K24896 6000 Broken Sound Parkway, NW Suite 300, Boca Raton, FL 33487 711 Third Avenue New York, NY 10017 an informa business 2 Park Square, Milton Park www.crcpress.com Abingdon, Oxon OX14 4RN, UK A ScieNce PUBLiSheRS BOOK Phylogeny, Anatomy and Physiology of Ancient Fishes TThhiiss ppaaggee iinntteennttiioonnaallllyy lleefftt bbllaannkk Phylogeny, Anatomy and Physiology of Ancient Fishes Editors Giacomo Zaccone Department of Food and Environmental Science University of Messina Messina Italy K. Dabrowski School of Environment & Natural Resources Ohio State University Columbus, OH USA Michael S. Hedrick Developmental Integrative Biology Research Cluster Department of Biological Sciences University of North Texas Denton, TX USA J.M.O. Fernandes Marine Genomics Research Group Faculty of Biosciences and Aquaculture University of Nordland Bodø Norway José M. Icardo Department of Anatomy and Cell Biology Faculty of Medicine University of Cantabria Santander SPAIN p, A SCIENCE PUBLISHERS BOOK GL--Prelims with new title page.indd ii 4/25/2012 9:52:40 AM Cover illustrations reproduced by kind courtesy of one of the editors, José M. Icardo. CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 © 2016 by Taylor & Francis Group, LLC CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works Version Date: 20150721 International Standard Book Number-13: 978-1-4987-0756-5 (eBook - PDF) This book contains information obtained from authentic and highly regarded sources. Reasonable efforts have been made to publish reliable data and information, but the author and publisher cannot assume responsibility for the validity of all materials or the consequences of their use. The authors and publishers have attempted to trace the copyright holders of all material reproduced in this publication and apologize to copyright holders if permission to publish in this form has not been obtained. If any copyright material has not been acknowledged please write and let us know so we may rectify in any future reprint. Except as permitted under U.S. Copyright Law, no part of this book may be reprinted, reproduced, transmitted, or utilized in any form by any electronic, mechanical, or other means, now known or hereafter invented, including photocopying, microfilming, and recording, or in any information stor- age or retrieval system, without written permission from the publishers. For permission to photocopy or use material electronically from this work, please access www.copy- right.com (http://www.copyright.com/) or contact the Copyright Clearance Center, Inc. (CCC), 222 Rosewood Drive, Danvers, MA 01923, 978-750-8400. CCC is a not-for-profit organization that pro- vides licenses and registration for a variety of users. For organizations that have been granted a photo- copy license by the CCC, a separate system of payment has been arranged. Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identification and explanation without intent to infringe. Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the CRC Press Web site at http://www.crcpress.com Preface Primitive fi shes, including the Agnathans as the fi rst representative of the vertebrates of today, have populated the aquatic environment for about the last 500 million years. Extant primitive species are essentially similar to their fossil counterparts, retaining a number of characteristics that no longer exist in living related species (or taxa). By studying the molecular, morphological and physiological characteristics of ancient fi shes we ought to analyze relevant episodes of our own evolutionary story. This is probably what makes this kind of study so appealing both to the public imagination and to many researchers from very different fi elds. In line with previous excellent compilations, like the one presented in Volume 26 of the Fish Physiology Series, year 2007, the objective of this work is to bring together the knowledge and experience of authors who are experts in phylogenetic relationships, morphology and physiology, in order to assemble a volume rich in information and new developments. In addition to excellent phylogenetic approaches, this volume combines studies on comparative anatomy, biology, biochemistry and molecular biology. The different chapters include the heart and the circulatory system, air respiration and the control of breathing, the digestive tract and gut development, the olfactory system and several other unusual features such as the presence of slime glands. Cyclostomes, lungfi shes, chondrosteans and holosteans constitute the main subjects of study. Teleosts are also protagonists of this work since, inevitably, exceptional biological strategies and specifi c morphological and physiological features have to be confronted with related behavior or characteristics (or the lack of them) exhibited by the more derived species. In this context, evolutionary relationships among the major actinopterygian and sarcopterygian lineages have not been conclusively resolved, and a consensus phylogeny for teleost is emerging today based on analysis of various subsets of actinopterygian taxa. All the chapters are state-of-the-art works on the topics under discussion. The list of subjects dealt with had to be pruned and some other topics had to be abandoned for various reasons. However, by engaging people from different fi elds and disciplines we sought to avoid overlooking interactions between the different levels of biological organization and to create a comprehensive view of the topics and animals studied. The view that emerges is that ancient vi Phylogeny, Anatomy and Physiology of Ancient Fishes fi shes may be very complex organisms. They are often contemplated as “relics” of evolution or, as it is more commonly said, living fossils. This is probably true, but relics do not survive through geological times. Ancient fi shes had the capability to adapt to changing environments and are able to thrive in present day conditions. The objective of research programs should not solely be for the purpose of increasing our knowledge base: our goal should be enhance the conditions for the preservation of these animals for generations to come. The present volume is intended to be useful to professionals and researchers interested in the diverse topics of study, to educators, to advanced students and, in general, to all of those sharing a biological interest in aquatic creatures. The editors are indebted to all the authors who generously expressed their willingness to cooperate, were able to complete their chapters and had the patience to bear with the inevitable delays inherent on a project of this kind. Additional thanks are due to the authors involved with the revision process. The Editors Contents Preface v 1. Phylogenetic Introduction 1 Hans-Peter Schultze 2. The Occurrence and Function of the NOS/NO System in the 19 Heart of the Eel and the African Lungfi sh Daniela Amelio, Filippo Garofalo, Sandra Imbrogno and Bruno Tota 3. Evolutionary Aspects on the Comparative Biology of Lungfi shes: 38 Emphasis on South-American Lungfi sh, Lepidosiren paradoxa Vera Maria Fonseca de Almeida-Val, Luciana Mara Lopes Fé and Derek Felipe de Campos 4. Developmental Physiology of the Australian Lungfi sh, 57 Neoceratodus forsteri Casey A. Mueller 5. Aestivation in African Lungfi shes: Physiology, Biochemistry and 81 Molecular Biology Shit F. Chew, Biyun Ching, You R. Chng, Jasmine L.Y. Ong, Kum C. Hiong, Xiu L. Chen and Yuen K. Ip 6. Anatomy of the Heart and Circulation in Lungfi shes 133 José M. Icardo, Bruno Tota and Yuen K. Ip 7. The Cardiac Outfl ow Tract of Primitive Fishes 151 Adrian C. Grimes 8. Control of Breathing in Primitive Fishes 179 Michael S. Hedrick and Stephen L. Katz 9. The Lung-Swimbladder Issue: A Simple Case 201 of Homology—Or Not? Markus Lambertz and Steven F. Perry 10. The Gut and Associated Organs in the African Lungfi sh 212 Protopterus annectens José M. Icardo viii Phylogeny, Anatomy and Physiology of Ancient Fishes 11. Morphology, Histology, and Functional Structure of the 233 Alimentary Canal of Sturgeon Ramón Carmona Martos, Cristina E. Trenzado Romero and Ana Sanz Rus 12. The Structural Organization in the Olfactory System of the 260 Teleosts and Garfi shes Michal Kuciel, Krystyna Żuwała, Eugenia Rita Lauriano, Leszek Satora and Giacomo Zaccone 13. Hagfi sh Slime and Slime Glands 272 Douglas S. Fudge, Timothy M. Winegard and Julia E. Herr Index 291 1 Phylogenetic Introduction Hans-Peter Schultze The term ‘Fishes’ is used either widely including all vertebrates with gills (Craniata or Vertebrata = agnathans + gnathostomes) or restricted to jaw- bearing vertebrates with gills (gnathostomes). The gnathostomes include the tetrapods, which is often not recognized. Tetrapods are a lineage within the Osteichthyes (Hennig 1983 introduced the term ‘Osteognathostomata’ for bony fi shes + tetrapods). In this book the term ‘Fishes’ is used in the wide sense, thus the term ‘ancient fi shes’ refers to agnathans and different gnathostomes (excellent overview of extant and fossil gnathostomes in Janvier 1996) excluding advanced actinopterygians (specifi cally advanced teleosts) and sarcopterygians (specifi cally tetrapods). Craniata or Vertebrata Craniates or Vertebrates are chordates with a division of the body in the head, trunk and tail (= regionalization). The name Craniata refers to the presence of a head. The name Vertebrata indicates the presence of vertebral elements, which are missing in hagfi shes. Thus earlier hagfi shes, Myxinoida, were considered more primitive than Petromyzontida and fossil agnathans, which were classifi ed together with the gnathostomes as vertebrates. A key character of Craniata and Vertebrata is the neural crest, which induces many of their features. The head includes a pair of eyes, a pair of nasal capsules, a brain, dorsal median organs (pineal- and parietal organ), neurohypophysis and adenohypophysis, two semicircular canals in the labyrinth and external Natural History Museum University of Kansas Lawrence, Kansas, USA. E-mail: [email protected]