Snake Creek Burial Cave and a review of the Quaternary mustelids of the Great Basin

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1 Great Basin Naturalist Volume 49 Number 2 Article Snake Creek Burial Cave and a review of the Quaternary mustelids of the Great Basin Emilee M. Mead Northern Arizona University, Flagstaff, Arizona Jim I. Mead Northern Arizona University, Flagstaff, Arizona Follow this and additional works at: Recommended Citation Mead, Emilee M. and Mead, Jim I. (1989) "Snake Creek Burial Cave and a review of the Quaternary mustelids of the Great Basin," Great Basin Naturalist: Vol. 49 : No. 2, Article 1. Available at: This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Great Basin Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact scholarsarchive@byu.edu, ellen_amatangelo@byu.edu.

2 The Great Basin Naturalist Published at Provo, Utah, by Bricham Young University ISSN Volume April 1989 No. 2 SNAKE CREEK BURIAL CAVE AND A REVIEW OF THE QUATERNARY MUSTELIDS OF THE GREAT BASIN EmileeM. Mead 13 and Jim I. Mead' 25 Abstract Snake Creek Burial Cave (SCBC), east central Nevada, is a unique paleontological deposit. The cave is the first natural trap excavated in the Great Basin and one of the few localities describing a valley-bottom community. The recovery of extinct Camelops sp. (camel) and Equus spp. (horse), in addition to radiometric dates, indicates at least some of the deposits to be of late Pleistocene age. Eight mustelid species have been identified from SCBC, including three species not previously reported from the late Rancholabrean of the Great Basin: Mustela nigripes (black-footed ferret), M. nivalis (least weasel), and Gulo gulo (wolverine). A review of late Pleistocene deposits indicates that there are more species of mustelids recovered from Snake Creek Burial Cave than from any other locality in the Great Basin. The Great Basin of western North America encompasses some 390,000 km 2 and, although centered in Nevada, also extends into several adjoining states (Fig. 1). This arid region is generally characterized by linear, north-south trending mountain ranges, separated by closed-drainage valleys (Hunt 1967). A number of archaeological and paleontological sites have been excavated in the Great Basin. Few, however, have contained lengthy, welldated, stratified sequences extending from the Pleistocene, across the critical Pleistocene-Holocene boundary, and into the Holocene. Futhermore, those few sites meeting these criteria have generally been restricted to the mountainous peripheries (Grayson 1987), leaving the valley-bottom communities largely undescribed. Since small mammals are generally sensitive to environmental change, their recovery in archaeological and paleontological sites can provide important paleoenvironmental data. Snake Creek Burial Cave (SCBC), White Pine County, east central Nevada, has recently been excavated; an estimated 30,000 fish, amphibian, reptilian, avian, and mammalian bones have been recovered. Of particular interest are the many carnivore remains, including numerous mustelids (Order Carnivora: Family Mustelidae; weasel family). This paper provides a brief description of SCBC, including recovered mustelids, as well as a timely review of other localities in the Great Basin reporting fossil or subfossil mustelids. A detailed osteological report of all carnivores recovered from SCBC is in progress. Additional reports describing the avian and remaining mammalian remains are forthcoming. Snake Creek Burial Cave Setting Snake Creek Burial Cave is a unique paleontological deposit. The natural trap cave, with a sinkhole depression and a 17-m, vertical, free-fall entrance (Fig. 2), is located at an elevation of 1,731 m on a small limestone 'Ralph M Bilby Research Center, Box 6013, Northern Arizona University, Flagstaff, Arizona department of Geology, Box Northern Arizona University, Flagstaff, Arizona Quaternary Studies Program, Northern Arizona University, Flagstaff, Arizona

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4 April 1989 Mead, Mead : Snake Creek Burial Cave 145 Fig. 2. Entrance to Snake Creek Burial Cave Recovered remains. Eight species of mustelids have been recovered from SCBC, including three species not previously reported from late Rancholabrean localities in the Great Basin: Mustela nigripes (blackfooted ferret), M. nivalis (least weasel), and Gulu gu/o (wolverine). The implications of these mustelid reports are discussed. The diversity of identified mustelid species from SCBC is higher than that from any other locality in the Great Basin. Table 1 provides a list of those localities in the Great Basin reporting fossil or subfossil mustelid remains. Reports from a majority of the sites include only one or two mustelid species (Hidden Cave and Smith Creek Cave both report six, the most after SCBC). However, some 18 localities are now reporting fossil or subfossil mustelids. Figure 1 illustrates the geographic locations of these 18 sites; the arbitrary site numbers correspond to the numbers assigned in Table 1. Also included is a review of the fossil and subfossil mustelid localities in the Great Basin. Descriptive Accounts Maries americana American marten Map localities. 2, 3, 5, 18. Discussion. Maries americana generally prefers mature conifer or mixed-forest stands with greater than 30% canopy cover, although meadows are often used in the summer if more food is available there. The American marten is an opportunistic feeder that takes advantage of local and seasonal abundances. While Clethrionomys spp. (red-backed vole), Microtus spp. (meadow vole), Lepus americanus (snowshoe hare), and Tamiasciurus spp. (tree squirrels) are favored foods, fruits and insects can play a significant role in diet (Ewer 1973, Strickland et al. etal. 1987). the 1982, Clark Martes americana is reported from four fossil localities in the central Great Basin. However, one of the two specimens from Bronco Charlie Cave (Spiess 1974) has been reassigned to M. nobilis (Grayson 1987) and is

5 146 Great Basin Naturalist Vol. 49, No. 2 Fig. 3. SCBC is located by the arrow, background. The valley bottom (pluvial Lake Bonneville) is clearly visible in the discussed in that section. The other specimen remains undetermined. Crystal Ball Cave, Deer Creek Cave, and Snake Creek Burial Cave are all outside the modern range of M. americana. Figure 6 identifies the fossil sites of M. americana, M. nobilis, and Maries sp. in the arid West and illustrates the modern geographic range of M. americana. The American marten currently ranges in the Sierra Nevada and the Bocky Mountains, but it does not occur on mountain ranges within the Great Basin. However, until more precise environmental and chronological guidelines are found that distinguish M. americana from M. nobilis localities, most paleoecological interpretations should be considered tentative. Maries nobilis Extinct noble marten Map localities. 2, 8, 17, 18. Discussion. Maries nobilis has been reported from three fossil localities in addition to the reassignment of the Bronco Charlie specimen (Fig. 6). Anderson (1970) provides measurements and morphological characters of the cranium and dentition used to distinguish the species. Anderson (1970: 85) states, "I do not believe that M. nobilis was related to M. americana, and that competition with the American marten, a warming climate, and perhaps the activities of man caused the extinction of the noble marten. Until recently, M. nobilis was thought to have become extinct at the end of the Pleistocene and perhaps to have adapted to cooler conditions. Grayson (1987), however, reports three localities in the arid West, two from the Great Basin, with noble marten remains dating into the late Holocene: (1) Dry Creek Bockshelter, southwestern Utah, 3,270 ± 110 yr B.P.; (2) Hidden Cave, Nevada, 3,600-3,700 yr B.P.; and (3) Bronco Charlie Cave, Nevada, 3,500 yr B.P. If we assume these dates are accurate in their associations, we see no clear reason why M. nobilis survived terminal Pleistocene environmental changes and then became extinct in the late Holocene. Clearly, additional research is required. Only at SCBC is there an indication that both species may have occurred together in the Great Basin.

6 April 1989 Mead, Mead: Snake Creek Burial Cave 147 /<2> ' to lower / cave orea Chimney Fig. 4. Plan view of SCBC with the excavation units enlarged. Martes sp. is reported from two localities, Hidden and Smith Creek caves (8, 17) (Fig. 6). The Hidden Cave specimens are all postcranial elements, and Grayson (1985) does not definitely distinguish them as either M. americana or M. nobilis. The Smith Creek Cave specimen was first identified by Goodrich (1965) as Martes sp. and further reported by Miller (1979), who made no further identification. Mustela erminea Ermine Map localities. 15, 17. Discussion. Ermines generally inhabit a variety of boreal habitats, although they tend to avoid dense coniferous forests and deserts. The diet of M. erminea consists primarily of small mammals, especially Microtus spp., Blarina spp. (short-tailed shrew), and Peromyscus spp. (deer mouse) (Ewer 1973,

7 148 Great Basin Naturalist Vol. 49, No. 2 Table 1. Fossil and subfossil mustelid localities reported from the Great Basin. Site numbers are arbitrary designations and correspond to the numbers on Figure 1. X = fossil/subfossil, in vicinity today; * = extirpated;! = extinct. «-p "H = Locality

8 April 1989 Mead, Mead: Snake Creek Burial Cave 149 HV

9 150 Great Basin Naturalist Vol. 49, No. 2 Fig. 8. Modern distribution of Mustela nivalis (shaded area); = Moonshiner Cave, Idaho (Anderson 1974); A = SCBC. localities are well outside the current distributional range of the least weasel (Fig. 8). Mustela vison Mink Map localities. 3, 8, 17. Discussion. The mink inhabits wetlands of all kinds including river banks, streams, and swamps. The diet of M. vison is quite variable by season and geographical location and can include insects, crustaceans, fish, amphibians, reptiles, and small mammals (Ewer 1973, Linscombe et al. 1982). Three fossil localities report M. vison in the central Great Basin, Crystal Ball Cave, Utah, and Hidden Cave and Smith Creek Cave in Nevada. The mink is known to currently inhabit the area near Hidden Cave (northwestern Nevada); however, it is extirpated from the Snake Bange (Crystal Ball and Smith Creek caves). The recovery of fossil M. vison from Crystal Ball and Smith Creek caves is not surprising due to the close proximity of these caves during the Pleistocene to either riparian (Smith Creek Cave) or lakeshore (Crystal Ball Cave) environments. With the recovery of M. nigripes from SCBC, and because of some osteological similarities with mink, all M. vison Fig. 9. Modern distribution map of Gulo gulo (shaded area). Dashed line indicates the range extension proposed bv Hall (1981). Fossil localities are indicated ( ); = SCBC material reported from the Great Basin should be reexamined. Mustela sp. is reported from five localities in the Great Basin (8, 9, 12, 13, 17). As discussed, the specimen reported from Last Supper Cave (Grayson 1988) may be assignable to either M. cf. erminea or M. cf. frenata, but it is too small to be either M. vison or M. nigripes. Gulo gulo (=luscus) Wolverine Map locality. 18. Discussion. SCBC is the only reported locality of Gulo in the central area of the Great Basin (Barker and Best 1976), although one was killed in 1900 in the Wasatch Mountains in Utah. Based solely upon this undated specimen, Hall (1981) extended the modern range of the genus into the Snake Bange (Fig. 9). However, since extinct fauna (Camelops and Equus) have been recovered from the surface of SCBC (cavers' backdirt, this report), presence alone cannot determine the age of the specimen. Without directly dating the SCBC specimen, it is impossible to unequivocally determine if the wolverine cranium is modern or fossil.

10 April 1989 Mead, Mead: Snake Creek Burial Cave 151 Table 2. Observed ranges, means, and standard deviations of modern Gulo gulo; measurements of the Snake Creek Burial Cave specimen; and observed ranges and means of Pleistocene Gulo specimens (Anderson 1977). All measurements in mm.

11 152 Great Basin Naturalist Vol. 49, No ). None of the eight reported fossil localities of S. putorius is outside the modern range. However, with the report of Brachyprotoma, an extinct skunk approximately the size of Spilogale, from Crystal Ball Cave, caution should he used in examining fossil skunk material from the Great Basin. Spilogale sp. is reported from two localities (13, 17); these specimens probably represent S. putorius. Discussion and Conclusions The Rancholabrean paleontological history of the Great Basin of western North America is largely unstudied. The region has provided a variety of diverse habitats through time, from valley to mountainous communities, including compressed zones resulting from mountain glaciations and pluvial lakes Bonneville and Lahontan in the Pleistocene. The Great Basin has been, and continues to be, a dynamic system recording fluctuating climatic conditions with lengthy paleobotanical and paleontological records. However, it has been only in the last few years that these records have been studied. Small mammals are sensitive to environmental change, and, therefore, their recovery from archaeological and paleontological sites can provide important paleoenvironmental data. SCBC has provided a unique situation in Great Basin paleontology, a deeply stratified, natural trap deposit representing a valleybottom community. The mustelid species diversity at SCBC exceeds any other reported locality in the Great Basin (Table 1) and includes three late Rancholabrean-age species not previously reported as fossil from the region: Mustela nigripes, M. nivalis, and Gtrfo gulo. The recovery of a Gulo cranium (Barker and Best 1976) from SCBC led Hall (1981) to extend the modern range of the genus into the Snake Range. Measurements of the SCBC specimen were compared with modern specimens and with Pleistocene wolverines from Alaska (Anderson 1977). We believe, based upon these measurements and on radiometric analyses of the cave fauna, that Gulo is not present today in eastern Nevada but, rather, that the specimen represents a fossil occurrence, the first in this part of the Great Basin. Several discoveries unique to SCBC have led us to further refine the hypotheses concerning vegetational communities in the valley bottoms of the central Great Basin during the last glacial. Brown (1971, 1978) proposed that, based upon the relictual appearance of boreal mammals on certain "island mountain ranges, the valley bottoms must have contained a continuous woodland or forest corridor from the Rocky Mountain "mainlands" to eastern Nevada. Wells (1983) examined Neotoma (packrat) middens and proposed that stands of the subalpine Finns longaeva (bristlecone pine) occurred down to 1,660 m elevation (as late as 11,880 yr B.P.), close to the high beach stands of Lake Bonneville, where the species probably mixed into a mosaic community with P. flexilis (limber pine). Thompson and Mead (1982) recovered a Ncotoma midden at 1,640 m elevation; from this record they hypothesized that subalpine stands of P. flexilis occurred at low elevations near the lake on rocky, calcareous substrates. Deep alluvial deposits in the valley bottoms (and away from Neotomo scavenging) were probably vegetated by shrub and/or meadow communities (Thompson and Mead 1982). Mead et al. (1982) analyzed a variety of faunas, including Smith Creek Cave (1,950 m elevation), from the nearby Snake Range. While many boreal and mountain species were recovered, the discoveries of Rancholabreanage C rotaphytus wislizenii ( = Gambclia; leopard lizard) and Phrynosoma platyrhinos (desert horned lizard) were anomalous. These two species indicate that an open, sparse vegetation community was in existence, possibly along the lake margins, a community not yet thoroughly examined (Mead et al. 1982). The recovery of Mustela nigripes from SCBC seems to corroborate the open, sparse community hypothesis, at least near the lakeshore. Mustela nigripes generally feeds on Cynomys, Spermophilus spp., and lagomorphs (rabbits and pikas), while the closely related M. eversmanni (Siberian polecat; European ferret) eats a variety of small mammals including Ochotona (pika), Microtus, and Marmota (Anderson et al. 1986). Although M. nigripes is currently a near obligate with the prairie dog, in the past the blackfooted ferret may have interacted with other colonial ground squirrels, any of which would require at least some open, possibly grassland

12 D April 1989 Mead, Mead : Snake Creek Burial Cave 153 areas. Interestingly, ground squirrels and lagomorphs (including pikas) are the most abundant species in the SCBC deposit. Is this near-shore area also the community used by the noble marten? SCBC is the only locality in the Great Basin to report the occurrence of both Martes americana and M. nobilis. KM. nobilis is a valid species, it is possible that these two species had overlapping ranges and yet occupied different niches. Unfortunately, the mystery still remains: How did these animals coexist for such a long period of time and what caused the abrupt extinction of M. nobilis 3,000-4,000 years ago? Further, Mustela nivalis currently occupies open woodlands, meadows, and cultivated field areas that during the Pleistocene were presumably in the valley bottoms away from rocky substrates. Assuming the behavior of the least weasel was similar in the past to its current behavior, we must conclude again that some open vegetation communities must have been in existence. We propose that the SCBC deposits and other scattered data (e.g., Crystal Ball Cave) imply that the valley-bottom vegetational communities of the central Great Basin adjacent to the Lake Bonneville beach areas were open, possibly with some grasslands. It is here, in what we would assume was probably a narrow zone along the lake region, that Mustela nigripes, M. nivalis, possibly Martes nobilis, and Brachyprotoma brevimala lived on other open-land species such as certain lagomorphs, colonial ground squirrels, and desert lizards. Further work in the valleybottom habitats will permit a more refined reconstruction. Acknowledgments Financial support for this project was provided by the National Geographic Society (# , JIM and EMM), the Geological Society of America (# , EMM), and the Quaternary Studies Program (Northern Arizona University) Mini-Grant Program (EMM). We are grateful to the Bureau of Land Management (BLM), particularly John Zancanella and Shaaron Netherton (Ely District Office) and Lynda Armentrout (Nevada State Office), for their assistance in obtaining permits and for their enthusiastic support of this project. We especially thank A. Cartwright, D. Cartwright, D. Dawson, S. Emslie, W. J. Marsh, J. W. Marsh, C. Mead, and P. Youngman for their assistance in the field. Bita, Chuck, Goodie, and Dave of Baker, Nevada, provided generous help throughout the project. We thank B. Bonnichsen and M. Sorg (Center for the Study of the First Americans, University of Maine), S. McFarlane (BLM), and D. Tuohy (Nevada State Museum). The American Museum of Natural History personnel (New York, New York) and the United States National Museum, particularly B. Fisher (Washington, D.C.), greatly aided our study of modern species. We appreciate the support of L. Agenbroad (Quaternary Studies Program, Northern Arizona University) and B. Foust and H. Hooper (Bilby Besearch Center, Northern Arizona University). Literature Cited AJKENS, C M Hogup Cave. University of Utah Anthropological Papers 93, Salt Lake City. Anderson. E Quaternary evolution of the genus Martes (Carnivora, Mustelidae). Acta Zoologica Fennica 130: A survey of the late Pleistocene and Holocene mammal fauna of Wyoming. Pages z';i M. Wilson, ed., Applied geology and archaeology: the Holocene history of Wyoming. Geological Survey of Wyoming, Rept. of Investigations No Pleistocene Mustelidae (Mammalia, Carnivora) from Fairbanks, Alaska. Bull. Mus. Comp. Zool. 148: Andekson. E. S C Forrest, T. W. Clark, and L. Richardson Paleobiology, biogeography, and systematica of the black-footed ferret, Mustela nigripes (Audubon and Bachman), Great Basin Nat. Mem. 8: Barker. M S, Jr., andt L. Best The wolverine (Cult) luscus) in Nevada. Southwestern Nat. 2: 133. BROWN, B The Conard Fissure, a Pleistocene bone deposit in northern Arkansas: with description of two new genera and twenty new species of mammals. Amer. Mus. Nat. Hist. 9: Brown, J H Mammals on mountaintops: nonequilibrium insular biogeography. Amer. Nat. 105: The theory of insular biogeography and the distribution of boreal birds and mammals. Great Basin Nat. Mem. 2: Clark, T W, E. Anderson, C. Douglas, and M. Strickland Martes americana. Mamm. Species 289: 1-8. Ewer, R. F The carnivores. Cornell University Fowler, D D, Press, Ithaca, New York. B Madsen, and E. M. Hattori Prehistory of southeastern Nevada. Desert Research Institute Publications in Social Sciences No. 6, Reno, Nevada.

13 and H and and 154 Great Basin Naturalist Vol. 49, No. 2 GODIN, A J Striped and hooded skunks. Pages in J. A. Chapman and G. A. Feldhamer, eds., Wild mammals of North America. The Johns Hopkins University Press, Baltimore. Goodrich, R. B The Quaternary mammalian microfaunal assemblage of Smith Creek Cave, Nevada. Unpublished thesis, California State University at Los Angeles. Grayson D. K The paleontology of Gateeliff Shelter: small mammals. Pages in D. H. Thomas, The archaeology of Monitor Valley 2. Gateeliff Shelter. Anthropological Papers of the American Museum of Natural History 59, New York The paleontology of Hidden Cave: birds and mammals. Pages in D. H. Thomas, ed.. The archaeology of Hidden Cave, Nevada. Anthropological Papers of the American Museum of Natural History 61, New York The biogeographic history of small mammals in the Great Basin: some observations on the last 20,000 years. J. Mammal. 68: Danger Cave, Last Supper Cave, and Hanging Rock Shelter: the faunas. Anthropological Papers of the American Museum of Natural History 66, New York. Grayson. D. K, P W. Parmalee Hanging Rock Shelter. Pages in D. K. Grayson, Danger Cave, Last Supper Cave, and Hanging Rock Shelter: the faunas. Anthropological Papers of the American Museum of Natural History 66, New York. Hall, E. R The mammals of North America. Vol. II. 2d ed. John Wiley and Sons, New York. Heaton, T. H Quaternary paleontology and paleoecology of Crystal Ball Cave, Millard County, Utah: with emphasis on mammals and description of a new species of fossil skunk. Great Basin Nat. 45: Howard. W. E., and R E. Marsh Spotted and hog-nosed skunks. Pages in J. A. Chapman and G. A. Feldhamer, eds., Wild mammals of North America. The Johns Hopkins University Press, Baltimore. Hunt, C B Physiography of the United States. W. H. Freeman and Company, San Francisco. Jennings, J. D Danger Cave. University of Utah Anthropological Papers 27, Salt Lake City. Kuchler, A W Potential natural vegetation map. USGS, scale 1:7,500,000. Kurten, B., and E. Anderson Pleistocene mammals of North America. Columbia University Press, New York. Lindzey. F. G Badger. Pages in J. A. Chapman and G. A. Feldhamer, eds., Wild mammals of North America. The Johns Hopkins University Press, Baltimore. Linscombe, G, N. Kinler. and R J Aulerich Mink. Pages in J. A. Chapman and G. A. Feldhamer, eds., Wild mammals of North America. The Johns Hopkins University Press, Baltimore. McGuire. K R Cave sites, faunal analysis, and big-game hunters of the Great Basin: a caution. Quaternary Res. 14: Mead. J I R S. Thompson, and T R Van Devender Late Wisconsinan and Holocene fauna from Smith Creek Canyon, Snake Range, Nevada. Trans. San Diego Soc. Nat. Hist. 20: MILLER, S J The archaeological fauna of four sites in Smith Creek Canyon. Pages in D. R. Tuohy and D. L. Rendall, eds., The archaeology of Smith Creek Canyon, eastern Nevada. Nevada State Museum Anthropological Papers 17, Carson City. Nowak R M. J L. Paradiso Walker's mammals of the world. Vol. II. 4th ed. The Johns Hopkins University Press, Baltimore. Spiess. A Faunal remains from Bronco Charlie Cave (26EK801), Elko County, Nevada. Pages in L. Casjens, The prehistoric human ecology of the southern Ruby Valley, Nevada. Unpublished dissertation, Harvard University, Cambridge, Massachusetts. Strickland. M A. C W Douglas, M. Novak, and N P Hunziger Marten. Pages in J. A. Chapman and G. A. Feldhamer, eds., Wild mammals of North America. The Johns Hopkins University Press, Baltimore. Svendsen. G E1982. Weasels. Pages in J. A. Chapman and G. A. Feldhamer, eds., Wild mammals of North America. The Johns Hopkins University Press, Baltimore. Thompson. R S, L. Benson, and E M Hattori A revised chronology for the late Pleistocene lake cycle in the central Lahontan Basin. Quaternary Res. 25: 1-9. Thompson, R S, J I Mead Late Quaternary environments and biogeography of the Great Basin. Quaternary Res. 17: Turnmire, K L An analysis of the mammalian fauna from Owl Cave One and Two, Snake Range, east-central Nevada. Unpublished thesis, University of Maine, Orono. Wells, P. V Paleobiogeography of montane islands in the Great Basin since the last glaciopluvial. Ecol. Monogr. 53: White, J A, G McDonald, E Anderson, and J. M. Soiset Lava blisters as carnivore traps. Pages in H. H. Genoways and M. R. Dawson, eds., Contributions in Quaternary vertebrate paleontology: a volume in memorial to John E. Guilday. Carnegie Museum of Natural History Special Publication 8, Pittsburgh. Unmodified mammal and bird re- Ziegler, A. C mains from Deer Creek Cave, Elko County, Nevada. Pages in M. E. Shutler and R. Shutler, Deer Creek Cave, Elko County, Nevada. Nevada State Museum Anthropological Papers 11, Carson City.