The Evolution of Phylogenetic Systematics 99778800552200227766558811__PPRRIINNTT..iinndddd ii 1122//0099//1133 55::3388 PPMM SPECIES AND SYSTEMATICS www.ucpress.edu/go/spsy The Species and Systematics series will investigate fundamental and practical aspects of systematics and taxonomy in a series of comprehensive volumes aimed at students and researchers in systematic biology and in the history and philosophy of biology. The book series will examine the role of descriptive taxonomy, its fusion with cyber- infrastructure, its future within biodiversity studies, and its importance as an empirical science. The philosophical consequences of classifi cation, as well as its history, will be among the themes explored by this series, including systematic methods, empirical studies of taxonomic groups, the history of homology, and its signifi cance in molecular systematics. Editor in Chief: Malte C. Ebach (University of New South Wales, Australia) Editorial Board Sandra Carlson (University of California, Davis, USA) Marcelo R. de Carvalho (University of São Paulo, Brazil) Darren Curnoe (University of New South Wales, Australia) Christina Flann (Netherlands Centre for Biodiversity Naturalis, The Netherlands) Anthony C. Gill (University of Sydney, Australia) Lynne R. Parenti (Smithsonian Institution, USA) Olivier Rieppel (The Field Museum, Chicago, USA) John S. Wilkins (University of Sydney, Australia) Kipling Will (University of California, Berkeley, USA) David M. Williams (The Natural History Museum, London, UK) René Zaragüeta i Bagils (University of Paris 6, France) 99778800552200227766558811__PPRRIINNTT..iinndddd iiii 1122//0099//1133 55::3388 PPMM The Evolution of Phylogenetic Systematics EDITED BY Andrew Hamilton UNIVERSITY OF CALIFORNIA PRESS Berkeley • Los Angeles • London 99778800552200227766558811__PPRRIINNTT..iinndddd iiiiii 1122//0099//1133 55::3388 PPMM University of California Press, one of the most distin- guished university presses in the United States, enriches lives around the world by advancing scholarship in the humanities, social sciences, and natural sciences. Its activities are supported by the UC Press Foundation and by philanthropic contributions from individuals and institutions. For more information, visit www.ucpress.edu. Species and Systematics, Vol. 5 For online version, see www.ucpress.edu. University of California Press Berkeley and Los Angeles, California University of California Press, Ltd. London, England © 2014 by The Regents of the University of California Library of Congress Cataloging-in-Publication Data The evolution of phylogenetic systematics / edited by Andrew Hamilton. pages cm — (Species and systematics ; v. 5) Includes bibliographical references and index. isbn 978-0-520-27658-1 (cloth : alk. paper) — isbn 978-0-520-95675-9 (ebook) 1. Cladistic analysis. 2. Biology—Classifi cation— Philosophy. I. Hamilton, Andrew, 1972- QH83.E96 2014 578.01′2—dc23 2013021973 Manufactured in the United States of America 23 22 21 20 19 18 17 16 15 14 10 9 8 7 6 5 4 3 2 1 The paper used in this publication meets the minimum requirements of ansi/niso z39.48–1992 (r 2002) (Permanence of Paper). Cover illustration: Chart of Evolution (1937) by Borgny Bay, Paleontological Museum, University of Oslo. Photo by Colin Purrington, courtesy of the Gould Library, Carleton College. 99778800552200227766558811__PPRRIINNTT..iinndddd iivv 1122//0099//1133 55::3388 PPMM Contents List of Contributors vii Introduction Andrew Hamilton 1 PART ONE. HISTORICAL FOUNDATIONS 15 1 Refl ections on the History of Systematics Robert E. Kohler 17 2 Willi Hennig’s Part in the History of Systematics Michael Schmitt 47 3 Homology as a Bridge between Evolutionary Morphology, Developmental Evolution, and Phylogenetic Systematics Manfred D. Laubichler 63 PART TWO. CONCEPTUAL FOUNDATIONS 87 4 Historical and Conceptual Perspectives on Modern Systematics: Groups, Ranks, and the Phylogenetic Turn Andrew Hamilton 89 99778800552200227766558811__PPRRIINNTT..iinndddd vv 1122//0099//1133 55::3388 PPMM vi | Contents 5 The Early Cladogenesis of Cladistics Olivier Rieppel 117 6 Cladistics at an Earlier Time Gareth Nelson 139 7 Patterson’s Curse, Molecular Homology, and the Data Matrix David M. Williams and Malte C. Ebach 151 8 History and Theory in the Development of Phylogenetics in Botany Brent D. Mishler 189 PART THREE. TECHNOLOGY, CONCEPTS, AND PRACTICE 211 9 Well-Structured Biology: Numerical Taxonomy’s Epistemic Vision for Systematics Beckett Sterner 213 10 A Comparison of Alternative Form-Characterization: Approaches to the Automated Identifi cation of Biological Species Norman MacLeod 245 11 The New Systematics, the New Taxonomy, and the Future of Biodiversity Studies Quentin Wheeler and Andrew Hamilton 287 Index 303 99778800552200227766558811__PPRRIINNTT..iinndddd vvii 1122//0099//1133 55::3388 PPMM Contributors Malte C. Ebach University of New South Wales [email protected] Andrew Hamilton University of Houston [email protected] Robert E. Kohler University of Pennsylvania [email protected] Manfred D. Laubichler Arizona State University [email protected] Norman MacLeod Natural History Museum, London University College London [email protected] Brent D. Mishler University of California, Berkeley [email protected] vii 99778800552200227766558811__PPRRIINNTT..iinndddd vviiii 1122//0099//1133 55::3388 PPMM viii | Contributors Gareth Nelson University of Melbourne [email protected] Olivier Rieppel The Field Museum orieppel@fi eldmuseum.org Michael Schmitt Ernst-Moritz-Arndt-Universitaet [email protected] Beckett Sterner University of Chicago [email protected] Quentin Wheeler Arizona State University [email protected] David Williams Natural History Museum, London [email protected] 99778800552200227766558811__PPRRIINNTT..iinndddd vviiiiii 1122//0099//1133 55::3388 PPMM Introduction ANDREW HAMILTON TRENDS AND TRADITIONS IN PHYLOGENETIC SYSTEMATICS In spring 2006 I met with the entomologist and systematic theorist Quentin Wheeler to discuss topics of common interest: species concepts, methodology in systematics, the recent history of phylogenetic classifi - cation, and biodiversity from a systematics perspective. The conversa- tion turned out to have a surprising twist. Quentin was founding a new research center at Arizona State University—the International Institute for Species Exploration (IISE)—that would have a focus on the history and philosophy of systematics. I jumped at the chance to direct this part of the Institute’s program because I saw that there was plenty of work to be done that would best be accomplished through the collaborative efforts of systematists, historians, and philosophers. With help from Manfred Laubichler, I set about recruiting scholars from several disciplines to participate in a workshop to help defi ne and articulate the agenda for the history and philosophy of systematics at the IISE as well as in my own work. Topics of discussion in the work- shop ranged from instrumentalism in contemporary systematics to the relationships between homology and monophyly, Willi Hennig’s theo- retical commitments, and on to how the history of recent and highly contested biology should be most productively pursued. No attempt has been made to reproduce the breadth of the workshop discussions here. Rather, I have narrowed the scope considerably by 1 99778800552200227766558811__PPRRIINNTT..iinndddd 11 1122//0099//1133 55::3388 PPMM