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AMERICAN MUSEUM NOVITATES Published by Number 1135 the AMERICAN MUSEUM of NATURAL HISTORY August 7, 1941 New York City

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AMERICAN MUSEUM NOVITATES Published by Number 1135 the AMERICAN MUSEUM of NATURAL HISTORY August 7, 1941 New York City AMERICAN MUSEUM NOVITATES Published by Number 1135 THE AMERICAN MUSEUM OF NATURAL HISTORY August 7, 1941 New York City THE OSTEOLOGY AND RELATIONSHIPS OF ARCHAEOMERYX, AN ANCESTRAL RUMINANT' BY EDWIN H. COLBERT PAGE INTRODUCTION ....................................................................... 1 MATERIALS UPON WHICH THE PRESENT STUDY IS BASED. 2 A REVIEW OF THE OSTEOLOGY OF Archaeomeryx optatus................................... 2 Analysis of the Diagnostic Characters of Archaeomeryx.................................. 2 The Comparative Osteology of Archaeomeryx.......................................... 3 The Skull....................................................................... 4 The Dentition.. 4 The Axial Skeleton ....................... 6 The Appendicular Skeleton. 7 The Fore-limb................................................................. 7 The Hind-limb................................................................ 7 The Relationships of Archaeomeryx................................................. 8 Tables of Measurements, Ratios and Indices........................................... 12 GENERAL DIscUssIoN ................................................................. 14 The Classification and Phylogeny of the Ruminants.................................... 14 CONCLUSIONS........................................................................ 22 BIBLIOGRAPHY ....................................................................... 23 INTRODUCTION The genus Archaeomeryx was first de- and fibular shafts are more primitive than scribed by Matthew and Granger in 1925, in any pecora, the upper incisors are still upon the basis of a large series of teeth, retained, the premolars are of primitive skulls and portions of skulls, skeletal ele- pattern, the molars brachyselenodont. It ments, and several associated skeletons, all lacks any of the various aberrant specializa- of which had been discovered in the Upper tions which exclude all of the Eocene genera Eocene Shara Murun formation of Mon- hitherto described from direct ancestry to golia by the Asiatic Expeditions of The the pecora and, as fortunately the principal American Museum of Natural History. In osteological and dental characters are deter- their original description, these authors minable from the exceptionally complete gave a detailed diagnosis of the type spe- material, the affinities of the genus can be cies, accompanied by figures of the denti- very satisfactorily appraised. So far as the tion. They also made a few remarks in higher ruminants are concerned, it affords this publication regarding the importance tangible and very convincing proof of the of this new form, as follows: theory of an Asiatic dispersal center."2 "This genus is of exceptional interest, Beyond the original description of the as it appears to be an approximate ances- genus, there has been very little notice tral type for the pecora. It has assumed taken of it in the literature, except for pass- the characteristic pecoran-traguline charac- ing references in the course of general dis- ter of the united naviculo-cuboid, but it cussions dealing with ruminant evolution. still retains the separate median pair and Since, because of its relatively early age and complete lateral pair of digits, the ulnar its generally unspecialized habitus, this is 1 Publications of the Asiatic Expeditions of The American Museum of Natural History. Contribu- 2 Matthew, W. D., and Granger, Walter, 1925. tion number 144. Amer. Mus. Novitates, No. 196, pp. 10-11. 2 AMERICAN MUSEUM NOVITATES [No. 1135 one of the most important of the ruminants tural ancestor for the pecorans. It is from an evolutionary standpoint, it has thought that such a detailed and compara- seemed advisable to make a detailed study tive osteological description, together with additional figures, will give to palaeontolo- of the genus as based upon the original ma- gists and zoologists an increased apprecia- terials, comparing it with other primitive tion and knowledge of this important form. ruminants in an effort to determine the The illustrations were made by Mr. John actuality of its position as a possible struc- C. Germann. MATERIALS UPON WHICH THE PRESENT STUDY IS BASED All of the specimens listed below were 20323, miscellaneous upper and found in the upper Eocene Shara Murun lower jaws and foot bones. formatiQn, at Ula Usu, Mongolia. All be- 20324, miscellaneous upper and long to the type species, Archaeomeryx lower jaws and foot optatus Matthew and Granger, 1925. bones. 20325, miscellaneous upper and Amer. Mus. No. 20311, type. Palate and man- lower jaws. dible. Numerous teeth and skeletal 20312, palate and lower jaws. elements, not numbered, repre- 20313, upper and lower jaws. sentative of at least 38 indi- 20314, upper and lower jaws. viduals. 20315, upper and lower jaws. 20316, portions of upper jaws; It may be seen that Archaeomeryx opta- right M2-3, left P4_M3. tus is known from a considerable series of 20317, upper and lower jaws. specimens, including literally dozens of den- 20318, upper and lower jaw titions. Tnus it is possible to present a .<.- ~~fragments. 20320, articulated skeleton. rather thorough study of the osteology of 20321, skull and jaws, part of this animal and to attempt a reconstruction skeleton, lumbars, pelvis of the skeleton upon a solid foundation of and part of hind limb. factual evidence, with the one exception 20322, skull and jaws, pelvis and hind limbs with ar- that in all cases the skulls are crushed, ticulated feet, fore limbs thereby making a restoration of the skull and feet. exceedingly difficult. A REVIEW OF THE OSTEOLOGY OF ARCHAEOMERYX OPTATUS ANALYSIS OF THE DIAGNOSTIC 3.-Dental formula 3/3, 1/1, 3/4, 3/3. OF 4.-Upper incisors well developed; upper ca- CHARACTERS Archaeomeryx nine of medium size; short post-canine diastema; The description by Matthew and Gran- three upper premolars; upper molars brachyo- ger, though brief, is remarkably complete, dont, quadritubercular, with strong parastyle and mesostyle and well-developed internal cingu- and there are but few important points in lum. the anatomy of this genus to be added to 5.-Lower incisors procumbent; lower canine what these authors have already so lucidly incisiform and in series with incisors; first lower described. On the basis of the original de- premolar caniniform and set apart by short dia- of the and the stemata from canine in front and from second scription genus characters lower premolar behind; lower premolars com- shown by the material at hand, a diagnosis pressed and trenchant, the last with a well- of the generic type may be presented as developed inner cusp and a posterior basin; lower follows. molars quadritubercular. 6.-Back rather long, with heavy vertebrae. Sacral vertebrae showing little fusion. Tail Archaeomeryx optatus very long. 1.-Of small size, approximately equal in this 7.-Radius and ulna separate. Carpals sepa- respect to the modern Tragulus. rate. Metacarpals separate, the lateral toes be- 2.-Skull hornless, with relatively small brain- ing complete but somewhat reduced. case, prominent occipital crest, and small orbit 8.-Pelvis not fused to sacrum. Fibula re- closed posteriorly. Mandible rather heavy. duced, only the proximal and distal ends being 19411 OSTEOLOGY OF ARCHAEOMERYX 3 Fig. 1. Comparison of the restored skull and jaw of (A) Archaeomeryx optatus Matthew and Gran- ger, based upon Amer. Mus. Ncs. 20311, 20320, 20321, 20322, with that of (B) Hypertragulus hes- perius Hay, Amer. Mus. No. 7918, type. Drawing of Hypertragulus adapted from Frick, 1937. Lateral views, natural size. retained. Astragalus completely artiodactyloid. and differences shown as it is compared Cuboid and navicular fused. Metatarsals sepa- rate, the lateral toes being complete but some- with certain important genera truly repre- what reduced. sentative of related families and subfamilies of traguloids.1 Thus, Archaeomeryx will be THE COMPARATIVE OSTEOLOGY OF compared so far as is possible with the Archaeomeryx primitive Eocene genus Amphimeryx, repre- In this comparative review of the oste- sentative of the family Amphimerycidae. of the will be ology Archaeomeryx, genus 1 The basis for the classificatory terms here used considered in the light of the resemblances will be elucidated on a subsequent page of this work. 4 AMERICAN MUSEUM NOVITATES [No. 1135 In comparing it with the hypertragulids, tive ruminants. Continuing, the orbit in the comparison will be largely with Hyper- Archaeomeryx is seemingly smaller in pro- tragulus itself, since this form is perhaps as portion to the size of the skull than is the typical as any member of the family. None case in Hypertragulus or Tragulus, as might of the protoceratids will be brought into be expected when comparing an Upper this comparative study, because they are so Eocene genus with forms of a later date. highly specialized as to be of little concern An interesting character of Archaeomeryx is to us in this connection. On the other the apparent closing of the orbit posteriorly, hand, comparisons with the true Tragulidae showing that this advanced ruminant char- will be necessary-with Gelocus as typical acter probably was established at a very of the subfamily Gelocinae, and particu- early date in the history of the suborder, larly with Tragu7us itself, as a central type even though in some later forms, such as representative of the subfamily Tragulinae. Hypertragulus, the orbit tends to remain In all of these comparisons, emphasis will open posteiiorly. be placed especially
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