The Lawrence Jacobsen Library
Books Received (Primate-Science/PrimateLit)

HUMAN EVOLUTION THROUGH DEVELOPMENTAL CHANGE

Edited by
Nancy Minugh-Purvis
and
Kenneth J. McNamara


The Johns Hopkins University Press 2002



CONTENTS


Foreword ix
F. Clark Howell


Preface xiii


List of Contributors xvii


What Is Heterochrony? 1
Kenneth J. McNamara


Part I: Evolution and Development


1. Evolutionary Developmental Biology: Where Embryos and Fossils Meet 7
Brian K. Hall


2. Shape and Stage in Heterochronic Models 28
David M. Alba


3. Multivariate Approaches to Development and Evolution 51
Gunther J. Eble


4. Are Some Heterochronic Transformations Likelier than Others? 79
Brian T. Shea


5. Sequential Hypermorphosis: Stretching Ontogeny to the Limit 102
Kenneth J. McNamara


6. Animal Domestication and Heterochronic Speciation: The Role of Thyroid Hormone 122
Susan J. Crockford


7. The Role of Heterochrony in Primate Brain Evolution 154
Sean H. Rice


Part II: The Evolution of Hominid Life-History Patterns


8. Brain Evolution by Stretching the Global Mitotic Clock of Development 173
Michael L. McKinney


9. Natural Selection and the Evolution of Hominid Patterns of Growth and Development 189
Nina J. Jablonski, George Chaplin, and Kenneth J. McNamara


10. Sexual Dimorphism and Ontogeny in Primates 207
Rebecca Z. German and Scott A. Stewart


11. Life-History Evolution in Miocene and Extant Apes 223
Jay Kelley


12. Dental Development and Life History in Hominid Evolution 249
Robert L. Anemone


13. An Assessment of Radiographic and Histological Standards of Dental Development in Chimpanzees 281
Kevin L. Kuykendall


14. Evolutionary Relationships between Molar Eruption and Cognitive Development in Anthropoid Primates 305
Sue Taylor Parker


Part III: The Evolution of Hominid Development


15. Enamel Microstructure in Hominids: New Characteristics for a New Paradigm 319
Fernando Ramirez Rozzi


16. Cranial Growth in Homo erectus 349
Susan C. Anton


17. Peramorphic Processes in the Evolution of the Hominid Pelvis and Femur 381
Christine Berge


18. Heterochrony and the Evolution of Neandertal and Modern Human Craniofacial Form 405
Frank L'Engle Williams, Laurie R. Godfrey, and Michael R. Sutherland


19. Adolescent Postcranial Growth in Homo neanderthalensis 442
Andrew J. Nelson and Jennifer L. Thompson


20. Between the Incisive Bone and Premaxilla: From African Apes to Homo sapiens 464
Bruno Maureille and Jose Braga


21. Heterochronic Change in the Neurocranium and the Emergence of Modern Humans 479
Nancy Minugh-Purvis


Glossary 499


Index 501



PREFACE


In recent years, biologists, paleontologists, and paleoanthropologists, separately and together, have come to recognize
development as an indispensable ingredient of evolutionary change. This would have been no surprise to our colleagues of
the nineteenth century, many of whom merged the phenomena of ontogeny and phylogeny into a single, intricate continuum.
But as the new synthesis began to reshape views of evolution in the 1920s and 1950s, genetic mechanisms were quickly
favored over developmental phenomena as the driving forces of evolution. Not until the close of the twentieth century did
an awareness resurface that the link between phenotype and genotype is development, that the ontogenetic perspective is
the new synthesis through which twenty-first century science will understand evolutionary change.


During the 1970s paleoanthropologists began to excavate museum archives as well as geological strata as interest grew in
the large number of long-ignored immature hominid remains languishing in museum storage. Typically more fragmentary than
the remains of adults, infant, juvenile, and adolescent hominid fossils posed the additional annoyance of not always
exhibiting the morphology seen in adults and so crucially needed for the typological studies traditionally characterizing
paleoanthropology. By the 1980s, however, as controversy over the nature of modern human origins fermented and the
necessity for studies beyond morphological analysis of adult phenotype became self-evident, interest in developmental
studies in paleoanthropology intensified.


During this same period, remarkable strides in molecular biology led to the emergence of the discipline of evolutionary
developmental biology. For the first time, evolutionary relationships between taxa, previously suspected on the basis of
paleontological research, could be tested in the laboratory and the ontogenetic links leading to the appearance of new
phenotypes could be demonstrated in the lab as well. Moreover, it became apparent that this generation of new phenotypes
this evolutionary patterning-was due to variations in developmental rate and timing, known as heterochrony, operating at
levels from molecules to organ systems.


In 1996, encouraged by Solomon Katz, former head of the Anthropology Section for the American Association for the
Advancement of Science (AMS), Nancy Minugh-Purvis and Michael McKinney organized a symposium, "Heterochrony: Merging
Evolutionary Perspectives in Biology, Paleontology, and Paleoanthropology," attempting further interdisciplinary
dialogue on heterochrony between evolutionary biologists from various fields. Twelve papers presented at the Baltimore
AAAS meeting discussed topics ranging from the effect of heterochronic patterning on the evolution of behavior to
morphogenesis. Even in the planning stages, the intensity and enthusiasm of exchanges between the participants indicated
the need for a book on the subject in which students of evolution from various subdisciplines could share their different
approaches to common problems.


When, in late 1997, other commitments prevented Mike McKinney from continuing as a coeditor, Ken McNamara enthusiastically
jumped into coeditorship. Several of the original participants were unable to contribute to this volume, but we are pleased
to have included some chapters by authors who were unable to attend the original meeting. This, we believe, has resulted
in a well-balanced examination of heterochronic theory and its application to current issues in hominid evolution.


The book is organized into three parts focusing on, first, more theoretical applications of heterochronic theory to
the hominid fossil record; second, the relationship of developmental change to aspects of hominid life history; and,
third, the role of heterochronic change in the evolution of Pliocene to late Pleistocene hominids. In Part I, "Evolution
and Development," after a brief introduction to heterochrony, Brian Hall introduces the reader to the biological
basis of heterochronic change and David Alba, Gunther Eble, Brian Shea, and Ken McNamara discuss theoretical frameworks
for heterochronic analysis. Part I ends with the contributions of Susan Crockford and Sean Rice, which look at biological
complexes that influence the evolution of behavior. Crockford provides a provocative, testable hypothesis on the possible
role of the thyroid in heterochronic patterning, and Rice looks at heterochronic modeling as it applies to the most
distinctively complex human feature-the brain.


Part II, "The Evolution of Hominid Life History Patterns," picks up where Part I left off. It begins with Mike
McKinney's engaging examination of human brain evolution, which is followed by Nina Jablonski, George Chaplin, and Ken
McNamara's chapter investigating the evolution of unique signal characteristics of the earliest hominids, such as
bipedalism, increased body size, and increased brain size. Rebecca German and Scott Stewart examine important, but often
neglected, questions concerning the ontogeny of sexual dimorphism and its role in hominid evolution. Exploiting the rich
dental fossil record, Jay Kelley examines the selective forces behind life-history evolution in Miocene apes in comparison
with living apes, including humans. Robert Anemone examines dental development as an indicator of life-history patterns in
early hominids, and Kevin Kuykendall discusses standards for assessing the dental evidence for life history in living
African apes. Sue Parker's chapter completes this section with an interesting model for the examination of comparative
cognitive development in primates, including hominids, based on extrapolations of neurological maturity from the evidence
for dental development.


Part III, "The Evolution of Hominid Development," provides a chronological survey of heterochronic change in
the hominid fossil record from the Pliocene to the late Pleistocene. Fernando Ramirez Rozzi's examination of differences
in dental development provides thought-provoking evidence for heterochronic diversity early in the hominid fossil record.
Christine Berge's chapter, as well as that of Andrew Nelson and Jennifer Thompson, provide rare glimpses of heterochrony
in the hominid postcranial complex-a difficult area of study, given the extremely rare and incomplete nature of immature
postcrania, but a record that they demonstrate is, nevertheless, a valuable source of developmental data. A developmental
study of the Homo erectus cranium by Susan Anton challenges some traditional interpretations of the Chinese and
Indonesian samples. Maureille and Braga provide an interesting study tracing developmental changes in the premaxillary
suture from early to later hominids. Frank Williams, Laurie Godfrey, and Michael Sutherland find considerable differences
in the growth patterning of the craniofacial complex of Neandertals and modern humans, providing an interesting contrast
to the final chapter of the book, in which Nancy Minugh-Purvis finds continuity of heterochronic change in the
neurocranium from Neandertals to modern humans in the Upper Pleistocene.


A few comments are needed regarding editorial practices in this volume. We have left the ever-contentious spelling of
the term for late Pleistocene archaic Europeans to the discretion of our individual authors. As a result, readers will
encounter Neandertal where authors have followed the convention of omitting the h, which was removed from the word thal
by modern German orthography; others, particularly English colleagues, prefer the term Neanderthal, which retains the h
persisting in the taxonomic nomen Homo sapiens neanderthalensis. Similarly, we have not been dogmatic editors in the
formal naming of the Kenyan hominid specimen KNM-WT 15000, formerly known colloquially as Turkana Boy, but now called
Nariokotome Boy. Some authors prefer to assign him to H. erectus, whereas others call him H. ergaster. There are also
some differences of opinion concerning heterochronic terminology (as there have been for over a hundred years). In
Chapter 2 David Alba discusses the variants in terminology in use at present. While we have our own predilections, we
refrain from wielding the editorial big stick but have tried to ensure that each author clearly explains the meaning of
the term that he or she is using. Our aim, of course, is to enlighten, not create confusion.


Finally, we sincerely thank all of our contributors, but especially F. Clark Howell for his thoughtful foreword and Mike
McKinney, who was instrumental in organizing the original AAAS conference and this book and whose title this volume bears.
We also thank the many outside reviewers who took the time to read and provide useful comment on the chapters contained
herein and the enthusiastic and capable editorial staff at Johns Hopkins University Press: Ginger Berman, early in this
project, and Wendy Harris, in the later stages. Ken McNamara also thanks Danielle West for her help with some of the
illustrations, and Nancy Minugh-Purvis thanks Doug Purvis for his love, support, and patience throughout this project.



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