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The Functional and Evolutionary Biology of Primates PDF

534 Pages·2007·35.641 MB·English
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The FUNCTIONAL and EVOLUTIONARY BIOLOGY PRIMATES of RUSSEll TUnlE editor The RlNCTIONAllnd EVOum 0N ARY II0lOOYo1 PRIMATES (cid:39)(cid:74)(cid:83)(cid:84)(cid:85)(cid:1)(cid:81)(cid:86)(cid:67)(cid:77)(cid:74)(cid:84)(cid:73)(cid:70)(cid:69)(cid:1)(cid:18)(cid:26)(cid:24)(cid:19)(cid:1)(cid:67)(cid:90)(cid:1)(cid:53)(cid:83)(cid:66)(cid:79)(cid:84)(cid:66)(cid:68)(cid:85)(cid:74)(cid:80)(cid:79)(cid:1)(cid:49)(cid:86)(cid:67)(cid:77)(cid:74)(cid:84)(cid:73)(cid:70)(cid:83)(cid:84) (cid:49)(cid:86)(cid:67)(cid:77)(cid:74)(cid:84)(cid:73)(cid:70)(cid:69)(cid:1)(cid:19)(cid:17)(cid:18)(cid:24)(cid:1)(cid:67)(cid:90)(cid:1)(cid:51)(cid:80)(cid:86)(cid:85)(cid:77)(cid:70)(cid:69)(cid:72)(cid:70) (cid:19)(cid:1)(cid:49)(cid:66)(cid:83)(cid:76)(cid:1)(cid:52)(cid:82)(cid:86)(cid:66)(cid:83)(cid:70)(cid:13)(cid:1)(cid:46)(cid:74)(cid:77)(cid:85)(cid:80)(cid:79)(cid:1)(cid:49)(cid:66)(cid:83)(cid:76)(cid:13)(cid:1)(cid:34)(cid:67)(cid:74)(cid:79)(cid:72)(cid:69)(cid:80)(cid:79)(cid:13)(cid:1)(cid:48)(cid:89)(cid:80)(cid:79)(cid:1)(cid:48)(cid:57)(cid:18)(cid:21)(cid:1)(cid:21)(cid:51)(cid:47) (cid:24)(cid:18)(cid:18)(cid:1)(cid:53)(cid:73)(cid:74)(cid:83)(cid:69)(cid:1)(cid:34)(cid:87)(cid:70)(cid:79)(cid:86)(cid:70)(cid:13)(cid:1)(cid:47)(cid:70)(cid:88)(cid:1)(cid:58)(cid:80)(cid:83)(cid:76)(cid:13)(cid:1)(cid:47)(cid:58)(cid:1)(cid:18)(cid:17)(cid:17)(cid:18)(cid:24)(cid:13)(cid:1)(cid:54)(cid:52)(cid:34) (cid:51)(cid:80)(cid:86)(cid:85)(cid:77)(cid:70)(cid:69)(cid:72)(cid:70)(cid:1)(cid:74)(cid:84)(cid:1)(cid:66)(cid:79)(cid:1)(cid:74)(cid:78)(cid:81)(cid:83)(cid:74)(cid:79)(cid:85)(cid:1)(cid:80)(cid:71)(cid:1)(cid:85)(cid:73)(cid:70)(cid:1)(cid:53)(cid:66)(cid:90)(cid:77)(cid:80)(cid:83)(cid:1)(cid:7)(cid:1)(cid:39)(cid:83)(cid:66)(cid:79)(cid:68)(cid:74)(cid:84)(cid:1)(cid:40)(cid:83)(cid:80)(cid:86)(cid:81)(cid:13)(cid:1)(cid:66)(cid:79)(cid:1)(cid:74)(cid:79)(cid:71)(cid:80)(cid:83)(cid:78)(cid:66)(cid:1)(cid:67)(cid:86)(cid:84)(cid:74)(cid:79)(cid:70)(cid:84)(cid:84) Copyright © 1972 by Wenner-Gren Foundation for Anthropological Research, Inc. 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Based on the 1970 Burg Wartenstein symposium on “functional and evolu- tionary biology of primates: methods of study and recent advances,” held July 18-26, 1970 at the European Conference Center of the Wenner-Gren Foundation for Anthropological Research. Originally published: Chicago : Aldine-Atherton, 1972. ISBN 978-0-202-36139-0 1. Primates—Congresses. 2. Evolution (Biology)—Congresses. 3. Human evolution—Congresses. I. Tuttle, Russell. QL737.P9F8 2007 599.8—dc22 2007022248 (cid:42)(cid:52)(cid:35)(cid:47)(cid:1)(cid:18)(cid:20)(cid:27)(cid:1)(cid:26)(cid:24)(cid:25)(cid:14)(cid:17)(cid:14)(cid:19)(cid:17)(cid:19)(cid:14)(cid:20)(cid:23)(cid:18)(cid:20)(cid:26)(cid:14)(cid:17)(cid:1)(cid:9)(cid:81)(cid:67)(cid:76)(cid:10) to THEODORE D. MCCOWN Introduction Burg Wartenstein Symposium No. 48 on Functional and Evolutionary Biology of Primates: Methods of Study and Recent Advances convened July 18-26, 1970, at the European Conference Center of the Wenner-Gren Foundation for Anthropological Research near Gloggnitz, Austria. Twenty one scientists participated in six daily sessions, and Professor Adolph H. Schultz honored us with an evening lecture on the history, progress, and prospects of primatological research. Invitations to the conference contained notice of the topics that I wished to be developed by each participant. The invitations also listed the following "statement of general purpose": The major objective of the conference is to bring several perspectives to bear on a variety of problems in evolutionary primatology. We hope to em phasize experimental design, quantification, and the strategies of the compara tive method and intensive longitudinal studies in morphology and behavior. Since the most convincing proofs of the validity of a particular technique or approach are the quality of results obtained through its application and the credibility of inferences that may be based on such studies, participants are asked to emphasize the results and implications of their research for basic theoretical problems. Subsequently, I wrote to each participant, suggesting more specifically the nature of the expositions that they might attempt on the problems selected for discussion at the conference. Five broad groups of basic subject matter on functional and evolutionary primatology were represented at the symposium: paleoprimatology; cranial morphology; comparative neurobiology and endocasts; postcranial mor phology; and aspects of behavior and ecology. In order to treat these five subject areas with an acceptable level of thoroughness, we were regrettably vii viii RUSSELL TUTTLE induced to omit detailed representation of certain other important disci plines, such as biomolecular primatology and prehistory which have been discussed at length in several previous conferences. Paleoprimatology provides the only direct evidence for detailing the phylogenetic histories of primate lineages. The chronology of evolutionary events is wholly dependent upon accurately dated fossils. But because of the fragmentary nature of most fossils, postdeposition perturbations at fossil sites, limited methods of recovery and analysis, the complexity and diversity of datable minerals and their associations with fossils, and many other com plicating factors, paleoprimatologists are somewhat limited on the empirical data that they can extract from available fossils. Thus, like other compara tive evolutionary biologists, they must depend heavily upon inference in constructing models and hypotheses on the course of evolution and the life habits of principals in the primate career. In order to establish broader bases of inference and more complete profiles on hypothetical form and functions in extinct species, paleoprimatologists often consult other branches of com parative evolutionary biology in addition to searching constantly for more fossils. The difficulties that beset the evolutionary primatologist who attempts to infer the mechanical and genetical factors that may be causally related to features of bones and teeth in extant primates are increased many-fold should he turn his attention to extinct species, upon which most kinds of experiments and behavioral observations cannot be conducted. Further, not only the incompleteness of certain fossils but also their sparse representation in available collections may limit the scope of information that may be re covered from the fossils themselves and the nature of quantitative compari sons between them and other taxa. Among evolutionary primatological sciences, investigations on cranial morphology occupy a preeminent position for the documentation of taxo nomic affinities among living and fossil species. Except for certain aspects of the masticatory apparatus, orbital dimensions, and perhaps some neuro cranial dimensions, many features of the skull evidence negligible detectable functional correlates. It is therefore commonly believed among evolutionary biologists that the postcranial skeleton may reflect more fully the functional and ecological particularities of a species than its skull can. Whereas selected features of primate skulls may be expressed quantita tively and persist in fossilized specimens, certain soft structures of the head, in particular the facial muscles, are generally difficult to measure in living forms and leave little impression on underlying bony features. Yet it is these perishable structures which are employed by taxonomists to distinguish colobine from cercopithecine monkeys. One of the greatest sets of challenges that face comparative primatologists and anthropologists is the precise description and evolutionary explanation of particular morphological features in carefully chosen series of primate Introduction ix brains. The evolution of the neurological bases of human speech, tool making capacities, and other symbolic behaviors also remain poorly eluci dated today. Two basic kinds of evidence may be brought to bear on problems of brain evolution in primate lineages, viz., results of comparative studies on brains of extant species and comparative studies on fossil and extant primate endocasts. Both approaches entail a great amount of painstaking effort in order to accumulate representative comparative and evolutionary series. Comparative neurobiologists and paleoneurobiologists must possess a broad knowledge of neurophysiology, naturalistic behavior, and ecology in living primates in order to explain the results of their studies in functional and evolutionary contexts. Whereas comparative neurobiologists studying extant primates may ex amine both external and internal neural features, the paleoneurobiologist is restricted to available surface features of naturally occurring fossil endocasts and endocasts prepared in the laboratory from fossil crania. There are no known instances of brains themselves being fossilized such that paleohis tological studies could be conducted on them. The occurrence of such fossil specimens is not to be anticipated due to the paucity of supporting connec tive tissues and concordant rapid postmortem degeneration of the brain. Studies on the postcranial morphology of primates and especially investi gations on primate locomotive systems have long been of central interest to evolutionary anthropologists and other biologists. During the past two decades, a profusion of experimental and analytical techniques have been developed and refined which not only permit but also necessitate their several employment in functional and evolutionary studies on primate loco motion and other motile behaviors. Studies on the morphology of extant and fossil primates, in view of biomechanical principles known to be operant in living systems, may gen erate credible hypotheses on the functions of selected parts of an individual and perhaps on certain aspects of its life habits. But functional models based on morphology alone are greatly limited in scope and often remain equivocal until they are confirmed, revised, or rejected on the basis of naturalistic behavioral observations, laboratory behavioral studies, or both. The increased number and intensity of behavioral studies on nonhuman primates, though unfortunately rarely focused on posture and locomotion, are providing somewhat firmer bases for functional interpretations of mor phological features than were possible during the more strictly anecdotal era of primate field studies. Direct observations of behavior and ecology are impossible for extinct organisms, so we must carefully delimit the nature of variations in the be havior and ecology of living primates and attempt to determine the extent to which uniformitarian arguments about these features might be employed to formulate refined models on the life modes of each fossil species.

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