Occurrence of the Musk Ox, Symbos Cavifrons, from Southeastern Idaho and Comments on the Genus Bootherium

Great Basin Naturalist

Volume 47 Number 2 Article 9

4-30-1987

Occurrence of the musk ox, Symbos cavifrons, from southeastern Idaho and comments on the genus Bootherium

Michael E. Nelson Fort Hays State University, Hays, Kansas

James H. Madsen Jr. Antiquities Section, Utah Division of State History, Salt Lake City

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Recommended Citation Nelson, Michael E. and Madsen, James H. Jr. (1987) "Occurrence of the musk ox, Symbos cavifrons, from southeastern Idaho and comments on the genus Bootherium," Great Basin Naturalist: Vol. 47 : No. 2 , Article 9. Available at: https://scholarsarchive.byu.edu/gbn/vol47/iss2/9

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 [email protected], [email protected]. OCCURRENCE OF THE MUSK OX, SYMBOS CAVIFRONS, FROM SOUTHEASTERN IDAHO AND COMMENTS ON THE GENUS BOOTHERIUM

Michael E. Nelson' and James H. Madsen, Jr."

Abstract.—A set of ovibovine horn cores collected from Pleistocene sediments in southeastern Idaho provides additional evidence for sexual dimorphism in the helmeted musk ox, Symhos cavifrons. Specimens previously assigned to Boothchum sargenti are placed in synonomy with Symbos cavifrons as sexual dimorphs (females). Bootherium bomhifrons is a \alid taxon and is probably not closely related to Symbos.

The taxonomic status of the e.xtinct musk figures and descriptions, pronounced that ox, Bootherium, has been questioned since Bootherium cavifrons (= Symbos cavifrons) Leidy (1852a, 1852b) proposed the name for was a male and B. bomhifrons a female. He two extinct species of ovibovines, B. bomh- then assigned these specimens to a new taxon, ifrons (Bos bomhifrons of Harland 1825, Wis- Ovihos priscus. Dawkins (1872) came to a sim- tar 1818) and B. cavifrons. Later, Dawkins ilar conclusion but chose to call the duo Ovi- (1867) and then Leid\' (1869) synonvmized hos cavifrons. Lydekker (1885, 1898) was also these extinct forms with the extant musk ox, convinced that B. bomhifrons and B. cav-

Ovihos. Osgood (1905a) erected the genus ifrons { =S. cavifrons) were sexual dimorphic Scaphoceros for a musk ox, S. ttjrreUi, col- forms of the same species. lected from the Yukon Territory of Canada but Osgood (1905a), as Allen (1913) so aptly later discovered that Scaphoceros was preoc- stated, cupied; he therefore replaced it with the

set forth the real facts of the case. . . . [S\\nce bombifrons name Symbos (1905b). In addition, Osgood and cavifrons have been considered by several authors as (1905a, 1905b) observed that Symhos cav- being not only congeneric, but conspecific, the establish- ifrons and Bootherium bomhifrons were dis- ment of a separate genus for each ma\' appear surprising. While it may be possible, from an examination of the tinct species and definitely separable from the figures only, to construct a hypothesis to the effect that extant Ovihos moschatus. Gidley (1908) cavifrons represents the male and bombifrons the female named a second species of Bootherium, B. of one species, it is inconceivable that any modern tax- sarg^enti, from a late-Pleistocene deposit in onomist would reach such a conclusion after comparing Michigan. the original types.

Taxonomists (see Kurten and Anderson Allen (1913) certainly agreed when he stated. 1980) now generally agree that all extant musk Upon carefully reading Osgood's paper I felt sure that he ox belong to Ovihos moschatus and that the had correctly soKed the problem. various proposed species q{ Symhos, e.g., S. tyrrelli Osgood, S. austraUs Brown, Liops zu- Evidently Hay (1924) did not believe the niensis Gidley (preoccupied, changed to problem had been solved and did "not accept Lissops zuniensis Gidley), and Gidleya zu- this opinion [of Allen and Osgood]." Hay be- niensis (Cossman) are all assignable to Syjnhos lieved that some of the 25 skulls of Symbos cavifrons (for a different opinion see McDon- cavifrons that he examined "must have been ald 1985). However, the taxonomic status of females. Bootherium has remained in question. Bison appalachicohis, originally named by Confusion surrounding Bootherium has Rhoads (1895), was assigned to Bootherium arisen from the supposed sex of various appalachicohis by Ray (1966a) and described Bootherium and Symbos skulls. Is as "closely related, if not conspecific" with Bootherium a female Sijmbos? Rutimever Bootherium sargenti. Bootherium nivicolens (1867), on the basis of Leidy's (1852a, 185'2b) was erected b\- Hay (1915) for a specimen from

Department of Earth Sciences and Sternberg Memorial Museum. Fort Hays State University. Hays, Kansas 67601-4099 -Antiquities Section, Utah Division of State Historv'. 300 Rio Grande, Salt Lake City, Utah 84101.

239 240 Great Basin Naturalist Vol. 47, No. 2

Fig. 1. Stjmbos cavifrons, UVP083; A, Dorsal view of horn cores. B, Posterodorsal view. Bar represents 10 cm.

Eschscholtz Bay, Alaska, while Hesse (1942) the specific taxonomy (not to mention the generic status) named Bootherium brazosis from Brazos o{ Bootherium is not yet well understood. County, Texas. The latter has been identified Nelson and Madsen (1978) documented the by Ray (1966b) as conspecific with Boother- existence of 21 individual musk ox from late- ium sargenti. Pleistocene sediments of Utah. They assigned Hibbard and Hinds (1960) reported. the four specimens of Bootherium to It is very likely that Bootherium is the female woodland Bootherium sp. indet., although Stokes and musk ox since all specimens ofSijmhos based on skulls are Hansen (1937) had previously assigned one considered those of bulls. specimen (BYUG 834) to B. bombifrons. Evidently Hibbard and Hinds examined the Harington (written communication 1978) type of B. sargenti and made their decision on synonymized Bootherium nivicolens with B. the basis of this specimen. It is not known if sargenti. He also believed that B. sargenti they intended to also include B. bombifrons was probably the female oi Symbos cavifrons as a female Symbos. because the differences between the speci- Semken et al. (1964) reviewed the litera- mens parallel the morphological differences ture and decided between male and female specimens of Ovi-

that Bootherium bombifrons represents a genus distinct bos moschatus. from Symbos. However, B. sargenti, the specimen exam- In Kurten and Anderson's (1980) summa- ined by Hibbard and Hinds (1960), may be a female rization of Pleistocene mammals of North Symbos. America, Bootherium sargenti and Symbos

Ray et al. (1967) excavated an ovibovine cranium eavifrons were synonymized, while the type from Saltville, Virginia, that was assigned to specimen of B. bombifrons was questionably Bootherium sp. indet., and stated that left as a separate species. April 1987 Nelson, Madsen: Fossil Musk Ox 241

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Fig. 2. Symbos cavifrons , UUVP8540, left lateral view. Bar represents 10 cm.

White (personal communication 1984, F;AM- Frick Collection, American Museum of Natural 1985) reviewed the musk ox of Idaho and con- Historv cluded that two cranial fragments of Bootherium collected from Pleistocene sedi- Collection Background ments in Idaho (IMNH 68001; LACM 6671) could not be assigned with confidence to es- In 1980 Chris and Jack Dukes of Salt Lake tablished species. However, the size of the City, Utah, brought a partial cranium with a cranial fragment (LACM 6671) collected from complete, although broken, set of horn cores

Minidoka Dam "suggests it may be referable to the Antiquities Section of the Utah Division " to B. sargenti. of State History. A laboratory assistant labeled the specimen "Bison, collected from Idaho" J. N. McDonald (Radford University) is studying a mummified specimen of and entered the specimens into the collec- Bootherium collected from a permafrost local- tions. In examining these collections during ity at Fairbanks Creek, Alaska (F:AM A- the winter of 1985, we recognized the horn 293-5286). Other than a preliminary report cores as ovibovine and not bison. The horn (McDonald 1984), little has been published cores resembled the numerous horn cores of on this interesting specimen. Symbos cavifrons in the Division of State His- Institutional abbreviations are as follows: tory and Utah Museum of Natural History collections, but the frontal area of the cranium

UVP- Utah Division of State Hist()r\ , .\nti<}uities Sec- lacked the characteristic exostosis and median tion, Vertebrate Collection UUVP- University of Utah, \'ertebrate PaleontoloR\ cranial sulcus of Symbos. Comparisons with Collection Utah specimens of Bootherium revealed no IMNH- Idaho Museum of Natural History close resemblances. This discovery prompted KUVP- Kansas University, Vertebrate Paleontoloe;\ a review of all Utah and Idaho musk ox speci- LACM- Los Angeles County Museum of Natural Historx- preparation of GRPM- Grand Rapids Public Museum mens and ultimately led to the ANSP- Academy of Natural Sciences of Philadelphia this report. TAMC- Te.xas A & M Uni\ersity Collection BYUVP, BYUG-Brigham Young University, Wrtebrate Paleontology Collection Age and Str.\tigr.\phy USU- Utah State University, Geology Collection USNM- United States National Museum of Natural His- Precise locality information and strati- tory graphic data were garnered after the discov- 242 Great Basin Naturalist Vol. 47, No. 2

Fig. 3. Bootheriumbomhifrons, USNM 215066 (cast), original ANSP 2994, right lateral view. Bar represents 10 cm. erer of record, Jerry Davis of Soda Springs, Beach section of McDonald and Anderson Idaho, was located. Davis collected the speci- (1975). men 3 March 1977 during the excavation of a Field relationships indicate that the new grain elevator shaft in American Falls, Idaho. musk ox specimen (UVP 083) was collected This elevator, located at the intersection of from the same sediments as the Dam l.f. and is Oregon Trail and Elevator Streets in Ameri- therefore a part of the Dam l.f. can Falls, has the following legal description: NEl/4, Sec 29, T7S, R31E, Power County, Systematic Paleontology Idaho. Order Artiodactyla The Dam local fauna (IMNH locality Family Bovidae 52002), described from a gravel quarry lo- Genus Symbos Osgood 1915 cated between the present American Falls Symbos cavifrons (Leidy) 1852 townsite and the east end of the American Falls Dam (Hay 1927, Gazin 1935, Hopkins Boothehum homhifrons Leidy 1852 (in part) Oiihos aitifron.s Dawkins 1883, Leidy 1869, Lydekker 1951, 1955, Hopkins, Bonnichsen, and 1885, McGee 1887 (in part) Fortsch 1969, Barton 1976, White 1985), was Bison appahicJucoIiis Rhoads 1895 collected approximately 200 m southwest of Ovibos cavifrons Hatcher 1902 the elevator construction site. Among other Scaphoccros tyrrcUi Osgood 1905 {Scaphoceros preoccupied) fossils collected by Gazin (1935), White (1985) Stjmbos tijrrclli Osgood 1905 Gidli'ijo zunicnsis Cossmann 1907 listed a number of specimens oi Symbos cav- Lissops zuniensis Gidle\' 1908 {Liops preoccupied) ifrons obtained from a pit in the American Symbos australis Brown 1908 Falls Formation. A radiocarbon date on in situ Bootlwrititn sar^cnti Gidley 1908 bone fragments from the locality vielded a Ovibos appalacliicolus Staudinger 1908, Allen 1913, Hay Kitts 1953 date of 26,500 ± 3,500 y.b.p. (Barton 1976). 1923, Frick 1937, Bootherium nivicolens Hay 1915 believed the l.f. was correl- White (1985) Dam Symbos promptus Hay 1920 ative with the "B layer" of Hopkins, Bonnich- Symbos convcxifrons Barbour 1934 sen, and Fortsch (1969) and the Rainbow Ovibos iiimntcus Frick 1937 April 1987 Nelson, Madsen: Fossil Musk Ox 243

Fig. 4. Sijmbos cavifrons, UUVP8540, dorsal view. Bar represents 10 cm.

Bootherium hrazosis Hesse 1942 low the orbits. This curvature contrasts with Bootherium appalachicohis Ray 1966 the horn core curvature seen in Bootherium Bootherium sp. indet. Nelson and Madsen 1978 (in part) honihifrons, where the horns flare only Bootherium sp. White 1985 (in part) slightly outward and then drop sharply down-

Materl\l. —UVP 083, cranial cap includ- ward and finally forward (Fig. 3). ing both horn cores. The horn cores in UVP 083 slightly overlap Description. —UVP 083 consists of a com- the frontals as on male Symbos skulls (Fig. 4). plete set of horn cores attached to the frontals However, there is an absence of an exostosis and the cranial cap (Fig. 1). Very little of the and cranial sulcus, and the area on the frontals cranium is preserved anterior or posterior to and parietals between the proximal borders of the attachment of the horn cores. The horn the horn cores is smooth (Fig. 1). A slightly cores curve outward, downward, and forward elevated anterior-posterior ridge extends with a high lateral flare and are very similar to along the midline of the frontals. the curvature on Symbos skulls that are A burr is neither present on the horn cores thought to be males. However, the horn core nor do the horn cores stand out on pedicles as tips of male Sytnbos skulls drop to a position in Bootherium. The horn cores are flattened that nears the base of the skull (Fig. 2). The on the dorsal surface near the proximal bor- horn cores on UVP 083 tend to flare out in a der. This condition is reflected in male Sym- lateral direction and do not extend much be- bos horn cores and stands in contrast with the 244 Great Basin Naturalist Vol. 47, No. 2

Fig. 5. Ovibos moschatits, unnumbered specimen, Alaska Fish and Game, (male) anterodorsal view. Bar represents 10 cm.

rounded horn cores ofBootherium. highly vacuolate bone. Posterior, the entire

The horn cores exhibit a sculpturing that is cranial roof is constructed of finely cancellous very similar, if not identical, to several male bone. This spongy bone is similar to that ob- Symbos specimens in the Utah Division of served on male Symbos skulls but absent in State History and Utah Museum of Natural specimens oiBoothchum bombifrons. History collections. Evident are a series of Discussion. —Harington (fide Kurten and grooves which tend to wrap around in a coun- Anderson 1980:332) believed the evidence terclockwise direction as they descend down supporting the female Symbos designation of the horn core. Although not continuous, the the type specimen of Bootherium sargenti grooves extend to the very tip of the cores quite compelling. He noted, according to

(Figs. 1, 4). Kurten and Anderson,

The thickness of the cranial roof does not the similar basic confirmation of the horn-cores, the vary appreciably anteriorly (62 mm) or poste- smaller size of the cranium, and the broad space between riorly (58 mm) from the horn cores. However, the horn-core bases parallels the differences between male and female Ovibos inoschatus. there is a distinct difference in the type of bone present in the cranial cap. Anterior to The sexual dimorphism in modern speci-

the horn cores, the cranium is composed of mens of Ovibos mosdiatus is clearly noted in April 1987 Nelson, Madsen: Fossil Musk Ox 245

Fig. 6. Ovibos moschatus, unnumbered specimen, Alaska Fish and Game, (female) anterodorsal view. Bar represents 10 cm.

cleaned skulls, although it is not apparent in stockier and somewhat shorter horn cores living specimens. In male specimens of Ovi- than UVP 083 but morphologically agrees in bos "the base of the horn-core is enlarged and all other details. The sculpturing is similar, expanded over a great part of the frontals and the cancellous bone is present, the horn cores parietals, on which large exostoses are devel- slightly overlap the frontals, the frontals are oped" (Lonnberg 1900). However, there is a smooth between the horn cores, and a slightly very characteristic medial groove that sepa- elevated mid-frontal ridge is present. Neas rates the horn cores (Fig. 5). In examining (personal communication 1985) believes that skulls of female O. moschatus, we have found the Natural Trap cave specimen may repre- that the horn cores are much more widely sent a female Symbos. separated at the bases and the frontals are The Wyoming (KUVP 61635) and Idaho nearly smooth between the horn cores. There specimens (UVP 083, LACM 6671) also ap- are no large exostoses developed (Fig. 6). In pear to be very similar, if not almost identical, addition, the horn cores of the females are not to the type oiBootherium sargenti from Mich- as massive as those of the male. This differ- igan (GRPM 114233101) described by Gidley ence then, between male and female O. (1908) (Figs. 8, 9). Therefore, we agree with moschatus specimens, is very similar to the the suggestions of Allen (1913), Hibbard and morphological differences noticed in speci- Hinds (1960), Semken, Miller, and Stevens mens of "Bootherium sargenti" and Symbos (1964), and Kurten and Anderson (1980) and cavifrons. hereby propose that Bootherium sargenti UVP 083 is almost identical to KUVP (GRPM 114233101), B. appalachicolus 61635, a set of horn cores and associated par- (ANSP 29, placed in synonymy with B. sar- tial skull from Natural Trap Cave in north- genti by Ray, 1966a), B. brazosis (TAMC western Wyoming (Neas, personal communi- 2553, placed in svnonymy with B. sargenti by cation 1981) (Fig. 7). This latter specimen has Ray, 1966b), B. nivicolens (USNM 2324, 246 Great Basin Naturalist Vol. 47, No. 2

Fig. 7. Stjmbos cavifrons, KUVP6135, anterodorsal view of horn cores. Bar represents 10 cm.

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I I

Fig. 8. "Bootherium sargenti" { = Symbos cavifrons), USNM 23488 (cast), original GRPM 11 423 3101, dorsal view. Bar represents 10 cm.

placed in synonymy with B. sargenti by Bootherium bombifrons is a valid taxon "pre-

Harington, written communication 1978), suming the type . . . does not represent an and Bootherium sp. (LACM 6671, White abnormal individual." A reexamination of the 1985) be included with Sijmbos cavifrons as Pleistocene specimens from Utah confirms sexual dimorphic forms. Other specimens the taxonomic validity of this determination. would also include KUVP 61635 (Neas, per- USU 3529 is a partial craniurn collected sonal communication 1985), BYUVP 9278 from the late-Pleistocene shoreline deposits (Nelson, unpublished information), and the of Lake Bonneville in northern Utah (Nelson Idaho specimen in this discussion, UVP 083. and Madsen 1978) (Fig. 10). Although the

skull is incomplete, there are sufficient diag-

nostic characters present to assign it to Status of Bootherium Bootheriu7n. The horn cores stand out from Kurten and Anderson (1980) and Harington the skull on a pedicel and display a distinct (written communication 1978) believe that burr at the proximal border. They are April 1987 Nelson, Madsen: Fossil Musk Ox 247

Fig. 9. "Bootheriwn sargenti" { = Syinbos cavifrons), USNM 23488 (cast), original GRPM 11 423 3101, anterodorsal view. Bar represents 10 cm.

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Fig. 10. Bootherium bombifrons, USU 3529, posterodorsal view. Bar represents 10 cm.

rounded in cross section and are unflattened and most importantly, there is a very abrupt dorsally near their bases. An exostosis and posteroventral slope of the dorsal outline of cranial sulcus are lacking, and the frontals the skull posterior to the horn cores. This between the horn cores are smooth. Perhaps, sloping area is a very diagnostic feature on the 248 Great Basin Naturalist Vol. 47, No. 2

Fig. 11. Bootherium bombifrons, USNM 215066 (cast), original ANSP 2994, posterodorsal view. Bar represents 10

Fig. 12. Bootherium bombifrons, BYUG 834, posterodorsal view. Bar represents 10 cm.

type specimen of B. bombifrons; the horn Although the ventral part of this skull is cores seem to be placed on the summit of the abraded, sufficient detail is present to permit skull very similar to the condition in goats and identification of the characteristic basioccipi- sheep, but unlike Syinbos (Fig. 11). tal-basisphenoid area. In Bootherium the April 1987 Nelson, Madsen: Fossil Musk Ox 249

K V

Fig. 13. Bootheriumsp. indet., IMNH 17124, right lateral view. Bar represents 10 cm.

proximal end of the basisphenoid is dorsally homhifrons. UUVP 8532, described by Nel- deflected from the basioccipital at a very low son and Madsen (1978), also appears angle (less than 20 degrees). In Symbos this assignable to B. homhifrons, but post mortem deflection is greater than 35 degrees (White, abrasion of the specimen makes this latter personal communication 1984). identification more tenuous. A second Bootherium skull from Utah A specimen, IMNH 17124, which was col- (BYUG 834) is quite similar to USU 3529 in lected from Bannock Gounty, Idaho, and de- most characteristics (Fig. 12). The single, ma- scribed by White (1985) as Bootherium sp., is jor difference is in the length of the sloping difficult to place specifically (Fig. 13). It ex- skull cap posterior to the horn core attach- hibits all of the characteristics of Bootherium ment. In BYUG 834 this length is only about homhifrons (see White 1985 for a descrip- 90% of the length of USU 3529. However, this tion), with the exception of the length of the difference may be attributable to post mortem skull cap posterior to the horn core attach- abrasion of the skull rather than to pathology, ment (Fig. 14). In IMNH 17124 the cranial individual variation, or sexual dimorphism. cap slopes abruptly downward at a much Both Utah specimens (BYUG 834 and USU greater angle than in B. homhifrons. In addi- 3529) of Bootherium may now be assigned tion, the length of the cranium posterior to the with certainty to B. homhifrons. They, but horn cores is 30% shorter than comparable especially USU 3529, are almost identical to specimens of B. homhifrons. This specimen the type specimen of B. homhifrons in size may represent a pathological individual, a and morphological characteristics. A second new species o( Bootherium, an individual vari- specimen of Bootherium in the Brigham ant, or a sexual opposite from that of the type Young University collections could not be lo- specimen of B. homhifrons. The gracile nature of cated (Nelson and Madsen 1978), but photo- this specimen suggests the latter choice, with

graphic evidence suggests that it too is B. IMNH 17124 representing a female. 250 Great Basin Naturalist Vol. 47, No. 2

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Fig. 14. A, Bootheriwn homhifrons, USNM 215066 (cast), original ANSP 2994, right donsolateral view; B, Bootherium sp. indet. , IMNH 17124, right dorsolateral view. Bar represents 10 cm.

Summary Corner (Nebraska State Museum) made specimens in their care available for study. We In summary, we assert that UVP 083 is also gratefully acknowledge the many discus- inseparable from Bootheriwn sargenti, which sions with other ovibovine enthusiasts includ- in turn is placed in synonymy with Syjnbos ing Jerry McDonald, John Neas, Dick cavifrons. These are sexual dimorphic forms, Harington, John White, Clayton Ray, Lee and it is likely that the typical S. cavifrons Stokes, and George Corner. The financial specimens are males whereas the "B. sar- support for this project was shared to some genti" forms are females. Bootherium homh- degree by the Utah Division of State History ifrons is a valid taxon and is probably not and DINOLAB, Salt Lake City, Utah, and closely related to S. cavifrons. Fort Hays State University, Hays, Kansas. Acknowledgments Literature Cited We could not have completed this paper Allen. A 191.3. Ontogenetic other variations without the counsel and help of John White J. and in musko.xen with and Clayton Ray. Robert Purdy of the Smith- a systematic review of the musko.x group, recent and extinct. Mem. Amer. Mus. Nat. sonian Institution graciously loaned us casts of Hist., n.s. 1: 103-226. "Bootherium sargenti" and B. homhifrons, Barton. J B 1976. A late Pleistocene local fauna from while John Neas of the University of Kansas American Falls, southeastern Idaho. Unpublished furnished a cast of the Natural Trap specimen. thesis, Brigham Young University, Provo, Utah. Dawkins, W. B. 1867. Ovibos moschatus (Blainville). Wade Miller (BYUG), Bill Akerston (IMNH), Proc. Roy. Soc. London 1.5: 516-517. Greg Ostrander (KUVP), Dave Liddell 1872. The British Pleistocene Mammalia. Part V. (USU), Charles Repenning (USGS), Tim British Pleistocene Ovidae, Ovibos moschatus. Smith (Alaska Fish and Game), and George Paleontographical Soc. 25: 1-30. April 1987 Nelson, Madsen: Fossil Musk Ox 251

Gazin. C. L 1935. Annotated list of" Pleistocene Mam- 1898. Wild oxen, sheep, and goats of all lands,

malia from American Falls, Idaho. J. Washington living and extinct. London. 318 pp. Acad. Sci. 25: 297-302. McDonald, J N 1984. An extinct muskox mummy from 1908. Descriptions of two new species of GiDLEY, J. W near Fairbanks, Alaska: a progress report. Pages Pleistocene ruminants of the genera OviJ)os and 148-152 in D. R. Klein, R. G. White, and S. Bootherium, with notes on the latter genus. Proc. Keller, eds.. Proceedings of the first international U.S. Nat. Mus. 34(1627): 681-684. muskox symposium. Biol. Pop. Univ. Alaska Spec. Harlan, R. 1825. Fauna Americana. Anthony Finley, Rep. 4. Philadelphia. 318 pp. McDonald. H. G., and E. Anderson 1975. A late Pleis- Hay, O. p. 1915. Contributions to the knowledge of the tocene fauna from southeastern Idaho. Tebiwa Pleistocene of North America. Proc. U.S. Nat. 18(1): 19-37. Mus. 48: 515-575.

Nelson, . Madsen, 1978. Late Pleis- 1924. The Pleistocene of the middle region of M E and J H Jr tocene musk oxen from Utah. Trans. Kansas Acad. North America and its vertebrated animals. 81: 277-295. Carnegie Inst. Washington Pub. 322A. 385 pp. Sci. 1927. The Pleistocene of the western region of Osgood, W. H. 1905a. ScapJwceros tijrelli, an extinct

North America and its vertebrated animals. Publ. ruminant from the Klondike Gravels. Smithsonian Carnegie Inst. Washington 322b. 346 pp. Misc. Coll. 48: 173-185.

Hesse. C J 1942. The genus Bootherium, with a new 1905b. Symbos, a substitute for Scaphoceros. record of its occurrence. Bull. Te.xas Arch. Paleon- Proc. Biol. Soc. Washington 18: 223-224. tol. Soc. 14: 77-87. Ray. C E 1966a. The identity of Bison appalachicolus. W., Hinds 1960. radiocarbon Hibbard. C. and F J A Notulae Naturae Acad. Nat. Sci. Philadelphia 384: date for a woodland musk o.x in Michigan. Pap. 1-7. Michigan Acad. Sci., Arts aixl Letters 45: 1966b. The status oi Bootherium brazosis. Texas 103-108. Mem. Mus. Pearce-Sellards Ser. 5: 1-7. Hopkins, M L 1951. Bison (Gi^antobison) latifrons and R\Y, C E., B N Cooper, and W S Benninghoff. 1967. Bison (Simobison ) allcni in southwestern Idaho. J. Fossil mammals and pollen in a late Pleistocene Mammal. 32: 192-197. deposit at Saltville, Virginia. Paleontol. 41: 1955. Skull of fossil camelid from American Falls J. 608-622. Bed area of Idaho. J. Mammal. 36(2): 278-282. Rhoads. S N 1895. Distribution of the American bison in Hopkins, M L , R Bonnichsen. and D E Fortsch 1969. The stratigraphic position and faunal associ- Pennsylvania, with remarks on a new fossil spe- ates of Bison (Gigantohison)latifrons in southeast- cies. Proc. Acad. Nat. Sci. Philadelphia 49: ern Idaho, a progress report. Tebiwa 12(1): 1-8. 483-502. Kurten. B., and E Anderson 1980. Pleistocene mam- RUTIMEYER. L 1867. Beitrage zu einer palaeontologis- mals of North America. Columbia University chen Geschichte der Wiederkaurer, Zunachst an Press, New York. 442 pp. Linne's Genus Bos. Verhandl. Naturf. Gesel. e.xtinct species Leidy, J. 1852a. Remarks on two crania of Basel. 4: 299-354. of ox. Acad. Nat. Sci. Philadelphia 6(3): 71. 1964. Semken. H A , Jr . B B Miller, and J B Stevens. 1852b. on the extinct species of American Memoir Late Wisconsin woodland musk oxen in associa- ox. Smithsonian Contrib. Knowledge 5: 1-20. tion with pollen and invertebrates from Michigan. 1869. The extinct mammalian fauna of Dakota and J. Paleontol. 38: 823-835. Nebraska, including an account of some allied Stokes, W. L.. andG H. Hansen 1937. Two Pleistocene forms from other localities, together with a synop- musk oxen from Utah. Proc. Utah Acad. Sci. 14: sis of the mammalian remains of North America. J. 63-65. Acad. Nat. Sci. Philadelphia, Ser. 2, 7: 23-472. White, 1985. Late Pleistocene musk oxen from Lonnberg. E 1900. On the structure and anatomy of the J A 64-71. muskox {Ovibos moschatiis). Proc. Zool. Soc. southern Idaho. Tebiwa 22: London. 686-718. WiSTAR, C 1918. An account of two heads found in the Lydekker. R 1885. Catalogue of the fossil Mammalia in morass, called Big Bone Lick, and presented to the British Museum, part 2, containing the order the Society, by Mr. Jefferson. Trans. Amer. Phil. Ungulata, suborder Artiodactyla. London. 314 pp. Socn.s. 1:375-380.