We reinvestigated the phylogeny of all 15 species of Marmota to resolve a conflict between 2 published analyses, one by Kruckenhauser et al. and another by Steppan et al., regarding the Olympic marmot (M. olympus) and to improve resolution in the genus. We acquired fresh samples of M. olympus, combined all available data on mitochondrial DNA (cytochrome b [Cytb] and ND3/ND4), new sequences for ND3/ND4, and 2,000 base pairs (bp) of the nuclear RAG1 gene. All analyses indicate that M. olympus is a much older, rather than more recent, offshoot of the widespread hoary marmot (M. caligata) or Vancouver Island marmot (M. vancouverensis). The mitochondrial data and some RAG1 results are largely congruent, but RAG1 differs on several points, including: the subgenus Marmota appears paraphyletic to Petromarmota, with reciprocally monophyletic Palearctic and Nearctic clades; and the long-tailed marmot (M. caudata) and Menzbier's marmot (M. menzbieri) are not sister species, suggesting mitochondrial introgression. Asia was colonized by Marmota from North America at approximately 4.6 million years ago (mya), followed by rapid diversification of several major lineages. M. olympus diverged from the M. caligata-M. vancouverensis lineage at approximately 2.6 mya, whereas M. vancouverensis and M. caligata diverged at only about 0.4-1.2 mya. M. olympus might have survived in isolation on the Olympic Peninsula in a nunatak refugium throughout a series of glacial maxima.

Cadmium, copper, iron, and zinc levels were measured in the kidneys of 115 grey wolves ( Canis lupus ) from Idaho, Montana and Alaska (United States), and from the Northwest Territories (Canada). No significant differences in the levels of iron or copper were observed between locations, but wolf kidneys from more northern locations had significantly higher cadmium levels (Alaska > Northwest Territories > Montana ≈ Idaho), and wolves from Alaska showed significantly higher zinc than other locations. Additionally, female wolves in Alaska had higher iron levels than males, and adult wolves in Montana had higher copper levels than subadults.

Three species of chipmunks, Tamias minimus, Tamias quadrivittatus and Tamias umbrinus, were studied on the eastern slope of the Front Range in Boulder and Larimer counties, Colorado. Tamias quadrivittatus and T. umbrinus are similar in size and appearance and are parapatric in distribution. The line of parapatry was found to occur at an elevation of 2,130 m in the study area, which is a revision of the 2,440 m previously reported in the literature. Tamias minimus differs in appearance and ecology from its two congeners and occurs sympatrically with them over a broad range of elevations and habitats. Overwinter survival rates for the populations are less than one third, and local population extinction or near-extinction is a common occurrence. Univariate and multivariate morphometric analyses are presented, as are electrophoretic omparisons of the three species. The three may be clearly diagnosed by a number of characters; no evidence of hybridization exists, although Front Range populations of T. umbrinus converge toward T. quadrivittatus in electrophoretic profile, as compared to other populations of the two species that are not in geographic contact. Analysis of sonagrams of the common vocalizations of the three species also shows that each has unique features. Possible mechanisms for the maintenance of parapatry are discussed. Throughout much of the mountainous region of western North America, one or more species of chipmunks (Tamias) can be found inhabiting most coniferous forest, woodland, shrub, and alpine tundra habitats. As many as 23 species and 69 subspecies of western chipmunks (subgenus Neotamias) are currently proposed. Levenson et al. (1985) reviewed the systematics of the genus, which also includes Tamias sibiricus (subgenus Eutamias), found across much of northern Asia, and Tamias striatus (subgenus Tamias), which is limited to deciduous forests of eastern North America. Considerable external phenotypic and ecological variability is displayed between related species and conspecific populations of Neotamias inhabiting different areas and in contact with different assemblages of congeners (see Patterson, 1981; Sullivan, 1985). The divergence of Neotamias and Tamias (see Levenson et al., 1985) about 10 to 13 million years ago in the late Miocene (Ellis and Maxson, 1979) resulted from the drying trend caused by mountain-building and the consequent rainshadow effect that occurred at various times across western North America between the early Eocene and late Pliocene (Axlerod, 1979). By the end of this period, the Great Plains was firmly established as an effective barrier between the two lineages. Phyletic divergence within Neotamias may have begun in the Pliocene with the isolation of two lineages on either side of the Great Basin (see Levenson et al., 1985). Again, during various glacial periods in the Pleistocene, the Great Basin was continuously forested or wooded (see Wells, 1983); speciation may have taken place during interglacials when mountain ranges became "forest islands." Present distributional patterns of mammalian species in the Great Basin suggest differential extinction in the relatively warm, dry post-glacial period (Brown, 1971; Grayson, 1987). Permanent barriers to dispersal between forest islands have existed in the West since the late Pleistocene (10,000 YBP; see Patterson, 1982), which may have contributed to the most recent HWESTERN URALIST 36(1):14-28 Bergstrom and Hoffmann-Tamias in the Front Range of Colorado pulse of speciation and intraspecific differentiation of isolated populations of Tamias. On the eastern slope of the Front Range of north-central Colorado, three species of Tamias occur. The smallest, Tamias minimus operarius (average weight of 44 g), is widespread, occurring in most habitats from lower montane open woodland-scrub to alpine tundra. Tamias umbrinus montanus (average weight of 59 g) occurs in all habitats except alpine tundra and the lowest band of montane woodland, about 500 m in elevational width. Tamias quadrivittatus (average weight of 62 g) is parapatric with T. umbrinus, generally occurring only in low-elevation habitats; however, each species occupies a much greater elevational range in regions where the other is absent (Armstrong, 1972). Tamias minimus is the most widely distributed of all North American Neotamias species (Hall, 1981) and is a member of the "boreo-cordilleran faunal element" of Armstrong (1972). Tamias m. operarius is the most widespread and most euryecious chipmunk in Colorado (Hall, 1981). Tamias quadrivittatus is a Coloradan autochthon with "Chihuahuan-like" distribution that probably evolved on the Colorado Plateau; it reaches its northernmost distribution in a narrow peninsular projection along the eastern slope of the Front Range in Larimer Co., Colorado (Armstrong, 1972). Other Chihuahuan species, including Peromyscus difficilis, Neotoma mexicana, and Spermophilus variegatus, share nearly identical distributions in this area, inhabiting only rocky, open woodland at foothills elevations. Tamias umbrinus is a Great Basin species with several major disjunct populations (Hall, 1981), the easternmost of which is Tamias u. montanus. The latter is parapatric with T. q. quadrivittatus, abutting its congener from the west and north in Colorado. Tamius u. montanus was first recognized as distinct from T. q. quadrivittatus by White (1953a, 1953b), based mainly on the morphology of the baculum. Parapatric distributions are the rule among western chipmunks (Findley, 1969; Hoffmann, 1974), with sympatry often occurring only in narrow overlap zones (e.g., Brown, 1971; Heller, 1971; Sheppard, 1972). When two or more species' ranges approach each other, there is some evidence of ecological, as well as morphological, shifts. Often morphological shifts involve convergence (Long and Cronkite, 1970; Armstrong, 1972; Patterson, 1981) and represent evolutionary responses to the particular assemblage of congeners present (Patterson, 1981). Morphological, biochemical, and ecological variability can be equally pronounced among populations of widespread species such as T. minimus (Sullivan, 1985). Tamias m. operarius in the Front Range is widely sympatric with both of its larger congeners, and it is not elevationally restricted as are other populations of T. minimus in proximity to different assemblages of Tamias (Heller, 1971; Sheppard, 1972; Patterson, 1981). This study documents the precise distributions of these three species of Tamias along a portion of the eastern slope of the Front Range. It also compares morphological, biochemical, and ecological attributes, including habitat distribution, relative abundance, and overwinter survival, of the populations in these areas. MATERIALS AND METHODS-Sixty-eight sites located along several elevational transects ranging from 1,700 m to 2,900 m were chosen for live trapping and habitat analysis (Fig. 1). Most of these sites were located within the Roosevelt National Forest in Larimer and Boulder counties, Colorado. They represented ecosystems ranging from grassland-lower montane ecotone to upper montane-subalpine ecotone (Marr, 1961). Chipmunks were observed but not trapped at 11 additional sites, representing mostly subalpine-alpine ecotones up to 3,200 m. Live trapping was conducted at each site for at least 2 weeks, with traps checked twice daily during the summers of 1981 and 1982. Additional trapping and observational records were obtained in the fall of 1980 and the summers of 1983 and 1984. Sherman live traps (3.5 by 3.5 by 23.0 cm) were baited with a mixture of rolled oats, scratch grain, and sunflower seeds and provided with cotton nesting material. Trapping grids varied from 24 to 36 stations with a trap spacing of 25 to 35 m and 1 trap/station. Two grids used as part of a longer study consisted of 64 and 120 stations, respectively. A minimum, therefore, of about 300 trap-days/site was maintained. Since the effective area trapped by each grid was hard to determine and likely varied in different habitats (Van Home, 1982), actual densities were not calculated, but rather a relative index of captures per unit trapping effort was used to represent abundance of the three chipmunk species on a scale of 0 to 4. In two cases, sight records of a species not captured were considered in the presence-absence data and caused a species' abundance rank to change from 0 to 1 for that site. Information recorded for each chipmunk captured included standard body measurements, weight, sex, age (juvenile or adult), molt condition, reproductive condition, vocalizations and behavior during handling, and period of day when captured. One hundred-eighty March 1991 15 The Southwestern Naturalist

Equus Linnaeus, 1758:73. Type species [Equus] caballus Linnaeus, 1758:73 (domestic), by original designation. Caballus Rafinesque, 1815:55. Substitute name for [Equus] caballus Linnaeus, 1758. Asinus Gray, 1825:244. Type species Equus vulgaris ( [Equus] asinus Linnaeus, 1758:73) by subsequent designation. Not Asinus Brisson, 1762:70 which is not Linnaean. Not Asinus Frisch, 1775, which is not Linnaean. Hippotigris Smith, 1841:321. Type species [Equus] zebra Linnaeus, 1758:74, by original designation. Zebra Allen, 1909:163. Type species Equus burchelli granti de Winton, 1896:319, by original designation. In legend to figures 7, 8d, and 8e (but not figures 3–6 and 9–19, where Equus is employed). Evidently a lapsus. Dolichohippus Heller, 1912:1. Type species Equus grevyi Oustalet, 1882:12, by original designation and monotypy. Microhippus Reichenau, 1915:152. Type species Equus przewalskii Polyakov, 1881:1, by monotypy. Not Microhippus Matschie, 1922:68, which was used in a subgeneric sense. Not Microhippus Pocock, 1948:764, which is preoccupied. Quagga Shortridge, 1934:397. Type species [Equus] quagga Gmelin, 1788:213, by monotypy. Onager Quinn, 1957:10. Type species [Equus] onager Boddaert, 1784:160, by original designation. Not Onager Brisson, 1762: 72, which is not Linnaean. Not Onager Zimmerman, 1780: 80, which was used in a subgeneric sense. Hemippus Dietrich, 1959:14. Type species Equus hemippus I. Geoffroy Saint-Hilaire, 1855a:1214, 1217, by monotypy. Hemionus Dietrich, 1959:14. Type species Equus hemionus Pallas, 1775:394, by original designation. Not Hemionus Cuvier, 1821:555, a nomen nudum (Trumler 1961). Not Hemionus Stehlin and Graziosi, 1935, which was used as a subgenus. Hydruntinus Radulesco and Samson, 1962:174. Type species Equus hydruntinus Regalia, 1904. Mesolithic to Neolithic subfossil.

Marmots are the largest ground squirrels and have been extensively studied by sociobiologists investigating the evolution of mammal societies. Being a member of the sciurid clade, traditionally considered inclined to convergence, they are also a group on which to test the hypothesis of sciurid propensity to homoplasy of osteological characters. In the present analysis, the dorsal and lateral surfaces of the cranium of all living marmot species are compared with geometric morphometric techniques. Phenetic groups are found which reflect the subgeneric classification of marmots and are consistent with previous morphometric analyses of the mandible and ventral cranium. Two species have distinctive morphologies and phenetic relationships not congruent with phylogeny. Marmota vancouverensis is highly divergent for osteological characters, fur colour and behaviour despite its young age and close genetic similarity to Marmota caligata. Its small population may represent a rare chance to study evolutionary processes during rapid allopatric speciation in mammals, but strong conservation efforts are required to preserve this unique component of the Vancouver Island biodiversity. Also, Marmota monax has distinctive cranial traits. These are possibly related to its long separate evolutionary history and unique ecology and behaviour. Size-related convergence is not evident in Marmota. When outgroup species are included, Spermophilus, Cynomys, Tamias, and Sciurus group together on one branch, Marmota on the other. This is best explained as a retention of the ancestral morphology in the smaller members of the Marmotini (Spermophilus, Cynomys, and Tamias) and the evolution of derived morphology in Marmota. Riassunto Le marmotte sono i piu grandi sciuridi viventi e sono state intensamente studiate dai sociobiologi nel tentativo di comprendere le dinamiche evolutive delle societa mammaliane. Questa linea di scoiattoli terricoli rappresenta, inoltre, un ambito per la verifica dell'ipotesi di propensione all'omoplasia nella forma delle ossa degli sciuridi. Nella presente indagine, le superifici dorsale e laterale del cranio di tutte le specie viventi di marmotte sono state analizzate e confrontate servendosi di tecniche di morfometria geometrica. I raggruppamenti fenetici scoperti rispecchiano la classificazione in sottogeneri delle marmotte e sono congruenti con i risultati di precedenti studi sulla morfologia della mandibola e del lato ventrale del cranio. Due specie, tuttavia, hanno caratteristiche peculiari e le loro relazioni fenetiche con gli altri membri del genere non-riflettono i rapporti filogenetici. Marmota vancouverensis, a dispetto della recente origine evolutiva e della somiglianza genetica con M. caligata, possiede caratteri osteologici fortemente divergenti, oltre a tratti del comportamento e colore della mantello unici tra le marmotte. La piccola popolazione di M. vancouverensis potrebbe rappresentare una rara opportunita di indagare i meccanismi evolutivi dei processi di speciazione rapida in allopatria. La sopravvivenza di questa componente unica della biodiversita dell'isola di Vancouver dipendera, tuttavia, da un'efficace e duratura politica conservazionista. Anche Marmota monax presenta caratteri craniometrici peculiari parallelamente ad aspetti singolari della sua ecologia ed etologia, ma, diversamente da M. vancouverensis, ha alle spalle una lunga storia evolutiva. Le marmotte non-sembrano, invece, caratterizzate da forme convergenti in specie di taglia simile. I rappresentanti dei generi Spermophilus, Cynomys, Tamias e Sciurus, adoperati come outgroup, si raggruppano insieme, ben separati dalle marmotte. Questo si puo spiegare con la conservazione di caratteri plesiomorfici nei membri di piccola taglia dei Marmotini (Spermophilus, Cynomys, and Tamias) e con l'evoluzione di tratti autoapomorfici nelle ben piu grosse marmotte.

We report the first record of the Inyo shrew, Sorex tenellus, from the central Great Basin. A single specimen was captured in Great Basin National Park at 3,000 m elevation in habitat dominated by Picea engelmanni. This constitutes a northeastern range extension of 300 km into east-central Nevada.

The architectonics of tenrec (Tenrecidae) spines has a convergent similarity to that in porcupine (Erethizontidae) quills and is also species specific. Like in Erethizontidae species, in tenrecs of the genus Hemicentetes using the tactics of active front defense, defensive spines of the trunk and crown fan are covered with an inverted cuticle, which serves as a particular harpoon. In Echinops and Setifer tenrecs using the tactics of passive defense (they can roll into a ball), spines of the trunk have no such cuticle. Spines of the stridulation organ in Hemicentetes are specialized to produce ultrasonic sounds due to rubbing against each other, and the spine structure reflects this peculiarity. The cuticle of trunk spines in Echipops telfairi is unique.

Although ground squirrels (Spermophilus) and prairie dogs (Cynomys) are among the most intensively studied groups of mammals with respect to their ecology and behavior, a well-resolved phylogeny has not been available to provide a framework for comparative and historical analyses. We used complete mitochondrial cytochrome b sequences to construct a phylogeny that includes all 43 currently recognized species in the two genera, as well as representatives of two closely related genera (Marmota and Ammospermophilus). In addition, divergence times for ground squirrel lineages were estimated using Bayesian techniques that do not assume a molecular clock. All methods of phylogenetic analysis recovered the same major clades, and showed the genus Spermophilus to be paraphyletic with respect to both Marmota and Cynomys. Not only is the phylogeny at odds with previous hypotheses of ground squirrel relationships, but it suggests that convergence in morphology has been a common theme in ground squirrel evolution. A well-supported basal clade, including Ammospermophilus and two species in the subgenus Otospermophilus, diverged from all other ground squirrels an estimated 17.5 million years ago. Between 10 and 14 million years ago, a relatively rapid diversification gave rise to lineages leading to marmots and to several distinct groups of ground squirrels. The Eurasian ground squirrels diverged from their North American relatives during this period, far earlier than previously hypothesized. This period of diversification corresponded to warming climate and spread of grasslands in western North America and Eurasia. Close geographic proximity of related forms suggests that most species evolved in or near their current ranges.

The systematics and distribution of Asiatic short-tailed shrews (genus Blarinella ) are confused. In this study, a taxonomic revision of Blarinella is presented. We investigated a sample of 98 specimens, 84 of which had complete skulls that were subjected to multivariate and univariate analyses. The results show that three species of Blarinella can be recognized. The species Blarinella quadraticauda occurs in Baoxin, Wenchuan, Daofu, and other localities in western Sichuan. Blarinella griselda and B. wardi , usually placed as subspecies of B. quadraticauda , are recognized as two valid species. The former has a wide distribution from southern Gansu to southern Yunnan, and northwestern Yunnan to northwestern Hubei. The latter is restricted in northern Burma and northwestern Yunnan. The range of B. quadraticauda does not overlap that of B. wardi , while the range of B. griselda partially overlaps that of B. wardi and probably includes the entire range of B. quadraticauda.

Many systematic relationships among Chinese white-toothed shrews of genus Crocidura are presently unresolved. In this paper, a taxonomic revision of Crocidura from Southern China is presented. We studied 338 specimens from Burma, China, Korea, Pakistan, Turkey, Middle and Central Asia, and Russia (Appendix I), 285 of which had complete skulls that were analyzed with principal component and discriminant analyses. Results indicated that 6 species of Crocidura can be recognized in South China. C. fuliginosa occurs in Southwestern and Eastern China, C. attenuata is broadly distributed throughout Southern China, and C. horsfieldii is restricted to the southern part of China. C vorax and C. rapax, usually placed as synonyms of the European C russula, are recognized as 2 valid species whose ranges overlap in Southwestern China. C. shantungensis of Eastern Asia extends to the northern part of Southern China and is distinct from C. suaveolens and C. gmelini of Middle and Central Asia, respectively.

There are 14 species of marmots distributed across the Holarctic, and despite extensive systematic study, their phylogenetic relationships remain largely unresolved. In particular, comprehensive studies have been lacking. A well-supported phylogeny is needed to place the numerous ecological and behavioral studies on marmots in an evolutionary context. To address this situation, we obtained complete cytochrome (cyt) b sequences for 13 of the species and a partial sequence for the 14th. We applied a statistical approach to both phylogeny estimation and hypothesis testing, using parsimony and maximum likelihood-based methods. We conducted statistical tests on a suite of previously proposed hypotheses of phylogenetic relationships and biogeographic histories. The cyt b data strongly support the monophyly of Marmota and a western montane clade in the Nearctic. Although some other scenarios cannot be rejected, the results are consistent with an initial diversification in North America, followed by an invasion and subsequent rapid diversification in the Palearctic. These analyses reject the two major competing hypotheses of M. broweri's phylogenetic relationships-namely, that it is the sister species to M. camtschatica of eastern Siberia, and that it is related closely to M. caligata of the Nearctic. The Alaskan distribution of M. broweri is best explained as a reinvasion from the Palearctic, but a Nearctic origin can not be rejected. Several other conventionally recognized species groups can also be rejected. Social evolution has been homoplastic, with large colonial systems evolving in two groups convergently. The cyt b data do not provide unambiguous resolution of several basal nodes in the Palearctic radiation, leaving some aspects of pelage and karyotypic evolution equivocal.

Severtzov's sheep (Ovis ammon severtzovi; Nasonov 1914) has a 2n = 56 diploid chromosome number and a karyotype consisting of two pairs of biarmed and 25 pairs of acrocentric autosomes, a large acrocentric X, and a minute biarmed Y. The G-banding patterns of the largest pair of biarmed chromosomes were identical to those of the largest biarmed chromosomes in all wild and domestic sheep of the genus Ovis. The banding patterns of the second pair of biarmed chromosomes were identical to the third pair of biarmed chromosomes in all sheep of the genus Ovis with 2n = 54 and to the third largest pair of chromosomes in the 2n = 52 karyotype of the Siberian snow sheep (O. nivicola). The G-banded karyotype of Severtzov's sheep is consistent with all subspecies of argali (O. ammon spp.) that have been karyotyped. Numerical ascription of acrocentric chromosome equivalents based upon the fundamental karyotype of Ovis that gave rise to the biarmed chromosomes of severtzovi are 1 and 3, and 5 and 11 for the largest and second largest biarmed chromosomes, respectively. Based upon diploid chromosome number and G bands, Severtzov's sheep should be considered a subspecies of argali and not a urial.

The distribution of the great gerbil (Rh. opimus) in China was analyzed on the basis of published data, museum collections, natural maps, and authors' field observations. A preliminary scheme was plotted for range regionalization, in which three parts have been isolated or, according to classification of Dubrovskiĭ and Kucheruk (1971), three regional complexes of autonomous groups of populations: Dzungarian, Beishanian and Alashanian. West of the main area of the great gerbil distribution, an isolated colony is located in the Ili River valley, connected with the Kazakhstan portion of the range (Ili regional complex of autonomous groups of populations). Additional field observations are required for more detailed description of the great gerbil range.

The distribution of the great gerbil (Rh. opimus) in China was analyzed on the basis of published data, museum collections, natural maps, and authors' field observations. A preliminary scheme was plotted for range regionalization, in which three parts have been isolated or, according to classification of Dubrovskii and Kucheruk (1971), three regional complexes of autonomous groups of populations: Dzungarian, Beishanian and Alashanian. West of the main area of the great gerbil distribution, an isolated colony is located in the Ili river valley, connected with the Kazakhstan portion of the range (Ili regional complex of autonomous groups of populations). Additional field observations are required for more detailed description of the great gerbil range.