The quaternary paleontology and paleoecology of crystal ball cave, millard county, utah: with emphasis on the mammals and the description of a new species of fossil skunk

Family Bovidae

Antilocapra americana

Material--Partial left maxilla with M1/,2/,3/ (BYUVP 7656).

Discussion--The M3/ was distinguished from Odocoileus by being very hypsodont and having a more pointed posterior end as in Antilocapra. It is identical in size and proportions to the largest male specimen of A. americana available for comparison and distinctly larger than the extinct Pleistocene antilocaprids.

Since A. americana presently lives around Gandy Mountain in small herds, it is not surprising to find it in the assemblage. But it is not well represented as a fossil, suggesting that Snake Valley has not always been the treeless desert that it is now. Since Odocoileus hemionus is the dominant artiodactyl in the fossil assemblage while Antilocapra americana is the dominant living artiodactyl in the area, Antilocapra americana must have become abundant in the area in Recent times at the expense of Odocoileus hemionus.

Ovis canadensis

Material--Posterior portion of lower right jaw with M/1,/2,/3 (LACM 123695) and posterior portion of lower left jaw with M/3 (LACM 123696, probably from the same individual), left magnum (BYUVP 7780).

Discussion--The molars of LACM 123695 and 123696 are distinctly larger and more robust than Ovis aries and are even slightly larger than Recent Ovis canadensis. This suggests that the jaws are Pleistocene rather than Recent in age because Harris and Mundel (1974) demonstrated that O. canadensis became reduced in size at the end of the Pleistocene.

Pleistocene bighorn sheep are commonly found in assemblages in the Great Basin (Hibbard and Wright 1956, Stokes and Condie 1961). Even in historic times they have been reported natively in the Snake Range (Durrant 1952, Hall 1946, 1981). O. canadensis was temporarily lost from the Snake Range but was reintroduced in the middle 1900s and presently thrives in the higher elevations (Mead et al. 1982). Shortly after this reintroduction, one young ram lived on Gandy Mountain for several months (J. C. Bates 1983 personal communication), but this is the only citing I know of at such a low elevation in the area.

Ovis Canadensis is the best represented ungulate in the Smith Creek Cave assemblage, and Oreamnos harringtoni is also well represented (Miller 1979). No Oreamnos material has been identified from Crystal Ball Cave, and Ovis is less represented than horse, camel, and deer. This difference between the two assemblages is probably because wild goats and sheep are mountainous animals and would rarely venture into Snake Valley. It may also represent the fact that Smith Creek Cave was a shelter for humans since many Ovis fossils found there appear butchered (Miller 1979).

Ovis cf. aries

Material--Right metacarpal and two first phalanges found associated (BYUVP 8300).

Discussion--These associated bones were found as float near the east entrance of Crystal Ball Cave, and their greasy appearance suggests that they are Recent. The length and shape of the metapodial demonstrates that it is of the genus Ovis, and it is slightly longer than the O. aries specimens to which it was compared but distinctly smaller than living O. canadensis. O. aries is now a common domestic animal in the area, and many roam on Gandy Mountain each winter (J. C. Bates 1984 personal communication).

Since this species is a Recent introduction from Europe, its presence has little significance to this study. It does show, however, that the smaller bones of large mammals are still being deposited in Crystal Ball Cave, probably by woodrats since gates on the cave entrances would keep out all but the smallest carnivores. These specimens were found just north of the east entrance, an area where woodrats and their nests are often found.

cf. Symbos cavifrons

Material--Second phalanx (BYUVP 7923), distal portion of second phalanx (BYUVP 7924), 2 partial second phalanges (BYUVP 7925, 7926), 2 distal portions of second phalanges (BYUVP 7921, 7922).

Discussion--These short, broad second phalanges compare best among living species to Ovibos moschatus but are slightly longer and narrower. BYUVP 7923 is the most complete specimen, missing only one side of the distal extension. It has a length of 42 mm, a proximal transverse width of 27 mm, and a proximal anteroposterior width of 26 mm. BYUVP 7924 has the same proximal measurements as BYUVP 7923, and BYUVP 7925 has a proximal anteroposterior width of at least 26 mm. The distal ends taper in such a way that they are hard to measure. The general shape of these second phalanges shows that they are from an animal more closely related to Ovibos than any other living bovid. Few phalanx measurements of Pleistocene oxen are available, but Nelson and Madsen (1980) and Stokes and Hansen (1937) reported abundant isolated Symbos cavifrons and Bootherium bombifrons crania from Lake Bonneville deposits, and McGuire (1980) reported Euceratherium from a late Pleistocene deposit in central Nevada.

Kurten and Anderson (1980) described Symbos cavifrons as being taller and more slender than Ovibos moschatus, and this description matches the difference between the Crystal Ball Cave specimens and Ovibos moschatus perfectly. Bootherium is smaller than Symbos and is thought by many to represent females or juveniles of that genus (Kurten and Anderson 1980, Nelson and Madsen 1980). Euceratherium was larger and more heavily built than Ovibos (Kurten and Anderson 1980), and a first phalanx illustrated by McGuire (1980) is far too big at the distal end to match the second phalanges from Crystal Ball Cave. So although no comparative material was available, both the description and known range of Symbos cavifrons make the Crystal Ball Cave specimens most referable to that species.

CONCLUSIONS

The Crystal Ball Cave assemblage is the first late Wisconsinan fauna reported from the state of Utah and represents the closest known terrestrial fossil deposit to Lake Bonneville. The assemblage differs from most other cave faunas by having its fossils far inside the cave where man and birds probably had no influence on what was deposited. As a result, the assemblage is better than average in representing the proportions of animals that lived in the area, but there are some obvious biases. Neotoma, always an animal of low density, was the second most abundant genus in the assemblage simply because it is one of the few animals that lives in the cave. But other than cave-dwelling species, the assemblage probably gives a fairly good record of the abundance of most groups, at least those which lived in the immediate vicinity of the cave. The assemblage, for example, contains a ratio of small mammals to large mammals and carnivores to herbivores that might be expected in a living community. One very strong bias is the size of bones in the assemblage that I have attributed to the limit of bone size that a wood rat can carry. Bones of large mammals were brought in after the carcasses deteriorated, as evidenced by the presence of only small isolated elements. This bias tends to make large species less represented in the assemblage than in the living community and very large species unrepresented. Proboscidian fossils are common in Lake Bonneville deposits (Nelson and Madsen 1980) but are unrepresented at Crystal Ball Cave, probably because there was no means to transport such large bones inside.

It is difficult to say if any other animals besides wood rats contributed to transporting fossils into the cave. No other rodents are known to transport bones as wood rats do. Small carnivores could have done so, but the low abundance of carnivore fossils in the assemblage suggests that none habitually used the cave as a home. The small size of the original cave entrance would have prevented the entry of any large mammals. Both the distance of the fossils inside the cave and the low abundance of birds compared to mammals suggests that birds did not transport any fossils in, and this is one of the main differences between Crystal Ball Cave and Smith Creek Cave (and most other cave deposits). Clearly no inorganic processes such as wind, water, or gravity could have been responsible for the fossil deposits since they are in fine dust in an isolated part of the cave where none of these forces had a magnitude capable of transporting bones.

Crystal Ball Cave has been accumulating fossils from at least 23,000 years ago to the present. Although some of the fossils are Recent, the assemblage as a whole shows dramatic differences from the present-day local fauna. The poor representation of many mammals that currently live in the area may be due to the shift from Neotoma cinerea to N. lepida as the wood rat that inhabited the cave, and it also suggests that the shift to the present climate occurred very recently in the history of the assemblage. Brachyprotoma, Smilodon, several species of Equus, Camelops, Hemiauchenia, and Symbos (or a closely related genus) are represented in the assemblage, all of which are now extinct. As mentioned earlier, there was a widespread extinction of large mammals at the close of the Pleistocene, the cause of which is under debate. This assemblage does not resolve that problem, but it does demonstrate that a marked climatic shift did take place contemporaneously with the extinctions, and this suggests to me that the extinctions were also a result of this climatic shift.

Equally as significant as the extinctions are the shifts in species ranges that the Crystal Ball Cave assemblage documents. The presence of Ondatra zibethicus and Mustela cf. vison, both of which require perennial water and are extirpated from the area, represent the drying of Lake Bonneville and perennial streams around Gandy Mountain. Ochotona princeps and Martes americana were extirpated from the Snake Range without replacement but still live at high elevations in nearby ranges. Marmota flaviventris, Cervus elaphus, and Ovis canadensis are represented in the assemblage but now inhabit only higher elevations in the Snake Range.

In other cases species now abundant at Gandy Mountain are unrepresented or poorly represented in the assemblage while their more boreal counterparts, now extirpated or rare in the area, are well represented as fossils. Among jack rabbits, Lepus californicus is presently the dominant species while L. townsendii, its more boreal counterpart, is by far the better represented species in the fossil assemblage. Among cottontails, Sylvilagus audubonii and S. nuttallii make up the present local fauna, but only S. nuttallii, the more northern species, is found in the assemblage. Lepus americanus, a functional cottontail (J. A. White 1984 personal communication) and a very boreal animal, is probably represented but is now extirpated from the Snake Range. Neotoma lepida, the only wood rat seen living in Crystal Ball Cave, is rare in the assemblage while N. cinerea, its more boreal counterpart, is one of the two most abundant fossil species. Vulpes vulpes is well represented in the cave assemblage but extirpated from the area while Urocyon cinereoargenteus, a more southern fox of similar size, now inhabits the area but is not found as a fossil.

Although the fossil assemblage differs dramatically from the present-day local fauna, it is not atypical of late Pleistocene assemblages in the region. Figure 9 shows the location of and table 10 compares the fauna of 10 late Pleistocene-Recent cave assemblages within 240 miles (400 km) of Crystal Ball Cave. The most unique feature of the Crystal Ball Cave assemblage is the presence of Brachyprotoma since it represents the first citing of the genus from the western United States and the first recovery of the new species herein named B. brevimala. Ondatra zibethicus was found in Crystal Ball Cave but not at the other localities, probably because of this cave's close proximity to Lake Bonneville. Symbos cavifrons may be present at Crystal Ball Cave but absent from the other assemblages for the same reason since it is most common in Lake Bonneville deposits.

Some interesting paleoecological information can be inferred from the differences between the Smith Creek Cave and Crystal Ball Cave assemblages in particular since they are close geographically but located in somewhat different habitats. Several species of Spermophilus have been recovered from Smith Creek Cave, but large numbers of a single species have been recovered from Crystal Ball Cave. This can probably be attributed to the greater habitat diversity at Smith Creek Cave, which is at the base of a high mountain. Among camels, Hemiauchenia is better represented at Smith Creek Cave but Camelops is better represented at Crystal Ball Cave. This suggests that Hemiauchenia favored higher and/or more rugged terrain than Camelops because Smith Creek Cave is located in the main Snake Range while Crystal Ball Cave is located in an outlier in Snake Valley. Of the non-camelid artiodactyls, Odocoileus hemionus is the best represented in the Crystal Ball Cave assemblage and Ovis canadensis is the best represented in the Smith Creek Cave assemblage. Oreamnos harringtoni fossils have been found in Smith Creek Cave but not in Crystal Ball Cave. Now Antilocapra americana is the best represented artiodactyl in Snake Valley, Odocoileus hemionus is the best represented artiodactyl in the Snake Range, Ovis canadensis is found only at high elevations in the Snake Range, and Oreamnos harringtoni is extinct. This suggests that these four artiodactyls can be placed in the following order of elevation preference starting at the highest: Oreamnos harringtoni, Ovis canadensis, Odocoileus hemionus, and Antilocapra americana. At the end of the Pleistocene, in rough terms, each of these species moved upward in elevation to fill the habitat of the next higher species. The one at the top went extinct; the one at the bottom became abundant. Differences of lesser magnitude between the Crystal Ball Cave and Smith Creek Cave assemblages must be dealt with more carefully because they may represent slight differences in the age of the deposits, biases in the mode of deposition, human intervention, or chance preservation. Identification of more material, especially at Smith Creek Cave, could make comparison of these two assemblages a very valuable paleoecological study.

The Crystal Ball Cave fauna, like many previously studied faunas, shows that a dramatic climatic shift occurred at the end of the Pleistocene and caused of many species to move northward in latitude and upward in elevation and to become extinct. This shift is particularly well expressed in the Crystal Ball Cave assemblage because its close proximity to Lake Bonneville made the drying trend very severe in the area. The Crystal Ball Cave fauna documents the previous ranges and abundances of many taxa that helps in reconstruction of details of the last Pleistocene ice age.
ACKNOWLEDGMENTS

This study was supervised by Wade E. Miller who helped in collecting and identifying specimens and preparing the manuscript. His insistence that every identification be backed by thorough research and explanation has made a lasting impression on me. Thanks is also due Kenneth L. Stadtman and Clyde L. Pritchett for providing comparative specimens and help in identification. Jerald C. and Marlene Bates of Gandy allowed me access to Crystal Ball Cave on many occasions and provided helpful information on the history and original condition of the cave, as well as the mammals that currently live in the immediate area. The Los Angeles County Museum generously loaned me the Crystal Ball Cave specimens in their possession so they could be included in this study.

Howard C. Stutz identified the plants, Lee F. Braithwaite the gastropods, and Stephen L. Wood the beetle. John A. White provided information valuable for identifying the lagomorphs. Elaine Anderson provided information helpful in evaluating the Brachyprotoma skull. Phillip M. Youngman provided unpublished information and measurements on Brachyprotoma specimens he recovered from Yukon Territory, Canada. Arthur H. Harris provided bone measurements for several species of horses. Theodore E. Downs gave me information on Pleistocene horses and measurements of Pleistocene camels. Jim I. Mead identified some of the bovid specimens and provided other helpful encouragement. Wade E. Miller, J. Keith Rigby, Morris S. Petersen, Lehi F. Hintze, Elaine Anderson, and Phillip M. Youngman made critical reviews of the manuscript. Thanks is especially due my wife, Julie, for help with collection and curation of specimens, gathering of literature, and preparation of the manuscript.

This study was funded by grants from the National Speleological Society and Associated Students of Brigham Young University, by a private donation from Herbert H. Gerisch, and by Brigham Young University research assistantships to the author. Publication costs were paid for by the Joseph M. and Jessie K. D. Savage Endowment of the Brigham Young University Monte L. Bean Museum and the Brigham Young University College of Physical and Mathematical Sciences.

REFERENCES CITED

Anderson, E. 1968. Fauna of the Little Box Elder Cave, Converse County, Wyoming. Univ. Colorado Stud. Earth Sci. Ser. 6:1-59.

________. 1970. Quaternary evolution of the genus Martes (Carnivora, Mustelidae). Acta. Zool. Fennica 130:1-132.

Bjork, P. R. 1970. The carnivora of the Hagerman local fauna (late Pliocene) of southwestern Idaho. Amer. Phil. Soc. Trans. 60(7):1-54.

Brattstrom, B. H. 1976. A Pleistocene herpetofauna from Smith Creek Cave, Nevada. South. California Acad. Sci. Bull. 75:283-284.

Brown, B. 1908. The Connard Fissure, a Pleistocene bone deposit in Northern Arkansas: with description of two new genera and twenty new species of mammals. Amer. Mus. Nat. Hist. Mem. 9(4):157-208.

Brown, J. H. 1971. Mammals on mountaintops: nonequilibrium insular biogeography. Amer. Nat. 105(945):467-478.

________. 1978. The theory of insular biogeography and the distribution of boreal mammals. Great Basin Nat. Mem. 2:209-227.

Bryan, A. L. 1979. Smith Creek Cave. In The archaeology of Smith Creek Canyon, Eastern Nevada. Tuohy, D. R. and D. L. Rendall (eds.) Nevada State Mus. Anthrop. Pap. 17:162-251.

Chamberland, R. V., and D. T. Jones. 1929. A descriptive catalog of Mollusca of Utah. Univ. Utah Biol. Ser. Bull. 1(1):1-203.

Cope, E. D. 1899. Vertebrate remains from the Port Kennedy bone deposit. Philadelphia Acad. Nat. Sci. J. 11:193-267.

Cragin, F. W. 1892. Observations on llama remains from Colorado and Kansas. Amer. Geol. 9:257-260.

Currey, D. R. 1982. Lake Bonneville: selected features of relevance to neotectonic analysis. U.S. Geol. Surv. Open-File Rept. 82-1070:1-29.

Dalquest, W. W. 1964. Equus scotti from a high terrace near Childress, Texas. Texas J. Sci. 16(3):350-358.

________. 1979. The little horses (genus Equus) of the Pleistocene of North America. Amer. Midland Nat. 101(1):241-244.

________, and J. T. Hughes. 1965. The Pleistocene horse, Equus conversidens. Amer. Midland Nat. 74(2):408-417.

Davies, W. E. 1960. Origin of caves in folded limestone. Nat. Speleo. Soc. Bull. 22:5-19.

Davis, W. M. 1930. Origin of limestone caverns. Geol. Soc. Amer. Bull. 41:475-628.

Durrant, S. D. 1952. Mammals of Utah. Univ. of Kansas Press, Lawrence, 1-549.

________, M. R. Lee, and R. M. Hansen. 1955. Additional records and extensions of known ranges of mammals from Utah. Univ. Kansas Mus. Nat. Hist. Pub. 9(2):69-80.

Finley, R. B., Jr. 1958. The wood rats of Colorado: distribution and ecology. Univ. Kansas Mus. Nat. Hist. Pub. 10(6):213-552.

Gazin, C. L. 1935. Annotated list of Pleistocene mammalia from American Falls, Idaho. Washington Acad. Sci. J. 25(7):297-302.

________. 1936. A study of the fossil horse remains from the Upper Pliocene of Idaho. U.S. Nat. Mus. Proc. 83(2985):281-320.

Gidley, J. W. 1900. A new species of Pleistocene horse from the Staked Plains. Amer. Mus. Nat. Hist. Bull. 13:111-116.

________. 1903. The freshwater Tertiary of northwestern Texas, American Museum expedition of 1899-1901. Amer. Mus. Nat. Hist. Bull. 19:617-635.

________, and C. L. Gazin. 1938. The Pleistocene vertebrate fauna from Cumberland Cave, Maryland. U.S. Nat. Mus. Bull. 171:1-99.

Gilbert, G. K. 1890. Lake Bonneville. U.S. Geol. Surv. Mon. 1:1-438.

Giles, E. 1960. Multivariate analysis of Pleistocene and Recent coyotes (Canis latrans) from California. Univ. California Geol. Sci. Pub. 36:369-390.

Goodrich, R. B. 1965. The Quaternary mammalian microfaunal assemblage of Smith Creek Cave, Nevada. Unpublished thesis. California State Univ., Los Angeles, 1-45.

Graham, R. W. 1981. Preliminary report on late Pleistocene vertebrates from the Selby and Dutton Archeological/Paleontological Sites, Yuma County, Colorado. Univ. Wyoming Contrib. Geol. 20(1):33-56.

Grayson, D. K. 1977. On the Holocene history of some northern Great Basin lagomorphs. J. Mammal. 58(4):507-513.

________. 1983. Small mammals. In The archaeology of Monitor Valley: 2, Gatecliff Shelter. Thomas, D. H. (ed.) Amer. Mus. Nat. Hist. Anthrop. Pap. 59(1):99-126.

Green, D. J. 1961. Introduction to the speleology of Utah and adjacent areas. Nat. Speleo. Soc. Salt Lake Grotto Tech. Notes 2(60):192-201.

Gruhn, R. 1961. The archaeology of Wilson Butte Cave, south-central Idaho. Idaho State Mus. Occas. Pap. 6.

Hall, E. R. 1936. Mustelid mammals from the Pleistocene of North America; with systematic notes on some Recent members of the genera Mustela, Taxidea and Mephitis. In Studies of Tertiary and Quaternary mammals of North America. Carnegie Inst. Washington Contrib. to Paleont. 473:41-119.

________. 1946. Mammals of Nevada. Univ. California Press, Berkeley, 1-710.

________. 1951. A synopsis of the North American lagomorpha. Univ. Kansas Mus. Nat. Hist. Pub. 5(10):119-202.

________. 1981. The Mammals of North America, 2nd Edition. John Wiley and Sons, New York, 1-1181.

Halliday, W. R. 1957. The caves of Gandy Mountain, an initial reconnaissance. Nat. Speleo. Soc. Salt Lake Grotto Tech. Notes 2(38):31-37.

Handley, C. O., Jr. 1959. A revision of American bats of the genera Euderma and Plecotus. U.S. Nat. Mus. Proc. 110(3417):95-246.

Harington, C. R. 1972. Extinct animals of Rampart Cave. Canadian Geog. J. 85(5):178-183.

Harper, K. T., D. C. Freeman, W. K. Ostler, and L. G. Klikoff. 1978. The flora of Great Basin mountain ranges: diversity, sources, and dispersal ecology. Great Basin Nat. Mem. 2:81-103.

Harrington, M. R. 1934. Ancient horses and ancient Men in Nevada. Masterkey 8:165-169.

Harris, A. H., and P. Mundel. 1974. Size reduction in bighorn sheep (Ovis canadensis) at the close of the Pleistocene. J. Mammal. 55(3):678-680.

Harris, A. H., and L. S. W. Porter. 1980. Late Pleistocene horses of Dry Cave, Eddy County, New Mexico. J. Mammal. 61(1):46-65.

Hay, O. P. 1915. Contributions to the knowledge of the mammals of the Pleistocene of North America. U.S. Nat. Mus. Proc. 48:515-575.

________. 1921. Descriptions of Pleistocene vertebrata, types of specimens of which are preserved in the United States National Museum. U.S. Nat. Mus. Proc. 59:617-638.

________. 1923. The Pleistocene of North America and its vertebrated animals from the States east of the Mississippi River and from the Canadian Provinces east of Longitude 95. Carnegie Inst. Washington Pub. 322:1-499.

Heaton, T. H. 1984. Preliminary report on the Quaternary Vertebrate fossils from Crystal Ball Cave, Millard County, Utah. Cur. Res. 1:65-67.

Hibbard, C. W. 1941. New mammals of the Rexroad Fauna from the Upper Pliocene of Kansas. Kansas Acad. Sci. Trans. 44:265-313.

________. 1944. Abnormal tooth pattern in the lower dentition of the jackrabbit, Lepus californicus deserticola (Mearns). J. Mammal. 25:64-66.

________. 1950. Mammals of the Rexroad Formation from Fox Canyon, Kansas. Univ. Michigan Mus. Paleontol. Contrib. 8(6):113-192.