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Please refer to the appropriate Press Permission Controller: <[email protected]> for authors in North America; <[email protected]> for authors in Australia; <[email protected]> for authors in the rest of the world. The Primate Fossil Record Edited by walter carl hartwig touro university PUBLISHED BY THE PRESS SYNDICATE OF THE UNIVERSITY OF CAMBRIDGE The Pitt Building, Trumpington Street, Cambridge, United Kingdom CAMBRIDGE UNIVERSITY PRESS The Edinburgh Building, Cambridge CB2 2RU, UK 40 West 20th Street, New York NY 10011-4211, USA 477 Williamstown Road, Port Melbourne, VIC 3207, Australia Ruiz de Alarco´n 13, 28014 Madrid, Spain Dock House, The Waterfront, Cape Town 8001, South Africa http://www.cambridge.org © Cambridge University Press 2002 This book is in copyright. Subject to statutory exception and to the provisions of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press. First published 2002 Printed in the United Kingdom at the University Press, Cambridge Typeface Joanna 10.25/12.5pt System Poltype“ [VN] A catalogue record for this book is available from the British Library Library of Congress Cataloguing in Publication data The primate fossil record / edited by Walter Carl Hartwig. p. cm. Includes bibliographical references and index. ISBN 0 521 66315 6 1. Primates, Fossil. I. Hartwig, Walter Carl, 1964– QE882.P7 P75 2002 569'.8 – dc21 2001037847 ISBN 0 521 66315 6 hardback Contents List of contributors [ix] Preface [xi] Acknowledgements [xii] Abbreviations [xiii] 1 Introduction to The Primate Fossil Record [1] walter carl hartwig 2 The origin of primates [5] david tab rasmussen The earliest primates and the fossil record of prosimians [11] 3 The earliest fossil primates and the evolution of prosimians: Introduction [13] herbert h. covert 4 Adapiformes: Phylogeny and adaptation [21] daniel l. gebo 5 Tarsiiformes: Evolutionary history and adaptation [45] gregg f. gunnell and kenneth d. rose 6 Fossil lorisoids [83] erica m. phillips and alan walker 7 Quaternary fossil lemurs [97] laurie r. godfrey and william l. jungers The origin and diversification of anthropoid primates [123] 8 The origin and diversification of anthropoid primates: Introduction [125] marian dagosto 9 Basal anthropoids [133] k. christopher beard 10 Platyrrhine paleontology and systematics: The paradigm shifts [151] alfred l. rosenberger 11 Early platyrrhines of southern South America [161] john g. fleagle and marcelo f. tejedor 12 Miocene platyrrhines of the northern Neotropics [175] walter carl hartwig and d. jeffrey meldrum vii viii " contents 13 Extinct Quaternary platyrrhines of the Greater Antilles 20 European hominoids [339] and Brazil [189] david r. begun ross d. e. macphee and ines horovitz 21 The hominoid radiation in Asia [369] jay kelley The fossil record of early catarrhines and Old World monkeys [201] 22 Middle and late Miocene African hominoids [385] steven c. ward and dana l. duren 14 Early Catarrhines of the African Eocene and Oligocene [203] The fossil record of human ancestry david tab rasmussen [399] 23 Introduction to the fossil record of human 15 The Pliopithecoidea [221] david r. begun ancestry [401] henry m. mchenry 16 The Victoriapithecidae, Cercopithecoidea [241] 24 Earliest hominids [407] brenda r. benefit and monte l. mccrossin tim d. white 17 Fossil Old World monkeys: The late Neogene 25 Early Genus Homo [419] radiation [255] holly dunsworth and alan walker nina g. jablonski 26 Migrations, radiations and continuity: Patterns in The fossil record of hominoid primates [301] the evolution of Middle and Late Pleistocene humans [437] 18 Perspectives on the Miocene Hominoidea [303] fred h. smith david r. pilbeam 19 Late Oligocene to middle Miocene catarrhines from References [457] Afro-Arabia [311] Historical figures index [519] terry harrison Taxonomic index [523] 10 " Platyrrhine paleontology and systematics: The paradigm shifts alfred l. rosenberger Prologue While I would not go so far as Fleagle (2000) did in endors- ing our position, the independent corroboration of many The risk I take in this essay is that of a critic. Ultimately, my facets of the cebid–atelid cladistic model (Rosenberger, intention is to shed more light on platyrrhine evolution and 1981b, 1984, 1992) more than confirms the branching on problem areas where we may have gone astray because of sequence for the living forms. It validates many of the method. Accomplishing this without being critical is diffi- characters used to generate the tree, and these are eminently cult. applicable to fossils. While debates about primate phylogeny are real, and Nearly the entire literature on platyrrhine higher scientifically healthy, a philosophical context veils the cir- phylogeny and systematics over the past 20 years is slanted cumstances. First, the posture of our contemporary system- toward cladistics. Rare is the paper that eschews PAUP atic literature is to advocate positions rather than elucidate (Phylogenetic Analysis Under Parsimony: Swofford, 1993), hypotheses by conjecture and refutation. Debates about but some of the best analyses of fossils (e.g., Meldrum & Kay, phylogeny are often entwined in a webof taxa and traits 1997a,onNuciruptor) prove not to need algorithms at all. I which requires an exegesis if it is to be properly understood. believe there are some systematic problems that can benefit Second, with automated tree-building, and a limited capac- from a judicious use of numerical cladistics. But these ity to objectively select the ‘‘truth’’ from among many po- methods are not easily applied to the broad sweep of mor- tential tree-solutions, we are flooded with hypotheses that phological characters we are used to dealing with in primate are methodologically immunized from rejection, so they are systematics. The promise that large data sets and parsimony presented as viable models for the sake of consistency. Peer algorithms would bring greater objectivity to systematics has review and editorial direction can easily change the status quo not been realized, for that pivotal series of decisions upon and move us toward a more constructive dialogue. which all else is based – character selection – is by definition My sense is that ‘‘knowing’’ the phylogeny of platyrrhines a subjective, idiosyncratic process, often rooted in experi- is within our grasp for several interconnected reasons. First, ence and training. the empirical evidence reveals a surprising number of long- Cladistics in a strict sense narrowly defines ‘‘phylogeny’’ lived lineages (Rosenberger, 1979b; Delson & Rosenberger, as a branching sequence, as recency of common ancestry, or 1984), which should make easier the jobof reconstructing a network of collateral relationships. Among fossil New history. Second, from an analytical perspective, the living World monkeys there is an excellent opportunity to find forms are known to comprise a number of high-level clades examples of true phyletic evolution, ancestral–descendant that are morphologically coherent, distinctive and derived in relationships. Their importance to the story of platyrrhine pattern, whose behavioral ecology is also fairly well known. evolution is being misread if ancestors and descendants are To stereotype them: pitheciins are dentally bizarre seed- simply labeled sister taxa without further inquiry. Several eaters; atelines are postcranially modified climbers; callit- likely generic lineages have already been identified (Delson richines are dentally specialized, small-bodied claw-clinging & Rosenberger, 1984; Setoguchi & Rosenberger, 1987; locomotors; cebines are large-headed, predaceous, frugivor- Rosenberger, 1979b). Indeed, the preponderance of long- ous omnivores. Thus, we are dealing for the most part with lived lineages seems to be a high-level evolutionary pattern what we might call a ‘‘shallow phylogeny’’, a radiation among platyrrhines. The fossils themselves beckon a broader where solid knowledge of the living can be extended to the set of questions, and methods suitable to a more inclusive past in order to maximize our interpretive capacity. phylogenetic enterprise. Neither cladistic analysis nor mol- A third reason why we should have confidence in the ecular systematics can help us retrieve the entire story. In my interpretation of fossil platyrrhines comes from the force of view, the non-automated approach to morphology and char- genetic evidence. Schneider & Rosenberger (1996) have acter analysis that pays particular attention to homology, stressed the satisfying congruence in the results of mor- polarity, character weighting, functional morphology, be- phological and molecular studies of platyrrhine