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Paleontological Institute, Russian a Cademy of Sciences Å À . V i s l o b o k o v a a n d  . À . T r o f i m o v Pal eontol ogi cal I nstitute, R ussi an A cadem y of Sciences C ont en t s V oL 3 6, Su p p L 5 , 2 0 02 T he su p p l em en t i s p u b l i sh ed o n l y i n E n g l i sh b y M A I K " N au k a i I n t er p er i o d i c a ' ( R u ssi a) . P a l eo n to l o g i c a l J o u r n a l I S S N 0 0 3 1- 0 30 1. w m o o u c mo x ñí ëðòÅê 1 S43 1 ÒÍ Å H I S T O R Y O F ST U D Y I N G A R C H A E O M E R 1X À Ì > T H E M A I N P R O B L E M S O F P H Y L O G E N Y O F T H E À Ê Ï Î Ð À Ñ ÒÚ× .À $43 1 CHAPTER 2 TAX ONOM IC REV IEW OF THE ARCHAEOMERYCIDAE CHAPTER 3 S44 1 OSTEOL OGY À1×?) OD ONTOL OGY OF ARCHAEOMERYX S44 1 SKUL L S44 1 CRA NIAL BONES S44 8 FACIA L BONES S455 DENT S ON S46 1 DIA ST EM ATA S465 ENA M EL ULTRA STRUCTURE S465 V ERTEBRA L COL UM N S466 S472 FOREL IM B BONES S472 HIND L IM B BONES S4 83 C H A PT E R 4 ì î è í î í ë÷ñ ò þ û ë ò. A NA LY SIS AND RECONSTRUCTION O F T H E Ì Î Ð Å O F L I F E O F À Ê Ñ Í À Å Î Ì Å Ê ÓÕ S494 M O R P H O F U N C T I O N A L F E A T U R E S O F SK U L L A N D P O S T C R A N I A L S K E L E T O N $ 494 R E C O N ST R U C T I O N O F À ÐÐÅ À Ê À Ì Ñ Å A N D Ì Î Ð Å O F L I F E S504 C H A P T E R 5 S506 T H E R O L E O F À ß Ñ Í À Å Î Ì Å ß ÓÕ I N T H E E V O L U T I O N O F T H E A R T I O D A C T Y L A A R C H A E O M E R YX Û Æ E A R L Y E U T H E R I A N S S506 A R C H A E O M E R YX Õ ß 0 E A R L Y U N G U L A T E S S50 8 A R C H A E 0 M E R 1X È Æ D I C H O B U N O I D S S509 Ò Í Å M A I N T R E N D S A N D PA I T E R N S O F R U M I N A N T E V O L U T I O N C O N C L U SI O N S S5 19 SU M M A R Y S5ãî A C K N O W L E D G M E N T S S5ãî R E F E R E N C E S S5ã 1 Dedicated to the memory of Wladimir Kowalevsky (1842— 1 883), the great founder of evolutionary paleotheriology "It can be proposed that some small species of Eocene ungulates can show us relationships about which now we have even ï î presumption." W .Î . K owalevsk v ( 1873 —1 8 7 ~ ~ Paleontological Journal, Vol. 36, Suppl. 5, 2002, pp. S429–S522. Original Russian Text Copyright © 2002 by Vislobokova, Trofimov. English Translation Copyright © 2002 by MAIK “Nauka /Interperiodica” (Russia). Abstract—The morphology of the skull and postcranial skeleton of the primitive ruminant artiodactyl Archae- omeryx optatus Matthew et Granger, 1925 from the Middle Eocene Shara Murun Formation of the Ula Usu locality in Inner Mongolia of China is described in detail. In addition to the type collection housed at the Amer- ican Museum of Natural History, extensive fossil material collected by the Joint Soviet–Chinese Paleontologi- cal Expedition in 1959 and stored at the Paleontological Institute of the Russian Academy of Sciences is stud- ied. The material of the type collection, including, among other fossils, one complete articulated and two frag- mentary skeletons, is supplemented by 14 additional skeletons. The appearance and ecology of Archaeomeryx are specified. It is shown that, in morphology and ecology, this animal was close to primitive eutherians. New mor- phological evidence of the early evolutionary stages of the suborder Ruminantia and the order Artiodactyla and comparisons with other primitive artiodactyls and ungulates show that the order Artiodactyla undoubtedly appeared much earlier than was registered in the fossil record and that its roots perhaps go back to the Late Cre- taceous. This conclusion supports the hypothesis first proposed by W. Kowalevsky in the 19th century. The adap- tatiogenesis and the major principles of macroevolutionary processes are traced based on Ruminantia evolution. INTRODUCTION Archaeomeryx is a unique extinct artiodactyl of the branches, which became extinct during the course of infraorder Tragulina that can be a key to understanding evolution. The key species (such as Archaeomeryx), the evolution and adaptatiogenesis of one of the major which occupy the basal position in large evolutionary eutherian branches. branches and have their descendants in the Recent fau- The study of Archaeomeryx is very important not nas, usually fall out of fossil record or are represented only for a better understanding of the early stages in the by rather fragmentary material. The study of such spe- evolution of the Ruminantia and for studying the evolu- cies is essential to the understanding of trends and the tion of the infraorder Pecora, which are among the principal patterns of evolution. dominant groups of modern mammals, but also for the Notwithstanding the fact that Archaeomeryx is usu- solution to a very difficult problem of the origin and ally involved in the consideration of the origin and rela- early evolution of the order Artiodactyla. Moreover, the tionships of ruminants, the evolutionary significance of study of this animal is of great importance for resolving this genus has been underestimated for a long time. such problems of general biology as the main trends Therefore, a detailed description of Archaeomeryx has and patterns of evolutionary developments of living not yet been performed. Today, almost 75 years after its creatures. first finding and more than 40 years after its second Archaeomeryx represented by a series of almost finding, we intend to improve this situation. complete skeletons attracted the attention of paleontolo- After the study performed by Colbert (1941), it was gists from its first fossil records found by an expedition generally accepted that Archaeomeryx was rather simi- of the American Museum of Natural History in the lar in structure and ecology to Recent chevrotains, in 1920s in the Middle Eocene Ula Usu locality, northern particular, the mouse deer of the genus Tragulus. Chev- China (Matthew and Granger, 1925a; Colbert, 1941; rotains are usually regarded as living fossils (see, e.g., Webb and Taylor, 1980; etc.). From the very beginning, Janis, 1984). Even the first researchers recognized the this genus has aroused exceptional interest because of primitive and unusual structure of chevrotains, the its relatively early age, the superb fossil material, and as smallest extant ruminants, which are more similar in a probable ancestral type for higher ruminants (Pecora) appearance to rodents than to deer or antelope. How- (Matthew and Granger, 1925a). ever, as compared to Archaeomeryx, the small and Currently, the data on the morphology and ecology primitive living chevrotains look like the acme of per- of Archaeomeryx and its role in the evolution of mam- fection. mals are essentially enlarged due to the study of an The main purposes of the present work are (1) to extremely abundant collection stored at the Paleonto- describe in detail the osteology and odontology of logical Institute of the Russian Academy of Sciences. Archaeomeryx, (2) to perform morphofunctional analy- These fossils were obtained by the Joint Soviet–Chi- sis of the main structures of this animal, (3) and to clar- nese Expedition during intensive excavations in the Ula ify the role of Archaeomeryx in the evolution of the sub- Usu locality in 1959. order Ruminantia and the order Artiodactyla. The finding of Archaeomeryx, extremely well pre- We analyzed the adaptatiogenesis of the earliest served by nature and belonging to one of the main ruminants, the relationships of archaeomerycids and streams of mammal evolution, is a very rare event. Pale- other primitive members of the order Artiodactyla, the ontologists usually deal with species that represent side relationships of archaeomerycids with certain early S429 S430 VISLOBOKOVA, TROFIMOV eutherians and the most primitive members of the American researchers (Matthew and Granger, 1925a; grandorder Ungulata.
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