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Ardipithecus Ramidus FULL COLLECTION Around the world, governments turn to AAAS, publisher of Science magazine, as an objective, multidiscipli- Science funding nary scientific authority toeducate public officials and judicial figures on today’s most pressing issues. Our Climate regulation goal is to promote informed policy decisions that benefit society. And this is just one of the ways that AAAS Human rights is committed to advancing science to support a healthy and prosperous world. Join us. Together we can make a difference. aaas.org/plusyou Ardipithecus ramidus CONTENTS Introduction and Author List Research Articles 5 Light on the Origin of Man 29 Ardipithecus ramidus and the Paleobiology of Early Hominids Editorial Tim D. White et al. 9 Understanding Human Origins 41 The Geological, Isotopic, Botanical, Bruce Alberts Invertebrate, and Lower Vertebrate Surroundings of Ardipithecus ramidus News Focus Giday Wolde Gabriel et al. 46 Taphonomic, Avian, and Small-Vertebrate 10 A New Kind of Ancestor: Indicators of Ardipithecus ramidus Habitat Ardipithecus Unveiled Antoine Louchart et al. 14 Habitat for Humanity 50 Macrovertebrate Paleontology and the Pliocene 15 The View From Afar Habitat of Ardipithecus ramidus Tim D. White et al. Authors’ Summaries 57 The Ardipithecus ramidus Skull and Its 18 Ardipithecus ramidus and the Paleobiology of Implications for Hominid Origins Early Hominids Gen Suwa et al. Tim D. White et al. 64 Paleobiological Implications of the 19 The Geological, Isotopic, Botanical, Ardipithecus ramidus Dentition Invertebrate, and Lower Vertebrate Gen Suwa et al. Surroundings of Ardipithecus ramidus 70 Careful Climbing in the Miocene: Giday WoldeGabriel et al. The Forelimbs of Ardipithecus ramidus and 20 Taphonomic, Avian, and Small-Vertebrate Humans Are Primitive Indicators of Ardipithecus ramidus Habitat C. Owen Lovejoy et al. Antoine Louchart et al. 78 The Pelvis and Femur of Ardipithecus ramidus: 21 Macrovertebrate Paleontology and the Pliocene The Emergence of Upright Walking Habitat of Ardipithecus ramidus C. Owen Lovejoy et al. Tim D. White et al. 84 Combining Prehension and Propulsion: The 22 The Ardipithecus ramidus Skull and Its Foot of Ardipithecus ramidus Implications for Hominid Origins C. Owen Lovejoy et al. Gen Suwa et al. 92 The Great Divides: Ardipithecus ramidus 23 Paleobiological Implications of the Reveals the Postcrania of Our Last Common Ardipithecus ramidus Dentition Ancestors with African Apes Gen Suwa et al. C. Owen Lovejoy et al. 24 Careful Climbing in the Miocene: 99 Reexamining Human Origins in Light The Forelimbs of Ardipithecus ramidus and of Ardipithecus ramidus Humans Are Primitive C. Owen Lovejoy C. Owen Lovejoy et al. See also related video, Science Podcast at 25 The Pelvis and Femur of Ardipithecus ramidus: www.sciencemag.org/ardipithecus/ The Emergence of Upright Walking C. Owen Lovejoy et al. 26 Combining Prehension and Propulsion: The Foot of Ardipithecus ramidus C. Owen Lovejoy et al. 27 The Great Divides: Ardipithecus ramidus Reveals the Postcrania of Our Last Common Ancestors with African Apes C. Owen Lovejoy et al. 28 Reexamining Human Origins in Light of Ardipithecus ramidus C. Owen Lovejoy . WHITE, 2008 T OPYRIGHT C CREDIT: CREDIT: AAAS is here. In America today, 1 in 3 individuals do not accept evolution.1 That’s why AAAS continues to play an important role in the effort to protect the integrity of science education. AAAS is hard at work ensuring that evolution Evolution continues to be taught in science classrooms, but we need your help. Join us. Together we can make a difference. aaas.org/plusyou/evolution 1. Pew Research Center for the People & the Press. May 2009, General Public Science Survey. INTRODUCTION Light on the Origin of Man CHARLES Darwin’s SEMINAL WORK ON THE ORIGIN OF SPECIES, These papers synthesize an enormous amount of data collected PUBLISHED 150 years ago next month, contains just one understated and analyzed over decades by the authors. Because of the scope of sentence on the implications of his theory for human evolution: these papers and the special broad interest in the topic of human “Light will be thrown on the origin of man and his history.” As evolution, we have expanded our usual format for papers and cover- Darwin implied in his introduction to The Descent of Man, he felt age. The papers include larger figures, tables, and discussions, and that those implications were obvious; he appreciated, as events the overview and two concluding papers provide extended introduc- quickly showed, that it would be only natural to look at evolution tions and analyses. foremost from our human perspective and contemplate what makes In addition, to aid understanding and introduce the main results us unique among other primates—our large brains and ability to of each paper, the authors provide a one-page summary of each communicate, to create, and to understand and investigate our his- paper, with an explanatory figure aimed at the general reader. Our tory and nature; our culture, society, and religion; the ability to run News Focus section, written by Ann Gibbons, provides further fast on two legs and manipulate tools; and more innovations that analysis and coverage, and it includes maps and a portrait of separate us from our primate relatives. the meticulous and at times grueling field research behind the Tracing our evolution and how we came to acquire these skills discoveries. Available online are a video interview and a podcast and traits, however, has been difficult. Genetic data now confirm with further explanations. that our closest living primate relative is the chimpanzee. We shared To accommodate this material and allow the full papers, this print and evolved from a common ancestor some 6 million or more years issue presents an Editorial, News coverage, the authors’ summaries, ago. But identifying our unique genes and other genetic differences and four papers in full: the overview paper and one key paper from between us and our primate cousins does not reveal the nature of each thematic group above. The other research papers, and of course that ancestor, nor what factors led to the genetic changes that un- all content, are fully available online. In addition, a special online derlie our divergent evolutionary paths. That requires a fossil record page (www.sciencemag.org/Ardipithecus/) links to several print and and enough parts of past species to assess key anatomical details. download packages of this material for AAAS members, research- It also requires knowing the habitat of early humans well, to deter- ers, educators, and other readers. mine their diet and evaluate what factors may have influenced their This collection, essentially an extra issue of Science in length, evolution through time. Many early human fossils have been found, reflects efforts by many behind the scenes. Every expert reviewer but with a few exceptions, these are all less than 4 million years old. evaluated, and improved, multiple papers, and several commented The key first several million years of human evolution have been on all 11 of them. The authors provided the summaries on top of poorly sampled or revealed. an already large writing and revision effort. Paula Kiberstis helped This issue presents 11 papers authored by a diverse international in their editing. The figures and art were drafted and improved by team (see following pages) describing an early hominid species, J. H. Matternes, Henry Gilbert, Kyle Brudvik, and Josh Carlson, Ardipithecus ramidus, and its environment. The hominid fossils as well as Holly Bishop, Nathalie Cary, and Yael Kats at Science. are 4.4 million years old, within this critical early part of human Numerous other Science copyediting, proofreading, and production evolution, and represent 36 or more individuals, including much of staff processed this content on top of their regular loads. Finally, the skull, pelvis, lower arms, and feet from one female. The papers special thanks go to the people of Ethiopia for supporting and facili- represent three broad themes. Five focus on different parts of the tating this and other research into human origins over many years, anatomy that are revealing for human evolution. These show that and for curating Ardipithecus ramidus for future research and for all Ardipithecus was at home both moving along trees on its palms and of us to admire. walking upright on the ground. Three characterize Ardipithecus’s Ardipithecus ramidus thus helps us bridge the better-known, habitat in detail, through analysis of the hosting rocks and thousands more recent part of human evolution, which has a better fossil of fossils of small and large animals and plants. These show that record, with the scarcer early human fossils and older ape fossils Ardipithecus lived and ate in woodlands, not grasslands. The first that precede our last common ancestor. Ardipithecus ramidus is a paper presents an overview, and it and the last two papers trace early reminder of Darwin’s conclusion of The Origin: human evolution and synthesize a new view of our last common an- cestor with chimps. One conclusion is that chimps have specialized There is grandeur in this view of life, with its several powers, having greatly since then and thus are poor models for that ancestor and for been originally breathed into a few forms or into one; and that, understanding human innovations such as our ability to walk. whilst this planet has gone cycling on according to the fixed law of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being, evolved. – BrookS HANSON www.sciencemag.org SCIENCE VOL 326 2 October 2009 Published by AAAS 5 The Authors Tim D. White Doris Barboni Laurent Bremond Human Evolution Research CEREGE (UMR6635 Center for Bio-Archaeology Center and Department CNRS/Université and Ecology (UMR5059 of Integrative Biology, Aix-Marseille), BP80, CNRS/Université 3101 Valley Life Sciences F-13545 Aix-en-Provence Montpellier 2/EPHE), Building, University Cedex 4, France.
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