The Biology of Gobies © 2012 by Taylor & Francis Group, LLC The Biology of Gobies Editors Robert A. Patzner Department Organismic Biology University of Salzburg Salzburg, Austria James L. Van Tassell Systematic Ichthyology Biology Department Hofstra University Hempstead, NY, USA Marcelo Kovačić Prirodoslovni Muzej Rijeka Lorenzov Prolaz 1 Rijeka, Croatia B.G. Kapoor Formerly Professor of Zoology The University of Jodhpur Jodhpur, India Science Publishers Jersey, British Isles Enfi eld, New Hampshire © 2012 by Taylor & Francis Group, LLC CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 © 2012 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: 20111205 International Standard Book Number-13: 978-1-4398-6233-9 (eBook - PDF) This book contains information obtained from authentic and highly regarded sources. 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Bloch, 1782-84 from Bloch’s Atlas Vol. 1, plate 107, Mergus Verlag, Melle, Germany Preface Th e Gobiiformes is one of the most diverse orders of the Percomorphs, with approximately 2000 species in 270 genera. Th ey represent 5 to 10% of all teleosts, with species occurring in marine, fresh, and brackish water environments. Both one of the smallest extant vertebrates, Trimmatom nanus, with a maximum length of 8.0 mm SL for gravid females, and the lightest weight vertebrate, Schindleria brevipinguis, with a maximum weight of about 2 mg, belong to the order Gobioidei. Some gobioids are amphidromous returning to their freshwater streams by traversing the face of 350 m waterfalls. Gobioids occupy a variety of marine habitats including open water, sandy or rocky habitats, tidepools, caves, sponges, and mudfl ats. Th ey are found at depths ranging from 0 to 1122 m (Karsten totoyensis) and in temperatures ranging from 7°C for Leucopsarion petersi to 40°C for Chlamydogobius eremius. Yet with all of this diversity and the important role they play in the ecology of regions a book about their biology has never been published. Th is book, the Biology of Gobies, contains the background information for others to join in on the study of this diverse group. Systematics within the Gobiiformes and their placement within the teleosts is not resolved. Morphological trees based on synapomorphies differ from those proposed by molecular trees, with some molecular trees not supported by any morphological synapomorphies. Th e history of morphology and molecular systematics is provided to give readers an understanding of the problems faced in resolving monophyletic clades. Th e current status of the systematics of each of the families is presented © 2012 by Taylor & Francis Group, LLC vi Preface by authors that have worked with systematics of those families for decades. Th e chapters on the study of morphological and molecular systematics, along with chapters on each of the families will help draw attention to the need for further studies in these areas. Gobies are unparalleled among vertebrates in their ability to diversify and adapt. Th ey excel in niche exploitation and microhabitat specialization which has led to rapid speciation and adaptive radiation. Th e section on zoogeography examines the genera and species of specifi c regions of the world including the Americas, north-eastern Atlantic, Mediterranean, southern Australia, New Zealand, Ponto-Caspian, and European freshwaters along with more generalized chapters on the diversity and distributions of marine and tropical freshwater gobies. Th e reproduction of gobies is unusual compared to most fi shes. Th ere is a large investment in parental care with demersal eggs guarded by the male. Th ey vary reproductively from gonochoric to hermaphrodite, monogamy to polygyny and promiscuity, some have short life spans and reproduce once only while others have longer life spans reproducing one or more times per year. Several chapters are devoted to reproductive strategies, reproductive organs, and the development of larvae and gobies. Gobies form a major component of coral reef and coastal fish communities throughout the tropics and subtropics; in coral reef habitats gobies account for over 50 % of the energy fl ow. Th e short lifespans and high levels of reproductive output result in rapid generational turnover and the production of a signifi cant component of the marine ichthyoplankton biomass. Symbiotic relationships of gobies as cleaners and in partnership with alpheid shrimp; paedomorphic species; mudskippers; and the role of gobies on coral reefs provide insight into the ecology of the group. Th irty-three authors from 15 countries have contributed to the book to provide a synopsis of our current knowledge and areas for further studies. Robert A. Patzner James L. Van Tassell Marcelo Kovačić B.G. Kapoor © 2012 by Taylor & Francis Group, LLC Contents Preface v List of Contributors xi Section 1: Systematics 1.1 A History of Gobioid Morphological Systematics 3 James L. Van Tassell, Luke Tornabene and Michael S. Taylor 1.2 Molecular Systematics of Gobioid Fishes 23 Lukas Rüber and Ainhoa Agorreta 1.3 Systematics of the Rhyacichthyidae 51 Helen K. Larson 1.4 Systematics of Odontobutidae 61 Akihisa lwata 1.5 Systematics of Butidae and Eleotridae 79 Christine Th acker 1.6 Systematics of Gobionellidae 87 Frank Pezold 1.7 Systematics of Oxudercinae 99 Edward Murdy 1.8 Systematics of Amblyopinae 107 Edward Murdy 1.9 Systematics of Sicydiinae 119 Philippe Keith and Clara Lord 1.10 Systematics of Gobiidae 129 Christine Th acker © 2012 by Taylor & Francis Group, LLC viii Contents Section 2: Zoogeography 2.1 Gobiiformes of the Americas 139 James L. Van Tassell 2.2 North-Eastern Atlantic and Mediterranean Gobies 177 Marcelo Kovačić and Robert A. Patzner 2.3 Diversity and Distribution of Marine, Euryhaline and Amphidromous Gobies from Western, Central and Southern Africa 207 Ulrich K. Schliewen 2.4 Th e Marine Temperate Gobioids of Southern Australia and the New Zealand Region 235 Helen K. Larson 2.5 Tropical Freshwater Gobies: Amphidromy as a Life Cycle 243 Philippe Keith and Clara Lord 2.6 Diversity and Distribution of Freshwater Gobies from the Mediterranean, the Black and Caspian Seas 279 Jörg Freyhof Section 3: General Biology 3.1 Gobies as Predator and Prey 291 C. Dieter Zander 3.2 Morphological Adaptations to Special Environments of Gobies 345 C. Dieter Zander 3.3 Morphological Organization and Variability of the Reproductive Apparatus in Gobies 367 Carlotta Mazzoldi, Robert A. Patzner and Maria B. Rasotto 3.4 Early Development of Gobies 403 Rita Borges, Cláudia Faria, Fátima Gil and Emanuel J. Gonçalves © 2012 by Taylor & Francis Group, LLC Contents ix Section 4: Ecology 4.1 Planktonic and Paedomorphic Gobioids 465 Mario La Mesa 4.2 Gobies on Coral Reefs 493 Juergen Herler, Philip L. Munday and Vanessa Hernaman 4.3 Gobies as Cleaners 531 Isabelle M. Côté and Marta C. Soares 4.4 Th e Partnership between Gobiid Fishes and Burrowing Alpheid Shrimps 559 Ilan Karplus and Andrew Richard Th ompson 4.5 Mudskippers: Front Runners in the Modern Invasion of Land 609 Atsushi Ishimatsu and Tomas T. Gonzales Index 639 Th e Editors 665 Color Plate Section 667 © 2012 by Taylor & Francis Group, LLC List of Contributors Agorreta Ainhoa Department of Zoology and Ecology, University of Navarra, Irunlarrea s/n, 31008 Pamplona, Spain and Department of Zoology, Th e Natural History Museum, Cromwell Road, SW7 5BD, London, UK. E-mail: [email protected] Borges Rita CCMAR, Centre of Marine Sciences, Universidade do Algarve, Campus de Gambelas 8005-139 Faro, Portugal and Eco-Ethology Research Unit, Instituto Superior de Psicologia Aplicada, Rua Jardim do Tabaco 34, 1149-041 Lisboa, Portugal. E-mail: [email protected] Côté Isabelle M. Department of Biological Sciences, Simon Fraser University, Burnaby BC, V5A 1S6, Canada. E-mail: [email protected] Faria Cláudia Instituto de Educação, Universidade de Lisboa, Campo Grande, Edifi cio C6, 1749-016, Lisboa, Portugal and Eco-Ethology Research Unit, Instituto Superior de Psicologia Aplicada, Rua Jardim do Tabaco 34, 1149-041 Lisboa, Portugal. E-mail: [email protected] Freyhof Jörg Leibniz-Institute of Freshwater Ecology and Inland Fisheries, Müggelseedamm 310, 12587 Berlin, Germany. E-mail: [email protected] © 2012 by Taylor & Francis Group, LLC xii List of Contributors Gil Fátima Aquário Vasco da Gama, Rua Direita do Dafundo, 1495-718 Cruz Quebrada- Dafundo, Portugal. E-mail: [email protected] Gonçalves Emanuel J. Eco-Ethology Research Unit, Instituto Superior de Psicologia Aplicada, Rua Jardim do Tabaco 34, 1149-041 Lisboa, Portugal. E-mail: [email protected] Gonzales Tomas T. Center for General Education, AMA International University, Bahrain, P.O. Box 18041, Manama, Kingdom of Bahrain. E-mail: [email protected] Herler Juergen Department of Th eoretical Biology, Morphology Section, Faculty of Life Sciences, University of Vienna, Althanstrasse 14, 1090 Vienna, Austria. E-mail: [email protected] Hernaman Vanessa Centre for Marine Studies, University of Queensland, St Lucia, Queensland 4072, Australia. E-mail: [email protected] Ishimatsu Atsushi Institute for East China Sea Research, Nagasaki University, Tairamachi 1551-7, Nagasaki 851-2213, Japan. E-mail: [email protected] Iwata Akihisa Laboratory of Ecology and Environment, Division of Southeast Asian Area Studies, Graduate School of Asian and African Area Studies, Kyoto University Yoshida, Shimoadachi-cho 46, Sakyo-ku, Kyoto 606-8501 Japan. E-mail: [email protected] Karplus Ilan Aquaculture Research Unit, the Institute of Animal Sciences, The Volcani Research Center, P.O. Box 6, Bet-Dagan 20250, Israel. E-mail: [email protected] © 2012 by Taylor & Francis Group, LLC