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Interdisciplinary Evolution Research 2 Emanuele Serrelli Nathalie Gontier Editors Macroevolution Explanation, Interpretation and Evidence Interdisciplinary Evolution Research Volume 2 Series editors Nathalie Gontier, Lisbon, Portugal Olga Pombo, Lisbon, Portugal About the Series The time when only biologists studied evolution has long since passed. Accepting evolution requires us to come to terms with the fact that everything that exists must be the outcome of evolutionary processes. Today, a wide variety of academic disciplines are therefore confronted with evolutionary problems, ranging from physics and medicine, to linguistics, anthropology and sociology. Solving evolutionary problems also necessitates an inter- and transdisciplinary approach, which is why the Modern Synthesis is currently extended to include drift theory, symbiogenesis, lateral gene transfer, hybridization, epigenetics and punctuated equilibria theory. The series Interdisciplinary Evolution Research aims to provide a scholarly platform for the growing demand to examine specific evolutionary problems from the perspectives of multiple disciplines. It does not adhere to one specific academic field, one specific school of thought, or one specific evolutionary theory. Rather, books in the series thematically analyze how a variety of evolutionary fields and evolutionary theories provide insights into specific, well-defined evolutionary problems of life and the socio-cultural domain. Editors-in-chief of the series are Nathalie Gontier and Olga Pombo. The Series is edited from within the Applied Evolutionary Epistemology Lab, more information on the lab is available at http://appeel.fc.ul.pt. More information about this series at http://www.springer.com/series/13109 Emanuele Serrelli · Nathalie Gontier Editors Macroevolution Explanation, Interpretation and Evidence 1 3 Editors Emanuele Serrelli Nathalie Gontier “Riccardo Massa” Department AppEEL, Centre for Philosophy of Science of Human Sciences Campo Grande University of Milano-Bicocca University of Lisbon Milan Lisbon Italy Portugal ISSN 2199-3068 ISSN 2199-3076 (electronic) Interdisciplinary Evolution Research ISBN 978-3-319-15044-4 ISBN 978-3-319-15045-1 (eBook) DOI 10.1007/978-3-319-15045-1 Library of Congress Control Number: 2014959855 Springer Cham Heidelberg New York Dordrecht London © Springer International Publishing Switzerland 2015 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made. Printed on acid-free paper Springer International Publishing AG Switzerland is part of Springer Science+Business Media (www.springer.com) Preface and Acknowledgments In the early 1980s of the twentieth century, which is now a little less than 40 years ago, and in response to the uproar that the introduction of Punctuated Equilibria theory had caused within evolutionary biology, John Maynard Smith invited pale- ontologists to join what he called, the “high table” of evolutionary theory. In the 1970s, Niles Eldredge and Stephen J. Gould provided convincing theoretical argu- ments as well as empirical data that the Modern Synthesis was incomplete because its founders had not adequately incorporated the study of life’s evolution above and beyond the differential distribution of genes across populations through time. They defined macroevolutionary research by arguing that species are real bio- logical individuals that have a clear beginning, lifespan, and ending in time, and furthermore proved that the fossil record evidences that species demonstrate long periods of stasis where no apparent morphological change occurs, which are inter- spersed by rapid speciation events that follow a branching pattern. When the fossil record does not give proof for intermediate life forms, then instead of assuming the incompleteness of the fossil record as Darwin and Neodarwinians did, they suggested to understand the gaps as data, and to postulate that no intermediates existed when none are found. Because species are understood as real biological individuals, speciation and extinction events are also understood as real phenom- ena that require scientific investigation. Researching morphological stasis, species and above species phenomena such as (mass) extinctions and speciation events, large-scale evolutionary trends, and major transitions across all domains of life; or mapping the various units, l evels, and mechanisms whereby life evolves as well as the hierarchical nature there exists between these units, levels, and mechanisms; require an epistemic incorporation of fields such as developmental biology, ecology, systematics, and b iophysics. Most of all, it requires a change in scholarly research attitudes, a willingness to transcend classic field-specific disciplinary boundaries that remain focused on reductionist, gene-centered theoretical accounts, in favor of complexity-focused, holistic epistemic stances. In recent years, the heated aura that surrounded these often polemic debates that associated with the introduction of micro- versus macroevolutionary stances v vi Preface and Acknowledgments appears to have cooled down a bit. The explosive entrance of macroevolutionary areas of research has cleared room for what we can characterize as an almost silent integration of the major claims first put forth by macroevolutionary scholars into standard evolutionary research. One can safely say that evolutionary biologists in both micro- as well as macro-oriented fields today accept that species are real bio- logical entities with an important evolutionary role to play, and research on the evolutionary and abiotic causes that underlie constraints, stasis, extinction, and speciation has never ranged so “high” at the “evolutionary table”. Indeed, these topics defined the talks of many of the lectures of the speakers that were invited to present their work at the Darwin 2009 bicentennial that was organized at the University of Chicago. One of us, Nathalie Gontier, attended that conference as an audience participant. What amazed me was that the topics introduced by macroevolutionary scholars ranged so high, while at the same time, little attention and credit was given to where these ideas stemmed from and in which context they had originated. During that same period, and with a grant from the European Marie Curie fellowships, I held a one-year visiting research position at the American Museum of Natural History in New York, under the supervision of Niles Eldredge, with the goal to perform theo- retical research on punctuated equilibria theory in particular, and how macroevolu- tionary theory in general is applicable within the sociocultural sciences. For many years, the Museum has set a crucial scene for the development of evolutionary the- ory, both in what regards its micro- as well as macroevolutionary stances, not in the least by employing scholars such as Niles Eldredge and Stephen J. Gould as well as Ernst Mayr. At that moment in time, by on the one hand reading these scholars’ important works that lie at the basis of micro- and macroevolutionary stances as well as on the other hand having the experience to attend the excellently organized Darwin commemoration where I had the chance to discuss many of these ideas with my contemporaries, I felt scholars from my generation had lost the roots of these significant ideas. From there grew the idea to edit a book on the matter, as well as to provide dissemination and outreach activities on the specificities of macro- evolutionary research outside the classic paleontological and biogeographical disciplines where they were first introduced, and to highlight how macroevolu- tionary-oriented scholars have contributed to a richer conceptualization as well as demonstration of life’s evolution. To obtain these goals, and back in Belgium, I started with writing out these ideas in the form of a grant proposal which I submitted to the John Templeton Foundation. The proposal, that had as goal to investigate the importance of both macroevolution as well as reticulate evolution for the Extended Synthesis within the field of evolutionary biology, as well as the impact these new areas of research have on the growing sociocultural evolutionary sciences, was accepted by the Foundation and was successfully executed at the Applied Evolutionary Epistemology Lab (http://appeel.fc.ul.pt), at the Portuguese University of Lisbon in 2012–2013. I hereby want to thank again the staff of the Templeton founda- tion and, in particular, Paul Wason as well as Kevin Arnold and Drew Rick-Miller, Preface and Acknowledgments vii my program officers. I am grateful to the project’s team, including Olga Pombo as well as Márcia Belchior, Francisco Carrapiço, Luís Correia, Larissa Mendoza Straffon, Marco Pina, and Emanuele Serrelli; and I want to express my gratitude toward the Portuguese Gulbenkian Foundation and Ciência Viva Agency for their enthusiasm and efforts in helping me to bring my ideas on outreach and knowl- edge dissemination into action during what turned out to be a crazily busy year. Together, we organized a session on how macroevolutionary theory transcends the Modern Synthesis for the 2013-meeting for the American Association for the Advancement of Science (http://appeel.fc.ul.pt/sub/eve/dir/aaas/aaas2013.html) that among the speakers had Alycia Stigall who also contributed to this volume; a session for the 2012-meeting of the American Anthropological Association that partially focused on cultural macroevolutionary theories and methodologies (http://appeel.fc.ul.pt/sub/eve/dir/aaa/aaa2012.html); two international evolu- tion schools for pre- and postdoctoral university students with modules on mac- roevolution (http://evolutionschool.fc.ul.pt) taught, among others, by Ilya Tëmkin and Folmer Bokma who also contributed to this volume as well as the confer- ence on Micro and Macro, Horizontal and Vertical Evolutionary Patterns (http:// evolutionarypatterns.fc.ul.pt); a public conference on evolution with special ses- sions for teenagers (http://evolutionconference.fc.ul.pt); we conducted several video interviews with the scholars who participated in the activities (https:// www.youtube.com/user/appeellisboa); and we concretized my multiple book ideas further, including this one on macroevolution. I am grateful to Emanuele for having accepted my invitation to co-edit the book. We are very much indebted and grateful to the numerous scholars we encoun- tered along our paths, who gave us advice and guidance, encouragement, and cri- tique. In particular, we want to thank Wallace Arthur, Quentin Atkinson, Michael Benton, Luis Borda de Agua, Rituparna Bose, Dan Brooks, Mike Bruford, Jorge Carneiro, Maurizio Casiraghi, Lounes Chikhi, Alex de Voogt, Charbel Niño El-Hani, Claus Emmeche, Douglas H. Erwin, Frietson Galis, Root Gorelick, Ronald Jenner, Fiona Jordan, Bruce Lieberman, Paulo Madruga, Octávio Mateus, Daniel McShea, Richard P. Meisel, Alex Mesoudi, Sérgio Moço, Ana Noronha, Samir Okasha, Mark Pagel, Octávio Paulo, Telmo Pievani, Anna Marie Prentiss, Luís Paulo N. Rebelo, Luis Mateus Rocha, Michael Ruse, Ana Sanchez, David Sepkoski, Artur Santos Silva, Rui Silveira, Andrej Spiridonov, Ian Tattersall, Inês Tomé, Derek Turner, Rosalia Vargas, Davide Vecchi, Emílio Rui Vilar, Tyler Volk, Matthew Joseph Walsh, Richard Watson, and Jon Umerez. We also want to cordially thank the whole Springer team, and in particular Anette Lindqvist and Sabine Schwarz. This book on macroevolution is neither a handbook for beginning scholars for which there exist numerous excellent works written by paleontologists, nor a work aimed toward philosophers and historians of science to exclusively highlight the history and epistemology of macroevolutionary ideas. This book highlights some of the most important research topics that macroevolutionary scholars have intro- duced into the evolutionary sciences, including debates on how microevolution differs from macroevolution; what the nature is of evolutionary stasis, extinction, viii Preface and Acknowledgments speciation; how we can define and measure evolutionary rates; how we can model biological hierarchies; how biophysics, ecology, evo-devo, genetics, and systemat- ics shed new light on life’s major patterns, trends and transitions, its origination, extinction and diversity; how macroevolutionary theory transcends biology and is applied within the sociocultural sciences; and how all the latter requires us to reconceptualize the very nature of evolutionary research in light of an extended synthesis. We are enormously grateful toward the authors who found the time to contrib- ute a chapter for the book, as well as the scholars who, behind the scenes, were willing to peer-review the chapters. We hope the reader will find as much enlight- enment on the subject of macroevolution as we did editing the volume. Nathalie Gontier Emanuele Serrelli Contents Macroevolutionary Issues and Approaches in Evolutionary Biology .... 1 Emanuele Serrelli and Nathalie Gontier Part I Macroevolutionary Explanations and Interpretations Can Modern Evolutionary Theory Explain Macroevolution? .......... 29 Douglas J. Futuyma Evolution as a Largely Autonomous Process ........................ 87 Folmer Bokma Visualizing Macroevolution: From Adaptive Landscapes to Compositions of Multiple Spaces ............................... 113 Emanuele Serrelli Toward a Natural Philosophy of Macroevolution .................... 163 Stanley N. Salthe Networks and Hierarchies: Approaching Complexity in Evolutionary Theory ......................................... 183 Ilya Tëmkin and Niles Eldredge Uniting Micro- with Macroevolution into an Extended Synthesis: Reintegrating Life’s Natural History into Evolution Studies ........... 227 Nathalie Gontier ix

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This book is divided in two parts, the first of which shows how, beyond paleontology and systematics, macroevolutionary theories apply key insights from ecology and biogeography, developmental biology, biophysics, molecular phylogenetics and even the sociocultural sciences to explain evolution in de
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