Evolutionary Biology VOLUME 21

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1 Evolutionary Biology VOLUME 21

2 A Continuation Order Plan is available for this series. A continuation order will bring delivery of each new volume immediately upon publication. Volumes are billed only upon actual shipment. For further information please contact the publisher.

3 Evolutionary Biology VOLUME 21 Edited by MAXK.HECHT Queens College of the City University of New York Flushing, New York BRUCE WALLACE Virginia Polytechnic Institute and State University Blacksburg, Virginia and GHILLEANT. PRANCE New York Botanical Garden Bronx, New York PLENUM PRESS. NEW YORK AND LONDON

4 The Library of Congress cataloged the first volume of this title as follows: Evolutionary biology. v New York, Appleton-Century-Crofts. v. illus. 24 cm annual. Editors: T. Dobzhansky and others. 1. Evolution-Period. 2. Biology-Period. I. Dobzhansky, Theodosius Grigorievich, QH366.AIE9 575' ISBN DOl / ISBN (e8ook) 1987 Plenum Press, New York Softcover reprint of the hardcover 1st edition 1987 A Division of Plenum Publishing Corporation 233 Spring Street, New York, N.Y All rights reserved No part of this book may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher

5 Contributors Jon E. Ahlquist Department of Biology and Peabody Museum of Natural History, Yale University, New Haven, Connecticut Present Address: Irvine Hall, Ohio University, Athens, Ohio Jean Chaline Laboratoire de Geodynamique Sedimentaire et Evolution Geobiologique, Laboratoire de Prehistoire et Paleoecologie du Quaternaire de L'Ecole Pratique des Hautes Etudes, and Institut des Sciences de la Terre, Universite de Dijon, Dijon, France Joel Cracraft Department of Anatomy, University of Illinois, Chicago, Illinois and Field Museum of Natural History, Chicago, Illinois Rob DeSalle Department of Genetics, Washington University Medical School, St. Louis, Missouri Lee Ehrman Division of Natural Sciences, State University of New York at Purchase, Purchase, New York L. Val Giddings Office of Technology Assessment, Congress of the United States, Washington, D.C Kenneth Y. Kaneshiro Hawaiian Evolutionary Biology Program, University of Hawaii at Manoa, Honolulu, Hawaii Sinzo Masaki Laboratory of Entomology, Faculty of Agriculture, Hirosaki University, Hirosaki 036, Japan v

6 vi Contributors P. A. Parsons Department of Genetics and Human Variation, La Trobe University, Bundoora, Victoria, 3083, Australia Bobb Schaeffer Department of Vertebrate Paleontology, American Museum of Natural History, New York, New York Frederick H. Sheldon Department of Biology and Peabody Museum of Natural History, Yale University, New Haven, Connecticut Present Address: Tiburon Center for Environmental Studies, San Francisco State University, Tiburon, Calfornia Charles G. Sibley Department of Biology and Peabody Museum of Natural History, Yale University, New Haven, Connecticut Present address: Tiburon Center for Environmental Studies, San Francisco State University, Tiburon, California S. E. Sultan Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, Massachusetts Alan R. Templeton Department of Biology, Washington University, St. Louis, Missouri Thomas J. Walker Department of Entomology and Nematology, University of Florida, Gainesville, Florida Marvin Wasserman Biology Department, Queens College of the City University of New York, Flushing, New York 11367

7 Preface Evolutionary Biology, of which this is the twenty-first volume, continues to offer its readers a wide range of original articles, reviews, and commentaries on evolution, in the broadest sense of that term. The topics of the reviews range from anthropology and behavior to molecular biology and systematics. In recent volumes, a broad spectrum of articles have appeared on such subjects as evolution of the bacterial genome, biochemical systematics in plants, a discussion of species selection, and development and evolution of the vertebrate limb. Articles such as these, often too long for standard journals, are the material for Evolutionary Biology. The editors continue to solicit manuscripts on an international scale in an effort to see that everyone ofthe many facets of biological evolution is covered. Manuscripts should be sent to anyone of the following: Max K. Hecht, Department of Biology, Queens College of the City University of New York, Flushing, New York 11367; Bruce Wallace, Department of Biology, Virginia Polytechnic Institute and State University, Blacksburg, Virginia 24061; or Ghillian T. Prance, New York Botanical Garden, Bronx, New York The Editors vii

8 Contents 1. The Significance of Asymmetrical Sexual Isolation 1 Lee Ehrman and Marvin Wasserman Introduction... 1 The Kaneshiro Models... 6 Selection for Nonsexual Behavior Traits... 6 Evidence from the Hawaiian Drosophila... 7 Other Evidence... 9 The Watanabe-Kawanishi Models Other Models Evidence from Drosophila arizonensis-drosophila mojavensis Case Evidence from Drosophila paulistorum Discussion References Comments on "The Significance of Asymmetrical Sexual Isolation" Rob DeSalle and Alan R. Templeton References The Significance of Asymmetrical Sexual Isolation and the Formation of New Species Kenneth Y. Kaneshiro and L. Val Giddings Introduction Other Models of Asymmetrical Sexual Isolation ix

9 x Contents The Watanabe-Kawanishi Model The Wasserman-Koepfer Model Empirical Data in Support of the Kaneshiro Model Male Choice or Female Choice? Mating Propensity and Sex Drive The Direction of Evolution in Hawaiian Drosophila Mating Asymmetries and Speciation Concluding Remarks References DNA Hybridization and Avian Systematics: Introduction 45 Max K. Hecht 4. DNA Hybridization and Avian Phylogenetics 47 Joel Cracraft Introduction From Organisms to Distances: The Data The Method of DNA Hybridization The Nature of the Data DNA Distances and Sources of Error From DNA Distances to Trees: The Analytical Methods What Kinds of Trees Are These? The Uniform Average Rate (UAR) Relaxing the Assumption of a UAR Evaluating DNA Tree Topologies Discussion and Interpretation DNA Distances and Their Phylogenetic Structure The Temporal Pattern of Avian Evolution Conflicts among Data: The Search for Congruence "The Problem of Phylogeny Has Been Solved": Conclusion References... 93

10 Contents xi 5. DNA Hybridization and Avian Phylogenetics: Reply to Cracraft Charles G. Sibley, Jon E. Ahlquist, and Frederick H. Sheldon References DNA Evolutionary Implications of Phenotypic Plasticity in Plants S. E. Sultan Introduction The Nature of Phenotypic Plasticity Analysis of Phenotypic Expression The Genotypic Basis of Plasticity The Expression of Plasticity Size and Allocation Leaf Morphology... " 139 Physiological Plasticity Biosynthetic Responses... " 141 Plastic Convergence Evolutionary Responses to Environmental Heterogeneity The Heterogeneity of the Plant Environment Theoretical Considerations Regarding Specialization and Plasticity Between-Population Response Evolutionary Response within Populations What Is Fitness in Plants? Conclusions References Deuterostome Monophyly and Phylogeny Bobb Schaeffer Introduction Deuterostomes

11 xii Contents Echinoderms Hemichordates Urochordates Cephalochordates Craniates Cleavage and Cell Movement Organizers and Mesodermal Induction Somites Ectoderm and Neural Induction Neural Crest Head Segmentation Endoderm and Gut Appendages Character Analysis Analysis and Comments Resume Abbreviations References Arvicolid Data (Arvicolidae, Rodentia) and Evolutionary Concepts Jean Chaline Introduction Arvicolids Paleo species (Morpho species) and Biospecies Morphological Data The Relativity of Morphospecies Analysis of Dental Characteristics Morphological Data and Arvicolid Phylogeny Origin of the Arvicolidae Arvicolid Paleontological History Chromosomal Data Biochemical Data Arvicolid Data and Speciation Speciation As a Spaciotemporal and Populational Phenomenon Speciation and Chromosomal Changes Speciation and Chromosomal, Biochemical, and Morphological Changes..., 269

12 Contents xlii Arvicolid Data and Phyletic Gradualism Phyletic Gradualism in Water Rats Phyletic Gradualism in Lagurines Phyletic Gradualism in Muskrats Phyletic Gradualism in Ellobines Other Cases of Phyletic Gradualism Arvicolid Data and Stasis Morphological Stasis Ecophenotypic Iterative Changes How Often? Relative Importance of Allopatric Speciation, Phyletic Gradualism, and Stasis among Arvicolids Allopatric Speciation, Phyletic Gradualism, Rates, and Stasis in Eurasiatic Arvicolidae Allopatric Speciation, Phyletic Gradualism, Rates, and Stasis in North American Arvicolidae Allopatric Speciation, Phyletic Gradualism, General Rates, and Stasis in Arvicolidae... " 290 Discussion: Arvicolid Data and Evolutionary Concepts Speciation Phyletic Gradualism Stasis Relationship between Speciation, Phyletic Gradualism, and Stasis: A Reconsideration of the Biological Species Concept The Role of Environment in Speciation Speciation and Modes of Evolution Arvicolid Data and Punctuated Equilibria Conclusion Summary References Evolutionary Rates under Environmental Stress P. A. Parsons Introduction Stress and Natural Selection Phenotypic Selection Heat Shock Proteins and Acclimatization

13 xiv Contents Variation under Stress Phenotypic versus Genotypic Recombination Mutation Mobile Genetic Elements Evolutionary Rates The Underlying Genotype Morphological Change and Ecological Stress Conclusions Summary References Cricket Life Cycles Sinzo Masaki and Thomas J. Walker Life Cycle Types Homodynamic Life Cycles Heterodynamic Life Cycles Eastern and Western Tendencies Comparison of Taxonomic Groups Regulation of Life Cycles Egg Stage Water Uptake Temperature Requirements for Embryogenesis Intensity of Diapause at High Temperature Diapause Termination by Low Temperature Tolerance to Cold Incidence of Diapause Genetic Control of Egg Diapause Nymphal Stage Number of Molts Effect of Temperature on Nondiapause Growth Group Effect on Development Photoperiodic Effect on Nymph Overwinterers Photoperiodic Effect on Egg Overwinterers Adult Stage Functional Division of Adult Life Wing Dimorphism Fecundity

14 Contents xv Geographical Adaptations Homodynamic versus Heterodynamic Life Cycles Egg versus Nymph Overwintering... ; Variation in Voltinism Linear Geographical Clines Saw-Toothed Patterns of Variation Polymorphism in Life Cycles Life Cycle and Speciation Seasonal Isolation Parallel Divergence of Life Cycles Climatic Speciation Model Evolutionary Potential in Life Cycle References Index

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