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Paleontological Contributions THE UNIVERSITY OF KANSAS PALEONTOLOGICAL CONTRIBUTIONS September 24, 1979 Paper 95 REVISION OF THE CAMELINAE (ARTIODACTYLA, TYLOPODA) AND DESCRIPTION OF THE NEW GENUS ALFORJAS 1 JESSICA A. HARRISON Museum of Natural History, University of Kansas Lawrence, Kansas 66045 ABSTRACT A cladistic analysis of the subfamily Camelinae indicates the presence of two major groups of camels in the Neogene. One group includes the extant lamines of South Amer- ica and their fossil relatives. The other group includes Came/us and its sister group, the gigantic North American genera Megatylo pus and Titanotylo pus. Alforjas taylori, a new genus and species of lamine camel, is described from the Edson local fauna (Hemphillian) of western Kansas. INTRODUCTION The Camelinae constitute one of the most from those of Webb, and are based upon a cla- widely distributed and abundant elements in the distic analysis of the group. Characters used in Neogene faunas of North and South America. In analysis are termed apomorphic (derived) or North America the fossil record of the Camelinae plesiomorphic (primitive), with an autapomor- may be traced from the relatively primitive mid- phic character unique to a particular taxon and a dle Miocene protolabidine camels through the synapomorphic character shared by more than later Cenozoic to the extant genera currently re- one taxon. For more detailed discussions of phy- stricted to Asia and South America. logenetic systematics, see Hennig (1966), Brun- Wortman (1898) produced one of the earliest din (1968), and Wiley (1976); for deductive revisions of the Camelinae, and subsequently sev- testing of hypotheses, see Popper (1968a, b). Mu- eral workers have revised all or part of the group. seum acronyms appearing in this paper are: Matthew left an extensive unpublished manu- KUVP, University of Kansas Museum of Natural script on the Camelidae (Osborn Library, Ameri- History, F:AM, Frick Collection, American Mu- American can Museum of Natural History) in addition to seum of Natural History, and AMNH, the phylogeny that he suggested in 1918. The Museum of Natural History. are in cen- most recent phylogeny of the Camelinae was Measurements.—All measurements The meas- proposed by Webb (1965) in his perceptive and timeters unless otherwise indicated. (I), canines detailed "Osteology of Cume/ops." The rela- urements of diastemata and incisors and molars (M) were taken tionships suggested in this paper differ somewhat (C), premolars (P), on the teeth themselves unless otherwise noted. Upper molars were measured for length along the Manuscript received March 9, 1979. 2 The University of Kansas Paleontological Contributions—Paper 95 occlusal surface of the ectoloph. Width equals seum of Natural History. I also appreciate access the maximum distance transverse to the long axis to the collections of the University of Kansas of the tooth. The mandibular symphysis was meas- Museum of Natural History and the University ured on the ventral surface from the posterior of Nebraska State Museum. Special thanks go to border of the first lower incisor to the posterior E. Manning and H. Galiano for their insight, end of the symphysis. Width of the lower molars encouragement, and patience. I also thank J. G. was measured transversely across the middle of Honey for helpful conversations on the proto- the anterior portion of each tooth. Mean and labidine camels. J. A. Breyer and B. E. Taylor standard deviation were calculated only for those critically read the manuscript. The expertise of elements with a sample size of four or more. G. Krochak facilitated the processing of several For additional meristic data, see tables in Harri- loans. This son (1979). project was supported in part by Acknowledgements.—I am indebted to B. E. National Science Foundation Grant DEB77- Taylor and R. H. Tedford for their advice and 15869, Geological Society of America Grant stimulating criticism as well as their permission 2202-77, and a Summer Fellowship from the to utilize fossil collections of the American Mu- Graduate School of the University of Kansas. RELATIONSHIPS OF THE CAMELINAE The cladogram in Figure 1 graphically shows discussed in regard to their polarity and distribu- inferred relationships of the Camelinae discussed tion among the Camelinae and outgroup taxa. in this paper. Characters appearing at nodes 1 Table 1 summarizes the distribution of several through 23 on the cladogram are listed and then of the characters utilized in this study. FIG. 1. Cladogram showing proposed relationships of the subfamily Camelinae. Harrison—Revision of the Camelinae 3 Node 1. The Synthetoceratinae (Fig. 1) are Node 9. Pliauchenia, Hemiauchenia, Palaeo- distinguished by the following synapomorphies: lama, and Lama share the following apomorphic A. Single-shafted rostral horn formed of fused characters: maxillary processes. A. Lacrimal vacuity reduced. B. Paired supraorbital horns derived from the B. Rostrum shortened. frontals. Node 10. Pliauchenia is primitive in all Node 2. Poebrotherium and the Camel; ,, le known characters relative to the remaining gen- are united by: era in the Lamini. A. Extreme reduction of metapodials II and V. Node 11. Hemiauchenia, Palaeolama, and B. Divergent distal metapodials III and IV. Lama are united by: C. Anteriorly, the vertebrarterial canal passes A. In and P 1 small. through base of neural arch; posteriorly, verte- B. 132 small. bral artery is confluent with neural canal. Node 12 . The autapomorphy distinguishing D. Tympanic bulla inflated and filled with Hemiauchenia is extreme elongation of the limbs spongy, canccllous bone. and cervical vertebrae. Poebrotheriurn is not distinguished by an autapo- Node 13. Palaeolama and Lama form a nat- morphy and is primitive relative to the Carnelinae ural group in sharing: for all known characters. A. P 1 absent. Node 3. The Camelinae are a monophyletic B. P I absent. group sharing: C. Maxillary fossa reduced. A. Buccinator fossa weak. D. Lower molars with moderately strong antero- B. Rostrum elongate. external style. Node 4. The Protolabidini have been char- Node 15. Lama differs from Palaeolama in acterized by Honey and Taylor (1978) as shar- having: ing: A. P3 absent. A. Anterior nares laterally expanded. B. Metacarpal length subequal to metatarsal B. Rostrum narrow. length. Node 5. The Camelinae exclusive of the Pro- C. Lower molars with strong anteroexternal style. tolabidini are united by: D. Lacrimal vacuity greatly reduced. A. Metacarpal length exceeds that of metatarsus. E. Nasals extremely retracted. B. Metapodials completely fused. F. 13 4 greatly reduced. C. P absent. Node 16. Alforjas is most closely related to Node 6. Procamelus is the sister taxon to the Camelops in sharing a greatly increased degree of remaining camelines. It retains several primitive hypsodonty in comparison to the other Lamini. characters, but has almost completed the loss of Width of the cheek teeth in Alforjas is reduced 1 2 . in relation to crown length and height, resulting Node 7. At Node 7 it becomes apparent that in a very slim, hypsodont dentition. two main tribes (see following classification) Node 17. Alforjas is primitive in all known comprise the bulk of the Camelinae. These two characters relative to Came/ops. tribes arc united by: Node 18. Came/ops is derived relative to A. 1 2 absent. Alforjas in having: B. P 2 absent. A. Cheek teeth extremely hypsodont. C. Po absent. B. In absent. D. Raised posterolateral edges on the proximal C. P I absent. end of the first phalanx. D. P 3 absent. E. Erythrocytes ovoid. E. Dorsal surface of mandibular condyle trans- Node 8. The Lamini are united by: versely concave. A. Configuration of the anterior end of the na- F. Suspensory ligament scar extends to center of sals. shaft and has a raised center. B. Lower molars with anteroexternal style (= Node 19. The Camelini, consisting of Came- llama buttress of Webb, 1965). lus and the giant North American forms, is TABLE 1 . —Summary of Character Distribution in the Camelinae and Outgroup Taxa. Character Camelinae Procameltts Alforjas Camelops 1. P lost lost lost 2. 1 2 very reduced lost lost 3. present present present 4. C' laterally compressed reduced; laterally very reduced; compressed laterally compressed 5. C, laterally compressed reduced ; laterally very reduced; compressed laterally compressed 6. 13' present present lost 7. 13 , present present lost 8. p2 present lost lost 9. P2 present lost lost 10. P 3 present internal crescent internal crescent incomplete incomplete 11. 13:, present present lost 12. P4 present present reduced 13. Anteroexternal style on absent weak moderate lower molars 14. Crown height moderately high moderately high extremely high 15. Premaxilla light to moderate light light 16. Lacrimal vacuity large very large 17. Maxillary fossa moderately large large 18. Nasals flattened arched very arched 19. Rostrum long long long 20. Zygomatic arch curved curved curved 21. Postglenoid foramen moderately large small small 22. Postglenoid process small small moderately small 23. Postglenoid facet on centrally positioned; centrally positioned; centrally positioned; mandibular condyle transversely elongated transversely elongated transversely elongated 24. Diastemal crest on tnandible sharp sharp very sharp 25. Angular process on mandible moderate; inflected small; inflected moderate; inflected 26. Dorsal surface of mandibular concave-convex concave-convex concave condyle 27. Metacarpal length vs. metc.>mett. metc.>mett. metc.>mett. metatarsal length 28. Metapodial elements fused fused fused
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