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Beach-Spawning Fishes Spawning Reproduction in an Endangered Ecosystem

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Beach-Spawning Fishes Spawning Reproduction in an Endangered Ecosystem Martin LIFE SCIENCE Beach- Spawning Beach- Fishes Fishes Beach-Spawning Spawning Reproduction in an Endangered Ecosystem Beach-spawning fishes from exotic locations on most continents of the world provide spectacular examples of extreme adaptations during the Fishes most vulnerable life cycle stages. The beauty, intriguing biology, and importance of these charismatic fishes at the interface between marine and terrestrial ecosystems have inspired numerous scientific studies. Adaptations of behavior, physiology, development, and ecology are Reproduction in an gathered together for the first time in this book. Beach-Spawning Fishes: Reproduction in an Endangered Endangered Ecosystem Ecosystem serves as a comprehensive guide to beach spawning, a charismatic animal behavior that is seen in a surprising number of teleost species. This unexpected form of reproduction provides a window into the ecology of coastal areas, the behaviors and physiology necessary for fishes and their eggs to adapt to terrestrial conditions, and the threats and challenges for conservation and management. Beach-spawning species include important forage fishes such as the capelin, exotic fishes such as the fugu puffer, and the spectacular midnight runs of the California grunion. K21560 Karen L. M. Martin 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 www.crcpress.com K21560_cover.indd 1 7/30/14 11:03 AM Beach- Spawning Fishes Reproduction in an Endangered Ecosystem Beach- Spawning Fishes Reproduction in an Endangered Ecosystem Karen L. M. Martin Pepperdine University Malibu, California, USA Boca Raton London New York CRC Press is an imprint of the Taylor & Francis Group, an informa business CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 © 2015 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: 20140716 International Standard Book Number-13: 978-1-4822-0800-9 (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 This book is dedicated to Doug Martin, always my favorite Pisces and the best catch of any day. Contents Preface.......................................................................................................................xi Acknowledgments .................................................................................................. xiii Introduction ..............................................................................................................xv About the Author ....................................................................................................xix Chapter 1 A Leap of Faith: The Evolution of Beach Spawning in Fishes ............1 1.1 Overview of Biogeography and Ecology of Beach-Spawning Fishes ............................................................1 1.2 Fish Families with Many Intertidal Species Also Have Many Species That Spawn on Beaches .....................................4 1.3 Some Lineages Indicate Independent Origins of Beach-Spawning Behavior ........................................................5 1.4 Some Teleost Fish Species Spawn on Beaches and Other Habitats ......................................................................................6 1.5 Lineages That Include Multiple Species of Beach- Spawning Fishes Show Multiple Independent Origins of This Behavior ........................................................................8 1.6 The Leap of Faith, Plastic Behavior, and Evolution ................ 11 References ..........................................................................................12 Chapter 2 Surf, Sand, and Beach: Physical Conditions of Shore Habitats for Fishes ............................................................................................ 17 2.1 Tidal Ebb and Flow Alter Habitat Conditions Rapidly and Predictably on Beaches ..................................................... 17 2.2 Some Fish Species Are Resident in the Intertidal Zones during All Parts of the Tidal Cycle, Others Visit Only at High Tide ................................................................................. 19 2.3 Beach Spawning for Fishes Usually Occurs during High Tides ...19 2.4 Beach-Spawning Behavior in Fishes May Not Involve Air Breathing ...........................................................................20 2.5 Beach Spawning Occurs on Specific Substrates ..................... 21 2.6 Few Species of Freshwater Fishes Spawn at the Water’s Edge ... 23 2.7 Air, Freshwater, and Seawater Have Very Different Properties as Respiratory Media .............................................24 2.8 Beach-Spawning Fishes Are Global in Distribution ...............26 References ..........................................................................................26 Chapter 3 Locals Only: Beach-Spawning Behavior in Resident Intertidal Fishes ...33 3.1 Many Teleosts Reproduce with Pelagic Eggs .......................... 33 3.2 Beach-Spawning Fishes Produce Demersal Eggs ...................34 vii viii Contents 3.3 Diverse Mating Behaviors and Mating Systems Are Seen in Beach-Spawning Fishes That Are Intertidal Residents ...... 35 3.4 Fishes That Reside in the Rocky Intertidal Zone Spawn on the Rocky Beach .................................................................37 3.5 Fishes in Estuaries Spawn on Beaches at the Water’s Edge or on Intertidal Mud Flats .............................................. 43 3.6 Summary of Beach Spawning by Resident Intertidal Fishes ..... 47 References ..........................................................................................48 Chapter 4 Vacation Sex: Subtidal Fishes That Make Spawning Migrations to the Beach ........................................................................................ 55 4.1 Some Species Migrate into the Rocky Intertidal Zone to Spawn ..................................................................................57 4.2 Some Species Migrate onto Gravel Beaches to Spawn ...........60 4.3 Some Fishes Migrate onto Sandy Beaches to Spawn ..............65 4.4 Some Fishes Spawn on Intertidal or Nearshore Vegetation .......67 4.5 Some Beach-Spawning Fishes Are Anadromous or Catadromous ............................................................................68 4.6 Some Fishes Appear to Be Transitioning toward Beach Spawning .................................................................................69 4.7 Summary for Fishes That Migrate from Other Habitats to Spawn on Beaches ................................................................... 71 References ..........................................................................................73 Chapter 5 Catching a Breath: Beach-Spawning Fishes and Air Breathing ........79 5.1 How Are Emergence and Air Breathing Beneficial for Beach-Spawning Fishes? ...................................................79 5.2 Intertidal Fishes Show Gradients of Amphibious Behavior and Air Breathing ....................................................80 5.3 Respiratory Structures for Amphibious Fishes Are Similar for Water and Air .................................................84 5.4 Respiratory Structures Undergo Modification as Conditions Change ..................................................................87 5.5 Metabolic Rate in Air during Emergence Is Similar to Metabolic Rate in Water for Many Amphibious Fishes ..........87 5.6 Terrestrial Activity Increases the Rate of Aerial Gas Exchange for Amphibious Fishes ............................................89 5.7 Many Amphibious Fishes Show Hypoxia Tolerance ...............90 5.8 Estuarine Fishes That Spawn
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