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Andreas Wanninger Editor Evolutionary Developmental Biology of Invertebrates Vol. 2 Lophotrochozoa (Spiralia) Evolutionary Developmental Biology of Invertebrates 2 Andreas Wanninger Editor Evolutionary Developmental Biology of Invertebrates 2 Lophotrochozoa (Spiralia) Editor Andreas Wanninger Department of Integrative Zoology University of Vienna Faculty of Life Sciences Wien Austria ISBN 978-3-7091-1870-2 ISBN 978-3-7091-1871-9 (eBook) DOI 10.1007/978-3-7091-1871-9 Library of Congress Control Number: 2015947925 Springer Wien Heidelberg New York Dordrecht London © Springer-Verlag Wien 2015 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifi cally the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfi lms 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 specifi c 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. Cover illustration: Scanning electron micrograph of an early trochophore larva of the tusk shell, Antalis entalis, a scaphopod mollusk. See Chapter 7 for details Printed on acid-free paper Springer-Verlag GmbH Wien is part of Springer Science+Business Media (www.springer.com) Pref ace The evolution of life on Earth has fascinated mankind for many centuries. Accordingly, research into reconstructing the mechanisms that have led to the vast morphological diversity of extant and fossil organisms and their evolution from a common ancestor has a long and vivid history. Thereby, the era spanning the nineteenth and early twentieth century marked a particularly groundbreaking period for evolutionary biology, when leading naturalists and embryologists of the time such as Karl Ernst von Baer (1792–1876), Charles Darwin (1809–1882), Ernst Haeckel (1834–1919), and Berthold Hatschek (1854–1941) realized that comparing ontogenetic processes between species offers a unique window into their evolutionary history. This revelation lay the foundation for a research fi eld today commonly known as Evolutionary Developmental Biology, or, briefl y, EvoDevo. While for many of today’s EvoDevo scientists the principle motivation for studying animal development is still in reconstructing evolutionary scenarios, the analytical means of data generation have radically changed over the centuries. The past two decades in particular have seen dramatic innovations with the routine establishment of powerful research techniques using micro- morphological and molecular tools, thus enabling investigation of animal development on a broad, comparative level. At the same time, methods were developed to specifi cally assess gene function using reverse genetics, and at least some of these techniques are likely to be established for a growing number of so-called emerging model systems in the not too distant future. With this pool of diverse methods at hand, the amount of comparative data on invertebrate development has skyrocketed in the past years, making it increas- ingly diffi cult for the individual scientist to keep track of what is known and what remains unknown for the various animal groups, thereby also impeding teaching of state-of-the-art Evolutionary Developmental Biology. Thus, it appears that the time is right to summarize our knowledge on invertebrate development, both from the classical literature and from ongoing scientifi c work, in a treatise devoted to EvoDevo. Evolutionary Developmental Biology of Invertebrates aims at providing an overview as broad as possible. The authors, all renowned experts in the fi eld, have put particular effort into presenting the current state of knowledge as comprehensively as possible, carefully weighing conciseness against level of detail. For issues not covered in depth here, the reader may consult additional textbooks, review articles, or web-based resources, v vi Preface particularly on well- established model systems such as Caenorhabditis elegans ( www.wormbase.org ) or Drosophila melanogaster ( www.fl ybase.org ) . Evolutionary Developmental Biology of Invertebrates is designed such that each chapter can stand alone, and most chapters are dedicated to one phylum or phylum-like taxonomic unit. The main exceptions are the hexa- pods and the crustaceans. Due to the vast amount of data available, these groups are treated in their own volume each (Volume 4 and Volume 5, respec- tively), which differ in their conceptual setups from the other four volumes. In addition to the taxon-based parts, chapters on embryos in the fossil record, homology in the age of genomics, and the relevance of EvoDevo for reconstructing evolutionary and phylogenetic scenarios are included in Volume 1 in order to provide the reader with broader perspectives of modern- day EvoDevo. A chapter showcasing developmental mechanisms during regeneration is part of Volume 2 . Evolutionary Developmental Biology of Invertebrates aims at scientists that are interested in a broad comparative view of what is known in the fi eld but is also directed toward the advanced student with a particular interest in EvoDevo research. While it may not come in classical textbook style, it is my hope that this work, or parts of it, fi nds its way into the classrooms where Evolutionary Developmental Biology is taught today. Bullet points at the end of each chapter highlight open scientifi c questions and may help to inspire future research into various areas of Comparative Evolutionary Developmental Biology . I am deeply grateful to all the contributing authors that made Evolutionary Developmental Biology of Invertebrates possible by sharing their knowledge on animal ontogeny and its underlying mechanisms. I warmly thank Marion Hüffel for invaluable editorial assistance from the earliest stages of this project until its publication and Brigitte Baldrian for the chapter vignette artwork. The publisher, Springer, is thanked for allowing a maximum of freedom during planning and implementation of this project and the University of Vienna for providing me with a scientifi c home to pursue my work on small, little- known creatures. This volume covers the animals that have a ciliated larva in their life cycle (often united as Lophotrochozoa), as well as the Gnathifera and the Gastrotricha. The interrelationships of these taxa are poorly resolved and a broadly accepted, clade-defi ning autapomorphy is lacking. Spiral cleavage is sometimes assumed as the ancestral mode of cleavage of this grouping, and therefore the clade is named Spiralia by some authors, although others prefer to extend the term Lophotrochozoa to this entire assemblage. Aside from the taxon-based chapters, this volume contains a chapter that highlights similari- ties and differences in the processes that underlie regeneration and ontogeny, using the Platyhelminthes as a case study. Tulbingerkogel, Austria Andreas Wanninger January 2015 Contents 1 Gnathifera. 1 Andreas Hejnol 2 Gastrotricha . 13 Andreas Hejnol 3 Platyhelminthes . 21 Teresa Adell, José M. Martín-Durán, Emili Saló, and Francesc Cebrià 4 Regeneration and Growth as Modes of Adult Development: The Platyhelminthes as a Case Study . 41 Francesc Cebrià, Emili Saló, and Teresa Adell 5 Cycliophora . 79 Andreas Wanninger and Ricardo Neves 6 Entoprocta . 89 Andreas Wanninger 7 Mollusca . 103 Andreas Wanninger and Tim Wollesen 8 Nemertea. 155 Jörn von Döhren 9 Annelida . 193 Christoph Bleidorn, Conrad Helm, Anne Weigert, and Maria Teresa Aguado 10 Phoronida . 231 Scott Santagata 11 Ectoprocta. 247 Scott Santagata 12 Brachiopoda . 263 Scott Santagata Index . 279 vii Gnathifera 1 Andreas Hejnol Chapter vignette artwork by Brigitte Baldrian. © Brigitte Baldrian and Andreas Wanninger. A. Hejnol Sars International Centre for Marine Molecular Biology , University of Bergen , Thormøhlensgate 55 , Bergen 5008 , Norway e-mail: [email protected] A. Wanninger (ed.), Evolutionary Developmental Biology of Invertebrates 2: Lophotrochozoa (Spiralia) 1 DOI 10.1007/978-3-7091-1871-9_1, © Springer-Verlag Wien 2015 2 A. Hejnol INTRODUCTION internal relationships using molecular data (Witek et al. 2008 ; Wey-Fabrizius et al. 2014 ). Figure 1.1 illustrates the likely phylogenetic The taxon Gnathifera was erected based on relationships as a consensus phylogeny that is morphological data by Ahlrichs ( 1995 , 1997 ). based on recent molecular as well as morpho- The taxon comprises the Gnathostomulida logical data. In all gnathiferans, fertilization is and Syndermata (which unites Rotifera, internal and the development direct. The para- Acanthocephala, Seisonida) (Fig. 1.1 ). With the sitic acanthocephalans have evolved additional discovery of Limnognathia maerski (Kristensen dispersal stages that allow infection and transi- and Funch 2000 ), the taxon Micrognathozoa has tions between hosts. been

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