Book Reviews 681

The Anatomical Waxes by Clemente Susini of the species of early hominids coexisted: therefore as a group University of Cagliari. (Bilingual Italian/English hominids were much more diverse during their evolutionary history than they are today. Much the same can be said for edition.) By LUIGI CATTANES and ALESSANDRO Theropithecus. The first fossil evidence is known from the RIVA. (Pp. 70; 25 colour plates; 80,000 lire; ISBN Hadar and Usno Formations in Ethiopia, and from the 88 7343 254 9.) University of Cagliari; Edizioni Koobi Fora Formation in Kenya, and the earliest remains date from around 3.6 ma (and are perhaps as old as 4 ma if della Torre. 1993. fossils from the Chemeron Formation are included). These This beautifully produced book is the 2nd edition of the fossil Theropithecus remains are assigned to 1 of 2 extinct catalogue of the University of Cagliari's small but valuable lineages; the modern gelada has no fossil record. In the first collection of anatomical wax models made by Susini and his of these lineages at least 2 time-successive species are coworkers. These were prepared in Florence between 1803 recognised; Theropithecus darti is the early species which is and 1805, and were the last of over 2000. The majority are replaced by Theropithecus oswaldi. This lineage spans the housed in the Museum of the Speculo in Florence and the time interval between 3.2 and 0.4 ma. The second lineage Josephinum in Vienna. A few are in other European and consists of a single species, Theropithecus brumpti, although Italian Anatomical Institutes. some regard T. brumpti as having evolved from the poorly The catalogue is preceded by an interesting history of the known species, Theropithecus baringensis. If so, the T. acquisition of the collection and a brief account of the brumpti lineage may be as old as 4 ma, but T. brumpti itself techniques employed in creating these superb models. The is known only from between 2.7 and 2 ma. Its geographical colour photographs by Alessandro Cadau and Alessandro distribution is restricted to sites within the Turkana basin in Riva and their reproduction are of the highest quality, and southern Ethiopia and northern Kenya. do justice to the artistry and outstanding skill of the The extinct species T. brumpti is one of the most unusual dissectors and modellers. Some of the details are not fossil primates known. It was at least as large as modern accurate, but those interested in the history of anatomy and chimpanzees and like them (and unlike T. oswaldi and the art of model-making will enjoy this book despite the rather modern gelada) it was adapted to tree-living. It was not a stilted English translation of the accompanying Italian 'small-object' feeder, like both T. oswaldi and the modern text. gelada, and its large gape suggests that T. brumpti was RUTH E. M. BOWDEN adapted to chewing on large hard objects such as fruits or tubers. Whereas males of the modern gelada weigh on Theropithecus: The Rise and Fall of a Primate Genus. average 20 kg, the size of the skulls and limb bones of T. oswaldi and T. brumpti suggest that males of the former Edited by NINA G. TABLONSKI. (Pp. xvi + 536; species may have weighed as much as 60 kg and males of the illustrated; £110 hardback; ISBN 0 521 41105 X.) latter species betweenA40 and 50 kg. These were substantial, Cambridge: Cambridge University Press. 1993. strongly built animals, equipped with impressive canine teeth. The artist's reconstruction on the dust sheet of the Just under 160 years ago the naturalist Ruppell gave the book captures very well the probable appearance of these genus name Theropithecus (meaning 'beast monkey') and early Theropithecus species. the species name gelada (which is the local Amharic name Fossils belonging to the 2 extinct Theropithecus lineages for these animals) to a species of baboon-like monkeys from are at least as common as hominids in the collections East Africa. Their habitat is not one we normally associated recovered from the major hominid fossil sites of East Africa. with primates, for the 'geladas' live in the gorges and How, then, do we account for their extinction? The escarpments of the Ethiopian highlands. shrinking of the forests associated with the gradual cooling Most gelada populations live at around 2000-3000 m, but and drying of the global, and particularly the African they have been observed at altitudes as high as 4500 m. climate, between 3 and 2 ma probably accounted for the Despite being close to the equator, habitats at these altitudes demise of T. brumpti. The fate of T. oswaldi is more difficult are cold (5°-10°) and wet. The geladas use the cliff-faces as to account for, but it is probable that the combination of sleeping sites and move onto the high plateaux to feed. Even their large body-size and a diet of low-energy foods was more unusually for primates, geladas are grazing animals their downfall. The modern geladas are probably a relic whose predominant food is the grasses which flourish in population closely related to, but not directly descended these cold, damp conditions. Geladas are related to modern from, the T. oswaldi lineage. baboon species with the genus Papio, and evidence from Nearly all of the 19 papers in this volume were presented molecular anthropology suggests that the ancestors of Papio at a conference about the evolutionary history of Thero- and Theropithecus parted company between about 7 and 32 pithecus that took place in Cambridge in 1990. The potential million years (ma) ago. Nonetheless, geladas are distinctive of Theropithecus to provide insights into the life and times of enough to continue to justify their inclusion in a separate our own ancestors was recognised by the editor of this genus. volume, Nina Tablonski, who organised the Cambridge So what is it about these peculiar monkeys that has meeting with Rob Foley. The breadth and high quality of prompted the publication of this handsome and valuable the papers are a credit to their perspicacity and hard work. volume? And does a book about the evolutionary biology of The papers fall into 4 groups, the largest of which is devoted a species of monkey justify a review in this journal? The to documenting the morphology and interpreting the unusual habitat and adaptations of Theropithecus gelada are functional anatomy of the extinct species of Theropithecus. interesting in their own right, but these creatures attract this The remaining 7 papers deal with the social organisation unusual degree of attention because the evolutionary history and the ecology of the modern geladas and the extent to of the genus Theropithecus was played out in much the same which their lifestyle can be extrapolated to the extinct geographical area, and during much the same time period, species. Some papers do explicitly what many others do as our own. implicitly: set out hypotheses about the factors which Human predecessors, or hominids, apparently emerged in shaped the evolutionary history of Theropithecus. Africa around 5-4 ma ago. At one time or another, up to 4 There is 'real' anatomy in abundance in this volume. It 682 Book Reviews ranges from detailed observations about dental devel- Theropithecus may have resembled that of Paranthropus opment, and tooth microwear, to the way that intramuscular species, known as the 'robust' australopithecines, among tendons within the masseter affect the size of gape. The the early hominids, there is, in Theropithecus, no evidence of cranium, jaws and teeth form the focus of the anatomical any increase in relative brain size of the sort that is found in interpretations, but the postcranial skeleton is not ignored. the early hominids. Jablonski, the editor, provides an Many new data are presented, but it is unfortunate that excellent overview as well as a useful commentary on the contributors were not encouraged to present the data evolution of the masticatory apparatus in Theropithecus, specimen by specimen, though the remains from each site and Lee and Foley provide a compelling reconstruction of are clearly set out in the appendix. A composite time-chart the ecological energetics of the extinct T. oswaldi. plotting the respective ages ofall the major specimens would This book would have benefitted from the inclusion of also have allowed readers to follow changes in variables, more authors prepared to make connections between such as tooth size, through time. hominid evolution and Theropithecus species. However, Only 2 papers, one by Foley and the other by Martin, much of what it does include is very worthwhile palaeo- seriously pursue the analogy between Theropithecus and biology which has been applied to the study of this group of hominid evolution. Foley restricts his analysis to the ways dietarily specialised, large-bodied, monkeys. This volume that hominids and Theropithecus may have been responsive gives some insight into why, in the USA, evolutionary to climatic perturbations. He concludes that it was the biology provides the research focus for some of the most frequency of climate fluctuations, and not the extent of the successful anatomy departments. Morphological studies are fluctuations themselves, which may have stimulated episodes alive and well thanks to an upsurge of interest in of species generation and extinction within the Hominidae evolutionary history. and Theropithecus. Martin reminds us that while the diet of BERNARD WOOD