HARVARD UNIVERSITY Library of the Museum of Comparative Zoology MCZ LIBRARY OCCASIONAL PAPERS AUG 2 1 1990 of the MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas NUMBER 137, PAGES 1-7 15 AUGUST 1990 PLEISTOCENE DIRE WOLF REMAINS FROM THE KANSAS RIVER WITH NOTES ON DIRE WOLVES IN KANSAS Xiaoming Wang1 The giant dire wolf, Canis {Aenocyon) dirus (Carnivora: Canidae), is the best known and most widely distributed canid in the Late Pleistocene of North America (Nowak, 1979). Up to 95 dire wolf localities throughout North America have been listed in Nowak (1979). The grey (timber) wolf {Canis lupus), a close extant relative of the dire wolf, generally has less extensive fossil records in the Pleistocene (57 localities; Nowak, 1979). Mainly limited to the Pleistocene Rancholabrean faunal age, the last record of the dire wolf is from the early Holocene (9,440 ± 760 B.P) from Brynjulfson Caves, Missouri (Kurten and Anderson, 1980). However dire wolf remains have been notably absent from eastern Kansas, though adjacent states and regions have well documented records (Missouri [Hawksley, 1963]; Nebraska [Voorhies and Corner, 1988]; and western Kansas [Hibbard, 1939, 1949]). In 1988, a partial right lower jaw of Canis dirus (KUVP 54032) was presented by Gary and Clodis Hunt of Bonner Springs, Kansas, to the Museum of Natural History at The University of Kansas. This specimen was collected from a Pleistocene sand bar on the west bank of the Kansas River, approximately 200 yards upstream from the Santa Fe railroad bridge, in the NE '/4, SW %, Sec. 32, T11S, R23E, Bonner Springs Quadrangle (U.S.G.S. 7.5' series topographic), south of Bonner Springs, Wyandotte County, Kan- 'Museum of Natural History and Department of Systematics and Ecology, The University of Kansas, Lawrence, Kansas 66045-2454. 2 OCCASIONAL PAPERS sas. A partial distal left humerus of the same species (KUVP 94008) was also donated, in the same year, by Mr. Ed Kost to the Museum of Natural History. The humerus was collected from a "large gravel bar" east of a bridge of Kansas Highway 7 over the Kansas River in the SW !4, Sec. 28, Tl IS, R23E, Edwardsville Quadrangle (U.S.G.S. 7.5' series), east of Bonner Springs. The two specimens may have been associated with the Newman Terrace assumed to be in the main valley of the Kansas River (Nelson and Neas, 1980). The terrace is believed to be Late Wisconsinan in age (Frye and Leonard, 1952). Numerous fossil remains have been found from sand bars along the Kansas River in eastern Kansas. Among the large mammals are mammoths (Mammuthus), American mastodont (Mammut), stag moose (Cervalces), elk (Cervus), caribou (Rangifer), white-tailed deer (Odocoileus), bison (Bison antiquus and B. bison), musk-oxen (Bootherium = Symbos), giant beaver (Castoroides), peccary (Mylohyus), sloth (Megalonyx jeffersoni), bear (Ursus), coyote (Canis latrans), mountain lion (Felis), and human (Homo) (Nelson and Neas, 1980). Because of the unconsolidated nature of the sediments and the constant washing by river water, this faunal list most likely represents a mixture of Pleistocene and Holocene vertebrate remains. The stratigraphic positions of most of the fossil remains are unknown. DESCRIPTIONS The mandible (KUVP 54032; Fig. 1 ) probably belongs to that of a mature adult judging from its modest wear on the occlusal surfaces of the teeth. The third premolar and first and second molars are well preserved. Parts of the alveoli for the canine and anterior two premolars are still intact. Wear has reduced the surface sculpturing of the bony part of the mandible; however, the enamel surfaces of the teeth are well-preserved. The deep cracking parallel to the bone fiber structure and flaking of the outer bone layer represent Stage II of weathering, as defined by Behrensmeyer (1978). indicating long surface exposure before burial. This individual (KUVP 54032) evidently lost its right fourth premolar early in life. In the place of the p4 is a 10 mm diastema, completely healed with no trace of an alveolus between p3 and m,. The di- astema is shorter than the 19.4 mm average length of p4 in C. d. dims (Kurten, 1984). This is probably the result of subsequent shifting of adjacent molars and premolars toward the vacant space. Similar tooth loss occurs in grey wolves and wolverines from the Pleistocene age Natural Trap Cave (Walker, MS; Rothschild and Martin, 1989), where it is the result of errors in bone manipulation. The p3 has a distinct posterior accessory cusp. The m , has a strong blade and a large shearing facet. The front edge of the paraconid of m, has an unique vertical wear facet (Fig. 1 ) not seen in any other dire wolf specimens examined. This long facet is probably due to the anterior shifting of m, toward DIRE WOLVES IN KANSAS ! 40 MM | Fig. 1 . Lateral view (lower) and partial crown view (upper) of KUVP 54032, Canis dims dims, from a Pleistocene Kansas River sand bar. the void left by the missing p4. The lower first molar had probably advanced far enough to occlude with the posterior edge of the upper third premolar. The talonid cusps of m, are almost completely worn. The m2 crown surface is also worn nearly flat. The only postcranial material of dire wolf from the Kansas River is a distal left humerus (KUVP 94008). The specimen is dark colored and heavily abraded, particularly around the medial and lateral epicondyles where the relatively fragile cancellous bone was badly eroded. The distance between medial and lateral epicondyles is approximately 49 mm. The posterior flange of the medial epicondyle, typical of canids, is also broken and lost. Both in terms of size and morphology, KUVP 94008 is similar to KUVP 72521 (a left mandible and partial postcranial skeleton) from Carrol Cave, Missouri. DISCUSSION Kurten (1984) differentiated two geographic subspecies of dire wolves, Canis dims dims from east of the Rocky Mountains and Canis dims guildayi from Mexico and the western coast (mainly the large sample from the Rancho La Brea tar pits) (Fig. 2). The eastern subspecies is said to differ from the western subspecies in its larger size, relatively longer distal limb bones, and a shorter P2. Although the differences between Kurten 's two subspecies are slight, it is probably justified considering that 24 geographic subspecies of grey wolves have been recognized in North America alone (Hall, 1981). 4 OCCASIONAL PAPERS Table 1. Selected measurements of Kansas River specimen (KUVP 54032) compared with those of both subspecies of Pleistocene dire wolves (data published in Kurten, 1984). RLB: Late Pleistocene Rancho La Brea tar pit samples. Measurement Sample // Mean SD Range Length p 3 C.d.guildayi (RLB) 20 16.3 1.00 14.1-18.7 KUVP 54032 1 17.5 — — C. d. dims (L. Wisconsin) 13 16.1 1.08 13.5-17.4 Width p, C.d.guildayi (RLB) 20 8.2 0.52 7.3-9.2 KUVP 54032 1 8.9 — — C. d. dims (L. Wisconsin) 14 7.7 0.86 6.7-9.0 Length m , C.d.guildayi (RLB) 33 35.5 1.57 32.9^10.2 KUVP 54032 1 37.9 — — C. d. dims (L. Wisconsin) 14 35.2 1.93 32.1-37.4 Width m, C.d.guildayi (RLB) 29 13.3 0.71 12.3-14.9 KUVP 54032 1 15.8 — — C. d. dims (L. Wisconsin) 22 13.4 1.13 11.8-16.3 Length m 2 C.d.guildayi (RLB) 25 13.3 0.79 11.7-15.5 KUVP 54032 1 13.3 — — C. d. dims (L. Wisconsin) 13 13.5 0.81 12.4-14.8 Width m. C.d.guildayi (RLB) 23 9.9 0.47 9.2-10.5 KUVP 54032 1 10.3 — — ■ C. d. dims (L. Wisconsin) 12 10.1 0.73 8.9-11.5 Despite the lack of diagnostic bony elements, the Kansas River specimen compares favorably in size with the Carrol Cave, Missouri specimen (KUVP 72521) and published measurements of the nominate subspecies by Kurten (1984) (Table 1 ). Most measurements of KUVP 54032 are either in the upper ranges, or slightly outside the upper limits, of the large eastern subspecies, C. d. dims. The overall dental morphology of KUVP 54032 is indistinguishable from either of the two subspecies (specimens examined: C. d. dims: KUVP 4613, 48135, 72521; C. d. guildayi: KUVP 222, 223). The Kansas River specimen is assignable to C. d. dims based on size. There are, however, exceptions to Kurten's (1984) subspecific taxonomy, notably a left mandible from Cragin Quarry, Meade Co., Kansas (KUVP 4613), and a few others. Kurten regarded them as "distal variants of this population." This is despite his realization that the measurements for the large C. d. dims were a mixture of widely separated geographical and temporal samples. In particular, the Cragin Quarry specimen is probably earlier than the Wisconsinan (Nowak [1984] listed it as from the Sangamonian). The Wisconsinan was the temporal interval upon which many of Kurten's DIRE WOLVES IN KANSAS • Canis dirus dims ■ Canis dims guildayi * Kansas River dire wolf ? Subspecies undetermined Fig. 2. Distribution of Pleistocene dire wolves, Canis dims in North America. Data from Graham (1987), Kurten (1984), Kurten and Anderson (1980), Nowak (1979), Voorhies and Corner ( 1 988 ), and Walker (1987). Shaded area represents the maximum advance of Wisconsinan glaciation (after Martin and Neuner, 1978). Subspecific determinations follow Kurten (1984). observations were based. Instead, the small Cragin Quarry specimen may represent an early transitional form between the presumed ancestral subspecies, Canis dims dirus, and its slightly smaller descendant, Canis dirus guildayi (Kurten, 1984). Unfortunately, teeth on KUVP 4613 are extremely worn, making dental comparisons difficult. However, the angular process of KUVP 4613 is flexed medially, in contrast to the relatively straight angular process of a Late Wisconsinan (12,777-21,370 B.P., Chomko and Gilbert, 1987) Wyoming specimen (KUVP 48130). Two of the adult Rancho La Brean 6 OCCASIONAL PAPERS specimens examined share a similar flexed condition. The above suggestions must remain speculative, until a more detailed study with larger samples is made. SUMMARY Dire wolves were present in the Late Pleistocene of eastern Kansas. Morphology of the eastern Kansas River specimens is suggestive of Canis dims dirus, supporting Kurten's (1984) contention that two distinct geographic subspecies of dire wolf populated North America. ACKNOWLEDGMENTS I thank Gary Hunt, Clodis Hunt, and Ed Kost for their generous donation of the specimens for this study. Larry D. Martin made important suggestions for the project. The manuscript was critically reviewed by Larry D. Martin and John Neas. I thank the two anonymous reviewers whose suggestions are greatly appreciated. LITERATURE CITED Behrensmeyer, A. K. 1978. Taphonomic and ecologic information from bone weath- ering. Paleobiology 4:150-162. Chomko, S. A. and B. M. Gilbert. 1987. The Late Pleistocene/Holocene faunal record in the northern Bighorn Mountains, Wyoming, pp. 394-^408. In: Graham, R. W., H. A. Semken, Jr. and M. A. Graham (eds.). Late Quaternary Mammalian Biogeography and Environments of the Great Plains and Prairies. Illinois State Mus. Sci. Pap., vol. 22. Springfield, Illinois. Frye, J. C. and A. B. Leonard. 1952. Pleistocene geology of Kansas. Bull. Geol. Surv. Kansas 99:1-230. Graham, R. W. 1987. Late Quaternary mammalian faunas and paleoenvironments of the Southwestern Plains of the United States, pp. 24-86. In: Graham, R. W., H. A. Semken, Jr. and M. A. Graham (eds.). Late Quaternary Mammalian Biogeography and Environments of the Great Plains and Prairies. Illinois State Mus. Sci. Pap., vol. 22. Springfield, Illinois. Hall, E. R. 1 98 1 . The Mammals of North America. 2nd ed. John Wiley & Sons, Inc., New York. 1:1-600+90,2:601-1181+90. Hawksley, O. 1963. The dire wolf in Missouri. Missouri Speleol. 5:63-72. Hibbard, C. W. 1 939. Notes on some mammals from the Pleistocene of Kansas. Trans. Kansas Acad. Sci. 42:463^179. Hibbard, C. W. 1949. Pleistocene stratigraphy and paleoecology of Meade County, Kansas. Contrib. Mus. Paleontol. Univ. Michigan 7:63-90. DIRE WOLVES IN KANSAS 7 Kurten, B. 1 984. Geographic differentiation in the Rancholabrean dire wolf (Canis dims Leidy ) in North America, pp. 2 1 8-227. In: Genoways, H. H. and M. R. Dawson (eds.). Contributions in Quaternary Vertebrate Paleontology: A Volume in Memorial to John E. Guilday. Spec. Pub. Carnegie Mus. Nat. Hist., No. 8. Pittsburgh, Pennsylvania. Kurten, B. and E.Anderson. 1980. Pleistocene Mammals of North America. Colum- bia University Press, New York, xvii+443 pp. Martin, L. D. and A. M. Neuner. 1978. The end of the Pleistocene in North America. Trans. Nebraska Acad. Sci. 6:117-126. Nelson, M. E. and J. Neas. 1 980. Pleistocene musk oxen from Kansas. Trans. Kansas Acad. Sci. 83(4):215-229. Nowak, R.M.I 979. North American Quaternary Canis. Univ. Kansas Mus. Nat. Hist. Monogr. 6:1-154. Rothschild, B. M. and L. D. Martin. 1989. Frequency of pathology in a large natural sample from Natural Trap Cave with special remarks on erosive disease in the Pleistocene. Symposium on Paleopathology, 49th Annual Meeting of Society of Vertebrate Paleontology. In press. Voorhies, M. R. and R. G. Corner. 1988. Canis dims: giant dogs of the ice age. Museum Notes, Univ. Nebraska State Mus. 72. Walker, D.N. 1987. Late Pleistocene/Holocene environmental changes in Wyoming: the mammalian record, pp. 334-392. In: Graham, R. W., H. A. Semken, Jr. and M. A. Graham (eds.), Late Quaternary Mammalian Biogeography and Environ- ments of the Great Plains and Prairies. Illinois State Mus. Sci. Pap., vol. 22. Springfield. Illinois. Walker, D. N. The wolves of Natural Trap Cave, Wyoming. Unpublished manuscript. UNIVERSITY OF KANSAS MUSEUM OF NATURAL HISTORY PUBLICATIONS The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 197 1 . Shorter research papers formerly published in the above series are now published as The University of Kansas Museum of Natural History Occasional Papers. 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