The Holocene

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The Holocene history of bighorn sheep (Ovis canadensis) in eastern Washington state, northwestern USA R. Lee Lyman The Holocene 2009; 19; 143 DOI: 10.1177/0959683608098958

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Downloaded from http://hol.sagepub.com at University of Missouri-Columbia on January 13, 2009 The Holocene 19,1 (2009) pp. 143–150

The Holocene history of bighorn sheep (Ovis canadensis) in eastern Washington state, northwestern USA R. Lee Lyman*

(Department of Anthropology, 107 Swallow Hall, University of Missouri-Columbia, Columbia MO 65211, USA)

Received 9 May 2008; revised manuscript accepted 30 June 2008

Abstract: Historical data are incomplete regarding the presence/absence and distribution of bighorn sheep (Ovis canadensis) in eastern Washington State. Palaeozoological (archaeological and palaeontological) data indicate bighorn were present in many areas there during most of the last 10 000 years. Bighorn occupied the xeric shrub-steppe habitats of the Channeled Scablands, likely because the Scablands provided the steep escape terrain bighorn prefer. The relative abundance of bighorn is greatest during climatically dry intervals and low during a moist period. Bighorn remains tend to increase in relative abundance over the last 6000 years. Bighorn were present in eastern Washington in the nineteenth century, but were largely locally extirpated by the twentieth century. A collection of prehistoric bighorn astragali that likely includes specimens spanning the terminal Pleistocene and the entire Holocene consists of specimens that are, with one exception, the same size as a collection of modern bighorn astragali from Wyoming. The single exceptionally large specimen falls in the middle of the size range of specimens from the late Pleistocene of northern Wyoming, suggesting that Washington’s prehistoric bighorn, like those from other areas, were larger than modern bighorn.

Key words: Biogeography, bighorn sheep, chronocline, Holocene, northwestern USA, Ovis canadensis catclawensis

Introduction Dalquest (1948: 405) speculated that bighorn had ‘in the past [occupied] the cliffs of the Columbia River Valley in eastern When Taylor and Shaw (1929: 31) produced their provisional list Washington’, including ‘the eastern or Columbia Plateau side of of land mammals of Washington State (USA), they noted that the [Columbia] river and therefore probably occupied the cliffs of Moses Coulee and Grand Coulee’. (Moses Coulee extends south- bighorn sheep (Ovis canadensis) were at that time ‘nearly extirpated’ within the state but that this ungulate had ‘apparently [once west through the centre of Douglas County. Grand Coulee is at the occurred] not only in the mountains and foothills [of the Cascade border of northwestern Grant County and the northeastern border Range], but also on the plains nearby’. They could add no evi- of Douglas County.) Unique for his time, Dalquest (1948: 405) dence to confirm Dice’s (1919: 20) earlier report that bighorn reported that ‘in caves along the Columbia River in [western] were thought to have previously occurred in the Blue Mountains Grant County, bones of sheep [had been] found in association with of the southeastern portion of the state. Booth (1947: 605) later stone arrowheads and other human artifacts’. Dalquest likely was indicated that bighorn still occurred in the Wallowa Mountains of referring to archaeological work taking place in the 1940s (eg, geographically adjacent northeastern Oregon, and though none Collier et al., 1942; Osborne et al., 1952; Swanson, 1962). had been seen in the Blue Mountains ‘in recent times, their former Dalquest and Hoffmeister (1948) described 37 skulls collected occurrence [there] is well substantiated’. He plotted two locations in 1889 near Mount Chopaka in northernmost central Okanogan of bighorn sheep in Columbia County but failed to indicate the County (Figure 1). Perhaps it was this large sample that type and source of the data for those locations. Booth (1947: 607) prompted Buechner (1960: 14) to report that bighorn ‘do not also suspected that bighorn may have once ‘been common on the seem to have been exceptionally abundant’ in Washington plains of [the Columbia Basin]’ (Douglas, Lincoln, Grant, Adams, except along the British Columbia border. He suggested that Franklin and Benton Counties; Figure 1) but could offer no evi- bighorn were likely absent from the lower Yakima Valley (in dence to validate his suspicion. central Benton County) and virtually all of the southeastern Whereas Booth (1947) suggested bighorn had once occupied fourth of Washington (Buechner, 1960). D.R. Johnson (1972) the ‘plains’ west of the Columbia River (the Yakima Valley?), questioned a historic report of bighorn in the extreme northeastern portion of Pend Oreille County. In his synopsis of bighorn sheep in Washington, R.L. Johnson (1983: 115–16) listed 18 *Author for correspondence (email: [email protected]) locations where bighorn sheep or their remains – typically an

Methods and materials

I consulted all reports known to me concerning the Holocene palaeozoology of eastern Washington, a politically defined geographic area bounded on the north by British Columbia, the east by Idaho and the south by Oregon. I chose the crest of the Cascade Range as the western boundary, and used county lines that follow or approximate the east–west drainage divide to define that boundary (Figure 1). I recorded the number of identified specimens (NISP) (Grayson, 1984; Lyman, 2008b) of bighorn sheep at each archaeological site location when such data were reported; some- times only the presence and not abundance of such remains was reported. A specimen is a bone or tooth or fragment thereof. Only those archaeological remains that were not modified into artefacts were recorded to avoid including bones and teeth that had been transported long distances from where animals were procured (Lyman, 1994); no palaeontological remains were recorded as none have been reported. Age of the remains was determined using stratigraphically associated radiocarbon ages, temporally diagnostic artefacts that were stratigraphically associated in the absence of the radiometric ages, or temporally distinct strata such as radiometrically dated volcanic tephra (eg, Lyman 2000a). For analytical purposes, standard zooarchaeological procedure was followed and the mid-point of the age range of deposits was used for statistical analysis (Lyman, 2003). Geographic locations of deposits that produced remains of bighorn were recorded by legal Figure 1 Map of eastern Washington State showing townships sam- description (township, range) and also plotted on a map. pled for palaeozological remains, townships where prehistoric bighorn Because sampling of eastern Washington deposits that contain remains have been identified and townships where bighorn have been mammalian remains has not been probabilistic across geographic historically reported. Faunal remains have been recovered from, but not space (Lyman, 2002, 2004b), efforts to determine the prehistoric studied in, some sampled townships. Five townships with historic range of bighorn or to monitor changes in the distribution of bighorn records have been archaeologically sampled but no bighorn remains must be tempered by knowledge that available samples may not be were identified. County lines and county names are shown for reference representative. The presence of remains of bighorn, particularly those of low socioeconomic value (minimal food value or useless as isolated skull someone had collected – were known to occur or tool material) such as phalanges that are unlikely to have been trans- had been found. One of the locations Johnson (1983) listed was ported far by prehistoric hunters (Lyman, 1994), may be viewed as an archaeological record (Thomson, 1962), but he did not report evidence of relatively local origin (< 10 km). The absence of all such records. The most recent synopsis of Washington State remains may reflect local absence of the species, lack of preserva- mammals does not include any reference to pre-1900 bighorn tion of bighorn remains or lack of recovery of bighorn remains as a (Johnson and Cassidy, 1997). result of minimal sampling (eg, Grayson, 1981; Lyman, 1994, 2002, In sum, the biogeographic record for bighorn in eastern 2004b, 2008a). Because bighorn were likely acquired by prehistoric Washington is of uneven quality; some of it is based on observed hunters within several kilometres of the site of bone and tooth dep- individuals, some on fortuitously collected specimens, some on osition, locations where bighorn remains have been recovered were hearsay. It is unlikely that the historic record will improve. mapped by township – a rectangular unit, usually 9.65 km × 9.65 Dalquest (1948) and Johnson (1983) identified another data km, used for legal geographic descriptions – rather than by individ- source that continues to improve – the palaeozoological (palaeon- ual recovery site. Each unique township that has produced remains tological and zooarchaeological) record. Subsequent to their of bighorn is noted regardless of how many locations or sites within reports of archaeological materials, excavations by professional that township have produced remains. archaeologists have resulted in the recovery of numerous bighorn In general, the older a deposit in eastern Washington, the less sheep remains from many archaeological deposits in eastern intensively and extensively it has been sampled (Lyman, 2000a, b, Washington. The prehistoric and early historic remains of animals 2004b). Thus I follow standard palaeozoological procedure recovered from palaeozoological excavations provide unprece- (Grayson, 1984) and track abundances of bighorn remains relative dented data for studying historic biogeography. to abundances of remains of other taxa across time and space. This In this paper I summarize the palaeozoological evidence for analytical protocol circumvents potential depositional, preserva- bighorn distribution and relative abundance in eastern Washington tional and sampling biases that may skew results (Lyman, 2008b). during the Holocene, or approximately the last 10 000 years in Nevertheless, detection of the influence of sample size gives rea- order to answer four questions. Were bighorn sheep present in son to be cautious. All ages are given as radiocarbon years before Washington State in pre-Columbian (pre-sixteenth century) present (RCYBP). times? If so, did their relative abundance shift coincident with climate change, as did that of bison (Bison sp.) and wapiti (Cervus elaphus) (Lyman 2004a, b)? Third, why were bighorn rare in east- Palaeoecology and ecology ern Washington in the late nineteenth and early twentieth centuries? Finally, are prehistoric bighorn bones from eastern Analysis of recent crania of bighorn (Wehausen and Ramey, 2000) Washington larger than those of modern bighorn, as has been suggests the distributions of bighorn subspecies postulated by observed elsewhere in North America? Cowan (1940) are incorrect. During the nineteenth and twentieth

Downloaded from http://hol.sagepub.com at University of Missouri-Columbia on January 13, 2009 R. Lee Lyman: Holocene history of bighorn sheep, Washington state 145 centuries (the likely time when specimens studied by Wehausen Table 1 Abundance (NISP, number of identified specimens) of and Ramey (2000) were deposited) all of eastern Washington State bighorn sheep remains and other artiodactyl remains in deposits was, apparently, occupied by Ovis canadensis canadensis rather containing bighorn sheep remains per 500 yr time interval in than O. c. californiana. Because modern subspecies designations eastern Washington State tend to be based on characters that do not preserve in the palaeozoological record, I do not explore the issue of which subspecies Age (RCYBP) Bighorn NISP Other Artiodactyl NISP was present prehistorically, with one notable exception. 0–500 299 2705 Eastern Washington witnessed changes in climate during the 501–1000 1221 3262 Holocene that might account for variation in the relative abundance 1001–1500 144 704 of bighorn during that time; forage quality and quantity have a 1501–2000 390 726 direct bearing on the size of bighorn populations (Festa-Bianchet, 2001–2500 33 513 1988a, b, c). As the Pleistocene ended and the Holocene began 2501–3000 137 3152 about 10 000 RCYBP, summers became warmer and drier 3001–3500 395 1337 (because of evaporation) and winters became extremely cold (cli- 3501–4000 55 290 matic history from Chatters, 1998). There was more grass and less 4001–4500 78 589 4501–5000 19 97 shrub vegetation in the Columbia Basin than today. About 8500 5001–5500 335 1441 RCYBP winters became warmer and wetter, but there was greater 5501–6000 12 127 aridity overall and grass decreased and sagebrush increased in 6001–6500 35 abundance. Between 5500 and 3500 RCYBP climates cooled and 6501–7000 14 moisture increased after 4500 RCYBP. After 3500 RCYBP cli- 7001–7500 1 164 mates remained cool and with abundant moisture, ground cover 7501–8000 00 increased. After 2500 RCYBP climates warmed and some shrub 8001–8500 00 steppe in the southeastern portion of the basin became more grassy. 8501–9000 00 Where the archaeological record of upland habitats is well 9001–9500 00 known, such as in western Wyoming and adjacent areas (eg, 9501–10 000 00 10 001–10 500 18 29 Grant, 1981; Frison et al., 1990; McGuire and Hatoff, 1991; Frison, 2004), bighorn sheep were regularly exploited by American Indians. Bighorn were procured by interception and ambush along seasonal migration routes, and by communal hunts from such settings, such as in Yakima, Chelan and Stevens coun- that utilized artificial drive lines that guided herds into elaborate ties, bighorn remains are not unusual (Figure 1). The prehistoric enclosures and traps constructed of logs and boulders. Where suf- record thus supplements the historic record for this species in ficiently well preserved, it is apparent that drive lines and enclo- mountainous areas. Perhaps more interesting is the fact that, as sures were constructed in such a way as to indicate intimate conjectured by Dalquest (1948), bighorn remains found in the cen- knowledge of bighorn behaviour, taking advantage of typical tral Columbia Basin east of the Columbia River indicate that predator-escape behaviour on a particular area of landscape bighorn sheep once roamed this part of the state. In this case the (Frison et al., 1990). These observations indicate two things perti- prehistoric record markedly expands the historic record. nent to this study. First, indirect (non-faunal) evidence of bighorn The oldest known bighorn remains in Washington State come may exist in the archaeological record. Second, archaeological from two sites. Seventeen bones and teeth (Lyman, unpublished remains of bighorn resulting from drives likely represent non- data, 2003) of bighorn recovered from Sentinel Gap (official state selective demographic cross-sections of bighorn populations. archaeological site number: 45KT1362) in eastern Kittitas County date to 10 200 RCYBP (Galm and Gough, 2000). A single humanly butchered bighorn bone was recovered from an undated stratum at Results Marmes Rockshelter (45FR50) in northeastern Franklin County (Figure 1). It was recovered from deposits stratigraphically below a Collier et al. (1942) and Osborne et al. (1952) were the first to stratum dated to about 10 000 RCYBP (Gustafson and Wegener, report the recovery of bighorn remains from archaeological sites in 1999: 84). This specimen was likely deposited between 11 200 and eastern Washington. Collier et al. (1942) listed two sites that pro- 10 000 RCYBP as the oldest known artefacts from the site fall in duced remains of bighorn and Osborne et al. (1952) listed two other this time span (Hicks, 2004). Bighorn remains have been recovered sites, but both research teams had only a vague notion of the age of from other deposits dating to the last 7500 years (Table 1). The the remains. In the subsequent four decades 87 (of 91 total) more absence of bighorn remains dated between 7500 and 10 000 archaeological sites have produced remains of bighorn (see Lyman, RCYBP is likely a result of inadequate sample sizes. 1995 and Parks, 2000, for partial summaries). These sites tend to be Artiodactyl remains present in palaeozoological collections concentrated along the Columbia and Snake Rivers where the under study here represent not only bighorn but deer (Odocoileus majority of archaeological excavations in eastern Washington have virginianus and O. hemionus), pronghorn (Antilocapra ameri- taken place (Lyman, 2002). As of mid-2007, archaeological and cana), wapiti (Cervus elaphus) and bison (Bison antiquus and B. palaeontological deposits in 206 townships had been sampled bison). All of these taxa of large mammal were frequent prey of (Figure 1). Not all sampled townships have produced mammalian prehistoric human hunters (Lyman, 2003); some taxa represented remains, and the remains collected from some townships have never in palaeozoological collections were not exploited by prehistoric been identified or described in the literature (Lyman, 2004b). The humans. The frequency of bighorn remains per 500 yr interval studied and published record indicates that individual sites in 55 correlates with the frequency of non-bighorn artiodactyl remains townships have produced remains of bighorn (Figure 1). per 500 yr interval (Pearson’s r = 0.702, P < 0.005), suggesting The historic records reported by Johnson (1983) and Johnson that the frequency of bighorn remains at any given time is a func- and Cassidy (1997) are largely restricted to mountainous terrain tion of sampling and analytical effort. Similarly, the frequency of (Figure 1). Archaeological excavation has to date been limited in bighorn remains per 500 yr interval correlates with both the fre- the boreal, subalpine and alpine habitats of the mountain ranges of quency of radiocarbon dates (r = 0.656, P < 0.01) and the eastern Washington. When faunal remains have been recovered frequency of radiocarbon-dated sites (r = 0.702, P < 0.005) per 500 yr

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such as those that have been found elsewhere (eg, Frison et al., 1990) would be indirect evidence for the prehistoric presence of bighorn in the area, as might prehistoric depictions of bighorn in the form of rock art (pictographs, petroglyphs) (eg, Keyser, 1992; Lothson, 1998). Eight assemblages of faunal remains containing bighorn sheep are post-Columbian in age (< 500 RCYBP). Of these, four seem to date to the nineteenth century (< 150 RCYBP). One of the lat- ter is adjacent to lower Grand Coulee, and two others are in southeastern Kittitas County. These are areas where historic records indicate bighorn were found in the last 200 or so years. Why were bighorn not more abundant and more ubiquitous at the end of the nineteenth century and beginning of the twentieth century? Archaeologists have often suggested several reasons for the historic low abundances of bighorn (Pippin, 1979; Frison, 2004). Figure 2 Relative abundance of bighorn remains among all artio- These include overhunting by American Indians equipped with dactyl remains in 500 yr bins (data from Table 1) rifles, overhunting by Euroamerican settlers, harsh winters and disease transfer from early historic livestock (Stelfox, 1971; Ebert, interval (data from Lyman, 2000b). The frequency of radiocarbon 1978). There were several harsh winters near the end of the nine- dates and the frequency of dated sites are both good proxy meas- teenth century that virtually wiped out local stockmen (Oliphant, ures of archaeological sampling intensity (Lyman, 2000a, b). Thus 1932). These winters could have impacted local bighorn popula- it would be unwise to infer anything about the absolute abundance tions as well, and if those populations were already depleted and of bighorn on the landscape, though the abundance of bighorn isolated from firearm hunting, then those winters may have been remains relative to the abundance of remains of other taxa can be the final cause of their extirpation. monitored (Grayson, 1984; Lyman, 2008b). All artiodactyl remains in assemblages of known age (N = 100) were lumped into temporal bins of 500 yr duration and the relative Size of bighorn (percentage) abundance of bighorn remains among all artiodactyls remains was calculated (Table 1, Figure 2). The abundance of During the late Pleistocene the high rugged mountainous terrain of bighorn remains increases slightly over time relative to the total western North America was occupied by a now extinct form of number of artiodactyl remains (Cochran’s test of linear trends Chi- bighorn sheep. Originally given species status by the palaeontolo- squared = 35.26, P < 0.0001). Causes of this increase are unclear. gists who first recognized it (Hibbard and Wright, 1956), some It may relate to the warmer and drier climate of the last 2500 palaeozoologists now consider the taxon a temporal subspecies – years. The relative abundance of bighorn remains is greater after Ovis canadensis catclawensis based on its larger than modern 2500 RCYBP (21.5%) than between 4500 and 2500 RCYBP average size (Harris and Mundel, 1974; Wang, 1984, 1988; (11.0%) when climates were cooler and more moist. Deep snow Emslie, 1986), although not everyone agrees with this assessment (>45 cm) is avoided by some bighorn populations today (Tilton (Gilbert and Martin, 1984; see especially Akersten et al., 2002). and Willard, 1982). If much of the moisture between 4500 and Detailed study of the morphometry of a large sample of skeletal 2500 RCYBP was snowfall, it may account for low bighorn abun- remains suggests the temporal subspecies designation is reason- dances at that time. able (Lawler, 1992, 1996), and that is the designation I use here Assemblages in which bighorn remains are abundant (> 50% of simply to designate the prehistoric large form. the total artiodactyl remains) are located in or adjacent to rugged The undated Moses Coulee Cave (45DO331) collection from terrain such as the deeply incised Snake River canyon (one col- southwestern Douglas County represents the largest collection of lection), Columbia River canyon between Douglas and Okanogan bighorn remains from any single site in eastern Washington, with counties (four collections) and between Douglas and Chelan coun- more than 2500 specimens reported, the majority of which were ties (one collection), Grand Coulee (two collections) and Moses accumulated by prehistoric humans who preyed on the sheep Coulee (one collection) (between Douglas and Grant counties). (Lyman, 1995). Temporally diagnostic projectile points suggest Only one collection (in southeastern Kittitas County) with abun- the cave was utilized by people during the entire span of the dant bighorn remains is not in such a topographic location. The Holocene. The bighorn remains likely were accumulated through- distribution of sites with abundant bighorn remains is significantly out this time period. How do the sizes of those bighorn remains associated with rugged terrain (Chi2 = 6.4, P < 0.025). It is likely compare with modern bighorn and to O. c. catclawensis? that the assemblages with relatively high abundances of bighorn To determine if any of the Moses Coulee Cave bighorn remains have those high abundances precisely because of the loca- remains were approximately the size of O. c. catclawensis, meas- tions of the sites that produced the remains. The settings of those urements of 15 dimensions of six kinds of well-preserved skele- sites include predator escape terrain, a favoured variable of tal parts were taken. The 214 skeletal parts (distal humerus, bighorn sheep habitats (Shackleton, 1985). proximal radius, proximal and distal metacarpal, distal tibia, All archaeological collections in eastern Washington that con- proximal and distal metatarsal, astragalus) provided 408 meas- tain remains of bighorn sheep derive from habitation sites rather urements. Lawler (1992) presented size data for the same 15 than procurement or kill sites. Thus far no clear evidence of drive dimensions from a sample of modern bighorn from Wyoming, lines or enclosure corrals or traps have been reported in uplands and for those same dimensions of the O. c. catclawensis speci- adjacent to the sites (but see Lothson, 1998), or elsewhere for that mens from Natural Trap Cave, Wyoming. Hockett and matter, but that is likely the result of sampling error. Virtually no Dillingham (2004) presented mean size data for ten of the dimen- archaeological research has, for example, been done in the Blue sions as represented by a small sample of O. c. catclawensis Mountains of southeastern Washington, and very little has been remains from Mineral Hill Cave, Nevada. A ratio diagram done in the uplands adjacent to the river canyons of the north- (Simpson, 1941) was constructed using the mean size of modern eastern portion of the state. Discovery of big game drive features bighorn bones as the standard (Figure 3). Mean size of O. c.

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Figure 3 Ratio diagram of bighorn sheep remains from Moses Coulee Cave (dots), Ovis canadensis catclawensis from Mineral Hill Cave (filled squares) and Natural Trap Cave (open squares) and modern bighorn from Wyoming (0.0 line). Points to the right of the 0.0 line are larger than an average modern bighorn, points to the left are smaller than an average modern bighorn. Some Moses Coulee Cave points do not show as a result of overlap. PB, proximal breadth (latero-medial); PD, proximal depth (antero-posterior); DB, distal breadth; DD, distal depth

catclawensis at Natural Trap Cave (from Lawler, 1992) and at bighorn (Student’s t = 3.066, P < 0.005, one-tailed test), and not Mineral Hill Cave (Hockett and Dillingham, 2004) were plotted significantly smaller than the average Natural Trap Cave specimen on the ratio diagram. Then, individual measurements of Moses (Student’s t = 0.313, P > 0.5). However, the distal depth of this Coulee Cave specimens were plotted on the graph. specimen is not significantly different than the average modern The ratio diagram (Figure 3) indicates several things. First, both bighorn (Student’s t = 0.32, P > 0.5) and is significantly smaller samples of O. c. catclawensis specimens are, on average, larger than the average Natural Trap Cave specimen (Student’s t = 2.677, than modern bighorn. Second, the Mineral Hill Cave O. c. cat- P < 0.01). Thus the metacarpal specimen cannot be confidently clawensis specimens are, on average, a bit smaller than the speci- assigned to O. c. catclawensis, though it might represent that form mens from Natural Trap Cave. Third, most specimens from Moses given the overlap in size of that form with modern bighorn Coulee Cave are approximately the same size as bones of modern (Lawler, 1992, 1996). bighorn from Wyoming. Fourth, most of the Moses Coulee Cave The astragalus seems to represent O. c. catclawensis. It not only specimens are smaller than the bones from Natural Trap Cave, and exceeds the mean size of specimens of this form recovered from many are smaller than those from Mineral Hill Cave. There are, Natural Trap Cave (Figure 3), but it falls well within the size range however, two specimens that deserve closer inspection. One distal of those specimens and far outside the range of modern bighorn. I metacarpal is close to the average size of the Natural Trap Cave measured two dimensions (greatest lateral length, distal breadth; specimens, and one astragalus exceeds the size of an average see von den Driesh, 1976) of 64 bighorn astragali (30 lefts, 34 Natural Trap Cave specimen. The distal breadth (DB) of the rights) from Moses Coulee Cave. I also measured the same two metacarpal specimen is significantly larger than the average modern dimensions on 61 astragali of modern bighorn sheep from

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Figure 4 Scatterplot of measurements of two dimensions of astragali from 61 modern Wyoming bighorn, 64 prehistoric bighorn from Moses Coulee Cave, Washington, and 78 late Pleistocene O. c. catclawensis from Natural Trap Cave, Wyoming. Arrow indicates possible O. c. catclawensis specimen from Moses Coulee Cave

Wyoming; measurements of these dimensions on 78 astragali from the early Holocene, available palaeozoological evidence from Natural Trap Cave in north-central Wyoming were taken suggests bighorn were present there during the last 10 500 from Lawler (1992). All three sets of measurements are plotted in (radiocarbon) years or so. The relative abundance of bighorn Figure 4, which indicates the majority of the astragali from Moses remains fluctuates over the last 4500 years in a manner related to Coulee Cave are modern in size. The placement of the large climatic change; there are more remains when climate was drier Moses Coulee Cave astragalus in Figure 3 is based on one dimen- and fewer remains when wetter. Bighorn remains tend to sion. Figure 4 shows that this specimen is considerably larger than increase overall relative to remains of other artiodactyls over the any modern bighorn and falls near the middle of the size range of last 6000 years. Bighorn were more abundant in eastern Natural Trap Cave’s O. c. catclawensis sample in two dimensions, Washington than other artiodactyls in areas where rugged pred- strengthening the taxonomic inference. This single specimen indi- ator-escape terrain was present. Available evidence is poor, but cates that at least one individual from Moses Coulee Cave is of a it seems likely that bighorn were rare in the nineteenth and early size similar to Ovis canadensis catclawensis. The terminal twentieth centuries as a result of firearm hunting, diseases intro- Pleistocene through Holocene history of the chronocline of west- duced by domestic livestock and perhaps an exceptionally harsh ern North American bighorn sheep is poorly known, but it would winter or two. Finally, some prehistoric bighorn were, like those not be unexpected, given that history for wapiti (Lyman, 2004b, found elsewhere, larger than modern bighorn. Learning why 2006) and bison (Lyman, 2004a), to find some larger than modern they, like many other ungulates that survived the terminal bighorn in Holocene deposits. I therefore assign the large astra- Pleistocene, became smaller during the Holocene should be a galus to O. c. catclawensis. research priority because the reasons may have implications for Specimens of O. c. catclawensis have been documented in conservation biology. Idaho (Akersten et al., 2002), Wyoming (Wang, 1984, 1988; Lawler, 1996), Colorado (Emslie, 1986; Wang and Neas, 1987), Nevada (Hockett and Dillingham, 2004), New Mexico (Harris and Acknowledgements Mundel, 1974), and Arizona (Hibbard and Wright, 1956). The O. c. catclawensis specimen from Moses Coulee Cave is the first Thanks to Danny Walker (University of Wyoming) for the loan such specimen reported for Washington State. of bighorn astragali, Bryan Hockett for the Mineral Hill Cave report, Jim Mead (Northern Arizona University) for a critical piece of information and Mark Lawler (Onondaga Community Conclusions College) for a copy of his thesis and a reprint. Comments on an early draft by D.K. Grayson facilitated the revision process, and Bighorn sheep were present in eastern Washington State during two anonymous reviewers identified several ways to improve pre-Columbian times. Indeed, assuming inadequate samples the discussion.

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