See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/262698724 Evolution and classification of Tragulina (Ruminantia, Artiodactyla) Article in Paleontological Journal · January 2001 CITATIONS READS 40 624 1 author: I. A. Vislobokova Russian Academy of Sciences 99 PUBLICATIONS 1,495 CITATIONS SEE PROFILE Some of the authors of this publication are also working on these related projects: Middle Pleistocene mammal faunas of Europe View project Entwicklung der Fauna und Flora im Oligozän und Miozän der Mongolei (FWF-Projekt: P-10505-GEO) View project All content following this page was uploaded by I. A. Vislobokova on 30 May 2014. The user has requested enhancement of the downloaded file. Å À . V i sl ob o k o v a Paleontological Institute, Russian Academy of Sciences, ul. Profsoyuznaya 123, Moscow, 117868 Russia (R ecei ved Òèï å 30, 2000) C on t en t s V o l . 3 5 , Su p p l . 2, 200 1 The supplement is published only in English by Ì À!Ê "Nauka/Interperiodica (Russia). Paleonrologi cal Journal ISSN 0031-0301. INTRODUCTION $ 69 L IM B BONES $ 106 Sc a p u l a a n d H u m er u s $ 107 Chapter 1 R a d i u s a n d U l n a $ 110 ÒÍ Å M A IN PROBL EM S OF CL A SSIFICATION M a n u s s » o A ND PHYL OGENY IN TRAGUL INES s v o P el v i s a n d F em u r S111 T i b i a a n d F i b u l a $11ã Chapter 2 P es COM PA RATIV E M ORPHOL OGY S7 7 $ 113 C h ap ter 3 E V O L U T I O N O F T R A G U L I N E S S K U L L $77 S k u l l S h a p e a n d P r o p o r t i o n s S77 ORIGIN ÿ ï S a g i t t a l a n d T em p o r a l C r est s S84 B asa l T r a g u l i n e G r o u p ÿ ï O r b i t a l R eg i o n S84 T r a g u l i n es a n d O t h er P r i m i t i v e A r t i o d a c t y l G r o u p s E t h m o i d a l F i ssu r e $ 86 P a l a t e S86 M A IN EVOL UT IONA RY T REN DS S119 O cc i p i t a l s S88 H er b ivor ou s A d ap t at ions ÿ ãî S p h en o i d $ 88 A d ap tat ion s of Sen so r y T em p o r al S90 and R esp i r at or y O r gan s ÿ ã~ P a r i et a l S94 C u r sor ial A d ap tat i on s ÿ ã1 F r o n t a l $ 94 PHY LOGENET IC PAT 7ERNS N a sa l S95 $ 122 T r a g u l o i d s a n d H y p er t r a g u l o i d s L ac r i m a l S95 ÿ ãã T r a g u l o i d s S] 24 J u g a l S96 H y p er t r a g u l o i d s $ 127 M a x i l l a S96 P a l a t i n e $96 Chapter 4 P r em a x i l l a $ 97 DISPERSA L OF TRAGUL INES S 129 M a n d i b l e $97 Chapter 5 CL A SSIFICATION OF TRAGUL INES D E N T I T I O N $ 99 D en t a l F o r m u l a $ 99 S14 1 I n c iso r s a n d C a n i n es S99 P r em o l a r s $ 99 A C K N OW L ED GM EN T S $ 14 1 M o l a r s $1î ç E n a m el D u r a b i l i t y $ 106 REFEREN CE S S 14 1 ~ Paleontological Journal, Vol. 35, Suppl. 2, 2001, pp. S69–S145. Original Russian Text Copyright © 2001 by Vislobokova. English Translation Copyright © 2001 by MAIK “Nauka /Interperiodica” (Russia). Abstract—The main evolutionary trends of the Tragulina are traced and the systematics and phylogeny of this taxon are revised based on the study of the comparative morphology of fossil and Recent species and on the estimation of the polarity and taxonomic value of features. Nine families are included in the Tragulina. The existence of two main traguline groups is confirmed, and the names Traguloidea and Hypertraguloidea are restored for these groups. The retention of many plesiomorphic characters shared with ancient eutherian mam- mals and the possession of very deep apomorphies distinguish tragulines from other artiodactyls and support a pre-Eocene origin of the Tragulina connected with the early radiation of artiodactyls. INTRODUCTION Tragulines are the most primitive extinct and extant their primitive characters. Some plesiomorphies are ruminants which flourished during the second part of shared with the Suiformes. For example, Tragulus jav- the Paleogene and gave rise to the pecorans at the anicus shares more muscle antigens with the Suiformes beginning of the Oligocene. The pecorans are one of the than with ruminants (Duwe, 1969). However, the stom- most numerous and widespread groups among Recent ach of Recent tragulids already has a true ruminant mammals. Hence, the study of tragulines helps us to reticulum and other ruminant characters (Boas, 1890) understand one of the most significant and intriguing which are not present in the Suiformes. chapters in the evolution of mammals. The main purposes of this work are (1) to clarify the According to the fossil record, the history of tra- evolutionary trends and relationships of tragulines gulines covers more than 50 million years from the using cranial, dental, and postcranial characters; and Middle Eocene to the present, but it seems that their (2) to revise the phylogeny and classification of the tra- history is much longer. The appearance of tragulines gulines. Special emphasis is made on the investigation was one of the earliest and most successful attempts at of character polarities and on the estimation of the tax- the development of herbivorous adaptations and strate- onomic level of features. gies to avoid predators. Even at the early stages of evo- lution, there was a rather wide spectrum of forms This research is based on abundant fossil material adapted to various plant foods. They inhabited all the from China and Mongolia that is housed in the Paleon- continents of the Northern Hemisphere. Most of them tological Institute of the Russian Academy of Sciences remained largely herbivorous, also feeding on some (PIN); on fossil tragulines from Europe, Asia, America, small animals (mollusks, insects, etc.), as living chev- and Africa; and on Recent tragulines from Asia and rotains do. Africa. These specimens are from the American Museum of Natural History (AMNH); Museum of Nat- Notwithstanding the more than 150 years that these ural History, London (BMNH); University of Kansas; ruminants have been studied, and the number of cladis- Zoological Institute of the Russian Academy of Sci- tic analyses that have been undertaken during the past ences; University of Hanoi; and the University of two decades, the taxonomy and phylogeny of the group Montpellier. remain disputable with respect to many questions. This concerns not only rare forms represented by fragmen- Scanning electron microscopy of the enamel ultra- tary material, but also many well-studied genera. Major structure of traguline molars and X-ray study were per- problems include the monophyly of the Tragulina and formed at the Paleontological Institute of the Russian the relationships, rank, and taxonomic position of most Academy of Sciences. groups. The anatomy, behavior, and habitats of Recent chev- Apart from the morphological data (Kaup, 1833; rotains were studied during a biological expedition to Gray, 1845; Milne-Edwards, 1864; Flower, 1867; Gar- Vietnam. rod, 1877; Carlsson, 1926; Flerov, 1931; Janis, 1984; etc.), there is genetic, immunological, and ethological In addition to the traguline material, the rich collec- evidence (Dubost, 1965; Todd, 1975) that distinguish tions of ancient ungulates in the American Museum of Recent tragulids from other ruminants and indicate Natural History were also used in this work. S69 S70 VISLOBOKOVA Chapter 1 THE MAIN PROBLEMS OF CLASSIFICATION AND PHYLOGENY IN TRAGULINES The rank determination and composition of the must possess a set of characters which makes it equiv- taxon Tragulina, proposed by Flower (1883) based on alent in rank to the acknowledged related taxa. the family Tragulidae, as well as the rank determination and composition of some other superspecies of tra- The objective renovation of phylogeny and taxon- guline taxa, is still controversial. The Tragulina has omy must be based on the revelation of the basal struc- been regarded as a suborder of the order Artiodactyla tures and main evolutionary trends (transmission of (Osborn, 1910; Colbert, 1941), an infraorder of the sub- heredity and main adaptation trends).