THE BIOLOGY OF PLETHODONTID SALAMANDERS

Transcription

1 THE BIOLOGY OF PLETHODONTID SALAMANDERS

2 THE BIOLOGY OF PLETHODONTID SALAMANDERS Edited by Richard C. Bruce Western Carolina University Cullowhee, North Carolina Robert G. Jaeger University of Southwestern Louisiana Lafayette, Louisiana and Lynne D. Houck Oregon State University Corvallis, Oregon Springer Science+Business Media, LLC

3 Library of Congress Cataloging-in-Publication Data The biology of plethodontid salamanders/edited by Richard C. Bruce, Robert G. Jaeger, and Lynne D. Houck p. cm. Includes bibliographical references. ISBN ISBN (ebook) DOI / Plethodontidae Congresses. I. Bruce, Richard C, II. Jaeger, Robert G. III. Houck, Lynne D. IV. Conference on the Biology of Plethodontid Salamanders (4th: 1998: Highlands, N.C.) QL668.C274 B '5 dc ISBN Proceedings of the Fourth Highlands Conference on the Biology of Plethodontid Salamanders, held June 12-14, 1998, in Highlands, North Carolina 2000 Springer Science+Business Media New York Originally published by Kluwer Academic / Plenum Publishers, New York in 2000 Softcover reprint of the hardcover 1st edition All rights reserved A C.I.P. record for this book is available from the Library of Congress 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

4 Professor Richard Highton. Photograph taken by Stevan J. Arnold during the Fourth Highlands Conference on the Biology of Plethodontid Salamanders.

5 FOREWORD The fourth Conference on the Biology of Plethodontid Salamanders was held at "The Mountain," in Highlands, North Carolina on June 12-14, Hosted by the Highlands Biological Station, and sponsored by the Highlands Biological Foundation, Inc., the conference afforded a state-of-the-art overview of these animals, as evidenced by the contents of the present volume and the credentials of the contributors. In the following pages, the reader will find the best current understanding of many aspects of plethodontid salamander evolution, systematics, development, morphology, life history, ecology, and field methodology. While the contents of this book consist of chapters developed from selected conference papers, their excellence is representative of the high overall quality of the conference presentations. The Highlands Biological Station is located on the Highlands Plateau in the southern Blue Ridge Mountains. As these mountains are a present center of plethodontid diversity, and are considered by some to be the center of origin of the group, the Highlands Biological Station has historically been important in the study of these animals. A list of visitors to the Station for the last 70 years would include a "who's who" of twentieth century North American herpetologists. The location and amenities of the Highlands Biological Station are unique. Within the city limits of Highlands at an elevation of nearly 1200 meters, the Station includes modern research laboratories, administrative offices, library facilities, as well as dormitory and living facilities. A nature center, nature trails, and Lake Ravenel offer diversions for researchers and Station visitors alike. Sites for studies of plethodontids are abundant and readily accessible. The town is virtually surrounded by the Nantahala National Forest. The Pisgah National Forest, Blue Ridge Parkway, and Great Smoky Mountains National Park are all within an hour and a half drive. The Highlands Biological Station is now an interinstitutional center of The University of North Carolina, administered by Western Carolina University. Originally, it was founded as a private institution in 1927 and remained so until The Highlands Biological Foundation was created to fund the activities of the Station, and remains an integral and vital partner in the advancement of scientific knowledge at the Station. Annually, the Foundation provides monies for research scholarships and grants-in-aid to students and others working at the Biological Station. Many graduate students investigating plethodontids have been supported in this way. Occasionally, it has been the privilege of the Foundation to sponsor special events, not the least of which has been the series of four symposia on plethodontid salamanders held at the Station. The first conference was held in 1972, with subsequent vii

6 viii Foreword meetings in 1982,1992, and It is with some understanding of advancing age that I admit to attending all but the first of these conferences. Each meeting was characterized by exchange of information and ideas, as well as the formation of new friendships and the renewal of old ones. Events such as these are the milestones by which we measure our academic lives. As I look at a series of group photos from these symposia, I see hair graying (or disappearing outright), and waistlines expanding (mine anyhow). But, fortunately, there are many new faces too, and each group is significantly larger. Such observations signify the importance of these conferences. The research of an everincreasing group of investigators over the years has and will continue to produce a vital and growing understanding of plethodontid salamanders. It has been my pleasure to attend these conferences and it has been the privilege of the Highlands Biological Foundation to sponsor them. It is the Foundation's hope that symposia will occur regularly in the future, and it is our wish to continue to support them. Production of a volume such as the one you are about to read requires not only financial support, but also the time and effort of a great many people. Thirtynine authors of 26 papers were involved, and the Foundation is appreciative of their efforts. In particular, we would like to recognize the hard work of editors Richard C. Bruce, Robert G. Jaeger, and Lynne D. Houck. Kluwer Academic/Plenum Publishers is to be thanked for its agreement to publish the volume. With continued research, cooperation, and communication, the process will be repeated several years from now. Andrew N. Ash, President The Highlands Biological Foundation, Inc.

7 PREFACE Lungless salamanders of the family Plethodontidae are not only fascinating animals in and of themselves, but have served as model organisms in a broad range of pioneering studies in systematics, ecology, behavior, genetics, and many other areas of evolutionary and comparative biology. The southern Appalachian Mountains represent a center of diversification of plethodontids-three of the four major subdivisions of the family occur here, species richness is high, and populations are often very dense. For these reasons, the area has long attracted specialists interested in plethodontid salamanders. From its very beginning in 1927, the Highlands Biological Station has played a major role in research efforts directed at these amphibians. Edwin Reinke, Claude Chadwick, Clifford Pope, Nelson Hairston, Sr., and Robert Gordon conducted research on salamanders at the Station in its early years, establishing a tradition that has been extended by legions of later herpetologists. Thus, it was only natural that in 1972 the Station initiated a series of conferences to promote dissemination of current research on plethodontids. The present volume represents a selection of papers deriving from the fourth plethodontid conference, held in June Although papers from the earlier conferences have appeared in a variety of formats, this is the first time that a formal conference volume has been published. It seemed appropriate to use the venue of the fourth conference to honor the research of our esteemed colleague, Richard Highton, on the occasion of his formal retirement from The University of Maryland. His ground-breaking research on molecular systematics and speciation in plethodontids dates from the late 1960s, when allozymes were first being used to address questions in these fields. TWo of us can remember that Dr. Highton's presentation at the very first plethodontid conference generated a lively discussion on the usefulness of allozymes in systematics. Accordingly, the long and productive career of Richard Highton was celebrated by beginning the fourth conference with a symposium in his honor. The symposium featured papers presented by specialists on amphibian systematics, and these papers constitute the first section of this volume. Other than the papers of the Highton Symposium, the organizers of the conference made no attempt to solicit contributions on specific topics in plethodontid biology. Thus, the remaining chapters represent a selection of the contributed papers and posters presented at the conference. We have grouped these contributions into two sections: the first a series of papers on a wide range of topics in evolutionary and comparative biology of plethodontids, the second a set of chapters more narrowly ix

8 x Preface focused on what we have called the behavioral ecology of small Plethodon (mainly P. cinereus). Whereas the former reflects the diversity of current research on plethodontid salamanders, the latter illustrates the usefulness of the smaller species of Plethodon as model organisms for investigation of a wide range of questions in behavioral ecology. We recognize that the coverage is not comprehensive; several areas of research are underrepresented or missing altogether. This is a consequence of the relatively unstructured nature of the conference itself. We trust, however, that the volume conveys to readers the current status of research on plethodontids, and will stimulate interest by researchers in furthering investigation of the many unresolved questions pertaining to the biology of these remarkable animals. Numerous individuals, too many to list individually, helped in the organization of the conference and the production of this volume, and we extend our appreciation to all of them. We do, however, wish to acknowledge the reviewers of chapters, whose cogent reviews have contributed immeasurably to the scientific quality of the volume. They are: Carl Anthony, Andrew Ash, Christopher Beachy, Paul Chippindale, Paul Cupp, Don Forester, Cary Guffey, James Hanken, Reid Harris, Todd Jackman, Elizabeth Jockusch, Allan Larson, Michael LeMaster, Dale Madison, Sharyn Marks, Jeremy Marshall, Alicia Mathis, Travis Ryan, David Sever, David Snyder, Judy Stamps, Stephen Tilley, Paul Verrell, and John Wiens. We are especially grateful to editors Adam Cohen and Mary Ann McCarra of Kluwer Academic/Plenum Publishers for their encouragement, support, and patience throughout. Finally, in commending the volume to our readers, we remind them that another plethodontid conference looms on the horizon, in the not-too-distant future, in a pleasing setting somewhere in the misty forests of the southern highlands. Richard C. Bruce, Western Carolina University Robert G. Jaeger, University of Southwestern Louisiana Lynne D. Houck, Oregon State University

9 CONTENTS Part I: Symposium in Honor of Professor Richard Highton 1. Points of View on Defining and Naming Species of Plethodontid Salamanders: A Symposium in Honor of Professor Richard Highton... 3 Robert G. Jaeger 2. Species, Descriptive Efficiency, and Progress in Systematics Darrel Frost 3. Geographic Protein Variation and Speciation in Salamanders of the Plethodon jordani and Plethodon glutinosus Complexes in the Southern Appalachian Mountains with the Description of Four New Species Richard Highton and Robert B. Peabody 4. Detecting Species Borders Using Diverse Data Sets: Examples from Plethodontid Salamanders in California David B. Wake and Elizabeth L. Jockusch 5. The Systematics of Desmognathus imitator Stephen G. Tilley 6. Species Boundaries and Species Diversity in the Central Texas Hemidactyliine Plethodontid Salamanders, Genus Eurycea Paul T. Chippindale 7. Systematics at the Tum of a Century Stevan 1. Arnold Part ll: Evolutionary and Comparative Biology of Plethodontids 8. Ethological Isolation and Variation in Allozymes and Dorsolateral Pattern between Parapatric Forms in the Desmognathus ochrophaeus Complex Louise S. Mead and Stephen G. Tilley xi

10 xii Contents 9. Phylogenetic Relationships within the Lowland Tropical Salamanders of the Bolitoglossa mexicana Complex (Amphibia: Plethodontidae) Mario Garcia-Paris, Gabriela Parra-Olea, and David B. Wake 10. Detecting Cryptic Species Using Allozyme Data Richard Highton 11. Sexual Size Dimorphism in the Plethodontidae Richard C. Bruce 12. Skull Development in Two Plethodontid Salamanders (Genus Desmognathus) with Different Life Histories Sharyn B. Marks Egg Recognition Cues and Maternal Behavior in Desmognathus ocoee Brian S. Masters and Don C. Forester 14. Southeastern U.S. Coastal Plain Habitats of the Plethodontidae: The Importance of Relief, Ravines, and Seepage 287 D. Bruce Means 15. Life History Evolution and Adaptive Radiation of Hemidactyliine Salamanders Travis J. Ryan and Richard C. Bruce 16. Population Cytogenetics of the Plethodontid Salamander Eurycea wilderae Stanley K. Sessions and Jodi L. Wiktorowski 17. Sperm Storage in Female Plethodontids with Especial Reference to the Desmognathinae David M. Sever 18. The Courtship of Plethodontid Salamanders: Form, Function, and Phylogeny Paul Verrell and Michelle Mabry Part III: Behavioral Ecology of Small Plethodon 19. Divergence of Trophic Morphology and Resource Use among Populations of Plethodon cinereus and P. hoffmani in Pennsylvania: A Possible Case of Character Displacement Dean C. Adams 20. Environmental Variation and Territorial Behavior in a Terrestrial Salamander John C. Maerz and Dale M. Madison

11 Contents xiii 21. Contributions to the Life History of the Redback Salamander, Plethodon cinereus: Preliminary Evidence for Kin Recognition Don C. Forester and Courtney L. Anders 22. Alternative Life Styles in a Terrestrial Salamander: Do Females Preferentially Associate with Each Other? Megan G. Peterson, Jennifer R. Gillette, Raelyn Franks, and Robert G. Jaeger 23. Pheromonal Attractions to Particular Males by Female Redback Salamanders (Plethodon cinereus) Jennifer R. Gillette, Sara E. Kolb, Joshua A. Smith, and Robert G. Jaeger 24. A Model of Alternative Mating Strategies in the Redback Salamander, Plethodon cinereus Robert G. Jaeger, Megan G. Peterson, and Jennifer R. Gillette 25. Cutaneous Microbial Flora and Antibiosis in Plethodon ventralis: Inferences for Parental Care in the Plethodontidae Richard M. Austin, Jr. 26. Monitoring Plethodon cinereus Populations: Field Tests of Experimental Coverboard Designs Margaret A. Carfioli, Harry M. TIebout III, Susan A. Pagano, Kristina M. Heister, and Fredric C. Lutcher Index 477