, 19.6; interorbital breadth, 12.3; length ofrostrum, 18.8; length of maxillary toothrow, 4.3; depth of crani­ Dipodomys ordii cupidineus um, 11.4. Goldman, 1924 Distribution. -Occurson the northern half of the Mesa Central, from southern Coa­ 1924. Dipodomys ordii cupidineus Goldman, J. huila, Nuevo Leon, and adjacent Tamau­ Washington Acad. Sci., 14:372, 19 September. lipas to northeastern Zacatecas and north­ Holotype. -Adult male, skin and skull, ern San Luis Potosi (Baumgardner and USNM 243093, from Kanab Wash, at Schmidly, 1981; Matson and Baker, 1986). 74 WILLIAMS ET AL.

Remarks. - The name fuscus, first given 20.35; nasal length, 14.85; interorbital to this taxon by Setzer (1949), was preoc­ breadth, 12.85. cupied by D. agilis fuscus Boulware, 1943. Distribution. - Known only from the sand Baumgardner and Schmidly (1981) provid­ dunes of Samalayuca in northern Chihua­ ed the most recent review of D. ordii dur­ hua, occupying an area approximately 48 ranti. Measurements were given by Setzer km in diameter (Anderson, 1972). (1949). Remarks. - Anderson (1972) listed means and ranges of measurements for nine spec­ imens. Anderson (1972) could find no sig­ Dipodomys ordii evexus Goldman, 1933 nificant differences, other than paler color, to distinguish extractus from nearby pop­ 1933. Dipodomys ordii evexus Goldman, J. ulations of D. ordii ordii. Washington Acad. Sci., 23:468, 15 October. Holotype. -Adult male, skin and skull, USNM 150990, from Salida, 7,000 ft, Chaf­ Dipodomys ordii fetosus fee County, Colorado; obtained by Merrit Durrant and Hall, 1939 Cary on 10 November 1907. Measurements ofholotype. - Total length, 1939. Dipodomys ordiifetosus Durrant and Hall, Mammalia, 3:14, March. 266; length oftail, 149; length ofhind foot, 42; greatest length ofskull, 38.9; occipitona­ Holotype. - Adult female, skin and skull, sal length, 37.3; width across bullae, 23.9; MVZ 48451, from 2 mi N ofPanaca, 4,800 length of nasals, 13.6; width of maxillary ft, Lincoln Co., Nevada; obtained by Ward arch at middle, 20.9; interorbital breadth, C. Russell on 24 June 1931. 12.7; length ofrostrum, 20.0; length ofmax­ Measurements ofholotype. - Total length, illary toothrow, 5.0; depth ofcranium, 11.7; 225; length oftail, 125; length ofhind foot, width of nasals (in front of incisors), 3.8; 41; length ofear, 13; greatest length ofskull, least width ofsupraoccipital, 3.0. 36.65; width across bullae, 23.85; breadth Distribution. - Valley of the upper Ar­ across maxillary arches, 20.25; nasal length, kansas River of south-central Colorado, 13.3; interorbital breadth, 11.9. above the Royal Gorge. Distribution. - Found in southeastern Remarks. -Setzer (1949) listed measure­ Nevada, in Lincoln Co., and southwestern ments of seven specimens. Utah in western Beaver and Millard coun­ ties. Remarks. -Setzer (1949) gave measure­ Dipodomys ordii extractus ments for four males and four females; Hall Setzer, 1949 (1946) listed measurements for six males and seven females. 1949. Dipodomys ordii extractus Setzer, Univ. Kansas Publ., Mus. Nat. Hist., 1:534,27 De­ cember. Dipodomys ordii fremonti Holotype. - Adult male, skin and skull, Durrant and Setzer, 1945 MVZ 76562, from I mi E Samalayuca, 4,500 ft, Chihuahua, Mexico; obtained on 15 May 1945. Dipodomys ordii fremonti Durrant and 1937 by William B. Richardson. Setzer, Bull. Univ. Utah, 35(26):21, 30 June. Measurements ofholotype. - Total length, Holotype. -Adult female, skin and skull, 244; length oftail, 132; length ofhind foot, CM 15661, from Torrey, 7,000 ft, Wayne 39; length ofear, 13; weight, 49.2 g; greatest Co., Utah; obtained on 19 July 1938 by W. length of skull, 38.6; width across bullae, F. and F. H. Wood. 24.35; breadth across maxillary arches, Measurements ofholotype. - Total length, TAXONOMY 75

246; length oftail, 132; length ofhind foot, 42; length of ear (crown, dry), 8; greatest 38; condylobasallength, 23.1; occipitonasal length of skull, 39.8; width across bullae, length, 30.3; length of nasals, 13.4; inter­ 25.5; length of nasals, 14.0; breadth across orbital breadth, 12.1; length ofbulla, 14.7; maxillary arches, 19.8; interorbital breadth, width of bulla, 9.6; depth of bulla, 12.1. 12.9; length ofrostrum, 19.8; length ofmax­ Distribution. - Known only from the vi­ illary toothrow, 4.4. cinity ofthe type locality in the upper reach­ Distribution. - Ranges through the Paint­ es ofthe Fremont River watershed in west­ ed Desert region of southeastern Utah, ex­ central Wayne Co., Utah (Setzer, 1949). treme southwestern Colorado, northwest­ Remarks. - Setzer (1949) listed measure­ ern New Mexico, and northeastern Arizona. ments for two males and two females. Remarks. -Setzer (1949) placed cleo­ mophila as a synonym of longipes. Hoff­ meister (1986) gave measurements for eight Dipodomys ordii inaquosus Hall, 1941 males and nine females from Arizona; Dur­ rant (1952) gave measurements for two 1941. Dipodomys ordii inaquosus Hall, Proc. males and five females from Utah. Bioi. Soc. Washington, 54:58, 20 May. Holotype. - Adult male, skin and skull, MVZ 73580, from 11 mi E, 1 mi N Jungo, Dipodomys ordii luteolus 4,200 ft, Humboldt Co., Nevada; obtained (Goldman, 1917) by Ward C. Russell on 26 July 1936. Measurements ofholotype. - Total length, 1917. Perodipus ordii luteolus Goldman, Proc. Bioi. Soc. Washington, 30:112, 23 May. 261; length oftail, 142; length ofhind foot, 41; length ofear, 12; weight, 49 g; greatest Holotype. -Adult male, skin and skull, length of skull, 37.55; width across bullae, USNM 160408, from Casper, Natrona Co., 24.15; breadth across maxillary arches, 20.4; Wyoming; obtained by Merritt Cary on 2 nasal length, 14.3; interorbital breadth, September 1903. 11.45. Measurements ofholotype. - Total length, Distribution. -Found in north-central 272; length oftail, 154; length ofhind foot, Nevada in southeastern Humboldt and 42; occipitonasallength, 38.1; width across northern Lander counties (Hall, 1946). bullae, 24.3; breadth across maxillary arch­ Remarks. - Hall (1946) gave measure­ es, 21.2; length of maxillary toothrow, 4.8; ments for six males and two females. least width of supraoccipital, 2.8. Distribution. - Occupies the northern Great Plains from western South Dakota to Dipodomys ordii longipes southern Nebraska, northeastern Colorado, (Merriam, 1890) and southeastern Wyoming (Hall, 1981; Setzer, 1949). 1890. Dipodops longipes Merriam, 1890, N. Remarks. -Armstrong (1972), Jones Amer. Fauna, 3:72, 11 September. (1964), Long (1965), and Setzer (1949) list­ 1933. Dipodomys ordii cleomophila Goldman, ed measurements. 1. Washington Acad. Sci., 23:469, 15 October. Holotype. - Young adult male, skin and skull, USNM 17703124639, from foot of Dipodomys ordii marsha/li Echo Cliffs, Painted Desert, Coconino Co., Goldman, 1937 Arizona; obtained by C. Hart Merriam on 22 September 1889. 1937. Dipodomys ordii marshalli Goldman, Measurements ofholotype. - Total length, Proc. Bioi. Soc. Washington, 50:223, 28 De­ 275; length oftail, 165; length ofhind foot, cember. 76 WILLIAMS ET AL.

Holotype. -Adult female, skin and skull, of southeastern and north-central New USNM 262655, from Bird Island, about Mexico and western Texas (Hall, 1981). 4,300 ft, Great Salt Lake, Tooele Co., Utah; Remarks. - Setzer (1949) listed measure­ obtained by William H. Marshall on 22 June ments for six male and two female topo­ 1937. types. Measurements ofholotype. - Total length, 273; length oftail, 123; length ofhind foot, 36; length ofear, 14; greatest length ofskull, Dipodomys ordii monoensis 35.5; width across bullae, 22.8; length of (Grinnell, 1919) nasals, 13.6; breadth across maxillary arch­ es, 20.0; interorbital breadth, 11.0; length 1919. Perodipus monoensisGrinnell, Univ. Cal­ of maxillary toothrow, 14.3; depth of cra­ ifornia Publ. Zool., 21 :46, 29 March. nium, 10.6; occipitonasallength, 35.0; width ofnasals (in front ofincisors), 3.4; least width Holotype. - Adult female, skin and skull, of supraoccipital, 2.6. MVZ 27002, from Pellisier Ranch, 5,600 ft, Distribution. - Found on Badger, Bird, 5 mi N Benton Station, Mono Co., Califor­ Carrington, and Stansbury islands, Great nia; obtained by Joseph Dixon on 21 Sep­ Salt Lake, and along the western edge of tember 1917. Great Salt Lake, north to Kelton, Box Elder Measurements ofholotype. - Total length, Co., Utah; and around the southern and 245; length oftail, 124; length ofhind foot, southeastern shores ofthe lake to the mouth 38; length ofear, 12; weight, 48.1 g; greatest of the Jordan River, Davis and Salt Lake length of skull, 37.15; width across bullae, counties, Utah. 23.40; breadth across maxillary arches, Remarks. - The skull ofthe holotype was 19.90; nasal length, 13.55; interorbital damaged sometime after Goldman's de­ breadth, 11.50. scription and after Poole and Schantz (1942) Distribution. - Found in the Mono Basin remarked on the condition ofthe skull. Set­ of Mono, Co., California and a wider area zer (1949) and Durrant (1952) gave mea­ ofwest-central Nevada, from Pyramid Lake surements for specimens of marshalli. and the Humboldt Sink on the north, south­ ward to Beatty, Nye Co., and, with the ex­ ception ofthe Reese River Valley, eastward Dipodomys ordii medius Setzer, 1949 over Nye Co. to the Quinn Canyon Moun­ tains (Hall, 1946, 1981). 1949. Dipodomys ordii medius Setzer, 1949, Remarks. - Hall (1946) gave measure­ Univ. Kansas Publ., Mus. Nat. Rist., 1:519, ments for 10 males and 3 females from Ne­ 27 December. vada, and Grinnell (1922) gave measure­ ments for 5 males and 5 females from Holotype. -Adult male, skin and skull, California. USNM 118526, from Santa Rosa, Guada­ lupe Co., New Mexico; obtained by Jason H. Grant on 5 October 1902. Measurements ofholotype. - Total length, Dipodomys ordii montanus Baird, 1855 256; length oftail, 146; length ofhind foot, 38; greatestlength ofskull, 38.7; width across 1855. Dipodomys montanus Baird, Proc. Acad. Nat. Sci. Philadelphia, 7:334, April. bullae, 24.9; length ofnasals, 13.7; breadth across maxillary arches, 21.0; length of Holotype. -Adult [sex unknown], skin maxillary toothrow, 4.8; depth ofcranium, and skull, ANSP 490/1631, from Fort Mas­ 12.0. sachusetts [now Fort Garland], Costilla Co., Distribution. - Found on the High Plains Colorado; obtained by F. Kreuzfeldt in 1853. TAXONOMY 77

Measurements ofholotype. - Total length, from Colorado; Goldman (1933) listed 247.7; length of tail, 133.4; length of hind measurements for 3 adult topotypes. foot, 38.1; greatest length of skull, 36.8; length of nasals, 12.8; breadth across max­ illary arches, 19.5; interorbital breadth, 12.5; Dipodomys ordii obscurus length ofrostrum, 19.5; length ofmaxillary (J. A. Allen, 1903) toothrow, 4.7. Distribution. - Found in the San Luis 1903. Perodipus obscurus J. A. Allen, Bull. Amer. Valley of south-central Colorado and ad­ Mus. Nat. Rist., 19:603, 12 November. jacent north-central New Mexico. 1939. Dipodomys ordii attenuatus Bryant, Oc­ Remarks. - External measurements and cas. Papers Mus. Zoo!., Louisiana State Univ., length of skull were converted from inches 5:65, 10 November. 1949. Dipodomys ordii idoneus Setzer, Univ. to mm from values published by Baird Kansas Pub!., Mus. Nat. Rist., 1:546,27 De­ (1858); measurements were not recorded cember. with the holotype and the skull is damaged such that a measure ofgreatest length could Holotype. - Adult male, skin and skull, not be taken by us. Armstrong (1972) pro­ AMNH 20957, from Rio Sestin, north­ vided statistics for 10 males and 6 females western Durango, Mexico; obtained by J. and Setzer (1949) published means and H. Batty on 13 April 1903. ranges of measurements for 11 males and Measurements ofholotype. - Total length, 11 females. 232; length oftail, 130; length ofhind foot, 35; length ofear (notch), 12.7; greatest length of skull, 35.6; width across bullae, 23.0; length of nasals, 12.8; breadth across max­ Dipodomys ordii nexilis Goldman, 1933 illary arches, 19.7; interorbital breadth, 12.4; length ofrostrum, 19.0; length ofmaxillary 1933. Dipodomys ordii nexilis Goldman, J. toothrow, 4.4. Washington Acad. Sci., 23:470, 15 October. Distribution. -Occurs on the northern Holotype. - Adult male, skin and skull, portion of the Mexican Plateau north of USNM 149938, from 5 mi W Naturita, northern Zacatecas and southern Coahuila Montrose Co., Colorado; obtained by Mer­ (Mesa del Norte), and the adjoining regions ritt Cary on 20 July 1907. ofthe Big Bend Basin and Rio Grande Plain Measurements ofholotype. - Total length, ofTexas (Baumgardner and Schmidly, 1981; 268; length oftail, 147; length ofhind foot, Bryant, 1939; Matson and Baker, 1986). 45; greatest length ofskull, 40.9; width across Remarks. - Baumgardner and Schmidly bullae, 25.9; length ofnasals, 14.0; breadth (1981) synonymized attenuatus and idoneus across maxillary arches, 21.2; interorbital with obscurus, to which they also referred breadth, 13.1; length ofrostrum, 19.6; length the northern populations previously as­ of maxillary toothrow, 5.0; depth of crani­ signed to durranti. Baumgardner and um, 12.1; occipitonasallength, 39.0; width Schmidly (1981) gave means and ranges of ofnasals (in front ofincisors), 4.3; least width measurements for samples of obscurus. of supraoccipital, 2.3. Distribution. - Found in southeastern Utah, between the Colorado and San Juan Dipodomys ordii oklahomae rivers, and in adjacent southwestern Colo­ Trowbridge and Whitaker, 1940 rado (Hall, 1981). Remarks. -Armstrong (1972) listed 1940. Dipodomys oklahomae Trowbridge and measurements of 10 males and 4 females Whitaker, J. Mamm., 21:343, 13 August. 78 WILLIAMS ET AL.

Holotype. - Young adult female, skin and commumtIes of southeastern Arizona, skull, USNM 265454, from north bank of southern New Mexico, Trans-Pecos Texas, South Canadian River, 2.25 mi S Norman, northern Chihuahua, and northeastern So­ Cleveland Co., Oklahoma; obtained by H. nora (Hall, 1981). L. Whitaker on 16 March 1934. Remarks. -Although Merriam (1890) Measurements ojholotype. - Total length, used the term duplicate type rather than 243; length oftail, 128; length ofhind foot, neotype, we believe his designation met the 41; greatest length ofskull, 38.4; occipitona­ essential criteria of the International Code sal length, 36.5; width across bullae, 23.8; ofZoological Nomenclature for designating length of nasals, 13.6; breadth across max­ a neotype, and so regard his duplicate type. illary arches, 21.6; interorbital breadth, 13.1; Anderson (1972) provided measurements length ofrostrum, 19.6; length ofmaxillary for 14 specimens from Chihuahua, Hoff­ toothrow, 4.8; depth ofcranium, 12.1; width meister (1986) gave statistics for measure­ ofnasals (in front ofincisors), 3.9; least width ments of 16 males and 10 females from of supraoccipital, 3.3; width of palate be­ Arizona, and Setzer (1949) listed measure­ tween first molars, 3.4; length of interpari­ ments for 5 male and 6 female topotypes. etal, 4.0; width of interparietals, 3.1. Distribution. -Known only from the South Canadian River Valley of central Dipodomys ordii paUidus Oklahoma from Canadian Co. eastward into Durrant and Setzer, 1945 Cleveland Co. Remarks. - Kennedy et al. (1980) ana­ 1945. Dipodomys ordii pallidus Durrant and lyzed intraspecific variation in anatomical Setzer, Bull. Univ. Utah, 35(26):24, 30 June. structure of oklahomae and other popula­ Holotype. -Adult male, skin and skull, tions of D. ordii from Oklahoma, and pro­ UU 3526, from Old Lincoln Highway, 18 vided lists of measurements. mi SW Orr's Ranch, Skull valley, 4,400 ft, Tooele Co., Utah; obtained by Stephen D. Durrant on 6 June 1940. Dipodomys ordii ordii Woodhouse, 1853 Measurements ojholotype. - Total length, 251; length oftail, 141; length ofhind foot, 1853. D[ipodomys). ordii Woodhouse, Proc. Acad. Nat. Sci. Philadelphia, 6:224. 45; occipitonasallength, 30.7; condylobasal length, 23.5; length of nasals, 13.0; length Holotype. -None designated, species of bulla, 15.1; width of bulla, 10.0; depth characterized from specimens obtained by of bulla, 12.2. Dr. Woodhouse at El Paso, El Paso Co., Distribution. -Occupies the lower-lying Texas. Neotype-Adult female, skin and valleys of west-central Utah in Juab, Too­ skull, USNM 18135125033, from El Paso, ele, and Millard counties (Durrant, 1952). El Paso Co., Texas; obtained on 11 Decem­ Remarks. -Durrant listed means and ber 1889 by Vernon Bailey (designated by ranges ofmeasurements for seven males and Merriam, 1890:45, as the "Duplicate type"; two females. see remarks below). Measurements ojneotype. - Total length, 240; length oftail, 134; length ofhind foot, Dipodomys ordii palmeri ~8; length ofear (crown), 7, (anterior base, (J. A. Allen, 1891) dry) 12; greatest length ofskull, 38.60; width 1891. Dipodops ordii palmeri J. A. Allen, Bull. across bullae, -; breadth across maxillary Amer. Mus. Nat. Hist., 3:276, 30 June. arches, 20.70; nasal length, 14.00; interor­ bital breadth, 12.40. Syntypes. - Two adult males, skins and Distribution. - Ranges through desert skulls, MCZ 5886,5887, from San Luis Po- TAXONOMY 79 tosi, San Luis Potosi, Mexico on 1 May and Distribution. - Known only from the type 1 September 1878 by Edward Palmer. locality in south-central Utah. Measurements ofsyntypes. - [5886, 5887; Remarks. - Hardy (1940) listed measure­ external measurements approximated by ments for two male and two female holo­ Allen (1891) from skins and reported as sin­ types. gle values] Tota11ength, 249; length oftail, 141; length of hind foot, 35; length of ear (notch, dry), 11.4; greatest length of skull, Dipodomys ordii priscus 36.0, 36.3; width across bullae, -, 23.0; Hoffmeister, 1942 length of nasals, 12.5, 13.3; breadth across maxillary arches, 19.1, 19.6; interorbital 1942. Dipodomys ordii priscus Hoffmeister, Proc. BioI. Soc. Washington, 55:167, 31 December. breadth, 12.8, 12.9; length ofrostrum, 17.5, 18.0; length ofmaxillary toothrow, 4.2, 4.3. Holotype. - Adult female, skin and skull, Distribution. - Occupies the southern MVZ 89119, from Kinney Ranch, 21 mi S portion of the Central Plateau of Mexico, Bittercreek, 7,100 ft, Sweetwater Co., Wy­ from northern San Luis Potosi and central oming; obtained by Donald T. Tappe on 16 Zacatecas, south to Hidalgo (Hall, 1981; September 1939. Matson and Baker, 1986). Measurements ofholotype. - Total length, Remarks. - Baumgardner and Schmidly 267; length oftail, 145; length ofhind foot, (1981) questioned the subspecific distinct­ 41; length ofear, 13; weight, 64.4 g; greatest ness ofpalmeri and durranti, based on mul­ length of skull, 39.5; width across bullae, tivariate analysis of anatomical structure. 24.45; breadth across maxillary arches, 20.7; Previous authors have listed the paper by nasal length, 14.3; interorbital breadth, 13.1. Allen (1881), as the first description of D. Distribution. -Located in the arid inter­ o. palmeri; Allen (1891) first named and mountain basins of southwestern Wyo­ described palmeri, noting that he had earlier ming, northwestern Colorado, and extreme (Allen, 1881) referred specimens ofthis sub­ northeastern Utah. species to Dipodomys phillipsii. Dalquest Remarks. - Armstrong (1972) gave mea­ (1953) gave means for measurements offive surements for one male and four females. males and nine females from San Luis Po­ Setzer (1949) provided means and ranges of tosi. measurements of 11 specimens.

Dipodomys ordii panguitchensis Dipodomys ordii pullus Anderson, 1972 Hardy, 1942 1972. Dipodomys ordii pullus Anderson, Bull. 1942. Dipodomys ordii panguitchensis Hardy, Amer. Mus. Nat. Hist., 148:317, 8 September. Proc. BioI. Soc. Washington, 55:90, 25 June. Holotype. - Adult female, skin and skull, Holotype. - Adult male, skin and skull, KU 73733, from El Rosario, 6,700 ft, Chi­ UU 4375, from 1 mi S Panguitch, 6,666 ft, huahua, Mexico; obtained on 28 July 1957 Garfield Co., Utah; obtained on 31 August by Sydney Anderson. 1940 by Ross Hardy. Measurements ofholotype. - Total length, Measurements ofholotype. - Total length, 254; length oftail, 147; length ofhind foot, 257; length oftail, 145; length ofhind foot, 40.5; length ofear, 15; weight, 57.3 g; great­ 41; length ofear (notch), 14; basal length of est length of skull, 36.95; width across bul­ skull, 25.2; width across bullae, 23.0; length lae, 24.10; breadth across maxillary arches, of nasals, 13.4; breadth across maxillary 21.70; nasal length, 13.20; interorbital arches, 20.0; interorbital breadth, 12.3; breadth, 13.40. length of maxillary toothrow, 4.7. Distribution. - Found in west-central 80 WILLIAMS ET AL.

Chihuahua, including the high valleys ofthe Measurements ofholotype. - Total length, Rio Papigochic, Laguna de los Mexicanos, 249; length oftail, 138; length ofhind foot, and Laguna de Bustillos (Anderson, 1972). 42; length ofear, 16; greatest length ofskull, Remarks. - Anderson (1972) listed means 37.80; width across bullae, 24.80; breadth and ranges of measurements for 51 speci­ across maxillary arches, 20.20; nasal length, mens. 13.10; interorbital breadth, 12.75. Distribution. -Occurs in the high desert ofeast-central Utah eastward into Colorado Dipodomys ordii richardsoni along the Colorado River Valley to Grand (J. A. Allen, 1891) Junction, Mesa Co. (Hall, 1981). Remarks. - Measurements were given by 1891. Dipodops richardsoni J. A. Allen, Bull. Armstrong (1972), Durrant and Setzer Amer. Mus. Nat. Hist., 3:277, 30 June. (1945), and Setzer (1949). Holotype. - Adult male, skin and skull, AMNH 302512345, from Beaver River [confluence of Cienquilla and Currumpaw Dipodomys ordii terrosus creeks, Sec. 32, T2N, R2E], Cimarron Co., Hoffmeister, 1942 Oklahoma; obtained by Jenness Richard­ son and John Rowley, Jr., on 26 October 1942. Dipodomys ordii terrosus Hoffmeister, Proc. Biol. Soc. Washington, 55:165, 31 De­ 1889. cember. Measurements of holotype. -Greatest length of skull, 40.4; width across bullae, Holotype. -Adult male, skin and skull, 25.0; length of nasals, 14.2; breadth across MVZ 93477, from Yellowstone River, 5 mi maxillary arches, 21.6; interorbital breadth, W Forsyth, 2,750 ft, Rosebud Co., Mon­ 11.9; length ofrostrum, 19.2; length ofmax­ tana; obtained on 2 June 1940 by J. R. Al­ illary toothrow, 5.2. corn. Distribution. -Occupies the west-central Measurements ofholotype. - Total length, Great Plains, from southwestern Nebraska, 266; length oftail, 143; length ofhind foot, to western Oklahoma and the Panhandle 43; weight, 71.8 g; occipitobasallength, 28.4; region of west Texas, and westward in width across bullae, 25.8; length of nasals, southeastern Colorado and northeastern 15.5; breadth across maxillary arches, 22.8; New Mexico (Hall, 1981). interorbital breadth, 13.2. Remarks. -Armstrong (1972), Jones Distribution. -Occupies the northwest­ (1964), Kennedy et al. (1980), and Setzer ern Great Plains, from southern Alberta and (1949) gave measurements for samples of Saskatchewan, southward through eastern richardsoni. Glass (1971) provided infor­ Montana, extreme southwestern North Da­ mation to amend the type locality and the kota, and northwestern South Dakota to year that the holotype was collected. northern Wyoming (Hall, 1981). Remarks. - Long (1965) provided mea­ surements for 3 males and 10 females; Set­ Dipodomys ordii sanrafaeli zer (1949) gave measurements of 5 adults. Durrant and Setzer, 1945

1945. Dipodomys ordii sanrafaeli Durrant and Dipodomys ordii uintensis Setzer, Bull. Univ. Utah, 35(26):26, 30 June. Durrant and Setzer, 1945 Holotype. - Adult female, skin and skull, UU 4612, from 1.5 mi N Price, 5,567 ft, 1945. Dipodomys ordii uintensis Durrant and Setzer, Bull. Univ. Utah, 35(26):27, 14 July. Carbon Co., Utah; obtained on 5 June 1940 by Ross Hardy and H. Higgins. Holotype. - Adult male, skin and skull, TAXONOMY 81

CM 11634, from Red Creek, 6,700 ft, 2 mi width across the maxillary arches averages N Fruitland, Duchesne Co., Utah; obtained from about 23.2 to 23.6 or greater; width by J. K. and M. T. Doutt on 15 August 1936. ofmaxillary arch, at middle, averages from Measurements ofholotype. - Total length, about 5.4 to 5.7 (Grinnell, 1922). 253; length oftail, 140; length ofhind foot, Comparisons. -Dipodomys panaminti­ 40; greatest length ofskull, 37.9; width across nus is most similar in size and proportions bullae, 23.3; length ofnasals, 13.5; breadth to some populations ofD. heermanni, which across maxillary arches, 20.3; interorbital occupies an allopatric range in central Cal­ breadth, 12.5; width ofrostrum, 3.9; basilar ifornia; panamintinus differs in having much length, 23.9. less black in the markings on the face, short­ Distribution. -Occupies sandy soils ofthe er ears, and having auditory bullae that are Uintah Basin in northeastern Utah. about half the volume of those of D. heer­ Remarks.-Durrant (1952) gave mea­ manni. From the sympatric five-toed spe­ surements for three adult males. cies, D. panamintinus can be distinguished by its considerably larger size, and propor­ tionately longer tail than D. ordii, and its Dipodomys ordii utahensis awl-shaped lower incisors in comparison to (Merriam, 1904) the chisel-shaped incisors of D. microps. Overall, D. panamintinus is nearly indistin­ 1904. Perodipus montanus utahensis Merriam, guishable externally from the allopatric spe­ Proc. Bioi. Soc. Washington, 17:143,30 June. cies, D. stephensi. It differs from stephensi in having smaller auditory bullae, longer na­ Holotype. -Adult male, skin and skull, sals, wider maxillary arches, and in other USNM 55115, from Ogden, Weber Co., proportions of the skull (Grinnell, 1922). Utah; obtained on 15 July 1893 by Vernon Distribution. -Occursin desert shrub and Bailey. arid woodland associations along the Measurements ofholotype. - Total length, boundary between California and Nevada, 260; length oftail, 150; length ofhind foot, from the Virginia Mountains, Washoe Co., 41; greatest length of skull, 37.50; width Nevada and the Honey Lake Valley, Lassen across bullae, 23.40; breadth across maxil­ Co., California, southward through the Mo­ lary arches, 20.15; nasal length, 13.75; in­ jave Desert to northeastern Los Angeles Co. terorbital breadth, 11.75. and Hesperia, San Bernardino Co., Califor­ Distribution. - From near the Idaho bor­ nia; it also occupies the slopes ofthe desert der southward in Utah along the eastern mountain ranges in this region. A disjunct margin of the Pleistocene Lake Bonneville population is found in extreme southern basin to northern Sevier Co.; ranges to the Nevada and adjacent California in Yucca west side ofthe Onaqui Mountains, Tooele associations of the Providence Mountain Co., on the west (Durrant, 1952). region (Hall, 1981). Remarks.-Durrant (1952) gave average Remarks. -Grinnell (1922) and Hall measurements of five specimens. (1946) gave measurements for samples of D. panamintinus from California and Ne­ vada, respectively. Populations of D. pan­ Dipodomys panamintinus amintinus and D. heermanni are similar in chromosome structure (Stock, 1974), and Diagnosis. - A relatively large, five-toed exhibit a high degree of genic similarity kangaroo rat with short ear pinnae, a broad (Patton et al., 1976), although Schnell et al. face, stout maxillary arch, and relatively (1978) determined that D. panamintinus was small auditory bullae; length of head and phenetically an outlier in terms ofany well­ body averages from about 123 to 128 mm; defined species group of Dipodomys. 82 WILLIAMS ET AL.

Dipodomys panamintinus argusensis vada and 10 adults of each sex from Cali­ Huey, 1945 fornia.

1945. Dipodomys mohavensis argusensis Huey, Trans. San Diego Soc. Nat. Hist., 10:131, 9 Dipodomys panamintinus leucogenys March. (Grinnell, 1919) Holotype. -Adult male, skin and skull, SDSNH 9552, from Junction Ranch, 5,725 1919. Perodipus leucogenysGrinnell, Univ. Cal­ ifornia Publ. Zool., 21:46, 29 March. ft, Argus Mountains, Inyo Co., California; obtained by Samuel G. Harter on 13 August Holotype. - Adult male, skin and skull, 1931. MVZ 26933, from Pellisier Ranch, 5,600 ft, Measurements ofholotype. - Total length, 5 mi N Benton Station, Mono Co., Califor­ 297; length oftail, 172; length ofhind foot, nia; obtained by Joseph Dixon on 20 Sep­ 44; length ofear, 13; greatest length ofskull, tember 1917. 40.2; width across bullae, 24.5; length of Measurements ofholotype. - Total length, nasals, 15.6; breadth across maxillary arch­ 310; length oftail, 176; length ofhind foot, es, 24.0; width ofmaxillary arch at middle, 48; length of ear (crown), 13; weight, 85.5 5.5; greatest width ofrostrum near end, 4.5. g; greatest length ofskull, 41.1; width across Distribution. -Known only from the vi­ bullae, 24.3; breadth across maxillary arch­ cinity of the type locality in the Argus es, 23.85; nasal length, 16.0; interorbital Mountains, Inyo Co., California. breadth, 13.05. Remarks. - Miller and Kellogg (1955) first Distribution. -Northeastern California treated argusensis as a subspecies ofD. pan­ and western Nevada from Lassen Co., Cal­ amintinus. ifornia and Washoe Co., Nevada, south­ ward along the state boundary to the Ex­ celsior Mountains, Nevada, then southward Dipodomys panamintinus caudatus in California through the upper Owens Val­ Hall, 1946 ley to the vicinity ofIndependence, Inyo Co. Remarks.-Grinnell (1922) gave average 1946. Dipodomys panamintinus caudatus Hall, measurements for 5 adults ofeach sex from Mammals of Nevada, p. 409, 1 July. California and Hall (1946) listed means and Holotype. - Adult female, skin and skull, ranges for 9 males and 10 females from Ne­ MVZ 80028, from 6 mi S Granite Well, vada. 3,800 ft, Providence Mountains, San Ber­ nardino Co., California; obtained on 18 De­ cember 1937 by F. Wallace Taber. Dipodomys panamintinus mohavensis Measurements ofholotype. - Total length, (Grinnell, 1918) 299; length oftail, 180; length ofhind foot, 43; length ofear, 16; weight, 68.8 g; greatest 1918. Perodipus mohavensis Grinnell, Univ. California Publ. Zool., 17:428, 25 April. length of skull, 40.2; width across bullae, 24.95; breadth across maxillary arches, Holotype.-Adult male, skin and skull, 22.55; nasal length, 15.0; interorbital MVZ 26835, from 0.5 mi E railway station breadth, 13.05. of Warren (about 5 mi N Mohave [Moja­ Distribution. -Chiefly found in Yucca as­ ve]), 3,275 ft, Kern Co., California; ob­ sociations ofthe Providence Mountains re­ tained on 27 March 1917 by Joseph Grin­ gion ofSan Bernardino Co., California and nell. adjacent Clark Co., Nevada. Measurements ofholotype. - Total length, Remarks. -Hall (1946) listed means and 305; length oftail, 178; length ofhind foot, ranges for 3 males and 3 females from Ne- 44, length ofear (crown), 12; weight, 88 g; TAXONOMY 83 greatest length of skull, 40.7; width across tive1y long tail with the dark dorsal and ven­ bullae, 25.0; length ofnasals, 15.8; width of tral stripes uniting in the distal third, and maxillary arch at middle, 5.2. usually a white, terminal tuft ofhairs on the Distribution. - Mojave Desert from near tail; auditory bullae relatively small, cra­ Lone Pine in the Owens Valley, southward nium flattened, maxillary region broad, and into the Antelope Valley in extreme north­ postrostra1 region nearly quadrate; maxil­ eastern Los Angeles Co., and to near Hes­ lary plate projecting posteriorly to level of peria, San Bernardino Co., California. second or third molar; rostrum relatively Remarks. -Grinnell (1922) listed means narrow and straight-sided; incisors slender and ranges of measurements for five adults in comparison to other species of Dipodo­ of each sex. mys (Jones and Genoways, 1975). Comparisons. - Dipodomys phillipsii can be distinguished from the other small- to medium-sized, four-toed kangaroo rats by Dipodomys panamintinus panamintinus the combination of a tail that has an all (Merriam, 1894) blackish distal third, with or without a white 1894. Perodipus panamintinus Merriam, Proc. terminal tuft; a hind foot less than 42 mm BioI. Soc. Washington, 9:114, 21 June. in length; and a relatively narrow skull, measuring less than 25.0 mm across the bul­ Holotype. - Adult male, skin and skull, lae. Itis most similar in size and appearance USNM 28566/40670, from head ofWillow to D. merriami, D. californicus and D. ela­ Creek [about 6,200 ft, about 3 mi NE Jack­ ass Spring], Panamint Mountains, Inyo Co., tor; both ofthe latter are larger, have broad­ California; obtained on 12 May 1891 by er rostra and heavier incisors, and are wide­ Edward W. Nelson. ly separated geographically. Other four-toed Measurements ofholotype. - Tota11ength, kangaroo rats with white tail tufts are D. 293; length oftail, 179; length ofhind foot, deserti, D. nelsoni, and D. spectabilis, all of 43; length of ear from anterior base (dry which are considerably larger than D. phil­ skin), 15; greatest length of skull, 39.75; lipsii, with larger hind feet and broader width across bullae, 24.50; breadth across skulls. From D. phillipsii, D. merriami dif­ maxillary arches, 22.80; nasallength, 15.55; fers most notably in coloration of the tail, interorbital breadth, 13.00. lacking an all-blackish distal one third, and Distribution. -Occupies a known area in lacking a prominent, all white tip. the Panamint Mountains measuring about Distribution. - Primarily occupies sandy 10 by 13 km in size, in the Upper Sonoran soils on the Mexican Plateau and adjacent and arid Transition life zones, in the vicin­ areas, from central Durango to northern ity of Jackass Spring, Inyo Co., California Oaxaca (Jones and Genoways, 1975). (Grinnell, 1922). Remarks. -Macrocolus halticus Wagner, Remarks. - Measurements for five adults 1845, Abhandl. K. Baier. Akad., 22:319, is ohach sex were listed by Grinnell (1922); a synonym of Dipodomys phillipsii, but its Merriam (1894a) listed average external type locality, other than Mexico, is un­ measurements for 16 specimens from the known; thus it is not listed in the synonymy type locality. of any of the subspecies of phillipsii. Gen­ oways and Jones (1971) provided a system­ atic review of D. phillipsii and gave means Dipodomys phillipsii and ranges of measurements for samples from throughout the geographic range ofthe Diagnosis. - A medium-sized kangaroo species. Jones and Genoways (1975) pro­ rat with four toes on the hind feet, a re1a- vided a review ofthe biology ofthe species. 84 WILLIAMS ET AL.

Dipodomys phillipsii oaxacae gave means and ranges ofmeasurements for Hooper, 1947 nine samples of ornatus.

1947. Dipodomys phillipsii oaxacae Hooper, J. Mamm., 28:48, 17 February. Dipodomys phillipsii perotensis Holotype. - Adult male, skin and skull, Merriam, 1894 UMMZ 88652, from Teotitl{m, 950 m, Oa­ xaca, Mexico; obtained on 15 September 1894. Dipodomys perotensis Merriam, Proc. BioI. Soc. Washington, 9:111, 21 June. 1944 by Helmuth O. Wagner. Measurements ofholotype. - Tota11ength, Holotype. - Adult female, skin and skull, 255; length oftail, 160; length ofhind foot, USNM 54285, from Perote, Vera Cruz, 37; length ofear (notch, dry), 10.5; greatest Mexico; obtained by Edward W. Nelson on length ofskull, 34.5; length ofnasals, 12.9; 21 May 1893. breadth across maxillary arches, 19.6; width Measurements ofholotype. - Total length, ofmaxillary arch at middle, 4.8; interorbital 265; length oftail, 162; length ofhind foot, breadth, 12.5; width ofnasals near end, 3.2; 40; length of ear (anterior base, dry), 14; greatest length ofinterparietal, 3.2; greatest greatest length of skull, 36.3; width across breadth of interparieta1s, 2.2. bullae, 22.8; length of nasals, 12.9; length Distribution. -Known from the type lo­ ofrostrum, 19.9; length ofmaxillary tooth­ cality and a single locality in southern Pueb­ row, 4.9; depth ofcranium, 11.2. 1a (Genoways and Jones, 1971). Distribution. - Known only from around Remarks. - This subspecies is distin­ Tlaxcala, west-central Veracruz, and from guished by small size and pale coloration. eastern Puebla (Genoways and Jones, 1971). Genoways and Jones (1971) listed mea­ Remarks.-Genoways and Jones (1971) surements. gave means and ranges ofmeasurements for three samples ofperotensis.

Dipodomys phillipsii ornatus Merriam, 1894 Dipodomys phillipsii phillipsii Gray, 1841 1894. Dipodomys ornatus Merriam, Proc. BioI. Soc. Washington, 9: 110, 21 June. 1841. Dipodomys phillipii [sic1Gray, Ann. Mag. Nat. Hist., ser. 1, 7:522, August. Holotype. -Adult female, skin and skull, USNM 57990, from Berriozabal, Zacate­ Holotype. -Adult male, skin and skull, cas, Mexico; obtained by Edward A. Gold­ BM(NH) 45.1580, from near Real del Mon­ man on 29 December 1893. te, Hidalgo, Mexico; obtained by John Phil­ Measurements ofholotype. - Total length, lips. 274; length oftail, 167; length ofhind foot, Measurements ofholotype. - Total length, 39; greatest length ofskull, 36.5; width across 292; length oftail, 165; length ofhind foot, bullae, 22.6; length ofnasals, 13.0; breadth 38. across maxillary arches, 21.4; interorbital Distribution. - Limited to the Valle de breadth, 13.6; length ofrostrum, 20.9; length Mexico and adjacent areas in Hidalgo and of maxillary toothrow, 4.8; depth of crani­ the Distrito Federal, Mexico (Genoways and um, 11.0. Jones, 1971). Distribution. -Ranges from central Du­ Remarks. - All ofthe skull except the up­ rango southeastward to Queretaro on the per and lower incsiors of the holotype are Mexican Plateau (Genoways and Jones, missing. See Genoways and Jones (1971) for 1971). a discussion ofcontroversy in the literature Remarks.-Genoways and Jones (1971) about the type locality. Coues (1875) dis- TAXONOMY 85 cussed the spelling of the specific epithet. antiquarius was a subspecies ofD. agilis (now Genoways and Jones (1971) listed means simulans). and ranges of measurements for a sample of phillipsii. Dipodomys simulans peninsularis (Merriam, 1907) Dipodomys simulans 1907. Perodipus simulans peninsularis Merri­ am, Proc. BioI. Soc. Washington, 20:79, 22 Diagnosis. -A medium-sized kangaroo July. rat with five toes on the hind feet, moder­ 1951. Dipodomys peninsularis pedionomus ately long ear pinnae, relatively narrow Huey, Trans. San Diego Soc. Nat. Hist., 11: breadth across the maxillary arches, and 60 247, 30 April. chromosomes. Length ofhead and body av­ 1951. Dipodomyspeninsularis eremoecus Huey, erages about 115 mm and length of hind Trans. San Diego Soc. Nat. Hist., 11 :248, 30 foot averages less than 42 mm; greatest April. length of skull averages less than 39.5 mm 1951. Dipodomys peninsularis australis Huey, and maxillary breadth averages less than Trans. San Diego Soc. Nat. Hist., 11:249, 30 21.0 mm. April. Comparisons. -Dipodomys simulans is 1962. Dipodomys antiquarius Huey, Trans. San Diego Soc. Nat. Hist., 12:477, 30 August. most similar to D. agilis, and must be dis­ tinguished from sympatric individuals ofD. Holotype. - Young adult male, USNM gravipes, and D. stephensi. From D. gra­ 139872, from Santo Domingo [Landing], vipes, simulans can be distinguished by the 28°51'N lat., 114°W long., Baja California, narrower breadth across the maxillary arch­ Mexico; obtained on 27 September 1905 by es, smaller hind feet, and larger ears. See Edward W. Nelson and Edward A. Gold­ account of agilis for comparison with that man. species; from stephensi, simulans differs in Measurements ofholotype. - Total length, being smaller with a narrower skull across 312; length oftail, 203; length ofhind foot, the maxillary arches and across the bullae; 45; greatest length of skull, [40.80]; width the bullae of stephensi are more globose in across bullae, 26.75; breadth across maxil­ shape. lary arches, 20.95; nasal length, [13.60­ Distribution. -Occupies coastal chapar­ tips broken]; interorbital breadth, 11.85. ral and grassland communities from the Los Distribution. -Occurs on the central and Angeles Basin and San Jacinto mountains southern portions of the Baja California of southern California, southward in Baja Peninsula, from the vicinity ofSan Fernan­ California to the vicinity ofBahia Almejas, do Mission, Baja California southward to Pacific Coast, Baja California Sur. the vicinity ofMagdalena and Almejas bays, Remarks. - Best et al. (1986) and Sullivan near lat. 24°30'N. and Best (in press) discussed the relation­ Remarks. -Best (1978) reported on sys­ ships ofD. agilis and D. simulans, and pro­ tematic investigations of D. s. peninsularis vided methods of distinguishing the two and its related subspecies in Baja California, species. Huey (1951) treated peninsularis as determining that D. peninsularis was not a species, and Best (1978) listed it as a sub­ specifically distinct from D. simulans and species ofD. agilis. Huey (1962) considered that D. antiquaris was indistinguishable from D. antiquarius, from Sierra Borja, Baja Cal­ D. s. pedionomus. Best (1983b) showed a ifornia, to be closest to D. stephensi, but close similarity in structure between Dipod­ Lackey (1967) could find no significant dif­ omys s. peninsularis and D. s. australis, ferences between samples ofantiquarius and whose geographic ranges are contiguous. The D. peninsularis. Stock (1974) postulated that current arrangement ofsubspecies and their 86 WILLIAMS ET AL. synonyms follows Sullivan and Best (in Baja California, Huey (1951) determined press). that specimens he referred to D. agilis lati­ maxillaris did not differ substantially from specimens from some populations ofsimu­ Dipodomys simulans simulans lans. Morphometric analyses by Best (1981, (Merriam, 1904) 1983b) showed little differentiation among some samples of martirensis, plectilis, and 1904. Perodipus streatori simulans Merriam, simulans. Best (1978) considered that D. Proc. BioI. Soc. Washington, 17:144, 14 July. paralius was indistinguishable from plectil­ 1904. Perodipus cabezonae Merriam, Proc. BioI. is. Soc. Washington, 17: 144, 14 July. 1925. Dipodomys agilis latimaxillaris Huey, Proc. BioI. Soc. Washington, 38:84, 26 May. Dipodomys spectabilis 1927. Dipodomys agilis martirensis Huey, Trans. San Diego Soc. Nat. Hist., 5:7, 20 February. Diagnosis. - One ofthe largest species of 1951. Dipodomys agilis plectilis Huey, Trans. kangaroo rats, with four toes on the hind San Diego Soc. Nat. Hist., 11 :240, 30 April. feet and with a tail terminating in a large, 1951. Dipodomysparalius Huey, Trans. San Di­ white tuft of hairs (tuft exceeds about 25 ego Soc. Nat. Hist., 11:241, 30 April. mm in length, typically about 40 mm) bor­ Holotype. - Adult female, skin and skull, dered proximally by a band of black hairs; USNM 33105/45103, from Dulzura, San lateral white stripes of tail present only on Diego Co., California; obtained on 24 No­ about the proximal half; underside of tail, vember 1891 by C. H. Marsh. proximal to white tuft, dark colored; head Measurements ofholotype. - Total length, and body length averaging from about 128 280; length oftail, 165; length ofhind foot, to 150 mm; the auditory bullae relatively 40; greatest length of skull, 38.00; width large and inflated; skull relatively wide across across bullae, 24.40; breadth across maxil­ the maxillary arches (breadth averaging from lary arches, 20.15; nasal length, 13.30; in­ about 25.0 to 26.3). terorbital breadth, 12.65. Comparisons. - Dipodomys spectabilis is Distribution. -On the north found in Pa­ most similar in size and appearance to D. cific coastal chaparral and grassland com­ nelsoni; see the account ofthe latter species munities generally below about 500 m in for diagnostic characters. It is also similar elevation in Los Angeles and San Bernar­ in size to the four-toed species, D. deserti, dino counties; also found on both the Pacific from which D. spectabilis can be distin­ anddesert slopes ofcoastal mountain ranges guished by the prominent black border to from San Gorgonio Pass, Riverside Co., the white tail tuft, the dark underside to its California, southward. Range extends tail, generally smaller hind feet (usually av­ through the coastal basins and mountains eraging from 47 to 52 in spectabilis and from of southern California southward on the 52 to 53.5 in deserti), and a less inflated slopes of the Sierra Juarez and Sierra San skull with a wider interorbital region and Pedro Martir and the Pacific coastal plains wider breadth across the maxillary arches of Baja California to the vicinity of Santa (Nader, 1978). See the accounts of D. cali­ Catarina and Santa Catarina Landing. fornicus and D. elator for other compari­ Remarks.-Grinnell (1922) noted only sons. slight and inconstant differences in speci­ Distribution. -Banner-tailed kangaroo mens ofD. a. agilis and simulans; the major rats occur in desert associations from south­ difference was the greater inflation of the central Arizona and northern Sonora, east­ bullae of simulans. After obtaining speci­ ward to the Trans-Pecos region of Texas; mens from several additional localities in they also extend from extreme north-central TAXONOMY 87

Arizona and the Four Corners Region ofthe Remarks. -According to Nader (1978), San Juan River Basin, northeastern Arizo­ the type locality is in Lincoln Co., not Chaves na, southward through lower-lying portions Co. as listed by Goldman. Nader (1978) ofNew Mexico; their range continues from found no consistent differences between northwestern Chihuahua, southward and samples of populations of clarencei and eastward through central Chihuahua to baileyi, and regarded clarencei as indistin­ northeastern Durango. A disjunct popula­ guishable from baileyi. Hall (198l) did not tion occurs south ofthe range ofD. nelsoni cite Nader's (1978) revision and did not ex­ in east-central Zacatecas, northeastern plain his reasoning for retention ofclarencei Aguascalientes, and western San Luis Po­ as a subspecies. We concur with Nader tosi. (1978). Nader (1978) provided ranges and Remarks. -Dipodomys nelsoni is treated means of measurements for D. s. baileyi. as a separate species herein despite evidence Hoffmeister (1986) reported on specimens of limited hybridization in a narrow zone from near Rainbow Lodge, Coconino Co., of contact with D. spectabilis in southern Arizona, approximately 112 km west and Chihuahua (Nader, 1978). The relation­ 80 km north of the populations in north­ ships between spectabilis and nelsoni, and western Arizona studied by Nader (1978). the relationship ofthe disjunct population, He tentatively assigned these specimens to cratodon, in central Mexico to other pop­ baileyi. ulations of D. spectabilis and to D. nelsoni require further study. Dipodomys spectabilis cratodon Merriam, 1907 Dipodomys spectabilis baileyi Goldman, 1923 1907. Dipodomys spectabilis cratodon Merriam, Proc. Biol. Soc. Washington, 20:75, 22 July. 1923. Dipodomys spectabilis baileyi Goldman, Holotype. - Adult male, skin and skull, Proc. Biol. Soc. Washington, 36:140, 1 May. USNM 78953, from Chicalote, Aguasca­ 1933. Dipodomys spectabilis clarencei Gold­ lientes, Mexico; obtained on 2 July 1896 by man, J. Washington Acad. Sci., 23:467,15 Oc­ Edward W. Nelson and Edward A. Gold­ tober. man. Holotype. -Adult male, skin and skull, Measurements ofholotype. - Total length, USNM 97185, from 40 mi W Roswell, 342; length oftail, 217; length ofhind foot, Chaves Co. [Lincoln Co.], New Mexico; ob­ 54; greatest length ofskull, 46.5; width across tained by Vernon Bailey on 13 June 1899. bullae, 29.7; length ofnasals, 15.6; breadth Measurements ofholotype. - Total length, across maxillary arches, 27.5; interorbital 385; length oftail, 283; length ofhind foot, breadth, 17.0; length ofrostrum, 25.2; length 58; greatest length ofskull, 47.4; width across of maxillary toothrow, 6.6; depth of crani­ bullae, 30.7; length ofnasals, 15.5; breadth urn, 14.2. across maxillary arches, 26.8; interorbital Distribution. -Occupies an area disjunct breadth, 15.6; length ofrostrum, 24.8; length from other populations of D. spectabilis in of maxillary toothrow, 6.3; depth of crani­ east-central Zacatecas, northeastern Aguas­ um, 14.5; least width ofsupraoccipital, 1.7. calientes, and western San Luis Potosi. Distribution. - Found from northeastern Remarks. -See the account ofD. nelsoni Arizona (where it no longer occurs; Hoff­ and the species diagnosis above concerning meister, 1986) through western, central, and the relationships of cratodon to other spec­ southeastern New Mexico, and the Trans­ tabilis group kangaroo rats. Dalquest (1953) Pecos area ofwestern Texas (Nader, 1978). and Nader (1978) listed measurements. 88 WILLIAMS ET AL.

Dipodomys spectabilis intermedius Distribution. - Found in south central Ar­ Nader, 1965 izona south of the Gila River, west of the Santa Catalina and Santa Rita ranges and 1965. Dipodomys spectabilis intermedius Na­ east from the Ajo Valley, southward into der, Proc. BioI. Soc. Washington, 78:50, 21 north central Sonora. July. Remarks.-Nader (1978) listed means Ho!otype. - Adult female, skin and skull, and ranges of measurements. Populations MVZ 82782, from 16.7 mi SW Bamori, ofthis subspecies are among the smallest of about 1,900 ft, Sonora, Mexico; obtained the species. by Seth B. Benson on 25 April 1938. Measurements ofho!otype. - Total length, Dipodomys spectabilis spectabilis 314; length oftail, 180; length ofhind foot, Merriam, 1890 47; length ofear, 17; weight, 105.7 g; great­ est length ofskull, 41.9; width across bullae, 1890. Dipodomys spectabilis Merriam, N. Amer. 27.1; breadth across maxillary arches, 24.8; Fauna, 4:46, 8 October. nasal length, 15.0; interorbital breadth, 15.65. Ho!otype. - Young adult male, skin and Distribution. - West central Sonora, skull, USNM 17886/24823, from Dos Ca­ Mexico, from Querobabi on the north, bezos [= Dos Cabezas], Cochise Co., Ari­ southward to Carbo and westward to a lo­ zona; obtained by Vernon Bailey on 22 No­ cality about 17 mi southwest of Bamori vember 1889. (Nader, 1978). Measurements ofho!otype. - Total length, Remarks. -Nader (1965) listed means 350; length oftail, 211; length ofhind foot, and ranges ofmeasurements for seven spec­ 52; length ofear (crown), 10, (anterior base, imens from the type locality. This race is dry skin), 16; greatest length of skull, 45.6; the smallest in size of the subspecies of D. width across bullae, 29.5; length of nasals, spectabilis. 15.9; interorbital breadth, 15.8; length of rostrum, 23.9; length ofmaxillary toothrow, 5.8; depth ofcranium, 13.3. Dipodomys spectabilis perb!andus Distribution. - Desert associations in Goldman, 1933 southeastern Arizona east ofthe Galiuro and Santa Rita ranges, northeastern Sonora, 1933. Dipodomys spectabilis perblandus Gold­ southwestern New Mexico, and northern man, J. Washington Acad. Sci., 23:466,15 Oc­ and central Chihuahua. tober. Remarks. -Nader (1978) and Anderson Ho!otype. - Adult female, skin and skull, (1972) gave means and ranges of measure­ USNM 17748/24689, from Calabasas, ments for samples of D. spectabilis specta­ about 3,500 ft, Santa Cruz Co., Arizona; bilis. obtained by Vernon Bailey on 27 October 1889. Measurements ofho!otype. - Total length, Dipodomys spectabilis zygomaticus 315; length oftail, 184; length ofhind foot, Goldman, 1923 48; greatest length ofskull, 42.9; width across bullae, 27.7; length ofnasals, 14.4; breadth 1923. Dipodomys spectabilis zygomaticus Gold­ across maxillary arches, 25.9; interorbital man, Proc. BioI. Soc. Washington, 36:140, 1 May. breadth, 15.2; length ofrostrum, 22.7; length of maxillary toothrow, 5.5; depth of crani­ Ho!otype. - Young adult male, skin and um, 13.1; width of nasals in front of inci­ skull, USNM 96432, from Parral [= Hi­ sors, 4.2; least width of supraoccipital, 2.3. dalgo del Parral], southern Chihuahua, TAXONOMY 89

Mexico; obtained by Edward A. Goldman D. simulans (distance across maxillary arch­ on 17 September 1898. es averages about 20.8 to 20.9 in simulans; Measurements ofholotype. - Total length, Lackey, 1967). 340; length oftail, 197; length ofhind foot, Distribution. -Confined to open, arid 54; greatest length ofskull, 46.1; width across grassland associations in a relatively small bullae, 30.6; length ofnasals, 16.2; breadth area in southern California, in the San Ja­ across maxillary arches, 27.1; interorbital cinto Valley and adjacent lowlands ofwest­ breadth, 16.4; length ofrostrum, 24.7; length em Riverside, southwestern San Bernardi­ of maxillary toothrow, 6.5; depth of crani­ no, and northwestern San Diego counties um, 13.5. (Bleich, 1977; Lackey, 1967); an apparently Distribution. -Occupies a range in south disjunct population occurs in north-central central Chihuahua and north central Du­ rango (Nader, 1978). San Diego Co., in the vicinity of Warner Remarks. -Anderson (1972) and Nader Springs (O'Farrell et al., 1986). (1978) gave means and ranges for measure­ Remarks. - Bleich (1977) reviewed pa­ ments of samples of D. s. zygomaticus. pers discussing the relationships of D. ste­ phensi to other species of Dipodomys; al­ though most authors agree that D. stephensi is related to the other broad-faced kangaroo Dipodomys stephensi rats ofthe heermannigroup, its interspecific relationships are unresolved. Diagnosis. -A medium-sized kangaroo rat with five toes on the hind feet, a wide face, reflected in broad maxillary arches, and relatively large and inflated auditory bullae Dipodomys stephensi (Merriam, 1907) with a unique, "globose" shape (Grinnell, 1922); length of head and body averages 1907. Perodipus stephensi Merriam, Proc. BioI. about 119 to 121 mm; width across the Soc. Washington, 20:78, 22 July. maxillary arches averages about 22.5 mm; 1962. Dipodomys cascus Ruey, Trans. San Di­ ego Soc. Nat. Rist., 12:479, 30 August. greatest breadth of skull, across the bullae, averages about 24.7 to 25.1 mm or more Holotype. - Adult male, skin and skull, (Lackey, 1967). USNM 186503, from San Jacinto Valley [a Comparisons. - Dipodomys stephensi is little W of Winchester, toward Menifee], similar to the allopatric species D. heer­ Riverside Co., California; obtained on 27 manni and D. panamintinus. From pana­ November 1885 by Frank Stephens. mintinus, stephensi is distinguished by its Measurements of holotype. -Greatest greater breadth across the auditory bullae length ofskull, [39.95]; width across bullae, (average 24.7 mm or greater in stephensi, 25.45; nasal length, [14.45]; interorbital 24.2 or less in panamintinus); the ratio of breadth, 12.30. interparietal length to width is 0.3 in ste­ Remarks. -No external measurements phensi and 0.6 inpanamintinus (see account were taken for the holotype, and its max­ of D. panamintinus and Lackey, 1967, for illary arches and tips ofnasals are missing. other diagnostic characters). Lackey (1967) According to Lackey (1967), the population stated that D. stephensi was more similar named cascus is similar to, but recognizably structurally to the southern subspecies ofD. distinct from the more northern population, heermanni than to other species; compared stephensi; Lackey did not recommend sub­ to D. heermanni tularensis, D. stephensi has specific recognition, however. The two pop­ a shorter basioccipital width and greater ju­ ulations apparently are currently allopatric. gal curvature. The face of D. stephensi is Lackey (1967) provided measurements and wider than the sympatric five-toed species, comparisons to other species. 90 WILLIAMS ET AL.

Dipodomys venustus that they are conspecific (see account ofD. elephantinus). D. venustus also is similar to Diagnosis. - A moderately large-sized, the northern populations of D. agilis and large-eared, five-toed kangaroo rat with a may be conspecific (Stock, 1974). Means relatively long tail, large feet, and dark col­ and ranges of measurements were given by oration; length of the ear pinna exceeded Grinnell (1922). only by D. elephantinus; face narrow and spread ofmaxillary arches usually less than 25.5 mm; auditory bullae large and back of Dipodomys venustus santiluciae skull wide; supraoccipital and interparietal Grinnell, 1919 very narrow; incisors relatively robust; na­ sals typically not flaring at end (Grinnell, 1919. Dipodomys santiluciae Grinnell, Proc. 1922). Bioi. Soc. Washington, 32:204, 31 December. Comparisons. - Dipodomys venustus can Holotype. - Adult male, skin and skull, be distinguished from all other kangaroo rats MVZ 29023, from I mi SW Jolon, Mon­ except D. elephantinus by having five toes terey Co., California; obtained by Joseph on the hind feet, long ears (length from crown Grinnell on 21 October 1918. in excess of 15.5 mm), a relatively long, Measurements ofholotype. - Total length, heavily crested tail averaging about 155% 315; length oftail, 119; length ofhind foot, oflength ofhead and body, and large body 46; length of ear (crown), 16; weight, 82 g; size; from typical individuals ofD. venustus, greatest length of skull, 42.5; width across D. elephantinus differs in being lighter in bullae, 25.8; length ofnasals, 15.5; breadth color, having longer ears (length ofear from across maxillary arches, 23.0; width ofmax­ crown in venustus averages less than about illary arch at middle, 5.3; width ofrostrum 16 mm), and more flaring nasals (see ac­ near end, 4.3. count of elephantinus for additional re­ Distribution. -Chaparral associations of marks). D. venustus differs from D. agilis in the coastal mountains, from the south end being slightly larger in most dimensions and ofMonterey Bay, Monterey Co., southward in having longer ears; length ofhind foot in to the Santa Lucia Mountains east ofMorro D. venustus averages 45 mm or more and Bay, San Luis Obispo Co., California. The less than 45 mm in D. agilis. Salinas Valley marks the eastern boundary Distribution. -Dipodomys venustus oc­ of the range. cupies chaparral communities on loose soils Remarks. - Grinnell (1922) provided in the coastal ranges of west-central Cali­ means and ranges ofmeasurements for two fornia. Its range extends from the Santa Cruz males and four females. Mountains and hills near the south end of San Francisco Bay in Santa Cruz and Santa Clara counties, eastward in the Diablo Range in western Stanislaus Co., thence southward Dipodomys venustus venustus along the outer coastal ranges to the Santa (Merriam, 1904) Lucia Range, San Luis Obispo Co., and in the inner coastal ranges (Diablo Range) to 1904. Perodipus venustus Merriam, Proc. Bioi. Soc. Washington, 17:142, 14 July. south of the boundary between San Benito and Fresno counties, in western Fresno Co. Holotype. -Adult male, skin and skull, (Grinnell, 1922; D. F. Williams, unpubl. USNM 51852, from Santa Cruz, Santa Cruz data). Co., California; obtained by G. B. Badger Remarks. -Some authorities have re­ on 12 March 1893. marked on the close similarity between D. Measurements ofholotype. - Total length, venustus and D. elephantinus and opined 339; length oftail, 211; length ofhind foot, TAXONOMY 91

46; greatest length of skull, 41.60; width hood; upper incisors grooved; knob of in­ across bullae, 24.75; breadth across maxil­ cisor pulp cavity on mandible at lower edge lary arches, 22.00; nasal length, 15.35; in­ of ascending ramus; incipient postorbital terorbital breadth, 12.10. process; center ofpalate between premolars Distribution. -Apparently disjunct pop­ not ridged; one pterygoid fossa apparent ulations occupy the Santa Cruz Mountains (anterior fossa nearly invisible); zygomatic and adjacent area west of the Santa Clara process of maxilla not much expanded; Valley, the Diablo Range in Santa Clara, masseter fossa separated from infraorbital Stanislaus, Merced, San Benito, and Fresno foramen by slight crest; posterior palantine counties, and the northern end of the Ga­ foramen located in palantine; auditory bul­ bilan Range in Monterey and San Benito lae more inflated than in other genera ofthe counties (Grinnell, 1922; Hall, 1981; un­ family- bullae meeting in symphysis across publ. data). ventral face ofbasisphenoid; tympanic bul­ Remarks. - Samples ofpopulations from la extends anterior to mandibular fossa (gle­ the Diablo Range are virtually indistin­ noid); ventral surface oftympanic bulla ex­ guishable from D. elephantinus. Grinnell tending below occlusal surface ofcheekteeth; (1922) gave ranges and means of measure­ obturator foramen of pelvis subtriangular ments for six males and two females from and much compressed dorso-ventrally; as­ the westernmost populations. tragalar-cuboid contact in foot and no con­ tact between navicular and calcaneum; tra­ pezium-scapholunar articulation; cervical Genus Microdipodops vertebrae mostly unfused; fewer caudal ver­ tebrae than other members of family, but 1891. Microdipodops Merriam, N. Amer. Fau­ individual vertebrae elongated; median na, 5:115, 30 July. ventral foramen in caudal vertebrae; tibia Type species. - Microdipodops mega­ and fibula fused for about 60% of their cephalus Merriam, 1891, N. Amer. Fauna, length; temporalis muscle reduced; M. ten­ 5:116, 30 July. sor fasciae latae insertion includes medial Diagnosis. - Size small, total length 130­ thigh; pallus with spines, urethral lappets, 180 mm and mass 10-17 g; body form rico­ and a dorsal groove; tip of phallus not up­ chetal; tail thickest near its midlength and turned; ventromedial prostrate gland ab­ only slightly longer than length ofhead and sent; baculum with large, bulbous basal part, body; tail with short hairs, not crested or tapering into a moderately upward-curving tufted at end; tail with fat deposit centered shaft; spermatozoa large with long, roughly at proximal one-third to one-half of tail; triangular head with rounded vertices, and body hairs relatively long, lax, and silky; no tail of medium length (Burt, 1960; Hafner large dermal sebaceous gland on back; soles and Hafner, 1983; Hall, 1941; Hatt, 1932; ofhind feet densely covered with long hair; Homan and Genoways, 1978; Quay, 1965; pes with five well-developed digits; manus Ryan, 1989; Wahlert, 1985; Wood, 1935). longer and more slender than other mem­ Remarks. - The subfamilial relationships bers of family; protoloph of P4 with one of Microdipodops have been controversial cusp; lophs ofP4 unite first at or near center since its description by Merriam (1891). oftooth; p4 with 5 or 6 cusps; molars with Wood (1935), in a major review ofhetero­ H-pattern; cusp patterns worn away quickly myid evolution, placed the kangaroo mice leaving occlusal surface ofcheek teeth as an in the subfamily Perognathinae, allying them island of dentine surrounded by enamel; with the silky pocket mice, Perognathus. cheek teeth extremely high-crowned and Another possible arrangement, in which with more than one root except M3/m3; kangaroo mice represent a distinct evolu­ roots of cheek teeth develop after adult- tionary line, was raised when Setzer (1949) 92 WILLIAMS ET AL. declined to place them in any recognized Comparisons. - Many of the characters subfamily. Reeder (1956) first placed them given in the key and diagnosis above are in the subfamily Dipodomyinae based on variable geographically. Individuals of the dental characters (see remarks in the diag­ two species from close geographic proxim­ nosis for Dipodomyinae for additional in­ ity should be compared to confirm identi­ formation). fication. In as far as is known, the diploid chromosome numbers are diagnostic (40 and 42; Hafner et aI., 1979). The dorsal tip Key to the Species of the tail of M. megacephalus is generally black, whereas in M. pallidus this area is the 1. Length of hind foot usually 25 mm or same color as the back. The shape of the less; anterior palatine foramina wide pos­ anterior palatine foramina and the point of teriorly and tapering to a sharp point an­ the posterior termination of the nasals ap­ teriorly; premaxillae terminating at, or pear to be the most useful cranial characters extending less than 1mm posterior to the for separating the species, but there is some posterior end of nasals; upper parts dark geographic variation in these traits in M. brownish or buffy with overwash of blackish; dorsal surface of tail blackish pallidus(Hall, 1941). distally and tipped in black . Distribution. - Found on a variety ofsub­ ...... Microdipodops megacephalus strates in the Great Basin region including 1'. Length of hind foot usually 25 mm or Nevada, southeastern Oregon, extreme more; anterior palatine foramina paral­ southwestern Idaho, west-central Utah, and lel-sided; premaxillae terminating pos­ extreme northeastern and east-central teriorly more than 1-2 mm beyond pos­ (Mono Co.) California. terior end ofnasals; upper parts pale, near Remarks. - The two species of kangaroo Light Pinkish Cinnamon (Ridgway, mice occur sympatrically, but areas ofsym­ 1912); top of tail distally approximately patry are not large. Hall (1941) reported same color as near base . sympatry at 12 localities. At one ofthese­ ...... Microdipodops pallidus Penoyer Valley, Nevada- he reported three hybrid individuals. Later examination of Species Accounts these and 89 additional specimens from the same area found only one individual that was structurally intermediate between the Microdipodops megacephalus species (Hafner et aI., 1979). Karyotypes and electrophoretic patterns, however, showed Diagnosis. -Anterior palatine foramina no evidence of hybridization. wide posteriorly and tapering to a sharp point anteriorly; nasals terminating poste­ riorly at, or almost at, the same level as the premaxillae; skull with less inflated auditory Microdipodops megacephalus albiventer bullae, thus a proportionally narrower skull; Hall and Durrant, 1937 upper incisors relatively curved; hind foot length averages about 23-35 mm; post-au­ 1937. Microdipodops pallidus albiventer Hall and Durrant, J. Mamm., 18:357, 14 August. ricular spot buffy; upper parts dark, brown­ ish or buffy, washed with blackish; hairs of Holotype. - Adult male, skin and skull, underparts grayish for most of length, but MVZ 52803, from Desert Valley, 5,300 ft, either subterminally white with buffy tips 21 mi W Panaca, Lincoln Co., Nevada; ob­ or lacking buffy and tipped with white (a tained on 30 May 1932 by Ward C. Russell. few populations have hairs white through­ Measurements ofholotype. - Total length, out their length); tail above tipped with 150; length of tail, 80; length of hind foot, black; diploid number 40. 24; length ofear, 9.5; weight, 11.6 g; greatest TAXONOMY 93 length of skull, 28.30; width across bullae, (1941) listed means and ranges ofmeasure­ 18.85; length ofnasals, 9.80; breadth across ments for 17 adult topotypes. maxillary arches, 11.90; interorbital breadth, 6.50; length of maxillary toothrow, 3.50; depth of cranium, 8.00. Microdipodops megacephalus atrirelictus Distribution. - Restricted to Desert Val­ Hafner, 1985 ley in central Lincoln Co., Nevada. Remarks. -Hall and Durrant (1937) de­ 1985. Microdipodops megacephalus atrirelictus scribed albiventer as a subspecies ofM. pal­ Hafner, Proc. Bioi. Soc. Washington, 98:3, 20 lidus because of its light coloration. They March. gave measurements for 16 adult topotypes. Holotype. - Adult female, skin, skull, and Hall (1941) classified it as a subspecies of skeleton, MVZ 160039, from 11 mi S, 44.2 M. megacephalus based upon cranial char­ mi W Riddle, 5,000 ft, Owyhee Co., Idaho; acters. M. m. albiventer, as currently de­ obtained on 8 October 1978 by John C. fined, is restricted to Desert Valley, Nevada, Hafner. although specimens from Coal Valley to the Measurements ofholotype. - Total length, west assigned to sabulonis approach them 169; length of tail, 93; length of hind foot, in coloration. 26; length ofear, 12; weight, 12.5 g; greatest length of skull, 29.27; width across bullae, 19.16; basal length of skull, 19.06; bullar Microdipodops megacephalus ambiguus length, 14.60; length of nasals, 10.35; Hall, 1941 breadth across maxillary arches, 11.62; in­ terorbital breadth, 6.69; greatest length of 1941. Microdipodops megacephalus ambiguus incisive foramina, 2.19; length ofmaxillary Hall, Field Mus. Nat. Hist., Zooi. Ser., 27:252, toothrow, 3.45; depth of cranium, 7.95. 8 December. Distribution. -Known only from south­ Holotype. -Adult male, skin and skull, western Owyhee Co., Idaho. MVZ 73840, from 1.25 mi N Sulphur, 4,050 Remarks. - This is among the most high­ ft, Humboldt Co., Nevada; obtained on 25 ly differentiated subspecies of Microdipo­ July 1936 by E. Raymond Hall. dops. It generally is larger than other sub­ Measurements ofholotype. - Total length, species ofM. megacephalus, and its dorsal 162; length of tail, 89; length of hind foot, pelage is nearly black. These kangaroo mice 25; length ofear, 9.8; weight, 13.2 g; greatest are isolated from other members ofthe ge­ length of skull, 29.00; width across bullae, nus by over 100 km of ground unsuitable 19.10; length ofnasals, 10.35; breadth across as habitat (Hafner, 1985). Hafner (1985) maxillary arches, 11.30; interorbital breadth, listed statistics for four individuals. 6.55; length of maxillary toothrow, 3.45; depth ofcranium, 8.10. Distribution. -Smoke Creek and Black Microdipodops megacephalus califomicus Rock deserts and the lower part of Hum­ Merriam, 1901 boldt River Valley in Nevada and extreme eastern Lassen Co., California. 1901. Microdipodops califomicus Merriam, Proc. Bioi. Soc. Washington, 14:128, 19 July. Remarks. - According to Hall (1941), specimens ofambiguus are widely variable, Holotype. - Young adult male, skin and particularly in external proportions and col­ skull, USNM 101227, from Sierra Valley, oration. This taxon is in contact with four near Vinton, Plumas Co., California; ob­ other subspecies. Hall (1941) found evi­ tained on 7 August 1900 by Walter K. Fish­ dence of integradation between ambiguus er. and each ofthese four other subspecies. Hall Measurements ofholotype. - Total length, 94 WILLIAMS ET AL.

158; length of tail, 91; length of hind foot, Microdipodops megacephalus medius 25; greatest length ofskull, 26.3; width across Hall, 1941 bullae, 17.7; length of nasals, 8.3; breadth across maxillary arches, 11.3; interorbital 1941. Microdipodops megacephalus medius Hall, breadth, 6.2; length of maxillary toothrow, Field Mus. Nat. Hist., Zool. Ser., 27:256, 8 3A; depth of cranium, 7.8; length of ros­ December. trum lOA. Holotype. - Adult female, skin and skull, Distribution. -Occurs in the intermon­ MVZ 73890, from 3 mi S Vernon, 4,250 ft, tane valleys ofeastern Plumas Co., Califor­ Pershing Co., Nevada; obtained on 28 July nia, and southern Washoe and Ormsby 1936 by E. Raymond Hall. counties, Nevada. Measurements ofholotype. - Total length, Remarks. - Hall (1941) found evidence 165; length of tail, 89; length of hind foot, of intergradation between califomicus and 25; length of ear, 10.5; greatest length of ambiguus at several points of contact be­ skull, 28.55; width across bullae, 18.65; tween their ranges, and therefore relegated length ofnasals, 10.10; breadth across max­ califomicus to subspecific status. Hall (1941) illary arches, 11.35; interorbital breadth, provided means and ranges of measure­ 6.25; length of maxillary toothrow, 3.55; ments for 5 adults and Hafner (1985) listed depth ofcranium, 7.85. statistics for measurements of 12 individ­ Distribution. -Southwestern Pershing uals. Co., Nevada. Remarks. - The only other subspecies with which medius is in contactis ambiguus. Microdipodops megacephalus leucotis From ambiguus, medius is distinguished by Hall and Durrant, 1941 its darker coloration and narrower skull re­ sulting from less inflated auditory bullae. 1941. Microdipodops megacephalus leucotis Hall Hall (1941) listed means and ranges ofmea­ and Durrant, The Murre1et, 22:6, 30 April. surements for 18 adult topotypes, and Haf­ Holotype. - Adult female, skin and skull, ner (1985) listed statistics for 12 individu­ UU 3525 from 18 miSWOrr'sRanch, 4,400 als. ft, Tooele Co., Utah; obtained on 6 June 1940 by Stephen D. Durrant. Measurements ofholotype. - Total length, Microdipodops megacephalus 142; length of tail, 75; length of hind foot, megacephalus 24; basa11ength (to anterior face ofincisor), Merriam, 1891 17.5; width across bullae, 19.2; length of nasals, 9.3; breadth across maxillary arches, 1891. Microdipodops megacephalus Merriam, N. Amer. Fauna, 5:116, 30 July. 11.3; interorbital breadth, 6.1. Distribution. - Restricted to the sand Holotype. -Adult male, skin and skull, dunes along the valley floors of the Bon­ USNM 24417/31823, from Halleck, Elko neville Basin in south-central Tooele Co., Co., Nevada; obtained 23 October 1890 by Utah. Vernon Bailey. Remarks. - This taxon is one ofthe palest Measurements ofholotype. - Total length, members ofthe genus, being palerthan many 157; length of tail, 83; length of hind foot, ;;opu1ati)ns ofM. pallidus. However, most 24; greatest length ofskull, 28A; width across cranial and external features ally leucotis bullae, 19.3; length of nasals, 9A; breadth with M. megacephalus (Hall, 1941). Later, across maxillary arches, 11.9; interorbital Hall (1981) remarked that leucotis might be breadth, 6.7; length of maxillary toothrow, a distinct species, but Hafner and Hafner 3A; depth of cranium, 7.9; length of ros­ (1983) found that genic and karyotypic ev­ trum, 10.6. id,;nce cEd not support specific status. Distribution. -Central and northeastern TAXONOMY 95

Nevada, from northern Elko Co. to north­ Co., Nevada; obtained on 22 May 1936 by ern Nye and Lincoln counties, and from Ward C. Russell. western Lander Co. eastward to near the Measurements ofholotype. - Total length, Utah border. 167; length of tail, 99; length of hind foot, Remarks. -Specimens from Eureka Co., 25; length ofear, 10; greatest length ofskull, Nevada, represent possible intergrades be­ 29.25; width across bullae, 19.25; length of tween megacephalus and nexus. There is a nasals, 10.40; breadth across maxillary broad zone ofintegradation with M. m. sa­ arches, 11.55; interorbital breadth, 6.60; bulonis in south-central Nevada. Hall (1941) length of maxillary toothrow, 3.30; depth gave means and ranges ofmeasurements for of cranium, 7.85. 10 adults, and Hafner (1985) listed statistics Distribution. -Found in Humboldt and for 12 individuals. Lander counties, Nevada. Remarks. - The subspecies nexus is sep­ arated from ambiguus to the west by a low Microdipodops megacephalus nasutus range ofmountains and there is a 75-mi gap Hall, 1941 to the eastward with no dark kangaroo mice. In color, nexus is intermediate between the 1941. Microdipodops megacephalus nasutus western, gray mice and the blackish mice to Hall, Field Mus. Nat. Hist., ZooI. Ser., 27:251, the east. Hall (1941) gave means and ranges 8 December. ofmeasurements for 20 adult topotyp~s ar.d Holotype. - Adult female, skin and skull, Hafner (1985) listed statistics for 12 indi­ MVZ 40439, from Fletcher, 6,098 ft, Min­ viduals. eral Co., Nevada; obtained on 22 July 1928 by Louise Kellogg. Measurements ofholotype. - Total length, Microdipodops megacephalus oregonus 158; length of tail, 88; length of hind foot, Merriam, 1901 25; greatest length of skull, 28.00; width across bullae, 18.25; length ofnasals, 10.10; 1901. Microdipodops megacephalus oregonus breadth across maxillary arches, 11.8; in­ Merriam, Proc. BioI. Soc. Washington, 14:127, terorbital breadth, 6.65; length ofmaxillary 19 July. toothrow, 3.40; depth ofcranium, 7.80. Holotype. - Young adult male, ~kn and Distribution. - Known only from the type skull, USNM 80128, from Wild Horse locality. Creek, 4 mi NW Alvord Lake, Harre)' Co., Remarks. - From polionotus, the only Oregon; obtained on 18 August 1896 by subspecies in contact with nasutus, nasutus Clark P. Streator. differs in having a longer body and shorter Measurements ofholotype. - T otal;~rtgth, tail, longer hind foot, and skull that is 153; length of tail, 88; length of hind foot, broader across the auditory bullae. Hall 24; greatest length ofskull, 27.8; widtb across (1941) listed means and ranges ofmeasure­ bullae, 18.1; length of nasals, 9.6; mteror­ ments for 4 adults. bital breadth, 6.6; length ofmaxillary tooth­ row, 3.5; depth of cranium, 7.8; length of rostrum, 11.0. Microdipodops megacephalus nexus Distribution. - Extends from southeast­ Hall, 1941 ern Oregon to Modoc and Lassen counties, in northeastern California, and Wasl:oe and 1941. Microdipodops megacephalus nexus Hall, Humboldt counties in northwestern Neva­ Field Mus. Nat. Hist., ZooI. Ser., 27:257, 8 da. December. Remarks.-Hall (1941) noted~ntegrada­ Holotype. -Adult male, skin and skull, tion between ambiguus and oregonus m in­ MVZ 7091 7, from 3 mi S Izenhood, Lander dividuals from Smoke Creek, WlshQ(' Co., 96 WILLIAMS ET AL.

Nevada, and the vicinity ofDenio, Oregon. S Benton Station, 5,200 ft, Mono Co., Cal­ From M. m. ambiguus, oregonus differs in ifornia; obtained on 10 July 1912 by Charles having grayer upper parts and a narrower D. Holliger. cranium. Hall (1941) listed means and rang­ Measurements ofholotype. - Total length, es of measurements for 13 adults and Haf­ 145; length of tail, 80; length ofhind foot, ner (1985) listed statistics for 11 individu­ 24; length of ear, 9; width across bullae, als. 18.35; breadth across maxillary arches, 12.10; interorbital breadth, 7.00; length of maxillary toothrow, 3.45. Microdipodops megacephalus paululus Distribution. - Found in the Mono Lake Hall and Durrant, 1941 Basin and head ofOwens Valley, Mono Co., California. 1941. Microdipodops megacephalus paululus Remarks. - The coloration of the upper Hall and Durrant, The Murre1et, 22:5, 30 April. parts of individuals ofpo!ionotus are quite variable (Hall, 1941). This taxon differs from Holotype. - Subadult male, skin and skull, other subspecies occurring along the Cali­ MVZ 74660, from Pine Valley, 0.5 mi E fornia-Nevada border in having less black headquarters building of the Desert Range on the dorsal surface of the distal portion Experiment Station, U.S. Forest Service, of the tail, although some black is present. T25S, R17W, Sec. 33, Salt Lake B. M., Mil­ Hall (1941) listed means and ranges ofmea­ lar Co., Utah; obtained on 17 July 1936 by surements for 20 adults. Stephen D. Durrant. Measurements ofholotype. - Total length, 147; length of tail, 76; length of hind foot, Microdipodops megacephalus sabulonis 24; length ofear, 11; greatest length ofskull, Hall, 1941 27.65; width across bullae, 18.30; length of nasals, 9.40; breadth across maxillary arch­ 1941. Microdipodops megacephalus sabulonis es, 11.45; interorbital breadth, 6.30; length Hall, Proc. BioI. Soc. Washington, 54:59, 20 of maxillary toothrow, 3.25; depth of cra­ May. nium, 7.70. Distribution. - Occurs in west-central Holotype. - Adult male, skin and skull, Utah in the Pine, White, and Snake valleys. MVZ 49381, from 5 mi SE Kawich P.O., Remarks. - Hall (1941) only provision­ 5,400 ft, Kawich Valley, Nye Co., Nevada; ally assigned specimens from Snake and obtained on 27 September 1931 by Robert White valleys to this subspecies because only T. Orr. two young specimens and one adult were Measurements ofholotype. - Total length, available from these areas, respectively. Hall 155; length of tail, 83; length of hind foot, (1981), however, maintained this arrange­ 25; length ofear, 10; weight, 11.3 g; greatest ment. Hall (1941) gave means and ranges length of skull, 27.90; width across bullae, of measurements for 9 adult topotypes. 19.05; length ofnasals, 9.85; breadth across maxillary arches, 11.35; interorbital breadth, 6.40; length of maxillary toothrow, 3.15; depth ofcranium, 7.80. Microdipodops megacephalus po!ionotus Distribution. -Occurs in south-central Grinnell, 1914 Nevada. Remarks. - The auditory bullae of spec­ 1914. Microdipodops polionotus Grinnell, Univ. imens ofsabulonis are proportionately much California Publ. Zool., 12:302, 15 April. inflated so that their greatest width exceeds Holotype. -Adult male, skin and skull, the basal length. Hall (1941) gave means MVZ 17031, from McKeever's Ranch, 2 mi and ranges of measurements for 9 adult TAXONOMY 97 topotypes. Hafner et al. (1979) listed means bullae, 19.00; length ofnasals, 9.75; breadth ± SD for 14 specimens from Penoyer Val­ across maxillary arches, 11.80; interorbital ley. breadth, 6.90; length ofmaxillary toothrow, 3.50; depth of cranium, 8.30. Distribution. -Occursin Railroad Valley, Microdipodops pallidus Nye Co., Nevada. Remarks. - This subspecies has the pale Diagnosis. -Anterior palatine foramina dorsal ground color of the species, but this parallel-sided; premaxillae extending more is overlaid with a frosting of black. Speci­ than 1-2 mm posterior to the posterior bor­ mens from east ofNew Revielle show slight der ofthe nasals; auditory bullae greatly in­ reddish tint ofM. p. ruficollaris. Hall (1941) flated; upper incisors relatively straight; listed means and ranges of measurements length ofhindfoot averages 25-27 mm; post­ for 12 adult topotypes. auricular spot pure white; upper parts near Light Pinkish Cinnamon (Ridgway, 1912) with a light overwash of buffy or blackish; Microdipodops pallidus pallidus tail about same color on dorsal side as the Merriam, 190 I dorsal body parts; tail lacks black tip; hairs of underparts white to their bases; diploid 1901. Microdipodops pallidus Merriam, Proc. chromosome number 42. BioI. Soc. Washington, 14:127, 19 July. Comparisons. -See account for M. 1926. Microdipodops megacephalus lucidus megacephalus. Goldman, Proc. BioI. Soc. Washington, 39:127, Distribution. - Found almost exclusively 27 December. on fine sands supporting some vegetation in 1927. Microdipodops megacephalus dickeyi the Great Basin region of west-central Ne­ Goldman, Proc. BioI. Soc. Washington, 40: 115, vada, extreme eastern Mono Co., California, 26 September. and as a disjunct population in Deep Spring Holotype. - Adult female, skin and skull, Valley, Inyo Co., California. USNM 93520, from Mountain Well [fide Remarks. - Pale kangaroo mice are much Vernon Bailey in litt.-Hall, 1941], Chur­ more restricted to fine, loose sands. Rela­ chill Co., Nevada; obtained on 11 May 1898 tionships with M. megacephalus are dis­ by Harry C. Oberholser. cussed in the account of that species. Measurements ofholotype. - Total length, 171; length oftail, 102; length ofhind foot, 25.5; greatest length of skull, 29.4; width Microdipodops pallidus ammophilus across bullae, 19.5; length of nasals, 9.0; Hall, 1941 breadth across maxillary arches, 12.2; in­ terorbital breadth, 6.7; length of maxillary 1941. Microdipodops pallidus ammophilus Hall, toothrow, 3.7; depth ofcranium, 8.1; length Field Mus. Nat. Hist., ZooI. Ser., 27:273, 8 of rostrum 11.6. December. Distribution. -Found in the lower, west­ Holotype. -Adult female, skin and skull, ern part of Nevada, Pershing Co., through MVZ 58208, from Railroad Valley, Able Fish Lake Valley to Oasis, and in the Deep Spring, 12.5 mi S Locks Ranch, 5,000 ft, Spring Valley, Inyo Co., California. Nye Co., Nevada; obtained on 29 July 1933 Remarks. - Integradation between rufi­ by E. Raymond Hall. collaris and pallidus is suggested by speci­ Measurements ofholotype. - Total length, mens from west ofMillers Wells, Esmeralda 162; length of tail, 90; length of hind foot, Co., Nevada. The specimens from Deep 26.2; length of ear, 11.0; weight, 14.6 g; Spring Valley are isolated from other pop­ greatest length ofskull, 28.55; width across ulations ofthe species, but are very similar 98 WILLIAMS ET AL. to specimens from Fish Lake Valley, Ne­ 25.5; length ofear, 12; weight, 9.5 g; greatest vada. Hall (1941) listed means and ranges length of skull, 28.01; width across bullae, of measurements for 20 adult topotypes; 19.53; length ofnasals, 10.14; breadth across Hafner (1985) gave statistics for measure­ maxillary arches, 12.49; interorbital breadth, ments of 14 individuals. 6.59; greatest length of incisive foramina, 2.23; length of maxillary toothrow, 3.35; depth of cranium, 8.15. Microdipodops pallidus purus Distribution. - Known only from the type Hall, 1941 locality, which is located atthe southern end ofRhodes Salt Marsh in Soda Spring Valley. 1941. Microdipodops pallidus purus Hall, Field Remarks. -Although restrictus is located Mus. Nat. Hist., Zooi. Ser., 27:273, 8 Decem­ in the western part of the range of the spe­ ber. cies, it is similar structurally to ruficollaris Holotype. -Adult male, skin and skull, to the east. Yet, restrictus differs karyotyp­ MVZ 52753, from 14.5 mi S Groom Baldy, ically from the eastern subspecies and is Lincoln Co., Nevada; obtained on 1 June similar to the taxa to the west (Hafner, 1985). 1932 by H. Robert Poultney. Hafner (1985) gave statistics for measure­ Measurements ofholotype. - Total length, ments of seven individuals. 160; length of tail, 88; length of hind foot, 26; length ofear, 10; weight, 12.9 g; greatest length of skull, 28.85; width across bullae, Microdipodops pallidus ruficollaris 19.85; length ofnasals, 10.00; breadth across Hall, 1941 maxillary arches, 12.45; interorbital breadth, 6.75; length of maxillary toothrow, 3.50; 1941. Microdipodos [misspelling ofMicrodipo­ depth of cranium, 7.90. dops]pallidus ruficollaris. Hall, Proc. BioI. Soc. Distribution. -Occursin the Emigrant and Washington, 54:60, 20 May. Desert valleys, Lincoln Co., Nevada. Holotype. -Adult female, skin and skull, Remarks. - Specimens from the type lo­ MVZ 49254, from 5 mi SE Kawich P.O., cality are the palest ofthe species. This tax­ 5,400 ft, Kawich Valley, Nye Co., Nevada; on also has a wide skull at the maxillary obtained on 25 September 1931 by Robert arches and a narrow interorbital region. The T. Orr. range ofpurus may not be disjunct from that Measurements ofholotype. - Total length, of ruficollaris, whiCll occurs close by to the 160; length of tail, 90; length of hind foot, west. Hall (1941) Lsted means and ranges 25; length ofear, 9; weight, 12.0 g; greatest of measurements for 20 adult topotypes. length of skull, 28.35; width across bullae, 19.35; length ofnasals, 9.85; breadth across maxillary arches, 12.15; interorbital breadth, Microdipodops pallidus restrictus 7.00; length of maxillary toothrow, 3.45; Hafner, 1985 depth ofcranium, 7.90. Distribution. - Ranges from the valleys of 1985. Microdipodops pale/dus restrictus Hafner, western Nye Co. eastward to western Lin­ Proc. 13;01. Soc. Wa"hir:gton, 98:6, 20 March. coln Co., Nevada. Holotype - Adult male, skin, skull, and Remarks. - This taxon is characterized by partial skeleton, MVZ 159970, from 8.9 mi reddish upper parts and a broad cinnamon­ S, I.? mi EMina, 4,400 ft, Mineral Co., colored collar. Hall (1941) found little vari­ Nevada; obtained on 2 August 1979 by John ation within this subspecies except for the C. Hafner. population in the Penoyer Valley. Hall Measurements ofholotype. - Total length, (1941) gave means and ranges of measure­ 158; lenfth of tail, 89; length of hind foot, ments for 10 adult topotypes. Hafner et al. TAXONOMY 99

(1979) gave means ± SD for 35 individuals the "central" ofcaudal vertebrae; two pter­ from the Penoyer Valley; and Hafner (1985) ygoid fossae present; auditory bullae mod­ gave statistics for measurements of 8 indi­ erately expanded; inflated interior of audi­ viduals. tory bullae filled with spongy trabeculae; mastoid bulla not appearing on dorsal sur­ face of skull and not projecting to postero­ Subfamily Heteromyinae Coues, 1875 dorsal plane ofocciput; lateral wall ofskull dorsal to external auditory meatus com­ Diagnosis. -Size medium to large, from posed entirely of squamosal; ventral plane 180 to 360 mm in total length; body form oftympanic bulla higher than occlusal plane quadrupedal and generally mouse- or rat­ ofcheek teeth; no stapedius muscle in mid­ like; tail not tufted or crested; locomotion dle ear; no fusion or broadening ofcervical scansorial with slight tendency to subrico­ chetal; hind limbs larger than forelimbs; pes vertebrae; scapula not prolonged posteri­ with five clawed digits; pes soles naked or orly; obturator foramen not triangular; no partly clothed with a scant covering ofshort articulation of trapezoid and scapholunar; hairs; pelage stiff or hispid; body hairs of astragalus articulating with cuboid; dorsal three types-straight, relatively long and surface of ectocuneiform not hourglass­ widened overhairs, wide, troughed over­ shaped; phallus without external spines and hairs, and a thin underfur of slightly wavy with 2-3 lobed urethral lappets; phallus long hairs; cuticular scales in trough ofhairs form relative to bacular length; baculum with a series oflongitudinal ridges; anterior face of swollen base and an upturned tip; male ac­ upper incisor either smooth or with shallow cessory reproductive glands variable-pre­ groove; molars progressively hypsodont, putial gland absent, ventromedial prostate rooted, and tuberculate; enamel cusp pat­ present or absent, anterior and dorsal pros­ tern ofcheek teeth lost to wear early in life; tates present or absent, ampullary present lophs of P4 unite first at lingual then labial or absent; vesicular glands elongate, hook­ side; protoloph of P4 with more than one shaped, and translucent; nasolabialis pro­ cusp; lophs of upper molars unite first at fundus pars maxillaris muscles originate lingual then labial side, surrounding a cen­ from infraorbital foramen; M. temporalis tral basin in a majority of species (most not reduced, origin far lateral, narrow, and prominent in M1); lophs ofp4 unite first at reaching back ofsquamosal; M. cleidomas­ lingual then labial side; stylids on any point toideus present; extensores breves muscles of p4, developing progressively in geologic present; M. abductor hallucis present; time; lophs of lower molars united first at lumbricales muscles not reduced or absent labial then lingual side; no foramina and (Burt, 1936; Hafner and Hafner, 1983; Gen­ usually no pit between m3 and the base of oways, 1973; Homan and Genoways, 1978; the coronoid process; palate extending be­ Merriam, 1889; Osgood, 1900; Ryan, 1989; yond level of M3; center ofpalate between Wahlert, 1985; Webster and Webster, 1975; cheek teeth not ridged; squamosal in broad Wood, 1935). contact with parietal on dorsal surface of Remarks. - Heteromyines are the most skull; squamosal not perforated by auditory murinelike members ofthe family, and are, bulla; anterior zygomatic root not 3reatly in most ways, more similar structurally to enlarged on joining lachrymal; ossification the ancestral heteromyids and geomyoids of orbital walls complete; incisive foramen than other living heteromyids. Of the two small; interorbital foramen preSeh'1t in or­ genera, Liomys shows more development bitosphenoid bone ventral to optic fora­ of subricochetal tendencies than Hetero­ men; masticatory and buccinator foramina mys. Genetic studies by Rogers (1990) sug­ united; stapedial and sphenofrontal foram­ gest that Heteromys and Liomys should be ina absent; no median ventral foramina in combined into one genus, perhaps contain- 100 WILLIAMS ET AL. ing three to five, or more, subgenera. One contributed much to elucidate the relation­ group would include species classified here ships of species within this genus. Herein, as Liomys, H. australis and an undescribed two subgenera are recognized, but their va­ species of Heteromys. lidity and relationships are questionable (see below). The subgenus Heteromys contains five species, whereas Xylomys is monotypic. Key to the Genera Additionally, Rogers (1990) showed the ex­ istence in Costa Rica ofan undescribed spe­ 1. Cheek teeth with medium-high crowns; H. lower premolar with 210phids; upper and cies related to anomalus; biochemically, lower premolars wider than last upper these two taxa are more closely linked with and lower molars; length of P4/greatest Liomys than with other species of Hetero­ length ofskull averages greater than 0.046; mys. The following key was adapted from posterior portion ofsoles ofhind feet with Schmidt et ai. (1989). We know couplet 4 sparse covering of hairs Liomys to be unreliable, but have not found key I'. Cheek teeth with high crowns; lower pre­ characters to distinguish all populations of molar with 3 or 410phids; upper and low- these species. er premolars wider than last upper and lower molars; length ofP4/greatest length of skull averages less than 0.043; poste­ Key to the Subgenera and Species rior portion of soles of hind feet usually naked (except H. gaumeri) .... Heteromys 1. Adult pelage harsh but bristles soft ... .. 2 I'. Adult pelage with numerous stiffbristles or spines Subgenus Heteromys 3 Genus Heteromys 2. Ears edged with white; premaxillary bones terminating posterior to nasals; known 1817. Heteromys Desmarest, Nouv. Diet. Hist. only from Costa Rica . Nat., 14:181, May. · Heteromys (Heteromys) oresterus 2'. Ears without white edges; premaxillary Type species. -Mus anomalus Thomp­ and nasals bones terminating at same lev- son, 1815, Trans. Linn. Soc. London, 11: el; known only from southeastern Chia­ 161, pI. 10. pas, Mexico and southwestern Guate­ Diagnosis. -Cheek teeth high crowned, mala ..... ".Heteromys (Xylomys) nelsoni with most complicated pattern in family; 3. Soles of hind feet hairy from posterior anterior cingulum in lower molars and a tubercle to heel; orange buff lateral line posterior cingulum in the upper molars broad and conspicuous . nearly as high as remainder ofcrown giving · '" " Heteromys gaumeri them three lophs before wear; upper inci­ 3'. Soles of hind feet naked posteriorly; lat­ sors asulcate; three or four lophids on lower eral line absent or when present never premolar; skull elongate; auditory region broad and conspicuous 4 4. Inner side offorearm clouded with dusky 5 uninflated; pelage hispid, consisting of stiff 4'. Inner side offorearm white . spines mingled with slender soft hairs; spines · Heteromys desmarestianus are flattened and anteriorly grooved; tail 5. Tail usually less than 140; slender hairs usually longer than head and body length; on dorsum clear grayish . soles of hind feet naked, except in H. gau­ · Heteromys australis meri; second digit ofhind foot with weakly­ 5'. Tail more than 140; slender hairs on dor- developed spoon-like claw; interpterygoid sum dull ochraceous buffy . fossa V-shaped anteriorly; adapted for · Heteromys anomalus scampering. Remarks. - The genus Heteromys was last Subgenus Heteromys reviewed by Goldman (1911). The research of Rogers and Schmidly (1982), Engstrom 1817. Heteromys Desmarest, Nouv. Diet. Hist. et ai. (1987), and Rogers (1989, 1990) has Nat., 14:181, May. TAXONOMY 101

Type species. -Mus anomalus Thomp­ Comparisons. - The range of this species son, 1815, Trans. Linn. Soc. London, 11: may contact that ofH. australis and H. des­ 161, pI. 10. marestianus. From both species, anomalus Diagnosis. - Pelage with numerous stiff may be distinguished by its larger ears, nar­ bristles or spines; inner sides of forelimbs rower interorbital region, and broader na­ white or slightly dusky; braincase more or sals. H. anomalus is larger than australis in less flat; parietals not extending laterally to head and body size. Compared with H. des­ the mastoids; posterior molars narrower marestianus, anomalus has dusky, instead than premolars; molars not complex but of white, inner forelimbs. possessing accessory enamel islands in pos­ Distribution. - Extreme eastern Panama, terior loops of upper molars and anterior western and northern Colombia, northern loops oflower molars; small posterior cin­ Venezuela, Trinidad, Tobago, and Marga­ gulum, which is closely compressed with rita Island. metaloph, present on last upper molar. Remarks. - Best (1992) gave external and Remarks. -Members ofthe subgenus are cranial measurements for this species. Rog­ widespread, covering the entire known dis­ ers (1990) indicated that this species oc­ tribution ofthe genus. This subgenus is cur­ curred in eastern Panama, near a locality rently recognized as containing five species, where H. australis was collected, but at a although Rogers (1990) stated that the cur­ slightly higher elevation. This specimen, rent taxonomy is incongruent with bio­ originally identified as H. australis, was gen­ chemical and chromosomal data. Goldman ically and morphologically most similar to (1911) and Hall (1981) divided this sub­ H. anomalus from eastern Venezuela (Rog­ genus into the anomalus species group con­ ers, 1990). This species also was found by taining anomalus and australis, and the des­ Rogers (1990) to be the sister taxon to sev­ marestianus species group which consisted eral taxa including Liomys, H. australis, and ofdesmarestianus, goldmani, and gaumeri. an undescribed species of Heteromys from Removal of gaumeri from the desmaresti­ Costa Rica. Engstrom et ai. (1987) reported anus species group was advocated by Eng­ no karyotypic variation for 20 specimens strom et ai. (1987) based on morphology from Venezuela; based on distribution these and karyology. Rogers (1990) advocated specimens would be referred to H. a. anom­ splitting the anomalus species group and in­ alus. Based on an examination by H. Gen­ cluding oresterus within the desmarestianus oways, one specimen from Palomino, Co­ species group of the subgenus Heteromys. lombia and two unregistered specimens from Bogota, Colombia in the British Museum Species Accounts (Natural History), are grayer and appear to have more spines than three additional Heteromys anomalus specimens from near Bogota that are ten­ tatively assigned H. australis. The former Diagnosis. -Size large for species of ge­ specimens are assigned to H. anomalus, al­ nus; adult head and body length generally though they might be desmarestianus. between 130 and 140 mm; tail longer than head and body length; tail dark above, light­ Heteromys anomalus anomalus er below, distinctly bicolored; dorsum dark (Thompson, 1815) mouse gray to grayish brown with a sprin­ kling ofochraceous hairs; venter white; lat­ 1815. Mus anomalus Thompson, Trans. Linn. eralline absent; inner forelimbs clouded with Soc. London, 11:161, pI. 10. dusky (except in H. anomalus jesupi); ears 1827. Heteromys thompsonii Lesson, Manuel de large; skull large; narrow interorbital region; Mammalogie, p. 264. bullae small; nasals broad; 2n = 60, FN = 1868. Perognathus bicolor Gray, Proc. Zool. Soc. 68. London, p. 202, May. 102 WILLIAMS ET AL.

1868. Heteromys melanoleucusGray, Proc. Zool. Holotype. - Adult female, skin and skull, Soc. London, p. 203, May. FMNH 18623, from El Panorama, Rio Au­ Holotype. -Adult ofunknown sex, poor­ rare, eastern shore ofLake Mamcaibo, Ven­ ly preserved skin and skull, BM(NH) un­ ezuela; obtained on 19 January 1911 by registered, from Trinidad. Wilfred H. Osgood and Stanley G. Jewett. Measurements ofholotype. - Interorbital Measurements ofholotype. - Total length, breadth, 8.9; length of nasals, 15.7; length 284; length oftail, 150; length ofhind foot, ofrostrum, 16.8; length ofmaxillary tooth­ 34; greatest length ofskull, 3::i.2; zygomatic row, 5.6. breadth, 16.1; interorbital breadth, 7.3; Distribution. -Northern and eastern length of nasals, 14.2; width of braiilcase, Venezuela and the islands of Trinidad and 14.8; length ofmaxillary toothrow, 5.6 (Os­ Margarita. good, 1912). Remarks. - We believe the holotype of Distribution. -Confined to the area east Mus anomalus and the cotypes of Perog­ of Lake Maracaibo. nathus bicolor and Heteromys melanoleucus Remarks. - Osgood (1912) described H. are the same taxon; the senior synonym is a. brachialis primarily because its upper ;nemb~rs Heteromys anomalus. The lectotype parts are paler than in other ofthe (BM[NH] 47.2.1.7) and lectoparatype species and because the front legs are nearly (BM[NH] 47.2.1.5) of P. bicolor are both white. The skull of brachialis was noted as juveniles with silky juvenile pelage and de­ being practically the same as those ofother ciduous premolars. The lectotype (BM[NH] populations ofH. anomalus. The :ype and 47.2.1.4) and lectoparatype (BM[NH] five other specimens were collected under 47.2.1.6) are both subadults with juvenile mayas and thorn shrubs that formed hedge­ pelage except on the head, neck, and rump like borders ofthe trails leading into El Pan­ and permanent premolars. Genoways's ex­ orama (Osgood, 1912). Specimens from the amination of the holotype indicates that region west of Lake Maracaibo (Handley, Gray apparently described juveniles from 1976) may be assignable to this subspecies. this series as P. bicolor and subadult and adult material as H. melanoleucus. Gray de­ scribed Liomys irroratus and L. albolim­ Heteromys anomalus hershkovitzi batus on a similar basis (Genoways, 1973: Hernandez-Camacho, 1956 111-112). These four specimens were all collected by Dyson; Gray described them as 1956. Heteromys (Heteromys) anolnalus hersh­ coming from Honduras but the specimen kovitzi Hernandez-Camacho, Lozania, 10:3, tags and the Museum register (Alston, 1879­ 26 April. 1882) indicate that they are from Venezue­ la. A note on the tag states "described and Holotype. -Adult female, lnstituto Car­ stand painted as from Honduras." External los Finlay, Bogota no. 2701, but now de­ measurements for this subspecies (including posited in the Instituto de Ciencias Natura­ melanoleucus) were given by Allen (1899). les, Bogota, from (according to Cabrera, Goldman (1911) gave measurements for five 1961:512) "volcanes, cerca C:'e la cabecera adult specimens from Trinidad. del corregimiento c,e Cordoba, municipio de Caparrapi, departmento de Cundina­ marca," 250 m; obtained 18 F~bruary 1944. Heteromys anomalus brachialis Measurements of holotype. -No mea­ Osgood, 1912 surements of the holotype are available. Distribution. - Known with certainty only 191~. Heteromys anomalus brachialis Osgood, from the type locality. Cabrera (1961) in­ Field Mus. Nat. Hist., Zool. Ser., 10 (5):54, 10 dicated that this subspecies is distrlbuted in January. the Magdalena River Valley ufthe Andean TAXONOMY 103 region of Colombia at least on the forested dusky, while the inner side of the forearm western slopes of the Eastern Cordillera. in H. desmarestianus is white. The dorsal pelage ofH. desmarestianus and H. anoma­ Heteromys anomalus jesupi Ius is usually sprinkled with ochraceous J. A. Allen, 1899 hairs; in H. australis these hairs are gray. The braincase of H. australis is inflated; it 1899. Heteromysjesupi J. A. Allen, Bull. Amer. is not inflated in H. anomalus and H. des­ Mus. Nat. Rist., 12:201,20 December. marestianus. Distribution. -H. australis is distributed Holotype.-Adu1t female, AMNH 15347, from eastern Panamainto northwestern Co­ from below Minca, 1,000 ft, Santa Marta lombia southward to northwestern Ecua­ District, Colombia; obtained on 30 July dor. It is also distributed eastward in Co­ 1899 by Herbert H. Smith. lombia to near Bogota. Measurements ofholotype. - Tota11ength, Remarks. - The karyotype for this spe­ 330; length oftail, 163; length ofhind foot, cies has not been reported. External and cra­ 33; greatest length ofskull, 36.2; zygomatic nial measurements were given by Best breadth, 16.4; interorbital constriction, 8.4; (1992). Rogers (1990) found that, genically, mastoid breadth, 5.3; length ofnasals, 15.0; H. australis was not closely related to H. length ofrostrum, 15.7; length ofmaxillary anomalus, but instead was more related to toothrow, 5.5. an undescribed species from Costa Rica. All Distribution. -Northern Colombia, the three taxa were more closely related to Lio­ mountain slopes of the Sierra Nevada de mys than to other species of Heteromys Santa Marta, sea level to 610 m. (Rogers, 1990). Goldman (1911), later fol­ Remarks. - External measurements for lowed by Hall (1981), placed australis and females ofthis taxon are greater than those anomalus in the anomalus species group. for females of either H. a. anomalus or H. Although the distribution of H. australis a. brachialis (Allen, 1899). Measurements contacts that of H. desmarestianus crassi­ for males are not reported. Osgood (1912) rostris in eastern Panama, H. australis is used the trinomial H. a. jesupi in comparing found at lower elevations. H. australis may this taxon tc H. (.10 brachialis. come into contactwith H. anomalus in Pan­ am?, and north-central Colombia. Heteromys australis Thomas, 1901 Heteromys australis australis Diagnosis. ~-Medium-sized species for Thomas, 1901 genus; head and body length less than 130 mm; tail equal to or slightly longer than 1901. Heteromys australis Thomas, Ann. Mag. head and body length; tail brownish above, Nat. Rist., ser. 7, 7: 194, February. lighter below, but not distinctly bicolored; 1912. Heteromys lomitensis J. A. Allen, Bull. Amer. Mus. Nat. Rist., 31:77,19 April. dorsum gray to blackish slate, grizzled with gray; venter white; inner sides offorelimbs Holotype. -Female, skin and skull, dusky; lateral line absent; ears without a BM(NH) 1.3.19.23, from St. Javier, 60 ft, white edging; skull short, broad; braincase northwestern Ecuador; obtained on 23 June inflated; auditory bullae small; zygoma an­ 1900 by G. Flemming and R. Miketta. teriorly spreading. Measurements ofholotype. - Total length, Comparisons. - This species can be dis­ 272; length oftail, 137; length ofhind foot, tinguished from H. anomalus and H. des­ 30; length ofear, 15; greatest length ofskull, marestian~s by its smaller size and shorter 34.5; zygomatic breadth, 16.0; interorbital tail (less than 140 mm); both of the above breadth, 9.2; mastoid breadth, 14.0; length have a taii length greater than 140 mm. In of nasals, 13.8; length of rostrum, 15.4; H. australis the inner side ofthe forearm is length of maxillary toothrow, 5.3. 104 WILLIAMS ET AL.

Distribution. - This subspecies is distrib­ Holotype. -Adult male, skin and skull, uted from northwestern Colombia south­ ANSP 19499, from Amagal, 1,000 ft, S of ward into northwestern Ecuador and east­ Guayabo Bay, Darien, Panama; obtained ward in Colombia to near Bogota. Elevations on 10 June 1938 by Oliver P. Pearson. range from 20 m to 2,750 m. The localities Measurements ofholotype. - Total length, in Colombia are intercordilleran. 275; length oftail, 144; length ofhind foot, Remarks. -Four specimens at the British 31; greatest length ofskull, 35.0; zygomatic Museum (Natural History) from localities breadth, 16.8; interorbital breadth, 9.0; near Bogota, Colombia may represent aus­ length of nasals, 14.4; length of maxillary tralis. Allen (1916) placed lomitensis in syn­ toothrow, 5.4 (Pearson, 1939). onymy with australis, noting that the dif­ Distribution. - Known only from the area ferences in pelage and coloration upon which near the type locality at elevations ranging the description oflomitensis were based were from 300 m to 610 m (Handley, 1966). probably accidental or ofa seasonal nature. Pearson (1939) noted that no specimens Goldman (1911) gave measurements for were collected near sea level. specimens from the type locality. Remarks. -According to Pearson (1939: 5), the nine specimens from the type locality Heteromys australis conscius were captured at elevations ranging from Goldman, 1913 300 m to 610 m in a "habitat ... charac­ terized by an enormous amount of rainfall 1913. Heteromys australis conscius Goldman, and a luxurious growth ofmany palms, con­ Smithsonian Misc. Co1l., 60(22):8, 28 Feb­ stituting an infinitely richer vegetation than ruary. found near Cana, the type locality of con­ scius." Pearson (1939) also gave averages Holotype. -Adult male, skin and skull, for external and cranial measurements for USNM 178699, from Cana, 2,000 ft; moun­ three adult female topotypes. tains of eastern Panama; obtained on 8 March 1912 by Edward A. Goldman. Measurements ofholotype. - Total length, Heteromys desmarestianus 260; length oftail, 133; length ofhind foot, 32; greatest length ofskull, 34.3; zygomatic Diagnosis. - Large-sized species for ge­ breadth, 16.0; interorbital breadth, 8.0; nus; head and body length generally be­ mastoid breadth, 13.8; length ofnasals, 13.7; tween 130 and 140 mm; tail longer than length ofrostrum, 15.2; length ofmaxillary head and body length; ears without white toothrow, 5.5; depth ofcranium, 8.7. edging; lateral line occasionally present, but Distribution. - Known from extreme never pronounced; dorsum gray to slaty eastern Panama and perhaps into north­ black with sprinkling of ochraceous hairs; western Colombia at elevations ranging from venter white; inner sides offorelimbs white; 150 m to 800 m. skull medium to large size; bullae small; in­ Remarks. -External measurements for terparietal variable; zygoma narrow or only two adult topotypes were given by Goldman slightly spreading anteriorly; base of bacu­ (1913). Specimens have been collected un­ lum one-third total bacular length; baculum der logs in the forest at low elevation slopes tapers to the slightly upturned and slightly of the Pirre Range (Goldman, 1920). laterally compressed tip; 2n = 60, FN = 67­ 86. Heteromys australis pacificus Comparisons. - This species may be dis­ Pearson, 1939 tinguished from H. nelsoni and H. oresterus by smaller body size, smaller ears, and by 1939. Heteromys australis pacificus Pearson, a greater development ofthe posterior por­ Notu1ae Naturae, Acad. Nat. Sci. Philadel­ tion ofthe toothrow. In addition, specimens phia, 6:4, 8 June. ofH. nelsoni and H. oresterus have soft, not TAXONOMY 105 stiff, bristles. Compared with H. gaumeri, Measurements ofholotype. - Total length, desmarestianus is larger and lacks the furred 302; length oftail, 154; length ofhind foot, soles ofthe hind feet; H. australis is smaller 35; length ofear, 19; greatest length ofskull, and the ochraceous hairs on the dorsum are 37.6; zygomatic breadth, 17.2; interorbital lacking. The ears ofH. anomalus are much breadth, 10.1; length of nasals, 17.0 (En­ larger than those of desmarestianus. ders, 1938). Distribution. - This species is distributed Distribution. - Southeastern Costa Rica southward from the southern Mexican andadjacent Chiriqui Province in west-cen­ province of Veracruz, including the south­ tral Panama at elevations from 1,150 m to ern Yucatan Peninsula, Belize, Guatemala, 1,600 m. Honduras, El Salvador, Costa Rica, Pana­ ma, and into northwestern Colombia. Remarks. - This is the most widely dis­ Heteromys desmarestianus crassirostris tributed species ofthe genus. Rogers (1989) Goldman, 1912 described standard karyotypes and used C-banding to evaluate the chromosomal 1912. Heteromys crassirostris Goldman, Smith­ variation within this species. Seven chro­ sonian Misc. CoIl., 60(2): 10, 20 September. mosomal forms are known for this species, each ofwhich exhibits no interpopulational Holotype. - Adult male, skin and skull, variation (Rogers, 1989). Mascarello and USNM 179016, from Mount Pirri, 5,000 ft, Rogers (1988) described G-banded chro­ near head ofRio Limon, Panama; obtained mosomes. Comparisons ofthe external, cra­ on 26 April 1912 by Edward A. Goldman. nial, and bacular morphology ofthis species Measurements ofholotype. - Total length, with Liomys were made by Genoways 267; length oftail, 132; length ofhind foot, (1973). Rogers and Schmidly (1982) placed 32.5; greatest length of skull, 33.0; zygo­ lepturus and longicaudatus in synonymy matic breadth, 15.8; interorbital constric­ with H. d. desmarestianus; H. temporalis tion, 9.4; mastoid breadth, 13.5; length of was recognized as a subspecies of desma­ nasals, 13.6; length ofrostrum, 15.1; length restianus. Goodwin (1969) treated nigricau­ of maxillary toothrow, 4.8; depth of crani­ datus as another synonym of H. d. desma­ um,8.9. restianus. Rogers (1990) found that H. Distribution. -Densely forested upper goldmani was conspecific with H. desma­ slopes of extreme eastern Panama into restianus, but did not remark on its sub­ northwestern Colombia at elevations from specific status. Herein we recognize it as a 1,350 m to 1,600 m. subspecies. Best (1992) gave external and Remarks. - This is the only subspecies of cranial measurements. desmarestianus with a distribution that ex­ tends into South America. Goldman (1920) placed this taxon as a subspecies of des­ Heteromys desmarestianus chiriquensis marestianus. External measurements for Enders, 1938 eight adult topotypes and cranial measure­ ments for five adults were given by Gold­ 1938. Heteromys desmarestianus chiriquensis man (1912). Enders, Proc. Acad. Nat. Sci. Philadelphia, 90: 141,20 September.

Holotype. -Adult male, skin and skull, Heteromys desmarestianus desmarestianus ANSP 17835, from Cerro Pando, a hill 4,000 Gray, 1868 ft in elevation situated between the Rio Chi­ riqui Viejo and its tributary, the Rio Col­ 1868. Heteromys desmarestianus Gray, Proc. orado, about 10 mi from the Post Office of Zool. Soc. London, p. 204, May. El Volcan, Chiriqui, Panama; obtained on 1868. Heteromys longicaudatus Gray, Proc. 20 August 1935 by Robert K. Enders. Zool. Soc. London, p. 204, May. 106 WILLIAMS ET AL.

1902. Heteromys griseus Merriam, Proc. BioI. anus. Measurements for three specimens are Soc. Washington, 15:42,5 March. given in Allen (1908). Distribution in north­ 1902. Heteromys goldmani lepturus Merriam, eastern Honduras was established by Ben­ Proc. BioI. Soc. Washington, 15:42,5 March. shoof et al. (1984). At this site, specimens 1928. Heteromys desmarestianus psakastus were collected in dry areas on the tops of Dickey, Proc. BioI. Soc. Washington, 41:10,1 hills in second growth and mature forest at February. elevations of 50 m and 100 m. These au­ 1956. Heteromys nigricaudatus Goodwin, Amer. Mus. Novit., 1791 :4, 28 September. thors also gave external measurements for three males and two females. Holotype. -Adult of unknown sex, skin and broken skull, BM(NH) 43.6.13.1 from Coban, Guatemala; obtained on an un­ Heteromys desmarestianus goldmani known date by an unknown collector. Merriam, 1902 Measurements of holotype. - Length of hind foot, 35; length of ear, 15; length of 1902. Heteromys goldmani Merriam, Proc. BioI. nasals 15.6; length of rostrum, 16.7; length Soc. Washington, 15:41,5 March. of maxillary toothrow, 5.3. Holotype. -Adult male, skin and skull, Distribution. - Humid forested mountain USNM 77576, from Chicharras, Chiapas; .;lopes and coastal plains ofsouthern Mexi­ obtained on 7 February 1896 by Edward W. co, Guatemala, Belize, and El Salvador. Nelson and Edward A. Goldman. Remarks. - The characteristics attributed Measurements ofholotype. - Total length, to H. longicaudatus are within the variation 347; length oftail, 199; length ofhind foot, (Ifseveral samples ofH. d. desmarestianus; the latter name was chosen by Rogers and 40; greatest length ofskull, 39.0; zygomatic Schmidly (1982) because both names were breadth, 17.4; interorbital breadth, 9.7; published simultaneously by Gray (1868). mastoid breadth, 16.0; length ofnasals, 15.8; .-\.lso see remarks in the species account length ofrostrum, 17.4; length ofmaxillary abovl~. toothrow, 5.6; depth of braincase, 10.0. Distribution. - Restricted to the heavily Heteromys desmarestianus fuscatus forested Pacific slope ofthe Sierra Madre in J. A. Allen, 1908 extreme southern Chiapas and adjacent Guatemala between 45 m and 1,860 m. 1908. Heteromys fuscatus J. A. Allen, Bull. Remarks. -Considerable morphologic Amer. Mus. Nat. Hist., 24:652, 13 October. geographic variation among three samples Holotype. -Adult male, skin and skull, from Chiapas, Mexico was found by Rogers \MNH 28451, from Tuma, Nicaragua; ob­ and Schmidly (1982). Based on morphol­ ,ained on 1 December 1907 by William B. ogy, Rogers and Schmidly (1982) recog­ ~ichardson. nized this taxon as a distinct species and Measurements ofholotype. - Total length, included H. goldmani in the H. desmares­ 300; length oftail, 150; length ofhind foot, tianus species group. Later, Rogers (1990) 30; greatest length ofskull, 36.4; zygomatic found that, based on genetic data, H. gold­ breadth, 16.9; interorbital constriction, 10.0; mani was indistinguishable from nearby mastoid breadth, 14.7; length ofnasals, 14.8; populations ofH. desmarestianus and pro­ length ofrostrum, 16.8; length ofmaxillary posed that H. goldmani be synonymized toothrow, 5.3. with H. desmarestianus. External and cra­ Distribution. -Southern, western, and nial measurements for this species were giv­ northeastern Honduras and central Nica­ en by Goldman (1911), Rogers and Schmid­ ragua. ly (1982), and Best (1992). Goldman (1951) Remarks. -Goldman (1920) recognized described collecting localities at two sites in this taxon as a subspecies of desmaresti- Chiapas. TAXONOMY 107

Heteromys desmarestianus panamensis Volcan de Chiriqui, 4,000 ft, Panama; ob­ Goldman, 1912 tained on 8 April 1901 by W. W. Brown, Jr. 1912. Heteromys panamensis Goldman, Smith­ Measurements ofholotype. - Total length, sonian Misc. ColI., 56(36):9, 19 February. 282; length oftail, 150; length ofhind foot, Holotype. - Adult male, skin and skull, 33; length ofear, 15; greatest length ofskull, USNM 171107, from Cerro Azul, 2,800 ft, 34.9; interorbital breadth, 9.4; mastoid near headwaters ofChagres River, Panama; breadth, 13.6; length ofnasals, 14.8; length obtained on 23 March 1911 by Edward A. ofrostrum, 15.6; length ofmaxillary tooth­ Goldman. row, 4.9. Measurements ofholotype. - Total length, Distribution. - Extreme southwestern 283; length oftail, 148; length ofhind foot, Panama. 35; greatest length ofskull, 34.4; zygomatic Remarks. - The type locality is at ap­ breadth, 16.9; interorbital constriction, 9.3; proximately 1,200 m. Goldman (1911) gave mastoid breadth, 14.8; length ofnasals, 13.9; external and cranial measurements for this length ofrostrum, 14.9; length ofmaxillary taxon. Later, Goldman (1920) recognized toothrow, 5.2; depth of cranium, 8.9. repens as a subspecies of desmarestianus. Distribution. - North-central Panama at elevations between 610 m and 910 m. Heteromys desmarestianus subaffinis Remarks. -Goldman (1920) placed this Goldman, 1937 taxon as a subspecies of desmarestianus. 1937. Heteromys desmarestianus subaffinis Heteromys desmarestianus planifrons Goldman, J. Washington Acad. Sci., 27(10): Goldman, 1937 420, 15 October. Holotype. - Adult male, skin and skull, 1937. Heteromys desmarestianus planifrons USNM 12904/38591, from Angostura, Goldman, J. Washington Acad. Sci., 27 (10): 418, 15 October. about 1,980 ft, southern side Rio Reven­ tazon, opposite Turrialba, Costa Rica; ob­ Holotype. -Adult female, skin and skull, tained in May 1876 by Jose C. Zeledon. USNM 250348, from San Geronimo, Pi­ Measurements of holotype. -Zygomatic rris, Costa Rica; obtained 12 April 1931 by breadth, 17.2; interorbital breadth, 10.0; C. F. Underwood. mastoid breadth, 14.5; length of maxillary Measurements ofholotype. - Total length, toothrow, 5.2. 303; length oftail, 169; length ofhind foot, Distribution. - Lowlands of northeastern 35; length ofear, 18; greatest length ofskull, Costa Rica. 37.8; interorbital breadth, 9.8; mastoid Remarks.-Goldman (1911) had earlier breadth, 15.7; length ofnasals, 16.2; length referred the specimens on which this sub­ ofrostrum, 17.8; length ofmaxillary tooth­ species is based to repens. This subspecies row, 5.6; depth of cranium, 9.3. appears to prefer relatively low elevations; Distribution. - Known only from the low­ the type locality is approximately 600 m. lands of western Costa Rica. The range ofthis subspecies is bordered by that of H. d. underwoodi; underwoodi is Heteromys desmarestianus repens smaller and is found at higher elevations. Bangs, 1902

1902. Heteromys repens Bangs, Bull. Mus. Heteromys desmarestianus temporalis Compo Zool., Harvard Univ., 39:45, April. Goldman, 1911

Holotype. -Adult female, skin and skull, 1911. Heteromys temporalis Goldman, N. Amer. MCZ 10356, from Boquete, southern slope Fauna, 34:26, 7 September. 108 WILLIAMS ET AL.

Holotype. -Adult female, skin and skull, and three females from Los Higuerones, Es­ USNM 63719, from Motzorongo, Vera­ cazu. He (Goodwin, 1943:2) noted that the cruz; obtained on 3 March 1894 by Edward type specimen was collected at an altitude W. Nelson and Edward A. Goldman. between 5,000 and 5,500 ft "in humid vir­ Measurements ofholotype. - Total length, gin country that is interspersed with patches 320; length oftail, 180; length ofhind foot, of maize as well as clearings for cattle." 37; greatest length ofskull, 39.2; zygomatic breadth, 17.8; interorbital breadth, 10.3; mastoid breadth, 15.5; length ofnasals, 15.7; Heteromys desmarestianus zonalis length ofrostrum, 17.7; length ofmaxillary Goldman, 1912 toothrow, 5.3; depth of cranium, 9.6. Distribution. - Forested eastern basal 1912. Heteromys zonalis Goldman, Smithson­ slopes ofmountains in central Veracruz be­ ian Misc. CoIl., 56(36):9, 19 February. tween 210 m and 460 m. Goldman (1951) Holotype. -Adult female, skin and skull, described the type locality. USNM 170976, from Rio Indio, near Ga­ Remarks. - This is the northernmost ex­ tun, Canal Zone, Panama; obtained on 15 tension of the genus. Rogers and Schmidly February 1911 by Edward A. Goldman. (1982) recognized temporalis as a subspe­ Measurements ofholotype. - Total length, cies ofdesmarestianus and presented exter­ 263; length oftail, 134; length ofhind foot, nal and cranial measurements. 35; greatest length ofskull, 36.3; zygomatic breadth, 16.8; interorbital breadth, 9.6; mastoid breadth, 14.5; length ofnasals, 13.6; Heteromys desmarestianus underwoodi length ofnostrum, 15.7; length ofmaxillary Goodwin, 1943 toothrow, 5.2; depth ofcranium, 9.5. Distribution. - Widely distributed in Pa­ 1943. Heteromys desmarestianus underwoodi nama from the northeastern part of the Goodwin, Amer. Mus. Novit., 1227:1, 22 country through central Panama and into April. eastern Panama. Holotype. -Adult female, skin and skull, Remarks. -Goldman (1920) recognized AMNH 131729, from Escazu, 7 mi SW San this taxon as a subspecies of desmaresti­ Jose, 3,000 ft, Province of San Jose, Costa anus. Rica; obtained on 28 April 1938 by C. F. Underwood. Measurements ofholotype. - Total length, Heteromys gaumeri 277; length oftail, 148; length ofhind foot, 30; length ofear, 15; greatest length ofskull, Diagnosis. - Size medium for genus; head 34.5; zygomatic breadth, 15.6; interorbital and body length less than 130 mm; tail lon­ breadth, 9.7; mastoid breadth, 14.3; length gerthan head and body, well-haired, grayish of nasals, 14.0; length of rostrum, 16.0; brown above, whitish below, and with ter­ length of maxillary toothrow, 5.2. minal tuft of hair; ears dusky, edged with Distribution. -Central Costa Rica at el­ dull white; dorsal pelage dark- to medium­ evations ranging from 1,350 m to 1,700 m. gray with orange-buff hairs; broad, ochra­ Remarks. - The range of this subspecies ceous lateral line, extending from the cheeks is located between that of H. d. planifrons to the base ofthe tail; venter and feet white; and H. d. subaffinis. H. d. underwoodi is soles offeet haired posterior to the last plan­ found at higher elevations and is smaller tar tubercle; skull medium-sized; large au­ than either ofthe two preceding subspecies. ditory bullae; lower premolar with three Goodwin (1943) presented external and cra­ lophs; upper and lower molars with three nial measurements for the type and one male lophs, enamel island formed between meta- TAXONOMY 109 loph (id) and cingulum disappearing with Holotype. -Adult male, skin and skull, wear; baculum with broad base approxi­ AMNH 12028 and 10461, from Chichen­ mately one-third the total bacular length; Itza, Yucatan, Mexico; obtained on 17 baculum with narrow oval shaft making up March 1896 by Frank M. Chapman. the remaining two-thirds ofthe length end­ Measurements ofholotype. - Total length, ing in a slightly upturned tip; 2n = 56, FN 292; length oftail, 162; length ofhind foot, = 76. 32; length ofear, 14.5; interorbital constric­ Comparisons. - This species can be dis­ tion, 8.9; length of nasals, 15.6; length of tinguished from all other species ofHetero­ rostrum, 16.4; length ofmaxillary toothrow, mys by the presence ofhair on the posterior 4.8. portion of the sole of the hind foot (hair is Remarks. - A review of this species was absent in other species) and a distinct, broad, given by Engstrom et al. (1987) and Schmidt ochraceous lateral line. Only the range of et al. (1989). External and cranial measure­ H. desmarestianus may overlap that of H. ments were presented by Best (1992), Eng­ gaumeri. Compared with H. desmaresti­ strom et al. (1987), Goldman (1911), and anus, H. gaumeri is smaller in both external Schmidt et al. (1989). This species exhibits and cranial measurements; has largc:r au­ little geographic variation, and populations ditory bullae; has a broader ochraceous lat­ are chromosomally monomorphic (Eng­ eralline; a well-haired tail with a terminal strom et al., 1987). Rogers (1990) found lit­ tuft; and a baculum with a narrow shaft. tle genic variation. Distribution. -Endemic to the Yucatan Peninsula at elevations from sea level to 100 Heteromys oresterus m. The range includes: northern Belize; El Peten, Guatemala; eastern Tabasco, Mexi­ Diagnosis. -Size very large for genus; co; and the Mexican states of Campeche, head and body greater than 140 mm; tail Quintana Roo, and Yucatan. This species length greater than or equal to length ofhead prefers tropical deciduous and subdecidu­ and body; dorsum blackish gray, grizzled ous-subperennial tropical rain forests and with ochraceous or buffY hairs; sides more thorn scrub forest (Engstrom et al., 1987; buffY than the dorsum; venter and feet white; Schmidt et al., 1989). Goldman (1951) de­ forelimbs more buffY than dorsum; hind scribed the localities for specimens collected limbs above and ankles dark gray; ears in the Mexican states of Campeche, Quin­ blackish with white edgings; tail black above tana Roo, and Yucatan. and white below except for black base and Remarks. - H. gaumeri is monotypic. white tip; premaxillae terminating posterior Based on morphologic, karyologic, and gen­ to nasals; palate narrow; 2n = 60, FN = 78. ic information, Engstrom et al. (1987) and Comparison. -See account of H. nelsoni Rogers (1990) recommended that H. gau­ for a comparison with that species. Only the meri be removed from the desmarestianus range of three subspecies of H. desmares­ species group and suggested that this taxon tianus (pIa nifrons, subaffinis, and under­ might warrant separate subgeneric recog­ woodi) are near the distribution ofthis spe­ nition. Extensive morphologic comparisons cies. From H. desmarestianus, H. oresterus ofthis species with species ofLiomys were may be identified by its larger body size; the made by Genoways (1973). soft, instead ofstiff, spines or bristles ofthe pelage; and the overall greater dimensions Heteromys gaumeri of the skull. Allen and Chapman, 1897 Distribution. - Known only from the provinces ofCartago and San Jose in central 1897. Heteromys gaumeri Allen and Chapman, Costa Rica at elevations from 1,800 m to Bull. Amer. Mus. Nat. Hist., 9:9, 23 February. 2,650 m. It appears to be restricted to cloud 110 WILLIAMS ET AL. forest in the western portion of the Tala­ Diagnosis. -Size large for genus; poste­ manca Range (Rogers and Rogers, 1992a). rior borders of nasals equal to posterior Remarks. - This is a monotypic species. margin ofpremaxillae; braincase arched, not Mascarello and Rogers (1988) and Rogers flat; palate narrow; rostrum not tapering an­ (1989) described the chromosomal varia­ teriorly; posterior molars equal to or broad­ tion. Rogers (1990) recommended that H. er than premolars; more intricate enamel oresterus be removed from the subgenus folds in posterior upper molars; posterior Xylomys and instead be included in the des­ cingulum on last upper molar as well de­ marestianus species group (desmarestianus veloped as either of the other two lophs, and oresterus) within the subgenus Hetero­ joined at center with metaloph; parietals ex­ mys. External and cranial measurement for tend laterally along lambdoidal crest, nearly this taxon were given by Harris (1932), Best reaching the mastoids; 2n = 42, FN = 72. (1992), and Rogers and Rogers (1992a). Remarks. - Xylomys currently is consid­ ered to be monotypic. Rogers (1989, 1990), based on chromosomal and genic data, Heteromys oresterus Harris, 1932 found that the two traditional species (H. oresterus, H. nelsoni) of Xylomys do not 1932. Heteromys oresterus Harris, Occas. Pa­ form a natural group. He recommended that pers Mus. ZooI., Univ. Michigan, 248:4, 4 Au­ H. oresterus be included within the des­ gust. marestianus species group; H. nelsoni may Holotype. -Adult male, skin and skull, be allied to this group, although the affinities UMMZ 64027, from El Copey de Dota, are not clear. Thus, the subgeneric ranking Cordillera de Talamanca, 6,000 ft, Costa of Xylomys is doubtful. Rica; obtained on 25 May 1931 by Austin Smith. Species Account Measurements ofholotype. - Total length, 357; length oftail, 174; length ofhind foot, Heteromys nelsoni 40; length ofear, 16; greatest length ofskull, 39.6; zygomatic breadth, 16.8; interorbital Diagnosis. -Size largest ofthe genus; ex­ breadth, 9.6; length ofnasals, 16.2; breadth ternal and cranial measurements the largest of braincase, 15.7; length of maxillary for the genus; head and body length greater toothrow, 5.2 (Harris, 1932). than 140 mm; tail longer than length ofhead Remarks. - This species is restricted to and body; dorsum mouse gray, darker along cloud forest habitats (Rogers and Rogers, the mid-dorsum; lateral line absent; inner 1992a). The type locality, which is about sides offore- and hind limbs dusky; venter 25 miles south ofCartago, is on "the Pacific white; ears large, blackish, without white watershed on the boundary between the edge; tail dusky above, whitish beneath ex­ subtropical and temperate zones" (Harris, cept tip which is dark all around; pelage 1932:1). Rogers and Rogers (1992a) re­ harsh, but bristles soft (also see subgeneric viewed the biology of this species. diagnosis). Comparisons. -Heteromys nelsoni dif­ fers from H. oresterus in overall greater size; Subgenus Xylomys the ears ofnelsoni lack the white edge found in oresterus; the nasals and premaxillae end 1902. Xylomys Merriam, Proc. BioI. Soc. Wash­ at the same level in nelsoni rather than hav­ ington, 15:43,5 March. ing the premaxillae longer than the nasals Type species. -Heteromys nelsoni Mer­ in oresterus; the tail ofnelsoni is dark-tipped, riam, 1902, Proc. BioI. Soc. Washington, whereas it is white-tipped in oresterus. Only 15:43, 5 March. H. d. goldmani occurs near the range ofH. TAXONOMY 111 nelsoni; H. nelsoni occurs at higher eleva­ 1881, in Allen, Bull. Mus. Compo Zoo1., tions, is larger in overall size, has soft, in­ 8:187, March. stead of stiff, bristles, and has wider pos­ Diagnosis. -Cheek teeth with medium­ terior molars. high crowns; upper incisors asulcate; only Distribution. - Known only from south­ two lophids on lower premolar; accessory ern Chiapas and western Guatemala. Spec­ enamel island on molars present only for imens have been collected in cloud forest at short period (visible only in unworn mo­ elevations ranging from 2,500 m to 2,800 lars); entostyle closely united to hypocone m, near Cerro Mozotol in southeastern so that Y-shape of median valley of upper Chiapas, Mexico, and from Volcan Taju­ premolar is poorly formed; auditory region mulco in western Guatemala (Rogers and uninflated; pelage hispid, consisting of stiff Rogers, 1992b). The habitat of the type lo­ spines mingled with slender soft hairs; soles cality was described by Goldman (1951). of hind feet sparsely haired; interpterygoid Remarks. - This is a monotypic species fossa V-shaped anteriorly; claw of second known only from four localities (Rogers and digit on hind foot spoon-like; adapted for Rogers, 1992b). The habitat ofthe type lo­ scampering. cality was characterized as humid, heavily Remarks. -Although Liomys is a com­ forested Pacific slopes of the Sierra Madre mon inhabitant of Mexico and Central (Goldman, 1951). America, Gray (1868) did not describe the first representative ofthe genus until 1868. The first species were described in the genus Heteromys nelsoni Merriam, 1902 Heteromys until Merriam (1902) described the genus Liomys just after the turn of the 1902. Heteromys nelsoni Merriam, Proc. BioI. 20th century. Goldman (1911) reviewed Soc. Washington, 15:43,5 March. both Liomys and Heteromys. Genoways Holotype. -Adult male, skin and skull, (1973) was the last author to review the ge­ USNM 77920, from Pinabete, 8,200 ft, nus Liomysand his arrangement is generally Chiapas; obtained on 11 February 1896 by followed in these accounts. Systematic re­ Edward W. Nelson and Edward A. Gold­ lationships among Liomys species, based on man. genic data, were investigated by Rogers Measurements ofholotype. - Total length, (1990). 358; length oftail, 195; length ofhind foot, 43.5; greatest length of skull, 40.6; inter­ orbital breadth, 9.3; mastoid breadth, 15.8; Key to the Species length of nasals, 15.8; length of maxillary 1. Five planter tubercles; pterygoid bones toothrow, 6.0; depth of braincase, 10.1. with broad wings; shaft ofbaculum oval Remarks. - Goldman (1911) suggested to tip; glans penis long (more than 75%) that this species represented a survivor of in comparison with baculum; FN of an ancestral group with more complicated chromosomes 60; upper parts greyish dentition. External and cranial measure­ brown, lateral stripe pale pinkish to buffy; ments for this taxon were given by Best occurring on Mexican Plateau and in ad­ (1992) and Rogers and Rogers (1992b). jacent areas ofnorthern and central Me­ xico, south as far as south-central Oaxaca ...... Liomys irroratus Genus Liomys 1'. Usually six plantar tubercles; pterygoid bones with narrow wings; shaft of bacu­ 1902. Liomys Merriam, Proc. BioI. Soc. Wash­ lum either flattened dorsoventrally or ington, 14:44, 5 March. flattened dorsoventrally and compressed laterally at some point; glans penis short Type species. -Heteromys alleni Coues, (less than 75%) in comparison with bac- 112 WILLIAMS ET AL.

ulum; FN ofchromosomes 48 or 56; up­ aging less than 33.5 mm); FN of chro­ per parts either reddish brown or choc­ mosomes 86; occurring from southern olate brown, or somewhat paler, with Oaxaca to central Costa Rica . either an ochraceous lateral stripe or lat­ ...... Liomys salvini eral stripe absent; not occurring on the 4'. Size large (greatest length of skull aver­ ~exican Plateau 2 aging over 34.5 mm); FN of chromo­ 2. Upper parts reddish brown with an somes 84 or less; occurring only in central ochraceous lateral stripe; interorbital re­ Panama Liomys adspersus gion broad in comparison with greatest length of skull; distal end ofthe shaft of the baculum with a laterally compressed Species Accounts ventral keel and just posterior to this re­ gion, the shaft is flattened dorsoventrally; Liomys adspersus tip of glans penis long when compared with its total length; FN ofchromosomes Diagnosis. - External and cranial mea­ 48; hairs on back not curled upward and surements large; premolars similar in struc­ not visible above spines; occurring along ture to those of L. salvini; baculum with the Pacific slope ofwestern ~exico as far large rounded base, shaft oval to a pointjust south as the vicinity ofTonahi, Chiapas, posterior to the slightly upturned tip where in the central valley of Chiapas, and in the southern half ofVeracruz ...... 3 it is dorsoventrally flattened; glans penis 2'. Upper parts chocolate brown to some­ medium-sized in comparison with length of what paler, lateral stripe absent; inter­ baculum, tip of glans short, glans highly orbital region narrow in comparison with sculptured and with deeply incised ventral greatest length ofskull; shaft ofbaculum folds; 2n = 56; FN probably 84; wings of dorsoventrally flattened just posterior to pterygoids narrow; six plantar tubercles; up­ slightly upturned tip, no laterally com­ per parts usually chocolate brown (some pressed ventral keel present; tip ofglans paler individuals may show grayish tones); penis short when compared with its total no lateral stripe; hairs on back curled up­ length; FN ofchromosomes 56; hairs on ward so as to be conspicuous above spines. back curled upward and visible above the Comparisons. -Specimens ofLiomys ad­ spines; occurring in Central America and spersus can easily be distinguished from into ~exico along the Pacific coast as far as the vicinity of Reforma, Oaxaca .... , 4 specimens ofL. salvini by their much larger 3. In southeastern Jalisco, size small (great­ external and cranial measurements. In both est length ofskull, 28.9 to 32.0 mm; spec­ species 2n = 56, but FN = 86 in salvini and imens approaching L. spectabilis in size FN = 84 in adspersus. occur in Guerrero and Oaxaca but these Distribution. -Central Panama princi­ are still slightly smaller and have pro­ pally on the Pacific versant. portionally deeper braincases); in Jalisco Remarks. - Liomys adspersus is mono­ hind foot rarely more than 30; laterally typic. This species is not sympatric with any compressed ventral keel on baculum other member ofthe genus; the distribution short, 0.85 to 1.25 mm; FN of chromo­ ofLiomys salvini is geographically nearest, somes 66; occurring along Pacific coast with the ranges ofthe two species separated of western ~exico, in the central valley of Chiapas, and in southern Veracruz ... by a distance of approximately 300 km...... Liomys pictus These two taxa have been shown to be close­ 3'. In southeastern Jalisco, size large (great­ ly related morphologically, karyotypically est length ofskull, 33.0 to 35.3 mm); hind (Genoways, 1973), and genically (Rogers, foot rarely less than 30 mm; laterally 1990). compressed ventral keel on baculum rel­ atively long, 1.30 mm; FN of chromo­ Liomys adspersus (Peters, 1874) somes 64; occurring only in southeastern Jalisco Liomys spectabilis 1874. Heteromys adspersus Peters, ~onatsb. 4. Size small (greatest length of skull aver- preuss. Akad. Wiss., Berlin, p. 357, ~ay. TAXONOMY 113

Holotype. - Young adult male, mounted ygoid bones are broad rather than narrow skin with skull, unknown number in Berlin in irroratus. The upper parts ofirroratus are Museum, from Panama. Type locality re­ grayish brown with a pale pinkish to buffy stricted to City of Panama by Goldman lateral stripe as opposed to upper parts red­ (1920). dish brown with an ochraceous lateral stripe Measurements ofholotype. - Total length, in pictus and spectabilis. 240; length of tail, 95; length of hind foot Distribution. - The range of Liomys ir­ 30. ' roratus includes the Mexican Plateau and Remarks. -Genoways (1973) found little adjacent areas from southern Chihuahua in variation among populations ofthis species. the northwest and southern Texas in the northeast to south-central Oaxaca in the south. It generally occurs to the east of the Liomys irroratus Sierra Madre Occidental, but also is found in various parts ofthe Transverse Volcanic Diagnosis. -A medium-sized spiny Belt. pocket mouse with the cranium relatively Remarks. -Hooper and Handley (1948) broad in comparison with length; protoloph presented a synopsis ofsubspecies ofirrora­ of upper permanent premolar composed of tus and analyzed the trends in geographic three discernible cusps; metaloph of upper variation within the species. Prior to the premolar with three cusps (hypocone larg­ study by Genoways (1973), three species est, metacone only slightly smaller than hy­ (bulleri, guerrerensis, and irroratus) were pocone); entostyle distinct but not widely recognized in the irroratus group by Gold­ separated from hypocone; re-entrant angle man (1911). However, Genoways (1973) re­ on labial side oflower premolar not united duced bulleri and guerrerensis to subspecific with median valley; baculum simple, with status while recognizing seven subspecies in large rounded base, oval-shaped shaft and the remaining species irroratus. The rela­ slightly upturned tip (extreme tip of bacu­ tionship of irroratus with the other species lum may ?e slightly laterally compressed); of Liomys has been little clarified by mor­ glans pems relatively long in comparison phologic, karyotypic, or genic data (Geno­ w~th length ofbaculum; 2n = 60, FN = 62; ways, 1973; Rogers, 1990). Liomys irrora­ wmgs of pterygoids relatively broad; five tus and L. pictus occur sympatrically in a plantar tubercles; upper parts grayish brown; zone from central Jalisco southward through lateral stripe, which is usually present, gen­ Michoacan to the region ofthe Balsas Basin erally pale pink to buff. and Sierra Madre del Sur of Guerrero and Comparisons. -Specimens of Liomys ir­ the Sierra Madre ofOaxaca. Liomys irrora­ roratus are most likely to be confused with tus is potentially sympatric with L. specta­ L. pictus or L. spectabilis. In areas of po­ bilis in southeastern Ja1isco, butthe two spe­ tential sympatry, L. irroratus is larger than cies have not been reported from the same L. pictus, but smaller than L. spectabilis. locality. The biology ofthis species was re­ There are five plantar tubercles in L. irrora­ viewed by Dowler and Genoways (1978). tus, whereas the other species have six (ex­ cept one population ofpictus). The karyo­ type of L. irroratus is composed of 60 chromosomes as opposed to 48 in L. pictus Liomys irroratus alleni (Coues, 1881) and L. spectabilis. The baculum ofirroratus 1881. Heteromys alleni Coues, in Allen, Bull. has an ovoid shaft to the slightly upturned Mus. Compo ZooI., 8:187, March. tip, whereas the baculum ofpictus and spec­ 1902. Liomys canus Merriam, Proc. BioI. Soc. tabilis has a terminal ventral keel that is Washington, 15:44, 5 March. laterally compressed and a section of the 1947. Liomys irroratus pullus Hooper, J. shaft that has been dorsoventrally flattened Mamm., 28:47, 17 February. posterior to the keel. The wings ofthe pter- 1948. Liomys irroratus acutus Hall and Villa- 114 WILLIAMS ET AL.

R., Univ. Kansas Publ., Mus. Nat. Hist., 1:253, small, subtriangular interparietal bone. L. i. 26 July. bulleri is potentially in contact with L. i. Holotype.-Probably an adult, sex un­ jaliscensis, although no direct intergrades known, skull in skin, MCZ 5889, from Ha­ have been found. cienda Angostura, Rio Verde, San Luis Po­ tosi; obtained on 26 February 1878 by Edward Palmer. Liomys irroratus guerrerensis Measurements of holotype.-No mea­ Goldman, 1911 surements are available. Distribution. - Extensive geographic area 1911. Liomys guerrerensis Goldman, N. Amer. on the Mexican Plateau north ofthe Trans­ Fauna, 34:62, 7 September. verse Volcanic Belt; also occurring in Sierra Holotype. - Subadult female, skin and Madre Oriental of Tamaulipas and Nuevo skull, USNM 127523, from Omilteme, Leon. Guerrero; obtained 17 May 1903 by Ed­ Remarks. - Populations ofL. i. alleni are ward W. Nelson and Edward A. Goldman. characterized by large size and a relatively Measurements ofholotype. - Total length, high percentage ofindividuals with the in­ 255; length oftail, 127; length ofhind foot, terparietal bone divided in halfand the pos­ 34; greatest length ofskull, 33.3; interorbital terior termination of nasals truncate. L. i. constriction, 8.6; mastoid breadth, 15.8; alleni intergrades with L. i. jaliscensis in an length of nasals, 12.5; length of rostrum, area from near Lago de Chapala in Jalisco 14.9; length of maxillary toothrow, 9.2. and Michoacan northward through Jalisco Distribution. - Pacific slope of the Sierra into extreme southern Zacatecas. Madre del Sur ofGuerrero. Remarks. - Until the study ofGenoways (1973), guerrerensis was considered to be a Liomys irroratus bulleri monotypic species of the irroratus-group. (Thomas, 1893) Genoways (1973) found specimens from Chilpancingo, Guerrero, that were inter­ 1893. Heteromys bulleri Thomas, Ann. Mag. mediate between guerrerensis and torridus. Nat. Hist., ser. 6, 11 :330, April. L. i. guerrerensis can be distinguished from Holotype. - Adult female, skin in alcohol all other subspecies of irroratus by larger with skull removed, BM(NH) 93.3.6.39, size and darker upper parts. from La Laguna, Sierra de Juanacatlan, Ja­ lisco; obtained in December 1892 by A. C. Buller. Liomys irroratus irroratus Measurements of holotype. -Greatest (Gray, 1868) length of skull, 34.1; zygomatic breadth, 16.7; interorbital constriction, 8.6; mastoid 1868. Heteromys irroratus Gray, Proc. Zool. Soc. breadth, 15.5; length ofnasals, 13.4; length London, p. 205, May. ofrostrum, 15.3; length ofmaxillary tooth­ 1868. Heteromys albolimbatus Gray, Proc. Zool. row, 5.9; depth of braincase, 10.0. Soc. London, p. 205, May. Distribution. - Known only from the 1956. Liomys irroratus yautepecus Goodwin, Amer. Mus. Novit., 1757:7,8 March. mountains of west-central Jalisco. Remarks. - Until the work of Genoways Holotype. -Subadult of unknown sex, (1973), bulleri was considered to be a mono­ skin and skull, BM(NH) 59.7.10.2, from typic species, but was placed as a subspecies Oaxaca; obtained on an unknown date by of irroratus in this study. L. i. bulleri is a A. Salle. Type locality restricted to Oaxaca, large-sized subspecies characterized by a Oaxaca, by Genoways (1973). TAXONOMY 115

Measurements of holotype. -Interobital Measurements ofholotype. - Total length, constriction, 9.1; length of nasals, 12.5; 231; length oftail, 114; length ofhind foot, length ofrostrum, 15.2; length ofmaxillary 30; greatest length ofskull, 31.0; zygomatic toothrow, 6.3; mastoid breadth, 15.2. breadth, 14.5; interorbital constriction, 8.0; Distribution. -Confined to central and mastoid breadth, 14.1; length ofnasals, 12.8; south-central Oaxaca. length ofmaxillary toothrow, 5.2; depth of Remarks. - This large-sized subspecies braincase, 8.6. was the first member of the genus to be Distribution. - Gulfcoastal lowlands from described (Gray, 1868). Besides large size, southern Texas to northern Puebla and cen­ this subspecies has a large percentage ofin­ tral Veracruz. dividuals with the posterior margin of the Remarks. - This small- to medium-sized nasals emarginate. subspecies intergrades with L. i. alleni for a considerable distance along the eastern foothills of the Sierra Madre Oriental and Liomys irroratus jaliscensis the eastern edge of the Mexican Plateau. (J. A. Allen, 1906) Some geographic variation within the sub­ species texensis was noted by Genoways 1906. Heteromys jaliscensis J. A. Allen, Bull. (1973). Specimens from Texas and northern Amer. Mus. Nat. Rist., 22:251, 25 July. Tamaulipas averaged larger than other sam­ Holotype. -Adult male, skin and skull, ples. The smallest specimens oftexensis, on AMNH 26325, from Las Canoas, 7,000 ft, the average, were from the vicinity ofEbano, Jalisco; obtained on 6 August 1905 by J. H. San Luis Potosi. Batty. Measurements of holotype. -Greatest length of skull, 31.6; interorbital constric­ Liomys irroratus torridus tion, 8.0; mastoid breadth, 14.1; length of Merriam, 1902 nasals, 11.6; length ofrostrum, 13.4; length of maxillary toothrow, 5.2; depth of brain­ 1902. Liomys torridus Merriam, Proc. BioI. Soc. case, 9.2. Washington, 15:45,5 March. Distribution. - Known from southern, 1902. Liomys torridus minor Merriam, Proc. BioI. Soc. Washington, 15:45,5 March. central northern Jalisco and extreme south­ 1903. Heteromys exiguus Elliot, Field Columbo ern Zacatecas and Nayarit. Mus., Zool. Ser., 3: 146, 20 March. Remarks. -L. i. jaliscensis is a medium­ sized subspecies that is located geographi­ Holotype. - Adult female, skin and skull, cally between two larger-sized subspecies, USNM 69645, from Cuicatlim, Oaxaca; ob­ alleni and bulleri. tained on 14 October 1894 by Edward W. Nelson and Edward A. Goldman. Measurements ofholotype. - Total length, Liomys irroratus texensis 242; length oftail, 134; length ofhind foot, Merriam, 1902 28; greatest length ofskull, 30.3; zygomatic breadth, 14.2; interorbital constriction, 7.9; 1902. Liomys texensis Merriam, Proc. BioI. Soc. mastoid breadth, 14.0; length ofnasals, 11.6; Washington, 15:44,5 March. length ofrostrum, 13.6; length ofmaxillary 1911. Liomys irroratus pretiosus Goldman, N. toothrow, 4.5; depth of braincase, 8.9. Amer. Fauna, 34:58, 7 September. Distribution. -South of the Transverse Holotype. - Adult female, skin and skull, Volcanic Belt in the Mexican states ofPueb­ USNM 58670, from Brownsville, Cameron la, Morelos, Guerrero, and northern Oa­ Co., Texa~; obtained on 19 February 1894 xaca. by J. Alden Loring. Remarks. - The subspecies torridus is 116 WILLIAMS ET AL. composed ofindividuals ofmedium to small annectens to a subspecies of L. pictus. The size for the species. The subspecies torridus species described by Goodwin (1956), L. appears to intergrade with the subspecies pinetorum, was placed as a junior synonym irroratus in central Guerrero and with the ofL. p. pictus. Genic data indicate that this subspecies alleni along the southern slope species is paraphyletic (Morales and Eng­ ofthe Transverse Volcanic Ble1t in Morelos strom, 1989; Rogers, 1990). The closest rel­ and Puebla, and in the lower country ofeast­ ative ofL. pictus in the genus is L. spectabilis ern Puebla. (Genoways, 1973; Rogers, 1990). These two species occur sympatrically in southeastern Jalisco at three localities. L. pictus is also Liomys pictus sympatric with L. salvini in coastal Oaxaca and Chiapas and with L. irroratus in a nar­ Diagnosis. - External and cranial mea­ row band from southern Jalisco through surements medium to small for the genus, Michoacan and Guerrero to Oaxaca. This although some populations (annectens) are is the only member of the genus that has relatively large in size; cranium relatively been reported as occurring sympatrically narrow in comparison with length; proto­ with more than one other species ofthe ge­ loph ofupper premolar generally appearing nus. The biology of this species was re­ to be composed ofa single cusp; three cusps viewed by McGhee and Genoways (1978). ofmetaloph connected by loph so as not to form discrete cones; hypocone largest cusp on metaloph; entostyle always connected to Liomys pictus annectens hypocone by loph; re-entrant angle on labial (Merriam, 1902) margin oflower premolar not reaching me­ dian valley; baculum long and with a small 1902. Heteromys annectens Merriam, Pmc. Bioi. Soc. Washington, 15:43,5 March. rounded base, distal end of shaft with ven­ tral keel that is laterally compressed and the Holotype. - Adult male, skin and skull, shaft dorsoventrally compressed posterior USNM 71510, from Pluma Hidalgo, Oa­ to terminal keel; 2n = 48; FN = 66; wings xaca; obtained on 18 March 1895 by Ed­ of pterygoids narrow; six plantar tubercles ward W. Nelson and Edward A. Goldman. on most specimens, although some individ­ Measurements ofholotype. - Total length, uals of L. p. planitarensis have only five; 300; length oftail, 165; length ofhind foot, upper parts reddish brown; lateral stripe 33; greatest length ofskull, 34.3; zygomatic generally ochraceous, but may be rather pale. breadth; 16.1; interorbital constriction, 7.9; Comparisons. - Liomys pictus is gener­ mastoid breadth, 14.7; length ofnasals, 14.7; ally smaller than L. spectabilis and this is length of maxillary toothrow, 5.3; depth of especially true in sympatric populations. The braincase, 8.9. hind foot of pictus is rarely more than 30 Distribution. - High elevations in the Si­ mm, but rarely less than 30 mm in spec­ erra Madre del Sur ofGuerrero and Oaxaca. tabiUs. In pictus, the FN = 66, whereas in Remarks. - Until the study by Genoways spectabiUs FN = 64. (1973), this taxon was considered a species Distribution. - West coast and adjacent distinct from L. pictus. However, there is slopes of Sierra Madre Occidental and Si­ clear evidence for intergradation between erra Madre del Sur from near Nogales, So­ the taxa in the vicinity of Candelaria, Oa­ nora, to Tonahl, Chiapas; central valley of xaca, along the lower slope of the Sierra Chiapas into adjacent Guatemala; across the Madre del Sur. L. p. annectens is distin­ Isthmus of Tehuantepec into southern Ve­ guished from other subspecies by larger size racruz on the east coast of Mexico. and darker color dorsally with a darker or­ Remarks. -Genoways (1973) reduced L. ange lateral stripe. TAXONOMY 117

Liomys pictus hispidus 1902. Liomys obscurus Merriam, Proc. BioI. Soc. (J. A. Allen, 1897) Washington, 15:48,5 March. 1902. Liomys phaeura Merriam, Proc. BioI. Soc. 1897. Heteromys hispidus J. A. Allen, Bull. Washington, 15:48,5 March. Amer. Mus. Hist., 9:56, 15 March. 1902. Liomys orbitalis Merriam, Proc. BioI. Soc. 1902. Liomys sonorana Merriam, Proc. BioI. Washington, 15:48, 5 March. Soc. Washington, 15:47,5 March. 1903. Heteromys paralius Elliot, Field Columb. 1906. Heteromys pictus escuinapae J. A. Allen, Mus., Zool. Ser., 3:233, 3 September. Bull. Amer. Mus. Nat. Hist., 22:221, 25 July. 1956. Liomyspinetorum Goodwin, Amer. Mus. Novit., 1791 :2, 28 September. Holotype. -Subadult female, skin and skull, AMNH 8333/6667, from Rancho El Holotype. -Adult female, skin and skull, Coloma, Compostela, Nayarit; obtained on BM(NH) 93.8.12.2, from San Sebastian, 11 February 1893 by A. C. Buller. 4,300 ft, Jalisco; obtained on 9 May 1893 Measurements of holotype. -Greatest by A. C. Buller. length of skull, 30.5; interorbital constric­ Measurements of holotype. -Zygomatic tion, 7.1; mastoid breadth, 13.9; length of breadth, 15.1; interorbital constriction, 7.6; nasals, 12:3; length ofrostrum, 13.4; length mastoid breadth, 13.7; length of maxillary of maxillary toothrow, 5.0. toothrow, 5.2; depth of braincase, 8.5. Distribution. - Northwestern Mexico Distribution. -Coastal western Mexico from near Nogales, Sonora, southward in from Santiago, Nayarit, southward through coastal Sinaloa and northern Nayarit and Jalisco, Colima, Michoacfm, Guerrero, Oa­ then inland as far as central Jalisco. xaca to Chiapas; central valley of Chiapas Remarks. - The subspecific name hispid­ and into and adjacent Guatemala; across us applies to populations along the north­ the Isthmus of Tehuantepec into southern western coast ofMexico that vary in a dinal Veracruz. fashion from medium-sized individuals in Remarks. -L. p. pictus is characterized Sonora and northern Sinaloa to relatively by a medium size, undivided interparietal small individuals in northern Nayarit. In bone, and a premaxillary bone terminating the vicinity of San BIas, Nayarit, hispidus posterior to the nasal bones. Goodwin (1956) intergrades with pictus from coastal areas to described L. pinetorum based upon a single the south. Specimens from the vicinity of specimen from near Tonala, Chiapas. Gen­ Tepic, Nayarit, and southeastward through oways (1973) showed that all characters of Nayarit to just south ofGuadalajara, Jalis­ this specimen fell within the range of vari­ co, are intermediate in size between pictus ation of L. p. pictus and placed pinetorum to the west and hispidus to the north, but as a junior synonym of pictus. Genoways possess cranial characters that ally them with (1973) described the intergradation ofpictus hispidus. and plantinarensis in Colima. The extent of morphological differentiation between pic­ tus andplantinarensis in Colima and Jalisco Liomys pictus pictus (Thomas, 1893) was assessed by Morales and Engstrom (1989). They (Morales and Engstrom, 1989: 1893. HeteromyspictusThomas, Ann. Mag. Nat. 2) suggested that, based on morphological Hist., ser. 6, 12:233, September. variation, "genetic exchange between the 1902. Liomys pictus rostratus Merriam, Proc. two subspecies is limited and that introgres­ BioI. Soc. Washington, 15:46, 5 March. sion is minimal." In this area ofcontact, L. 1902. Liomys pictus isthmius Merriam, Proc. BioI. Soc. Washington, 15:46,5 March. p. pictus has an interparietal length greater 1902. Liomys veraecrucis Merriam, Proc. BioI. than 4 mm, a hindfoot length greater than Soc. Washington, 15:47,5 March. 28 mm, and is found in more mesic areas. 118 WILLIAMS ET AL.

Liomys pictus plantinarensis especially short; protoloph of permanent Merriam, 1902 upper premolar appears to be composed of one cusp, metaloph composed ofthree and 1902. Liomys plantinarensis Merriam, Proc. sometimes four cusps, metacone of meta­ Bioi. Soc. Washington, 15:46, 5 March. loph sometimes larger than hypocone, en­ 1904. Liomys parviceps Goldman, Proc. Bioi. tostyle distinctly separated from other cusps Soc. Washington, 17:82,21 March. ofmetaloph; re-entrant angle on labial mar­ Holotype. -Adult female, skin and skull, gin of lower premolar connected with me­ USNM 45630/33595, from Platanar, Jalis­ dian valley; baculum with large rounded co; obtained on 4 April 1892 by Edward W. base, shaft oval to pointjust posterior to the Nelson and Edward A. Goldman. slightly upturned tip where it is dorsoven­ Measurements ofholotype. - Total length, trally flattened; 2n = 56; FN = 86; wings of 202; length oftail, 102; length ofhind foot, pterygoids narrow; six planter tubercles; up­ 26; greatest length ofskull, 29.4; interorbital per parts grayish brown to deep chocolate constriction, 7.4; mastoid breadth, 13.0; brown; no lateral stripe; hairs on back curled length of nasals, 11.8; length of maxillary so as to be conspicuous above spines. toothrow, 5.0. Comparisons. -Liomys salvini can be Distribution. - Interior drainage basins distinguished from L. pictus by its smaller and river systems of southeastern Jalisco, overall size. The entostyle on the upper pre­ eastern Colima, central Michoacan, and molar is more distinctly separated from oth­ northern Guerrero. er cusps in salvini than pictus and the re­ Remarks. - Members ofthe subspecies L. entrant angle on the labial margin of the p. plantinarensis are characterized by small lower premolar reaches the median valley size, high percentage ofindividuals with the in specimens of salvini but not pictus. The posterior border ofthe interparietal notched, baculum of salvini lacks the laterally com­ and a low percentage ofindividuals with the pressed distal keel found in pictus. The 2n termination ofthe nasal bones truncate. The ofL. salvini is 56, whereas that ofL. pictus number of plantar tubercles is usually con­ is 48. The upper parts of salvini are choc­ stant in species of Liomys, but the devel­ olate brown or paler and lack the lateral opment ofthe sixth tubercle is highly vari­ stripe whereas in pictus the upper parts are able in northern populations of L. p. reddish brown with an ochraceous lateral plantinarensis. Morales and Engstrom stripe. The hairs on the back of specimens (1989) found this taxon to be morphologi­ ofsalvini are curled upward so that they are cally distinct from L. p. pictus and a mono­ visible above the spines, but not in speci­ phyletic lineage within the pictus species­ mens ofpictus. group. They suggested that specific status Distribution. -Occurs along the Pacific may be warranted. Similar results were coast and adjacent mountain slopes from found by Rogers (1990) using genic data. In Reforma, Oaxaca, southward through Chia­ the area of contact, L. p. plantinarensis is pas, Guatemala, Honduras, EI Salvador, smaller and inhabits drier areas, whereas L. Nicaragua to Monte Rey, Costa Rica; also p. pictus is larger and is found in more mesic in the Caribbean drainage of central and areas (Morales and Engstrom, 1989). eastern Guatemala. Remarks. - Before the revision by Gen­ oways (1973), five species were recognized Liomys salvini in the crispus-group. Genoways (1973) rec­ ognized two species-salvini and adsper­ Diagnosis. - External and cranial mea­ sus-in a group that was renamed the sal­ surements small for the genus, tail being vini -group. He reduced crispus to a TAXONOMY 119 subspecies of L. salvini and placed L. an­ the slopes ofthe Guatemalan highlands on thonyi and L. heterothrix as a junior syn­ the southern coast of Guatemala. onym of L. s. salvini. Goodwin (1946) had earlier placed L. vulcani as a subspecies of L. salvini, which Genoways continued to Liomys salvini salvini (Thomas, 1893) follow. Morphologic, genic, and karyotypic data indicate that L. salvini is most closely 1893. Heteromys salvini Thomas, Ann. Mag. related to L. adspersus (Genoways, 1973; Nat. Rist., ser. 6,11:331, April. Rogers, 1990). L. salvini occurs sympatri­ 1893. Heteromys salvini nigrescens Thomas, cally only with L. pictus among the four Ann. Mag. Nat. Rist., ser. 6, 12:234, Septem­ other species in the genus. Carter and Gen­ ber. oways (1978) reviewed the biology of this 1902. Liomys heterothrix Merriam, Proc. BioI. species. Soc. Washington, 15:50, 5 March. 1932. Liomys anthonyi Goodwin, Amer. Mus. Novit., 528:2, 23 May. Liomys salvini crispus Merriam, 1902 1938. Liomys salvini aterrimus Goodwin, Amer. Mus. Novit., 987:4, 13 May. 1902. Liomys crispus Merriam, Proc. BioI. Soc. Holotype. -Adult male, skin and skull, Washington, 15:49,5 March. BM(NH) 75.2.27.35, from Duenas, Saca­ 1902. Liomys crispus setosus Merriam, Proc. tepequez, Guatemala; obtained on 31 July BioI. Soc. Washington, 15:49,5 March. 1873 by Osbert Salvin. Holotype. - Adult male, skin and skull, Measurements of holotype. -Greatest USNM 75105, from Tonahi, Chiapas; ob­ length of skull, 32.8; zygomatic breadth, tained on 7 August 1895 by Edward W. 15.0; interorbital constriction, 7.3; mastoid Nelson and Edward A. Goldman. breadth, 14.1; length ofnasals, 12.5; length Measurements ofholotype. - Total length, ofrostrum, 14.2; length ofmaxillary tooth­ 210; length of tail, 99; length of hind foot, row, 5.4; depth of braincase, 9.3. 27.5; greatest length of skull, 31.8; inter­ Distribution. -Guatemala (along the orbital constriction, 6.4; mastoid breadth, southern coast, in the highlands around 13.8; length of nasals, 12.2; length of ros­ Guatemala City, and in the valleys of the trum, 13.5; length of maxillary toothrow, Rio Negro and Rio Motagua as far as San 5.0; depth of braincase, 9.0. Pedro Sula, Honduras), most of El Salva­ Distribution. -Coastal areas of south­ dor, south-central Honduras, north-central eastern Oaxaca, Chiapas, and southwestern and central Nicaragua, Isla de Ometepe in Guatemala from Reforma, Oaxaca, in the Lake Nicaragua, extreme southwestern Nic­ north to vicinity of Mazatenago, Guate­ aragua, and western and central Costa Rica. mala, in the south. Remarks. -L. s. salvini is characterized Remarks. - The taxon crispus was con­ by large external and cranial measurements, sidered to be a distinct species by Goldman at least some individuals in each population (1911) and subsequent authors until the with the interparietal bone divided, and a work ofGenoways (1973). He considered it low percentage of individuals having the to be a subspecies of salvini characterized posterior margin ofthe interparietal deeply by short total length and length oftail, high notched. Genoways (1973) reduced two rec­ percentage ofindividuals with the posterior ognized species (anthonyi Goodwin, 1932, margin of the interparietal bone deeply and heterothrix Merriam, 1902) and two notched, and no individuals with the inter­ recognized subspecies (nigrescens Thomas, parietals divided. The subspecies crispus and 1893, aterrimus Goodwin, 1938) to junior salvini appear to come into contact along synonyms of salvini. 120 WILLIAMS ET AL.

Liomys salvini vulcani related to L. pictus as indicated by mor­ (J. A. Allen, 1908) phologic, genic, and karyotypic data (Gen­ oways, 1973; Rogers, 1990). These two taxa 1908. Heteromys vulcani J. A. Allen, Bull. Amer. occur sympatrically at three locations in Nat. Rist., 24:652, 13 October. southeastern Jalisco. It is potentially sym­ Holotype. -Adult female, skin and skull, patric with L. irroratus, but the two have AMNH 28315, from Volclin de Chinan­ not been collected together. L. spectabilis is dega, about 4,000 ft, Chinandega, Nicara­ monotypic. gua; obtained on 7 May 1909 by William B. Richardson. Liomys spectabilis Genoways, 1971 Measurements ofholotype. - Total length, 220; length oftail, 110; length ofhind foot, 1971. Liomys spectabilis Genoways, Occas. Pa­ 25; interorbital constriction, 6.8; mastoid pers Mus. Nat. Rist., Univ. Kansas, 5:1, 18 breadth, 13.5; length ofnasals, 11.1; length June. ofrostrum, 12.5; length ofmaxillary tooth­ row, 4.9. Holotype. -Adult male, skin and skull, Distribution. -Confined to western Nic­ KU 96051, from 2.2 mi NE Cont1a, 3,850 aragua on the volcanoes that make up the ft, Jalisco; obtained on 20 September 1963 Cordillera los Marrabios and the lowland to by Percy L. Clifton. the west of them, west of Lake Managua, Measurements ofholotype. - Total length, on the Meseta de los Pueblos west of Lake 280; length oftail, 142; length ofhind foot, Nicaragua, and on Isla de Zapatera. 31; length ofear, 17; greatest length ofskull, Remarks. - L. s. vulcani, which was rec­ 35.1; zygomatic breadth, 16.3; interorbital ognized as a distinct species until the work constriction, 8.2; mastoid breadth, 15.1; ofGoodwin (1946), is characterized by gen­ length of nasals, 14.0; length of rostrum, erally smaller size, especially cranially. In­ 16.0; length of maxillary toothrow, 5.0; tergradation between vulcani and salvini was depth of braincase, 8.6. detected in specimens from 8 km N Las Remarks. - Because only 21 specimens of Maderas, Nicaragua, and the taxa also ap­ this species are known, very little infor­ pear to be in contact between La Celera and mation is available on variation in the spe­ San Pedro near the southern edge of the cies (Genoways, 1973). Meseta de los Pueblos ofNicaragua. Subfamily Perognathinae Coues, 1875 Liomys spectabilis Diagnosis. -Size small to medium, from Diagnosis. -External and cranial mea­ 100 to 230 mm in total length and 5 to 60 surements large; premolar resembles that of g in mass; body form quadrupedal and gen­ L. pictus; baculum long with a small round­ erally mouselike; locomotion scansorial to ed base; distal end ofshaft with ventral keel subricochetal; hind limbs considerably larg­ that is laterally compressed; shaft dorso­ er than forelimbs; hind feet and legs longer ventrally compressed posterior to terminal than typical murids and cricetids; pes with keel; 2n = 48; FN = 64; wings ofpterygoids five clawed digits; pes soles naked or partly narrow; six plantar tubercles; upper parts clothed with a scant covering ofshort hairs; reddish brown; lateral stripe ochraceous. body hairs oftwo types-straight, relatively Comparisons. -See accounts for L. irro­ long overhairs and a thin underfur ofshort, ratus and L. pictus. often slightly curved hairs; upper incisors Distribution. -Confined to southeastern strongly grooved; molars progressively Jalisco. brachydont, rooted, and tuberculate; enam­ Remarks. - L. spectabilis is most closely el pattern ofcheek teeth persisting into late TAXONOMY 121 life of individual, but lost by wear in older Wahlert, 1985; Webster and Webster, 1975; individuals; lophs ofP4 unite first at or near Wood, 1935). center oftooth; protoloph ofP4 usually sin­ Remarks. - This diagnosis was devel­ gle-cusped; lophs ofupper molars unite first oped primarily from information about Re­ at the lingual margin and progress to buccal cent species. As defined above, the subfam­ margins; those ofp4 unite at center oftooth, ily Perognathinae includes the genera presenting an X-pattern; lophs oflower mo­ Perognathus and Chaetodipus, but not Mi­ lars united first at buccal margin and pro­ crodipodops (see remarks in the account of gressively at center of tooth, sometimes Dipodomyinae). Wood (1935:88) amended forming an H-pattern; no foramina and usu­ the name, Perognathidinae, given to the ally no pit between m3 and the base of the subfamily by Coues (1875). coronoid process; palate broad, extending beyond level of M3; center of palate be­ tween cheek teeth not ridged; squamosal in Key to the Genera broad contact with parietal on dorsal sur­ face of skull; squamosal not perforated by 1. Sole ofhind foot naked; pelage relatively auditory bulla; anterior zygomatic root not coarse, often with stiff, spine-like bristles on rump; stiff, coarse hairs usually pro­ greatly enlarged on joining lachrymal; large ject across anterior margin ofear pinna; orbital nonossification usually including antitragus of ear pinna lobed; mastoid ethmoid foramen; incisive foramen small; bulla usually not projecting posteriorly one prominent pterygoid fossa (the anterior beyond plane ofocciput; postero-medial fossa nearly invisible); auditory bullae border of mastoid bulla usually project­ greatly expanded, mastoid bullae appearing ing as a distinct indentation into the su­ on dorsal surface of skull; inflated interior praoccipital; interparietal width equal to of mastoid and tympanic bullae filled with or greater than interorbital breadth .... spongy trabeculae; no stapedius muscle in ...... Chaetodipus middle ear; no fusion ofcervical vertebrae; 1'. Posterior one-third to one-halfofsole of in so far as is known, no median ventral hind foot with sparse covering of short hairs; pelage relatively fine and soft, nev- foramina in the "central" of caudal verte­ er with stiff, spine-like bristles on rump; brae; scapula not prolonged posteriorly; ob­ no long, stiff, coarse hairs projecting across turator foramen not triangular; no articu­ anterior margin of ear pinna; antitragus lation of trapezoid and scapholunar; of ear pinna usually not lobed; mastoid astragalus articulating with cuboid; cuboid bulla projecting posteriorly beyond the with long process between calcaneum and plane ofthe occiput; postero-medial bor- navicular, meeting astragalus; dorsal sur­ der of mastoid bulla not projecting as a face of ectocuneiform hour-glass-shaped; distinct indentation into the supraoccip­ phallus with gently upcurved tip, and with ital; interparietal width nearly always less or without external spines and urethral lap­ than interoribital breadth .... Perognathus pets; baculum relatively long and slender with a swollen base and an upturned tip; tip of baculum usually simple, but one species Genus Chaetodipus with an ornate, trifid tip; nasolabialis pro­ fundus pars maxillaris muscles of dual or­ 1889. Chaetodipus Merriam, N. Amer. Fauna, 1:5,25 October. igin, from lateral zygoma and infraorbital foramen; M. cleidomastoideus present; ex­ Type species. - Perognathus [Chaetodi­ tensores breves muscles present; M. abduc­ pus] spinatus Merriam, 1889, N. Amer. tor hallucis present; lumbricales muscles not Fauna, 1:5, 25 October. reduced or absent (Burt, 1936; Hafner and Diagnosis. -Size small to medium, total Hafner, 1983; Homan and Genoways, 1978; length from about 150 to 230 mm, weight Merriam, 1889; Osgood, 1900; Ryan, 1989; from about 15 to 60 g; body form quadru- 122 WILLIAMS ET AL. pedal and scansorial with slight tendency to Hafner, 1983; Homan and Genoways, 1978; ricochetal; tail usually relatively long, length Ryan, 1989; Wood, 1935). averaging greater than length of head and Remarks. -Merriam (1889) includedjor­ body (except C hispidus); tail with distinct mosus in the subgenus Perognathus, an ar­ distal, dorsal crest and terminal tuft ofhairs rangement that endured until Patton et al. or without crest and terminal tuft; tail some (1981) demonstrated conclusively that it shade ofbrown or buffy above, whitish be­ should be classified as a Chaetodipus; Haf­ low; sole ofhind foot naked; earpinna short, ner and Hafner (1983) formally elevated rounded, and with a lobed antitragus; coarse, Chaetodipus to generic rank. In the follow­ long hairs project from in front of anterior ing key, presence or absence of spines (ac­ margin ofear pinna to partly cover opening; tually stiff bristles) are not infallible char­ usually a small, whitish spot present below acters, as juvenile and molting animals often the external auditory meatus; dorsal sur­ exhibit no obvious spines, individuals of faces generally some shade ofbuffy, brown­ some species normally having spines are ish, or grayish; dorsal parts usually tinged found without spines, and individuals of with black; usually a clear, buffy lateral stripe species normally lacking spines are found without blackish tinge; undersides usually with spines. whitish; hairs relatively short to medium in length and straight; texture ofhair relatively harsh; hairs flattened in cross section; hairs have dorsal trough; cuticular scales of Key to the Subgenera and Species troughed hairs extending laterally across 1. Tail without distal crest and tuft oflong trough, not forming longitudinal ridges; stiff, hairs at tip; length oftail usually shorter spine-like hairs present or absent in dorsal than length of head and body . pelage; mastoid bullae not extending be­ ·. .. Chaetodipus (Burtognathus) hispidus yond the plane of the occiput (except C I'. Tail with distal crest and long tuft of baileyi and Cjormosus, whose mastoid bul­ hairs at tip; length of tail averaging lae extend slightly beyond the occiput); tym­ greater than length of head and body . panic bullae relatively widely spaced ante­ · .... Chaetodipus (Chaetodipus) ..... 2 riorly on the ventral surface of the skull; 2. Rump with mix oflong, stiff, whitish or supraoccipital with strong lateral indenta­ blackish, bristly hairs or spines and nor- tions of mastoid bullae; interparietals not mal hairs 9 2'. Rump without stiffbristles or spines in compressed and wider than interorbital pelage 3 breadth; phallus relatively long; soft tissue 3. Posterior end of mastoid bullae pro­ ofphallus extends about one-halfthe length jecting slightly beyond the posterior ofthe baculum; phallus lacking urethral lap­ plane of the occipitals 4 pets; rim ofterminal crater ofphallus forms 3'. Posterior end ofmastoid bullae anterior vent-like urethral opening; baculum rela­ to or about level with posterior plane of tively long, with only slightly enlarged prox­ occipitals 5 imal end, and slender, strongly upturned 4. Size moderate, length ofhead and body distal end, or shaft thick and straight with generally less than 90 mm; length ofhind trifid tip; vesicular glands of male short, foot usually less than 26 mm; width of round or bulb-shaped, and yellow- to gray­ interparietals averaging 5.9 mm or less, rarely ranging to 6.5 . colored (pinkish and granular in fresh spec­ ·...... Chaetodipus formosus imens); head ofsperm approximating a tri­ 4'. Size large, length ofhead and body gen­ angle, with acute vertices; enamel patterns erally 90 mm or more; length of hind ofcheek teeth less persistent with age than foot usually 26 mm or greater; width of Perognathus; tendon at origin of M. rectus interparietals averaging 6.1 mm or femoris Y-shaped (Hall, 1981; Hafner and greater Chaetodipus baileyi TAXONOMY 123

5. Ear pinna large and rounded, length from River in southern Nevada, California, notch usually 10 mm or greater ..... or the Baja California peninsula ..... 11 · Chaetodipus artus 10'. Not occurring west ofthe Colorado Riv- 5'. Ear pinna short to moderate size, length er in southern Nevada, California, or from notch usually less than 10 mm .. 6 the Baja California peninsula...... 12 6. Tail thinly haired and with small crest; 11. Broad lateral stripe of buffY (yellowish occurring along the coastal lowlands of with strong orangish tone) contrasting western Mexico, from southern Sonora with brownish-appearing dorsal parts, to northern Nayarit; in southern part of shading to more blackish on the rump; range, broad lateral stripe of buffY or skull relatively arched dorsally . yellowish contrasting with brownish­ · Chaetodipus fal/ax appearing dorsal parts; in northern Si­ 11'. Lateral stripe narrow and pale yellow­ naloa and southern Sonora, lateral stripe ish-gray, or obsolete; upper parts narrow or faint Chaetodipus pernix brownish to pale buffY-yellow and not 6'. Tail with moderate to large crest; not shading to more blackish on rump; skull occurring south ofsouthern Sonoraalong relatively flattened dorsally . the Pacific coast of mainland Mexico; · Chaetodipus spinatus lateral stripe on sides between fore- and 12. Length ofhead andbody averages about hind limbs faint or absent 7 77 to 80 mm; tail heavily crested and 7. Dorsal color dull gray with buffY wash pelage of upper parts relatively coarse (no blackish overwash); buffy lateral or harsh; spines on rump usually mod­ stripe narrow, but distinct; occurring erate in number and easily detected; in only on the Mexican Plateau in San Luis western Texas and Chihuahua, dorsal Potosi and possibly eastern Zacatecas wing of premaxilla extending distinctly ·...... Chaetodipus lineatus beyond posterior tip of nasal . 7'. Dorsal color some shade ofbrownish or · Chaetodipus nelsoni yellowish-gray with faint to strong over­ 12'. Length ofhead and body averaging from wash or admixture ofblackish; ranging about 70 to 75 mm; tail moderately cre­ widely, including northern and western ated and pelage ofupper parts not mark­ San Luis Potosi 8 edly coarse; spines on rump often sparse 8. Occurring only on the Baja California and difficult to detect; in western Texas peninsula and its islands; tail faintly an­ and Chihuahua, dorsal wing ofpremax­ nulated in fresh specimens and only illa terminating only slightly posterior slightly longer than length of head and to tip of nasal " Chaetodipus intermedius body; interoribital breadth averaging 13. Occurring only west of the Colorado greater than 25.8% of occipitonasal River in California and Baja California length, except for some insular popu- ...... " 14 lations Chaetodipus arenarius 13'. Occurring only on the Pacific slopes and 8'. In Baja California, only occurring on the plains ofthe coastal mainland ofMexi- Gulf plains in the northeastern corner, co 15 from San Felipe north; elsewhere rang­ 14. A few, weakly developed bristles on ing widely in the creosote bush deserts rump; length ofear usually less than 10 of the U.S. and Mexico; tail consider­ mm; occurs only in the Cape region of ably longer than length ofhead and body Baja California Sur . and annulated in life; interorbital · Chaetodipus arenarius breadth averaging less than 25% of oc­ 14'. Strong, whitish spines on rump; length cipitonasallength in most populations, ofear usually 10 mm or greater; occurs including those in Baja California .... in California and the central highlands ·...... Chaetodipus penicillatus of northern Baja California . 9. Ear pinna long, usually measuring more ·...... Chaetodipus californicus than 9 mm 13 15. Dorsal wing of premaxilla extending 9'. Ear pinna short, usually measuring less distinctly beyond posterior tip ofnasal; than 9 mm 10 length of mastoid bullae averaging less 10. Occurring only west of the Colorado than 6.0 mm Chaetodipus artus 124 WILLIAMS ET AL.

15'. Dorsal wing of premaxilla terminating bead on frontal; and other diagnostic char­ about even with posterior end ofnasal; acters of the genus Chaetodipus. length of mastoid bullae averaging Remarks. - Burtognathus is monotypic. greater than 6.5 mm . The unique characters ofBurtognathus sug­ ...... Chaetodipus goldmani gest an early divergence and long, indepen­ dent evolution from species ofChaetodipus (Burt, 1936; Hafner and Hafner, 1983; Mer­ Subgenus Burtognathus riam, 1889; Patton et aI., 1981). Burtog­ Hoffmeister, 1986 nathus was named by Hoffmeister (1986) in recognition of William H. Burt, for his pi­ 1986. Burtognathus Hoffmeister, Mammals of oneering work, "Bacula ofNorth American Arizona, Univ. Arizona Press, p. 247, 31 July. Mammals" (Burt, 1960), in which he de­ Type Species. -Perognathus hispidus scribed the baculum of C. hispidus. Al­ Baird, 1858, Mammals, in Repts. Expl. Surv. though the name Burtognathus literally ... , 8(1):421, 14 July. means "Burt jaw" or "Burt face," we be­ Diagnosis. - Body large for genus and tail lieve Dr. Burt, who coined the name "Bai­ relatively short; length oftail slightly shorter ley's pocket-faced fornicator" (Perognathus or about the same as length of head and baileyi fornicatus), would see both the hu­ body, both average about 100-115 mm; mor and honor in his patronym. mass averages from about 35 to 60 g; no crest or long tuft at the end ofthe tail; pelage harsh, with flattened, troughed hairs, cres­ Species Account cent-shaped in cross section; trough ofhairs deep; hairs not differentiated into spines or Chaetodipus hispidus bristles on rump; frontal bone ofskull with strong supraorbital bead; squamosal por­ Diagnosis. -See diagnosis of subgenus. tion ofzygomatic arch usually free ofor only Comparisons. - Chaetodipus hispidus is loosely attached to auditory bulla; supraoc­ distinguished from all other species of cipital does not indent into mastoid bulla; Chaetodipus by its large body size and short, mastoid bulla small; baculum relatively long noncrested tail; length of head and body (14-18 mm) and straight with a unique, tri­ generally ranges from about 95 to 115 mm. lobed tip (two ventral and one dorsal lobes); In body size, C. hispidus is similar only to diploid karyotype of 34 biarmed chromo­ C. baileyi; hispidus differs in being some­ somes. what larger, and having longer, broader in­ Comparisons. - Burtognathus differs from terparietals, smaller mastoid bullae, beaded other genera and subgenera of pocket mice supraorbital ledges, and a short, noncrested in the unique, three-lobed tip ofits baculum tail. (the bacula of others taper to simple tips). Distribution. - Chaetodipus hispidus oc­ From the subgenus Chaetodipus, Burtog­ cupies the Great Plains from south-central nathus differs further in having a supraor­ North Dakota southward, east ofthe Rocky bital bead on the frontal; squamosal portion Mountains and generally west of the Mis­ ofzygomatic arch free ofmeatal part ofbul­ souri River and Ozark Plateau through cen­ la (or at best a loose attachment); smaller tral New Mexico, virtually all ofTexas, east­ bullae; and absence of an indentation into ern and central Chihuahua, extreme mastoid bulla by the supraoccipital. Bur­ northeastern Durango, northeastern Coa­ tognathus can be distinguished from Perog­ huila, northern Nuevo Leon, and south­ nathus by a variety ofcharacters, including central Tamaulipas; on the east, extending larger size, hispid pelage of flattened, to west-central Louisiana; on the west, ex­ troughed hairs, less inflated mastoid and tending across southern New Mexico to tympanic bullae, presence of supraorbital southeastern Arizona and probably extreme TAXONOMY 125 northeastern Sonora; on the south, an ap­ 1894. Perognathus latirostris Rhoads, Amer. parently disjunct population occupies the Nat., 28:185, February. central plateau of Mexico, ranging from 1894. Perognathus conditi J. A. Allen, Bull. southeastern Coahuila and probably south­ Amer. Mus. Nat. Hist., 6:318, 7 November. western Nuevo Leon on the northeast, and Holotype. - Adult female, skin and skull, east-central Durango on the northwest, USNM 94011544, from Trego Co., Kansas; southward through eastern San Luis Potosi, obtained on 17 October 1884 by A. B. Ba­ Guanajuato, to Mexico on the east, and ker. through most ofZacatecas and eastern Jalis­ Measurements ofholotype. - Total length, co on the west (Hall, 1981). 205; length oftail, 105; length ofhind foot, Remarks.-Glass (1947) reviewed the 26; length of ear (crown, dry), 5.5; occipi­ species exclusive ofthe Mexican subspecies, tonasal length, 32.0; basilar length, 26.6; zacatecae, and gave average measurements basilar length ofHensel, 23.7; width across and diagnoses for subspecies. mastoid bullae, 15.7; length ofinterparietal, 4.5; width ofinterparietals, 8.0; least inter­ Chaetodipus hispidus hispidus mastoid distance, 9.0. (Baird, 1858) Distribution. -Occupiesthe northern and western Great Plains and Chihuahuan Des­ 1858. Perognathus hispidus Baird, Mammals, ert area, ranging from south-central North in Repts. Expl. Surv.... ,8(1):421, 14 July. Dakota southward through western and Holotype. -Adult female, skin and skull, central South Dakota and Nebraska, west­ ern Kansas, Oklahoma, and Texas; on the USNM 577/1696, from Charco Escondido, Tamaulipas, Mexico; obtained in 1853 by west, from extreme southeastern Montana, eastern Wyoming and Colorado, central and D. N. Couch. southern New Mexico, southeastern Ari­ Measurements of holotype. - Length of zona, and west-central Chihuahua to ex­ head and body, 79; length of tail (terminal treme north-central Durango (Hall, 1981). portion broken om, 72; length ofhind foot, Also known from a disjunct area near Camp 21.5; length ofear (crown, dry), 4.5; occip­ Verde, Yavapai Co., Arizona. itonasal length (approximated by Merriam Remarks. - Measurements were given by from broken skull), 27.5; width across mas­ Anderson (1972), Armstrong (1972), Cock­ toid bullae, 14.5; length ofinterparietal, 3.8; rum (1952), Hoffmeister (1986), Jones width of interparietals, 7.0; least intermas­ (1964), and Long (1965). Hoffmeister (1986) toid distance, 8.0. compared a sample of 15 specimens from Distribution. - Ranges from near the Arizona with a sample of 14 paradoxus from Oklahoma boundary in north-central and the Great Plains, as well as with other sam­ eastern Texas and from west-central Loui­ ples. Because he found that the Arizona siana southward to northeastern Coahuila sample was distinguishable from the Great and south-central Tamaulipas. Remarks.-Baker (1956) listed means and Plains sample, he believed that conditi was ranges for measurements offour males and a valid subspecies. four females from Coahuila; Osgood (1900) gave average measurements for three spec­ imens. Chaetodipus hispidus spilotus (Merriam, 1889)

Chaetodipus hispidus paradoxus 1889. Perognathus paradoxus spilotus Merriam, (Merriam, 1889) N. Amer. Fauna, 1:25,25 October. 1904. Perognathus hispidus maximus Elliot, 1889. Perognathus paradoxus Merriam, N. Field Columbian Mus., Publ. 87, Zool. Ser., Amer. Fauna, 1:24, 25 October. 3:253. 126 WILLIAMS ET AL.

Holotype. -Adult female, skin only, to, and Mexico; on the west, through most USNM 186514, from Gainesville, Cook [= of Zacatecas and eastern Jalisco. Cooke] Co., Texas; obtained on 8 October Remarks. - Baker (1956) listed measure­ 1886 by G. H. Ragsdale. ments ofone specimen from Coahuila, and Measurements ojholotype. - Total length, Genoways and Jones (1973) gave measure­ 196; length of tail, 95; length of hind foot, ments for an adult male from Zacatecas. 23; length of ear (crown, dry), 5. Distribution. -Ranges from southeastern Nebraska southward through east-central Subgenus Chaetodipus Kansas and eastern and central Oklahoma to extreme north-central Texas along the 1889. Chaetodipus Merriam, N. Amer. Fauna, Oklahoma border. 1:5,25 October. Remarks.-Merriam (1889), while not Type species. - Perognathus [Chaetodi­ designating it as a cotype, characterized cra­ pus] spinatus Merriam, 1889, N. Amer. nial features ofspilotus using USNM 23096, Fauna, 1:5, 25 October. a young adult female from the type locality, Diagnosis. -Characters as for genus ex­ obtained on 24 September 1888 by G. H. cept baculum tapers to a simple, non-lobed Ragsdale; its measurements, listed by Mer­ tip; no supraorbital bead or ridge on frontal; riam (1889), are: occipitonasallength, 31.0; squamosal portion ofzygomatic arch firmly basilar length, 25.2; basilar length of Hen­ attached to meatal part of auditory bulla; sel, 23.0; width across mastoid bullae, 14.5; supraoccipital forms indentation into mas­ length of interparietals, 4.2; width of inter­ toid bulla; diploid number ofchromosomes parietals, 7.3; least intermastoid distance, in so far as is known 36 to 56. 8.6. Cockrum (1952) gave measurements for Comparisons. -See accounts of Burtog­ two males and three females from Kansas. nathus and Perognathus for comparison with those taxa. Remarks. - The subgenus Chaetodipus as Chaetodipus hispidus zacatecae applied here, differs from the subgenus of (Osgood, 1900) Merriam (1889) as it was applied by him and subsequent authorities except Hoff­ 1900. Perognathus hispidus zacatecae Osgood, meister (1986). N. Amer. Fauna, 18:45,20 September. Holotype. - Young adult female, skin and skull, USNM 91877, from Valparaiso, Za­ Species Accounts catecas, Mexico; obtained by Edward A. Goldman on 16 December 1897. Chaetodipus arenarius Measurements ojholotype. - Total length, 211; length oftail, 105; length ofhind foot, Diagnosis. -A small species of Chaeto­ 27.5; occipitonasal length, 30.2; basilar dipus, with a tail that is longer than length length ofHensel, 22.5; interorbital breadth, ofhead and body, small ears, and with rel­ 7.0; width across mastoid bullae, 15.0; length atively soft pelage usually lacking stiffbris­ ofinterparietal, 4.0; width ofinterparietals, tIes or spines; a clear, buffy lateral stripe is 8.0; length of nasals, 12.0. usually absent; length of head and body Distribution. - Found from southeastern ranges from about 66 to 80 mm; length of Coahuila and probably southwestern Nue­ tail ranges from about 70 to 103 mm; length vo Leon on the northeast, and east-central ofear ranges from about 7 to 10 mm. Durango on the northwest, southward Comparisons. - Chaetodipus arenarius is through eastern San Luis Potosi, Guanajua- most similar in general appearance to the TAXONOMY 127 other small, soft-haired species, C. penicil­ tween the karyotypes of dalquesti and C. latus and C. pernix; arenarius differs from arenarius. penicillatus in being smaller, having a less annulated appearing tail in life, and broader skull; pernix differs from arenarius in hav­ Chaetodipus arenarius albescens ing a well developed lateral stripe of clear (Huey, 1926) buff. From other species ofChaetodipus oc­ curring on the baja California Peninsula (c. 1926. Perognathus arenarius albescens Huey, baileyi, californicus, formosus, fallax, and Proc. BioI. Soc. Washington, 39:67, 30 July. spinatus), C. arenarius can be distinguished by its smaller size; it further differs from Holotype. -Adult male, skin and skull, SDSNH 5103, from San Felipe, Baja Cali­ californicus, fallax, and spinatus by lacking fornia, Mexico; obtained by Laurence M. stiff bristles or spines in the pelage of the Huey on 23 March 1926. rump and flanks, although weakly devel­ Measurements ofholotype. - Total length, oped bristles or spines may be found rarely; 170; length of tail, 95; length of hind foot, ofthe sympatric soft-haired species, the hind 22; length of ear (crown), 5; weight, 15 g; foot measures less than 25 mm in arenarius condylobasal length, 23.2; interorbital and greater than 25 mm in baileyi; length breadth, 6.0; length of maxillary toothrow, ofear generally measures greater than 9 mm 3.0; width across mastoid bullae, 12.3; length informosus and less than 9 in arenarius; the of nasals, 8.8. mastoid bullae offormosus project slightly Distribution. - Known from sandy soils beyond the plane of the occipitals at the along the Gulf of California coast in the back ofthe skull, whereas those ofarenarius vicinity of the type locality in northeastern do not extend to the posterior plane of the Baja California. occipitals. Remarks.-Huey (1926) listed measure­ Distribution. -Generallylimited to sandy ments for 10 adults. soils on the Baja California Peninsula, from near De Mara's Well, Laguna Salada, near the international boundary in northeastern Baja California, and San Quintin on the Pa­ Chaetodipus arenarius albulus cific coast on the northwest, southward to (Nelson and Goldman, 1923) near the southern cape in Baja California Sur; not known from the eastern, gulf slope 1923. Perognathus penicillatus albulus Nelson from about El Barril, Baja California south and Goldman, Proc. BioI. Soc. Washington, to near La Paz, Baja California Sur, nor from 36: 159, 1 May. coastal areas south of La Paz (Hall, 1981). Holotype. -Adult male, skin and skull, Remarks. - We have treated Chaetodipus USNM 146864, from Magdalena Island, dalquesti (Roth, 1976) as a synonym of C. Baja California Sur, Mexico; obtained by arenarius arenarius based on studies by M. Edward W. Nelson and Edward A. Gold­ A. Bogan (in litt.), although Bogan's inves­ man on 3 December 1906. tigations left subspecific status of dalquesti Measurements ofholotype. - Total length, unresolved. Presence or absence ofstiffbris­ 155; length of tail, 83; length of hind foot, ties or spines on the rump is not constant 22; greatest length ofskull, 22.8; interorbital within other species ofChaetodipus, such as breadth, 6.1; length of maxillary toothrow, C. intermedius and C. penicillatus (Hoff­ 3.4; width across mastoid bullae, 12.0; length meister and Lee, 1967), and alone does not ofinterparietal, 3.4; width ofinterparietals, indicate specific status for dalquesti. Hafner 7.0; length of nasals, 8.7; zygomatic width, and Hafner (1983) found no differences be- 11.2. 128 WILLIAMS ET AL.

Distribution. -Found only on Magdalena length of skull, 26.1; basilar length, 18.0; Island, Baja California Sur. interorbital breadth, 6.6; length ofmaxillary Remarks.-Nelson and Goldman (1923) toothrow, 3.8; width across mastoid bullae, listed means and ranges of measurements 13.2; length of interparietal, 3.8; width of of nine adults. interparietals, 7.3; length ofnasals, 9.4; zy­ gomatic width, 13.1; length ofdiastema, 6.4. Distribution. - Known only from Santa Chaetodipus arenarius ambiguus Margarita Island, Baja California Sur, off (Nelson and Goldman, 1929) the west coast of the Baja California Pen­ insula. 1929. Perognathus arenarius ambiguus Nelson Remarks. -Osgood (1907) gave averages and Goldman, Proc. BioI. Soc. Washington, and extremes of measurements of nine to­ 42:108,25 March. potypes. Holotype. - Young adult male, skin and skull, USNM 140011, from Yubay, 30 mi Chaetodipus arenarius arenarius SE Calamahue [= Calamajue], 2,000 ft, Baja (Merriam, 1894) California; obtained on 18 September 1905 by Edward W. Nelson and Edward A. Gold­ 1894. Perognathus arenarius Merriam, Proc. man. Calif. Acad. Sci., ser. 2, 4:461, 25 September. Measurements ofholotype. - Total length, 1976. Perognathus dalquesti Roth, J. Mamm., 150; length of tail, 88; length of hind foot, 57:562, 27 August. 22; greatest length ofskull, 22.6; interorbital Holotype. -Adult female, skin and skull, breadth, 6.2; length of maxillary toothrow, CAS 99, from San Jorge, near Comondu, 3.2; width across mastoid bullae, 12.0; length Baja California Sur, Mexico; obtained by ofinterparietal, 2.8; width ofinterparietals, Walter E. Bryant on 17 March 1889. 5.2; length of nasals, 8.3; width of nasals, Measurements of holotype. -[External 2.3. measurements from dry skin] Total length, Distribution. -Occupies sandy soils in the 136; length of tail, 70; length of hind foot, central Baja California Peninsula, from near 20; length ofear (anterior base), 7; greatest Chapala southward to the middle ofthe Viz­ length of skull, 22.5; occipitonasal length, caino Desert in Baja California Sur (Huey, 23.0; basilar length, 15.3; interorbital 1964). breadth, 6.2; width across mastoid bullae, Remarks.-Nelson and Goldman (1929) 12.0; length of interparietal, 3.5; width of listed average and extremes of measure­ interparietals, 6.4; length ofnasals, 8.8; zy­ ments of five adults. gomatic breadth anteriorly, 11.0. Distribution. - Found on sandy soils from near the cape region northward to the south­ Chaetodipus arenarius ammophilus ern part of the Vizcaino Desert on the Pa­ (Osgood, 1907) cific coast ofthe Baja California peninsula, in Baja California Sur. 1907. Perognathus penicillatus ammophilus Os­ Remarks.-Roth (1976:564) character­ good, Proc. BioI. Soc. Washington, 20:20, 23 ized dalquesti as being larger, with a longer, February. darker colored tail than arenarius, and hav­ Holotype. -Adult male, skin and skull, ing "distinct rump spines," similar to C. USNM 146859, from Santa Margarita Is­ intermedius, but "not as stout." Hafner and land, Baja California Sur, Mexico; obtained Hafner (1983) stated that the karyotype of on 20 November 1905 by Edward W. Nel­ dalquesti was identical to that ofC. arenari­ son and Edward A. Goldman. us. Banks (1964) listed means and ranges of Measurements of holotype. -Greatest measurements of 8 males and 12 females TAXONOMY 129 from Todos Santos. Total length of skull, Distribution. - Known from sandy areas listed by Merriam (1894c) differs from oc­ on the western side of Laguna Salada in cipitonasal length reported by Osgood northeastern Baja California. (1900). Remarks.-According to Huey (1939b), this subspecies may range northward as far as a few miles into the state of California, Chaetodipus arenarius helleri although the species has not been taken in (Elliot, 1903) the United States.

1903. Perognathus helleri Elliot, Field Colum­ bian Mus., Pub!. 74, Zoo!' Ser., 3:166, 7 May. Chaetodipus arenarius paralios Holotype. -Adult male, skin and skull, (Huey, 1964) FMNH 10355, from San Quentin [= San Quintin], Baja California, Mexico; obtained 1964. Perognathus arenarius paralios Huey, Trans. San Diego Soc. Nat. Hist., 13:113, 15 by Edmund Heller on 26 July 1902. January. Measurements ofholotype. - Total length, 159; length of tail, 83; length of hind foot, Holotype. - Adult male, skin and skull, 20.5; length of ear, 8; occipitonasallength, SDSNH 15542, from [El] Barril, lat. 23.0; basilar length of Hensel, 14.0; inter­ 28°20'N, on the Gulf of California, Baja orbital breadth, 6.0; length of maxillary California, Mexico; obtained by Laurence toothrow, 3.0; width across mastoid bullae, M. Huey on 23 March 1947. 11.5; length ofnasals, 7.5; width ofrostrum, Measurements ofholotype. - Total length, 4.0; zygomatic breadth, 11.5; greatest width 150; length of tail, 81; length of hind foot, of brain case, 10.5; palatal arch to alveoli 22; length ofear (crown), 5; greatest length ofincisors, 8.5. of skull, 24.0; interorbital breadth, 6.3; Distribution. - Known from sandy soils length of maxillary toothrow, 2.9; width on the San Quintin Plain on the Pacific coast across mastoid bullae, 13.0; length ofnasals, of northern Baja California. 8.9. Remarks. - Elliot (1903) listed extremes Distribution. -Occurs on sandy soils on for 15 specimens. the GulfofCalifornia slope from near Bahia Los Angeles southward to near the type lo­ cality at £1 Barril, Baja California. Chaetodipus arenarius mexicalis Remarks. -No other measurements are (Huey, 1939) available for this subspecies.

1939. Perognathus arenarius mexicalis Huey, Trans. San Diego Soc. Nat. Hist., 9:57,31 Au­ Chaetodipus arenarius sabulosus gust. (Huey, 1964) Holotype. -Adult female, skin and skull, SDSNH 12127 from Los Muertos Canyon 1964. Perognathus arenarius sabulosus Huey, fan at Gaskill's Tank, near Laguna Salada, Trans. San Diego Soc. Nat. Hist., 13:114, 15 January. lat. 32°27'N, long. 115°53'W, Baja Califor­ nia, Mexico; obtained on 23 November 1936 Holotype. - Adult male, skin and skull, by Laurence M. Huey. SDSNH 5300, from mainland on S side Measurements ofholotype. - Total length, Scammon's Lagoon, Baja California Sur, 176; length oftail, 102; length ofhind foot, Mexico; obtained by Laurence M. Huey on 23; length ofear (crown), 5; greatest length 22 May 1926. ofskull 23.7; interorbital breadth, 6.1; length Measurements ofholotype. - Total length, of maxillary toothrow, 3.2; width across 165; length of tail, 91; length of hind foot, mastoid bullae, 12.0; length of nasals, 8.5. 21; length ofear (crown), 5; greatest length 130 WILLIAMS ET AL. of skull, 23.5; interorbital breadth, 6.1; nia Sur, Mexico on 16 February 1906 by length of maxillary toothrow, 2.9; width Edward W. Nelson and Edward A. Gold­ across mastoid bullae, 12.3; length ofnasals, man. 8.8. Measurements ofholotype. - Total length, Distribution. - Found on sandy ground 167; length of tail, 95; length of hind foot, from Bahia Santa Rosalia, Baja California, 22; greatest length ofskull, 24.8; interorbital southward through the western Vizcaino breadth, 6.3; length of maxillary toothrow, Desert to the vicinity of Laguna San Igna­ 3.2; width across mastoid bullae, 12.6; length cio. ofinterparietal, 3.7; width ofinterparietals, Remarks. - There are no published mea­ 6.4; length of nasals, 9.4; width of nasals, surements other than those ofthe holotype. 2.3. Distribution. - Known from sandy areas in the desert basin in the vicinity ofLa Paz, Chaetodipus arenarius siccus on the Gulf coast of Baja California Sur. (Osgood, 1907) Remarks. - Nelson and Goldman (1929) listed means and extremes for external mea­ 1907. Perognathus penicillatus siccus Osgood, surements of 10 adult topotypes. Banks Proc. Bioi. Soc. Washington, 20:20, 23 Feb­ (1964) listed means and ranges ofmeasure­ ruary. ments for 13 specimens ofeach sex. Holotype. -Adult male, skin and skull, USNM 146890, from Ceralbo [= Cerralvo] Chaetodipus artus Island, Baja California Sur, Mexico; ob­ tained by Edward W. Nelson and Edward Diagnosis. -A species of Chaetodipus of A. Goldman on 13 February 1906. medium-large size, with weakly developed Measurements of holotype. -Greatest spines present or absent on the rump, rel­ length of skull, 25.9; basilar length, 17.7; atively large, rounded ears, and a tail of interorbital breadth, 6.6; length ofmaxillary about average length for the genus; skull toothrow, 4.0; width across mastoid bullae, with broad, nearly elliptically-shaped inter­ 13.8; length of interparietal, 3.6; width of parietals and small mastoid bullae; baculum interparieta1s, 7.2; length ofnasals, 8.9; zy­ relatively long with a strongly curved tip. gomatic width, 12.6; length ofdiastema, 6.0. Comparisons. - Chaetodipus artus shares Distribution. - Known only from Cerral­ parts of its geographic range with C. gold­ vo Island in the Gulf of California, Baja mani, to which it is most similar in ap­ California Sur. pearance, and with C. baileyi and C. pernix. Remarks. - Osgood (1907) listed means No single character ofskin or skull will sep­ and extremes of external measurement of arate all artus from all goldmani (Anderson, 10 topotypes. Banks (1964) gave means and 1964), but C. artus typically differs from C. ranges of measurements for 18 males and goldmani in being slightly smaller; having 17 females. a less hairy tail; fewer and more weakly­ developed rump spines; darker color, in­ cluding the ear pinnae; shorter, more Chaetodipus arenarius sublucidus strongly curved baculum; smaller, narrower (Nelson and Goldman, 1929) skull; smaller mastoid bullae; and in having the ascending process ofthe premaxillae ex­ 1929. Perognathus arenarius sublucidus Nelson tending posteriorly beyond the end of the and Goldman, Proc. Bioi. Soc. Washington, nasals a distance equal to or greater than 42:109,25 March. the least breadth ofone nasal bone (in gold­ Holotype. -Adult male, skin and skull, mani the premaxillae barely extends pos­ USNM 146896, from La Paz, Baja Califor- teriorly beyond the end of the nasal). C. TAXONOMY 131 pernix is considerably smaller than artus and erate length, and soft, spineless pelage; skull lacks any spines on the rump; interorbital large and robust with narrow interparietals breadth of pernix is less than 5.8 mm, and relatively large mastoid bullae that pro­ whereas it is greater than 5.8 in artus (Hall, ject slightly posteriad beyond the plane of 1981). Total length ofadult C. baileyi gen­ the occipitals; width ofinterparietals about erally exceeds 200 mm, whereas artus rarely equal to or less than interorbital breadth; measures more than 200 mm; length ofhind dorsal and lateral color grayish washed with foot is 26 mm or more in baileyi and rarely yellowish. exceeds 25.5 mm in artus (Anderson, 1964; Comparisons. - Chaetodipus baileyi is ex­ Hall, 1981); the mastoid bullae of baileyi ceeded in length of head and body only by are relatively large; and baileyi always lacks some C. hispidus, from which it can be dis­ spines in the pelage. tinguished by its grayish color and its long, Distribution. - Found along the Pacific crested tail (the tail ofhispidus is not crested coastal slope of western Mexico, from the and is shorter than length ofhead and body); barrancas of southwestern Chihuahua and it can be distinguished from all other Chae­ northwestern Durango westward to south­ todipus species lacking spines in the pelage ern Sonora and southward to northern Na­ by its larger size (average length ofhead and yarit (Hall, 1981). body greater than 90 mm), longer hind foot Remarks. -Refer to Anderson (1964) for (generally exceeding 26 mm), larger mastoid characteristics distinguishing C. artus and C. goldmani. Anderson (1972) listed means bullae, and grayish rather than brownish and ranges of measurements for 13 speci­ color of upper parts (discounting the yel­ mens from Chihuahua. lowish tinge present on both the grayish­ and brownish-colored species). Distribution. - Chaetodipus baileyi occu­ Chaetodipus artus (Osgood, 1900) pies rocky hillsides and brushy, desert as­ sociations from extreme southwestern New 1900. Perognathus artus Osgood, N. Amer. Fau­ Mexico westward across southern Arizona na, 18:55,20 September. to extreme southeastern California, and Holotype. -Adult female, skin and skull, southward throughout most ofthe Baja Cal­ USNM 96298, from Batopilas, Chihuahua, ifornia peninsula, and along the lowlands of Mexico; obtained by Edward A. Goldman Sonora to northwestern Sinaloa (Hall, 1981). on 6 October 1898. Remarks. - Patton et al. (1981) reviewed Measurements ofholotype. - Total length, the patterns ofbiochemical and karyotypic 197; length oftail, 109; length ofhind foot, variation in C. baileyi, noting that the pop­ 24; greatest length of skull, 26.30; width ulations west ofthe Colorado River in Cal­ across bullae, 12.55; breadth across maxil­ ifornia and the Baja California peninsula, lary arches, 11.95; nasal length, 10.10; in­ and east ofthe Colorado River in Arizona, terorbital breadth, 5.95. New Mexico and mainland Mexico formed Remarks. -Osgood listed means for two relatively distinct units. Both geograph­ measurements of three specimens. Ander­ ic units showed relatively high uniformity son (1964) gave means and ranges of mea­ in biochemical and karyotypic traits (Pat­ surements for 14 samples of mixed ages. ton, 1977) within their units and reduced levels ofsimilarity between units. They not­ ed that the biochemical differentiation was Chaetodipus baileyi generally unrelated to the current intraspe­ cific taxonomy based on structural features. Diagnosis. -A large species of Chaeto­ The species has never had a comprehensive dipus with a strongly crested tail of mod- systematic review. 132 WILLIAMS ET AL.

Chaetodipus baileyi baileyi 25; greatest length ofskull, 29.0; interorbital (Merriam, 1894) breadth, 6.9; length of maxillary toothrow, 4.5; width across mastoid bullae, 14.3; length 1894. Perognathus baileyi Merriam, Proc. Acad. ofinterparietal, 3.4; width ofinterparietals, Nat. Sci. Philadelphia, 46:262, 27 September. 6.1; length of nasals, 10.2; width of nasals, 1928. Perognathus baileyi domensis Goldman, 2.9; zygomatic breadth, 15.5. Proc. BioI. Soc. Washington, 41:204, 18 De­ cember. Distribution. -Found on the Pacific slope of the cape region of Baja California Sur, Holotype. -Adult female, skin and skull, from near San Jose de Comondu on the USNM 17838/24775, from Magdalena, So­ north to near Todos Santos on the south. nora, Mexico; obtained by Vernon Bailey Remarks. - Nelson and Goldman (1930) on 3 November 1889. listed measurements for three adults; Burt Measurements ofholotype. - Total length, (1932) gave ranges and means for measure­ 210; length oftail, 122; length ofhind foot, ments of six specimens. 27; length of ear (anterior base, dry), 11; greatest length of skull, 29.5; basal length, Chaetodipus baileyi fornicatus 24.0; basilar length of Hensel, 20.5; length (Burt, 1932) of nasals, 11.5; greatest zygomatic breadth (posteriorly), 15.5. 1932. Perognathus baileyifornicatus Burt, Trans. Distribution. - Ranges from the Colorado San Diego Soc. Nat. Rist., 7: 164, 31 October. river in southwestern Arizona, the southern Holotype. -Adult male, skin and skull, edge ofthe Mogollon Plateau in central Ar­ CIT 50289, from Monserrate Island, lat. izona, and southwestern New Mexico 25°38'N, long. 11lo02'W, Gulf of Califor­ southward in the foothills ofthe Sierra Ma­ nia, Baja California Sur, Mexico; obtained dre Occidental and the coastal plain of So­ by William H. Burt on 16 December 1931. nora to northern Sinaloa (Hall, 1981; Hoff­ Measurements ofholotype. - Total length, meister, 1986). 202; length oftail, 108; length ofhind foot, Remarks. -Hoffmeister (1986) listed sta­ 26; length of ear (crown), 7, (notch, dry), tistics for measurements of 13 samples from 8.9; greatest length of skull, 29.1; basal Arizona. Merriam (1894b) listed average length, 24.6; interorbital breadth, 6.9; length external measurements for eight topotypes. of maxillary toothrow, 4.2; length of mas­ Hoffmeister (1986) considered domensis to toid bulla, 8.8; width across mastoid bullae, be indistinguishable from baileyi; Goldman 14.3; length of interparietal, 3.8; width of (1928) noted that domensis mainly differed interparietals, 6.6; length of nasals, 11.1. from baileyi in its more palely-colored up­ Distribution.-Known only from Mon­ per parts. serrate Island, GulfofCalifornia, Baja Cal­ ifornia Sur. Chaetodipus baileyi extimus Remarks. - Burt (1932) gave means and (Nelson and Goldman, 1930) ranges for measurements of 10 specimens. The holotype is now housed in the UCLA 1930. Perognathus baileyi extimus Nelson and collections. Goldman, J. Washington Acad. Sci., 20:223, 19 June. Chaetodipus baileyi hueyi Holotype. - Adult female, skin and skull, (Nelson and Goldman, 1929) USNM 146672, from Tres Pachitas, 700 ft, 36 mi S La Paz, Baja California Sur, Mexi­ 1929. Perognathus baileyi hueyi Nelson and co; obtained by Edward W. Nelson and Ed­ Goldman, Proc. BioI. Soc. Washington, 42: 106, 25 March. ward A. Goldman on 25 December 1905. Measurements ofholotype. - Total length, Holotype. - Young adult female, skin and 198; length oftail, 107; length ofhind foot, skull, SDSNH 5220, from San Felipe, TAXONOMY 133 northeastern Baja California, Mexico; ob­ Holotype. -Adult male, skin and skull, tained by Laurence M. Hueyon 17 April SDSNH 14470, from San Borja Mission, 1926. near lat. 28°45'N, Baja California, Mexico; Measurements ofholotype. - Total length, obtained by Laurence M. Huey on 13 Oc­ 196; length oftail, 106; length ofhind foot, tober 1941. 24; greatest length ofskull, 27.5; interorbital Measurements ofholotype. - Total length, breadth, 6.4; length of maxillary toothrow, 212; length oftail, 121; length ofhind foot, 3.7; width across mastoid bullae, 14.3; length 25; length ofear (crown), 6; greatest length ofinterparietal, 4.4; width ofinterparietals, of skull, 29.7; interorbital breadth, 7.0; 5.7; length of nasals, 10.5; width of nasals, length of maxillary toothrow, 4.1; width 2.5; zygomatic breadth, 14.5. across mastoid bullae, 15.6; length ofnasals, Distribution. -Found on the desert slopes 7.0. of the coastal mountains of southern San Distribution. - Ranges over the middle Diego Co., California eastward to the Col­ portion ofthe Baja California peninsula from orado River and southward in Baja Cali­ Onyx, near 1at. 30oN, in Baja California, fornia to the vicinity of San Felipe on the southward to Bahia Concepcion, Baja Cal­ Gulf of California. ifornia Sur. Remarks. -No other measurements are Remarks. - There are no published mea­ available. surements other than for the ho10type.

Chaetodipus baileyi rudinoris Chaetodipus baileyi insularis (Elliot, 1903) (Townsend, 1912) 1903. Perognathus baileyi rudinoris Elliot, Field 1912. Perognathus baileyi insularis Townsend, Columbian Mus., Publ. 74, Zool. Ser., 3: 167, Bull. Amer. Mus. Nat. Hist., 31:122,14 June. 7 May. Holotype. - Male, skin and skull, USNM 1903. Perognathus knekus Elliot, Field Colum­ 198410, from Tiburon Island, Gulf of Cal­ bian Mus., Publ. 74, Zool. Ser., 3:169, 7 May. ifornia, Sonora, Mexico; obtained on 13 Holotype. - Adult male, skin and skull, April 1911 by H. E. Anthony. FMNH 10329, from San Quentin [= San Measurements ofholotype. - Total length, Quintin], Baja California, Mexico; obtained 212; length oftail, 118; length ofhind foot, by Edmund Heller, on 24 July 1902. 27; greatest length of skull, 28.50; width Measurements ofholotype. - Total length, across bullae, 15.10; breadth across maxil­ 232; length oftail, 128; length ofhind foot, lary arches, 13.15; nasal length, 11.25; in­ 27; length ofear, 11.5; occipitonasallength, terorbital breadth, 6.55. 31.0; basal length of Hensel, 22.0; interor­ Distribution. - Known only from Tibu­ bital breadth, 6.5; length ofmaxillary tooth­ ron Island, GulfofCalifornia, Sonora, Me­ row, 4.0; width across mastoid bullae, 15.0; xico. width ofinterparietals, 6.5; length ofnasals, Remarks. - Townsend (1912) listed av­ 10.0; width ofnasals (anterior), 3; width of erage external measurements ofthree spec­ rostrum, 4.0; zygomatic breadth, 16.0; dis­ imens; Burt (1932) listed means and ex­ tance from palatal arch to alveolus of in­ tremes ofmeasurements ofeight specimens. cisor, 12.0; greatest width of basioccipital between bullae, 5.0. Distribution. - Found on the coastal plain Chaetodipus baileyi mesidios around San Quintin and the slopes of the (Huey, 1964) southern foothills of the Sierra San Pedro Martir, southward to the region of E1 Mar­ 1964. Perognathus baileyi mesidios Huey, Trans. mol and San Fernando Mission, in Baja Cal­ San Diego Soc. Nat. Hist., 13: 112, 15 January. ifornia. 134 WILLIAMS ET AL.

Remarks. - Except for the holotypes of western slope of the Sierra Nevada and rudinoris and knekus, no measurements thence northward to Placer Co. (Hall, 1981). have been published. Remarks. - Chaetodipus califomicus has never been reviewed systematically; little information is available on structural and Chaetodipus califomicus biochemical variation, and the published records ofoccurrence suggest errors in tax­ Diagnosis. -A Chaetodipus species of onomic assignment to subspecies. medium-large size, with a relatively long, strongly crested tail, numerous stiff spines on the rump and flanks, and a relatively long Chaetodipus califomicus bensoni ear pinna; mastoid bulla relatively small and (von Bloeker, 1938) interparietal broad; ratio of lengths of tail to head and body ranges from about 1.15 1938. Perognathus califamicus bensani von to 1.55; length ofear usually greater than 9 Bloeker, Proc. BioI. Soc. Washington, 51:197, 23 December. mm (range 9-14), and width of interpari­ etals usually greater than 8.1 mm. Holotype. - Subadult male, skin and skull, Comparisons. - Chaetodipus califomicus MVZ 81579, from Stonewall Creek, 1,300 is most similar in general appearance to oth­ ft, 6.3 mi NE Soledad, Monterey Co., Cal­ er medium-sized species ofChaetodipus with ifornia; obtained on 16 June 1937 by Jack spines on the rump; it can be distinguished C. von Bloeker, Jr. from C. fallax by its longer ears (the length Measurements ofholotype. - Total length, of the ear of fallax usually is less than 9 197; length oftail, 107; length ofhind foot, mm), slightly larger size, relatively longer 29; length ofear, 12; greatest length ofskull, tail, and broader interparietal; from C. spi­ 26.7; interorbital breadth, 6.5; length of natus, C. califomicus differs in having fewer mastoid bulla, 7.9; width across mastoid spines, a conspicuous stripe ofbuffy-colored bullae, 13.85; length of interparietal, 4.0; fur on the sides (lateral stripe), a relatively width ofinterparietals, 7.6; length ofnasals, longer tail, longer ear (length 8.5 mm or less 10.3; width of rostrum, 4.7. in spinatus), and wider interparietals; cali­ Distribution. -Occupies chaparral and fomicus is larger, has a relatively longer tail, arid shrub communities in the Diablo Range larger ears, and more spines on the rump of west-central California, from eastern than the allopatric species, C. intermedius Stanislaus Co., on the north, southward to and C. nelsoni; califomicus differs from the the Temblor Range in Kern and San Luis allopatric species, C. goldmani, in having Obispo counties; ranging on the west to the wider parietals, a longer tail and, in typical Salinas Valley, Monterey Co. individuals, more spines on the rump; from Remarks. - Means and ranges of mea­ the allopatric species, C. artus, califomicus surements of10 adults and 5 subadults were is distinguished by its longer tail and greater listed by von Bloeker (1938). number of spines. Distribution. -Occurs mostly in chapar­ ral and less often in annual grassland and Chaetodipus califomicus bemardinus desert shrub communities ofcoastal south­ (Benson, 1930) western North America, ranging from south of San Francisco Bay southward in the 1930. Peragnathus califamicus bemardinus coastal ranges west ofthe Central Valley to Benson, Univ. California PubI. ZooI., 32:449, 6 September. the Sierra San Pedro Martir, Baja Califor­ nia, Mexico; in California, also ranges east­ Holotype. -Adult male, skin and skull, ward in the Transverse Range to the south- MVZ 44094, from 2 mi E Strawberry Peak, TAXONOMY 135

5,750 ft, San Bernardino Mountains, San type; Osgood (1900) gave measurements for Bernardino Co., California; obtained on 19 five adults. September 1920 by Laurence M. Hueyand Donald R. Dickey. Measurements ofholotype. - Total length, Chaetodipus californicus dispar 227; length oftail, 131; length ofhind foot, (Osgood, 1900) 28; length ofear, 12; greatest length ofskull, 28.65; interorbital breadth, 7.15; length of 1900. Perognathus cali/ornicus dispar Osgood, mastoid bulla, 8.3; width across mastoid N. Amer. Fauna, 18:58,20 September. bullae, 13.3; length of interparietal, 3.95; width ofinterparietals, 7.5; length ofnasals, Holotype. - Adult male, skin and skull, 11.35; width of rostrum, 4.8. USNM 32116/43928, from Carpenteria, Distribution. - Known from the slopes of Santa Barbara Co., California; obtained by the San Gabriel and San Bernardino Moun­ Edward W. Nelson on 19 December 1891. tains in Los Angeles and San Bernardino Measurements ofholotype. - Total length, counties, California. 218; length oftail, 120; length ofhind foot, Remarks. -Benson (1930) listed means 27; greatest length of skull, 28.40; width and ranges for measurements ofeight spec­ across bullae, 13.95; breadth across maxil­ lary arches, 12.70; nasal length, 11.80; in­ imens. terorbital breadth, 6.50. Distribution. -Occupies coastal chapar­ Chaetodipus californicus californicus ral communities in California, from San Luis (Merriam, 1889) Obispo, San Luis Obispo Co., California, southward to Santa Monica, Los Angeles 1889. Perognathus cali/ornicus Merriam, N. Co. Amer. Fauna, 1:26, 25 October. Remarks. - The population ranging along 1889. PerognathusarmatusMerriam, N. Amer. the western Sierra Nevada foothills between Fauna, 1:27, 25 October. Placer and Fresno counties (Hall, 1981) is Holotype. -Adult female, skin and skull, excluded here from dispar on geographic USNM 186506, from Berkeley, Alameda grounds. Benson (1930) gave statistics for Co., California; obtained on 8 November measurements of 15 specimens. 1888 by T. S. Palmer and Charles A. Keeler. Measurements ofholotype. - Total length, 186; length of hind foot, 24; length of ear Chaetodipus californicus femoralis (crown), 6; occipitonasallength, 26.7; bas­ (J. A. Allen, 1891) ilar length, 20.5; basilar length of Hensel, 18.1; width across mastoid bullae, 13.4; 1891. Perognathus (Chaetodipus)femoralis J. A. Allen, Bull. Amer. Mus. Nat. Hist., 3:281, 30 length of interparietal, 4.3; width of inter­ June. parieta1s, 8.5; least intermastoid breadth, 8.7. Holotype. - Adult male, skin and skull, Distribution. -Occurs in the hills around AMNH 3386/2659, from Dulzura, San Di­ San Francisco Bay, California, from the San ego Co., California; obtained on 12 Feb­ Francisco peninsula on the northwest and ruary 1891 by C. H. Marsh. Mt. Diablo on the northeast, southward to Measurements ofholotype. - Total length, Portola, San Mateo Co., and Gilroy, Santa 241.3; length of tail, 152.4; length of hind Clara Co. foot (dry), 27.5; length of ear, 9.5; greatest Remarks.-Merriam (1889) listed mea­ length of skull, 27.4; interorbital breadth, surements for two specimens (holotype of 6.7; length ofmaxillary toothrow, 3.8; width armatus included) in addition to the holo- across mastoid bullae, 13.6; length ofnasals, 136 WILLIAMS ET AL.

10.0; zygomatic breadth, 14.0; length ofros­ ifornia, Mexico; obtained by Edmund Hell­ trum, 11.5. er on 6 October 1902. Distribution. -Occupies chaparral com­ Measurements ojholotype. - Total length, munities on the Pacific slope in San Diego 232; length oftail, 136; length ofhind foot, Co., California, ranging southward in the 27; length of ear, 14; occipitonasallength, central mountain range (Hanson Laguna 27.0; basilar length of Hensel, 18.0; inter­ Mountains) of northern Baja California to orbital breadth, 6.0; width across mastoid the northwestern slopes of the Sierra San bullae, 13.0; length ofnasals, 10.0; width of Pedro Martir. nasals (anterior), 3.0; posterior width ofna­ Remarks. -Benson (1930) listed mea­ sals, 2.0; width of rostrum, 5; zygomatic surements of 20 specimens. breadth, 13.0; distance from palatal arch to alveolus of incisor, 10.0. Distribution. -Occurs in the Sierra San Chaetodipus califomicus marinensis Pedro Martir, Baja California. (von Bloeker, 1938) Remarks. -No other published measure­ ments are available for this subspecies. The 1938. Perognathus californicus marinensis von description of color suggests an animal in B1oeker, Proc. BioI. Soc. Washington, 51:199, juvenile pelage; characterization ofthe sub­ 23 December. species was primarily on the basis of its Holotype. -Adult male, skin and skull, grayer color compared to Jemoralis. MVZ 81550, from Indian Harbor, 50 ft, 1.5 mi S Marina, Monterey Co., California; ob­ tained by J. C. von Bloeker, Jr., on 27 May Chaetodipus califamicus ochrus 1937. (Osgood, 1904) Measurements ojholotype. - Total length, 203; length oftail, 114; length ofhind foot, 1904. Perognathus californicus ochrus Osgood, 26; length ofear, 14; greatest length ofskull, Proc. BioI. Soc. Washington, 17:128,9 June. 28.1; interorbital breadth, 6.6; length of Holotype. - Young adult female, skin and mastoid bulla, 7.9; width across mastoid skull, USNM 130348, from Santiago bullae, 13.5; length of interparietal, 3.8; Springs, 16 mi SW McKittrick, Kern Co., width ofinterparietals, 8.1; length ofnasals, California; obtained on 30 July 1903 by Lu­ 11.3; width of rostrum, 5.15. ther J. Goldman. Distribution. -Occupies coastal chapar­ Measurements ojholotype. - Total length, ral communities in California, from the 196; length oftail, 117; length ofhind foot, north end of the Gabilan Range and Mon­ 25; greatest length of skull, 27.05; width terey Bay, southward to Morro Bay. across bullae, 13.50; breadth across maxil­ Remarks.-Means and ranges of mea­ lary arches, 12.45; nasal length, 10.15; in­ surements for 12 adult and subadult spec­ terorbital breadth, 6.15. imens were listed by von Bloeker (1938). Distribution. - Found in desert shrub, chaparral, annual grassland, oak woodland, and pine communities of the Transverse Chaetodipus califamicus mesopolius ranges and southeastern slopes ofthe south­ (Elliot, 1903) ern Sierra Nevada, California; ranging from the southern Temblor Range and Carrizo 1903. Perognathus femoralis mesopolius Elliot, Plain, San Luis Obispo Co., south and east­ Field Columbian Mus., PubI. 74, ZooI. Ser., ward to eastern Tulare Co. on the desert 3:168,7 May. slope of the Tehachapi Mountains and Si­ Holotype. -Adult female, skin and skull, erra Nevada, and northward on the western FMNH 10374, from Penon [= Piiion], 5,000 slope of the Sierra Nevada to Placer Co. ft, San Pedro Martir Mountains, Baja Cal- Remarks. -Osgood (1904) listed statis- TAXONOMY 137 tics for external measurements of 10 young subspecies of C. fallax herein, based pri­ adult topotypes. There are no published marily on the work of C. L. Blount (in litt. measurements for skulls ofochrus. The sub­ and pers. comm.). species was characterized entirely on the ba­ sis ofcolor. Chaetodipus fallax anthonyi (Osgood, 1900) Chaetodipus fallax 1900. Perognathus anthonyi Osgood, N. Amer. Diagnosis. -A medium-sized species of Fauna, 18:56,20 September. Chaetodipus with relatively coarse pelage, a Holotype. -Adult female, skin and skull, moderate number ofspines on the rump, a USNM 81058, from South Bay, Cerros [= clear, buffy lateral stripe, a relatively short Cedros] Island, Baja California, Mexico; ear pinna, and a long, crested tail; mastoid obtained on 29 July 1896 by A. W. An­ bullae relatively large; length of head and thony. body ranging from about 80 to 90 mm; Measurements ofholotype. - Total length, length of tail ranging from about 88 to 120 168; length of tail, 92; length of hind foot, mm; width ofinterparietals usually less than 23.5; occipitonasa1 length, 25.4; basilar 8.0mm. length ofHensel, 17.4; interorbital breadth, Comparisons. - Chaetodipus fallax is 6.0; width across mastoid bullae, 12.9; length most similar to C. califomicus in general ofinterparietal, 2.6; width ofinterparieta1s, appearance (see account ofthe latter species 5.8; length of nasals, 10.2. for distinguishing characteristics), and also Distribution. - Known only from Isla resembles other species ofChaetodipus with Cedros, Pacific Ocean, Baja California. spines on the rump. From C. spinatus, fal­ Remarks. - C. fallax anthonyi is charac­ lax differs in having fewer spines (spines on terized as being smaller than typical fallax, the flanks are less numerous and conspic­ but is similar in size to the subspecies, C. f uous), and a more conspicuous lateral stripe inopinus, on the mainland opposite Cedros of clear buff. Other species of Chaetodipus Island. with spines in the pelage (artus, goldmani, intermedius, and nelsoni) have geographic ranges allopatric tofallax, andgenerally have Chaetodipus fallax fallax fewer and more weakly-developed spines; (Merriam, 1889) the ear pinnae of artus and goldmani are longer (generally greater than 9 mm); typical 1889. Perognathus fallax Merriam, N. Amer. Fauna, 1: 19, 25 October. specimens offallax are larger than typical specimens of intermedius and nelsoni. Holotype. - Adult male, skin and skull, Distribution. -Found mainly on the Pa­ USNM 15889/22684, from San Bernardino cific slopes and the basins of southwestern [Reche Canyon, 3 mi SE Colton, 1,250 £1], California and Baja California, but also oc­ San Bernardino Co., California; obtained by cupies the desert slopes ofthe southern Cal­ Frank Stephens on 21 April 1887. ifornia mountain chain; ranges from Oro Measurements ofholotype. - Total length, Grande and Twenty-nine Palms, San Ber­ 183; length oftail, 104; length of hind foot nardino Co., in the Mojave Desert, south­ (dry), 24.0; length of ear (crown, dry), 6; ward and westward to the Sierra Vizcaino occipitonasal length, 26.7; basilar length, in extreme northwestern Baja California Sur; 20.8; basilar length of Hensel, 18.7; width also occurs on Cedros Island off the Pacific across mastoid bullae, 13.8; length ofinter­ coast of Baja California. parietal, 3.7; width of interparietals, 8.1; Remarks.-Huey (1 960a) provided brief least intermastoid distance, 8.4. discussions of the subspecies in Baja Cali­ Distribution. -Occupies the basins and fornia. Chaetodipus anthonyi is treated as a slopes on the Pacific side ofthe mountains 138 WILLIAMS ET AL. ofsouthern California, from the base ofthe Measurements ofholotype. - Total length, San Bernardino Mountains, Los Angeles 191; length oftail, 105; length ofhind foot, Co., southward to Ensanada, Baja Califor­ 24; length ofear, 6; greatest length ofskull, nia along the coast, and inland to the north­ 26.7; interorbital breadth, 6.9; length of ern end ofthe Sierra San Pedro Martir near maxillary toothrow, 3.6; width across mas­ Cerro San Matias, Baja California. toid bullae, 14.7; length of nasals, 9.8. Remarks. -Osgood (1900) gave average Distribution. - Found primarily on the external measurements for six individuals coastal mesas ofBaja California, from south and average cranial measurements for three of Santo Tomas southward to El Rosario. individuals. Remarks. - No measurements have been published for specimens other than the ho­ lotype. Chaetodipus fallax inopinus (Nelson and Goldman, 1929)

1929. Perognathus fallax inopinus Nelson and Chaetodipus fallax pallidus Goldman, Proc. BioI. Soc. Washington, 42:110, (Mearns, 1901) 25 March. 1901. Perognathusfallax pallidus Mearns, Proc. Holotype. -Adult male, skin and skull, Biol. Soc. Washington, 14:135,9 August. USNM 81059, from Turtle Bay [= Bahia Tortugas; also known as Bahia San Barto­ Holotype. - Adult female, skin and skull, lome], Baja California Sur, Mexico; ob­ USNM 61007, from Mountain Spring, half­ tained by A. W. Anthony on 1 August 1896. way up E slope ofCoast Range Mountains, Measurements ofholotype. - Total length, on the Mexican Boundary Line, in San Di­ 180; length oftail, 104; length of hind foot ego Co., California; obtained on 16 May (dry), 23; greatest length of skull, 25.5; in­ 1894 by Edgar A. Mearns. terorbital breadth, 6.2; length of maxillary Measurements ofholotype. - Total length, toothrow, 3.5; width across mastoid bullae, 188, length of tail, 98; length of hind foot, 13.2; length of interparietal, 4.3; width of 23.7; length ofear, 7; greatest length ofskull, interparietals, 6.6; length ofnasals (median 27.20; width across bullae, 13.85; breadth line), 9.3; width of nasals, 2.5. across maxillary arches, 12.70; nasal length, Distribution. - Found along the Pacific 10.40; interorbital breadth, 6.65. coast ofthe Baja California Peninsula, from Distribution. -Southern margin of the near Punta San Carlos and Desembarcadero Mojave Desert, California, along the north­ Santa Catarina, Baja California, southward ern slopes ofthe San Bernardino Mountains to Bahia Tortugas, Baja California Sur. and the western edge ofthe Colorado Des­ Remarks. - Other than those of the ho­ ert, south to the Mexican boundary. lotype, there are no published measure­ Remarks. - Mearns (1901) listed means ments for inopinus. and ranges ofexternal measurements for six adult females, and characterized pallidus as fallaxfallax Chaetodipus fallax majusculus being exactly like C. in size and (Huey, 1960) cranial characters, the only difference being its lighter gray color. 1960. Perognathus fallax majusculus Huey, Trans. San Diego Soc. Nat. Hist., 12:418, 1 February. Chaetodipus fallax xerotrophicus (Huey, 1960) Holotype. - Adult male, skin and skull, SDSNH 15952, from San Quintin, Baja 1960. Perognathus fallax xerotrophicus Huey, California, Mexico; obtained by Laurence Trans. San Diego Soc. Nat. Hist., 12:419. 1 M. Huey on 29 June 1947. February. TAXONOMY 139

Holotype. - Adult male, skin and skull, es to formosus and differ in size and ap­ SDSNH 8310, from 2 mi NE Chapala, Baja pearance (see accounts of those species for California, Mexico; obtained on 15 October details). 1930 by Laurence M. Huey. Distribution. -Occupies desert slopes of Measurements ofholotype. - Total length, the middle and southern Great Basin, Mo­ 210; length oftail, 127; length ofhind foot, jave and Colorado deserts, and the Gulf 24; length ofear, 6; greatest length ofskull, coastal slope of the Baja California penin­ 26.8; interorbital breadth, 6.2; length of sula. Ranges on the northwest from the maxillary toothrow, 3.7; width across mas­ Honey Lake Valley in extreme eastern Las­ toid bullae, 14.0; length of nasals, 10.4. sen Co., California, southward through Distribution. - Found at inland sites in western Nevada, and on the northeast, from Baja California, from the central Sierra San near the western and southern shores ofthe Pedro Martir south to near Rosarito. Great Salt Lake, Utah, southward through Remarks. - No measurements, other than western Utah and extreme eastern Nevada those of the holotype, are available. to south-central Utah and northwestern Ar­ izona, north of the Grand Canyon of the Colorado River; thence across the southern Chaetodipus formosus one-third ofNevada and southward through southeastern California and the eastern slope Diagnosis. -A medium-sized species of ofthe Baja California peninsula to near Santa Chaetodipus, with soft, spineless pelage, rel­ Rosalia, in northeastern Baja California Sur. atively large ear pinnae and long tail, large Remarks. - Refer to Hafner and Hafner mastoid bullae and narrow interparietals; (1983) and Patton et al. (1981) for detailed mastoid bullae projecting slightly posteriad discussions ofthe systematic position of C. beyond the plane ofthe occipitals; tympanic formosus. bullae nearly meeting over the basisphenoid anteriorly; length of ear is usually greater than 9 mm; width of interparietals usually averages less than 5.9 mm and rarely ranges Chaetodipus formosus cinerascens to about 6.5 mm. (Nelson and Goldman, 1929) Comparisons. - Chaetodipus formosus is generally similar in appearance to most oth­ 1929. Perognathusformosus cinerascens Nelson er species of Chaetodipus without spines. and Goldman, Proc. BioI. Soc. Washington, 42:105,25 March. Refer to the account of arenarius for com­ parison with that species. From C. penicil­ Holotype. - Young adult female, skin and latus, formosus can be distinguished by its skull, MVZ 37685, from San Felipe, north­ finer and longer pelage, more heavily haired eastern Baja California, Mexico; obtained and crested tail, longer ear, narrower inter­ by Chester C. Lamb on 10 April 1926. parietals, and larger mastoid bullae; the Measurements of holotype. --Total mastoid bullae ofpenicillatus do not extend length, 155; length oftail, 75; length ofhind posteriad to the plane ofthe occipitals. From foot, 22.8; weight, 15.8 g; greatest length of C. baileyi, formosus can be distinguished by skull, 24.3; interorbital breadth, 6.1; length its smaller size, narrower interparietals, and of maxillary toothrow, 3.4; width across shorter hind foot (length ofhind foot offor­ mastoid bullae, 13.6; length ofinterparietal, mosus averages less than 25 mm and rarely 3.5; width of interparietals, 5.0; length of ranges to 26, the hind foot of baileyi mea­ nasals, 9.3; width of nasals, 2.2. sures 26 mm or greater and averages 27 or Distribution. - Known from an area along more). Other species of Chaetodipus with the Gulf slope of central Baja California, spineless pelage (some artus, hispidus, linea­ from San Felipe southward to near El Mar­ tus, pernix) have allopatric geographic rang- mol. 140 WILLIAMS ET AL.

Remarks. - No additional measurements Holotype. - Adult female, skin and skull, for cinerascens are available. MVZ 78555, from 2 mi W Smith Creek Cave, 6,300 ft, Mt. Moriah, White Pine Co., Nevada; obtained on 18 June 1937 by Lee Chaetodipus formosus formosus W. Arnold. (Merriam, 1889) Measurements ofholotype. - Total length, 190; length oftail, 103; length ofhind foot, 1889. PerognathusformosusMerriam, N. Amer. 23; length ofear, 12; weight, 21. 5 g; greatest Fauna, 1: 17, 25 October. length of skull, 22.30; width across bullae, 1956. Perognathus formosus domisaxensis 13.70; breadth across maxillary arches, Cockrum, J. Washington Acad. Sci., 46:131, 12.35; nasal length, 11.00; interorbital 7 May. breadth,7.05. 1956. Perognathus formosus melanocaudus Distribution-Ranges over the basin of Cockrum, 1. Washington Acad. Sci., 46:132, 7 May. the Pleistocene Lake Bonneville in north­ western Utah and northeastern Nevada. Holotype. -Adult male, skin and skull, Remarks. - Hall (1946) gave means and USNM 186508, from St. George, Washing­ ranges of measurements for four females ton Co., Utah; obtained by Vernon Bailey from the type locality. on 9 January 1889. Measurements ofholotype. - Total length, Chaetodipus formosus infolatus 195; length oftail, 111; length ofhind foot, (Huey, 1954) 26; length ofear (crown, dry), 6; occipitona­ sal length, 27.5; basilar length, 21.1; basilar 1954. Perognathus formosus infolatus Huey, length ofHensel, 19.2; width across mastoid Trans. San Diego Soc. Nat. Hist., 12:1, 1 bullae, 14.6; length of interparietal, 4.0; March. width of interparietals, 5.8; least intermas­ Holotype. -Adult male, skin and skull, toid distance, 6.0. SDSNH 15664, from 7 mi W San Francis­ Distribution. -Occupies south-central and quito Bay, lat. 28°30'N, GulfofCalifornia, southwestern Utah and northwestern Ari­ Baja California Sur, Mexico; obtained by zona; in Utah, found near St. George east­ Laurence M. Huey on 3 April 1947. ward to the Colorado River; in Arizona, Measurements ofholotype. - Total length, ranges north and west ofthe Colorado Riv­ 187; length oftail, 104; length ofhind foot, er. 25; length of ear (crown), 6; occipitonasal Remarks. -Cockrum (1956) listed mea­ length, 27.5; interorbital breadth, 6.6; length surements for 6 adults under the two sub­ of maxillary toothrow, 3.8; width across specific synonyms listed above; Durrant mastoid bullae, 14.6; length ofnasals, 10.2. (1952) gave statistics for measurements of Distribution. -Occupies the southern 7 adult female topotypes; and Hoffmeister portion ofthe range ofthe species, from near (1986) gave statistics for measurements of El Marmol, Baja California, southward to 4 samples ranging from 15 to 29 individuals near Santa Rosalia, northeastern Baja Cal­ each. Hoffmeister (1986) considered domi­ ifornia Sur. saxensis and melanocaudus to be synonyms Remarks. -No other measurements are offormosus. available for infolatus.

Chaetodipus formosus incolatus Chaetodipus formosus melanurus (Hall, 1941) (Hall, 1941)

1941. Perognathusformosus incolatus Hall, Proc. 1941. Perognathus formosus melanurus Hall, Bioi. Soc. Washington, 54:56, 20 May. Proc. Bioi. Soc. Washington, 54:57, 20 May. TAXONOMY 141

Holotype. - Adult female, skin and skull, northeastern Baja California, ranging from MVZ 73442, from 6 mi E California bound­ near the type locality on the northwest and ary, 4,000 ft, lat. 40 0 28'N, Washoe Co., Ne­ near Needles, San B~rnardino Co., on the vada; obtained by E. Raymond Hall on 7 northeast, southward on the Gulf slope of July 1936. Baja California to the east-central Sierra San Measurements ofholotype. - Total length, Pedro Martir. 199; length oftail, 108; length ofhind foot, Remarks. - No other published measure­ 25.3; length ofear, 11; weight, 21.2 g; great­ ments for mesembrinus are known. Length est length ofskull, 27.20; width across bul­ ofskull given by Elliot (1904) suggests either lae, 14.40; breadth across maxillary arches, an error in measuring or recording, or a ju­ 14.40; nasal length, 11.00; interorbital venile specimen. breadth, 7.00; length ofmaxillary toothrow, 3.95; depth ofcranium, 3.55. Distribution. - Extreme northeastern Chaetodipus formosus mohavensis California in the eastern Honey Lake Val­ (Huey, 1938) ley, Lassen Co., and northwestern Nevada in the Smoke Creek desert and adjacent ar­ 1938. Perognathusformosus mohavensis Huey, Trans. San Diego Soc. Nat. Hist., 9:35, 21 No­ eas; in Nevada, from southwestern Hum­ vember. boldt Co. southward to the Truckee River near Toulon, Pershing Co., and westward Holotype. - Adult male, skin and skull, to near Reno, Washoe Co. SDSNH 11317, from Bonanza King Mine, Remarks. - Hall (1946) listed means and Providence Mountains, San Bernardino Co., ranges for measurements of 10 males and California; obtained on 4 April 1935 by 11 females. Laurence M. Huey. Measurements ofholotype. - Total length, 181, length oftail, 100; length ofhind foot, 24; length of ear (crown), 8; occipitonasal Chaetodipus formosus mesembrinus length, 25.8; interorbital breadth, 7.0; length (Elliot, 1904) of maxillary toothrow, 3.7; width across mastoid bullae, 13.8; length of nasals, 9.5. 1904. Perognathus mesembrinus Elliot, Field Distribution. Columbian Mus., Pub!. 87, Zoo!' Ser., 3:251, -Occupies the Mojave Des­ 7 January. ert area of southeastern California, and western Nevada from the Walker River Holotype. -Adult male, skin and skull, southward and eastward to the extreme FMNH 11784, from Palm Springs, River­ southwestern comer of Utah. side Co., California; obtained by Edmund Remarks. - Hall (1946) listed means and Heller on 24 February 1903. ranges for measurements offour males and Measurements ofholotype. - Total length, seven females from Nevada; Durrant (1952) 195; length oftail, 114; length ofhind foot, listed measurements for one male and two 23; length of ear, 11; total length of skull, females from Utah. Specimens from north­ 21; basilar length ofHensel, 18; interorbital western Arizona, assigned to this subspecies breadth, 7.0; length of maxillary toothrow, by Hall (1981) were considered to be C. f 4.0; length of mastoid bulla, 9.0; width formosus by Hoffmeister (1986). across mastoid bullae, 14.0; length ofnasals, 9.4; zygomatic breadth, 13.0; greatest pa­ rietal width, 10.5; palatallength, 10.0; length Chaetodipus goldmani of mandible, 12.5; length of mandibular toothrow, 3.5. Diagnosis. -A medium-large species of Distribution. -Occupies the Colorado Chaetodipus with coarse pelage with few Desert region ofsoutheastern California and short, stout spines on the rump, long, 142 WILLIAMS ET AL. rounded ear pinna, a moderately long and Measurements ofholotype. - Total length, strongly crested tail, and a conspicuous, 202; length oftail, 108; length ofhind foot, buffy-colored lateral stripe; baculum com­ 28; length ofear (anterior base), 11; greatest paratively long and tip weakly curved. length of skull, 28.80; width across bullae, Comparisons. -Chaetodipus goldmani is 14.10; breadth across maxillary arches, most likely to be confused with C. artus, 13.00; nasal length, 10.70; interorbital with which it shares a portion of its geo­ breadth, 6.25. graphic range (see account ofartus for com­ Remarks. -Osgood (1900) listed means parison). C. goldmani is sympatric with no ofskull measurements for three specimens; other species of Chaetodipus with spines in Anderson (1964, 1972) and Straney and the pelage, although its range approaches Patton (1980) provided statistics for mea­ that ofC. intermedius; intermedius is small­ surements of several samples. er, with a relatively longer tail, smaller hind foot, and shorter ear pinnae (less than 9 mm). From the potentially sympatric spe­ Chaetodipus intermedius cies without spines in the pelage, C. gold­ mani differs as follows: size smaller than C. Diagnosis. -A smaller than average to baileyi, with relatively smaller mastoid bul­ medium-sized species of Chaetodipus, with lae, and wider interparietals; size larger than relatively weak spines usually present on the C. pernix, hind foot significantly longer rump, a long, crested tail, relatively short (length averages greater than 24 mm in gold­ ear pinna, conspicuous, buffy lateral stripe, mani), and tail much more strongly crested; wide interparietals, and narrow skull, mea­ size larger, pelage coarser, and lateral stripe sured at the anterior point ofthe zygomatic more conspicuous than in C. penicillatus. arches; head and body length varies from Distribution. -Occupies thornscrub and about 70 to 77 mm; width ofinterparietals short-tree forest associations in the Rio Ya­ ranges from about 7.2 to 8.0 mm; anterior qui drainage of extreme northeastern So­ zygomatic breadth ranges from about 10.5 nora, Mexico, southward through the coast­ to 11.8 mm (Anderson, 1972; Weckerly and al plains of Sonora to northern Sinaloa; on Best, 1985). the east, ranges in the barrancas of the Pa­ Comparisons. - Chaetodipus intermedius cific slope of the Sierra Madre Occidental is most often confused with C. penicillatus, into southwest-central Chihuahua (Patton, which is similar in size and proportions, and 1969a; Hall, 1981). with which it is broadly sympatric. C. in­ Remarks. - Patton (1969a) documented termedius differs from penicillatus in having the distribution of six allopatric chromo­ rump spines present (they may be rarely ab­ somal races of C. goldmani; the species is sent individually and seemingly so when an­ currently monotypic. Straney and Patton imals are molting); a shorterhind foot (length (1980) documented geographic variation in usually averages 22 mm or less in inter­ structure. medius and over 22 in penicillatus); nar­ rower rostrum; narrower breadth across the anterior portion of the zygomatic arches Chaetodipus goldmani (Osgood, 1900) (Anderson, 1972); and anterior extension of the supraoccipital, between the mastoid and 1900. Perognathus goldmani Osgood, N. Amer. interparietal, is shorter and comes to an Fauna, 18:54,20 September. acute angle anteriorly (in penicillatus it is Holotype. -Adult female, skin and skull, longer, broader, and squared off at the an­ USNM 96673, from Sinaloa, Sinaloa, Me­ terior end; Hoffmeister and Lee, 1967). C. xico; obtained by Edward A. Goldman on intermedius is probably most closely related 15 February 1899. to C. nelsoni, C. artus, and C. goldmani, the TAXONOMY 143 latter two being larger than and having al­ provided statistics for measurements of 16 lopatric geographic ranges to intermedius; males and 14 females. from nelsoni, intermedius differs in being slightly smaller, with slightly finer, softer pelage with fewer rump spines, and in hav­ Chaetodipus intermedius beardi ing shorter interparietals and a narrower Weckerly, Gennaro, and Best, 1988 rostrum. Other species ofChaetodipus found within the geographic range of intermedius 1988. Chaetodipus intermedius beardi Weck­ are C. baileyi, C. formosus (possible mar­ erly, Gennaro, and Best, Southwestern Nat., ginal sympatry), and C. hispidus; all three 33:100, 30 March. are larger and lack spines in the pelage. Holotype. -Adult female, skin and skull, Distribution. - Chaetodipus intermedius ENMU 8919, from 26 mi N, 15.5 mi E is broadly distributed in rocky habitats in Engle, Socorro Co., New Mexico; obtained the southwestern deserts ofNorth America, by Floyd W. Weckerly on 23 August 1982. ranging from south and east ofthe Colorado Measurements ofholotype. - Total length, River in extreme south-central Utah through 157; length of tail, 91; length of hind foot, most of western and southern Arizona to 19; length ofear, 7.7; greatest length ofskull, northern Sonora, central and southern New 23.5; width across mastoid bullae, 12.8; na­ Mexico, Trans-Pecos Texas, and northern sal length, 8.2; interorbital breadth, 6.2; and central Chihuahua (Hall, 1981). greatest depth ofcranium, 8.7; rostral width, Remarks. - Chaetodipus intermedius 3.6; nasal width, 2.1; interparietal length, generally is limited to rocky areas and sev­ 3.7; interparietal width, 7.1; length ofmax­ eral black colored subspecies, confined to illary toothrow, 3.6; length of mandible, relatively small, isolated lava flows, have 10.6. been described. The species has never been Distribution. -Occurs only on the Pedro reviewed systematically. Armendariz lava field in Sierra and Socorro counties, New Mexico. Remarks. - Weckerly et al. (1988) pro­ Chaetodipus intermedius ater vided measurements for four adult topo­ (Dice, 1929) types in addition to the holotype, and means and ranges ofmeasurements for samples of 1929. Perognathus intermedius ater Dice, Oc­ 68 adult males and 82 adult females. cas. Papers Mus. Zool., Univ. Michigan, 203: 2, 19 June. Holotype. - Adult male, skin, skull, and Chaetodipus intermedius crinitus body skeleton, UMMZ 58474, from Mal­ (Benson, 1934) pais Spring, 15 mi W Three Rivers, Otero Co., New Mexico; obtained on 17 July 1927 1934. Perognathus intermedius crinitus Benson, by Lee R. Dice. Proc. Biol. Soc. Washington, 47:199, 2 Octo­ ber. Measurements ofholotype. - Total length, 167; length of tail, 86; length of hind foot, Holotype. -Adult male, skin and skull, 20; length of ear (notch), 8.0; weight, 14.3 MVZ 55883, from 2.6 mi W Wupatki Ru­ g; occipitonasal length, 21.4; interorbital ins, Coconino Co., Arizona; obtained by breadth, 6.3; width across mastoid bullae, Louise Kellogg on 8 October 1932. 12.3; length of nasals, 9.6. Measurements ofholotype. - Total length, Distribution. - Known from the Carrizo­ 177; length oftail, 101; length ofhind foot, zo lava beds in western Lincoln and Otero 23.5; length ofear, 5; weight, 13.5 g; greatest counties, New Mexico (Hall, 1981). length of skull, 25.3; interorbital breadth, Remarks. -Weckerly and Best (1985) 6.35; length of mastoid bulla, 7.5; width 144 WILLIAMS ET AL. across mastoid bullae, 12.9; length ofinter­ and central Chihuahua; in New Mexico, parietal, 3.35; width ofinterparietals, 7.25; ranges northward in the Rio Grande low­ length ofnasals, 9.35; width ofrostrum, 4.45. lands to near Albuquerque; in Texas, ranges Distribution. -Occurs in north-central east to about western Winkler and Ward and northwestern Arizona and south-cen­ counties; in Chihuahua, ranges southward tral Utah; ranging the south side ofNavajo to near Ojinaga and westward to near Casas Mountain, south and east of the Colorado Grandes and 5 mi W San Francisco (Hall, River and north ofthe Mogollon Rim; mar­ 1981). ginallocalities on the east are Aztec Creek, Remarks. - Hoffmeister (1986) noted that river mile 68.5, Glen Canyon, San Juan Co., the type locality for nigrimontis, a dark-col­ Utah; Moa Ave and Walnut Tank (10 mi ored but not black population, was in a zone N Angell Augusta), Coconino Co., Arizona of intergradation between intermedius and (Hall, 1981). phasma. Hoffmeister (1986) gave statistics Remarks. - Benson (1934) listed averages for measurements offour samples from Ar­ and ranges of measurements for 15 males; izona; Weckerly and Best (1985) gave sta­ Hoffmeister (1986) gave statistics for mea­ tistics for measurements oftwo samples (68 surements of samples of 9 and 13 individ­ males, 82 females; 11 males, 16 females) uals. from New Mexico; Anderson (1972) gave statistics for external measurements ofthree samples and cranial measurements for one Chaetodipus intermedius intermedius sample from Chihuahua; Genoways et al. (Merriam, 1889) (1977) listed measurements for two males and three females from Texas. 1889. Perognathus intermedius Merriam, N. Amer. Fauna, 1:18,25 October. 1889. Perognathus obscurus Merriam, N. Amer. Fauna, 1:20, 25 October. Chaetodipus intermedius lithophilus 1933. Perognathus intermedius nigrimontis (Huey, 1937) Blossom, Occas. Papers Mus. Zool., Univ. Michigan, 265:1, 21 June. 1937. Perognathus intermedius lithophilus Huey, Trans. San Diego Soc. Nat. Hist., 8:355, 15 Holotype. - Adult male, skin and skull, June. USNM 186509, from Mud Spring, Mohave Co., Arizona; obtained by Vernon Bailey on Holotype. -Adult male, skin and skull, 26 February 1889. SDSNH 11211, from Porto [Puerto] Liber­ Measurements ojholotype. - Total length, tad, summit ofrocky hill 1.5 mi NNW fresh 183; length oftail, 106; length ofhind foot, water spring on beach, Sonora, Mexico; ob­ 21; length of ear (crown, dry), 4.5; occipi­ tained by Laurence M. Huey on 5 February tonasal length, 24.4; basilar length, 19.0; 1935. basilar length ofHensel, 16.6; width across Measurements ojholotype. - Total length, mastoid bullae, 13.5; length ofinterparietal, 166; length of tail, 91; length of hind foot, 3.0; width ofinterparietals, 7.8; least inter­ 19; length of ear (crown), 5; occipitonasal mastoid distance, 8.0. length, 23.5; interorbital breadth, 6.2; length Distribution. - Ranges from the Colorado of maxillary toothrow, 3.4; width across River in western Arizona southward and mastoid bullae, 12.7; length of nasals, 9.3. eastward across Arizona south of the Mo­ Distribution. - Known only from the type gollon Rim (but not including the lower Col­ locality in northwestern Sonora. orado River Valley or the western Gila Riv­ Remarks. - Huey (1937) gave means and er Valley) to southern and central New ranges of measurements for eight speci­ Mexico, Trans-Pecos Texas, and northern mens, including the holotype. TAXONOMY 145

Chaetodipus intermedius minimus orbital breadth, 5.8; length of maxillary (Burt, 1932) toothrow, 3.4; width across mastoid bullae, 12.3; length of interparietal, 2.6; width of 1932. Perognathus penicillatus minimus Burt, interparietals, 6.1; length of nasals, 8.5. Trans. San Diego Soc. Nat. Hist., 7:164, 31 Distribution. -Occurs in westernmost October. Arizona along the Colorado River, from near Holotype. - Adult male, skin and skull, Hoover Dam southward, extending east­ CIT 50424, from Turner's Island, lat. ward in the Gila River Valley to Phoenix 28°43'N, long. 112°19'W, Gulf of Califor­ and southward to Organ Pipe National nia, Sonora, Mexico; obtained on 31 De­ Monument; also known from a small area cember 1931 by William H. Burt. in northwestern Sonora and southwestern Measurements ofholotype. - Total length, Arizona encompassed by the Pinacate 162; length of tail, 97; length of hind foot, Mountain lava flows and in Sonora in the 20; length of ear (crown), 5, (notch, dry), Tinajas and Tule ranges (Hall, 1981; Hoff­ 6.2; greatest length of skull, 23.9; basal meister, 1986). length, 19.5; interorbital breadth, 6.2; length Remarks. - Hoffmeister (1986) found that of maxillary toothrow, 3.5; length of mas­ specimens of pinacate did not differ from toid bulla, 7.1; width across mastoid bullae, other samples ofphasma except for darker 12.5; length of interparietal, 3.6; width of color. Hoffmeister (1986) gave statistics for interparietals, 7.1; length of nasals, 9.2. four samples ofphasma from Arizona. Distribution. - Known only from the type locality. Remarks. - Hoffmeister and Lee (1967), Chaetodipus intermedius rupestris in their review of Chaetodipus penicillatus, (Benson, 1932) decided that minimus was a subspecies of C. intermedius; in so far as is recorded, min­ 1932. Perognathus intermedius rupestris Ben­ imus is represented only by the holotype. son, Univ. Califomia Publ. Zool., 38:337, 14 The holotype is now in the UCLA collec­ April. tions. Holotype. - Young adult male, skin and skull, MVZ 50595, from that part of lava beds nearest Kenzin, Dona Ana Co., New Chaetodipus intermedius phasma Mexico; obtained on 24 October 1931 by (Goldman, 1918) Annie M. Alexander. Measurements ofholotype. - Total length, 1918. Perognathus intermedius phasma Gold­ 169; length of tail, 92; length of hind foot, man, Proc. BioI. Soc. Washington, 31:22, 16 20.5; length of ear (crown), 4; weight, 12.9 May. g; occipitonasal length, 23.9; frontonasal 1933. Perognathus intermedius pinacate Blos­ length, 16.0; interorbital breadth, 5.8; length som, Occas. Papers Mus. Zool., Univ. Mich­ ofmastoid bulla, 7.8; width across mastoid igan, 273:4, 31 October. bullae, 12.4; distance between stylomastoid Holotype. - Adult male, skin and skull, foramina, 9.7. USNM 203003, from Tinajas Atlas, 1,400 Distribution. - Known only from the Af­ ft, Gila Mountains, Yuma Co., Arizona; ob­ ton Lava Flow, Dona Ana Co., south-cen­ tained by Edward A. Goldman on 23 No­ tral New Mexico. vember 1913. Remarks. - Benson (1932) gave measure­ Measurements ofholotype. - Total length, ments of2 paratypes and Weckerly and Best 165; length of tail, 97; length of hind foot, (1985) listed statistics for measurements of 20.5; greatest length of skull, 23.0; inter- 50 males and 55 females. 146 WILLIAMS ET AL.

Chaetodipus intermedius umbrosus (Dalquest, 1951). From C. nelsoni, it is dis­ (Benson, 1934) tinguished by its dull gray rather than dark brownish color and the lack ofspines in the 1934. Perognathus intermedius umbrosus Ben­ pelage; from C. penicillatus, lineatus differs son, Proc. BioI. Soc. Washington, 47:200, 2 in being grayer without the strong, black October 1934. overwash of sympatric penicillatus, and in Holotype. -Adult male, skin and skull, its larger size and larger and broader skull MVZ 55964, from Camp Verde, Yavapai (Dalquest, 1951). C. lineatus may also be Co., Arizona; obtained on 3 October 1932 found with C. hispidus, which is much larger by Louise Kellogg. with a relatively short, non-crested tail. Measurements ofholotype. - Total length, Distribution. - Known only from a small 173; length of tail, 99; length of hind foot, area in southwestern San Luis Potosi and 23; length ofear (crown), 5; weight, 16.7 g; extreme southeastern Zacatecas (Dalquest, greatest length of skull, 25.5; interorbital 1951; Hall, 1981; Matson and Baker, 1986). breadth, 6.55; length ofmastoid bulla, 7.95; Remarks-The status of C. lineatus has width across mastoid bullae, 13.3; length of been questioned by several mammalogists, interparietal, 3.6; width of interparietals, primarily because ofthe inability ofothers 7.65; length ofnasals, 9.6; width ofrostrum, to capture specimens clearly referable to this 4.35. species and because ofits great similarity to Distribution. -Occurs in central Arizona, C. nelsoni, except for its color and lack of from the Verde Valley, Yavapai Co., on the rump spines. Variability in presence or ab­ southeast, northwestward to near Drake, in sence of rump spines has been noted for a extreme south-central Coconino Co. (Hoff­ number ofspecies ofChaetodipus, including meister, 1986). those that typically have spines (e.g., C. ar­ Remarks. -Hoffmeister (1986) restricted tus, C. intermedius, molting and/or juve­ the distribution of umbrosus, assigning niles of several species), and species nor­ specimens from the southern and eastern mally without spines (e.g., C. penicillatus, portion of the range previously attributed Hoffmeister and Lee, 1967). Furthermore, to umbrosus (Hall, 1981) to intermedius in­ there is no definite distinction between the stead. Hoffmeister (1986) gave statistics for normal, coarse, troughed overhairs, weak a sample of50 individuals from near Camp bristles, and stiffer spines ofall Chaetodipus Verde; Benson (1934) gave means and ex­ spp. (Homan and Genoways, 1978). Three tremes of measurements of 7 adult males. species ofsimilar-sized Chaetodipus sharing the same area would be unusual. Specimens of C. lineatus may be spineless individuals Chaetodipus lineatus of C. nelsoni. That they are also dull gray rather than the normal dark brownish sug­ Diagnosis. -A medium-sized species of gests that a simple mutation or age is re­ Chaetodipus with a long, crested tail; color sponsible for the differences. However, this is dull-gray dorsally, consisting ofa mix of matter only can be resolved by additional gray and blackish and lined with buffy, with study. a narrow, buffy lateral stripe; no spines on the rump; skull essentially the same as C. nelsoni; length of head and body averages around 76 to 79 mm; length oftail averaging Chaetodipus lineatus (Dalquest, 1951) from about 95 to 98 mm. Comparisons. - Chaetodipus lineatus 1951. Perognathus lineatus Dalquest, J. Wash­ ington Acad. Sci., 41:362, 14 November. shares its geographic range with C. nelsoni and C. penicillatus, both ofwhich have been Holotype. -Adult male, skin and skull, taken in the same traplines with lineatus LSUMZ 5253, from 1 km S Arriaga, San TAXONOMY 147

Luis Potosi, Mexico; obtained on 21 Sep­ larger animal with a relatively short, non­ tember 1950 by Walter W. Dalquest. crested tail and without spines on the rump. Measurements ofholotype. - Total length, Distribution. - Ranges from southeastern 175; length of tail, 98; length of hind foot, New Mexico, southward through south­ 23; length ofear, 8; greatest length ofskull, central Chihuahua, the eastern one-half of 25.20; width across bullae, 14.00; breadth Durango (except for the northeastern cor­ across maxillary arches, 11.90; nasal length, ner), extreme northeastern Jalisco, Zacate­ 8.80; interorbital breadth, 6.55. cas, and extreme east-central Jalisco; on the Remarks. - Dalquest (1951) listed means east, ranges to south-central Nuevo Leon, for measurements ofeight males and seven southwestern Tamaulipas, and central San females. Luis Potosi (Hall, 1981). Remarks. - The relationship between C. nelsoni and C. lineatus appears to be close; Chaetodipus nelsoni lineatus may not be a valid species, but rath­ er a name applied to variant specimens of Diagnosis. - A medium-sized species of nelsoni (see account of C. lineatus). Aside Chaetodipus with a long, crested tail, rela­ from C. lineatus, C. nelsoni appears to be tively coarse hair with spines on the rump, most closely related to C. artus, C. gold­ and short ear pinnae (8 mm or less); skull mani, and C. intermedius, the former two moderately narrow across anterior portion having allopatric geographic ranges to nel­ ofzygomatic arches; interparietals relative­ soni. ly long; length of head and body averages between about 76 and 80 mm; length oftail averages about 98 to 110 mm; length of Chaetodipus nelsoni canescens interparietal averages 3.7 mm (range 3.4 to (Merriam, 1894) 3.9); width of interparietals ranges from about 6.7 to 8.1 mm (mean 7.5); anterior 1894. Perognathus (Chaetodipus) intermedius zygomatic breadth averages 11.9 mm (range canescens Merriam, Proc. Acad. Nat. Sci. Phil­ 11.0 to 12.9). adelphia, 46:267, 27 September. Comparisons. - Chaetodipus nelsoni is 1938. Perognathus collis Blair, Occas. Papers most likely to be confused with C. penicil­ Mus. Zoo!., Univ. Michigan, 381:1, 20 June. latus, which is similar in size and propor­ 1938. Perognathus collis popei Blair, Occas. Pa­ tions and with which it is broadly sympat­ pers Mus. Zoo!., Univ. Michigan, 381 :3, 20 ric. C. nelsoni differs from penicillatus in June. usually having rump spines, narrower ros­ Holotype. - Young adult male, skin and trum, and narrower breadth across the an­ skull, USNM 51016, from Jara1, Coahuila, terior portion ofthe zygomatic arches (An­ Mexico; obtained by Clark P. Streator on derson, 1972). In nelsoni, the anterior 14 January 1893. extension ofthe supraoccipital, between the Measurements ofholotype. - Total length, mastoid and interparietal, is shorter and 193; length oftail, 117; length ofhind foot, comes to an acute angle anteriorly (in pen­ 22; length ofear (anterior base, dry), 8; oc­ icillatus it is longer, broader and squared off cipitonasal length, 25.0; basilar length of at the anterior end). C. nelsoni typically has Hensel, 17.5; interorbital breadth, 6.1; width more spines and a broader interorbital re­ across mastoid bullae, 13.5; length ofinter­ gion and longer interparietals than artus parietal, 3.7; width of interparietals, 7.2; (Anderson, 1972). C. goldmani is larger with length of nasals, 9.3. relatively smaller mastoid bullae. See ac­ Distribution. - Ranges from southeastern count of intermedius for comparison with New Mexico, near the boundary with Texas, that species. The only other potentially southward through eastern Chihuahua to sympatric species is C. hispidus, a much northeastern Durango and southern Coa- 148 WILLIAMS ET AL. huila; probably also occurs in extreme west­ measurements ofthree specimens from the central Nuevo Leon; easternmost localities type locality. in Texas are Sheffield, Pecos Co., and Com­ stock, Val Verde Co. (Hall, 1981). Remarks. -Anderson (1972) gave statis­ Chaetodipus penicillatus tics for measurements of14 specimens from Chihuahua, and Baker (1956) listed mea­ Diagnosis. -A medium-sized species of surements (means, ranges) for 5 males and Chaetodipus, with a relatively long, crested, 8 females from Coahuila. and sparsely-haired tail, short ear pinnae, no spines on the rump, and a buffy lateral stripe that is comparatively faint or absent; Chaetodipus nelsoni nelsoni proximal two-thirds oftail relatively sparse­ (Merriam, 1894) ly haired and with annulated pattern; an­ terior extension of the supraoccipital be­ 1894. Perognathus (Chaetodipus) nelsoni Mer­ tween interparietal and mastoid relatively riam, Proc. Acad. Nat. Sci. Philadelphia, 46: broad, straplike, and squared at the anterior 266, 27 September. end; mastoid bulla relatively small, but in­ Holotype. -Old adult female, skin and terparietals not comparatively broader; av­ skull, USNM 50214, from Hacienda La erage length of head and body ranges from Parada, San Luis Potosi, Mexico; obtained about 77 to 87 mm; average length of tail by Edward W. Nelson on 19 August 1892. ranges from about 88 to 101 mm; length of Measurements ofholotype. - Total length, ear averages 8 mm or less. 190; length oftail, 105; length ofhind foot, Comparisons. - Chaetodipus penicillatus 24; length of ear (anterior base, dry), 8; is similar in general appearance to other greatest length ofskull, 26.05; width across small and medium-sized species of Chae­ bullae, 13.00; breadth across maxillary todipus without spines on the rump, and to arches, 12.30; nasal length, 10.05; interor­ C. intermedius and C. nelsoni, both ofwhich bital breadth, 6.75. normally have spines (accounts ofthe latter Distribution-Ranges from south-central two species have comparisons). See ac­ Chihuahua southward through the eastern counts of C. arenarius, C. baileyi, and C. one-half of Durango (except for the north­ formosus for methods ofdistinguishing those eastern corner), extreme northeastern Jalis­ species from C. penicillatus. C. penicillatus co, Zacatecas (except montane forests in also is generally similar to C. pernix; peni­ western part), and extreme east-central Ja­ cillatus differs in being larger, lacking a con­ lisco; on the east, ranges to south-central spicuous, buffy lateral stripe, and having a Nuevo Leon, southwestern Tamaulipas, and larger crest of hairs on the tail, interorbital central San Luis Potosi (Hall, 1981; Matson breadth averaging 5.9 mm orgreater (versus and Baker, 1986). less than 5.8 in pernix), and longer mastoid Remarks. -Alvarez (1963) listed means bullae usually measuring more than 7.5 mm and ranges of measurements for nine spec­ rather than less (Anderson, 1972; Hall, 1981; imens from Tamaulipas. Baker (1956) gave Hoffmeister and Lee, 1967). means and ranges ofmeasurements for sev­ Distribution. - Nearly always associated en males and two females from Coahuila. with creosote bush (Larrea spp.) commu­ Dalquest (1953) listed means of measure­ nities in the deserts of southwestern North ments for 10 males and 10 females from America, ranging from southeastern Cali­ San Luis Potosi. Genoways and Jones (1973) fornia and northwestern Baja California, gave statistics for seven adults from Zaca­ eastward and southward through southern tecas, and Osgood (1900) listed means for Nevada to extreme southwestern Utah, cen- TAXONOMY 149 tral and southern Arizona south ofthe Mo­ surements and diagnoses for each subspe­ gollon Rim, all ofSonora except the south­ cies. Although generally considered to lack ern and southeastern one-fifth, southern spines on the rump, Hoffmeister and Lee New Mexico, the Trans-Pecos region of (1967) noted that a few individuals from western Texas, and the desert portions of scattered localities in Arizona and many in­ the Mexican Plateau, including the northern dividuals in a population from the Graham and eastern halves ofChihuahua, all but the Mountains, Arizona, had spines. northeastern segment of Coahuila, eastern Durango, northeastern Zacatecas, the northern half of San Luis Potosi, and the Chaetodipus penicillatus angustirostris southern extreme ofNuevo Leon; may also (Osgood, 1900) occur in the extreme southwestern portion ofTamaulipas (Hall, 1981). 1900. Perognathus penicillatus angustirostris Remarks. -Patton (1969b) and Patton et Osgood, N. Amer. Fauna, 18:47,20 Septem­ aI. (1981) found three chromosomal and ber. biochemical races of C. penicillatus which Holotype. -Adult male, skin and skull, corresponded to the groups, based on geo­ USNM 73881, from Carriso [= Carrizo] graphic variation of structure, documented Creek, Colorado Desert, Imperial Co., Cal­ by Hoffmeister and Lee (1967). These "ge­ ifornia; obtained by A. W. Anthony on 31 netic" races did not correspond precisely to March 1895. Hoffmeister and Lee's taxonomic assign­ Measurements ofholotype. - Total length, ments, however. The locations ofthe three 191; length of tail, 105; length of hind foot major races are: Sonora and eastern Arizona (dry), 24.4; greatest length of skull, 27.10; (pricei and the eastern segment of penicil­ width across bullae, 13.30; breadth across latus); Chihuahuan Desert (eremicus and maxillary arches, 12.90; nasal length, 11.10; atrodorsalis); Mojave and Colorado desert interorbital breadth, 6.40. areas of California, Baja California, and Distribution. -Found in southeastern western Arizona (angustirostris, western California, from near Peck's Butte, Los An­ Arizona segment ofpenicillatus). C. p. ste­ geles Co., southward to the region around phensi and sobrinus ofthe northern Mojave San Felipe, Baja California on the Gulfslope, Desert, California and southern Nevada and and eastward to the Colorado River at Fort Utah, respectively, have not been studied Yuma, Imperial Co., California. electrophoretically or karyotypically. C. p. Remarks. -Length offoot (dry) listed on seri of Tiburon Island has the same karyo­ the tag ofthe holotype was 23.9 mm. Based type as pricei from the mainland ofSonora on research of Patton (1969b) and Patton (Patton, 1969b). Individuals ofthe first two et al. (1981), the western Arizona popula­ genetic races have been taken at one locality tion of C. penicillatus penicillatus probably near the boundary ofthe Sonoran and Chi­ should be allied with angustirostris rather huahuan deserts, on the Deming Plain of than with populations of penicillatus from southwestern New Mexico, without evi­ central Arizona. dence of hybridization. Some evidence of hybridization of the first and third genetic races has been found, however (Patton et Chaetodipus penicillatus atrodorsalis aI., 1981). The significance ofthese findings (Dalquest, 1951) to the specific status of the three races has not yet been determined. Hoffmeister and 1951. Perognathus penicillatus atrodorsalis Lee (1967) reviewed C. penicillatus system­ Dalquest, J. Washington Acad. Sci., 41:362, atically and provided statistics for mea- 14 November. 150 WILLIAMS ET AL.

Holotype. -Adult male, skin and skull, Chaetodipus penicillatus penicillatus LSUMZ 5226, from 7 km W Presa de Gua­ (Woodhouse, 1852) dalupe, San Luis Potosi, Mexico; obtained by WalterW. Dalquest on 12 October 1950. 1852. Perognathus penecillatus [sic] Wood­ Measurements ofholotype. - Total length, house, Proc. Acad. Nat. Sci. Philadelphia, 169; length of tail, 96; length of hind foot, 6:200, December. 22; length ofear, 8; greatest length ofskull, Holotype. - Type specimen not specified 24.90; width across bullae, 12.80; breadth by Woodhouse, but later regarded as adult across maxillary arches, 12.05; nasal length, male, mounted skin, skull, and baculum, 9.90; interorbital breadth, 5.90. USNM 2676/37437 (Merriam, 1889; Os­ Distribution. - Known from northern and good, 1900), from San Francisco Moun­ eastern San Luis Potosi and southern Nuevo tains, New Mexico (later thought to be Leon (Hall, 1981). northeast side ofSan Francisco Mountains, Remarks. - Dalquest (1951) listed means Coconino Co., Arizona); type locality sub­ for 9 adult males and 12 adult females from sequently fixed as: 1 mi SW Parker, Yuma the vicinity of the type locality; Dalquest Co., Arizona (Hoffmeister and Lee, 1967). (1953) listed means for 8 males from a sec­ Measurements ofholotype. - Total length, ond locality in San Luis Potosi. 204; length oftail, 115; length ofhind foot, 25.5; occipitonasal length, 27.55; interor­ Chaetodipus penicillatus eremicus bital breadth, 6.6; length ofmaxillary tooth­ (Mearns, 1898) row, 3.9; length ofmastoid bulla, 8.5; width across mastoid bullae, 13.7; length ofnasals, 1898. Perognathus (Chaetodipus) eremicus 10.9. Mearns, Bull. Amer. Mus. Nat. Hist., 10:300, Distribution. - Known from west-central 31 August. Arizona, from the Colorado River Valley in Holotype. -Adult female, skin and skull, the vicinity ofToprock, Mohave Co., south USNM 21052/36094, from Fort Hancock, to near Yuma, Yuma Co., and eastward El Paso [now in Hudspeth] Co., Texas; ob­ south of the Mogollon Rim to San Carlos tained by Edgar A. Mearns on 27 June 1893. Reservoir, Gila County; the southern Measurements ofholotype. - Total length, boundary generally parallels the course of 163; length of tail, 83; length of hind foot, the Gila River, although populations as­ 22.1; basilar length of Hensel, 17.5; inter­ signed to this subspecies are found at some orbital breadth, 6.4; width across mastoid localities several miles south of the river bullae, 12.6; length of interparietal, 3.0; (Hall, 1981; Hoffmeister, 1986; Hoffmeister width ofinterparietals, 7.0; length ofnasals, and Lee, 1967). 9.8. Remarks. - The karyotypic and bio­ Distribution. - Ranges from southern New chemical information reported by Patton Mexico, southward through Trans-Pecos (1969b) and Patton et al. (1981) suggests Texas (Winkler Co. on the northwest and that western Arizona populations ofC. pen­ Val Verde Co. on the southeast) and the icillatus penicillatus are allied with popu­ desert regions ofChihuahua, Coahuila, Du­ lations ofangustirostris from California and rango, Zacatecas, and the western comer of Baja California. San Luis Potosi (Hall, 1981). Remarks. - Anderson (1972) listed statis­ Chaetodipus penicillatus pricei tics for measurements of 18 specimens from (J. A. Allen, 1894) Chihuahua, Genoways et al. (1977) gave measurements for 4 specimens from Texas, 1894. Perognathus pricei J. A. Allen, Bull. Amer. Mus. Nat. Hist., 6:318, 7 November. and Baker (1956) gave means and ranges of measurements for 8 adult males and 5 adult Holotype. - Subadult male, skin and skull, females from Coahuila. AMNH 8359/6685, from Oposura [= Moc- TAXONOMY 151 tezuma], Sonora, Mexico; obtained by B. C. and ranges of measurements of 18 speci­ Condit on 31 May 1894. mens. Measurements ofholotype. - Total length, 157; length of tail, 90; length of hind foot, 23; length ofear, 7.5; greatest length ofskull, Chaetodipus penicillatus sobrinus 23.0; basilar length, 18; interorbital breadth, (Goldman, 1939) 5.5; length ofmaxillary toothrow, 3.5; width across mastoid bullae, 11.5; length ofinter­ 1939. Perognathus penicillatus serosus Gold­ parietal, 4.0; width of interparietals, 8.0; man, Proc. Biol. Soc. Washington, 52:34, 11 length of nasals, 7.7; zygomatic breadth, March. 11.5; length of rostrum, 9.6. 1939. Perognathus penicillatus sobrinus Gold­ Distribution. -Found across southern man, J. Mamm., 20:257, 15 May. Renaming Arizona, generally south ofthe Gila River, of serosus Goldman, 1939. southward to the vicinity ofthe Rio Mayo, Holotype. - Adult male, skin and skull, near Navajoa, Sonora, and eastward to USNM 27598/39697; from sand flat along southwestern New Mexico near Deming Virgin River, 7 mi above Bunkerville, Clark (Hall, 1981; Patton et al., 1981). Co., Nevada [considered to be Mohave Co., The boundary between Remarks. - pricei Arizona by Hardy, J. Mamm., 30:435, 17 and in southwestern New Mexico eremicus November 1949]; obtained on 9 May 1891 is not established in detail; Patton et al. by Vernon Bailey. (1981) reported capturing specimens ofboth Measurements ofholotype. - Total length, subspecies, based on chromosomes and tis­ 202; length oftail, 102; length ofhind foot, sue allele patterns, from the same traplines 26; occipitonasal length, 27.6; interorbital near Deming, Luna Co., New Mexico. breadth, 6.4; length of maxillary toothrow, 4.0; width across mastoid bullae, 13.5; length ofinterparietal, 3.2; width ofinterparietals, Chaetodipus penicillatus seri 7.5; length of nasals, 11.1; zygomatic (Nelson, 1912) breadth, 14.2. Distribution. - Ranges from the extreme 1912. Perognathus penicillatus goldmani Town­ southwestern corner of Utah and extreme send, Bull. Amer. Mus. Nat. Hist., 31:122,14 northwestern corner ofArizona to near Da­ June. vis, Mohave Co., southward and westward 1912. Perognathus peniciUatus seri Nelson, Proc. through southern Nevada (Hall, 1981). Biol. Soc. Washington, 25:116, 29 June. Re­ Remarks. - The state in which the type naming of goldmani Townsend, 1912. locality is found has been disputed (Hoff­ Holotype. -Male, skin and skull, USNM meister and Lee, 1967). Hall (1946) gave 198411, from Tiburon Island, Gulf ofCal­ means and ranges of measurements of 15 ifornia, Sonora, Mexico; obtained by H. E. males and 7 females from Nevada, listed as Anthony on 13 April 1911. sobrinus, and 10 males and 10 females listed Measurements ofholotype.-Total length, as penicillatus. Hoffmeister (1986) gave sta­ 171; length of tail, 90; length of hind foot, tistics for six males and six females from 23.0; greatest length of skull, 24.80; width Arizona. across bullae, 11.90; breadth across maxil­ lary arches, 11.25; nasal length, 9.60; inter­ orbital breadth, 6.10. Chaetodipus penicillatus stephensi Distribution. - Restricted to Tiburon Is­ (Merriam, 1894) land, Gulf of California, Sonora. Remarks. - Hoffmeister and Lee (1967) 1894. Perognathus (Chaetodipus) stephensi listed the type specimen as AMNH 31845; Merriam, Proc. Acad. Nat. Sci. Philadelphia, itis as given above. Burt (1932) gave means 46:267,27 September. 152 WILLIAMS ET AL.

Holotype. - Adult male, skin and skull, fers in being much smaller (see also the ac­ USNM 27774/39873, from Mesquite Val­ count ofbaileyi). Refer to the account ofC. ley, NW arm Death Valley, -13 ft, Inyo artus for characters distinguishing that po­ Co., California; obtained by Frank Stephens tentially sympatric species. on 6 April 1891. Distribution. -Occurs on the coastal Measurements ofholotype. - Total length, plains ofwestern Mexico, from central So­ 177; length of tail, 96; length of hind foot, nora southward to northern Nayarit (Hall, 21; length of ear (anterior base, dry), 7.5; 1981). occipitonasallength, 22.7; basilar length of Remarks. - Patton et al. (1981) noted ex­ Hensel, 16.0; interorbital breadth, 6.0; width tensive differences in the karyotypes of C. across mastoid bullae, 12.0; length ofinter­ pernix pernix and C. pernix rostratus (2n = parietal, 3.0; width of interparietals, 6.7; 36 or 38 and 52, respectively) and a cor­ length of nasals, 9.0. responding biochemical difference oflesser Distribution. - Known only from the magnitude. Little data are available on northern end of Death Valley, Inyo Co., structural variation. California (Hoffmeister and Lee, 1967). Remarks. - Hoffmeister and Lee (1967) noted that the specimens ofstephensi avail­ Chaetodipus pernix pernix able to them were too few and too variable (1. A. Allen, 1898) in size to characterize adequately the sub­ species or determine its relationships to an­ 1898. Perognathus pernix J. A. Allen, Bull. gustirostris and penicillatus. Amer. Mus. Nat. Rist., 10:149, 12 April. Holotype. -Adult male, skin and skull, BM(NH) 98.3.2.126, from Rosario, Sina­ Chaetodipus pernix loa, Mexico; obtained by P. O. Simmons on 22 February 1897 [field numberofcollector, Diagnosis. -A smaller than average spe­ 139]. cies of Chaetodipus, with spineless pelage, Measurements ofholotype. - Total length, a long, small-crested tail and a conspicuous, 165; length of tail, 90; length of hind foot, buffy lateral stripe; relatively small mastoid 22; length ofear, 9; greatest length ofskull, bullae, wide interparietals, and wide ros­ 25.0; basal length, 19.6; interorbital breadth, trum; anterior extensions of supraoccipital 6.0; width across mastoid bullae, 12.0; length narrow and ending in a more or less acute of nasals, 8.5. point between the mastoid bulla and the Distribution. - Ranges from north-cen­ interparietal; length ofhead and body rang­ tral Sinaloa, near Pericos, southward to es from about 68 to 78 mm; length of tail northern Nayarit. from about 94 to 97 mm. Remarks. -Osgood (1900) gave average Comparisons. - Chaetodipus pernix is measurements for three specimens from the most likely to be confused with C. penicil­ type locality. latus, with which it shares the northern por­ tion ofits range; see account ofpenicillatus for comparison. C. pernix is also similar in appearance to the allopatric species, C. ar­ Chaetodipus pernix rostratus enarius (see account ofthat species for com­ (Osgood, 1900) parison). C. pernix differs from the sym­ 1900. Perognathus pernix rostratus Osgood, N. patrie species, C. goldmani, by being much Amer. Fauna, 18:51,20 September. smaller, having a smaller crest on the tail, and lacking spines on the rump. From the Holotype. - Young adult male, skin and sympatric species, C. baileyi, C. pernix dif- skull, USNM 95818, from Camoa, Rio TAXONOMY 153

Mayo, Sonora, Mexico; obtained by Ed­ Distribution. -Generally found on rocky, ward A. Goldman on 28 October 1898. desert slopes and other rocky areas in the Measurements ofholotype. - Total length, Colorado Desert ofsoutheastern California 162; length of tail, 94; length of hind foot, and the Baja California peninsula; ranges 23.5; greatest length of skull, 23.30; width from the extreme southern tip of Nevada across bullae, 11.40; breadth across maxil­ along the Colorado River, and from near lary arches, 11.15; nasal length, 8.75; inter­ Palm Springs, Riverside Co., California, orbital breadth, 5.45. southward in Baja California to Cabo San Distribution. -Ranges from near Tecori­ Lucas, Baja California Sur; on the Baja Cal­ pa, in south-central Sonora, southward to ifornia peninsula, not recorded from the near Guamuchil, north-central Sinaloa cooler Pacific coastal plains north ofthe Si­ (Hall, 1981). erra de la Laguna in the cape region, but Remarks. -Osgood (1900) listed average found on Magdalena and Margarita islands measurements ofthree specimens from the offofthe Pacific coast ofBaja California Sur type locality. (Hall, 1981; Huey, 1964). Remarks. - Populations ofthis species are Chaetodipus spinatus found on most of the islands on the Baja California peninsula side ofthe GulfofCal­ Diagnosis.-A medium- to medium-large ifornia, as well as two islands on the Pacific sized species of Chaetodipus, with a long, side of the peninsula. With one exception, crested tail, small ear pinnae, numerous each of these populations is considered to spines on the rump and often on the flanks, be a different subspecies, resulting in the and without, or nearly without, a lateral relatively large number ofsubspecies for C. stripe ofbuffy color; skull comparatively flat spinatus. The species has never been re­ dorsally; skull relatively narrow; mastoid viewed systematically. bullae small; interparietals moderate in width; length ofhead and body ranging from about 75 to 95 mm; length ofear from about Chaetodipus spinatus broccus 5 to 8.5 mm; width ofinterparietals ranges (Huey, 1960) from about 7.0 to 8.4 mm; length ofmastoid bulla ranges from about 6.4 to 8.2 mm. 1960. Perognathus spinatus broccus Huey, Trans. San Diego Soc. Nat. Hist., 12:410, 1 February. Comparisons. - Chaetodipus spinatus is generally similar in appearance to C. cali­ Holotype. - Adult male, skin and skull, fornicus and C. fallax, both of which may SDSNH 6891, from San Ignacio, lat. be marginally sympatric with spinatus; and 27°l7'N, Baja California Sur, Mexico; ob­ to C. intermedius, which is found on the tained by Laurence M. Huey on 18 March opposite side (eastern) ofthe Colorado Riv­ 1928. er from spinatus, along the Arizona-Cali­ Measurements ofholotype. - Total length, fornia boundary. C. spinatus differs from 204; length oftail, 118; length ofhind foot, these species in lacking or having a weakly 22; length ofear (crown), 5; greatest length developed, buffy, lateral stripe (see accounts of skull, 25.6; interorbital breadth, 6.6; of califomicus and fallax); from interme­ length of maxillary toothrow, 3.4; width dius, spinatus also differs in having more across mastoid bullae, 13.1; length ofnasals, numerous and more strongly developed 9.8. rump spines and usually flank spines as well, Distribution. - Ranges over the northern and in its larger size; other potentially sym­ three-fourths of the peninsula in Baja Cal­ patrie species ofChaetodipus all lack spines ifornia Sur, generally on the slopes of the in the pelage (arenarius, baileyi, formosus, Sierra de la Giganta. and penicillatus). Remarks. - Huey (1960b) gave means of 154 WILLIAMS ET AL. measurements of five males from the type Distribution. - Found only in Isla Mejia, locality. Gulf of California, Baja California. Remarks. -Burt (1932) and Banks (1967) gave means and ranges ofmeasurements for Chaetodipus spinatus bryanti 4 and 27 specimens, respectively. (Merriam, 1894)

1894. Perognathus bryanti Merriam, Proc. Cal­ ifornia Acad. Sci., ser. 2, 4:458, 25 September. Chaetodipus spinatus guardiae (Burt, 1932) Holotype. -Adult male, skin and skull, CAS 551/594, from San Jose Island, Gulf 1932. Perognathus spinatus guardiae Burt, ofCalifornia, Baja California Sur; obtained Trans. San Diego Soc. Nat. Hist., 7:165, 31 by Walter E. Bryant on 2 May 1892. October. Measurements ofholotype. - Total length, Holotype. -Adult male, skin and skull, 216; length oftail, 127; length ofhind foot, CIT 50495, from Puerto Refugio, north end 25; length ofear, 11; greatest length ofskull, Angel de la Guardia [= Guarda] Island, 30 26.65; width across bullae, 12.90; breadth ft, GulfofCalifornia, Baja California, Mexi­ across maxillary arches, 12.30; nasal length, co; obtained on 7 January 1932 by William 10.65; interorbital breadth, 6.7; length of H. Burt. maxillarytoothrow, 3.55; depth ofcranium, Measurements ofholotype. - Total length, 3.50. 164; length of tail, 89; length of hind foot, Distribution. - Known only from Isla San 22; length of ear (crown), 5, (notch, dry), Jose, Baja California Sur. 6.7; greatest length of skull, 24.1; basal Remarks.-Benson (1930) gave means length, 19.6; interorbital breadth, 6.1; length and ranges of measurements of 7 adults, of maxillary toothrow, 4.0; length of mas­ Burt (1932) listed means and ranges ofmea­ toid bulla, 6.9; width across mastoid bullae, surements for 23 specimens, and Osgood 12.1; length of interparietal, 3.0; width of (1900) gave means for 3 individuals, in­ interparietals, 7.0; length of nasals, 9.7. cluding the holotype. Distribution. -Occurs only on Isla Angel de la Guarda, Gulf ofCalifornia, Baja Cal­ Chaetodipus spinatus evermanni ifornia. (Nelson and Goldman, 1929) Remarks. -Burt (1932) listed means and extremes ofmeasurements of36 specimens; 1929. Perognathus evermanni Nelson and Banks (1967) gave means and ranges of Goldman, Proc. BioI. Soc. Washington, 42:111, measurements of 12 specimens. The holo­ 25 March. type is now in the UCLA collections. Holotype. -Adult male, skin and skull, CAS 3937, from Mejia Island, near north end of Angel de la Guardia [= Guarda] Is­ Chaetodipus spinatus lambi land, Gulf of California, Baja California; (Benson, 1930) obtained by V. W. Owen on 28 June 1921. Measurements ofholotype. - [External 1930. Perognathus spinatus lambi Benson, Univ. California PubI. ZooI., 32:452, 6 September. measurements from dry skin] Total length, 156; length of tail, 80; length of hind foot, Holotype. - Young adult female, skin and 20.7; greatest length of skull, 24.2; inter­ skull, MVZ 42938, from San Gabriel, Es­ orbital breadth, 5.6; length of maxillary piritu Santo Island, GulfofCalifornia, Baja toothrow, 3.4; width across mastoid bullae, California Sur, Mexico; obtained by Chester 11. 7; length of interparietal, 3.0; width of C. Lamb on 9 January 1929. interparietals, 6.5; length of nasals, 9.6; Measurements ofholotype. - Total length, width of nasals, 2.5. 175; length oftail, 105; length ofhind foot, TAXONOMY 155

23; length ofear (crown), 6; greatest length tained by Richard C. Banks on 22 October of skull, 24.85; interorbital breadth, 6.1; 1964. length of mastoid bulla, 6.85; width across Measurements ofholotype. - Total length, mastoid bullae, 11.85; length of interpari­ 169; length of tail, 93; length of hind foot, etal, 3.6; width ofinterparietals, 7.25; length 20; length ofear, 8; weight, 13.4 g; greatest of nasals, 9.85; width of rostrum, 3.9. length of skull, 23.9; interorbital breadth, Distribution. - Known only from Isla Es­ 6.1; length ofmaxillary toothrow, 3.5; width piritu Santo, Gulf of California, Baja Cali­ across mastoid bullae, 12.2; length ofnasals, fornia Sur. 9.4; depth of skull, 7.7. Remarks. -Benson (1930) and Burt Distribution. -Occurs on South and North (1932) listed means and ranges ofmeasure­ San Lorenzo islands, GulfofCalifornia, Baja ments for five and eight adults, respectively. California. Remarks. - Banks (1967) gave means and Chaetodipus spinatus latijugularis ranges of measurements for 20 specimens. (Burt, 1932) South San Lorenzo Island is listed on recent Mexican maps as Isla San Lorenzo; North 1932. Perognathus spinatus latijugularis Burt, San Lorenzo Island (28°42'N, 112°57'W) is Trans. San Diego Soc. Nat. Hist., 7:168, 31 also known as Isla Las Animas. October. Holotype. -Adult male, skin and skull, Chaetodipus spinatus magdalenae (Osgood, 1907) Donald R. Dickey collection (UCLA) 18020, from San Francisco Island, lat. 24°50'N, 1907. Perognathus spinatus magdalenae Os­ long. 1l0034'W, Gulf of California, Baja good, Proc. Bioi. Soc. Washington, 20:21, 23 California Sur, Mexico; obtained on 19 February. March 1930 by H. H. Sheldon. Measurements ofholotype. - Total length, Holotype. - Adult female, skin and skull, 188; length oftail, 110; length ofhind foot, USNM 146102, from Magdalena Island, 22; length ofear (notch, fresh), 9, (dry), 7.1; Pacific Ocean, Baja California Sur, Mexico; greatest length of skull, 25.7; basal length, obtained by Edward W. Nelson and Edward 21.2; interorbital breadth, 6.7; length of A. Goldman on 25 November 1905. maxillary toothrow, 4.0; length of mastoid Measurements of holotype. -Greatest bulla, 7.8; width across mastoid bullae, 12.7; length of skull, 26.4; basilar length, 17.9; length of interparietal, 3.7; width of inter­ interorbital breadth, 6.9; length ofmaxillary parietals, 7.5; length of nasals, 9.3. toothrow, 4.3; width across mastoid bullae, Distribution. - Known only from Isla San 12.6; length of interparietal, 3.6; width of Francisco, GulfofCalifornia, Baja Califor­ interparietals, 8.1; length ofnasals, 10.5; zy­ nia Sur. gomatic width, 12.8; length ofdiastema, 6.0. Remarks.-Burt (1932) gave means and Distribution. - Known only from Isla ranges for measurements ofnine specimens. Magdalena, Pacific Ocean, Baja California Sur. Remarks. -Osgood (1907) listed external Chaetodipus spinatus lorenzi measurements (means, extremes) for 10 (Banks, 1967) topotypes.

1967. Perognathus spinatus lorenzi Banks, Proc. Bioi. Soc. Washington, 80:101, 28 July. Chaetodipus spinatus macrosensis (Burt, 1932) Holotype. -Adult male, skin and skull, SDSNH 19901, from South San Lorenzo 1932. Perognathus spinatus macrosensis Burt, Island, lat. 28°36'N, long. 112°5l'W, Gulf Trans. San Diego Soc. Nat. Hist., 7:166, 31 of California, Baja California, Mexico; ob- October. 156 WILLIAMS ET AL.

Holotype. - Adult male, skin and skull, sured fresh); no other data on measure­ CIT 50604, from San Marcos Island, lat. ments are available for margaritae. 27°13'N, long. l12°05'W, Gulf of Califor­ nia, Baja California Sur, Mexico; obtained on 18 January 1932 by William H. Burt. Chaetodipus spinatus occultus (Nelson, 1912) Measurements ofholotype. - Total length, 174; length oftail, 102; length ofhind foot, 1912. Perognathus spinatus nelsoni Townsend, 22; length of ear (crown), 5, (notch, dry), Bull. Amer. Mus. Nat. Hist., 31:122,14 June. 7.8; greatest length of skull, 25.4; basal 1912. Perognathus spinatus occultus Nelson, length, 20.9; interorbital breadth, 6.2; length Proc. BioI. Soc. Washington, 25:116, 29 June. of maxillary toothrow, 3.6; length of mas­ Renaming of nelsoni Townsend, 1912. toid bulla, 7.5; width across mastoid bullae, 12.5; length of interparietal, 3.6; width of Holotype. -Male, skin and skull, USNM interparietals, 7.1; length of nasals, 9.8. 198409, from Carmen Island, Gulf of Cal­ Distribution. - Known only from Isla San ifornia, Baja California Sur, Mexico; ob­ Marcos, GulfofCalifornia, Baja California tained by H. E. Anthony on 3 April 1911. Sur. Measurements ofholotype. - Total length, Remarks. -Burt (1932) gave means and 170; length of tail, 88; length of hind foot, ranges of measurements of 17 topotypes. 24.5; greatest length of skull, 26.05; width The holotype is now in the UCLA collec­ across bullae, 12.35; breadth across maxil­ tions. lary arches, 12.20; nasal length, 10.50; in­ terorbital breadth, 6.45. Distribution. - Known only from Isla del Chaetodipus spinatus margaritae Carmen, GulfofCalifornia, Baja California (Merriam, 1894) Sur. Remarks.-Townsend (1912) gave aver­ 1894. Perognathus margaritae Merriam, Proc. age external measurements oftwo males and California Acad. Sci., ser. 2, 4:459, 25 Septem­ one female; Burt (1932) listed means and ber. ranges ofmeasurements for five specimens. Holotype. -Adult female, skin and skull, CAS 90, from Santa Margarita Island, Pa­ Chaetodipus spinatus oribates cific Ocean, Baja California Sur, Mexico; (Huey, 1960) obtained on 2 March 1889 by Walter E. Bryant. 1960. Perognathus spinatus oribates Huey, Measurements ofholotype. - [External Trans. San Diego Soc. Nat. Hist., 12:409, 1 February. measurements from dry skin] Total length, 170; length oftail, 102; length ofhind foot, Holotype. -Adult male, skin and skull, 22.5; length of ear (anterior base), 8.5; oc­ SDSNH 18742, from San Fernando Mis­ cipitonasal length, 25.9; basilar length of sion, lat. 30° N, Baja California, Mexico; Hensel, 18.0; interorbital breadth, 6.5; width obtained by Laurence M. Huey on 27 Feb­ across mastoid bullae, 12.0; length ofinter­ ruary 1958. parietal, 3.7; width of interparietals, 8.0; Measurements ofholotype. - Total length, length of nasals, 10.3. 192; length oftail, 112; length ofhind foot, Distribution. -Known only from Isla 22; length ofear (crown), 5; greatest length Santa Margarita, Pacific Ocean, Baja Cali­ of skull, 24.9; interorbital breadth, 6.5; fornia Sur. length of maxillary toothrow, 3.5; width Remarks. -Merriam (1894c) gave exter­ across mastoid bullae, 12.8; length ofnasals, nal measurements of two specimens (mea- 9.1. TAXONOMY 157

Distribution. -Occupies rocky terrain on Holotype. - Adult male, skin and skull, the lower slopes of the central mountain SDSNH 8450, from 25 mi N Punta Prieta, mass of the northern Baja California pen­ lat. 29°24'N, long. l14°24'W, Baja Califor­ insula; found in the Sierra San Pedro Martir nia, Mexico; obtained by Laurence M. Huey and the Sierra San Miguel, southward to the on 26 October 1930. vicinity of La Ramona, northeast of Santa Measurements ofholotype. - Total length, Catarina, all in Baja California. 194; length oftail, 112; length ofhind foot, Remarks.-Huey (1960b) listed averages 21; length ofear (crown), 5; greatest length for measurements of five specimens. of skull, 25.2; interorbital breadth, 5.9; length of maxillary toothrow, 3.3; width across mastoid bullae, 12.3; length ofnasals, Chaetodipus spinatus peninsulae 9.5. (Merriam, 1894) Distribution. - Ranges over rocky areas of the central Baja California peninsula, from 1894. Perognathus spinatus peninsulae Merri­ near San Augustin, Baja California, south am, Proc. California Acad. Sci., ser. 2, 4:460, to near Santa Gertrudis Mission, inland, and 25 September. El Barril on the Gulf coast. Holotype. - Young adult female, skin and Remarks. - Banks (1967) listed means and skull, CAS 274, from San Jose del Cabo, ranges of measurements for 22 specimens Baja California Sur, Mexico; obtained on and Huey (1930) gave means of measure­ 11 September 1891 by Walter E. Bryant. ments of 5 topotypes. Measurements ofholotype. - Total length, 198; length oftail, 107; length ofhind foot, 23; length of ear (anterior base, dry), 9; Chaetodipus spinatus pullus greatest length ofskull, 26.60; width across (Burt, 1932) bullae, 12.80; breadth across maxillary arches, 13.25; nasal length, 10.05; interor­ 1932. Perognathus spinatus pullus Burt, Trans. bital breadth, 6.60; length of maxillary San Diego Soc. Nat. Hist., 7:166, 31 October. toothrow, 3.50; depth of cranium, 3.55. Distribution. -Occupies the Baja Califor­ Holotype. -Adult male, skin and skull, nia peninsula in the cape region from ap­ CIT 50324, from Coronados Island, lat. proximately the level of Bahia de la Paz to 26°06'N, long. 111018'W, Gulf of Califor­ the southern end, on both coasts (Hall, nia, Baja California Sur, Mexico; obtained 1981 ). by William H. Burt on 20 December 1931. Remarks. -Benson (1930) listed means Measurements ofholotype. - Total length, and ranges ofmeasurements 00 specimens, 192; length oftail, 111; length ofhind foot, Huey (1930, 1960b) listed means of 12 and 23; length ofear (crown), 6 (notch, dry), 8.2; 5specimens, respectively, from different lo­ greatest length of skull, 25.8; basal length, calities, and Osgood (1900) gave means for 21.3; interorbital breadth, 6.4; length of 4 specimens. maxillary toothrow, 3.6; length of mastoid bulla, 7.3; width across mastoid bullae, 12.6; length of interparietal, 4.1; width of inter­ Chaetodipus spinatus prietae parietals, 7.8; length of nasals, 10.0. (Huey, 1930) Distribution. - Found only on Isla Coro­ nados, Gulf of California, Baja California 1930. Perognathus spinatus prietae Huey, Trans. Sur. San Diego Soc. Nat. Hist., 6:232, 24 Decem­ Remarks. -Burt (1932) listed means and ber. ranges of measurements for seven speci- 158 WILLIAMS ET AL. mens. The holotype is now in the UCLA length of interparietal, 3.6; width of inter­ collections. parietals, 7.6; length of nasals, 10.1. Distribution. -Occurs only on Isla Dan­ zante Primero, GulfofCalifornia, Baja Cal­ Chaetodipus spinatus rufescens ifornia Sur. (Huey, 1930) Remarks. -Burt (1932) provided means and extremes of measurements of seven 1930. Perognathus spinatus rufescens Ruey, specimens. The holotype is now in the Trans. San Diego Soc. Nat. Rist., 6:231, 24 UCLA collections. December. Holotype. - Adult male, skin and skull, Chaetodipus spinatus spinatus SDSNH 7446, from mouth of Palm Can­ (Merriam, 1889) yon, Borego Valley, San Diego Co., Cali­ fornia; obtained by Laurence M. Huey on 1889. Perognathus spinatus Merriam, N. Amer. 10 November 1929. Fauna, 1:21, 25 October. Measurements ofholotype. - Total length, Holotype. -Adult male, skin and skull, 194; length oftail, 112; length ofhind foot, USNM 186516, from lower Colorado Riv­ 21; length ofear (crown), 5; greatest length er, 25 mi below [S] the Needles, San Ber­ of skull, 25.2; interorbital breadth, 5.9; nardino Co., California; obtained by Vernon length of maxillary toothrow, 3.3; width Bailey on 23 March 1889. across mastoid bullae, 12.3; length ofnasals, Measurements ofholotype. - Total length, 9.5. 179; length oftail, 104; length ofhind foot, Distribution. -Rocky, desert slopes ofthe 21; length of ear (crown, dry), 3.5; occipi­ southern California coastal ranges, from near tonasal length, 23.7; basilar length, 18.8; Palm Springs, Riverside Co., south at least basilar length ofHensel, 16.5; width across to the Mexican Boundary (Hall, 1981). mastoid bullae, 12.2; length ofinterparietal, Remarks.-Huey (1930, 1960b) listed 3.5; width ofinterparietals, 7.3; least inter­ means for measurements ofeight specimens mastoid distance, 8.0. from San Diego Co., California. Distribution. - Ranges along the Colora­ do River, from Granite Springs in extreme southern Clark Co., Nevada, south to near Chaetodipus spinatus serosus Fort Yuma, Imperial Co., California, and (Burt, 1932) southward on rocky slopes in northeastern Baja California to the vicinity ofSan Felipe 1932. Perognathus spinatus serosus Burt, Trans. on the Gulf of California. San Diego Soc. Nat. Rist., 7:167, 31 October. Remarks.-Huey (1930, 1960b) and Os­ Holotype. -Adult male, skin and skull, good (1900) listed means for five and four CIT 50307, from Danzante Island [= Isla specimens, respectively. Danzante Primero], 1at. 25°47'N, long. 111°ll'W, GulfofCalifornia, Baja Califor­ nia Sur, Mexico, on 17 December 1931 by William H. Burt. Genus Perognathus Measurements ofholotype. - Total length, 187; length oftail, 104; length ofhind foot, 1839. PerognathusWied, Nova Acta Phys.-Med. Acad. Caesar. Leop.-Carol., 19(1 ):368. 22; length ofear (crown), 5, (notch, dry), 7; greatest length of skull, 25.7; basal length, Type Species. -Perognathus fasciatus 21.4; interorbital breadth, 6.7; length of Wied, 1839, Nova Acta Phys.-Med. Acad. maxillary toothrow, 3.6; length of mastoid Caesar. Leop.-Carol., 19(1 ):368. bulla, 7.2; width across mastoid bullae, 12.3; Diagnosis. -Size small, total length from TAXONOMY 159 about 100 to 200 mm, weight from about Remarks. - This diagnosis is based on 5 to 30 g; body form quadrupedal and scan­ Recent species. The genus Perognathus in­ sorial; hind limbs longer than forelimbs; tail cludes as synonyms Cricetodipus Peale, 1848 relatively short, length usually averaging less (type species C. parvus), a name that also to slightly more than length of head and was subsequently applied, for a time, to some body; tail without prominent distal, dorsal species ofDipodomys; Abromys Gray, 1868 crest or terminal tuft ofhairs, although sev­ (type species A. lordi, a synonym ofP. par­ eral species have a slight crest and terminal vus), and Dtognosis Coues, 1875 (type spe­ tuft (pencil); tail some shade of brown or cies D. /ongimembris). Considerable taxo­ buffy above, whitish below; proximal one­ nomic confusion arose as a result of S. F. fourth to one-half of sole ofhind foot with Baird, E. Coues, J. E. Gray, T. R. Peale, and sparse covering of short hairs; ear pinna others misapplying the names Perognathus short and rounded, and without a lobed an­ fasciatus, Perognathus hispidus, Abromys titragus except in P. parvus and P. a/tico/a; lordi, and Cricetodipus parvus. Merriam antero-lateral edge of ear pinna without (1889) and Osgood (1918) were largely re­ covering of long, coarse hairs; generally a sponsible for clarifying the nomenclature contrasting, light buffy area on the head and stabilizing the taxonomy ofpocket mice. around ear pinna (post-auricular patch), and As conceived herein, the genus Perognathus a small, whitish spot below the external au­ does not include the hispid-haired species ditory meatus; dorsal surfaces generally of Chaetodipus nor Perognathus formosus some shade of buffy or brownish, usually Merriam, which has been shown to be a tinged with black; usually a clear, buffy lat­ species of Chaetodipus (Homan and Gen­ eral stripe without blackish tinge; under­ oways, 1978; Patton et aI., 1981). sides usually whitish; in general, hairs rel­ Most ofthe structural characteristics that atively short, soft, and oval to flattened in are of value in distinguishing species are cross section; hairs lack dorsal trough ex­ related to size, proportions, and color, all cept, in so far as is known, P. amp/us; lon­ ofwhich exhibit considerable individual and ger, stiff, spine-like hairs never present in geographic variation, making identification pelage of dorsal and lateral surfaces; mas­ keys based on skins and skulls cumbersome. toid bullae usually extending beyond the plane ofthe occiput; tympanic bullae nearly meeting anteriorly on the ventral surface of Key to the Species the skull; exoccipital without strong lateral indentations of mastoid bullae; interpari­ 1. Antitragus of ear pinna lobed 2 1'. Antitragus of ear pinna not lobed 3 etals compressed and narrower than inter­ 2. Inner surface ofear pinna with white or orbital breadth; phallus relatively short in yellowish hairs; distal one-third of tail length; soft tissue of phallus extends about with blackish hairs dorsally; mastoid two-thirds of the length of the baculum; bulla forms small, sharp indentation in phallus lacks external spines but has ure­ exoccipital; occurring only in southern thral lappets; baculum relatively short in California in the Transverse ranges and length, with swollen, bulbous proximal end the San Bernardino Mountains . and slender, slightly upturned distal end; ...... Perognathus alticola vesicular glands of male reproductive sys­ 2'. Hairs ofinner surface ofear pinna buffy tem elongated and tube-like with hooked rather than white or yellowish; distal end and translucent; head of spermatozoa one-third oftail with mix ofsooty brown and black hairs dorsally; mastoid bulla approximating a triangle, with rounded ver­ with little or no indentation into exoc­ tices; tendon at origin of M. rectus femoris cipital; occurring on the Columbia Pla­ fan-shaped (Hafner and Hafner, 1983; Hall, teau, in the Great Basin, and the eastern 1981; Homan and Genoways, 1978; Ryan, slopes ofthe Sierra Nevada . 1989; Wood, 1935)...... Perognathus parvus 160 WILLIAMS ET AL.

3. Length of tail averages less than length yellow (olivaceous) tone; length of in­ ofhead and body (a few individuals have terparietal generally less than 3.0 mm; tails longer than length of head and length of head and body averaging body); not occurring west ofeastern Utah greater than 68 mm; not occurring south or west-central Arizona, but found in and east of western Nebraska on the Sonora along the Gulf coast 4 Great Plains and southeastern Colorado 3'. Length of tail averages greater than along the eastern front of the Rocky length ofhead and body (a few individ­ Mountains Perognathus fasciatus uals have tails shorter than head and 7'. Dorsal yellowish color with yellowish­ body); not occurring east ofcentral Ar- orange tone; length ofinterparietal gen­ izona and south-central Utah 9 erally greater than 3.0 mm; length of 4. Interparietal length less than 2.9 mm; head and body averaging less than 66 length of tail less than 66 mm (mean mm; not occurring north and west ofa less than 60 mm); interorbital breadth line from approximately the east base averages 4.7 mm or less and maximum of the Rocky Mountains in northern less than 5.0; width ofinterparietals less Colorado, north and east through south- than 4.2 mm, averaging 3.6 or less. . .. 5 central North Dakota . 4'. Interparietal length greater than or equal · Perognathus jlavescens to 2.9 mm; length oftail averages great­ 8. Dorsal yellowish color with an olive­ er than 60 mm; interorbital breadth 4.5 yellow (olivaceous) tone; least interbul­ mm or more, mean 4.8 or greater; width lar distance (on dorsal surface of skull) ofinterparietals 3.15 mm or more, mean averaging 4.3 mm or more; width offirst 4.5 or greater 6 upper molar 1.16 mm of less (mean 5. Length oftail averages 56 mm or more; 1.10); not occurring south ofthe Uintah interorbital breadth averages 4.5 or Basin of Utah and Colorado . more; width of interparietals averages · Perognathus fasciatus 3.3 mm or more; pelage sleek, not no­ 8'. Dorsal yellowish color with yellowish­ ticeably lax; dorsal color yellowish tinged orange tone; least interbullar distance with blackish from black-tipped hairs; averaging less than 4.0 mm, generally slight contrast between the darker mid­ not exceeding 4.3; width of first upper dorsal and lighter dorso-lateral color; molar 1.13 mm or more (mean 1.22); post-auricular light spot relatively small not occurring north ofthe Uintah Basin ...... Perognathus merriami of Colorado and Utah . 5'. Length of tail averages 55 mm or less; · Perognathus jlavescens interorbital breadth averages 4.5 mm or 9. Not occurring in California 10 less; width ofinterparietals averages 3.3 9'. Occurring in California 11 or less; pelage lax, not sleek in appear­ 10. Greatest length of skull generally less ance; dorsal color buffy-yellow with a than 23.0 mm; frontonasal length av­ pinkish hue and a tinge ofblackish from eraging less than about 15 mm; length black-tipped hairs; dark mid-dorsal area ofhind foot 20 mm or less (mean 19 or contrasting markedly with lighter dor­ less) Perognathus longimembris so-lateral color; post-auricular light spot 10'. Greatest length ofskull generally greater relatively large Perognathus jlavus than 23.0 mm; frontonasal length av­ 6. Occurring on the Great Plains, or in the eraging greater than 15.1 mm; length of northern Chihuahuan Desert region in hind foot 19 mm or more (mean greater southern Arizona and New Mexico, than 20) Perognathus amplus western Texas, and northern Chihuahua.. 7 11. Occurring in central California in the 6'. Occurring in the intermountain pla­ San Joaquin and Sacramento valleys, teaus and basins of south-central and north of the Tehachapi Mountains ... southwestern Wyoming, eastern Utah, · Perognathus inornatus southern and western Colorado, eastern 11 '. Occurring in southern California from Arizona, and New Mexico 8 the Tehachapi Mountains southward 12 7. Dorsal yellowish color with an olive- 12. Occipitonasal length of adults (perma- TAXONOMY 161

nent upper premolar with moderate to Distribution. -Occupies arid shrub and heavy wear) generally greater than 22.5 forest communities in south-central Cali­ mm Perognathus inornatus fornia, in the Transverse Ranges of Kern 12'. Occipitonasal length of adults (see and Los Angeles counties, and the San Ber­ above) generally less than 22.1 mm .. nardino Mountains, San Bernardino Co...... Perognathus longimembris Remarks. - Perognathus alticola is close­ ly related to, and perhaps only subspecifi­ cally distinct from, P. parvus. The members Species Accounts of the parvus species group exhibit rela­ tively great diversity in chromosome struc­ Perognathus alticola ture (Williams, 1978a) and biochemical variation (Sulentich, 1983). Sulentich (1983) Diagnosis. -Size medium-large for ge­ considered P. a. alticola to be specifically nus, total length from about 130 to 180; distinct from P. a. inexpectatus, primarily greatest length ofskull from about 23 to 26 on the basis of the size and shape of the mm; size small compared to most subspe­ interparietals. Greater variation, however, cies of P. parvus; length of tail equal to or is seen in size and proportions ofthe inter­ slightly longer than length ofhead and body; parietals ofP. parvus (for example, those of tail slightly crested for distal one-third; up­ P. p. bullatus are extremely compressed per side of tail shading to blackish toward compared to those ofadjacent populations tip; antitragus ofear pinna lobed; inner side ofP. p. clarus in eastern Utah; D. Williams, of ear pinna with white or yellowish hairs; unpubl. data), and variation in size and auditory bullae relatively small; posterior shape of the interparietals in P. flavescens border of mastoid bulla about even with and P. fasciatus was shown to be strongly posterior extent of supraoccipital; mastoid related to degree ofbullar inflation and sig­ bulla forms small indentation in border of nificantly correlated with degree of envi­ exoccipital on back of skull; interparietal ronmental aridity (Williams, 1978b; Wil­ compressed compared to most subspecies liams and Genoways, 1979). ofP. parvus; ascending branches ofsupraoc­ cipital relatively broad; interorbital region broad; phallus and baculum, in so far as is Perognathus alticola alticola known, relatively short compared to P. par­ Rhoads, 1894 vus, and relatively short, compared to head and body length, for genus (Burt, 1936, 1960; 1894. Perognathus altieolus Rhoads, Proc. Acad. Nat. Sci. Philadelphia, 45:412, 27 January. Osgood, 1900; Rhoads, 1894; Sulentich, 1983). Holotype. -Adult male, skin and skull, Comparisons. - Perognathus alticola is ANSP 1615, from Squirrel Inn, near Little most similar to P. parvus, from which it can Bear Valley, 5,500 ft, San Bernardino be distinguished only by its smaller average Mountains, San Bernardino Co., California; size, having white or yellowish hairs on the obtained on 22 September 1893 by R. B. inner surface ofthe pinna, having a tail with Herron. more blackish hairs on the dorsal surface of Measurements ofholotype. - Total length, its distal one-third, and by having a more 157; length of tail, 77; length of hind foot, pronounced indentation ofthe mastoid bul­ 20; length of ear (crown), 5 (external mea­ la into the border of the exoccipital. P. al­ surements from dry skin); basilar length of ticola can be distinguished from all other skull, 16.0; width across bullae, 12.5; inter­ species ofPerognathus by its lobed antitra­ orbital breadth, 6.0; length of nasals, 8.6; gus and more prominent crest on the distal length ofmandible, 10.5; height ofcoronoid portion of the tail. process, 4.6. 162 WILLIAMS ET AL.

Distribution. - Known only from arid ly). This, together with the more inflated ponderosa pine communities in the vicinity bullae and compressed interparietals, and of Little Bear Valley and Strawberry Peak, the shorter phallus and baculum of P. al­ San Bernardino Mountains, San Bernardino ticola inexpectatus, led him to conclude that Co., California. inexpectatus was specifically distinct from Remarks. - The specific epithet, alticolus, both P. parvus and P. alticola. He did not constructed by Rhoads (1894), apparently have data on biochemical variation or phal­ referred to the mountainous home of this lic structure ofP. alticola alticola, however. species (altus = high; cola = inhabiting). His Although we concur that individuals of alteration of the word cola to conform in inexpectatus are distinct from P. parvus gender to Perognathus was in error, how­ samples from adjacent geographic regions, ever. Cola is not based on the root, col; the we think it best to retain the current taxo­ latter is a Latin prefix to be used before r nomic arrangement until information on and meaning "with" or "together." Osgood other populations of P. parvus and P. alti­ (1900) amended the spelling without com­ cola is available. Sulentich (1983) provided ment, although Rhoads' spelling is still used means and ranges of measurements for 20 by some. speCImens.

Perognathus alticola inexpectatus Perognathus amplus Huey, 1926 Diagnosis. -A medium-sized species of 1926. Perognathus alticola inexpectatus Huey, Perognathus with a relatively long tail, large Proc. Bioi. Soc. Washington, 39:121, 27 De­ hind feet, inflated mastoid bullae, and com­ cember. pressed interparietals; head and body length Holotype. -Adult male, skin and skull, normally ranges between about 71 and 88 SDSNH 5724, from 14 mi W Lebec, 6,000 mm; length oftail generally ranges between ft, Kern Co., California; obtained by George about 72 and 88, and the ratios oflength of G. Cantwell on 28 August 1926. tail to head and body range between about Measurements ofholotype. - Total length, 0.88 and 1.33; length ofhind foot generally 176; length of tail, 97; length of hind foot, measures 20 mm or more; width of skull 22; greatest length ofskull, 25.9; width across varies from about 12.8 to 15.2 mm and bullae, 13.8; breadth across maxillary arch­ width ofinterparietal from 3.2 to 4.45 mm. es, 13.0; length of nasals, 10.0; interorbital Comparisons. - Perognathus amplus is breadth, 6.2; length of maxillary toothrow, most similar to P. inornatus and P. longi­ 3.4. membris. No single character ofskin or skull Distribution. -Occupies arid shrub-steppe is known that will distinguish all individuals communities in the Tehachapi Mountains of P. amplus from these other species; am­ of south-central California, from the vicin­ plus averages larger than inornatus in most ity of Tehachapi Pass, Kern Co., on the dimensions, has a relatively longer and more northeast to the vicinity ofMt. Pinos, Ven­ penciled tail and an interorbital breadth in tura Co. on the northwest, and Elizabeth excess of 5.1 mm (average between about and Quail lakes, Los Angeles Co., on the 5.3 and 5.6), whereas interorbital breadth south (unpubl. data). in inornatus averages between about 4.75 Remarks. - Sulentich (1983) found rela­ and 4.9, and individuals rarely exceed 5.1 tively great biochemical differences between mm. P. amplus differs from P. longimem­ P. a. inexpectatus and P. parvus olivaceus bris in larger average size and in having a and P. p. xanthonotus (Rogers' similarity wider upper premolar; greatest length of coefficients of0.404 and 0.402, respective- skull generally exceeds 23.2 mm in amplus TAXONOMY 163 and less than 23.1 in /ogimembris; length of Distribution. -Occupies central and hind foot usually measures 20 mm or more southwestern Arizona, from Fort Verde, in amp/us and 19.5 or less in /ogimembris. Yavapai Co., southeastward to San Carlos P. amp/us is also similar in size and general Lake, Gila Co., andwestward to near Signal, appearance to some subspecies ofP. jlaves­ Mojave Co., thence southward through cens; the tail ofamp/us is longer relative to central Yuma Co. into coastal Sonora to the head and body length and is more penciled, vicinity of Puerto Libertad. the interparietal is narrower, and the length Remarks. - Hoffmeister (1986) placed ro­ ofhind foot is greater (20 mm or more ver­ tundus andjacksoniin the synonymy ofam­ sus 20 or less). p/us, and provided statistics for measure­ Distribution. -Occurs as apparently dis­ ments of 190 specimens in 6 samples. junct populations in north-central and west­ Benson (1933) gave measurements of one central and southwestern Arizona, and ad­ topotype of amp/us; Goldman (1933) gave jacent areas in northwestern Sonora (Hall, measurements offour adults from Yavapai 1981 ). Co.; and Goldman (1932a) listed average Remarks. - The relationships among the measurements for five adults from Yuma taxa ofthe /ongimembris species group (the Co. nominate forms, amp/us, inornatus, and /ongimembris) are poorly known and a new taxonomy probably will be required when Perognathus amp/us cineris these relationships are clarified. Hoffmeis­ Benson, 1933 ter(1986) discussed ways to distinguish am­ p/us from /ogimembris in Arizona. 1933. Perognathus amp/us cineris Benson, Proc. Biol. Soc. Washington, 46:109, 27 April. 1933. Perognathus amp/us ammodytes Benson, Proc. Biol. Soc. Washington, 46: 110, 27 April. Perognathus amp/us amp/us Osgood, 1900 H%type. -Adult male, skin and skull, MVZ 55771, from near the Wupatki Ruins, 1900. Perognathus amp/us Osgood, N. Amer. Wupatki National Monument (about 27 mi Fauna, 18:32, 20 September. NE Flagstaff), Coconino Co., Arizona; ob­ 1932. Perognathus amp/us rotundus Goldman, tained by Annie M. Alexander on 12 Oc­ J. Washington Acad. Sci., 22:387, 19 July. tober 1932. 1933. Perognathus amp/us jacksoni Goldman, Measurements ofh%type. - Total length, 1. Washington Acad. Sci., 23:465, 15 October. 197; length oftail, 107; length ofhind foot, H%type. -Adult male, skin and skull, 20; length of ear (crown), 4; weight, 10.3 g; USNM 34626/46711, from Fort Verde, Ya­ greatest length of skull, 23.45; interorbital vapai Co., Arizona; obtained by J. Alden breadth, 5.8; length of mastoid bulla, 8.85; Loring on 26 June 1892. width across mastoid bullae, 12.85; length Measurements ofh%type.- Total length, ofinterparietal, 2.3; width ofinterparietals, 155; length of tail, 80; length of hind foot, 3.8; length ofnasals, 8.7; width ofrostrum, 20; greatest length of skull, 25.05; occipi­ 4.0. tonasallength, 23.65; interorbital breadth, Distribution. - Found south of the Colo­ 5.20; length of maxillary toothrow, 3.55; rado River, from near Navajo Spring, Echo width across maxillary toothrows, 5.20; Cliffs, Coconino Co., on the north, south­ length of mastoid bulla, 9.85; width across ward along the Little Colorado River Valley mastoid bullae, 13.90; length of interpari­ to the vicinity of Wupatki National Mon­ etal, 2.70; width of interparietals, 3.20; ument, Coconino Co., Arizona. length ofnasals, 9.40; width ofnasals, 2.00; Remarks. -Hoffmeister (1986) consid­ width of rostrum, 3.95. ered ammodytes to be indistinguishable 164 WILLIAMS ET AL. from cineris except for the darker color of serve (near Northeast Station), 35 mi S Tuc­ populations in the vicinity of the type lo­ son, about 4,000 ft, Pima Co., Arizona' ob­ cality. Hoffmeister (1986) listed statistics for tained on 3 August 1930 by Walter P. Ta~lor. measurements of samples of 20 and 32; Measurements ofholotype. - Total length, Benson (1933) gave measurements for 7 155; length of tail, 84; length of hind foot, specimens from the vicinity ofWupakti Na­ 20; greatest length of skull, 24.35; occipi­ tional Monument. tonasallength, 23.35; interorbital breadth 5.30; length of maxillary toothrow, 3.30~ width across maxillary toothrows, 4.20; Perognathus amplus pergracilis length of mastoid bulla, 9.00; width across Goldman, 1932 mastoid bullae, 13.35; length of interpari­ etal, 3.15; width of interparietals, 3.35; 1932. Perognathus amplus pergracilis Gold­ length ofnasals, 9.20; width ofnasals, 2.25; man, J. Washington Acad. Sci., 22:387,19 July. width of rostrum, 3.55. Distribution. - Desert areas ofsouth-cen­ Holotype. -Adult male, skin and skull, tral Arizona, south ofthe Gila River in Pin­ USNM 227528, from Hackberry, 3,500 ft, al Co. (and probably southeastern Maricopa Mohave Co., Arizona; obtained on 14 Sep­ Co.), southward east of Organ Pipe Cactus tember 1917 by Edward A. Goldman. National Monument to 13 mi W of Cabor­ Measurements ofholotype. - Total length, ca, Sonora (Hoffmeister, 1986). 143; length of tail, 80; length of hind foot, Remarks.-Goldman (1932b) gave av­ 21; greatest length of skull, 22.60; occipi­ erages and ranges of measurements for 10 tonasal length, 22.00; interorbital breadth, adult topotypes; Hoffmeister (1986) gave 5.35; length of maxillary toothrow, 3.50; statistics for measurements of samples of width across maxillary toothrows, 4.25; 14 and 26 individuals. length of mastoid bulla, 7.85; width across mastoid bullae, 12.40; length of interpari­ etal, 2.90; width of interparietals, 3.80; Perognathus fasciatus length ofnasals, 8.65; width ofnasals, 1.95; width of rostrum, 2.65. Diagnosis. -A medium-sized species of Distribution. -Occurs in desert associa­ Perognathus, with a relatively short, non­ tions in northwestern Arizona, from south penciled tail and olivaceous (olive-yellow) of the Grand Canyon, Mohave Co., south­ tone to the dorsal color; skull with relatively ward to near Wikieup, Mohave Co. small auditory bullae and wide interorbital Remarks. - Hoffmeister (1986) gave region. Width of lower premolar relatively measurements and associated statistics for wider than other species of Perognathus, samples of 17,49, and 38; Goldman (1932a) ranging from 0.52 to 0.77 mm (mean, 0.66). listed measurements for 5 adult topotypes, Length ofhead and body averages between and Benson (1933) gave statistics for mea­ about 68.9 and 70.2 mm; length oftail av­ surements of 18 specimens. erages between about 59.3 and 64.3 mm; interorbital breadth ranges between 4.5 and 5.6 mm with means ranging from about 4.9 Perognathus amplus taylori to 5.1; length ofmastoid bulla averages from Goldman, 1932 about 7.6 to 8.5 mm (Williams and Geno­ ways, 1979). 1932. Perognathus amplus taylori Goldman, J. Comparisons. - Perognathus fasciatus is Washington Acad. Sci., 22:488, 19 October. most similar to some populations ofPerog­ Holotype. - Adult female, skin and skull, nathus flavescens, from which it can be dis­ USNM 250533, from Santa Rita Range Re- tinguished by its unique, olivaceous dorsal TAXONOMY 165 color. Comparisons of cranial characters oming; obtained on 14 May 1897 by J. Al­ must be made by subspecies: P. fasciatus den Loring. callistus differs from P. jlavescens caryi by Measurements ofholotype. - Total length, smaller size, shorter interparietal, narrower 135; length of tail, 63; length of hind foot, rostrum, and wider interbullar region; P. 18; greatest length of skull, 22.80; occipi­ fasciatus fasciatus differs from Great Plains tonasallength, 22.65; interorbital breadth, populations ofP. jlavescens by being larger, 5.05; length of maxillary toothrow, 3.10; with a narrower interorbital breadth, larger width across maxillary toothrows, 4.45; auditory bullae, shorter interparietals, and length of mastoid bulla, 8.55; width across larger molariform teeth. P. fasciatus is sig­ mastoid bullae, 13.10; length of interpari­ nificantly larger with less inflated auditory etal, 2.70; width of interparietals, 4.65; bullae and wider interbullar region than P. length ofnasals, 8.10; width ofnasals, 2.35; flavus ofthe Great Plains area. From P. par­ width of rostrum, 3.55. vus, P. fasciatus is distinguished by its sig­ Distribution. -Occupies desert and steppe nificantly smaller size (greatest length ofskull grassland associations in the Uintah, Bridg­ less than 25 versus greater than 25.5 mm) er, and Great Divide basins and contiguous and non-penciled tail. No other species of areas of Colorado, Utah, and Wyoming Perognathus occurs within or near the geo­ (Williams and Genoways, 1979). graphic range of P. fasciatus (Williams and Remarks. - Most populations included in Genoways, 1979). P. fasciatus litus prior to the review by Wil­ Distribution. -Occupies grasslands ofthe liams and Genoways (1979) were regarded northern Great Plains from southeastern by them as inseparable from P. f callistus; Alberta, southern Saskatchewan, and south­ however litus is a synonym of P. fasciatus western Manitoba southward through fasciatus. northwestern Nebraska and east-central Colorado along the east flank of the Rocky Mountains to Huerfano Co. On the west, Perognathus fasciatus fasciatus ranging to central Montana, western Wyo­ Wied, 1839 ming, and the Uintah Basin ofnortheastern Utah and northwestern Colorado (Williams 1839. Perognathus fasciatus Wied, Nova Acta and Genoways, 1979). Phys.-Med., Acad. Caesar. Leop.-Carol., 19: Remarks. - Williams (1978a), redefined 369. thefasciatus species group ofOsgood (1900) 1893. Perognathus infraluteus Thomas, Ann. to exclude P. jlavus; thus it consists of P. Mag. Nat. Hist., ser. 6, 11:406, May. fasciatus and P. jlavescens. Williams and 1911. Perognathus fasciatus litus Cary, Proc. Genoways (1979) gave means and ranges of Biol. Soc. Washington, 24:61, 22 March. measurements for four samples of P. fas­ 1940. Perognathus jlavescens olivaceogriseus ciatus and provided a comprehensive, sys­ Swenk, Missouri Valley Fauna, 3:6, 5 June. tematic review of the species. Neotype. -Adult male, skin and skull, USNM 168599, from Buford, Williams Co., North Dakota; obtained on 6 May 1910 by Perognathus fasciatus callistus H. E. Anthony. Osgood, 1900 Measurements ofneotype. - Tota11ength, 140; length of tail, 66; length of hind foot, 1900. Perognathus callistus Osgood, N. Amer. 18; greatest length of skull, 23.35; occipi­ Fauna, 18:28, 20 September. tonasallength, 23.35; interorbital breadth, Holotype. - Adult male, skin and skull, 5.30; length of maxillary toothrow, 3.20; USNM 88245, from Kinney Ranch [about width across maxillary toothrows, 4.45; 22 mi S Bitter Creek], Sweetwater Co., Wy- length of mastoid bulla, 7.70; width across 166 WILLIAMS ET AL. mastoid bullae, 11.90; length of interpari­ ulations of P. m. gilvus; from P. merriami etal, 2.65; width of interparietals, 4.90; and P. jlavus, P. jlavescens can be distin­ length ofnasals, 8.75; width ofnasals, 2.15; guished by its wider interparietals (width width of rostrum, 4.00. averages greater than 4.0 mm in Great Plains Distribution. -Occupies light, sandy soils populations of jlavescens versus less than in the northern Great Plains, from near Sas­ 3.6 in P. merriami and P. jlavus); P. jlavus katoon, Saskatchewan, southward to near generally has a darker color dorsally due to La Veta, Huerfano Co., Colorado, and from numerous black-tipped guard hairs, and its near the North Dakota-Minnesota bound­ post-auricular patch appears larger and con­ ary westward through Wyoming, central trasts more strongly with surrounding areas Montana, and southeastern Alberta (Wil­ than in P. jlavescens; P. jlavescens generally liams and Genoways, 1979). averages larger, with a longer tail and hind Remarks. - There is no information on foot than P. merriami and P. jlavus; differ­ the disposition of the holotype of Wied ences in size are most pronounced in sym­ (1839). Williams and Genoways (1979) pre­ patric populations from the intermountain sumed that it was lost and therefore desig­ basins of southwestern North America. P. nated a neotype from near the type locality jlavescens is marginally sympatric with P. (upper Missouri River near junction with parvus in eastern Utah; jlavescens is smaller the Yellowstone River, near Buford, Wil­ (length ofhind foot and greatest skull length liams Co., North Dakota). They chose a less than 21 and 25.5 mm injlavescens, re­ specimen different from the "duplicate type" spectively, versus greater than 21 and 25.5 of Merriam (1889) because the specimen mm in parvus) and lacks the small crest on chosen by Merriam was not from the orig­ the distal, dorsal one-third of the tail (pen­ inal type locality and because the right, low­ cil) exhibited by parvus. P. jlavescens is mar­ er premolar exhibited a unique anomaly. ginally sympatric with P. amp/us in north­ central Arizona and possibly sympatric with Perognathus jlavescens P. /ongimembris in north-central Arizona and south-central Utah;jlavescens has a rel­ Diagnosis. -A small- to medium-sized atively shorter and less penciled tail than species of Perognathus with a tail typically amp/us or /ongimembris (ratio oflengths of shorter than length ofhead and body (ratio tail to head and body averages 0.95 or less oflengths oftail to head and body averages for jlavescens versus greater than 1.10 for from about 0.86 to 0.97), relatively broad amp/us and /ongimembris); amp/us is gen­ interparietals, and wide interorbital region. erally larger, with a length of hind foot 20 Dorsal yellowish color lacks an olivaceous mm orgreater versus 20 orless injlavescens; tone. Length ofhead and body ranges from interparietal width averages wider in jla­ about 60 to 73 mm; length of tail ranges vescens than amp/us from nearby popula­ from 56 to 73 mm; interorbital breadth tions (3.9 mm or greater versus about 3.5). ranges from about 4.5 to 5.6 mm; interpa­ From /ongimembris, nearby populations of rietal width varies from 3.15 to 5.4 mm jlavescens differ in larger average body size, (Williams, 1978b; Williams and Genoways, reflected in most external and cranial mea­ 1979). surements (Williams, 1978b; Williams and Comparisons. - Perognathus jlavescens is Genoways, 1979; D. Williams, unpubl. most similar to Perognathus fasciatus (see data). account offasciatus for distinguishing char­ Distribution. -Occupies sandy soils in acteristics). Great Plains populations exhib­ grassland and desert communities of the it similarity in external size and color to Great Plains and intermountain basins of some P. merriami merriami and some pop- west-central North America, ranging from TAXONOMY 167 southeastern North Dakota south and east Perognathus jlavescens apache nearly to the Mississippi River in southern Merriam, 1889 Minnesota; on the east extending from Min­ nesota southwesterly through the extreme 1889. Perognathus apache Merriam, N. Amer. northwestern comer of Missouri, central Fauna, 1:14,25 October. Kansas and central Oklahoma; on the west, 1918. Perognathus apache cleomophila Gold­ extending southwesterly from southeast man, Proc. BioI. Soc. Washington, 31:23, 16 May. North Dakota to extreme southeastern Wy­ oming and then southward through eastern Holotype. -Adult male, skin and skull, Colorado and New Mexico to western Tex­ USNM 186504, from near Kearn's Canyon, as; also occupies the intermountain basins Navajo Co., Arizona; obtained by Jere Sul­ and plateaus, from the Uintah Basin of livan on 22 May 1888. northeastern Utah and northwestern Col­ Measurements ofholotype. - Total length, orado, southward to northeastern Arizona 140; length of tail, 68; length of hind foot, and most ofcentral and western New Mex­ 18.5; greatest length of skull, 24.05; occip­ ico, extending westward in the northern itonasallength, 23.95; interorbital breadth, Chihuahuan Desert to near Willcox, Ari­ 5.10; length of maxillary toothrow, 3.20; zona, and southward to the vicinity ofCasas width across maxillary toothrows, 4.60; Grandes, Chihuahua (Williams, 1978b). length of mastoid bulla, 8.45; length of in­ Remarks. - Perognathus jlavescens ex­ terparietal, 2.50; width of interparietals, hibits extreme geographic variation in col­ 3.75; length ofnasals, 8.70; width ofnasals, or, size, bullar inflation, constriction ofthe 2.45; width of rostrum, 4.65. post-cranial region between the mastoid Distribution. - Found on loose sandy soils bullae, and relative size ofthe rostrum and in arid grassland and open woodland com­ nasal bones, all of which are strongly cor­ munities in northeastern Arizona, north and related with environmental temperature and east ofthe Mogollon rim, west ofthe Chuska moisture (Williams, 197 8b). This great Mountains, and east of the Coconino Pla­ variation makes it difficult to characterize teau, northward into southeastern Utah east and distinguish this species from other Pe­ ofthe Colorado River and south ofthe San rognathus. Williams (1978b) provided a Juan River (Williams, 1978b). systematic review of the intermountain Remarks. -Williams (1978b) treated populations, formerly designated as P. cleomophila as a synonym ofapache. Mer­ apache, and gave means and ranges ofmea­ riam (1889) listed the type locality as in surements for 17 samples; Williams and Apache Co. Genoways (1979) provided measurements for an additional sample from the northern Great Plains. Reed and Choate (1986) re­ Perognathus jlavescens caryi viewed geographic variation in the Great Goldman, 1918 Plains populations and gave statistics for 1918. Perognathus apache caryi Goldman, Proc. measurements of six samples, representing BioI. Soc. Washington, 31:24, 16 May. the four subspecies occupying the Great Plains. Hoffmeister (1986) remarked that Holotype. -Adult male, skin and skull, the evidence developed by Williams (I978b) USNM 148206, from 8 mi W Rifle, Garfield did not unequivocally prove that apache and Co., Colorado; obtained by Merritt Cary on flavescens were conspecific, and chose to 4 October 1906. treat apache as a distinct species. However, Measurements ofholotype. - Total length, no evidence reviewed suggests recognition 154; length of tail, 73; length of hind foot, oftwo species. 21; greatest length of skull, 25.15; occipi- 168 WILLIAMS ET AL.

tonasallength, 25.15; interorbital breadth, ored subspecies is easily confused with P. 5.60; length of maxillary toothrow, 3.50; jlavus on the basis ofexternal features. Ac­ width across maxillary toothrows, 4.65; cording to Reed and Choate (1986) cock­ length of mastoid bulla, 9.00; width across rumi only differs from samples ofjlavescens mastoid bullae, 13.40; length of interpari­ from Kansas in darker color. etal, 3.35; width of interparietals, 4.15; length ofnasals, 9.30; width ofnasals, 2.30; width of rostrum, 4.35. Perognathus jlavescens copei Distribution. - Typically found on sandy Rhoads, 1894 substrates in semiarid grassland and pin­ yon-juniper associations, from the Uintah 1894. Perognathus copei Rhoads, Proc. Acad. Basin of Utah and Colorado, southward Nat. Sci. Philadelphia, for 1893, 46:404, 27 through western Colorado and eastern Utah January. north of the San Juan River, thence south Holotype. -Adult male, skin and skull, and eastward in northwestern New Mexico ANSP 1612, from Staked Plains, near Mo­ and southwestern Colorado from the San beetie, Wheeler Co., Texas; obtained by E. Juan Basin to the Rio Grande Valley near D. Cope on 26 August 1893. Vale Verde, Socorro Co.; on the east to the Measurements ofholotype. - Total length, upper Pecos River Valley, San Miguel Co., 114; length of tail, 49; length of hind foot, New Mexico (Williams, 1978b). 15; greatest length ofskull, 19.5; interorbital Remarks. - Williams provided means and breadth, 4.5; width across mastoid bullae, ranges of measurements for samples of P. 10.0; length of nasals, 7.0; length of man­ jlavescens caryL dible, 9.7; height of coronoid process from angle, 4.2; ratio of length to breadth of in­ terparietal, 0.62. Perognathus jlavescens cockrumi Distribution. - Sandy desert and arid Hall, 1954 grassland associations of western Oklaho­ ma, western Texas, and eastern New Mex­ 1954. Perognathus jlavescens cockrumi Hall, Univ. Kansas Pub!., Mus. Nat. Hist., 7:589, ico. 15 November. Remarks. - The structural differences be­ tween P. f copei and P. f jlavescens are Holotype. -Adult female, skin and skull, relatively slight and the variation appears KU 13045, from 4.5 mi NE Danville, Har­ to be largely clinal, a situation that may not per Co., Kansas; obtained on 1 December warrant recognition of copei as a distinct 1939 by Sam Tihen. subspecies (Williams, 1978b; D. Williams, Measurements ofholotype. - Total length, unpubl. data). 114; length of tail, 51; length of hind foot, 17; condylobasal length, 18.5; interorbital breadth, 4.7; length of maxillary toothrow, 3.1; length of mastoid bulla, 6.8; width Perognathus jlavescens jlavescens across mastoid bullae, 10.5; width ofinter­ Merriam, 1889 parietals, 4.3. 1889. Perognathus fasciatus jlavescens Merri­ Distribution. - Known from sandy soils am, N. Amer. Fauna, 1:11,25 October. from central Kansas southward to west-cen­ tral Oklahoma (Hall, 1981). Holotype. -Adult male, skin and skull, Remarks. - Hall (1954) characterized the USNM 186507, from Kennedy, Cherry Co., holotype as subadult, but by the classifica­ Nebraska; obtained on 11 June 1888 by tion scheme of Williams (1978b), it would Vernon Bailey. be adult (age class 4). This small, dark-col- Measurements ofholotype. - Total length, TAXONOMY 169

136; length of tail, 73; length of hind foot, New Mexico, southward to the Samalayuc­ 17; length of ear (crown, dry), 4; greatest ca Sands and Casas Grandes, Chihuahua; length ofskull, 22.10; occipitonasa11ength, on the southwest, extending to Willcox Pla­ 22.10; interorbital breadth, 5.25; length of ya, Arizona, and on the southeast, at least maxillary toothrow, 3.00; width across to E1 Paso Co., Texas (Williams, 1978b). maxillary toothrows, 4.45; length of mas­ Remarks. - Populations of melanotis toid bulla, 7.25; width across mastoid bul­ from the White Sands ofthe Tularosa Basin lae, 11.85; length ofinterparietal, 3.10; width are whitish dorsally and ventrally; color, ofinterparietals, 5.20; length ofnasals, 7.90; however, is extremely variable geographi­ width ofnasals, 2.20; width ofrostrum, 3.75. cally, depending upon the color ofthe local Distribution. - Occupies sandy soils ofthe substrate and indirectly on the amount of west-central Great Plains, from southwest­ annual precipitation (Williams, 1978b). ern North Dakota through southwestern Kansas; on the west to extreme southeastern Wyoming and southward east ofthe Rocky Perognathus jlavescens perniger Mountain front at least to Pueblo Co., Col­ Osgood, 1904 orado. Remarks. -Armstrong (1972), Jones 1904. Perognathus jlavescens perniger Osgood, (1964), Reed and Choate (1986), Williams Proc. Biol. Soc. Washington, 17: 127, 9 June. (l978b), and Williams and Genoways (1979) Holotype. -Subadult female, skin and listed means and ranges of measurements skull, USNM 57725, from Vermillion, Clay for samples of P. f jlavescens. Co., South Dakota; obtained on 22 August 1889 by G. S. Agersborg. Measurements ofholotype. - Total length, Perognathus jlavescens melanotis 140; length of tail, 68; length of hind foot, Osgood, 1900 17 (dry); greatest length ofskull, 21.05; oc­ cipitonasal length, 21.05; interorbital 1900. Perognathus apache melanotis Osgood, breadth, 4.90; length ofmaxillary toothrow, N. Amer. Fauna, 18:27,20 September. 3.10; width across maxillary toothrows, 1929. Perognathus gypsi Dice, Occas. Papers 4.15; length of mastoid bulla, 6.40; width Mus. Zool., Univ. Michigan, 203:1, 19 June. across mastoid bullae, 10.70; length of in­ Holotype. -Adult female, skin and skull, terparietal, 2.55; width of interparietals, USNM 97416, from Casas Grandes, Chi­ 4.70; length ofnasals, 7.50; width ofnasals, huahua, Mexico; obtained on 21 May 1899 2.00; width of rostrum, 3.70. by Edward A. Goldman. Distribution. -Occupies sandy soils in Measurements ofholotype. - Total length, grassland associations in the northeastern 133; length of tail, 65; length of hind foot, sector ofthe species range, from southeast­ 19.5; greatest length of skull, 22.20; occip­ ern North Dakota and southern Minnesota itonasallength, 22.20; interorbital breadth, southward to northeastern Iowa, and west­ 5.10; length of maxillary toothrow, 2.85; ward and southward to central South Da­ width across maxillary toothrows, 4.20; kota and eastern Nebraska (Hall, 1981). length of mastoid bulla, 7.60; width across Remarks. - Typical specimens of perni­ mastoid bullae, 11.75; length of interpari­ ger are larger than typical specimens of lI:tal, 2.60; width of interparietals, 3.95; cockrumi, but both share the characteristic length ofnasals, 8.25; width ofnasals, 2.25; of dark color, which is associated with the width of rostrum, 3.65. high precipitation in the eastern sector of Distribution. - Sandy areas in desert and the geographic range of the species. Jones arid grassland associations from Gran Qui­ (1964) gave external and cranial measure­ vira and the San Augustine Plains ofcentral ments for a sample of perniger from Ne- 170 WILLIAMS ET AL. braska; Reed and Choate (1986) gave sta­ stricted, with short, narrow interparietals; tistics for measurements for a grouped length ofhead and body averages about 60 sample of 20. mm or less; length oftail averages less than 57 mm and rarely exceeds 60; interparietal length averages from about 2.3 to 2.5 mm Perognathus flavescens relictus and rarely exceeds 2.9; width of interpari­ Goldman, 1938 etals averages between about 3.0 and 3.3 mm and rarely exceeds 3.9. 1938. Perognathus apache relictus Goldman, J. Comparisons. -Po flavus is most similar Mamm., 19:495, 14 November. to P. merriami; they may occasionally hy­ H%type. -Adult male, skin and skull, bridize at a few localities. Generally, flavus USNM 150768, from Medano Springs differs from merriami in having: a shorter Ranch, 7,600 ft, 15 mi NE Mosca, Alamosa tail; longer, softer, and laxer pelage; darker, Co., Colorado; obtained on 2 November more contrasting mid-dorsal color with a 1907 by Merritt Cary. pinkish rather than yellowish or yellowish­ Measurements ofh%type. - Total length, orange hue; larger post-auricular spots; rel­ 137; length of tail, 68; length of hind foot, atively shorter tail; larger, more inflated 19.0; greatest length of skull, 22.70; occip­ auditory bullae; narrower interorbital and itonasallength, 22.70; interorbital breadth, interparietal widths; and wider P4. No sin­ 5.45; length of maxillary toothrow, 3.15; gle set of characters is known that will dis­ width across maxillary toothrows, 4.20; tinguish all P. flavus from all P. merriami. length of mastoid bulla, 8.00; width across Comparisons of individuals from areas of mastoid bullae, 12.40; length of interpari­ sympatry show P. f flavus and P. m. gilvus, etal, 3.10; width of interparietals, 3.75; respectively, to average: bullar lengthloc­ length ofnasals, 7.90; width ofnasals, 2.35; cipitonasallength, 0.388, 0.359; tail length/ width of rostrum, 3.60. total length, 0.452, 0.488; distance across Distribution. - In so far as is known, oc­ mastoid bullae, 11.93, 11.44 mm; width of cupies sandy substrates in the San Luis Val­ P4, 0.90, 0.82 mm. See accounts of P. fas­ ley, south-central Colorado (Williams, ciatus and P.flavescens for comparisons with 1978b). those sympatric species. P. flavus is also Remarks.-Williams (1987b) gave means sympatric with P. amp/us and P. /ongimem­ and ranges of measurements for 17 adults, bris; it differs from those species in having and Armstrong (1972) listed measurements an absolutely and relatively shorter, non­ for 6 adult males. penciled tail; from amp/us, P. flavus also differs in being significantly smaller (length of hind foot rarely exceeds 18 mm versus Perognathus flavus rarely less than 19); from /ongimembris, fla­ vus differs in smaller average size, smaller Diagnosis. - The smallest species of Pe­ hind feet, narrower interorbital width, and rognathus, with the shortest tail and small­ significantly wider upper molars. P. flavus est ratio oflengths oftail to head and body is significantly smaller and has a relatively (ratio averaging less than 0.9); dorsal color shorter, non-penciled tail than the allopatric is heavily overlain with blackish-tipped hairs species P. a/tico/a, P. inornatus, and P. par­ in most subspecies, contrasting sharply with vus. the clear buffY post-auricular patch and a Distribution. - Perognathus flavus occu­ narrow line without black tinge on the side pies a broad range in the west-central and adjacent to the white underparts; the pos­ southweatem Great Plains, intermountain terior region of the cranium has relatively basins, and central plateau ofMexico; a dis­ large auditory bullae and is relatively con- junct population is found along the coastal TAXONOMY 171 plain of the Gulf of California in Sonora. so fragmented that it precludes taking stan­ Its geographic range extends from eastern dard measurements. Hoffmeister (1986) Wyoming and western Nebraska, south­ gave statistics for measurements of 10 spec­ ward through western Texas and western imens; Osgood (1900) gave average external Coahuila on the Mexican Plateau to eastern measurements for 10 individuals and cra­ Jalisco, Morelos, Puebla, and tiny area of nial measurements for 3 individuals. east-central Veracruz; on the west, it ranges across New Mexico and southern Colorado to southeastern Utah and west-central Ar­ Perognathus jlavus bunkeri izona both north and south ofthe Mogollon Cockrum, 1951 Mountains (Hall, 1981). 1951. Perognathus flavus bunkeri Cockrum, Remarks. - Wilson (1973) treated P. Univ. Kansas Publ., Mus. Nat. Hist., 5:205, merriami merriami and P. m. gilvus as sub­ 15 December. species of P. jlavus. Anderson (1972) and Lee and Engstrom (1991) presented struc­ Holotype. -Adult female, skin and skull, tural and genic evidence that jlavus and KU 11716, from Conrad Farm, 1 mi E merriami are not conspecific. A few indi­ Coolidge, Hamilton Co., Kansas; obtained viduals of P. f jlavus and P. m. gilvus ap­ by F. Parks and Claude W. Hibbard on 1 parently hybridize at a few localities in July 1936. southeastern New Mexico, complicating Measurements ofholotype. - Total length, identification. Williams (1971, 1978a) 104; length of tail, 44; length of hind foot, placed P. jlavus with P. merriami in the jla­ 16; length of ear, 7; occipitonasal length, vus species group, separate from the fascia­ 20.6; frontonasal length, 14.0; interorbital tus group to whichjlavus and merriami had breadth, 4.5; length of maxillary toothrow, been assigned by Osgood (1900) and sub­ 3.3; length of mastoid bulla, 8.0; width sequent researchers. across mastoid bullae, 12.1; width ofinter­ parietals, 3.2. Distribution. -Occurs in the west-central Perognathus jlavus bimaculatus Great Plains from eastern Colorado though Merriam, 1889 western Kansas, and south to western Okla­ homa. 1889. Perognathusflavus bimaculatus Merriam, Remarks. -Armstrong (1972) listed N. Amer. Fauna, 1:12,25 October. measurements for seven individuals from Holotype. - Adult male, skin and skull, Colorado. USNM 8455/23789, from Fort Whipple, Yavapai Co., Arizona; obtained on 21 May 1865 by Elliott Coues. Perognathus jlavus jlavus Measurements ofholotype. - Total length, Baird, 1855 117; length of tail, 40; length of hind foot, 17; length ofear (crown, dry), 4. 1855. Perognatus [sic]flavus Baird, Proc. Acad. Nat. Sci. Philadelphia, 7:332. Distribution. -Occupies the plateau re­ gion ofwest-central Arizona, from the south Holotype. - Sex and age unknown, skin, rim of the Grand Canyon of the Colorado USNM 14811130, from El Paso, El Paso River in central Coconino Co., southward Co., Texas; obtained by 1. H. Clark in 1851 around the west rim of the Mogollon Pla­ (see remarks below). teau to near Prescott, Yavapai Co., and Measurements ofholotype. - Total length, westward to the Aubrey Valley in western 102.1; length of tail, 50.8; length of hind Coconino Co. (Hall, 1981). foot, 15.7. Remarks. - The skull of the holotype is Distribution. - Ranges from the extreme 172 WILLIAMS ET AL. northern Panhandle of Texas westward 18; length ofear (crown, dry), 4; nasa11ength, through most ofnorthern New Mexico, ex­ 7.70; interorbital breadth, 4.65. cluding the San Juan Basin, and southward Distribution. -Occupies arid grassland in central New Mexico through extreme and woodland associations in the volcanic western Texas and northern and eastern region around the San Francisco Mountains Chihuahua to north-central Durango; also and Mogollon Plateau near Flagstaff, north­ extends westward across southern New ward west of the Little Colorado River to Mexico to southeastern Arizona, south of near the south rim ofthe Grand Canyon of the Gila River and east of the Santa Cruz the Colorado River, all in Coconino Co., River, and probably into northeastern So­ Arizona. nora (Hall, 1981). Remarks. - The skull of the holotype is Remarks.-Some measurements given by broken, preventing the measuring ofseveral Baird (1858), in a more complete descrip­ traits. Hoffmeister (1986) gave statistics for tion of the species, differ from the original a sample of 20, and Osgood (1900) listed description. According to Merriam (1889) average measurements for 3 individuals. the holotype was lost; he designated USNM 4328/5047 (adult male) from Mason, Texas as a duplicate type, but noted that Mason Perognathus flavus fuscus was about 400 miles east of £1 Paso, and Anderson, 1972 remarked that "it would not be strange if 1972. Perognathusjlavusfuscus Anderson, Bull. future collections show the £1 Paso animal Amer. Mus. Nat. Hist., 148:304,8 September. to be different from the one here described." The duplicate type is from the easternmost Holotype. -Adult male, skin and skull, area ofintergradation between P. merriami KU 81168, from 2 mi W Minaca, 6,900 ft, gilvus and P. m. merriami, and clearly does Chihuahua; obtained by Charles A. Long not represent P. f flavus. Anderson (1972) on 2 July 1959. gave means and ranges ofmeasurements for Measurements ofholotype. - Total length, 9 specimens from Chihuahua, Hoffmeister 103; length of tail, 45; length of hind foot, (1986) listed statistics for measurements of 16; length of ear, 6; weight 7.4 g; greatest 14 specimens from Arizona; Williams length of skull, 20.40; width across bullae, (1971) gave statistics for measurements for 11.70; breadth across maxillary arches, 30 adults from Bernalillo Co., New Mexico; 10.30; nasal length, 7.60; interorbital and Wilson (1973) gave means ± SD for 37 breadth, 4.40. specimens. Distribution. - Known from the upper valley ofthe Rio Papigochic and the water­ shed ofthe Laguna Bustillos above 6,000 ft Perognathus flavus fuliginosus elevation, west-central Chihuahua, Mexico Merriam, 1890 (Anderson, 1972). Remarks. -Anderson (1972) gave means 1890. Perognathus fuliginosus Merriam, N. and ranges of measurements for five indi­ Amer. Fauna, 3:74, 11 September. viduals. Holotype. -Immature male, skin and skull, USNM 17708124644, from cedar Perognathus flavus goodpasteri belt, northeast ofSan Francisco Mountain, Hoffmeister, 1956 7,000 ft, Coconino Co., Arizona; obtained by Vernon Bailey on 4 October 1889. 1956. Perognathusjlavus goodpasteri Hoffmeis­ Measurements ofholotype. - Tota11ength, ter, Proc. BioI. Soc. Washington, 69:55, 21 116; length of tail, 58; length of hind foot, May. TAXONOMY 173

Holotype. - Adult male, skin and skull, east ofthe Little Colorado River and south UIMNH 6312, from 2.75 mi NW Springer­ and east of the Colorado River, but occurs ville, Apache Co., Arizona; obtained by west ofthe Little Colorado River near Wu­ Charles A. McLaughlin on 16 August 1953. patki National Monument (Hoffmeister, Measurements ofholotype. - Total length, 1986). 107; length of tail, 54; length of hind foot, Remarks. -Hoffmeister (1986) gave sta­ 17; length of ear (notch), 7; occipitonasal tistics for measurements for a sample of23 length, 20.5; basilar length, 14.4; interor­ individuals. bital breadth, 4.4; length ofmaxillary tooth­ row, 3.3; width across mastoid bullae, 12.2; length of interparietal, 3.1; width of inter­ Perognathus jlavus medius parietals, 3.1; length ofnasals, 7.2; width of Baker, 1954 nasals, 2.0. Distribution. -Occupies grassland asso­ 1954. Perognathus flavus medius Baker, Univ. ciations in east-central Arizona, ranging Kansas Publ., Mus. Nat. Hist., 7:343,15 Feb­ from near Springerville through Snowflake ruary. to south ofHolbrook on the northeast Mo­ Holotype. - Adult female, skin and skull, gollon Plateau, and also occurs on the south KU 48583, from 1 mi S, 6 mi E Rin60n de side of the Mogollon Plateau near Nash Romos, 6,550 ft, Aguascalientes, Mexico; Creek Reservoir, Navajo Co. (Hoffmeister, obtained on 14 July 1952 by Rollin H. Ba­ 1986). ker. Remarks. -Hoffmeister (1986) gave sta­ Measurements ofholotype. - Total length, tistics for a sample of 10 individuals. 111; length of tail, 52; length of hind foot, 17; occipitonasal length, 20.7; frontonasal length, 13.9; interorbital breadth, 4.5; length Perognathus jlavus hopiensis of maxillary toothrow, 3.0; length of mas­ Goldman, 1932 toid bulla, 7.8; width ofinterparietals, 2.9. Distribution. -Occurs on the plateau of 1932. Perognathus flavus hopiensis Goldman, north-central Mexico, from southeastern Proc. Biol. Soc. Washington, 45:89, 21 June. Durango and southeastern Coahuila south­ Holotype. - Adult female, skin and skull, ward through San Luis Potosi to Aguasca­ USNM 248014, from Oraibi, 6,000 ft, Hopi lientes, extreme northern and eastern Jalis­ Indian Reservation, Navajo Co., Arizona; co, and Guanajuato. obtained by George G. Cantwell on 5 June Remarks. -Baker (1954) listed statistics 1927. for measurements of 17 adults. Measurements ofholotype. - Total length, 115; length of tail, 50; length of hind foot, 15; occipitonasallength, 19.7; interorbital Perognathus jlavus mexicanus breadth, 4.4; length of maxillary toothrow, Merriam, 1894 2.9; width across mastoid bullae, 12.0; length ofinterparietal, 2.5; width ofinterparietals, 1894. Perognathus flavus mexicanus Merriam, 3.2; length of nasals, 7.0; width of nasals, Proc. Acad. Nat. Sci. Philadelphia, 46:265, 27 September. 2.2. Distribution. -Occurs from southeastern Holotype. - Young adult male, skin and Utah, mostly south of the San Juan River, skull, USNM 50714, from Tlalpam, Valley and extreme southwestern Colorado in of Mexico, Distrito Federal, Mexico; ob­ Montezuma Co., southward through north­ tained by Edward W. Nelson on 4 Decem­ eastern Arizona; in Arizona occurs mostly ber 1892. 174 WILLIAMS ET AL.

Measurements ofholotype. - Total length, Holotype. - Adult female, skin and skull, 118; length of tail, 55; length of hind foot, KU 38402, from 4 mi W, 2 mi S Guada­ 17.5; length ofear (from anterior base, dry), lajara, 5,100 ft, Jalisco, Mexico; obtained 6; greatest length ofskull, 21.00; width across by J. R. Alcorn on 15 June 1950. bullae, 11.90; breadth across maxillary Measurements ofholotype. - Total length, arches, 10.20; nasal length, 6.90; interor­ 113; length of tail, 52; length of hind foot, bital breadth, 4.60. 17; occipitonasal length, 20.9; frontonasal Distribution. -Occupies south-central length, 13.9; interorbital breadth, 4.6; length Mexico from Queretaro and Hidalgo south­ of maxillary toothrow, 3.2; length of mas­ ward to Morelos, Puebla, and west-central toid bulla, 7.9; width across mastoid bullae, Veracruz (Baker, 1954). 12.2; width ofinterparietals, 3.3. Remarks. -Baker (1954) gave means and Distribution. - Known from the plateau extremes of measurements for 27 speci­ region ofcentral and eastern Jalisco and ex­ mens. treme southern Zacatecas (Baker, 1954; Matson and Baker, 1986). Remarks. - Baker (1954) listed means and Perognathus j/avus pallescens ranges of measurements for 15 specimens. Baker, 1954

1954. Perognathusjlavus pallescens Baker, Univ. Perognathus j/avus piperi Kansas PubI., Mus. Nat. Hist., 7:345, 15 Feb­ Goldman, 1917 ruary. 1917. Perognathusjlavuspiperi Goldman, Proc. Holotype. - Adult male, skin and skull, BioI. Soc. Washington, 30:148, 27 July. KU 40298, from 1 mi SW San Pedro de las Colonias, 3,700 ft, Coahuila, Mexico, on 9 Holotype. - Adult male, skin and skull, February 1951 by J. R. Alcorn. USNM 168650, from 23 mi SW Newcastle, Measurements ofholotype. - Total length, Weston Co., Wyoming; obtained on 25 May 107; length of tail, 52; length of hind foot, 1910 by S. E. Piper. 16; greatest length ofskull, 20.5; frontonasal Measurements ofholotype. - Total length, length, 13.7; interorbital breadth, 4.5; length 113; length of tail, 51; length of hind foot, of maxillary toothrow, 3.0; length of mas­ 17; greatest length ofskull, 22.0; interorbital toid bulla, 8.0; width across mastoid bullae, breadth, 4.6; length of maxillary toothrow, 11.9; width ofinterparietals, 2.8. 3.5; width across mastoid bullae, 12.4; length Distribution. -Occupies portions of the ofinterparietal, 1.6; width ofinterparietals, Chihuahuan Desert in central and western 2.7; length of nasals, 8.1. Coahuila, southeastern Chihuahua, north­ Distribution. - Known from the northern eastern Durango, and extreme northern Za­ Great Plains in eastern Wyoming and west­ catecas (Hall, 1981; Matson and Baker, ern Nebraska (Hall, 1981). 1986). Remarks. -Jones (1964) gave measure­ Remarks. - Baker (1954) gave measure­ ments for four specimens from Nebraska. ments for four specimens from Coahuila.

Perognathus j/avus sanluisi Hill, 1952 Perognathus j/avus parviceps Baker, 1954 1942. Perognathus jlavus sanluisi Hill, Amer. Mus. Novitates, 1212:1,7 December. 1954. Perognathusjlavus parviceps Baker, Univ. Kansas PubI., Mus. Nat. Hist., 7:344, 15 Feb­ Holotype. - Adult female, skin and skull, ruary. AMNH 137669, from 9 mi E of Center, TAXONOMY 175

7,580 ft, Saguache Co. (20 mi NNW ofAla­ interorbital breadth relatively narrow com­ mosa, Alamosa Co.), Colorado; obtained by pared to other longimembris-group species; Peter E. Crowe on 18 August 1940. width of interparietal highly variable, gen­ Measurements ofholotype. - Total length, erally broader than other longimembris­ 109; length of tail, 46; length of hind foot, group species; dorsal profile of skull rela­ 15; length of ear (notch), 6; greatest length tively flat; length ofhead and body averages of skull, 20.0; basilar length, 14.1; interor­ from about 70 to 75 mm; ratio of lengths bital breadth, 4.5; length ofmaxillary tooth­ oftail to head andbody averages from about row, 3.1; width across mastoid bullae, 11.7; 1.02 to 1.07; interorbital breadth ranges length of interparietal, 2.6; width of inter­ from about 4.75 to 5.7 mm (means from parietals, 3.1; zygomatic breadth, 9.7; length about 4.9 to 5.1); width of interparietal of nasals, 7.5; length of rostrum, 8.2. ranges from about 3.3 to 4.7 mm, averaging Distribution. -Occurs only in the San Luis from about 3.6 to 3.9. Valley of south-central Colorado. Comparisons. -Perognathus inornatus is Remarks. -Armstrong (1972) gave sta­ most similar to P. amplus and P. longi­ tistics for measurements ofseven males and membris (see account of amplus for meth­ four females. ods ofdistinguishing that species). The sim­ ilarities between P. inornatus and longimembris are great, but generally P. in­ Perognathus jlavus sonoriensis ornatus averages slightly larger with a rel­ Nelson and Goldman, 1934 atively shorter and less penciled tail; the interorbital region ofinornatus is relatively 1934. Perognathus j/avus sonoriensis Nelson and narrower, but there is broad overlap in ab­ Goldman, J. Washington Acad. Sci., 24:267, solute measurements. P. inornatus is poten­ 15 June. tially sympatric with P. alticola, and pos­ sibly also with P. parvus; from these two Holotype. -Adult female, skin and skull, species, inornatus differs in having a less­ USNM 250885, from Costa Rica Ranch, penciled tail, smaller average size (hind foot lower Rio Sonora, Sonora, Mexico; ob­ generally less than 21 versus 21 mm ormore); tained on 13 December 1932 by Vernon relatively larger mastoid bullae and greater Bailey and Frederic Winthrop. mastoid breadth, and relatively narrower Measurements of holotype. -Length of interparietals. From the allopatric species, hind foot, 15; occipitonasallength, 19.3; in­ P. fasciatus, P. jlavescens, P. jlavus, and P. terorbital breadth, 4.2; length of maxillary merriami, inornatus differs in having a toothrow, 2.8; width across mastoid bullae, slightly penciled tail averaging longer than 11.7; length of interparietal, 3.0; width of length of head and body (the tails of the interparietals, 3.2; length of nasals, 6.5; other species lack pencils and average short­ width of nasals, 2.0; zygomatic breadth er in length than length of head and body). (posteriorly), 10.3. Distribution. -Occupies arid, annual Distribution. -Known only from the grassland, savanna, and desert shrub asso­ coastal plains ofcentral Sonora (Hall, 1981). ciations of west-central California, ranging Remarks. -No other published infor­ from the upper Sacramento Valley, Tehama mation is known. Co., southward through the San Joaquin and Salinas Valleys and contiguous areas to the Mojave Desert, Los Angeles, Kern, and ex­ Perognathus inornatus treme western San Bernardino counties; also occupies the Tehachapi mountains and the Diagnosis. -A medium-sized species of foothills ofthe western Sierra Nevada below Perognathus, with a slightly penciled tail av­ about 2,000 ft (Hall, 1981; D. Williams, eraging longer than length ofhead and body; unpubl. data). 176 WILLIAMS ET AL.

Remarks. - The literature ofP. inornatus quin Valley floor and the Tehachapi Moun­ shows great confusion about the taxonomy tains remains to be determined. of longimembris and inornatus (e.g., Os­ Remarks. - Pocket mice of this complex good, 1918), and the systematic relation­ from the eastern side of the San Joaquin ships among populations of these two spe­ Valley in Fresno, Madera, and Stanislaus cies. These relationships are not adequately and the floor ofthe valley in Fresno, Kings, clarified; at least two and possibly three spe­ and Kern counties have 50 chromosomes cies probably are currently included under and differ structurally from those of the the name P. inornatus. With one exception, western edge ofthe San Joaquin Valley and we retain the current taxonomy for longi­ areas to the west in Kern, San Luis Obispo, membris group pocket mice from central Fresno, San Benito, Merced, San Joaquin, California, although our unpublished infor­ and Alameda counties. Individuals of the mation suggests a different taxonomy will latter populations have 56 chromosomes be required when studies are completed. and are generally larger, with relatively lon­ ger tails. A single individual from Lake Co. had 60 chromosomes; no others from the Perognathus inornatus inornatus Sacramento Valley region have been stud­ Merriam, 1889 ied cytologically (Williams, 1978a; D. Wil­ liams, unpubl. data). Largely on the basis 1889. Perognathus inornatus Merriam, N. Amer. of the consistent chromosomal difference, Fauna, 1: 15, 25 October. populations in the foothills of the western Holotype. -Adult male, skin (in alcohol) edge of the San Joaquin Valley and nearby and skull, USNM 13394123790 from Fres­ mountains are assigned to P. inornatus ne­ no, Fresno Co., California; obtained by glectus rather than inornatus in deference to Gustav Eisen (no date; cataloged 6 February Hall (1981). 1882). Measurements of holotype. - [Externals taken by Merriam from alcoholic specimen Perognathus inornatus neglectus before skinning out skull, except ear from Taylor, 1912 notch] Total length, 137; length of tail, 71; length of hind foot, 18.5; length of ear 1912. Perognathus longimembris neglectus Tay­ (crown), 4, notch 6.6; greatest length ofskull, lor, Univ. California Publ. Zool., 10:155,21 May. 22.80; occipitonasallength, 22.35; interor­ bital breadth, 4.60; length of maxillary Holotype. - Adult male, skin and skull, toothrow, 3.10; width across maxillary MVZ 14526, from McKittrick, 1,111 ft, toothrows, 4.05; length of mastoid bulla, Kern Co., California; obtained on 18 May 8.70; width across mastoid bullae, 12.35; 1911 by H. S. Swarth and W. L. Chandler. length ofinterparietal, 2.80; width ofinter­ Measurements ofholotype. - Total length, parietals, 3.55; length ofnasals, 8.20; width 157; length of tail, 77; length of hind foot, of nasals, 2.05; width ofrostrum, 3.40. 22; length ofear, 6; greatest length ofskull, Distribution. - Ranges through the Sac­ 25.75; interorbital breadth, 5.75; length of ramento Valley, from Tehama Co. on the mastoid bulla, 8.85; width across mastoid north to the coastal mountains in eastern bullae, 13.85; length of interparietal, 2.80; Lake Co. on the west, southward along the width of interparietals, 3.65; length of na­ eastern side and floor of the San Joaquin sals, 9.60; width of rostrum, 3.85. Valley to near its southern end. The specific Distribution. -Occurs in the hills and status ofpocket mice ofthe inornatus com­ piedmont slopes along the western edge of plex from the western side of the San Joa- the San Joaquin Valley from near Suisun TAXONOMY 177

Bay, Contra Costa Co., southward to the ley, from near Soledad, southward at least Mojave Desert, western San Bernardino and to Hog Canyon, Monterey Co. northeastern Los Angeles counties; and Remarks. - The only differences we can westward in the Panoche Valley, San Benito find between the type specimens and as­ Co., the Carrizo Plain, San Luis Obispo Co., signed specimens ofP. longimembris psam­ and the upper Cuyama Valley in Santa Bar­ mophilus and P. inornatus sillimani of von bara and San Luis Obispo counties. Bloeker (1937) are due to age. Nearly all of Remarks. -See remarks above under P. von Bloeker's (1937) specimens of psam­ inornatus inornatus and below under P. in­ mophilus are juveniles and subadults, and ornatus psammophilus. We consider indi­ nearly all of his specimens of sillimani are viduals from the Carrizo Plain to be indis­ adults; the type series of the latter are old tinguishable from those near the type locality adults (age class 5 ofWilliams, 1978b). On on the western edge ofthe San Joaquin Val­ the basis of page precedence, sillimani is a ley. This arrangement differs from that of junior synonym ofpsammophilus. P. inor­ von Bloeker (1937). The taxonomic assign­ natus psammophilus is most similar to P. ment ofpopulations from the extreme west­ inornatus neglectus; we have assigned spec­ ern floor ofthe San Joaquin Valley and the imens from the Carrizo Plain that von Salinas River Valley is undecided. Speci­ Bloeker (1937) treated as sillimani and mens from the Mojave Desert are assigned psammophilus to P. inornatus neglectus, and to neglectus on the basis ofsimilar structure. consider it likely that psammophilus will Taylor (1912) gave means ofmeasurements prove to be a synonym of neglectus. for an unknown number of individuals of each sex. Perognathus longimembris

Perognathus inornatus psammophilus Diagnosis. -A small species of Perogna­ von Bloeker, 1937 thus with a relatively long, slightly penciled tail, medium-sized interparietals, and nar­ 1937. Perognathus iongimembris psammophi­ row upper premolar. Ratio oflength oftail ius von Bloeker, Proc. BioI. Soc. Washington, to head and body nearly always exceeds 1.03, 50:153,10 September. ranging to about 1.40; length of hind foot 1937. Perognathus inornatus sillimani von ranges from about 15 to 20 mm (means Bloeker, Proc. BioI. Soc. Washington, 50: 154, between about 18 and 19); occipitonasal 10 September. length rarely exceeds 22.0 mm (population Holotype. -Subadult female, skin and means always less than 22.0); width of up­ skull, MVZ 74681, from west side ofArroyo per premolar ranges from about 0.75 to 0.95 Seco Wash, 150 ft, 4 mi S Soledad, Mon­ mm. terey Co., California; obtained on 5 June Comparisons. - Perognathus longimem­ 1936 by Jack C. von Bloeker, Jr. bris is most similar to P. amplus and P. Measurements ofholotype. - Total length, inornatus (see accounts of those species for 130; length of tail, 62; length of hind foot, comparisons). P. longimembris is similar in 18; length ofear, 7; greatest length ofskull, size to P. flavescens, but differs in having a 21.10; interorbital breadth, 4.85; length of tail longer than length ofhead and body and mastoid bulla, 7.35; width across mastoid with a slight pencil; from P. alticola and P. bullae, 11.55; length of interparietal, 2.55; parvus, P. longimembris differs in being sig­ width of interparietals, 4.50; length of na­ nificantly smaller in nearly all dimensions; sals, 7.50; width of rostrum, 3.50. length of hind foot rarely exceeds 20 mm Distribution. -Occupies the Salinas Val- in longimembris whereas the hind feet of 178 WILLIAMS ET AL. alticola and parvus usually exceed 20 mm. lat. 31 0 52' N, long. 116 0 06' W, Baja Cal­ From P. merriami, P. longimembris differs ifornia, Mexico; obtained on 10 June 1927 in larger size with a relatively and absolutely by Laurence M. Huey. longer tail; length of tail in longimembris Measurements ofholotype. - Total length, typically exceeds 66 mm whereas in mer­ 135; length of tail, 71; length of hind foot, riami, the tail rarely measures as long as 66 17; length of ear, 5; weight, 8.7 g; greatest mm. See account ofP.jlavus for comparison length of skull, 22.0; interorbital breadth, with that species. 5.4; length ofmaxillary toothrow, 2.8; width Distribution. -Occupies desert, shrub­ across mastoid bullae, 13.2; length ofnasals, steppe, and open, arid woodland commu­ 8.0. nities of the Great Basin, southward from Distribution. - Known from the western southeastern Oregon, extreme southwestern base ofthe Sierra Juarez, from near the type Idaho, and western Utah; the Mojave Des­ locality on the north, south to El Valle de ert and Tehachapi Mountains, southern la Trinidad (Hall, 1981). California coastal basins, Colorado Desert, Remarks. - Like other populations of P. and adjacent areas along the Colorado River longimembris from the southwestern part of in south-central Utah, Arizona, and Cali­ its range, the mastoid bullae are greatly in­ fornia; also extends into northern Baja Cal­ flated; whether it is subspecifically distinct ifornia and around the northern perimeter from adjacent subspecies is equivocal, how­ ofthe GulfofCalifornia in Sonora; and oc­ ever. Huey (1939a) listed measurements for curs as apparently disjunct populations in five males and three females. Pinal and Maricopa counties, south-central Arizona, and around Bahia Kino in west­ Perognathus longimembris arizonensis central Sonora (Hall, 1981). Goldman, 1931 Remarks. - Distribution records for P. longimembris from west-central California 1931. Perognathus longimembris arizonensis in the Sacramento Valley, most or all ofthe Goldman, Proc. Biol. Soc. Washington, 44: 134, San Joaquin Valley, Salinas Valley, and on 17 October. the Carrizo Plain are based on misidentified 1935. Perognathus longimembris arcus Benson, specimens of P. inornatus. The northern Univ. California Publ. Zool., 40:451,31 De­ distributional limit of P. longimembris in cember. central California has not been determined, 1939. Perognathus longimembris virginis Ruey, but appears to coincide approximately with Trans. San Diego Soc. Nat. Rist., 9:55,31 Au­ the Transverse ranges, from which popu­ gust. lations are known. The identity of small Holotype. - Adult female, skin and skull, pocket mice on the northern slopes of the USNM 250032, from 10 mi S Jacobs Pools, Transverse ranges in the San Joaquin Valley Houserock Valley, north side of Marble is uncertain (unpubl. data). Canyon of Colorado River, 4,000 ft, Co­ conino Co., Arizona; obtained on 17 June 1931 by Edward A. Goldman. Perognathus longimembris aestivus Measurements ofholotype. - Total length, Huey, 1928 137; length of tail, 79; length of hind foot, 18.5; greatest length of skull, 22.05; occip­ 1928. Perognathus longimembris aestivus Ruey, itonasallength, 21.35; interorbital breadth, Trans. San Diego Soc. Nat. Rist., 5:87, 18 Jan­ 4.95; length of maxillary toothrow, 3.00; uary. width across maxillary toothrows, 4.05; Holotype. -Adult male, skin and skull, length of mastoid bulla, 8.50; width across SDSNH 6110, from Sangre de Cristo, Valle mastoid bullae, 12.15; length ofinterpari­ San Rafael on western base ofSierra Juarez, etal, 3.15; width of interparietals, 3.10; TAXONOMY 179 length ofnasals, 8.00; width ofnasals, 2.10; Springs, Riverside Co., southward to near width ofrostrum, 3.35; zygomatic breadth, the Mexican boundary in western Imperial 10.9. Co. (unpubl. data). Distribution. -Occurs in extreme south­ Remarks. -Huey (1939a) listed mea­ western and south-central Utah and north­ surements for five males and five females. western and north-central Arizona generally north ofthe Colorado River, and in south­ eastern Nevada (Hall, 1981; Hoffmeister, Perognathus longimembris bombycinus 1986). Known from south (east) ofthe Col­ Osgood, 1907 orado River only from an area near Rain­ bow Bridge, San Juan Co., Utah, and the 1907. Perognathus bombycinus Osgood, Proc. Bioi. Soc. Washington, 20:19, 23 February. vicinity of Page, Coconino Co., Arizona (Hall, 1981; Hoffmeister, 1986). H olotype. - Adult male, skin and skull, Remarks. - Hoffmeister (1986) consid­ USNM 136123, from Yuma, Yuma Co., ered virginis not to be subspecifically dis­ Arizona; obtained on 18 March 1905 by tinct from arizonensis, and he found the two Edward A. Goldman. specimens known of arcus to provide little Measurements ofholotype. - Total length, justification for recognition as a subspecies. 134; length of tail, 79; length of hind foot, Goldman (1931) listed mean external mea­ 18.5; greatest length of skull, 21.55; occip­ surements for four topotypes. Hall (1946) itonasallength, 20.40; basilar length, 14.5; gave means and ranges ofmeasurements for interorbital breadth, 4.50; length of maxil­ 10 individuals of each sex from Nevada; lary toothrow, 2.95; width across maxillary Hoffmeister (1986) gave statistics for 201 toothrows, 3.40; length of mastoid bulla, specimens in four samples from Arizona. 8.60; width across mastoid bullae, 11.7; length of interparietal, 2.5; width of inter­ parietals, 2.6; length ofnasals, 7.3; width of Perognathus longimembris bangsi nasals, 1.80; width of rostrum, 2.90; zygo­ Mearns, 1898 matic width, 9.8; diastema, 5.1. Distribution. -Found along the lower 1898. Perognathus longimembris bangsi Mearns, Colorado River Valley in southeastern Cal­ Bull. Amer. Mus. Nat. Hist., 10:300, 31 Au­ ifornia and southwestern Arizona from near gust. Parker Dam on the north, and around the 1900. Perognathus panamintinus arenicola Ste­ northern end of the Gulf of California in phens, Proc. Bioi. Soc. Washington, 13:151, northwestern Sonora and northeastern Baja 13 June. California. Isolated populations extend Holotype. - Adult female, skin and skull, eastward in southwestern Arizona to near AMNH 5304, from Palm Springs, Colorado Phoenix and Casa Grande. In Baja Califor­ Desert, Riverside Co., California; obtained nia, extends to near San Felipe, and in So­ on 13 April 1896 by E. C. Thurber. nora, to near the Pinacate Lava Flows (Hall, Measurements ofholotype. - Total length, 1981). 138; length of tail, 80; length of hind foot, Remarks.-Huey (1939a) listed mea­ 19; greatest length ofskull, 20.6; interorbital surements ofthree males and four females. breadth, 4.8; length of maxillary toothrow, 3.4; width across mastoid bullae, 11.4; length Perognathus longimembris brevinasus of nasals, 7.7; length of rostrum, 9.5; zy­ Osgood, 1900 gomatic breadth, 9.8. Distribution. - Occurs in the Colorado 1900. Perognathus panamintinus brevinasus Desert region west of the Imperial Valley, Osgood, N. Amer. Fauna, 18:30,20 Septem­ in southern California, from north ofPalm ber. 180 WILLIAMS ET AL.

Holotype. -Adult female, skin and skull, tocene Lake Bonneville in eastern Nevada USNM 186515, from San Bernardino, San and western Utah, from near Kelton, Box Bernardino Co., California; obtained on 2 Elder Co., Utah, southward to near Pruss May 1885 by Frank Stephens. Lake, Millard Co., Utah (Hall, 1981). Measurements ofholotype. - Total length, Remarks. -Durrant (1952) gave mea­ 124; length of tail, 66; length of hind foot surements for six males and three females (dry), 17.4; greatest length of skull, 20.65; from Utah; Hall (1946) listed measure­ occipitonasal length, 20.15; interorbital ments for five females and three males from breadth, 5.05; length ofmaxillary toothrow, Nevada. 3.00; width across maxillary toothrows, 3.85; length of mastoid bulla, 8.00; width across mastoid bullae, 11.80; length of in­ Perognathus longimembris internationalis terparietal, 3.10; width of interparietals, Huey, 1939 4.20; length ofnasals, 7.20; width ofnasals, 1.90; width of rostrum, 3.35. 1939. Perognathus longimembris internation­ Distribution. -Occupies the arid coastal alis Huey, Trans. San Diego Soc. Nat. Hist., 9:47,31 August. basins of southern California in grassland and coastal sage associations, from approx­ Holotype. -Adult male, skin and skull, imately Burbank and San Fernando, Los SDSNH 11972, from Baja California side Angeles, Co., on the northwest, to San Ber­ of the International Boundary at Jacumba, nardino, San Bernardino Co., and Cabazon, San Diego Co., California; obtained by Hemet, and Aguanga, Riverside Co., on the Laurence M. Huey on 24 April 1936. east; the southern limit is near the northern Measurements ofholotype. - Total length, boundary ofSan Diego Co. (Huey, 1939a). 141; length of tail, 78; length of hind foot, Remarks. -Osgood (1900) gave average 19; length ofear (crown), 5; greatest length measurements for three topotypes; Huey of skull, 21.9; interorbital breadth, 5.2; (1939a) gave measurements offive individ­ length of maxillary toothrow, 3.1; width uals of each sex. across mastoid bullae, 12.6; length ofnasals, 7.5. Distribution. - Known from La Puerta Perognathus longimembris gulosus and San Felipe valleys of San Diego Co., Hall, 1941 California and adjacent area in Baja Cali­ fornia (Hall, 1981). 1941. Perognathus longimembris gulosus Hall, Remarks.-Huey (1939a) gave measure­ Proc. BioI. Soc. Washington, 54:55, 20 May. ments for five adults ofeach sex. This sub­ Holotype. - Adult female, skin and skull, species seems to represent a form structur­ MVZ 78764, from near [0.25 mi S] Smith ally intermediate between the coastal basin Creek Cave, 5,800 ft, Mount Moriah, White subspecies, brevinasus andpacificus, and the Pine Co., Nevada; obtained by Lee W. Ar­ inland desert forms, bangsi and aestivus, a nold on 4 June 1937. situation to be expected on the basis of its Measurements ofholotype. - Total length, intermediate geographic position. Subspe­ 132; length of tail, 72; length of hind foot, cific recognition is equivocal. 17; length of ear, 7; weight, 8.9 g; occipi­ tonasallength, 21.1; frontonasallength, 14.2; Perognathus longimembris kinoensis interorbital breadth, 5.3; length ofmaxillary Huey, 1935 toothrow, 3.0; length of mastoid bulla, 8.2; width across mastoid bullae, 12.5. 1935. Perognathus longimembris kinoensis Distribution. -Occurs in desert associa­ Huey, Trans. San Diego Soc. Nat. Hist., 8:73, tions along the western margin ofthe Pleis- 24 August. TAXONOMY 181

Holotype. -Adult male, skin and skull, occipitonasal length, 20.30; interorbital SDSNH 11300, from Bahia Kino (northern breadth, 4.90; length ofmaxillary toothrow, end ofthe sand dune peninsula that borders 3.00; width across maxillary toothrows, the bay and forms the northern arm of the 3.90; length of mastoid bulla, 7.50; width estuary), Sonora, Mexico; obtained by across mastoid bullae, 11.45; length of in­ Laurence M. Huey on 26 February 1935. terparietal, 2.85; width of interparietals, Measurements ofholotype. - Total length, 4.35; length ofnasals, 7.10; width ofnasals, 135; length of tail, 80; length of hind foot, 1.75; width of rostrum, 3.65. 17; length ofear (crown), 4; greatest length Distribution. -Occurs in the Mojave of skull, 20.7; interorbital breadth, 4.6; Desert region and Transverse ranges of length of maxillary toothrow, 2.6; width southeastern California, from near Inde­ across mastoid bullae, 11.4; length ofnasals, pendence in the Owens Valley on the north­ 7.2. east, to near Mount Pinos on the southwest, Distribution. -Known only from the vi­ and the Providence Mountains on the cinity ofthe type locality on the GulfCoast southeast. ofcentral Sonora, Mexico. Remarks. -Specimens from the north of Remarks. -Like other Perognathus liv­ the Transverse ranges in west-central Cali­ ing in relatively humid environments, the fornia, assigned to this species (Hall, 1981), bullae are less inflated and the posterior cra­ are P. inornatus. The northern distribution­ nial region less constricted than populations al limits of P. longimembris longimembris from arid environments. This subspecies are not known. The type locality is in a po­ appears to be little differentiated from bom­ sition where either or both P. longimembris bycinus from farther north. and P. inornatus neglectus may occur, but no pocket mice have been taken from there in this century. The holotype may not have Perognathus longimembris longimembris come from Ft. Tejon, but rather from some­ (Coues, 1875) where in the surrounding country, perhaps the Mojave Desert side of the Transverse 1875. o[tognosis]. longimembris Coues, Proc. ranges (Osgood, 1918). Bole (1937) gave Acad. Nat. Sci. Philadelphia, 27:305, 31 Au­ gust. measurements of 17 specimens. 1904. Perognathus elibatus Elliot, Field Colum­ bian Mus., Publ. 87, Zool. Ser., 3:252, 7 Jan­ uary. Perognathus longimembris nevadensis 1904. Perognathus pericalles Elliot, Field Co­ Merriam, 1894 lumbian Mus., Publ. 87, Zool. Ser., 3:252, 7 January. 1894. Perognathus nevadensis Merriam, Proc. Acad. Nat. Sci. Philadelphia, 46:264, 27 Sep­ Holotype. -Adult female, skin (in alco­ tember. hol) and skull, USNM 9856/37356, from Fort Tejon, Tehachapi Mountains, Kern Holotype. - Adult male, skin and skull, Co., California; obtained by John Xantus USNM 54828, from Halleck, East Hum­ sometime in 1857 or 1858. boldt Valley, Elko Co., Nevada; obtained Measurements ofholotype. - [Total length by Vernon Bailey on 4 July 1893. determined from length of head and body Measurements ofholotype. - Total length, given by Coues (1875) plus length of tail 127; length of tail, 72; length of hind foot, taken from specimen in alcohol; other ex­ 19; length of ear (anterior base, dry), 7; ternal measurements made from specimen greatest length ofskull, 21.40; occipitonasal in alcohol.] Total length, 121; length oftail, length, 20.80; interorbital breadth, 5.05; 63; length of hind foot, 16.1; length of ear length of maxillary toothrow, 2.90; width (notch), 5.7; greatest length ofskull, 20.55; across maxillary toothrows, 4.15; length of 182 WILLIAMS ET AL. mastoid bulla, 7.90; width across mastoid Perognathus longimembris panamintinus bullae, 12.05; length of interparietal, 2.25; Merriam, 1894 width of interparietals, 4.10; length of na­ sals, 8.05; width of nasals, 1.75; width of 1894. Perognathus longimembris panamintinus rostrum, 3.20. Merriam, Proc. Acad. Nat. Sci. Philadelphia, Distribution. -Occupies Great Basin des­ 46:265, 27 September. ert associations from the southeastern cor­ Holotype. - Adult male, skin and skull, ner of Oregon, southwestward to the Sur­ USNM 27767/39866, from Perognathus prise Valley, Modoc Co., California, and Flat, 5,200 ft, Panamint Mountains, Inyo southward in northern and Central Nevada; Co., California; obtained by Vernon Bailey in Nevada, south to Smiths Creek Valley on 16 April 1891. and eastward to Halleck and near Eureka Measurements ofholotype. - Total length, (Hall, 1946, 1981). 152; length of tail, 83; length of hind foot, Remarks. - Hall (1946) gave statistics for 20; greatest length of skull, 22.80; occipi­ measurements of 10 males and 5 females. tonasal length, 22.10; interorbital breadth, 5.45; length of maxillary toothrow, 3.10; width across maxillary toothrows, 4.25; Perognathus longimembris pacificus length of mastoid bulla, 8.15; width across Mearns, 1898 mastoid bullae, 12.35; length of interpari­ etal, 3.00; width of interparietals, 4.05; 1898. Perognathus pacificus Mearns, Bull. Amer. length ofnasals, 8.55; width ofnasals, 2.00; Mus. Nat. Rist., 10:299, 31 August. width of rostrum, 3.55. 1932. Perognathus longimembris cantwelli von Distribution. - Occupies Great Basin Bloeker, Proc. Biol. Soc. Washington, 45:128, Desert associations in western Nevada and 9 September. southeastern California, ranging from Quinn River Crossing, Humboldt Co., Nevada Holotype. - Adult female, skin and skull, south into Clark Co., Nevada and westward USNM 61022, from the shore ofthe Pacific in Inyo Co., California to the Panamint Ocean at Mexican boundary monument No. Mountains (Hall, 1981). 258, San Diego Co., California; obtained on Remarks. - Hall (1946) gave statistics for 12 July 1894 by Edgar A. Mearns. measurements of 10 males and 7 females Measurements ofholotype. - Total length, from Nevada. Bole (1937) listed measure­ 113; length of tail, 53; length of hind foot, ments of 20 specimens from California. 15.5; length ofear, 5; greatest length ofskull, 19.20; occipitonasallength, 19.00; interor­ bital breadth, 4.65; length of maxillary toothrow, 2.85; width across maxillary Perognathus longimembris pimensis toothrows, 3.75; length of mastoid bulla, Huey, 1937 7.15; width across mastoid bullae, 10.85; length ofinterparietal, 2.30; width ofinter­ 1937. Perognathus longimembris pimensis Ruey, parietals, 3.70; length ofnasals, 7.05; width Trans. San Diego Soc. Nat. Rist., 8:355, 15 June. of nasals, 1.85; width of rostrum, 3.10. Distribution. - Occurs on the coastal Holotype. - Adult male, skin and skull, plains of southern California, from near El SDSNH 12579, from 11 mi W Casa Gran­ Segundo, Los Angeles Co., southward to de, Pinal Co., Arizona; obtained by Lau­ near Tiajuana, Baja California, Mexico. rence M. Huey on 22 May 1937. Remarks. -Huey (l939a) listed mea­ Measurements ofholotype. - Total length, surements for 10 males and 6 females. 144; length of tail, 83; length of hind foot, TAXONOMY 183

18; length of ear (crown), 4; greatest length Holotype. -Adult male, skin and skull, of skull, 21.2; interorbital breadth, 5.0; MVZ 74668, from 1 mi W Kennedy Mead­ length of maxillary toothrow, 2.7; width ows, 6,000 ft, S Fork Kern River, Tulare across mastoid bullae, 12.2; length ofnasals, Co., California; obtained on 14 August 1936 7.4. by William B. Richardson. Distribution. -South-central Arizona, Measurements ofholotype.-Total length, from near Marinette, Maricopa Co., on the 127; length of tail, 68; length of hind foot, north to near Casa Grande, Pinal Co., on 19; length of ear (notch), 7; occipitonasal the southeast and Gila Bend, Pinal Co., on length, 21.0; frontonasal length, 14.6; in­ the southwest. terorbital breadth, 5.2; length of mastoid Remarks. - This population is apparently bulla, 8.2; width across mastoid bullae, 11.7; isolated from others of P. longimembris. distance between stylomastoid foramina, 10.5. Distribution. -Known only from the vi­ Perognathus longimembris salinensis cinity ofthe type locality in the upper valley Bole, 1937 of the Kern River in eastern Tulare Co., California (Hall, 1981). 1937. Perognathus longimembris salinensis Bole, Remarks. -Richardson (1937) gave Sci. PubIs., Cleveland Mus. Nat. Hist., 5(2):3, means and extremes for measurements of 4 December. five subadult and adult specimens. Aside from slightly darker color, this subspecies Holotype. -Adult male, skin and skull, differs from P. longimembris longimembris UMMZ 121257, from 1 mi N Salt Camp, only in negligibly smaller size ofthe mastoid 1,060 ft, west edge salt lake, Saline Valley, bullae and related structure, differences that Inyo Co., California; obtained by P. N. may reflect the immaturity of most of the Moulthrop on 29 March 1934. type series. Measurements ofholotype. - Total length, 130; length of tail, 74; length of hind foot, 17.5; occipitonasal length, 19.5; basilar length, 13.9; interorbital breadth, 4.9; length Perognathus longimembris venustus of maxillary toothrow, 3.1; width across Huey, 1930 mastoid bullae, 11.6; length ofinterparietal, 2.0; width of interparietals, 3.6; length of 1930. Perognathus longimembris venustus Huey, nasals, 7.5; width of rostrum, 2.1. Trans. San Diego Soc. Nat. Hist., 6:233, 24 Distribution. - Known only from the Sa­ December. line Valley, Inyo Co., California. Holotype. -Adult female, skin and skull, Remarks. - The holotype was originally SDSNH 8196, from San Agustin, lat. 30oN, CMNH 6242 (Cleveland Museum of Nat­ long. l15°W, Baja California, Mexico; ob­ ural History). Bole (1937) listed measure­ tained on 4 October 1930 by Laurence M. ments for 20 specimens. Huey. Measurements ofholotype. - Total length, 130; length of tail, 78; length of hind foot, Perognathus longimembris tularensis 19; length of ear (crown), 5; greatest length Richardson, 1937 of skull, 21.8; interorbital breadth, 5.0; length of maxillary toothrow, 3.3; width 1937. Perognathus longimembris tularensis across mastoid bullae, 12.3; length ofnasals, Richardson, J. Mamm., 18:510, 22 Novem­ 7.9. ber. Distribution. - Known only from the type 184 WILLIAMS ET AL. locality in north-central Baja California or small set of characters that will reliably (Hall, 1981). distinguish all P. m. gilvus from all P. f Remarks.-Huey (1939a) gave measure­ jlavus. ments for three specimens. We see no sig­ nificant differences in the measurements listed by Huey (1939a) for venustus and aes­ Perognathus merriami gilvus tivus, leaving only the "much darker color" Osgood, 1900 of the former as apparently diagnostic, a situation that may not warrant taxonomic 1900. Perognathus merriami gilvus Osgood, N. separation. Amer. Fauna, 18:22,20 September. Holotype. -Adult male, skin and skull, USNM 35939/48273, from Eddy, Eddy Co., Perognathus merriami New Mexico; obtained by B. H. Dutcher on 18 September 1892. Diagnosis. -A small pocket mouse, with Measurements ofholotype. - Total length, length of head and body seldom exceeding 118; length of tail, 58; length of hind foot, 62 mm, and with a relatively short tail; ratio 16.5; greatest length of skull, 21.00; width oflength oftail to head and body averaging across bullae, 11.75; breadth across maxil­ 0.82-0.95; dorsal color yellowish or yellow­ lary arches, 10.00; nasal length, 7.25; inter­ ish-orange with a slight blackish tinge and orbital breadth, 4.65. little contrast between the mid-dorsal and Distribution. -Occurs in western Texas dorso-lateral areas; buffy post-auricular spot and southeastern New Mexico, southward relatively small; auditory bullae moderately through eastern Chihuahua into extreme inflated; rostrum proportionately broad and northern and western Coahuila, Mexico short. (Hall, 1981). Comparisons. -See account of P. jlavus Remarks. - Anderson (1972) listed ways for comparison with that species. Size ofdistinguishing P. m. gilvus and P. f jlavus smaller, tail shorter, and interparietals not and included a ratio diagram for this pur­ as wide as P.jlavescens. P. merriami is sym­ pose, although he had measurements for patric only with the preceding two species only two P. m. g. from Chihuahua. Wilson of Perognathus. See accounts of the other (1973) gave statistics for measurements of species for additional comparisons. 42 specimens from throughout the geo­ Remarks. - P. merriami was considered graphic range, and Baker (1956) gave mea­ a species distinct from P. jlavus until Wilson surements for 4 individuals from western (1973), based on morphology, treated them Coahuila. as conspecific. The distribution ofP. m. gil­ vus is geographically intermediate to the main range ofP. jlavus and P. m. merriami, Perognathus merriami merriami and is in many ways structurally interme­ J. A. Allen, 1892 diate to P. m. merriami and P. f jlavus, giving the impression that the two taxa 1892. Perognathus merriami J. A. Allen, Bull. broadly hybridize. However, genic analysis Amer. Mus. Nat. Rist., 4:45, 25 March. shows distinct genotypes for P. m. gilvus and 1896. Perognathus mearnsi J. A. Allen, Bull. Amer. Mus. Nat. Rist., 8:237, 21 November. P. f jlavus from several localities of sym­ patry, and only a few probable hybrids from Holotype. - Adult male, skin and skull, one locality in southeastern New Mexico AMNH 4145/3177, from Brownsville, (Lee and Engstrom, 1991). On this basis, Cameron Co., Texas; obtained by F. B. Lee and Engstrom (1991) recommended the Armstrong on 10 August 1891. two nominate forms be recognized as dis­ Measurements ofholotype. - Total length, tinct species. There is no known character 124; length of tail, 58; length of hind foot, TAXONOMY 185

18; greatest length ofskull, 20.3; interorbital into southwestern Montana, southeastern breadth, 4.6; length of maxillary toothrow, Wyoming, and northeastern Utah; also 2.9; width across mastoid bullae, 11.2; length ranges through all but the southern tip of of nasals, 7.2; length of rostrum, 7.7. Nevada and eastward through the western Distribution. - Ranges from central Texas two-thirds of Utah, and southward into near the Oklahoma boundary, southward to northwestern Arizona north of the Grand eastern Coahuila, north and central Nuevo Canyon ofthe Colorado River (Hall, 1981). Leon, and central Tamaulipas. Remarks. - Perognathus parvus and the Remarks.-Wilson (1973) gave means closely related P. alticola comprise the par­ and ranges of measurements for 57 speci­ vus species group; herein we treat Perog­ mens from throughout the range ofthe sub­ nathus xanthonotus as a subspecies of P. species; Baker (1956) listed measurements parvus. The subspecies and species of the for 10 adults from eastern Coahuila. parvus group are poorly known; a system­ atic review of the group is called for.

Perognathus parvus Perognathus parvus bullatus Diagnosis. - The largest species ofPerog­ Durrant and Lee, 1956 nathus, with a slightly crested and tufted tail 1956. Perognathus parvus bullatus Durrant and (pencil) that averages longer than length of Lee, Proc. BioI. Soc. Washington, 69:183,31 head and body; distal, dorsal one-third of December. tail with a mix of buffy, brownish, and blackish hairs; antitragus ofear pinna lobed; Holotype. -Adult male, skin and skull, hairs ofinner surface ofpinna not white or UU 8771, from Ekker's Ranch, Robbers yellowish; interparietal relatively short; Roost, 25 mi (airline) E ofHanksville, 6,000 mastoid breadth relatively narrow and pos­ ft, Wayne Co., Utah; obtained by John terior cranial region relatively unconstricted Bushman on 18 May 1951. by the mastoid bullae; mastoid bulla forms Measurements ofholotype. - Total length, slight indentation in lateral border of ex­ 160; length of tail, 85; length of hind foot, occipital on back ofskull; interorbital region 22; length ofear, 8; greatest length ofskull, broad; baculum relative to head and body 25.40; width across bullae, 14.00; breadth length long for species group, but about av­ across maxillary arches, 11.80; length ofna­ erage for genus (Burt, 1960; Osgood, 1900; sals, 9.15; interorbital breadth, 6.10. Su1entich, 1983). Distribution. - Known from southeastern Comparisons. -Perognathus parvus is Utah in an area bounded by the San Rafael most similar to P. alticola; see account of (Huntington Creek tributary), Green, Col­ the latter species for comparisons. P. parvus orado, and Fremont rivers, in Emery and can be distinguished from all other species Wayne counties (Hall, 1981). of Perognathus by its larger size, lobed an­ Remarks. -Durrant and Lee (1956) gave titragus, and more prominent crest on the statistics for measurements of 19 speci­ distal portion of the tail. mens. Distribution. -Occupies steppe and open, arid shrub and woodland communities in the Columbia River Basin, Great Basin, and Perognathus parvus clarus contiguous areas in western North America; Goldman, 1917 ranges from sagebrush communities in the Okanogan Valley of south-central British 1917. Perognathus parvus clarns Goldman, Proc. BioI. Soc. Washington, 30:147, 27 July. Columbia, southward through eastern Washington, Oregon, and California; east­ Holotype. -Adult male, skin and skull, ward across the Snake River Plains ofIdaho USNM 178939, from Cumberland, Lincoln 186 WILLIAMS ET AL.

Co., Wyoming; obtained by Stanley G. Jew­ length of mastoid bulla, 9.40; width across ett on 18 May 1912. mastoid bullae, 14.45; length of interpari­ Measurements ofholotype. - Total length, etal, 3.30; width of interparietals, 4.80; 181; length of tail, 84; length of hind foot, length ofnasals, 11.25; width ofnasals, 2.60; 22; greatest length of skull, 25.35; occipi­ width of rostrum, 4.30. tonasallength, 25.25; interorbital breadth, Distribution. -Occurs on the Columbia 5.75; length of maxillary toothrow, 3.70; Plateau of central Washington, generally width across maxillary toothrows, 4.95; north and east of the Columbia River and length of mastoid bulla, 8.35; width across north and west of the Snake River on the mastoid bullae, 13.15; length of interpari­ south, and south ofthe Columbia River on etal, 3.45; width of interparieta1s, 5.70; the north; recorded from west of the Co­ length ofnasals, 10.00; width ofnasals, 2.45; lumbia River only from near Wenatchee, width of rostrum, 4.30. Chelan Co., Washington (Hall, 1981). Distribution. -Occupies the northeastern Remarks. - Merriam (18 94b) indicated sector ofthe species' range, in southeastern that the holotype was a young adult; its cheek Idaho, southwestern Montana, southwest­ teeth show considerable wear (age class 5 of ern Wyoming, and northeastern Utah (Hall, Williams, 1978a), indicating that it is old, 1981). not young. Osgood (1900) gave average ex­ Remarks. - Williams (1978a) considered ternal measurements for 5 adult topotypes, specimens from the Uintah Basin ofnorth­ Broadbooks (1954) listed statistics for 52 eastern Utah to belong to this subspecies specimens by age and sex, and Dalquest rather than P. p. olivaceous; in either case (1948) listed means for external measure­ there is little to distinguish the two subspe­ ments of 21 males and 9 females. cies except color, which varies considerably over the range of the two subspecies. Long (1965) listed means and ranges ofmeasure­ Perognathus parvus idahoensis ments for eight males from the type locality Goldman, 1922 and one male and one female from 21 mi W of Rock Springs, Wyoming. 1922. Perognathus parvus idahoensis Goldman, Proc. BioI. Soc. Washington, 35:105, 17 Oc­ tober. Perognathus parvus columbianus Holotype. -Adult male, skin and skull, Merriam, 1894 USNM 236394, from Echo Crater, 20 mi SW Arco, Blaine [Butte] Co., Idaho; ob­ 1894. Perognathus columbianus Merriam, Proc. tained by Luther 1. Goldman on 14 June Acad. Nat. Sci. Philadelphia, 46:26], 27 Sep­ 1921. tember. Measurements ofholotype. - Total length, Holotype. -Adult male, skin and skull, 189; length oftail, 102; length ofhind foot, USNM 27351139450, from Pasco, Plains of 24; greatest length of skull, 27.40; occipi­ the Columbia, east side ofColumbia River tonasal length, 27.40; interorbital breadth, near mouth of Snake River, Franklin Co., 5.90; length of maxillary toothrow, 3.85; Washington; obtained by Clark P. Streator width across maxillary toothrows, 5.15; on 9 May 1891. length of mastoid bulla, 9.00; width across Measurements ofholotype. - Total length, mastoid bullae, 14.25; length of interpari­ 187; length of tail, 96; length of hind foot, etal, 3.60; width of interparietals, 5,50; 23; greatest length of skull, 27.60; occipi­ length ofnasals, 10.55; width ofnasals, 2.75; tonasal length, 27.60; interorbital breadth, width of rostrum, 4.35. 6.00; length of maxillary toothrow, 4.10; Distribution. -Occurs on lava soils in width across maxillary toothrows, 5.15; south-central Idaho, on a portion of the TAXONOMY 187

Snake River Plain and slopes of volcanos skull, 25.5; width across bullae, 13.9; nasal in the vicinity of Craters of the Moon Na­ length, 9.4; interorbital breadth, 6.0. tional Monument (Hall, 1981). Distribution. - Ranges from near the in­ Remarks. - The type locality, 20 mi SW ternational boundary in south-central Brit­ Arco, is in Butte Co., rather than Blaine Co. ish Columbia, southward in north-central and eastern Washington to west-central Ida­ ho in Nez Perce Co., and south eastern Perognathus parvus laingi Washington in Asotin Co.; generally ranges Anderson, 1932 north of the Columbia River in north-cen­ tral Washington and east of the Columbia 1932. Perognathus laingi Anderson, Bull. Natl. Basin in eastern Washington (Hall, 1981). Mus. Canada, 70:100,24 November. Remarks. - The holotype was originally mounted; external measurements provided Holotype. -Adult male, skin and skull, by Merriam (1889) were estimated from the NMC 9200, from Anarchist Mountain, near dry skin. The occipital and maxillary regions Osoyoos-Bridesville summit, about 8 mi E of the skull of the holotype are broken. ofOsoyoos Lake, about 3,500 ft, lat. 49 0 08' 0 Anderson (1932) listed statistics for mea­ N, long. 119 32' W, British Columbia; ob­ surements of 24 males and 12 females. tained by Hamilton Laing on 29 August Broadbooks (1954) listed statistics for mea­ 1928. surements by age and sex for 29 specimens Measurements ofholotype. - Total length, from British Columbia and Washington, and 191; length oftail, 102; length ofhind foot, Dalquest (1948) gave means for external 25; occipitonasallength, 27.0; basal length measurements of 29 males and 10 females. of Hensel, 20.0; interorbital breadth, 5.0; Rhoads (1894) supplemented the extremely width across mastoid bullae, 14.5; length of scanty description of Gray, and provided interparietal, 4.0; width ofinterparietals, 5.0; measurements for a "duplicate type." length ofnasals, 10.0; length ofincisive fo­ ramina, 2.0. Distribution. -Occupies the upper Oka­ Perognathus parvus mollipilosus nogan Basin of south-central British Co­ Coues, 1875 lumbia, from Ashcroft on the northwest to near Osoyoos on the southeast (Hall, 1981). 1875. P[erognathus]. mollipilosus Coues, Proc. Remarks. -Anderson (1932) gave mea­ Acad. Nat. Sci. Philadelphia, 27:296, 31 Au­ surements for three specimens. gust. Holotype.-Female, USNM 7251, from Fort Crook [about 2 mi NE ofBurgettville], Perognathus parvus lordi Shasta Co., California; obtained by J. Feil­ (Gray, 1868) nero Measurements of holotype. -[Measure­ 1868. Abromys lordiGray, Proc. Zool. Soc. Lon­ ments from specimen in alcohol, before don, p. 202, May. skinning] Total length, 145; length of tail, Holotype. -Age and sex unknown, 81.3; length ofhind foot, 20.3; length ofear, BM(NH) 62.12.12.19, obtained from Brit­ 7.6 (see remarks). ish Columbia by J. K. Lord. Distribution. -Occupies the Modoc Pla­ Measurements ofholotype. -[External teau and adjacent areas ofnortheastern Cal­ measurements and greatest length of skull ifornia, and south-central Oregon in the from Merriam, 1889] Total length, 155; Klamath Basin; ranges from Crater Lake length oftail, 81; length ofhind foot, 21.8; National Park, Oregon on the north, to length ofear (crown), 6.5; greatest length of Edgewood, Siskiyou Co., California on the 188 WILLIAMS ET AL. west, and Susanville, Lassen Co., California cept, according to Durrant (1952) and Hall on the southeast (Hall, 1981). (1981) for populations in Emery, Duchesne, Remarks. -Coues (1875) listed the cata­ and Uintah counties, east of the Wasatch log number for the holotype but did not Range. indicate the museum. Merriam (1889) re­ Remarks. - Durrant (1952) gave statistics marked that the specimen should be in the for measurements of 5 males and 4 females U.S. National Museum but it could not be from central Utah, and Hall (1946) provid­ found and the number was wrong. Robert ed measurements for 10 individuals ofeach Fisher (in litt.) noted that USNM 7251 was sex from Nevada. Williams (1978a) as­ a Sealopus aquatieus that also was missing. signed specimens from the Uintah Basin, Osgood (1900) gave average measurements Duchesne Co., Utah, to P. p. clarus. for three topotypes. Perognathus parvus parvus Perognathus parvus olivaeeus (Peale, 1848) Merriam, 1889 1848. Cricetodipus parvus Peale, Mammalia and ornithology, in U.S. Expl. Exped.... , 8:53. 1889. Perognathus olivaceus Merriam, N. Amer. 1858. Perognathus monticola Baird, Mammals, Fauna, 1: 15, 25 October. in Repts. Expl. Surv.... , 8(1):422, 14 July. 1889. Perognathus olivaceus amoenus Merriam, N. Amer. Fauna, 1:16,25 October. Holotype. -Juvenilefemale [age assumed 1900. Perognathus parvus magruderensis Os­ from the description], Exploration Expe­ good, N. Amer. Fauna, 18:38, 20 September. ditions Collections, USNM [no information 1939. Perognathus parvus plerus Goldman, J. recorded concerning holotype and assumed Mamm., 20:352, 14 August. to be lost or destroyed-originally pre­ Holotype. -Adult male, skin and skull, served in alcohol]; assumed to be from The USNM 186511, from Kelton, near N end Dalles, Wasco Co., Oregon. Great Salt Lake, Box Elder Co., Utah; ob­ Measurements ofholotype. - Total length, tained by Vernon Bailey on 24 October 106; length of tail, 58; length of hind foot, 1888. 20.5. Measurements ofholotype. - Total length, Distribution. - Ranges from southeastern 184; length oftail, 101; length ofhind foot, Washington, south ofthe Snake River, east 23; greatest length of skull, 26.70; occipi­ ofthe Columbia River, and west ofthe Blue tonasal length, 26.70; interorbital breadth, Mountains, through eastern Oregon to the 6.10; length of maxillary toothrow, 3.70; extreme northwestern comerofNevada, and width across maxillary toothrows, 4.55; in southwestern Idaho along the lowlands length of mastoid bulla, 8.90; width across ofthe Snake and Salmon rivers; in Oregon, mastoid bullae, 13.30; length of interpari­ ranges from the eastern edge ofthe Cascade etal, 2.95; width of interparietals, 5.15; Mountains southward east ofthe Winter and length ofnasals, 10.65; width ofnasals, 2.75; Warner ranges (Hall, 1981). width of rostrum, 4.40. Remarks. - The original description ob­ Distribution. - Occupies the Great Basin, viously was ofa very young animal. Its ju­ its range extending from extreme south-cen­ venile characteristics led to considerable tral Idaho southward through all but the taxonomic confusion during the 1800's be­ extreme northwestern comer and extreme tween Perognathus and Dipodomys. Rhoads southern portion of Nevada; into east-cen­ (1894) and Osgood (1900) were primarily tral California from near Lake Tahoe, responsible for determining the identity of southward to Inyo Co.; through western Crieetodipus parvus of Peale. Hall (1946) Utah west ofthe central mountain axis, ex- listed measurements of 4 specimens from TAXONOMY 189

Nevada, Anderson (1932) listed means and fornia; obtained by H. A. Carr on 27 June ranges of measurements for 6 specimens 1911. from Oregon, and Da1quest (1948) gave Measurements ofholotype. - Tota11ength, means ofexternal measurements of3l males 170; length of tail, 85; length of hind foot, and 9 females from Washington. 22.5; greatest length of skull, 25.40; inter­ orbital breadth, 6.00; length ofmastoid bul­ la, 8.10; width across mastoid bullae, 12.95; Perognathus parvus trumbullensis length ofnasals, 9.10; width ofrostrum, 4.55. Benson, 1937 Distribution. -Occupies the eastern des­ ert slope ofthe northern Tehachapi Moun­ 1937. Perognathus parvus trumbullensis Ben­ tains, Kern Co., California; known range son, Proc. BioI. Soc. Washington, 50:181-182, extends from Indian Wells Canyon on the 28 October. north to Horse Canyon on the south (un­ Holotype. - Young adult male, skin and pub!. data). skull, MVZ 60929, from Nixon Spring, Remarks. - We concur with Sulentich 6,250 ft, Mount Trumbull, Mohave Co., (1983) that available evidence does not sup­ Arizona; obtained on 26 May 1933 by An­ port recognizing xanthonotus as a species nie M. Alexander. separate from P. parvus. The ranges of P. Measurements ofholotype. - Tota11ength, parvus olivaceous and P. parvus xanthonotus 174; length of tail, 90; length of hind foot, are probably continuous along the eastern 23; length ofear, 9; greatest length ofskull, front ofthe Sierra Nevada. Sulentich (1983) 26.85; interorbital breadth, 5.85; length of listed statistics for measurements of 20 mastoid bulla, 9.25; width across mastoid specimens. bullae, 13.90; length of interparietal, 3.45; width of interparieta1s, 4.80; length of na­ Perognathus parvus yakimensis sals, 10.45; width of rostrum, 4.00. Broadbooks, 1954 Distribution. -Occupies the high plateau country of northwestern Arizona, north of 1954. Perognathus parvus yakimensis Broad­ the Colorado River in Mohave and Coconi­ books, J. Mamm., 35:96, 10 February. no counties, and adjacent areas of south­ Holotype. - Young adult male, skin, skull, central and southwestern Utah in Washing­ and baculum, UMMZ 95978, from Rocky ton, Kane, and Garfield counties (Hall, Flat (or Rocky Prairie), 16 mi NW Naches, 1981 ). 3,800 ft, Yakima Co., Washington; ob­ Remarks.-Benson (1937) gave means tained by Harold E. Broadbooks on 27 Au­ and ranges of measurements for 20 adult gust 1947. males; Hoffmeister (1986) listed statistics Measurements ofholotype. - Tota11ength, for measurements of a sample of 18 indi­ 174; length of tail, 91; length of hind foot, viduals from Arizona. 24; length ofear, 8; weight, 17.2 g; greatest length of skull, 25.9; width across bullae, 13.1; breadth across maxillary arches, 11.9; Perognathus parvus xanthonotus nasal length, 11.2; interorbital breadth, 5.8. Grinnell, 1912 Distribution. -Occurs in sagebrush and open, arid ponderosa pine forests in south­ 1912. Perognathus xanthonotus Grinnell, Proc. central Washington, west of the Columbia BioI. Soc. Washington, 25:128, 31 July. River and east of the Cascade Mountains; Holotype. - Adult male, skin and skull, ranges from the Kittitas Valley, Kittitas Co., MVZ 16154, from Freeman Canyon, 4,900 on the north, southward to the Columbia ft, east slope Walker Pass, Kern Co., Ca1i- river, thence westward to the vicinity of 190 WILLIAMS ET AL.

Dallesport, Klickitat Co. (Broadbooks, notes by the collector. Bulletin of the Museum of Comparative Zoology, 8:183-189. 1954). --. 1891. Notes on new and little-known North Remarks. -Broadbooks listed statistics American mammals, based on recent additions to for measurements of 49 specimens by age the collection ofmammals in the American Museum of Natural History. Bulletin of the American Mu­ and sex. Hooper (1977) indicated that the seum of Natural History, 3:263-310. type locality was 17 mi NW Naches. --. 1899. NewrodentsfromColombiaandVen­ ezuela. Bulletin ofthe American Museum ofNatural History, 12:195-218. Acknowledgments --. 1908. Mammals from Nicaragua. Bulletin of the American Museum ofNatural History, 24:647­ 670. Troy L. Best, Kerry S. Kilburn and Duke S. --. 1916. List ofmammals collected in Colom­ Rogers reviewed portions ofthe manuscript and bia by the American Museum of Natural History provided helpful suggestions for its improve­ Expeditions, 1910-1915. Bulletin of the American ment. Troy also provided information on his "in Museum of Natural History, 35:191-238. press" manuscripts that allowed us to keep abreast ALLEN, J. A., AND F. M. CHAPMAN. 1893. On a col­ lection of mammals from the Island of Trinidad, ofsome taxonomic changes. Susan D. Williams with descriptions of new species. Bulletin of the provided invaluable editorial assistance by American Museum of Natural History, 5:203-234. proofreading and verifying bibliographic and no­ ALSTON, E. R. 1876. On the classification ofthe order menclatural data and measurements of type Glires. Proceedings ofthe Zoological Society ofLon­ don, 1876:61-98. specimens. Donald F. Hoffmeister kept us ap­ --. 1879-1882. Biologia Centrali-Americana. praised of the progress of his book, "Mammals Mammalia. Published for the Editors by R. H. Por­ of Arizona," in which he proposed several tax­ ter, London, 220 pp. onomic changes. The following curators and col­ ALVAREZ, T. 1963. The Recent mammals ofTamau­ lection managers provided information on, or lipas, Mexico. University of Kansas Publications, Museum of Natural History, 14:363-473. took measurements of, listed holotypes: Sydney ANDERSON, R. M. 1932. Five new mammals from Anderson, American Museum of Natural His­ British Columbia. National Museum ofCanada Bul­ tory; Ted Daeschler, The Academy of Natural letin, 70:99-119. Sciences, Philadelphia-Dipodomys h. heerman­ ANDERSON, S. 1964. The systematic status ofPerog­ nathus artus and Perognathus go/dmani (Rodentia). ni; Robert D. Fisher, National Museum ofNat­ American Museum Novitates, 2184:1-27. ural History- D. heermanni tularensis, D. o. or­ --. 1972. Mammals ofChihuahua taxonomy and dii (neotype), D. o. utahensis, Perognathusjlavus distribution. Bulletin of the American Museum of mexicanus; Mark S. Hafner, Louisiana State Natural History, 148:149-410. University, Museum of Zoology-Po [Chaeto­ ARATA, A. A. 1964. The anatomy and taxonomic significance ofthe male accessory reproductive glands dipus] penicillatus atrodorsalis, P. [C.] lineatus; ofmuroid rodents. Bulletin ofthe Florida State Mu­ Thor Holmes, University ofKansas, Museum of seum, 9:1-42. Natural History- D. ordii pullus, P. jlavusfuscus; ARMSTRONG, D. M. 1972. Distribution ofmammals Paula Jenkins, British Museum (Natural Histo­ in Colorado. Monograph, University ofKansas Pub­ lications, Museum of Natural History, 3:1-415. ry)-Abromys [Perognathus] lordi; Philip Myers, BAIRD, S. F. 1858. Mammals, in Explorations and University of Michigan, Museum of Zoology­ surveys, to ascertain the most practical and econom­ P. parvus yakimensis; Eric A. Richard, Univer­ ical route for a railroad from the Mississippi River sity ofUtah, Utah Museum ofNatural History­ to the Pacific Ocean. United States Department of D. o. celeripes, D. o. sanrafaeli, P. parvus bul­ the Interior, Washington, D.C., 8(1):1-757. BAKER, R. H. 1954. The silky pocket mouse (Perog­ latus; Robert M. Timm, University of Kansas, nathus flavus) ofMexico. University ofKansas Pub­ Museum ofNatural History. We extend our ap­ lications, Museum of Natural History, 7:339-347. preciation to these persons and the curators and --. 1956. Mammals of Coahuila, Mexico. Uni­ curatorial staffs who allowed us access to their versity ofKansas Publications, Museum of Natural History, 9: 125-335. holotype collections and provided so generously BANKS, R. C. 1964. The mammals ofCerralvo Island, of their time and hospitality. Baja California. Transactions of the San Diego So­ ciety of Natural History, 13:397-404. --. 1967. A new insular subspecies ofspiny pock­ Literature Cited et mouse (Mammalia; Rodentia). Proceedings ofthe Biological Society of Washington, 80:101-104. ALLEN, J. A. 1881. List ofmammals collected by Dr. BAUMGARDNER, G. D. 1989. Refinement ofthe type Edward Palmer in northeastern Mexico, with field- locality and distribution of Dipodomys compactus TAXONOMY 191

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HAFNER, J. C. 1978. Evolutionary relationships of mals in the Museum of Zoology, the University of kangaroo mice, genus Microdipodops. Journal of Michigan. Occasional Papers ofthe Museum ofZo­ Mammalogy, 59:354-366. ology, University of Michigan, 680:1-21. --. 1985. New kangaroo mice, genus Microdi­ HOOPER, E. T., AND C. O. HANDLEY, JR. 1948. Char­ podops (Rodentia: Heteromyidae), from Idaho and acter gradients in the spiny pocket mouse, Liomys Nevada. Proceedings of the Biological Society of irroratus. Occasional Papers of the Museum of Zo­ Washington, 98: 1-9. ology, University of Michigan, 514:1-34. HAFNER, 1. c., AND M. S. HAFNER. 1983. Evolution­ HUEY, L. M. 1926. The description of a new sub­ ary relationships ofheteromyid rodents. Great Basin species of Perognathus from Lower California with Naturalist Memoirs, 7:3-29. a short discussion ofthe taxonomic position ofother HAFNER, M. S. 1982. A biochemical investigation of peninsular members of this genus. 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