m x h'v pc ^ g pi ^^£5^ g pi 5" m >TITUTION ^ NOIinillSNI~~NVINOSHlllAIS S3 1 HVH 8 tl“u B R AR I ES^SMITHSONIAN^NSTITUTION NOI i,s 1 . J 1 ... .A J& 1 I M* s X i 1 I I s | v^gasa; I HVH a !1 2 Li BRAR I ES^SMITHSONIAN*" INSTITUTION NOIlfUllSNI NVINOSHiliAIS^SH I HVH 0 n LI E — tn ~ tO “ tfi ZZ 1 O ” N' Q.msS X o sTITUTION 2 NOlinillSNI^NVINOSHlIWS^Sa 8 HVaa n J LI B RAR I ES^SMITHSONIAN^SNSTSTUTIONf NQf 2 r” > 2 r° z . ■ L. 2 O xSvnH/X ” V<>. o “ /T^aSN O ^s^Sv,. ™ y ^-mw 5 Xs ^le£^ / > 2 ^ 2 > § • m z <0 * z ^ to * z navaan libraries smithsonian institution NonniiiSNi nvinoshiiiais S3iavaan_i! co 5 to = _.. to — _ 2 to e \i H V o STITUTION^ NOlinilJLSNI^NVINOSHllWS^SB IHVaail^LIBRARI ES^SMUHSONIAN^ INSTITUTION ^ NO 2 /gx “ 2 5 /^3^\ 2 nrk J t\ i_ vplI\ ^.-. Js^. .-.x 1— ./v^iiFK A _ 4 CO OiVmnoUMIMli moiiawiiwix ”NVIN0SHllWS W S3iMVHaH LI B R AR I ES^SMITHSONIAN INSTITUTION 2 w z 5 iOi# lC)l«( g X gy&cJ X 5 '4x ! ' s ^ S \ ^SMITHSONIAN^ INSTlTUTiQM^MOSIfUIJtSN! NVINOSHIIWS^SI I H VH 9 11 V) = w ^ w u m w u ^ C/3 NOiinillSNi NVIN0SH1IWS a'<» ^ O ^ p ^5 ^ ” ~ ~ s | s§ p’*’ ^ 2 >’ 2 NVIN0SH1IMS OT S3 VIS 8 Sl^LI B RAR 8 ES^SMITHSONiAN ^INSTITUTION LIBRARIES SMITHSONIAN CO UJ i- / NOULfUilSNI NVINOSHlItNS LIBRARIES SMITHSONIAN C/> 2 S3 y& ^ 'NOlinillSNI^NVINOSHIlWS 0 ’ z (/) SMITHSONIAN INSTITUTION NOIlfULLSNI NVINOSHIIWS S3JHViJ0f1 LIBRARIES SMITHSONIAN r* _ z •" z f 4-03 /fifiifjKMv ? /fffiX ro 2 oo 3 PfrT q|£\ > }Jo| L\ k „ a >W 33 pi \^ocx «" m *" m z j ’nVINOSHIIWS^SS I HVHan“u BRAR I ES^SMITHSONiAN” INSTITUTION NQlinillSNI NVIN0SH1IIAIS >. to z w . z v.- w z < S .,< a^,. 5 XSSw'sv < .^\o 1 ^ _ a > N ^2ii£2' < ^ 2g W SMITHSONIAN^INSTITUTION »\ Palaeontology Published by The Palaeontological Association • London Price £25-50 VOLUME 32 • PART 3 OCTOBER 1989 THE PALAEONTOLOGICAL ASSOCIATION The Association was founded in 1957 to promote research in palaeontology and its allied sciences. COUNCIL 1989-1990 President : Dr J. D. Hudson, Department of Geology, University of Leicester, Leicester LEI 7RH Vice-Presidents'. Dr M. Romano, Department of Geology, University of Sheffield, Sheffield S3 7HF Dr P. W. Skelton, Department of Earth Sciences, The Open University, Milton Keynes MK7 6AA Treasurer : Dr M. E. Collinson, Department of Biology, King’s College, London W8 7AH Membership Treasurer : Dr H. A. Armstrong, Department of Geology, University of Newcastle, Newcastle upon Tyne NE1 7RU Institutional Membership Treasurer : Dr A. W. Owen, Department of Pure and Applied Geology, University of Glasgow, Glasgow G12 8QQ Secretary : Dr P. Wallace, The Croft Barn, Church Street, East Hendred, Oxon 0X12 SLA Circular Reporter : Dr D. Palmer, Department of Geology, Trinity College, Dublin 2 Marketing Manager: Dr C. R. Hill, Department of Palaeontology, British Museum (Natural History), London SW7 5BD Public Relations Officer: Dr M. J. Benton, Department of Geology, University of Bristol, Bristol BS8 1RJ Editors Dr M. J. Benton, Department of Geology, University of Bristol, Bristol BS8 1RJ Dr J. E. Dalingwater, Department of Environmental Biology, University of Manchester, Manchester M13 9PL Dr D. Edwards, Department of Geology, University of Wales College of Cardiff, Cardiff CF1 3 YE Dr C. R. C. Paul, Department of Earth Sciences, University of Liverpool, Liverpool L69 3BX Dr P. A. Selden, Department of Extra-Mural Studies, University of Manchester, Manchester M13 9PL Dr P. D. Taylor, Department of Palaeontology, British Museum (Natural History), London SW7 5BD Other Members Dr J. A. Crame, Cambridge Dr G. B. Curry, Glasgow Dr E. A. Jarzembowski, Brighton Dr R. A. Spicer, Oxford Overseas Representatives Argentina: Dr M. O. Mancenido, Division Paleozoologia invertebrados, Facultad de Ciencias Naturales y Museo, Paseo del Bosque, 1900 La Plata. Australia: Dr K. J. McNamara, Western Australian Museum, Francis Street, Perth, Western Australia 6000. Canada: Prof S. H. Williams, Department of Earth Sciences, Memorial University, St John’s, Newfoundland A1B 3X5. China: Dr Chang Mee-mann, Institute of Vertebrate Palaeontology and Palaeoanthropology, Academia Sinica, P.O. Box 643, Beijing. Dr Rong Jia-yu, Nanjing Institute of Geology and Palaeontology, Chi-Ming-Ssu, Nanjing. France: Dr J.-L. Henry, Institut de Geologie, Universite de Rennes, Campus de Beaulieu, Avenue du General Leclerc, 35042 Rennes Cedex. Iberia : Prof F. Alvarez, Departamento de Geologia, Universidad de Oviedo, C/. Jesus Arias de Velasco, s/n. 33005 Oviedo, Spain. Japan: Dr I. Hayami, University Museum, University of Tokyo, Hongo 7-3-1, Tokyo. New Zealand: Dr R. A. Cooper, New Zealand Geological Survey, P.O. Box 30368, Lower Hutt. Scandinavia: Dr R. Bromley, Fredskovvej 4, 2840 Holte, Denmark. U.S.A. : Prof A. J. Rowell, Department of Geology, University of Kansas, Lawrence, Kansas 66044. Prof N. M. Savage, Department of Geology, University of Oregon, Eugene, Oregon 97403. Prof M. A. Wilson, Department of Geology, College of Wooster, Wooster, Ohio 44691. W. Germany : Prof F. T. Fursich, Institut fur Palaontologie, Universitat, D8700 Wurzburg, Pliecherwall 1 MEMBERSHIP Membership is open to individuals and institutions on payment of the appropriate annual subscription. Rates for 1989 are: Institutional membership. £50 00 (U.S. $90) Student membership £1 1-50 (U.S. $20) Ordinary membership £2100 (U.S. $38) Retired membership . £10-50 (U.S. $19) There is no admission fee. Correspondence concerned with Institutional Membership should be addressed to Dr A. W. Owen, Department of Pure and Applied Geology, The University, Glasgow G12 8QQ. Student members are persons receiving full- time instruction at educational institutions recognized by the Council. On first applying for membership, an application form should be obtained from the Membership Treasurer: Dr H. A. Armstrong, Department of Geology, University of Newcastle, Newcastle upon Tyne NE1 7RU. Subscriptions cover one calendar year and are due each January; they should be sent to the Membership Treasurer. All members who join for 1989 will receive Palaeontology , Volume 32, Parts 1-4. Enquiries concerning back numbers should be directed to the Marketing Manager. Non-members may subscribe, and also obtain back issues of Vol. 29 onwards, at cover price through Basil Blackwell Ltd, Journal Subscription Department, Marston Book Services, P.O. Box 87, Oxford OX2 0DT, England. For issues before Vol. 29 contact the Marketing Manager. Cover: Ammonite ( Grossouvria ) from the Oxford Clay (Jurassic) of Woodham, Bucks., containing geopetal pyrite stalactites. Direct print from a thin section; pyrite white. Diameter of specimen approximately 10 mm. (See Sedimentology , 29, 639-667, 1982.) A CLIMATIC EXPLANATION FOR PATTERNS OF EVOLUTIONARY DIVERSITY IN UNGULATE MAMMALS by CHRISTINE M. JANIS Abstract. The radiation of ruminant artiodactyls, bovids in particular, that characterized the latter part of the Neogene, appeared to be at the expense of the hindgut-fernrenting ungulates (perissodactyls, proboscideans, and hyracoids), that showed a corresponding decrease in diversity and total numbers. However, climatic and vegetational changes may have been the cause for this decline, rather than direct competition with ruminants. The Tertiary change in relative diversity of hindgut fermenters, from initially more than 50% of the ungulate fauna to only 25-30%, occurred during the late Eocene and early Oligocene in higher latitudes, and in the mid- Miocene in lower latitudes. In both cases, this change was correlated with a climatic shift from low to high seasonality. Subsequently, the relative abundance of hindgut fermenters remained more or less constant in all latitudes until the end of the Pleistocene. The radiation of the ruminant artiodactyls appears to have taken place at the expense of less specialized selenodont artiodactyls such as anthracotheres, oreodonts, and traguloids, that were the first artiodactyls to show an increase in diversity after the late Eocene reduction in numbers of hindgut fermenters. The pattern of evolutionary diversity of ungulate mammals through the Tertiary period has long been seen as a success story for the Order Artiodactyla. The radiation and diversification of ruminant artiodactyls during the Neogene, reaching its apogee with the supreme Plio-Pleistocene success of the Family Bovidae, is often contrasted with that of the Order Perissodactyla. Perissodactyls reached their maximum species diversity in the late Eocene, and thereafter showed a decline in abundance and family diversity that was more or less synchronous with the onset of the diversification of the ruminating artiodactyls (members of the Suborders Tylopoda and Ruminantia). This contrasting pattern of radiation between the two orders is usually attributed to competitive interaction and ecological replacement of an inferior group by a better adapted one, the key cited feature being the forestomach site of fermentation in the ruminants, contrasted with the hindgut site of fermentation in the caecum and colon of the perissodactyls (e.g. Simpson 1953; Van Valen 1971; Stanley 1974). This simple pattern of artiodactyl radiation versus perissodactyl decline is more complex than has previously been assumed. Perissodactyls reached their maximum diversity at the family level in the late Eocene with 13 families, 3 of which were extinct by the end of the Eocene, and a further 6 were extinct by the middle Miocene (text-fig. 1). However, the remaining perissodactyl families did not show a continuing Neogene evolutionary pattern of steady decline in diversity and abundance. The Rhinocerotoidea had a maximal diversity in the later Oligocene and early Miocene, and rhinocerotids were abundant throughout the Miocene. The Equidae had their maximal diversity in the late Miocene, the Chalicotheriidae had a moderate diversification during the Neogene (only suffering extinction during the Pleistocene) and, while the Tapiriidae never displayed a great diversification, they persisted little changed from the Oligocene to the present day (Janis 1984). All living perissodactyls, derived from a common early Eocene or late Palaeocene ancestry, are hindgut fermenters, and it thus seems parsimonious to assume that all extinct members of this order (which all possess lophed cheek teeth indicative of a folivorous diet) had a similar type of digestive physiology. Only two out of a total of 37 Tertiary artiodactyl families (Bovidae and Cervidae) have a | Palaeontology, Vol. 32, Part 3, 1989, pp. 463-481. | © The Palaeontological Association 6 PAL 32 464 PALAEONTOLOGY, VOLUME 32 M.Y.B.P. RECENT PLEISTOCENE PLIOCENE 5.6 12 MIOCENE M 16 24 OLIGOCENE 31 37 40 EOCENE M 51 57 5--S - • o ^ I -C srlh s r l ° _ o — T5 E " E 'CONDYLARTHRA" < < £ 5 O