OUP user menu

Revision of the subspecies of the American pika, Ochotona princeps (Lagomorpha: Ochotonidae)

David J. Hafner, Andrew T. Smith
DOI: http://dx.doi.org/10.1644/09-MAMM-A-277.1 401-417 First published online: 16 April 2010

Abstract

Historically, a large number of taxonomic forms has been recognized within Nearctic pikas (Lagomorpha: Ochotonidae; Ochotona), including up to 13 species and 37 subspecies. After 1965, 2 species and 37 forms have been recognized: the monotypic O. collaris of Alaska, British Columbia, Yukon, and Northwest Territories, and O. princeps, with 36 subspecies spread throughout western Canada and the western United States. The 36 subspecies of O. princeps have been distinguished by subtle differences, particularly in pelage coloration and body size, within the highly fragmented distribution of the species on isolated “islands” of cool, rocky habitat. However, molecular phylogenetic studies (allozyme electrophoresis and sequencing of both mitochondrial and nuclear genomes) indicate the existence of 5 phylogenetic lineages within O. princeps. The cohesiveness of each lineage has been reinforced during glacial stages by introgressive hybridization among currently isolated populations within each lineage. In contrast, the low level of cranial variation and lack of consistent differentiation in cranial characters, pelage coloration, or body size among the 5 lineages indicates idiosyncratic interlocality variation due to high inbreeding within highly isolated populations, genetic drift, and possibly selection for common traits. Examination of allozymic, morphological, and nuclear DNA data indicates previous introgressive hybridization among several of the lineages, probably associated with contact during the Last Glacial Maximum. Herein we characterize morphometric variation between and among O. collaris (n = 164) and O. princeps (n = 1,999) and revise the subspecific taxonomy of O. princeps to 5 subspecies based on molecular phylogenetic lineages, at least 3 of which are known to possess a unique dialect in the short call: O.p. princeps (Northern Rocky Mountains), O. p. fenisex (Coast Mountains and Cascade Range), O. p. saxatilis (Southern Rocky Mountains), O. p. schisticeps (Sierra Nevada and Great Basin), and O. p. uinta (Uinta Mountains and Wasatch Range of central Utah). These 5 subspecies represent evolutionarily meaningful units for consideration of possible management applications if populations of O. princeps are imperiled by human activities.

Key words
  • allozymes
  • mitochondrial DNA
  • morphology
  • nuclear DNA
  • Ochotona collaris
  • Ochotona princeps
  • pika
  • subspecies
  • systematics
  • taxonomy

Recognition of subspecies within a geographically widespread species has typically involved an iterative process of accumulation of names as new forms are discovered and described and reduction in names as forms are compared morphologically and synonymized based on similarity. Inevitably, species with highly fragmented distributions and closed breeding structure, which can lead to inbreeding and genetic drift, are more likely to include a higher number of subspecies, particularly if drift-induced morphological change obfuscates shared similarities among isolated populations. But if the subspecific classification is to have evolutionary meaning, as envisioned by Lidicker (1960, 1962) and expanded by Endler (1977), geographic variation must be evaluated in an evolutionary context to reveal phylogeo-graphic patterns that could be hidden beneath random and idiosyncratic variation in ecophenotypically plastic characters. By identifying these evolutionary units as subspecies, the subspecies takes on enhanced importance for evolutionary studies and conservation management.

For example, diversification within the well-studied pocket gophers (Geomyidae) is often noted as being overrepresented at the subspecific level (Hafner et al. 2008; Hoffmeister 1986; Patton and Smith 1990; Rios and Álvarez-Castañeda 2007). Botta's pocket gopher (Thomomys bottae) occurs over much of western North America (Hall 1981) in restricted patches of soil reflecting its fossorial lifestyle, and like other geomyid species displays often flamboyant variation in size (correlated with soil friability and vegetative cover—Hafner et al. 2008; Hoffmeister 1986) and pelage coloration (strongly correlated with soil coloration—Krupa and Geluso 2000). As a result, 195 different subspecies of T. bottae have been named based on morphological criteria. Subsequent investigations (most incorporating genetic criteria) have reduced the number of subspecies of T. bottae in select regions: Arizona, 41 subspecies to 14, a 66% reduction (Hoffmeister 1986); California, 46 subspecies to 15, a 67% reduction (Patton and Smith 1990); and Baja California, 6 subspecies to 1, an 83% reduction (Rios and Álvarez-Castañeda 2007). Similarly, Hafner et al. (2008) used morphometric, karyotypic, and mitochondrial DNA (mtDNA) and nuclear DNA (nDNA) sequence data to reduce the number of recognized subspecies in another pocket gopher, Cratogeomys castanops, from 26 to 4 (85% reduction), with 2 of those 4 included in a separate species, C. goldmani. It is likely that further application of genetic data would result in synonymy of many of the remaining subspecies of geomyids.

The American pika (Ochotona princeps) shares several major demographic characteristics with pocket gophers, characterized by a highly fragmented distribution of precinctive, inbreeding populations distributed over a large geographic area. The species is widely distributed over the Intermountain West of North America, spanning the western contiguous United States and southwestern Canada, yet often is highly restricted to remote patches of talus habitat within generally cool, mesic, and usually montane habitat (Hafner 1993; Smith and Weston 1990). Consequently, previously undetected populations continue to be discovered (Beever et al. 2008; Hafner 1994; Millar and Westfall, in press). Also like pocket gophers, pikas exhibit extensive geographic variation in pelage coloration (Smith and Weston 1990), ranging from light buffy to nearly black. Describing the distinguishing characters of 25 subspecies of O. princeps, Howell (1924) relied heavily on pelage coloration (all 25 subspecies), while mentioning overall body size or skull shape in less than one-half and one-third (respectively) of the subspecies. The combination of distinctive coloration, extreme geographic isolation, and subtle cranial differences among populations that continue to be discovered has resulted in recognition of as many as 12 species and 36 subspecies within currently recognized O. princeps.

Recent analyses of mtDNA and nDNA sequence data from throughout the distribution of Nearctic pikas (O. princeps and O. collaris) have clarified the major phylogeographic patterns within O. princeps (Galbreath et al. 2009, in press). In this paper we review molecular genetic variation within Nearctic pikas (allozymes, mtDNA sequence, and nDNA sequence) and geographic patterns of dialects in the short call vocalization, undertake a cranial morphometric analysis of geographic variation within O. princeps relative to O. collaris, and revise the subspecific taxonomy of O. princeps. In general, we use the biological species concept in that we recognize reciprocally monophyletic lineages while emphasizing molecular similarities as indicative of recent or occasional long-distance gene flow among currently isolated populations, and recognize evidence of past introgressive hybridization as indicative of potential gene flow between currently isolated lineages.

In the century following the initial naming of Nearctic pikas (Richardson 1828), 13 species and 25 subspecies were described. The last full revision of the American pika was conducted by Howell (1924), who recognized 3 species: a monotypic O. collaris, O. princeps (16 subspecies), and O. schisticeps (9 subspecies). Subsequently, Miller (1936) synonymized O. schisticeps under O. princeps, based primarily upon the study of Borell (1931), and the contemporary content of O. princeps includes 36 subspecies (Hall 1981; Smith and Weston 1990). In the most recent comprehensive treatment of O. princeps Hoffmann and Smith (2005) delineated the 36 subspecies into 5 main groups following the allozyme-based genetic study by Hafner and Sullivan (1995). These are listed in the“Synonym”section of Hoffmann and Smith (2005) as representing groups from the 1) Northern Rockies (= northern Rocky Mountains), 2) Central Rockies, 3) Southern Rockies, 4) Sierra Nevada-Great Basin, and 5) Cascades. Select subspecies were highlighted in bold in this treatment, but these taxa were not specifically identified as overarching subspecies, and this summary did not constitute a revision of the species. The senior author of this treatment viewed the charge to delineate geographically distinct groups of subspecies, not to formulate new content for the species, despite the introductory statement of the editors (Wilson and Reeder 2005:xxxiii) that“Currently recognized subspecies are listed in boldface type, followed by their junior synonyms.”Because of this statement and because other contributing authors did use bolded names in the“Synonym”section of their accounts to delineate subspecies, the Web site for Mammal Species of the World (http://www.bucknell.edu/MSW3/browse.asp?id=13500064; accessed 20 August 2009) has interpreted the bolded taxa as identifying five subspecies of O. princeps representing the 5 groupings outlined above, in order: princeps, figginsi, saxatilis, schisticeps, and taylori. Compounding this taxonomic confusion, these 5 geographic groupings did not accurately represent the genetic units defined by Hafner and Sullivan (1995), who did not recognize a separate Central Rockies geographic unit (figginsi). Also, the appropriate name for a Cascades form would be O. p. fenisex (not taylori), because the type locality of taylori is within the Sierra Nevada lineage.

Hafner and Sullivan (1995) examined allozymic patterns at 26 presumptive genetic loci among 56 populations of O. princeps and a reference outgroup composed of specimens from 3 Alaskan populations of O. collaris. They found low overall allozymic variation in all examined populations, likely due to a combination of the demographic pattern of the species—highly fragmented metapopulations (Gilpin 1991; Smith 1987) with a closed breeding structure—and, in some regions, derivation of current populations from a genetically depauperate source population. They found strong support for 5 major groups within O. princeps. Four of these were geographically structured groups (Fig. 1) that corresponded with subspecies that were originally described as full species: Northern Rocky Mountains (NRM; O. p. princeps), Southern Rocky Mountains (SRM; O. p. saxatilis), Sierra Nevada (SN; O. p. schisticeps), and Cascade Range (CR; O. p. fenisex). A 5th group included 8 populations, considered as“contact sites,”that exhibited mixtures of alternate fixed or major alleles of neighboring units; 5 populations were fully monomorphic and 3 populations were polymorphic for such alleles. Based on allozymic mixture in geographically intermediate metapopulations, Hafner and Sullivan (1995) concluded that no indication existed that O. princeps may contain cryptic species (Glover et al. 1977) but noted that the“evolutionarily volatile system”dictated by the demographics of O. princeps renders them potentially vulnerable to natural and human-induced change in habitat, which would be expected to be quite different for each geographic group. For example, in wetter, higher-elevation areas human impact that decreases vegetative cover and increases- erosion might artificially maintain open talus and prolong population persistence, whereas the same impacts might adversely affect lower-elevation populations in drier regions by removing limited food resources.

Fig. 1—

Geographie distribution of Ochotona princeps indicating 5 genetic units identified in allozyme, mitochondrial DNA, and nuclear DNA studies (gray outlines), and grouped localities used in morphometric analyses (numbered as in Appendix I). Localities used in morphometric transects are labeled as follows: I (localities 2–4), II (5–7), III (45–48), IV (8–10, 17), V (12–16, 19), and VI (42, 49–52). Past introgressive hybridization between subspecies, prior to the Last Glacial Maximum (LGM), occurred at (1) Mt. McLoughlin in southern Oregon and (2) the Aquarius Plateau in southern Utah. More recent, post-LGM introgression is indicated (3) along the Colorado River of central Colorado and (4) across southern British Columbia. Localities from which dialects have been described are indicated by letters (as in Table 3); localities A and B (O. collaris) are in Alaska and Yukon Territory, respectively.

Galbreath et al. (2009) evaluated sequence variation in the mtDNA cytochrome-b (Cytb) gene and part of the D-loop (total of 1,668 base pairs [bp]) in the control region among 37 populations of O. princeps. In addition to confirming the monophyly of populations of O. princeps relative to O. collaris (designated outgroup), the resulting chronogram revealed 5 well-supported clades whose divergence predated the Last Glacial Maximum (LGM). Four of these corresponded to the geographic lineages previously identified by allozyme analysis (Hafner and Sullivan 1995), and the 5th was composed of 3 populations from central Utah (CU; Uinta Mountains and Wasatch Range) that had been included in the contact sites group based on allozymes. Only 1 of the other 5 contact site populations (all from outside of Utah) was included in this study; it grouped with neighboring populations in the Southern Rocky Mountains lineage. Galbreath et al. (2009) found the 5 lineages (Fig. 1) to be associated with mountain systems rather than individual“sky islands,”suggesting maintenance of regional cohesion via gene flow promoted by elevational shifts during glacial maxima.

Galbreath et al. (in press) examined sequence variation in mtDNA (Cytb and D-loop) and nDNA (mast cell growth factor, —550 bp; and protein kinase C iota, ∼660 bp) among 64 populations of O. princeps, including those from both Hafner and Sullivan (1995) and Galbreath et al. (2009), and again using O. collaris to root the phylogeny. The resulting phylogeny identified the same 5 major lineages associated with mountain systems (Fig. 1), with phylogeographic concordance between mtDNA and nDNA markers. All populations previously considered contact sites by Hafner and Sullivan (1995) were assigned unambiguously to the 5 lineages. Moreover, inspection of allozymic data from Hafner and Sullivan (1995) revealed that 6 of the 8 contact sites (4 of which were assigned to the CU lineage) exhibited fixation of shared, ancestral alleles rather than any evidence of past gene flow. However, the 2 remaining sites did exhibit low-level mixtures of fixed alleles associated with neighboring lineages, evidently the result of introgression following secondary contact during the LGM. Evidence for LGM gene flow between previously diverged lineages also was detected in the nDNA markers along the transition between the NRM and SRM lineages. Comparison of allozyme complements, mtDNA haplotypes, nDNA alleles, cranial morphology, and vocalization dialects along the same contact revealed broadscale concordance but fine-scale discordance. This was interpreted as a consequence of shifting biogeographical barriers, historical gene flow along the contact zone, and differences in relative rates of evolution and lineage sorting for alternative markers (Galbreath et al., in press).

At least 9 distinct acoustic signals have been described from O. princeps (Conner 1985). The short call is given in a variety of social and alarm situations and is most often given when the animal is perched on rocks (Broadbooks 1965; Conner 1982; Ivins and Smith 1983; Somers 1973). As noted by Hafner and Sullivan (1995), at least 3 of the phylogenetic lineages (NRM, SRM, and SN) are characterized by distinct vocal dialects in the short call. Although Conner (1982) defined this variation in intraspecific calls as geographic variation instead of dialects, the pattern of vocalizations in fact conforms to the definition of avian vocal dialects (Marler 1960; Nottebohm 1969). Pikas are able to discriminate between different dialects of the short call (Conner 1983), which appears on day 12 in young captive pikas and is structurally identical to future adult calls (i.e., it is not learned—Conner and Whitworth 1985). Somers (1973) described distinct dialects in the NRM (Wyoming, Utah, and Colorado) and SRM (Colorado and New Mexico) lineages, separated by the Colorado River in central Colorado. Galbreath et al. (in press) detailed the regionally concordant but locally discordant patterns of allozyme complements, cranial morphology, mtDNA haplotypes, nDNA alleles, and dialects along the Colorado River. Conner (1982) described an additional dialect, distinct in comparison to the SRM dialect, from populations of the SN lineage in California and Utah. Although short calls have not been analyzed from the CR or CU lineages, Broadbooks (1965) remarked that a greater difference was apparent between short calls of O. princeps from the Big Horn Mountains of Wyoming (NRM lineage) and the Cascade Range in Washington (CR) than between the latter and O. collaris of Alaska. Trefry and Hik (in press) described regional variation in the short call of O. collaris in Alaska and Yukon, Canada, and found it differed primarily in shorter duration from that of O. princeps in the Rocky Mountains of Alberta, Canada. The 4 dialects thus far described (Fig. 2) differ primarily in fundamental frequency and duration. In addition, the short call of the SN lineage is usually of multiple notes, that of the SRM lineage usually is a single note, and those of the NRM lineage and O. collaris invariably are of a single note.

Fig. 2—

Dialects of the short call in Nearctic species of Ochotona (O. collaris and genetic units of O. princeps). Means of data from Conner (1982), Somers (1973), and Trefry and Hik (in press). Letters conform to Fig. 1 and Table 3.

The short call appears to function primarily in social contexts (primarily territorial behavior—Conner 1984), and only 5% of the total calls analyzed by Svendsen (1979) involved disturbances caused by something other than pikas. However, Ivins and Smith (1983) demonstrated that pikas discriminate between predatory and nonpredatory species (calling more frequently and for longer duration in response to predators) and responded differently to weasels (Mustela frenata and M. erminea) than to pine martens (Martes americana). Specifically, pikas delay alarm calls in response to weasels, which actively hunt pikas in talus. They noted at least 3 occasions in which weasels pursued pikas after hearing 1 short call. During collection of pikas throughout their geographic distribution (D. J. Hafner, pers. obs.) pikas commonly appeared on rocky perches apparently in response to pika-like short calls emitted by the collector, and on at least a dozen occasions weasels appeared within approximately 1 m of the collector in response to these calls. Response of pikas to avian predators, in contrast, has not been studied, although short call responses to hawks (Buteo sp.—Broadbooks 1965), large birds, and airplanes (Krear 1965; Svendsen 1979) have been noted anecdotally. During fieldwork (D. J. Hafner, pers. obs.) predation or attempted predation on pikas by common ravens (Corvus corax) and Cooper's hawks (Accipiter cooperii) were observed. Similarly, the flight of any large bird (including hawks, ravens, or vultures) could be tracked across talus slopes by the progressive short calls of pikas (D. J. Hafner and A. T. Smith, pers. obs.). Alertness to potential avian predators, coupled with the long amount of time spent during vulnerable surveillance on rocky perches (more than one-half of the time an animal is active on the surface—Smith and Ivins 1984), suggests a selective basis for the extensive geographic variation in pelage coloration within O. princeps. Although the degree of matching between pelage and talus coloration has not been quantified, pikas of darkest coloration are from black basalt talus (e.g., O. p. nigrescens of the Jemez Mountains of New Mexico, O. p. goldmani of lava beds southern Idaho, and O. p. fumosa of the volcanic Cascade Range in Oregon), and the lightest occur on lighter-colored sedimentary shale, sandstone, limestone, and quartzite (e.g., O. p. lutescens of the Rocky Mountains in Alberta and O. p. nevadensis of the Ruby Mountains of northeastern Nevada). A collection of digitized images from 35-mm color slides of 948 O. princeps from 56 populations between New Mexico and British Columbia showing dorsal pelage of the freshly collected specimen, a gray card (18% reflectance), and the talus substrate upon which the pika was shot is available for inspection from D. J. Hafner.

Materials and Methods

We examined a total of 2,163 specimens of Nearctic pikas (O. collaris, n = 164; O. princeps, n = 1,999) from collections throughout North America (Appendix I). Fourteen mensural characters were recorded from adult specimens, including 2 external measurements (total length [TL] and hind-foot length [HF]) recorded directly from the skin tag and 12 cranial measurements measured with handheld calipers to the nearest 0.01 mm: greatest skull length (GL), zygomatic breadth (ZB), bullar breadth (BB), least interorbital breadth (LIB), nasal length (NL), rostral breadth (RB; taken at the premaxilla–maxilla junction), bullar length (BL), diastema length (DL), length of molar toothrow (MTR), width of palatal bridge (PB = length of bony palate; Figs. 3a and 3b), width of P3 (WP3), and width of P4 (WP4).

Fig. 3—

Comparative appearance of the palatal bridge (outlined in circles) in extreme examples of Ochotona princeps schisticeps (a; NMMNH 852, adult male, from California: Siskiyou Co.; Red Butte, Mt. Shasta) and most populations of other subspecies of O. princeps (b; NMMNH 863, adult male, O. p. fenisex, from Oregon: Lane Co.; 5 mi. N, 14 mi. E McKenzie Bridge), c) Distribution of measurements (mean ± SD) of palatal bridge width for males of O. p. schisticeps (white bars; n = 246, 1.74 ± 0.31), O. p. princeps (gray bars; n = 382, 2.15 ± 0.40), and the remainder of O. princeps (black bars with white outlines; n = 460, 2.52 ± 0.44). All 3 group means are significantly different from one another (P < 0.001; Tukey pairwise mean comparisons).

Populations were grouped into 129 regional localities based on geographic proximity without crossing obvious geographic discontinuities or currently recognized subspecific boundaries. The only exception is that all specimens of the monotypic O. collaris were combined into a single group (group 1) for analysis (Fig. 1; Appendix I). Obvious geographic discontinuities included gaps of >5 km between continuous montane habitat that were low-elevation, arid ecosystems, and major rivers. The maximum dispersal distance observed for individual pikas is approximately 3 km (Tapper 1973), and Hafner (1994) estimated that post-Xerithermal (6,000 years) recolo-nization of sites >20 km distant was unlikely.

All morphometric analyses were carried out using SYSTAT version 7.0 (Wilkinson 1997). Raw data were standardized (X̄ = 0, SD = 1) to reduce the effects of individual size variation, and Lilliefors test was used to test for normality of the transformed data. We used multivariate analysis of variance (MANOVA) to test the null hypothesis of no significant difference between sexes and among groups, and we used both principal component analysis and direct discriminant function analysis of standardized morphometric data to determine if groups could be separated without or with (respectively) an a priori hypothesis of group membership in clades identified by molecular genetic (allozyme, mtDNA, and nDNA) analyses. Tukey pairwise comparisons of unstandardized variables were used to assess differences among external and cranial dimensions of taxa relative to overall means for O. princeps. Tukey pairwise comparison of standardized group-population means was used to identify populations that are morphologically similar at P > 0.95. Agglomerative, polythetic hierarchical cluster analysis was used to examine similarities among short calls, and linear regression analysis was used to evaluate correlation among selected cranial characters and characteristics of the short call.

Results and Discussion

Molecular phylogenetic pattern.—Analyses of allozyme complements, mtDNA haplotypes, and nDNA alleles (Galbreath et al. 2009, in press; Hafner and Sullivan 1995) were concordant in delineating 5 lineages within O. princeps that are reciprocally monophyletic relative to O. collaris. Subsequent to divergence from O. collaris, the CR and possibly the SN lineages diverged prior to the nearly simultaneous divergence of the NRM, SRM, and CU lineages (Galbreath et al., in press), and this lineage diversification has proceeded through at least 2 glacial-interglacial cycles (Hafner and Sullivan 1995). Evidence of past and ongoing introgressive hybridization among pairs of lineages was detected in comparative allozyme complements, shared nDNA alleles, and incongruent patterns among all 3 genetic measures at zones of past or current contact between lineages (Galbreath et al., in press; Hafner and Sullivan 1995). Subspecific names that would apply to these 5 genetic lineages (Fig. 1) are O. p. princeps (NRM), O. p. fenisex (CR), O. p. saxatilis (SRM), O. p. schisticeps (SN), and O. p. uinta (CU).

Morphometric pattern.—Results of a MANOVA revealed significant secondary sexual dimorphism in all external and cranial variables except PB (P < 0.01; multivariate Wilk's lambda, Pillai trace, and Hotelling-Lawley trace tests all with P < 0.001), with males larger than females, and sexes subsequently were treated separately. For all subsequent analyses of geographic variation, patterns were similar between sexes. Previously, Millar (1971), Brunson (1973), and Smith (1981) detected only slight secondary sexual size dimorphism (males larger than females), and crania of O. princeps > 1 year of age exhibited no size differences (Weston 1981; Wiseley 1973).

A MANOVA based only on males revealed significant differences among populations for 9 of 14 variables (P < 0.05; multivariate Wilk's lambda, Pillai trace, and Hotelling-Lawley trace tests all with P < 0.001). Inspection of a plot of the first 2 principal components of a principal component analysis of grouped populations (not shown) could not discern any distinct groups related to either the 36 described subspecies of O. princeps or the 5 genetic lineages but instead indicated broad overlap among all groups. At a more detailed geographic level and in agreement with these results, Coots (1972) found no morphological distinction between 2 subspecies in Oregon (O. p. brunnescens and O. p. fumosa, both in the CR lineage). Because of the lack of delineation of groups based on morphometric data, populations instead were grouped a priori according to the 5 lineages identified by molecular analyses (Fig. 1) and subjected to discriminant function analysis.

Results of a discriminant function analysis indicated either 2 (males) or 3 (females) functions with eigenvalues >1. In both cases character loadings on the 1st function (DFI) for all variables were high and positive, indicating a significant influence of overall size (Table 1). Minimum convex polygon plots of the grouped locality means on the first 2 functions (Fig. 4) revealed broad overlap, particularly among O. collaris, O. p. fenisex, O. p. saxatilis, and O. p. uinta, with O.p. princeps bridging the gap between these former taxa and O. p. schisticeps. Percentage of individuals correctly identified to subspecies within O. princeps (averaged for males and females) based on posterior classification was >95% only between O.p. fenisex and O.p. schisticeps and between O.p. uinta and O. p. schisticeps', all other comparisons of neighboring subspecies ranged from 81% (between O. p. princeps and O. p. saxatilis) to 86% (between O. p. princeps and O. p. schisticeps) correctly classified. Galbreath et al. (in press) have mapped the local discordance between molecular and morphometric patterns along the Colorado River of central Colorado. We evaluated the other broad geographic area of overlap along 2 west-east transects in southern British Columbia (between O.p. fenisex and O.p. princeps, Fig. 1) using box diagrams and Tukey pairwise mean comparisons (Fig. 5). Stepped clinal variation generally corresponding to molecular delineations is apparent in the northernmost transect (Fig. 5a), but smoother clinal variation is evident in the southernmost transect where geographically intermediate populations genetically identified as O. p. princeps exhibit a morphology more similar to that of O. p. fenisex.

View this table:
Table 1

Component loadings for morphometric variables, and eigenvalues and total variance explained for the first 2 components of a discriminant function analysis based on means of standardized data for 129 grouped populations of Ochotona princeps. Explanations for abbreviations of variables are provided in text.

VariableComponent loadings
MalesFemales
IIIIII
TL0.648−0.2310.5070.219
HF0.766−0.0150.535−0.165
GL0.9450.0820.926−0.014
ZB0.8940.1910.844−0.214
BB0.6580.1630.587−0.272
LIB0.4620.3820.267−0.590
NL0.7830.1370.7540.103
RB0.8740.0990.8010.015
BL0.368−0.6790.5030.184
DL0.8070.2750.776−0.169
MTR0.860−0.0210.7990.066
PB0.6880.2770.397−0.532
WP30.602−0.6170.5290.588
WP40.634−0.5350.4690.607
% total variance explained53.4611.2742.0711.56
Eigenvalue7.491.585.891.62
Fig. 4—

Distribution (minimum convex polygon) of scores along discriminant functions I (DFI) and II (DFII) of a discriminant function analysis of standardized means of morphometric characters for grouped populations (Appendix I) of a) males of Ochotona princeps (n = 1,167) and b) females of O. princeps (n = 939) assigned to subspecies based on genetic groupings (Fig. 1).

Compared to mean values of morphometric variables for O. princeps (Table 2), O. collaris is significantly smaller (P < 0.05) in 4 or 5 of 14 characters (TL, NL, RB, WP3, and WP4) and significantly larger in 6 others (ZB, BB, LIB, BL, DL, and PB), describing a substantial shape difference between the crania of the 2 species. In contrast, variation among the subspecies of O. princeps is principally in size: O. p. uinta, O. p. fenisex, and O. p. saxatilis are each significantly larger than the species' means for 13, 8 or 9, and 8 or 9 of 14 characters, respectively, O. p. schisticeps is significantly smaller than the means for 11 characters, and O. p. princeps differs only in being smaller for WP3.

View this table:
Table 2

Comparison of means of external and cranial measurements for male (upper of each pair) and female (lower) Ochotona collaris and subspecies of O. princeps with overall means of O. princeps. Values smaller than mean for O. princeps are in parentheses; significance (Tukey pairwise mean comparison) is indicated by * (P < 0.05) and ** (P < 0.01). Explanations for abbreviations of variables are provided in text.

TaxonnTLHFGLZBBBLIBNLRBBLDLMTRPBWP3WP4
0. collaris86(180.9)**31.343.5521.99**17.92**5.62**(13.45)**(6.95)*12.20**10.36**(8.54)2.51**(2.46)**(2.56)**
71(179.4)**30.542.6021.66**17.74**5.53**(13.11)**(6.76)**12.03**10.11**(8.50)2.51**(2.41)**(2.51)
0. princeps1,081187.030.743.5121.5017.605.3114.407.0811.939.988.582.212.622.70
870184.930.442.8621.2517.425.2614.216.9611.809.788.512.182.612.68
0. p. princeps381(186.6)31.0(43.49)(21.48)17.615.3514.547.10(11.92)10.06(8.55)(2.15)(2.59)**(2.69)
280(184.5)30.6(42.76)(21.19)(17.32)5.2714.276.9611.839.82(8.47)(2.11)(2.56)**(2.67)
0. p. fenisex249193.6**31.8**44.06**21.78**17.665.34(14.27)7.21**(11.85)(9.86)8.78**2.53**2.66*2.73
178193.4**31.9**43.56**21.59**17.65**5.28(14.10)7.09**(11.74)(9.73)8.70**2.50**2.66**2.70
0. p. saxatilis150(185.8)(30.6)44.40**21.89**17.755.49**15.01**7.25**(11.90)10.61**8.67*2.47**(2.61)2.70
147(180.8)**(30.0)*43.74**21.55**17.52**5.43**14.84**7.13**(11.66)*10.44**8.61*2.36**(2.60)2.67
0. p. schisticeps243(180.1)**(29.1)**(42.03)**(20.82)**(17.38)**(5.08)**(13.65)**(6.76)**11.94(9.40)**(8.27)**(1.74)**(2.60)(2.68)
206(180.4)**(29.0)**(41.44)**(20.67)**(17.23)**(5.08)**(13.52)**(6.64)**11.88(9.14)**(8.22)**(1.74)**(2.59)(2.67)
0. p. uinta58193.1**31.4*45.20**22.23**17.825.48*15.63**7.42**12.32**10.68**8.97**2.58**2.81**2.78**
59187.0**30.543.97**21.84**17.57*5.42**15.00**7.22**11.99**10.31**8.84**2.57**2.74**2.74**
Fig. 5—

Box plots of scores along discriminant function I (DFI) for 4 transects of grouped populations of males of Ochotona princeps across a) northern and b) southern British Columbia and Alberta, Canada (population numbers and I–VI as in Fig. 1). Sample sizes are shown in parentheses immediately above box plots. Central horizontal line marks the median, box includes the central 50% of values, whiskers indicate the range (* = outside values, ◯ = far outside values). Horizontal bars above box plots indicate group means not significantly different (P > 0.05) based on Tukey pairwise mean comparisons. Genetic assignment of grouped population is shown in parentheses below box plots: F = O. p. fenisex; P = O. p. princeps.

Vocalization dialects.—The dialects of the short call in Nearctic pikas thus far described are distinct in terms of fundamental frequency and duration (Table 3; Fig. 2), and the dialect in O. p. schisticeps is further distinguished by multiple notes. Short calls of O. p. fenisex and O. p. uinta have not been described in detail. Perhaps the most obvious cranial character distinguishing subspecies of O. princeps is the relative length of the bony palate (Fig. 3). It is smallest in O. p. schisticeps (i.e., mostly a soft palate), intermediate in O. p. princeps, and largest in O. p. fenisex, O.p. saxatilis, and O.p. uinta (no significant difference among means for these 3 subspecies). A significant (P = 0.002) and positive overall correlation exists between mean width of the palatal bridge and mean short call frequency for those grouped populations for which dialects have been described (Fig. 6). However, the correlation is not significant within subspecies with >2 samples (P = 0.068 in O. p. saxatilis and P = 0.235 and a negative correlation in O. p. princeps). Although we found an overall positive and significant (P = 0.011) correlation between width of the palatal bridge and body size (as indicated by greatest skull length), that correlation remains significant only within O. p. saxatilis (P = 0.025). Finally, an overall positive but nonsignificant correlation (P = 0.222) exists between call frequency and body size, which is reversed within O. p. princeps (a negative correlation; P = 0.016). The frequency of the short note may be functionally related to the length of the bony palate, with lower frequencies associated with softer palates (which perhaps function much like a loose reed in a reed instrument) and secondarily influenced by overall body size (again, as in the lower pitch of larger wind instruments). Of the 3 short calls described thus far (Conner 1982; Somers 1973), the fundamental frequencies range from 511 to 537 Hz for O. p. schisticeps, the smallest of the subspecies with the narrowest palatal bridge; 400 to 550 Hz for O. p. princeps, a larger subspecies with a palatal bridge of intermediate width; and 930 to 1,332 Hz for O. p. saxatilis, a larger subspecies with a wide palatal bridge (Table 2).

View this table:
Table 3

Mean values for fundamental frequency and duration of short calls, mean width of the palatal bridge (PB), and predicted fundamental frequency (in parentheses; based on PB) in regionally grouped populations of Ochotona collaris and subspecies of O. princeps. Populations are lettered as in Figs. 1, 2, and 6.

TaxonPopulationState or provinceFundamental frequency (Hz)Call duration (ms)Palatal bridge width (mm)
O. collarisAAlaskaa7131432.48
O. collarisBYukon Territorya8481082.54
O. p. princepsCAlbertaa7283112.01
O. p. princepsDWyomingb5403512.08
O. p. princepsEWyomingb4003732.26
O. p. princepsFWyomingb4863222.23
O. p. princepsGColoradob4703392.20
O. p. princepsHColoradob4703671.87
O. p. princepsIColoradob4533642.44
O. p. princepsJColoradob4573552.27
O. p. saxatilisKColoradob1,1121742.52
O. p. saxatilisLColoradob1,0961902.41
O. p. saxatilisMColoradob1,0692092.57
O. p. saxatilisNColoradob1,0141692.42
O. p. saxatilis0Coloradob9301962.19
O. p. saxatilisPNew Mexicob1,1741592.60
O. p. saxatilisQNew Mexicobc1,2361912.71
O. p. saxatilisRNew Mexicob9542122.15
O. p. saxatilisSUtahb8831432.58
O. p. schisticepsTCaliforniac5371301.80
O. p. schisticepsUUtahc5111501.61
British Columbia,(920)2.53
Washington,
O. p. fenisexOregon
O. p. uintaUtah(951)2.58
  • a Trefry and Hik, in press.

  • b Somers 1973.

  • c Conner 1982.

Fig. 6—

Relationship between mean values for width of palatal bridge (PB) and fundamental frequency of the short call for grouped populations of Ochotona collaris and subspecies of 0. princeps (Tables 2 and 3). PB is significantly correlated with frequency (r2 = 0.402, n = 21, P = 0.002) but not with call duration (r2 = 0.065,n = 21, P = 0.263). Dotted line indicates a linear relationship (y = 618.96x – 645.95). Black dots indicate predicted means of frequencies for O. p. fenisex and O.p. uinta, for which dialects have not been described.

In contrast to fundamental frequency, which appears to be directly related to structure of the palate, the other characteristics of the short call (call duration and number of notes) may be learned in pikas. Pikas call more frequently and for longer duration in response to predators than to nonpredators and delay calling and call less frequently in response to weasels than to martens (Ivins and Smith 1983), indicating contextual variation in employment of the short call.

Phenetic similarity among described dialects in O. collaris and subspecies of O. princeps is not concordant with perceived evolutionary relationships among these taxa. The short call dialects of O. collaris and O.p. saxatilis are quite similar, as are the predicted dialect frequencies of those taxa and the dialects of O. p. fenisex and O. p. uinta, whereas the dialect of O.p. princeps is most distinct on the basis of call duration. Hierarchical cluster analysis of call frequency and duration, using single linkage and Euclidian distance (not shown), linked the dialects of O. collaris and O.p. saxatilis in 1 cluster and O.p.princeps and O.p. schisticeps in the other cluster. In contrast, Galbreath et al. (in press) found that O. collaris and O. princeps diverged 1st and O. p. fenisex and perhaps O. p. schisticeps diverged next, followed by more recent and simultaneous divergence of O. p. princeps, O. p. saxatilis, and O. p. uinta. Trefry and Hik (in press) used a playback experiment to reject an acoustic adaptation hypothesis (i.e., that calls transmit with less degradation across their own species' habitat than the habitat of their congener) to explain differences between the short calls of O. collaris and O.p. princeps. Call characteristics may reflect either selection related to some other common environmental factor or simply idiosyncratic differences accumulated via genetic drift during lineage isolation.

Taxonomic conclusions.—Molecular phylogenetic analyses are concordant in recognizing 5 phylogeographic groups within O. princeps. In contrast, variation in cranial and external mensural characters appears to be quantitative, mostly due to variations in size, and (based on the principal component analysis) is idiosyncratic rather than related to phylogeographic groups. We conclude that molecular genetic characters more likely reflect deeper phylogenetic history, whereas other characters examined reflect more recent population-genetic processes. For example, initial variation in palatal breadth and dialect likely resulted from genetic drift; selection might have influenced pelage coloration (matching talus coloration) and perhaps body size (energetics); and variation in other cranial characters might have resulted primarily from inbreeding and subsequent drift. To best reflect evolutionary lineages in the trinomial we therefore synonymize a majority of the 36 recognized subspecies of O. princeps under the 5 major phylogeographic lineages.

Ochotona princeps (Richardson, 1828) American Pika (Synonymy under subspecies)

Geographic range.—Patchily distributed in cool, rocky habitat (usually in montane situations, but occasionally in lower-elevation ecosystems) from Coast Mountains and Rocky Mountains of central British Columbia and Alberta from 54°N latitude south through the Cascade Range, Sierra Nevada, Wasatch Range, Uinta Mountains, and southern Rocky Mountains to 36°N latitude, and isolated mountains neighboring those major ranges and throughout the Great Basin regional desert. Elevational range is from near sea level (along the Pacific coast in British Columbia and along the Columbia River) to the highest peaks (>4,000 m) of the western United States and Canada. The upper elevational limit of pikas has not been determined. Hafner (1993) described both latitudinal and longitudinal gradients in current elevational lower limits (increasing to both the south and east), and gross environmental limits, the most critical of which appears to be short summers (<20–30 days/year above 35°C), in agreement with environmental limits described by Grinnell (1917) and Smith (1974). Near its southern geographic limits in the Sierra Nevada the species rarely occurs below 2,500 m (Smith and Weston 1990; but see Millar and Westfall, in press), and near its southeastern geographic limit in basalt talus of the Jemez Mountains of New Mexico (population 128; Fig. 1) populations persist at ≥3,000 m. Basalt talus, particularly in massive lava flows, can provide lower-elevation refugia of cool habitat due to its thermal insulating qualities. For example, the lower limits of the Jemez populations are 500 m lower than those in the adjacent Sangre de Cristo Mountains (population 129), and populations in lava flows of Lava Beds National Monument (northern California; population 85) and Craters of the Moon National Monument (southern Idaho; population 68) occur 120 m and 570 m lower, respectively, than the predicted elevation based on latitude and longitude (Hafner 1993).

Description.—Similar to O. collaris, its closest sister taxon (Lissovsky et al. 2007; Niu et al. 2004) in a Holarctic clade (Northern Group of Yu et al. [2000]) that also includes O. hyperborea, O. alpina, and O.pallasi, each of which may represent separate species groups (Lissovsky et al. 2007). Smith and Weston (1990) described the general characteristics of the species, which differs from O. collaris in having buffy (versus white) underparts; lacking the grayish collar found on the shoulders of O. collaris; and having a narrower skull with a longer, broader nose, shorter auditory bullae, and larger upper premolars (Smith and Weston 1990; Weston 1981,1982; Table 2).

Ochotona princeps princeps (Richardson, 1828)

Lepus (Lagomys)princeps Richardson, 1828:520. Type locality “Rocky mountains.”Further restricted by Preble (1908:198) to “near the sources of Elk (Athabaska) River,”Alberta, Canada.

Ochotona princeps: Trouessart, 1897:648. First use of current name combination.

Ochotona cuppes Bangs, 1899:40. Type locality “Monishee Divide, Gold Range, B[ritish]. C[olumbia, Canada]., altitude 4000 feet.”

Ochotona levis Hollister, 1912:57. Type locality “Chief Mountain [= Waterton] Lake, [Glacier Co.,] Montana.”Ochotona figginsi Allen, 1912:103. Type locality “Pagoda

Peak, Routt County [= Rio Blanco Co.], Colorado.”Ochotona princeps lutescens Howell, 1919:105. Type locality

“Mt. Inglesmaldie, near Banff, Alberta.”[Ochotona princeps]princeps: Howell, 1919:105. First use of current name combination.

Ochotona uinta lemhi Howell, 1919:106. Type locality “Lemhi Mountains, 10 miles west of Junction, [Lemhi Co.,] Idaho.”

Ochotona uinta ventorum Howell, 1919:106. Type locality “Fremont Peak, Wind River Mountains, [Fremont Co.,] Wyoming.”

Ochotona uinta nevadensis Howell, 1919:107. Type locality “Ruby Mountains, southwest of Ruby Valley P.O., [Elko Co.,] Nevada, altitude 10,500 feet.”

Ochotona princeps levis: Howell, 1924:16.

Ochotona princeps lemhi: Howell, 1924:16.

Ochotona princeps ventorum: Howell, 1924:18.

Ochotona princeps nevadensis: Howell, 1924:21.

Ochotona princeps figginsi: Howell, 1924:21.

Ochotona princeps saxatilis: Howell, 1924:23. Part (only populations north of the Colorado River); not Ochotona saxatilis Bangs, 1899.

Ochotona princeps cuppes: Howell, 1924:27.

Ochotona schisticeps goldmani Howell, 1924:40. Type locality “Echo Crater, Snake River Desert, [Butte Co.,] Idaho (20 miles southwest of Arco).”

Ochotona princeps howelli Borell, 1931:306. Type locality “summit of Smith Mountain, Adams Co., Idaho, altitude 7500 feet (near head of Bear Creek, south end of Seven Devils Mountains).”

Ochotona princeps clamosa Hall and Bowlus, 1938:335. Type locality “N rim Copenhagen Basin, 8400 ft., Bear Lake County, Idaho.”

Ochotona princeps goldmani: Hall and Bowlus, 1938:337.

Ochotona princeps saturatus Cowan, 1954:23. Type locality “Mount Huntley in Wells Gray Park, B[ritish] Columbia].,”Canada; incorrect gender concordance.

Ochotona princeps wasatchensis Durrant and Lee, 1955:2. Type locality “10 miles above lower powerhouse, road to Cardiff Mine, Big Cottonwood Canyon, Salt Lake County, Utah.”

Ochotona princeps obscur a Long, 1965:538. Type locality “Medicine Wheel Ranch, 28 miles east of Lovell, 9000 ft., Big Horn County, Wyoming.”

O[chotona].p[rinceps].saturata: Hall and Kelson, 1959:248.

Geographic range.—Patchily distributed in cool, rocky habitat (usually in montane situations, but occasionally in lower-elevation ecosystems such as lava beds) from the northern Rocky Mountains of central British Columbia and Alberta through Idaho and Montana, with geographically isolated populations in the Little Belt and Belt mountains of Montana, the Big Horn Mountains of Wyoming, the Ruby Mountains of Nevada, the Wasatch Range of Idaho and Utah, and the Park Range and Front Range of Colorado north of the Colorado River.

Description.—Size and cranial features average for the species, except with a significantly narrower P3; pelage color variable. Broad regional introgression with neighboring subspecies (O. p. fenisex in British Columbia and O. p. saxatilis in Colorado) is indicated by clinal variation in cranial morphology, discordance among morphological and various genetic data sets, or both. Short call (described for populations in Alberta, Wyoming, and Colorado—Somers 1973; Trefry and Hik, in press) characterized by a single note of relatively low fundamental frequency (400–728 Hz) and long duration (0.276-0.407 s).

Ochotona princeps fenisex Osgood, 1913

Lagomys minimus Lord, 1863:98. Type locality “Ptarmigan Hill.”Further restricted by Howell (1924:28) to “near head of Ashnola River, Cascade Range, British Columbia,” Canada; preoccupied by Lagomys minimus Schinz, 1821.

Ochotona fenisex Osgood, 1913:80. Replacement name for Lagomys minimus Lord, 1863, preoccupied.

Ochotona fenisex brunnescens Howell, 1919:108. Type locality “Keechelus, [Kittitas Co.,] Washington.” Ochotona fenisex fumosa Howell, 1919:109. Type locality “Permilia Lake, west base Mt. Jefferson, [Marion Co.,] Oregon.”

Ochotona princeps fenisex: Howell, 1924:28. First use of current name combination.

Ochotona princeps brooksi Howell, 1924:30. Type locality “Sicamous, British Columbia,”Canada.

Ochotona princeps brunnescens: Howell, 1924:31.

Ochotona princeps fumosa: Howell, 1924:33.

Ochotona princeps septentrionalis Cowan and Racey, 1947:102. Type locality “Itcha Mountains, British Columbia, altitude 6500 feet,”Canada.

Ochotona princeps littoralis Cowan, 1954:22. Type locality “Hagensborg, B[ritish]. C[olumbia].,”Canada.

Geographic range.—Patchily distributed in cool, rocky habitat (usually in montane situations, but occasionally in lower-elevation ecosystems such as along the Columbia River) from the Coast Mountains and Cascade Range from central British Columbia to southern Oregon.

Description.—Size and cranial features large for the species; pelage color variable. Broad regional introgression with neighboring O. p. princeps in British Columbia is indicated by clinal variation in cranial morphology and discordance among morphological and various genetic data sets, and past introgression (probably during LGM contact) with O.p. schisticeps is indicated by allozyme data in the population at Mt. McLaughlin (population 37; Fig. 1) in southern Oregon (Hafner and Sullivan 1995). Short call said to be more similar to that of O. collaris than to that of O. p. princeps from Wyoming (Broadbooks 1965), but it has not been described in detail.

Ochotona princeps saxatilis Bangs, 1899

O. saxatilis Bangs, 1899:41. Type locality “Snowy Range, Montgomery, Park Co., Colorado.”Part; only populations of this taxon south of the Colorado River.

O. nigrescens Bailey, 1913:133. Type locality “Jemez Mts., [Sandoval Co.,] New Mexico (alt. 10,000 feet)”; further restricted by Bailey (1931:67) to “Jemez Mountains, [Sandoval Co.], N[ew]. Mexfico]., at 10,000 feet altitude on Goat Peak at the head of Santa Clara Creek.”

O. saxatilis incana Howell, 1919:107. Type locality “Pecos Baldy, [Santa Fe Co.], New Mexico, altitude 12,000 feet.”

O. princeps saxatilis: Howell, 1924:23. First use of current name combination.

O. princeps incana: Howell, 1924:25.

O. princeps nigrescens: Howell, 1924:26.

O. princeps lasalensis Durrant and Lee 1955:4. Type locality “Warner Ranger Station, 9,750 feet, La Sal Mountains, Grand County, Utah.”

Geographic range.—Patchily distributed in cool, rocky habitat (usually in montane situations) in the southern Rocky Mountains south of the Colorado River (Front Range, San Juan Mountains, and Sangre de Cristo Range) and nearby isolated highlands including the La Sal Mountains of southwestern Utah, Grand Mesa of Colorado, and Jemez Mountains of New Mexico.

Description.—Size and cranial features large for the species; pelage color variable. Broad regional introgression with neighboring O. p. princeps in Colorado is indicated by discordance among morphological and various genetic data sets (Galbreath et al., in press). Short call (Conner 1982; Somers 1973) characterized by a single note (occasionally 2 or 3) of relatively high fundamental frequency (883–1,332 Hz) and short duration (0.143–0.212 s).

Ochotona princeps schisticeps (Merriam, 1889)

Lagomys schisticeps Merriam, 1889:11. Type locality “Donner [= Summit, Placer Co.], California.”

Ochotona schisticeps: Merriam, 1899:99.

Ochotona cinnamomea Allen, 1905:121. Type locality “Briggs [= Britt's] Meadows, (alt. 11,000 ft.). Beaver Range, Beaver County, Utah” 5 miles by road W Puffer Lake, according to Hardy (1945).

Ochotona albatus Grinnell, 1912:125. Type locality “near Cottonwood Lakes, 11,000 feet, Sierra Nevada Mountains, Inyo County, California”; incorrect gender concordance.

Ochotona taylori Grinnell, 1912:129. Type locality “Warren Peak, 9000 ft. alt., Warner Mts., Modoc Co., Calif[ornia].”

Ochotona schisticeps muiri Grinnell and Storer, 1916:6. Type locality “9300 feet altitude near Ten Lakes, Yosemite National Park, Tuolumne County, California.”

Ochotona schisticeps albatus: Grinnell and Storer, 1916:6. Incorrect gender concordance.

Ochotona schisticeps sheltoni Grinnell, 1918:429. Type locality “11,000 feet altitude, near Big Prospector Meadow, White Mountains, Mono County, California.”

Ochotona schisticeps jewetti Howell, 1919:109. Type locality “head of Pine Creek, near Cornucopia, south slope Wallowa Mts., Baker County, Oregon.”

Ochotona schisticeps fuscipes Howell, 1919:110. Type locality “Brian Head, Parawan Mts., [Iron Co.], Utah.”

Ochotona schisticeps taylori: Howell, 1924:39.

Ochotona schisticeps albata: Howell, 1924:44.

Ochotona schisticeps cinnamomea: Howell, 1924:46.

Ochotona princeps tutelata Hall, 1934:103. Type locality “Monitor Mountains, Greenmonster Canyon, 8150 feet, Nye County, Nevada.”

Ochotona princeps muiri: Hall, 1934:103.

Ochotona princeps cinnamomea: Hall, 1934:103.

Ochotona princeps schisticeps: Miller, 1936:175. First use of current name combination.

O[chotona].p[rinceps]. fuscipes: Hall and Hayward, 1941:108.

Ochotona princeps sheltoni: Hall, 1946:592.

Ochotona princeps albata: , 1951:127.

Ochotona princeps jewetti: 1951:130.

Ochotona princeps taylori: Hall, 1951:133.

Geographie range.—Patchily distributed in cool, rocky habitat (usually in montane situations, but occasionally in lower-elevation ecosystems) of the southern Cascades, Klamath Mountains, and Warner Mountains of California, throughout the Sierra Nevada of California and Nevada, and isolated highlands throughout the Great Basin of Nevada, eastern Oregon (north to the Blue Mountains), and southwestern Utah.

Description.—Size and cranial features small for the species in virtually all characters except bullar length, which is average for the species (and so relatively large in this subspecies); pelage color variable. Past introgression (probably during LGM contact) with O.p. fenisex and O.p. uinta is indicated by allozyme data in the population of O. p. fenisex at Mt. McLaughlin in southern Oregon and the population of O. p. uinta on the Aquarius Plateau of southern Utah (Hafner and Sullivan 1995). Short call (at least for single populations in California and Utah—Conner 1982) usually (87%) characterized by multiple notes (up to 4) of relatively low fundamental frequency (511–537 Hz) and short duration (0.13–0.15 s).

Ochotona princeps uinta Hollister, 1912

Ochotona uinta Hollister, 1912:58. Type locality “Uintah Mountains, [Summit Co.], Utah.”Further restricted by Howell (1924:19) to “Uinta Mountains, near head of east fork of Bear River, [Summit Co.], Utah.”

Ochotona princeps uinta: Howell, 1924:19. First use of current name combination.

Ochotona princeps utahensis Hall and Hay ward, 1941:107. Type locality “2 miles west of Deer Lake, Garfield County, Utah.”

Ochotona princeps moorei Gardner, 1950:344. Type locality “1 mile northeast of Baldy Ranger Station, Manti National Forest, altitude 10,000 feet, Sanpete County, Utah.”

Ochotona princeps barnesi Durrant and Lee, 1955:6. Type locality “Johnson's Reservoir, 8,800 feet, 15 miles north of Loa (Fishlake Plateau), Sevier County, Utah.”

Geographic range.—Patchily distributed in cool, rocky habitat (usually in montane situations) in the Uinta Mountains and Wasatch Range of central Utah.

Description.—Size and cranial features large for the species; pelage color variable. Past introgression (probably during LGM contact) with O. p. schisticeps is indicated by allozyme data in the population on the Aquarius Plateau of southern Utah (Hafner and Sullivan 1995). Short call has not been described.

Acknowledgments

We thank curators and collections managers at the following institutions for making accessible museum specimens in their care: Carnegie Museum of Natural History; University of Kansas, Museum of Natural History; University of Utah, Utah Museum of Natural History; Brigham Young University, Monte L. Bean Life Science Museum; California Academy of Sciences; Humboldt State University; Harvard University, Museum of Comparative Zoology; University of California, Berkeley, Museum of Vertebrate Zoology; Provincial Museum of Alberta; Portland State University, Collection of Vertebrates; Royal Ontario Museum; Texas Tech University, Museum of Texas Tech University; University of British Columbia, Cowan Vertebrate Museum; University of Puget Sound, James R. Slater Museum of Natural History; American Museum of Natural History; Royal British Columbia Museum; Washington State University, Charles R. Conner Museum; Denver Museum of Nature and Science; The Field Museum; Natural History Museum of Los Angeles County; Canada Museum of Nature; University of Alberta, Museum of Zoology; University of California, Los Angeles, Dickey Collection; University of Idaho, Bird and Mammal Museum; United States National Museum of Natural History; University of Washington, Thomas Burke Memorial Washington State Museum; New Mexico Museum of Natural History; Oregon State University, Museum of Natural History; Oregon State University, Department of Fisheries and Wildlife Mammal Collection; San Diego Natural History Museum; and San Diego Society of Natural History. This research was supported by National Science Foundation grant BSR-8615003 to DJH.

Appendix I

Specimens examined

A total of 2,163 specimens from the following collections in the United States and Canada (abbreviations as in Hafner et al. [1997]) was examined during this study: American Museum of Natural History (AMNH); Brigham Young University, Monte L. Bean Life Science Museum (BYU); California Academy of Sciences (CAS); Carnegie Museum of Natural History (CM); Canada Museum of Nature (CMN); Washington State University, Charles R. Conner Museum (CRCM); Denver Museum of Nature and Science (DMNH); The Field Museum (FMNH); Humboldt State University (HSU); University of Kansas, Museum of Natural History (KU); Natural History Museum of Los Angeles County (LACM); Harvard University, Museum of Comparative Zoology (MCZ); University of New Mexico, Museum of Southwestern Biology (MSB); University of California, Berkeley, Museum of Vertebrate Zoology (MVZ); New Mexico Museum of Natural History (NMMNH); Oregon State University, Museum of Natural History (OSMNH); Oregon State University, Department of Fisheries and Wildlife Mammal Collection (OSUFW); Provincial Museum of Alberta (PMA); University of Puget Sound, James R. Slater Museum of Natural History (PSM); Portland State University, Collection of Vertebrates (PSU); Royal British Columbia Museum (RBCM; formerly BCPM); Royal Ontario Museum (ROM); San Diego Natural History Museum (SDNHM); San Diego Society of Natural History (SDSNH); Texas Tech University, Museum of Texas Tech University (TTU); University of Alberta, Museum of Zoology (UAMZ); University of British Columbia, Cowan Vertebrate Museum (UBC); University of California, Los Angeles, Dickey Collection (UCLA); University of Idaho, Bird and Mammal Museum (UIDA); University of Utah, Utah Museum of Natural History (UMNH); United States National Museum of Natural History (USNM); and University of Washington, Thomas Burke Memorial Washington State Museum (UWBM; formerly BMWSM). Collection acronyms and specimen catalog numbers are listed within group localities (numbered as in Fig. 1) for each taxon; boldface type indicates group locality number, with sample sizes in parentheses (male, female, and undetermined sex if a 3rd number is provided), and a general identifier of locality.

Ochotona collaris (n = 164; 86, 71, 7)

1 (86, 71, 7) ALASKA (HSU 4950–4953; MVZ 37593, 57345, 57346; PSM 5078, 5079; FMNH 24249; CMN 30650–30655, 40302–40305, 40307–40315, 40317, 40318, 50386–50390; USNM 36297, 36298, 131258, 131261, 131265, 131266, 131268, 131270, 131275, 131276, 131280, 131285, 131288, 131298, 131304, 131314, 131322, 131323, 131326, 131328, 131329, 131331, 131336, 131340, 131341, 131343, 131890, 148589–148593, 157225–157230, 175128, 241741– 241743, 244057, 271691–271695, 512800, 512802; NMMNH 1307, 1313, 1316–1318, 1321–1323, 1326, 1328, 1329, 1332, 1337, 1340, 1343, 1345). CANADA: British Columbia (KU 29090, 29095, 29099; BCPM 6085, 6089–6092, 6094); Northwest Territories (AMNH 127808, 127810; CMN 30302–30304, 46541; UAMZ 8498); Yukon (ROM 22978; CMN 17328, 17802, 17803, 17824, 17828, 17829, 17831, 18120, 29410, 29414, 29416, 30647, 30648, 31161, 31163, 31165–31167, 31170, 31171, 31174, 31175, 31201–31203, 35316, 35319, 35324, 35329–35331, 44998, 45000, 45002, 45003, 45005, 45007, 45009, 45012, 45013; UAMZ 6704, 6705, 6762; USNM 134936, 134939).

Ochotona princeps fenisex (n = 439; 251, 178, 10)

CANADA: British Columbia; 2 (3, 2) Kimsquit (CMN 16544, 16562, 16738, 16751, 16761). 3 (2, 1)Hagensborg (CMN 15823, 15825, 15826). 4 (23, 10) Stuie (CMN 15623, 15633, 15634, 15707, 15708, 15741, 15743, 15850, 15862, 15864–15868, 15871, 15950–15952, 16658, 16667, 16668, 16673, 16682; NMMNH 978, 979, 983, 984, 986, 988, 989, 991, 995, 998). 5 (6,1) Itcha Mountains (CMN 5906; NMMNH 999, 1000, 1003, 1005, 1008, 1009. 6 (3,1) Kleena Kleene (LACM 9914, 9915; CMN 28755, 28756). 7 (8, 5) Hanceville (MVZ 53704, 53705; BCPM 5332; CMN 22881, 22882, 28753, 28757–28760, 28767, 28774, 28776). 8 (6, 0) Fawn Bluff (CMN 14230, 14434, 14442, 14443, 14445, 14605). 9 (9, 3) Lillooet (BCPM 859, 860, 862, 2408, 2409, 2411–2415; CMN 2728). 10 (23,12,1) Alta Lake (MVZ 54535; UBC 334, 339, 441, 478, 504, 1084, 1797–1799, 3619, 5911, 5912, 5914–5916, 5919–5924, 5926–5932, 5934, 5935; BCPM 1555–1558; CMN 28773). 11 (15, 8, 2) Sicamous (BCPM 851, 852, 2090–2092, 2094, 2095, 2867–2869; CMN 155, 156; USNM 67129–67133, 67136, 67148, 69272, 69273, 69275, 69278, 69279, 69281. 12 (6, 6) Westwold (NMMNH 933, 935–939, 941, 945–947, 951, 952). 13 (9, 12, 1) Okanagan (MVZ 77493; PSM 5620–5624, 5626; BCPM 831, 832, 834, 836, 840–842, 849, 850, 853; USNM 94303, 94304, 203215, 230778, 233625). 14 (0, 2, 1) Tulomeen (ROM 29065, 29067; BCPM 846). 15 (3, 2) Manning Park (BCPM 5179–5181, 5484, 5486). 16 (8, 7) Hedley (and adjacent WASHINGTON: Okanogan Co.) (MVZ 113825; CRCM 47165, 53174, 76331, 81583; LACM 10532; CMN 8939; USNM 235186–235189, 235220, 235222, 235223; SDSNH 16893).

WASHINGTON: Whatcom Co.; 17 (8, 9) Mt. Baker (and adjacent CANADA: British Columbia) (MVZ 134812, 134814; CRCM 649, 650, 652; CMN 7867; USNM 88704, 88705, 234784, 234950; BMWSM 30824, 30939; NMMNH 1011, 1015, 1016, 1020, 1022). Ferry Co.; 18 (2, 1) Sherman Creek Pass (MVZ 84513, 84514, 134794). Okanogan Co.; 19 (5, 8, 1) Glacier (and adjacent Chelan Co.) (CRCM 651, 8143, 8144; USNM 230094–230096; NMMNH 1027, 1028, 1033, 1035–1037, 1040, 1048). King Co.; 20 (11, 8) Tye (and adjacent Chelan Co. and Snohomish Co.) (KU 22400, 22404; MVZ 134777–134787, 134789, 134790; USNM 230091, 234215; BMWSM 9868, 12224). Kittatas Co.; 21 (4, 2) Keechelus (and adjacent King Co.) (MVZ 54536, 134791; UCLA 8637; PSM 11035; USNM 227258, 227259). Chelan Co.; 22 (14, 13) Wenatchee (and adjacent Kittatas Co.) (KU 22401, 22403; MVZ 134809–134811, 134815, 134816; PSM 2147, 8724; CRCM 81383; USNM 41538, 41539, 41541, 41542, 41544, 90539, 90541, 90542, 90544, 90545, 91129–91132, 91134, 273575; BMWSM 18373). Pierce Co.; 23 (15, 13, 2) Mt. Rainier (and adjacent Lewis Co. and Yakima Co.) (PSM 610–612, 4186, 4187, 10321; LACM 10662; UCLA 9617; USNM 89840, 89841, 89843, 232717, 233154, 233164, 233166, 233167, 233169–233171; BMWSM 15246, 30272, 30274, 30415, 30811; SDSNH 16881, 16883, 16884, 16886–16888). Skamania Co.; 24 (10, 7) Mt. St. Helens (and adjacent Clark Co. and Cowlitz Co.) (MVZ 94671–94674, 94676–94680; PSM 11965, 11966; USNM 90760–90762, 90765, 90766, 274449). 25 (1, 0) Sawtooth Mtn. (MVZ 87864). Yakima Co.; 26 (2, 1, 1) Mt. Adams (USNM 226756, 226757, 274005, 274006).

OREGON: Multnomah Co.; 27 (2, 6, 2) Mt. Hood (and adjacent Clackamas Co.) (HSU 4356, 4360; PSU 853, 854, 1258, 1657, 2826, 2827; PSM 10362; USNM 80081). Linn Co.; 28 (1, 0) Sweet Home (OSMNH 2680). 29 (0, 1) Carpenter Mtn. (PSM 16647). 30 (7, 3) Mt. Jefferson (and adjacent Jefferson Co.) (MVZ 96086; USNM 91136, 91138, 91140–91144; SDSNH 16878, 16879). Lane Co.; 31 (19, 19) McKenzie Bridge (PSU 909, 910, 1324–1328, 1331, 1388, 1392; PSM 11967–11969, 14503, 16646, 19122–19124; USNM 204866, 204867, 204869, 272367, 272368; NMMNH 862–864, 866, 867, 869; OSMNH 3628, 6120, 6125; OSUFW 3633, 3634, 6131, 7099, 7683, 7910). 32 (1, 0) Waldo Lake (PSM 11970). Deschutes Co.; 33 (8, 3) Bend (MVZ 84067–84073; OSMNH 6115; OSUFW 6126, 6127; SDSNH 16877). 34 (1, 0) Elk Lake (OSMNH 1340). 35 (2,2) Paulina Lake (USNM 204871; SDSNH 16872, 16873, 16876). Klamath Co.; 36 (13, 8) Crater Lake (MVZ 31075, 79631, 79634–79636, 109102; UCLA 8098, 8099, 8109, 8117; USNM 79811, 80071, 80074, 80076, 80077, 80080, 80586–80588, 88073, 274862). Jackson Co.; 37 (1, 2) Mt. McLoughlin (NMMNH 1211, 1212, 1214).

Ochotona princeps princeps (n = 680; 382, 282, 16)

CANADA: British Columbia; 38 (3, 3) Indianpoint Lake (MVZ 42613, 42614, 44716, 44717, 44719; BCPM 1200). 39 (5, 8) Raft Mountain (NMMNH 955, 958, 960–962, 966–968, 970, 974–977). 40 (20, 8) Monashee Mtns. (MCZ 3603, 7389; MVZ 65185, 77497; ROM 29069, 29071, 29074; BCPM 2096, 2098, 2100, 2101, 2105, 2106, 2108, 2109, 2111–2116; CMN 12775, 30848; UCLA 17230; USNM 68800, 68801, 249078, 290746). 41 (5, 2) Golden (NMMNH 917, 920, 923, 925, 926, 928, 929). 42 (1, 2, 2) Assiniboine (AMNH 141786, 141790, 141792–141794). 43 (3, 0) Cranbrook (BCPM 854; CMN 28766, 28768). 44 (20, 9) Nelson Range (and adjacent IDAHO: Boundary Co., MONTANA: Lincoln Co., and WASHINGTON: Pend Oreille Co.) (MVZ 134477–134482; PSM 3778, 4191; CRCM 40114, 59194, 80736, 81393, 81576; CMN 951, 10012, 10128; USNM 66677–66683, 68798, 68799, 91191, 91194, 236452, 272359). Alberta; 45 (1, 2) Burns Mtn. (UAMZ 5397, 5399, 5400). 46 (25,22,1) Henry House (CM 22905, 22907, 22913, 22914; MVZ 54532; PMA 749318, 824723; AMNH 122661, 122662; CMN 10780, 10781, 10783, 10798, 10813, 10814, 10816, 10817, 16854, 18770, 23786–23790; USNM 75457–75461, 75465, 75467–75469, 81529, 81530, 81532, 81534, 81537–81542, 81548, 81555, 81559, 81560, 174392). 47 (3, 7, 1) Sunwapta Pass (CMN 18736, 18744; UAMZ 3598, 3601–3606, 4232, 4233). 48 (1, 0) Ptarmigan Lake (AMNH 35146). 49 (5, 2) Egypt Lake (AMNH 141773–141775, 141778, 141785; CMN 10934, 18697). 50 (21, 11, 1) Banff (MCZ 10767, 10768; AMNH 95189, 95190; FMNH 5852, 6991, 6992, 8589, 8592; CMN 10857, 10870, 10882, 16030, 16182, 26516, 26518, 41137; USNM 68803, 68810, 108650, 108651; NMMNH 894–897, 899, 905, 907, 909, 911, 914–916). 51 (5,11,2) Coleman (PMA 84181, 86301; ROM 22382, 22383, 22390, 22391, 22407, 22411–22415; AMNH 125343, 125344, 125346; UAMZ 1479, 1481, 5401).

MONTANA: Glacier Co.; 52 (10, 2) Waterton Lake (and adjacent CANADA: Alberta and British Columbia) (PMA 263100, 263101, 826398, 826399; CMN 4650; UAMZ 575; USNM 22241, 72768–72770, 72772, 72777). Powell Co.; 53 (10, 4) Ovando (NMMNH 602, 603, 606–610, 612, 613, 615, 616, 618–620). Ravalli Co.; 54 (4, 2) Darby (KU 34063; USNM 169153, 169158; NMMNH 596–598). Unspecified county; 55 (0, 1) Belt Mtns. (USNM 189001). Cascade Co.; 56 (6, 6) Neihart (USNM 232759, 232761; NMMNH 573, 579, 581–584, 587–589, 592). Beaverhead Co.; 57 (5, 1) Dillon (CM 22145, 22146, 22372; MVZ 107162–107164). Park Co.; 58 (4, 1) Emigrant (MCZ 7780; USNM 225887, 225888, 253997, 253998). Carbon Co.; 59 (10, 4) Red Lodge (and adjacent Park Co.) (KU 37991; MVZ 107166–107169; USNM 66763, 169745, 169746, 169785; NMMNH 562–564, 568, 569).

IDAHO: Shoshone Co.; 60 (14, 9,1) Mullan (MVZ 54856, 54857, 54859, 78062, 78064, 78069–78072, 78074–78078, 78080, 78081, 78083; UIDA 237602, 40634, 40637, 40638, 74011, 74012, 74014). Adams Co.; 61 (17, 12) Black Lake (KU 45687–45690, 45692, 45695–45698, 45701–45705, 45709; CRCM 78409, 78450–78452; USNM 74786; NMMNH 622, 625, 626, 629, 631, 632, 634, 636, 638, 641). Idaho Co.; 62 (1, 0, 4) Buffalo Hump (MVZ 88853; FMNH 90611–90613; UIDA 1489). 63 (0, 3) Salmon (ROM 31356, 77615; AMNH 176526). Valley Co.; 64 (6, 7) Salmon River Mtns. (UIDA 1493; USNM 30904, 30961, 30963, 31057, 31058, 31204, 31205, 31209, 31210, 31212, 31214, 507437). Custer Co.; 65 (22, 11) Stanley (and adjacent Blaine Co., Boise Co., and Valley Co.) (KU 18739; MVZ 54864–54872, 54876, 54877, 72074–72077, 72079–72082, 72085, 72086; AMNH 38272, 38274; CRCM 79604; USNM 265907; NMMNH 644, 646, 648, 649, 654, 660; OSMNH 3629). 66 (3, 4) Lost River Range (MVZ 72065, 72067, 72071, 72072; USNM 30826, 30902, 30903). Blaine Co.; 67 (3, 4) Ketchum (KU 27419; MCZ 8501, 8502; PSM 3923, 3924; USNM 31455, 31785). Butte Co.; 68 (4, 8) Craters of the Moon (MVZ 78084–78088; USNM 236406–236408, 236410; NMMNH 666, 668, 669). Bear Lake Co.; 69 (12, 10, 1) Copenhagen Basin (and adjacent Franklin Co. and UTAH: Rich Co.) (KU 132510, 132601; UMNH 21160–21662; MVZ 78090, 78092, 78094–78100, 78103, 78107; USNM 543119; NMMNH 672, 674, 675, 678, 679, 681).

NEVADA: Elko Co.; 70 (17, 23) Ruby Mtns. (KU 45710–45712, 45715–45721, 45723; MVZ 120742, 120743, 120745–120748, 120751; USNM 94212, 94213; NMMNH 256, 260, 262–264, 266, 270, 272, 273, 275, 276, 278, 280).

UTAH: Salt Lake Co.; 71 (8, 7) Alta (UMNH 4787, 4789, 4790, 8254, 9967, 10638, 13580, 13582, 13584; MVZ 116732, 116733; NMMNH 299, 302, 307, 309).

WYOMING: Big Horn Co.; 72 (12, 10, 1) Big Horn Mtns. (KU 20878, 20880–20882, 32914, 32915, 32917–32919; MCZ 32706; USNM 168940, 168942; NMMNH 538–542, 544–546, 548, 554, 560). Teton Co.; 73 (33, 20) Teton Range (and adjacent Fremont Co. and Sublette Co.) (KU 15862, 15865, 15866, 15871, 15877, 15881, 15884, 15886–15888, 37994; MSB 705; MVZ 89241–89244, 89246; AMNH 138120; USNM 169920, 170283, 170298–170302, 170383–170386, 176776, 176778–176780, 176839, 190230, 298486; NMMNH 502, 505, 506, 508, 513–517, 521, 522, 524, 528, 532, 534, 535, 537). Lincoln Co.; 74 (13, 11) Salt River Range (and adjacent Sublette Co.) (KU 15891, 15893, 15896, 15898, 15906, 15919, 132511, 132512, 132515–132522; USNM 55174, 55176, 55177, 176920, 176974, 176975, 177098, 177100). Carbon Co.; 75 (18, 11) Encampment (KU 26600, 26601, 26603, 45708, 91037, 132513, 132514, 132524–132526; USNM 176456; NMMNH 476–478, 480–483, 485–488, 490, 492, 493, 495, 497^99). Albany Co.; 76 (16, 3) Centennial (KU 17594, 17597, 21865, 132528, 132529; MVZ 89247, 89249, 89250, 89252, 89254, 89255; USNM 176457; NMMNH 455, 459–462, 469, 470).

COLORADO: Jackson Co.; 77 (4, 1) Mt. Zirkel (DMNH, 1075, 1077–1080). Garfield Co.; 78 (12,20,2) Trappers Lake (CM 20195, 20196, 20604; KU 20886–20888, 20893, 70017; MVZ 81979; AMNH 32721, 124389, 124391; DMNH 337, 5798; NMMNH 405–409, 413, 418, 419, 421–427, 429–432, 434).

Ochotona princeps schisticeps (n = 460; 244, 206, 10)

OREGON: Wallowa Co.; 79 (11, 12) Wallowa Mtns. (MCZ 32705; MVZ 84046, 84047, 84052–84054, 84057–84063, 84066; USNM 90768, 90771, 90773, 208352, 208354; OSUFW 6128; SDNHM 16904–16906). Baker Co.; 80 (15,10) Blue Mtns. (AMNH 40547–40549, 40551; CRCM 81585, 81586; USNM 79197, 79198, 207676, 208343–208345, 208348, 209577, 209578, 209581, 209582, 209584; NMMNH 1051–1053, 1059, 1061; SDNHM 16903; SDSNH 16900). Lake Co.; 81 (1, 0) Cougar Peak (USNM 246058). 825,1, 1) Hart Mtn. (PSM 7956; USNM 205238; SDNHM 16862, 16864, 16866, 16869, 16871). Harney Co.; 83 (6, 6, 2) Steens Mtn. (USNM 215998, 216001, 216003, 216005–216008; NMMNH 871, 875–878; SDNHM 16867, 16868).

CALIFORNIA: Siskiyou Co.; 84 (1, 2) Goose Nest Mtn. (MVZ 27927; USNM 97910, 97912). 85 (4, 0) Lava Beds National Monument (MVZ 68985–68987, 70497). 86 (6, 7) Mt. Shasta (MVZ 3327, 65435, 86882; USNM 95098, 95542–95544; NMMNH 852, 854–856, 859, 860). Modoc Co.; 87 (9, 4) Warner Mtns. (CAS 5069; MVZ 11292, 36908, 36912–36914, 36917, 36920, 36924; LACM 4404, 4409; NMMNH 847, 851). Lassen Co.; 88 (1, 0) Madeline Plains (USNM 89120). 89 (5, 8, 1) Mt. Lassen (and adjacent Shasta Co. and Tehama Co.) (MVZ 34158, 34932, 36883–36885, 36889, 36890, 36897, 36898, 37616, 39771; USNM 95728, 95729, 98137). Plumas Co.; 90 (1, 0) Prattville (USNM 98136). Placer Co.; 91 (9, 7, 1) Donner (MVZ 25561; AMNH 1305, 1306; UCLA G252–G254, G259, G273; UIDA 237604; USNM 55544, 95583, 95585, 100246, 100249, 100250, 100464, 186517). El Dorado Co.; 92 (10, 4) Pyramid Peak (CAS 17460; MCZ 9290, 9294, 9295, 9298; MVZ 11999, 12000, 12004, 103443, 165868, 165869; UCLA G160, G161, G166). Alpine Co.; 93 (15, 9) Ebbetts Pass (UMNH 14162; CAS 7647, 7648; HSU 3649, 3652–3656, 3658; USNM 68457, 68458; NMMNH 833–835, 838, 840, 845). Mono Co.; 94 (41, 34) Yosemite (and adjacent El Dorado Co.) (KU 10731; HSU 3539, 3540; MVZ 22903, 22906, 22907, 22912, 22914, 22916, 22917, 22920, 22921, 23462, 23465, 23468, 23473, 23476, 23478, 23480, 24145, 32942, 105631, 105637, 105640, 105644, 105645, 105647, 105648, 105650–105653, 105655, 105657, 105658, 105660, 105661, 135056–135061, 135063, 135064, 135067, 135071, 135073, 135074, 135079–135086; UCLA 9936, 9945, 9949; USNM 42067, 109181, 109182, 109270, 109407, 109408, 110344, 110347, 110348, 244756, 259064, 264536; SDNHM 6714–6715; SDSNH 21655). 95 (19, 15, 1) White Mtns. (and adjacent Inyo Co. and NEVADA: Esmeralda Co.) (HSU 5775; MVZ 27553, 27555–27557, 27559–27561, 27564–27566, 27569–27574, 27578–27582, 59396, 119302, 119303; LACM 3900, 3902, 3903; USNM 41420, 41422, 41425, 41526; NMMNH 819, 820, 824). Fresno Co.; 96 (1, 3) Bullfrog Lake (MVZ 25315, 25316, 25319, 25320). Inyo Co.; 97 (22, 22) Mt. Whitney (and adjacent Tulare Co.) (HSU 5908; MCZ 8482, 8483; MVZ 16223, 17543, 17546, 17547, 17552, 17553, 17555, 17557, 17558, 30095, 99289, 99290, 109100; AMNH 138968, 138970, 138971; LACM 32347; USNM 41246, 42297, 42686–42690, 42692, 42694–42698, 42703, 274858–274860; NMMNH 828–831).

NEVADA: Humboldt Co.; 98 (2, 3) Pine Forest Range (and adjacent Washoe Co.) (MVZ 66659, 66660, 74273, 74274, 96767). Mineral Co.; 99 (0, 4) Wassuk Range (MVZ 107330, 107332, 107335, 107337). Churchill Co.; 100 (6, 7) Desatoya Mtns. (and adjacent Lander Co.) (HSU 5144–5148, 5150–5152; MVZ 64529–64533). Lander Co.; 101 (3, 5) Shoshone Mtns. (HSU 5297–5299; MVZ 64537, 64538, 64540, 64542, 64544). 102 (4, 0) Toiyabe Range (USNM 93471, 93472, 93651, 208917). Nye Co.; 83 (3, 10) Toquima Range (MVZ 58497, 58500–58503, 58505–58507, 58509, 58515; NMMNH 250, 255). 104 (4,3) Monitor Range (MVZ 58516, 58517, 58519, 58521–58523, 58525).

UTAH: Iron Co.; 105 (16, 14, 3) Cedar City (CM 14729, 14733, 14737, 14740; UMNH 1426, 8122, 8124, 8126, 20374; BYU 1375, 1376, 1486, 1488, 2140, 2608, 2609, 2611; MVZ 41415, 41418, 47648, 70284, 113632; AMNH 124066, 124067; USNM 158091, 158094–158096, 158098; NMMNH 221, 226–228). Beaver Co.; 106 (23, 16, 1) Beaver (CM 14720, 14722; UMNH 17007, 17008, 17013, 17016–17018, 17021, 17023, 17026–17029; MVZ 41419, 41421–41424; AMNH 28733, 124740; USNM 158080, 158082, 158085–158087, 158089, 158093, 158499; NMMNH 235, 237, 240, 242, 246, 247). Sevier Co.; 107 (1, 0) Monroe Peak (UMNH 17003).

Ochotona princeps uinta (n = 120; 58, 59, 3)

UTAH: Summit Co.; 108 (18, 25, 3) Uinta Mtns. (and adjacent Daggett Co., Duchesne Co., Summit Co., Uinta Co., and Wasatch Co.) (CM 9461, 9462, 9465, 9469, 9472, 9479, 9485, 9486, 12021, 12022, 12026–12028, 12030, 12033, 14181, 19284; KU 38089; UMNH 615, 622, 4808, 7411, 18815, 18819, 18820, 18823, 18827–18829, 27226; BYU 1304; MCZ 18562–18564; MVZ 41429; USNM 9750, 25608, 25609; NMMNH 281, 285, 291–295, 298). San Pete Co.; 109 (5, 6) Gunnison (UMNH 10633, 10634, 10646, 10647, 28554, 28558; USNM 248436; NMMNH 310, 313, 316, 324). Sevier Co.; 110 (13, 20) Richfield (UMNH 8138, 8140, 8142, 8143, 8248, 8249, 8252, 9364, 9366, 9368, 10640, 10641, 10648, 10882–10884, 13473, 21747, 23323; PSM 9329, 9330; NMMNH 327–329, 333, 334, 336, 338–340, 343, 345, 346). Wayne Co.; 111 (4, 0) Thousand Lake Mtn. (NMMNH 1217, 1220, 1222, 1224). Garfield Co.; 112 (18, 8) Aquarius Plateau (and adjacent Wayne Co.) (NMMNH 881, 882, 884–886, 888, 890, 893; UMNH 8128, 8129, 8232, 8234, 8236–8239, 8242, 8243, 9369–9372; BYU 1595, 1611; MVZ 95273; USNM 287776).

Ochotona princeps saxatilis (n = 300; 150, 143, 7)

UTAH: San Juan Co.; 113 (10, 17, 1) La Sal Mtns. (and adjacent Grand Co.) (CM 16025, 16026, 20201, 20202, 20608; UMNH 6409, 11148, 11150, 11153–11157, 13583, 14636–14639, 21745, 21746; USNM 149966; NMMNH 349, 354, 356–358, 360, 365).

COLORADO: Larimer Co.; 114 (4, 6) Rocky Mountain National Park (KU 12654, 12656, 25150; MVZ 84148, 84149; AMNH 36139; USNM 74052, 74054–74056). Clear Creek Co.; 115 (26, 23) Front Range (and adjacent Boulder Co., Gilpin Co., Grand Co., Park Co., and Summit Co.) (CM 22167; AMNH 36140; DMNH 1369, 1373, 1378, 1379, 1381, 3055, 3057, 3058, 3071, 3072, 3074, 3075, 3077, 3080, 3346, 3348, 3640, 4000–4002, 4007–4010, 4013, 4017, 4371, 4893, 4894, 4896, 4897, 4926; FMNH 11243, 11248; CMN 28007, 28008; NMMNH 436, 438–441, 445, 446, 448, 450–452). Park Co.; 116 (8, 9) Alma (and adjacent Summit Co.) (MCZ 2703; MVZ 6915; ROM 19656; DMNH 1258, 1261, 2169, 4365, 4369, 5598, 5600, 5601, 5604, 7101; USNM 205789, 205791–205793). Lake Co.; 117 (5, 4) Leadville (and adjacent Pitkin Co.) (KU 113637, 113639, 113642, 113644–113647; DMNH 3289, 3652). Gunnison Co.; 118 (3, 5, 1) Gothic (MCZ 7712; MVZ 120340, 125655, 125656; UAMZ 5305, 5354; USNM 303592, 485418, 485420). Delta Co.; 119 (8, 6, 1) Grand Mesa (and adjacent Mesa Co.) (KU 59672–59674; PSM 9345; FMNH 90618–90621; NMMNH 367–370, 372, 373, 375). Chaffee Co.; 120 (12, 15) Garfield (and adjacent Gunnison Co.) (MVZ 125654; DMNH 3906, 3907; NMMNH 63–66, 68, 74, 77, 83, 86, 380, 381, 385, 386, 390, 393–397, 399, 401–404). El Paso Co.; 121 (2, 4, 1) Pikes Peak (NMMNH 685, 687, 689, 696, 698, 700). Saguache Co.; 122 (5, 6) La Garita Mtns. (UMNH 17626, 17627, 17632–17640). San Juan Co.; 123 (16, 8, 2) Silverton (and adjacent Ouray Co.) (AMNH 128312; DMNH 5463, 5465–5467; USNM 48193, 56755, 56757–56760, 56762, 56764–56771; NMMNH 1062, 1065, 1066, 1069, 1072, 1073). Huerfano Co.; 124 (8, 11) Cuchara (KU 59676; NMMNH 92–98, 100, 106–108, 110, 116, 117, 701, 702, 704, 708). Conejos Co.; 125 (8, 8, 1) Cumbres Pass (and adjacent Archuleta Co. and NEW MEXICO: Rio Arriba Co.) (MSB 53996, 53998, 54000; MVZ 116343, 116344, 116346, 116347; DMNH 886, 1065, 1067, 1465, 1522, 1529; USNM 157559; NMMNH 1206–1208).

NEW MEXICO: Taos Co.; 126 (10, 7) Twining (MSB 1435, 1437, 27721; TTU 2368, 2370; USNM 130970, 133469; NMMNH 119, 122, 131, 132, 135, 729, 734–737). Rio Arriba Co.; 127 (1, 0) Truchas Peak (USNM 128740). Sandoval Co.; 128 (12, 6) Jemez Mtns. (and adjacent Los Alamos Co.) (MSB 31222, 31613, 31614; USNM 147976; NMMNH 50, 165, 170, 1750–1753, 1757–1759, 1761, 1762, 1765, 1766). Santa Fe Co.; 129 (12, 8) Tesuque (and adjacent San Miguel Co.) (KU 58956; CAS 11105; TTU 16790; USNM 128912–128914; NMMNH 155, 156, 159, 160, 162, 164, 709–711, 719, 722, 724, 727, 728).

Footnotes

  • Associate Editor was Burton K. Lim.

Literature Cited

View Abstract