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Part II - Organ structure, function, and behavior
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- By Fred Anapol, Professor in the Department of Anthropology (adjunct in Biological Sciences) and the Director of the Center for Forensic Science University of Wisconsin–Milwaukee, Rebecca Z. German, Professor in Biological Sciences University of Cincinnati, Nina G. Jablonski, Chair and Curator of Anthropology California Academy of Sciences, Charles Oxnard
- Edited by Fred Anapol, University of Wisconsin, Milwaukee, Rebecca Z. German, University of Cincinnati, Nina G. Jablonski, California Academy of Sciences, San Francisco
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- Shaping Primate Evolution
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- 10 August 2009
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- 20 May 2004, pp 97-98
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Summary
My own earliest studies tried to understand functional adaptation in the locomotor system, primarily of bones and joints, through morphometrics. As a preliminary, however, I did attempt to find out what I could of the muscles that moved the bone–joint unit. In those days all we did was dissect muscles and measure them through relative lengths, directions of pull, relative weights, and frequencies of (what were called in those days) muscular anomalies. Limited though such studies were, they were incredibly time-consuming. In all, I dissected 52 shoulders representing 28 primate species, 145 arms and forearms representing 27 primate species, and 167 hips and thighs in 33 primate species. That took many years. Yet even such primitive data provided useful initial information about muscles in relation to the respective bone–joint units.
But it was always clear to me, and I wrote about it without ever doing it, that better muscular studies would be necessary. Such studies would need to understand much more about muscular architecture than simply relative muscle weights, much more about muscle activity and functions through studies of living muscle than just anatomical inferences, much more about movements, postures, and overall behaviors of the living animals than just simple classifications of locomotion.
However, I certainly did not, in those early days, envisage the possibility of studies of the biomechanics of the jaws and teeth and, further, the biomechanics of food being masticated by them. And I knew nothing at all of functional adaptations in the brain.
A few examples of such investigations follow in this section. Again, my students and colleagues have gone so much further than was possible for me. Yet notwithstanding, stimulated by them, I have been able in these latter years to enter some of these areas myself.
Index
- Edited by Fred Anapol, University of Wisconsin, Milwaukee, Rebecca Z. German, University of Cincinnati, Nina G. Jablonski, California Academy of Sciences, San Francisco
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- Shaping Primate Evolution
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- 10 August 2009
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- 20 May 2004, pp 420-426
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2 - The ontogeny of sexual dimorphism: the implications of longitudinal vs. cross-sectional data for studying heterochrony in mammals
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- By Rebecca Z. German, Department of Biological Sciences, University of Cincinnati, Cincinnati, OH 45221-0006, USA
- Edited by Fred Anapol, University of Wisconsin, Milwaukee, Rebecca Z. German, University of Cincinnati, Nina G. Jablonski, California Academy of Sciences, San Francisco
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- Shaping Primate Evolution
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- 10 August 2009
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- 20 May 2004, pp 11-23
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Introduction
As the name suggests, studies of sexual dimorphism began with a focus on morphological differences between the sexes (Darwin 1871). Current use of the term “dimorphism” and current studies of sexual dimorphism have expanded to include ecological, behavioral, and physiological differences between the sexes (Harvey and Clutton-Brock, 1985). Charles Oxnard (1987) brought his unique quantitative perspective to the investigation of sexual dimorphism, showing that studies, particularly quantitative studies, of differences between the sexes in morphology are meaningful and not outdated. His work has provided inspiration for this chapter, which examines the role that data and analysis play in understanding evolution. As Oxnard identified multiple dimorphisms among taxa along morphological axes, this study examines heterochronic variation among taxa to show that different ontogenetic trajectories produced analogous multiple dimorphisms. Crucial to Oxnard's work, and to the results presented here, are matches among question, data, and method.
Studies of sexual dimorphism and growth
Most research addressing questions of growth and sexual dimorphism examines the ontogeny of that dimorphism, focusing on how growth produces adult differences (see German and Stewart, 2002 for review). A slight shift in focus to the sexual dimorphism of ontogeny, or the way the sexes grow, will generate alternative questions, centering on the differences in growth itself (e.g., Watts and Gavan, 1982; Glassman et al., 1984; Coelho, 1985; Watts, 1986).
Growth in mammals has a number of distinguishing characteristics that make comparisons of growth between two groups difficult, whether they are sexes or species. Mammalian growth is nonlinear.
Preface: shaping primate evolution
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- By Fred Anapol, Professor in the Department of Anthropology (adjunct in Biological Sciences) and the Director of the Center for Forensic Science University of Wisconsin–Milwaukee, Rebecca Z. German, Professor in Biological Sciences University of Cincinnati, Nina G. Jablonski, Chair and Curator of Anthropology California Academy of Sciences
- Edited by Fred Anapol, University of Wisconsin, Milwaukee, Rebecca Z. German, University of Cincinnati, Nina G. Jablonski, California Academy of Sciences, San Francisco
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- Shaping Primate Evolution
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- 10 August 2009
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- 20 May 2004, pp xv-xvi
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The last half-century has witnessed a dramatic improvement in our understanding of the relationship between form and function in biology. This phenomenon has been fueled by innovations in many fields, from molecular biology to mechanical engineering and multivariate statistics. The importance of “big thinkers” has also been critical – people who can understand and synthesize information from diverse technical fields and then apply and integrate such information in the context of organismal biology. The theoretical and substantive innovations brought about by “big thinkers” such as Charles E. Oxnard have transformed the last half-century of zoology and anthropology.
Charles Oxnard's fascination with size and form in primates shaped not only his career, but also the careers of students and colleagues. This volume is a tribute to how that fascination has influenced numerous others. Many of the highly regarded, internationally known contributors to this volume participated in a recent symposium (American Association of Physical Anthropologists, April 2000) honoring Professor Oxnard for his many and various contributions to the study of primate evolutionary morphology. Each contributor has been influenced strongly by Professor Oxnard and each contribution elaborates on the analysis of the form–function–behavior triad in a unique and compelling way.
This book is diverse both in the topics covered and in the range of levels of biological organization that are addressed, from the cellular level (Jouffroy and Médina) to the evolution of primate ecology (Fleagle and Reed).
Contents
- Edited by Fred Anapol, University of Wisconsin, Milwaukee, Rebecca Z. German, University of Cincinnati, Nina G. Jablonski, California Academy of Sciences, San Francisco
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- Shaping Primate Evolution
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- 10 August 2009
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- 20 May 2004, pp ix-xi
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List of contributors
- Edited by Fred Anapol, University of Wisconsin, Milwaukee, Rebecca Z. German, University of Cincinnati, Nina G. Jablonski, California Academy of Sciences, San Francisco
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- Shaping Primate Evolution
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- 10 August 2009
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- 20 May 2004, pp xii-xiv
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Part I - Craniofacial form and variation
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- By Fred Anapol, Professor in the Department of Anthropology (adjunct in Biological Sciences) and the Director of the Center for Forensic Science University of Wisconsin–Milwaukee, Rebecca Z. German, Professor in Biological Sciences University of Cincinnati, Nina G. Jablonski, Chair and Curator of Anthropology California Academy of Sciences, Charles Oxnard
- Edited by Fred Anapol, University of Wisconsin, Milwaukee, Rebecca Z. German, University of Cincinnati, Nina G. Jablonski, California Academy of Sciences, San Francisco
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- Shaping Primate Evolution
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- 10 August 2009
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- 20 May 2004, pp 9-10
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The study of craniofacial form and variation has always been one of the most important areas for those interested in shaping primate evolution. Skulls were the most frequently collected specimens in museums. Skull parts, especially teeth, are most frequently found in the fossil record. Skulls and teeth are easily examined in the living. The bones of the face allow some estimation of how their owners appeared. Appearance and change in appearance as produced by medical and dental technologies have profound effects upon individual well-being. All these are good reasons why this is one of the most critical of anatomical regions.
At the same time, however, skulls, faces, jaws, and teeth are the most complex region of the body. More, perhaps, than in any other region, do a number of completely different functions have to be integrated in its structure. The genetics underlying cranium, face, jaw, and teeth are even now not well known and clearly far more complicated than the postcranium. The development and growth of the head depends upon complex mechanisms and processes, many of which have only been elucidated in the last two decades. In evolutionary terms, the “head problem” in chordates, reflecting at the same time both very ancient and very recent elements, has always been more difficult to understand than, say, the equivalent trunk problem or limb problem (which problems do not even rate quotation marks).
Frontmatter
- Edited by Fred Anapol, University of Wisconsin, Milwaukee, Rebecca Z. German, University of Cincinnati, Nina G. Jablonski, California Academy of Sciences, San Francisco
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- Shaping Primate Evolution
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- 10 August 2009
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- 20 May 2004, pp i-viii
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Part IV - Theoretical models in evolutionary morphology
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- By Fred Anapol, Professor in the Department of Anthropology (adjunct in Biological Sciences) and the Director of the Center for Forensic Science University of Wisconsin–Milwaukee, Rebecca Z. German, Professor in Biological Sciences University of Cincinnati, Nina G. Jablonski, Chair and Curator of Anthropology California Academy of Sciences, Charles Oxnard
- Edited by Fred Anapol, University of Wisconsin, Milwaukee, Rebecca Z. German, University of Cincinnati, Nina G. Jablonski, California Academy of Sciences, San Francisco
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- Shaping Primate Evolution
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- 10 August 2009
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- 20 May 2004, pp 279-280
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Though I had always been interested in the use of mathematical methods of one kind or another, in the earlier days these interests were almost always for data analysis. The models of function with which we worked in the early days were simplistic in the extreme, scarcely deserving of the term “model” – in fact they were little more than functional anatomical inferences. At later stages I used models for assessing mechanical efficiency, but these mainly employed what is now a quite old-fashioned technique – photoelastic analysis – and they were extremely limited – e.g., to two dimensions only, and only to isotropic situations. Only much more recently, in collaboration with others (e.g., O'Higgins), have I come to use better modeling methods (such as finite elements).
However, in a completely surprising way, through a stimulus applied by Sydney Brenner and his invitation to present a model of mtDNA evolution (which I did not do) at a workshop hosted by the International Institute for Advanced Studies in Kyoto, I have come to be involved in true evolutionary modeling (mimicking species evolution and individual lineages) with Dr. Ken Wessen. Though not included as a section in this book, Dr. Wessen's work (in which I have been pleased to share) is reported in my own final chapter.
At this point, however, it is an especial pleasure to recognize, through the following sections, kinds of modeling that I scarcely envisaged in those earlier days. Thus the following chapters on modeling the origins and mechanics of bipedalism, and the mechanics of mastication, are extremely relevant.
Shaping Primate Evolution
- Form, Function, and Behavior
- Edited by Fred Anapol, Rebecca Z. German, Nina G. Jablonski
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- 10 August 2009
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- 20 May 2004
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Shaping Primate Evolution is an edited collection of papers about how biological form is described in primate biology, and the consequences of form for function and behavior. The contributors are highly regarded internationally recognized scholars in the field of quantitative primate evolutionary morphology. Each chapter elaborates upon the analysis of the form-function-behavior triad in a unique and compelling way. This book is distinctive not only in the diversity of the topics discussed, but also in the range of levels of biological organization that are addressed from cellular morphometrics to the evolution of primate ecology. The book is dedicated to Charles E. Oxnard, whose influential pioneering work on innovative metric and analytic techniques has gone hand-in-hand with meticulous comparative functional analyses of primate anatomy. Through the marriage of theory with analytical applications, this volume will be an important reference work for all those interested in primate functional morphology.
Part V - Primate diversity and evolution
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- By Fred Anapol, Professor in the Department of Anthropology (adjunct in Biological Sciences) and the Director of the Center for Forensic Science University of Wisconsin–Milwaukee, Rebecca Z. German, Professor in Biological Sciences University of Cincinnati, Nina G. Jablonski, Chair and Curator of Anthropology California Academy of Sciences, Charles Oxnard
- Edited by Fred Anapol, University of Wisconsin, Milwaukee, Rebecca Z. German, University of Cincinnati, Nina G. Jablonski, California Academy of Sciences, San Francisco
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- Shaping Primate Evolution
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- 10 August 2009
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- 20 May 2004, pp 351-352
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Once again, though my earlier work was clearly aimed at understanding functional adaptations in specific bone–joint–muscle units, it also equally clearly led into interests in primate diversity and evolution. I was not, of course, a primate taxonomist, never having had the requisite training. I never worked in field situations (being allergic to high temperatures, heavy rainfall, high altitudes, mosquitoes, leeches, etc). I therefore never participated in field observations of living species or in field discoveries of fossils. And though I was never formally educated in mathematics and statistics, and never capable myself of making advances in them, I was always a user who was interested in how such methods could be applied to data. Especially was I interested in the kinds of questions that the above methods and data might answer.
I have thus remained enormously interested in all such studies carried out by others – indeed, the data of others were essential to some of the investigations that I myself made. Mainly working with colleagues, however, I may have been perhaps the first to apply full multivariate statistical analyses to the data of field observation, of the niche. Likewise, through colleagues and students, I may have been amongst the first to use morphometric methods as tools to go beyond the data themselves, to seek correlations with behavior, with the niche, with development, and with evolution. The following chapters on the niche, on cladistics, and on the development of morphometrics itself carry all this so much further. These studies (and many others not represented in this book) go so very far beyond what I originally envisaged.
Part III - In vivo organismal verification of functional models
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- By Fred Anapol, Professor in the Department of Anthropology (adjunct in Biological Sciences) and the Director of the Center for Forensic Science University of Wisconsin–Milwaukee, Rebecca Z. German, Professor in Biological Sciences University of Cincinnati, Nina G. Jablonski, Chair and Curator of Anthropology California Academy of Sciences, Charles Oxnard
- Edited by Fred Anapol, University of Wisconsin, Milwaukee, Rebecca Z. German, University of Cincinnati, Nina G. Jablonski, California Academy of Sciences, San Francisco
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- Shaping Primate Evolution
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- 10 August 2009
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- 20 May 2004, pp 227-228
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Though it would appear from my early work that I primarily used anatomical inference for the “functional” explanations for the morphological, especially morphometric, adaptations that I saw in my studies, I nevertheless was always interested in the “real” biomechanics, interested in testing, that is, the hypotheses resulting from anatomical inference. The best that we could do at the time for the shoulder was to rely on the pioneering electromyographic studies on the human shoulder carried out by Inman, Saunders, and Abbott during the Second World War. Doing electromyography on nonhuman primates seemed such a long shot in those days.
Yet in fact my first research grant (from the US Public Health Service as it was called then), awarded in 1962 while I was still in the UK, aimed in part to design an implantable telemetric device for recording electromyographic and strain-gauge information from freely moving primates. Stanley Salmons was the research fellow employed on that grant for that purpose. But of course, only a few weeks' reading was enough to demonstrate that we could not do it – it just was not possible to make the device small enough for that purpose in those days. Stanley Salmons, however, did go on to design the “buckle transducer'” a first device for measuring tension in tendons, so all was not lost. He is today Professor of Biomedical Engineering at the University of Liverpool and has been engaged in these latter years in studies making the latissimus dorsi muscle into an adjunct pump in cases of cardiac insufficiency.
Ontogenetic allometry in the locomotor skeleton of specialized half-bounding mammals
- Andrew R. Lammers, Rebecca Z. German
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- Journal of Zoology / Volume 258 / Issue 4 / December 2002
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- 13 November 2002, pp. 485-495
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- December 2002
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Specialization for a locomotor behaviour may affect limb bone morphology throughout ontogeny. Ontogenetic development of the limb skeletons of two mammalian species, which are behaviourally specialized for the half-bounding gait (Chinchilla lanigera and Oryctolagus cuniculus), were compared to two similarly-sized species which are not specialized for half-bounding (Rattus norvegicus and Monodelphis domestica). Limb bone lengths and anteroposterior diameters (mediolateral diameters for the radius and metacarpal) were measured from radiographs taken throughout the ontogeny of each species. Body mass was also measured repeatedly during growth. Bone measurements were regressed against body mass, as well as forelimb bone length vs serially homologous hindlimb bone length, bone length vs total limb length and bone length vs width. Similar comparisons were made among adults of each species using ratios. Although there were many significant differences among species, overall there were few consistent differences in adult scaling ratios or ontogenetic allometry slopes between specialized and generalized groups. Adult specialized half-bounders had significantly narrower tibiae and metatarsals than the gait-generalized runners. Specialized half-bounders usually had similar slopes for hindlimb length vs width ontogenetic comparisons, but the non-specialized species did not group together. However, there were two patterns that occurred among all four species: (1) hindlimb bone lengths nearly always grew faster than the serially homologous forelimb bone lengths in all species; (2) proximal elements usually increased in length proportionally faster than distal elements. In conclusion, small mammals may share strong developmental constraints that govern their relative growth rates. It is also likely that there are different selective pressures on juveniles and adults, but that these selective pressures may not be different between specialized and unspecialized runners during ontogeny.