Invertebrate Medicine E-Book

• Herausgeber: John Wiley & Sons
• Kategorie: Fachliteratur
• Sprache: Englisch

Seitenzahl: 1557

Veröffentlichungsjahr: 2010

Beschreibung

Invertebrate Medicine, Second Edition offers a thorough update to the most comprehensive book on invertebrate husbandry and veterinary care. Including pertinent biological data for invertebrate species, the book's emphasis is on providing state-of-the-art information on medicine and the clinical condition. Invertebrate Medicine, Second Edition is an invaluable guide to the medical care of both captive and wild invertebrate animals. Coverage includes sponges, jellyfish, anemones, corals, mollusks, starfish, sea urchins, crabs, crayfish, lobsters, shrimp, hermit crabs, spiders, scorpions, and many more, with chapters organized by taxonomy. New chapters provide information on reef systems, honeybees, butterfly houses, conservation, welfare, and sources of invertebrates and supplies. Invertebrate Medicine, Second Edition is an essential resource for veterinarians in zoo animal, exotic animal and laboratory animal medicine; public and private aquarists; and aquaculturists.

Leseprobe

Table of Contents

Cover

Title page

Copyright page

DEDICATION

CONTRIBUTORS

FOREWORD

PREFACE

ACKNOWLEDGMENTS

Chapter 1 INTRODUCTION

Chapter 2 SPONGES

Natural History and Taxonomy

Anatomy and Physiology

Environmental Disorders

Infectious Diseases

Analgesia, Anesthesia, and Surgery

Chapter 3 COELENTERATES

Introduction

Natural History and Taxonomy

Anatomy and Physiology

Environmental Diseases and Preventive Medicine

Infectious Diseases

Analgesia, Anesthesia, and Surgery

Diagnostic Techniques

Treatment Protocols

Transport and Handling

Chapter 4 CORAL REEF AQUARIUM HUSBANDRY AND HEALTH

History and Introduction

Factors Affecting Coral Reef Aquarium Health

Risks Associated with Reef Aquaria

Infections of Reef Systems

Stress and Other Indicators for Corals

Chapter 5 TURBELLARIANS

Natural History and Taxonomy

Anatomy and Physiology

Environmental Disorders and Preventive Medicine

Infectious Diseases

Neoplasia

Miscellaneous Disorders

Analgesia, Anesthesia, and Surgery

Drug Action Models

Treatment Protocols and Formulary

Chapter 6 GASTROPODS

Natural History and Taxonomy

Anatomy and Physiology

Environmental Disorders and Preventive Medicine

Infectious Diseases

Miscellaneous Disorders

Anesthesia, Analgesia, and Surgery

Treatment Protocols and Formulary

Additional Resource

Chapter 7 CEPHALOPODS

Introduction

Natural History and Taxonomy

Anatomy and Physiology

Environmental Disorders/Preventative Medicine

Infectious Diseases

Neoplasia

Miscellaneous Disorders

Analgesia, Anesthesia, and Surgery

Treatment Protocols and Formulary

Chapter 8 BIVALVES

Introduction

Basic Anatomy and Physiology

Behavior

Bivalves as Biological Filters

Bivalves as Bioindicators

Culture

Health Requirements

Chapter 9 ANNELIDS

Introduction

Polychaetes

Oligochaetes

Hirudineans

Chapter 10 HORSESHOE CRABS

Natural History and Taxonomy

Anatomy and Physiology

Environmental Diseases

Infectious Diseases

Diagnostic Techniques

Health Management

Anesthesia and Surgery

Chapter 11 SPIDERS

Introduction

Taxonomy and Natural History

Anatomy and Physiology

Environmental Disorders

Infectious Diseases and Parasites

Neoplasia

Miscellaneous Disorders

Anesthesia and Surgery

Examination and Treatment

Drugs and Therapeutic Agents

Legislation

Resources

Terms

Chapter 12 SCORPIONS

Introduction

Natural History and Taxonomy

Anatomy and Physiology

Medical Conditions

Restraint

Wound Treatment

Infectious Diseases

Venom Collection

Chapter 13 CRUSTACEANS

Introduction

Natural History and Taxonomy

Anatomy and Physiology

Diseases Affecting Many Crustaceans

Diseases of Shrimp

Diseases of Lobsters

Diseases of Crabs

Diseases of Hermit Crabs

Diseases of Crayfish

Diseases of Other Crustaceans

Neoplasia

Diagnostic Techniques for Crustacean Diseases

Diseases of Special Concern

Management of Crustacean Diseases

Analgesia, Anesthesia and Surgery

Euthanasia

Treatments and Formulary

Chapter 14 MYRIAPODS (CENTIPEDES AND MILLIPEDES)

Natural History and Taxonomy

Anatomy and Physiology

Legal Aspects of Keeping

Environmental Disorders and Preventive Medicine

Husbandry

Handling

Infectious Diseases

Neoplasia

Miscellaneous Disorders

Anesthesia, Analgesia, and Surgery

Diagnostic Techniques

Therapeutics

Chapter 15 INSECTS

Introduction

Natural History and Taxonomy

Anatomy and Physiology

Environmental Disorders and Preventive Medicine

Infectious Diseases

Neoplasia

Miscellaneous Disorders

Analgesia, Anesthesia, and Surgery

Treatment Protocols and Formulary

The Future

Acknowledgments

Some Websites Relating to Insects and their Study

Chapter 16 HONEYBEES

Introduction

Natural History and Taxonomy

Anatomy

Introduction to Honeybee Pathology

Environmental Disorders

Viruses and Honeybees

Bacterial Diseases

Parasitic Diseases

Fungal Disease

Predators and Enemies of Colonies and Hives

Acknowledgments

Chapter 17 BUTTERFLY HOUSES

Introduction

Purpose Value and Justifications

Legal Status of Butterfly Houses

Design and Management

Breeding

Staff

Health and Safety and Risk Assessment

Disease Exclusion and Control

Education, Research, and Conservation Activities

Other Types of Invertebrate Management

Acknowledgments

Additional Resource

Chapter 18 NEMATODES

Natural History and Taxonomy

Anatomy and Physiology

Environmental Disorders and Preventive Medicine

Infectious Diseases

Neoplasia

Miscellaneous Disorders

Anesthesia, Analgesia, and Surgery

Miscellaneous

Treatment Protocols and Formulary

Chapter 19 CHAETOGNATHS (ARROWWORMS)

Natural History and Taxonomy

Anatomy

Physiology

Environmental Disorders

Infectious Diseases

Neoplasia

Miscellaneous Disorders

Analgesia, Anesthesia, and Surgery

Treatment Protocols and Formulary

Chapter 20 ECHINODERMS

Natural History and Taxonomy

Anatomy and Physiology

Environmental Disorders and Preventive Medicine

Infectious Diseases

Neoplasia

Miscellaneous Conditions

Clinical Procedures

Treatment Protocols

Human and Animal Health Hazards

Chapter 21 UROCHORDATES

Introduction

Natural History and Taxonomy

Anatomy and Physiology

Infectious Diseases

Cup Cell Disease

Miscellaneous Conditions

Chapter 22 DIAGNOSTIC TECHNIQUES AND SAMPLE HANDLING

Introduction

Sample Collection from Living Specimens

Sample Collection

Summary

Resources

Fixatives and Solutions

Chapter 23 LAWS, RULES, AND REGULATING AGENCIES FOR INVERTEBRATES: U.S. AND INTERNATIONAL

Part I: United States of America

Introduction

Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES)

Endangered Species Act (ESA)

Magnuson–Stevens Fishery Conservation and Management Act (MSFCMA)

The Lacey Act (LA)

U.S. Department of Agriculture (USDA)

U.S. Fish and Wildlife Service (USFWS)

National Marine Fisheries Service (NMFS)

State and Local Regulations

Animal Welfare Act and Animal Welfare Regulations: Licenses and the IACUC

World Organization for Animal Health (OIE)

Summary

Part II: International

Introduction

Definitions

Specific Areas of Law Relevant to Invertebrates

Conservation: Species Protection

Other Areas of Law

Conclusion

Chapter 24 NEOPLASIA

Introduction

Understanding Invertebrate Neoplasia

Selected Invertebrate Neoplasms

Treatment

Chapter 25 EUTHANASIA

Chapter 26 INVERTEBRATE ANIMAL WELFARE

Introduction

Captive Care

Pain

Euthanasia

Acknowledgments

Chapter 27 INVERTEBRATE HEALTH ISSUES IN CONSERVATION

Introduction

Why Conserve Invertebrates?

Which Invertebrates to Conserve, and How

Regulation and Legislative Input

Wider Perspective

Management Planning: Threats to Invertebrates

Ex situ Conservation

Discussion

Chapter 28 SOURCES OF INVERTEBRATES AND SUPPLIES

Introduction

Anesthetic/Chemical Suppliers

Diagnostic Laboratories

Dietary Supplies

Environmental Monitoring

General Supplies

Husbandry/Exhibit Supplies

Sources of Invertebrates

Veterinary/Diagnostic Supplies

Chapter 29 REPORTABLE DISEASES OF INVERTEBRATES

Introduction

Diseases of Invertebrates Notifiable to the OIE

Diseases of Invertebrates Reportable to the USDA

Index

Color Plates

This edition first published 2012 © 2012 by John Wiley & Sons, Inc.

Chapter 4 remains with the U.S. government.

First Edition ©2006 Blackwell Publishing Ltd.

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Library of Congress Cataloging-in-Publication Data

Invertebrate medicine / editor, Gregory A. Lewbart.—2nd ed.

p. cm.

 Rev. ed. of: Invertebrate medicine / Gregory A. Lewbart. 2006.

 Includes bibliographical references and index.

 ISBN-13: 978-0-8138-1758-3 (hardcover : alk. paper)

 ISBN-10: 0-8138-1758-7

 1. Invertebrates—Diseases. 2. Veterinary medicine. I. Lewbart, Greg. II. Lewbart, Greg. Invertebrate medicine.

 SF997.5.I5L49 2012

 639'.4—dc23

2011018146

A catalogue record for this book is available from the British Library.

This book is published in the following electronic formats: ePDF 9780470960776; Wiley Online Library 9780470960806; ePub 9780470960783; Mobi 9780470960790

DEDICATION

To my wife, Diane Deresienski, for her constant love and support during the process of nurturing this work, and as my quarter-century-plus partner and teammate on this challenging and exciting journey named life!

CONTRIBUTORS

Robert S. Bakal, DVM, MS

Wake Forest, NC

Ilze K. Berzins, PhD, DVM

The John G. Shedd Aquarium

Michael S. Bodri, MS, VMD, PhD

Department of Biology

North Georgia College and State University

Robert A. Bullis, DVM, MS

Florida Keys Community College

John R. Chitty, BVetMed, CertZooMed, CBiol, MIBiol, MRCVS

Strathmore Veterinary Clinic

Larry S. Christian, BS

Department of Clinical Sciences

College of Veterinary Medicine

North Carolina State University

John E. Cooper, DTVM, FRCPath, FIBiol, FRCVS

Department of Veterinary Medicine

University of Cambridge

Margaret E. Cooper, LLB, FLS

Faculty of Veterinary Medicine

University of Nairobi, Kenya

and

DICE

The University of Kent

and

UK Comparative Pathology, Forensic and Wildlife Health Services

Flavio Corsin, MS, PhD

International Collaborating Centre for Aquaculture & Fisheries Sustainability (ICAFIS)

Vietnam Fisheries Society (VINAFIS)

Daniel S. Dombrowski, MS

College of Veterinary Medicine

North Carolina State University

and

North Carolina Museum of Natural Sciences

Laura Foster

Mayfair Animal Hospital

Cary, NC

Fredric L. Frye, DVM, MS, CBiol, FIBiol

La Primavera Farm

Amy L. Hancock, VMD

Marine Biological Laboratory

Craig A. Harms, DVM, PhD, Dipl ACZM

Department of Clinical Sciences

College of Veterinary Medicine

North Carolina State University

Center for Marine Sciences and Technology

Mac Law, DVM, PhD, Dipl ACVP

Department of Population Health and Pathobiology

College of Veterinary Medicine

North Carolina State University

Wade Lehmann, PhD

Office of Science & Technology, Office of Water

U.S. Environmental Protection Agency

Jay Levine, DVM, MPH

Aquatic Animal Epidemiology and Conservation Genomics Laboratory

Population Health and Pathobiology

College of Veterinary Medicine

North Carolina State University

Gregory A. Lewbart, MS, VMD, Dipl ACZM

Department of Clinical Sciences

College of Veterinary Medicine

North Carolina State University

Michael J. Murray, DVM

Monterey Bay Aquarium

Timothy R. New, PhD

Department of Zoology

La Trobe University

Edward J. Noga, MS, DVM

South Eastern Aquatechnologies, Inc.

Esther C. Peters, PhD

Department of Environmental Science and Policy

George Mason University

Romain Pizzi, BVSc, MS, CertZooMed, FRES, MRCVS, MACVSc(Surg)

Zoological Medicine Ltd.

Taylor L. Reynolds DVM, Dipl ACVP

Johns Hopkins School of Medicine

Joseph M. Scimeca, DVM, PhD

Research Development and Administration-SIUC

Southern Illinois University

Stephen A. Smith, DVM, PhD

Department of Biomedical Sciences and Pathobiology

Virginia/Maryland Regional College of Veterinary Medicine

Virginia Polytechnic Institute and State University

Roxanna Smolowitz, DVM

Aquatic Diagnostic Laboratory

Roger Williams University

Michael K. Stoskopf, DVM, PhD, Dipl ACZM

Department of Clinical Sciences

College of Veterinary Medicine

North Carolina State University

Bernard Vallat, DVM, PhD

Director General

World Organisation for Animal Health

Nicolas Vidal-Naquet, DVM

DIE Apiculture-Pathologie Apicole

Trevor T. Zachariah, DVM, MS

College of Veterinary Medicine

University of Illinois at Urbana-Champaign

FOREWORD

This is a very auspicious year for the publication of this second edition, being the 250th anniversary of the world’s first veterinary school, which was founded in Lyon, France, in 1761. Since then, many veterinarians have made substantial contributions to the world’s knowledge of animal health and welfare.

The publication of this second edition continues this proud tradition, with diverse subject material that illustrates the diversity and scope of the veterinary profession. The editor has succeeded, once again, in bringing together a group of scientists, primarily veterinarians, from institutions with established and respected expertise in the fields of zoological and aquatic animal medicine. The result is an even more comprehensive veterinary textbook, which addresses the care of common terrestrial and aquatic invertebrates that are kept as pets, displayed in aquariums and zoos, used for research, or consumed by humans.

Veterinarians play an increasingly important and diverse role in society. They address the health needs of every species of animal and play a critical role in environmental protection, food safety, animal welfare, and public health. In the last 250 years, when society has identified a need for veterinary advice, veterinarians have responded to the challenge and taken steps to develop the knowledge and skills required to provide appropriate veterinary care. The authors of this textbook evidence this “can-do” attitude. The contributors have developed specialized expertise in the clinical assessment and medical treatment of the diverse range of invertebrate species covered in this edition.

A key role of the World Organisation for Animal Health (OIE) is developing standards, such as the Aquatic Animal Health Code and the Manual of Diagnostic Tests for Aquatic Animals, which are recognized under the World Trade Organization Agreement on the Application of Sanitary and Phytosanitary Measures to ensure safe trade in aquatic animals and their products. The development of these OIE standards depends on the input of veterinarians and other biologists with expertise in aquatic animal health. Similarly, a publication of this type is only possible because of the availability of veterinarians with specialized knowledge, in this case, on invertebrate species.

In keeping with 2011 as the “Year of the Veterinarian,” this publication showcases the versatility of the veterinary profession in meeting the challenges posed by contemporary society. This textbook is an impressive collection of current knowledge on invertebrate medicine. It will serve as a valuable resource to graduate and undergraduate veterinarians and care providers who work with this diverse range of species.

Dr. Bernard Vallat

Director General

World Organisation for Animal Health (OIE)

Paris, February 2011

PREFACE

For many decades, invertebrates have been kept as pets, displayed in aquariums and zoos, used for research, and consumed. Maintaining live invertebrates in captivity has become more sophisticated and popular in recent years. Arthropod zoos and insectariums, jellyfish exhibits, and captive living coral reefs are relatively commonplace today but were rare or nonexistent 25 years ago. Despite this popularity, diversity, and economic importance, veterinary medicine has traditionally paid relatively little attention to this huge segment (over 95% of the earth’s animal species) of the animal kingdom.

My own interest in invertebrates started on a family trip to Campobello Island in New Brunswick, Canada, nearly four decades ago. Finding a sand dollar test on the beach was an exciting moment for a young boy, as was the subsequent quest to identify it. Years later, I found myself studying invertebrate zoology at Gettysburg College; my professor was Dr. Robert D. Barnes. After exciting field trips with Dr. Barnes to the Duke University Marine Laboratory in Beaufort, North Carolina, and the Bermuda Biological Station (now the Bermuda Institute of Ocean Science) in St. Georges, Bermuda, I was definitely hooked! These educational experiences were a pivotal point in my life and I hold a special place in my heart and mind for Dr. Barnes and his inspirational teaching.

I began thinking about working on a veterinary text for invertebrates in the early 1990s and was very happy (and just a little apprehensive) when I signed a contract to edit the first edition of this text with Blackwell Publishing Professional (at the time Iowa State University Press) in 2001. The book was published in 2006, and after several years, it became clear there was a need for a follow-up edition that would capture some of the exciting and dynamic changes and advances occurring in the field. The editors at Wiley-Blackwell were instrumental in the pursuit of this edition and agreed that an updated text would be a welcome addition to the market.

This book is the product of a concerted effort by a group of dedicated and experienced authors on the topic of invertebrate animal medicine. This is not an invertebrate zoology text and is by no means comprehensive with regard to the anatomy, physiology, natural history, and taxonomy of the myriad of invertebrate taxa. This is a veterinary and husbandry text about invertebrate animals that includes pertinent biological data as well as state-of-the-science information pertaining to medicine and the clinical condition. It is my hope that this book will be a valuable guide to those charged with the medical care and well-being of both captive and wild invertebrate animals.

At the North Carolina State University College of Veterinary Medicine (NCSU-CVM), my students and colleagues continually inspire me. Nearly a decade ago, the veterinary students started the Invertebrate Medicine Club, and we have offered an intensive 1-week elective course on invertebrate medicine for the past 8 years. The NCSU-CVM administration has been extremely encouraging of these efforts, and I am grateful and fortunate to be working in such a rich, supportive environment.

Invertebrates and their health management are a good fit with the widely adopted and held philosophy of “one health” or “one medicine,” which pertains to all animal species, including humans. It is my hope that this text will contribute to the advances and understanding of this approach to global health and well-being.

I am very excited about invertebrate animal medicine and hope you will join me in this excitement. There is much work to be done in this realm in which the opportunities are truly endless.

Gregory A. Lewbart, MS, VMD, Dipl ACZM

Professor of Aquatic Animal Medicine

Department of Clinical Sciences

North Carolina State University

College of Veterinary Medicine

ACKNOWLEDGMENTS

A project like this could not come to fruition without the help, support, and assistance of many individuals.

For their instruction, inspiration, guidance, support, and mentoring during my student years, I thank all of my mentors and professors, but, in particular, Donald Abt, Robert Barnes, Philip Bookman, Dale Dickey, John Gratzek, Louis Leibovitz, William Medway, Trish Morse, Nathan “Doc” Riser, Ralph Sorensen, and Richard Wolke. Doc was my master’s thesis advisor and I am especially grateful for his knowledge, wisdom, and patience. I am fortunate to be associated with the North Carolina State University College of Veterinary Medicine (NCSU-CVM), a fine, progressive institution of higher learning. The school’s founding Dean, Terrence Curtin, is a valued mentor, good friend, and true inspiration. His drive and progressive vision helped create this wonderful academic treasure. I am grateful to all of my NCSU-CVM friends and colleagues. Warwick Arden, Oscar Fletcher, Elizabeth Hardie, Craig Harms, Edward Noga, Elizabeth Stone, and Michael Stoskopf have been especially supportive.

I also collectively thank the veterinary students I have worked with, both at the NCSU-CVM and from other colleges of veterinary medicine. They are the bright future of our profession, and I know some days they teach me more than I teach them. Dan Dombrowski, one of this book’s authors, deserves special mention as an inspiration to all those interested in keeping and caring for the earth’s invertebrates.

The following individuals were generous in providing case material, images, suggestions, and other assistance: Genevieve Anderson, Rich Aronson, Herman Berkhoff, Charles Bland, Shane Boylan, J. Brock, Mike Buchal, Rick Cawthorn, James Clark, Angelo Colorni, Liza Dadone, Diane Deresienski, Chris Eads, David Engel, Kate Freeman, Carlton Goldthwaite, Stacey Gore, Steve Gschmeissner, Malcolm Hill, Mike Holliman, William Holmes, C. Hooper, Sarah Joyner, Andy Kaufmann, Kelly Krell, Rita Krickl, Wade Lehmann, Monique L’Hostis, Douglas Mader, Stuart May, Gavin Merrifield, H. Millar, Melissa Miner, Jim Moore, Alf Nilsen, Hendrik Nollens, Melanie Rembert, David Rotstein, Clay Rouse, Christophe Roy, Johanna Sherrill, Jerry Stevens, Cinamon Vann, Peter van Zuuren, Nicole Webster, and Randy Wilder. Eric Borneman and Richard Fox were especially generous in providing images.

I am grateful to Brenda Bunch and Alison Schroeer, both outstanding biological illustrators, who provided high-quality images and were tolerant of my numerous requests and deadlines. I also thank the helpful personnel of the NCSU-CVM Biomedical Communications Department.

I am very grateful to the talented group of authors who constructed this diverse collection of chapters; there would be no Invertebrate Medicine without their hard work, dedication, and commitment to the project. As the book’s editor, I take full responsibility for any errors or omissions.

The folks at Wiley-Blackwell have been exceptional through this entire process. They are true professionals and a pleasure to work with. I specifically acknowledge David Rosenbaum, the person who “signed me up” for the first edition; Cheryl Garton, who helped in the early stages of the first edition; Justin Jeffryes, who was instrumental in securing this edition; Antonia Seymour, the editorial director supervising this project; Susan Engelken, my accurate and efficient managing editor, who made certain everything was in order for the presses; Carrie Horn, my production editor; and, finally, Nancy (Simmerman) Turner, my initial managing editor, who was always there for this book and me. Thanks Nancy!

Stephanie Sakson did a tremendous job of facilitating the copyediting and deserves special mention. She and Toppan Best-set Premedia Limited’s meticulous editorial skills have greatly improved the book.

I really don’t think I could have completed this book or the previous edition without the dedicated and selfless help of Shane Christian during the past 9 years. He is simply the most reliable and competent person I know, and his efforts allowed me the time to focus on this project. Thanks Shane!

Finally, I am so very grateful for the love and support I consistently receive from my wife, Diane Deresienski. Her advice is always sound. Her patience never thin. Her ideas consistently good. Her hugs always warm. And she’s a top-notch veterinarian whom I deeply admire and respect.

G.A.L.

Chapter 1

INTRODUCTION

Gregory A. Lewbart

This book, the second edition of Invertebrate Medicine, represents a concerted effort by a group of dedicated authors on the topic of invertebrate animal medicine. It has been substantially expanded from the first edition to reflect the tremendous growth of the pertinent literature and work that is being accomplished in the fields of invertebrate animal medicine, disease investigation, conservation, husbandry, and animal welfare. With the increased merging of animal and human medicine (“one health” or “one medicine”); the growing importance of invertebrates in biomedical research; and the many ongoing efforts to preserve habitat, protect at-risk species, and learn more about how invertebrates help define and connect ecosystems, this new edition is both warranted and timely.

Six new chapters (“Coral Reef Systems,” “Butterfly Houses,” “Honeybees,” “Conservation and Preservation,” “Welfare,” and “Sources of Supplies”) have debuted and three first edition appendices (“Neoplasia,” “Euthanasia,” and “Reportable Diseases”) have been expanded into full chapters. The “Laws and Regulations” chapter has been broadened, with the addition of a new author, to include more international information.

This is not an invertebrate zoology text and is by no means comprehensive with regard to the anatomy, physiology, natural history, and taxonomy of the myriad of invertebrate taxa. This is a veterinary text about invertebrate animals. It includes pertinent biologic information as well as the state-of-the-science information pertaining to medicine and the clinical condition.

What sort of topic is invertebrate medicine? And what exactly are invertebrates? Ruppert and Barnes (1994) have said that the invertebrates are a group of unrelated taxa that share no universal “positive” traits. Undergraduate and graduate courses are dedicated to invertebrate zoology or even to specific parts of this topic, such as entomology, malacology, or protozoology. Simply put, the invertebrates are a collection of animals, comprising more than 95% of the earth’s species, unified by the lack of a vertebral column.

Relatively recent and interesting genetic research has produced some intriguing ideas about intertaxonomic relationships regarding invertebrates and vertebrates. With the complete genetic sequencing for a number of species, including the fruit fly (Drosophila melanogster), an important biomedical research nematode (Caenorhabditis elegans), humans (Homo sapiens), and more recently an annelid worm (Platyneries dumerlii), some theories on animal evolution and phylogenetic relationships are being revisited and revised (Telford, 2004; Wolf et al., 2004; Federov and Federova, 2006). The more traditional “Coelomata hypothesis,” linking animals (many invertebrates and all vertebrates) possessing a coelom, may be giving way to the “Ecdysozoa hypothesis,” which places molting invertebrates, such as arthropods and introvertans (nematodes and other pseudocoelomates), in the same broad taxonomic group (Telford, 2004). Based largely on the comparison of small nonprotein-coding DNA segments called introns, researchers have learned that annelid worms may be more closely related to humans than they are to either insects or nematodes (Raible et al., 2005).

Detailed accounts and discussion of these theories, and invertebrate taxonomy and phylogeny in general, are beyond the scope of this medical text. Still, this is an active area of sophisticated research, the results and subsequent deliberations of which are worth being aware of for the clinician or caregiver charged with invertebrate animal health, welfare, and husbandry.

Depending on the text or investigator, there are currently over 30 recognized phyla of invertebrates (not including the protozoans). Many of these might be considered obscure, but for no better reason than they may contain few species, microscopic representatives, or lack obvious economic value to humans. In reality, each phylum and its members are important to the diversity and survival of the planet, even if the group is only studied by a small number of investigators. Unfortunately, very little is known about the veterinary aspects of many of these taxa, and writing a comprehensive text for all invertebrate phyla would currently be a daunting and somewhat inefficient task. Consequently, I have elected to include the most economically important and “visible” metazoan taxonomic groups. Exclusively parasitic taxa (e.g., trematodes, cestodes, and acanthocephalans) are only touched upon. Table 1.1, which lists the major taxonomic groups (along with brief descriptions) that do not have their own chapter, has been included in an effort to remind readers of the diversity of the invertebrate animal kingdom. Table 1.2 provides a snapshot of animal diversity with regard to the number of described species and habitat. I encourage interested readers to obtain one or more of the general invertebrate zoology texts listed under the section “General Invertebrate Zoology Resources,” where detailed descriptions of the various groups in Table 1.1 and throughout this book can be found.

Table 1.1. Invertebrate phyla and major classes not reviewed in this book

Taxonomy and descriptions largely from Ruppert et al. (2004).

Placozoa: A monotypic phylum containing only the species Trichoplax adhaerens. This primitive amoeboid metazoan is flattened, less than 3 mm in diameter, and exhibits extracellular digestion of detritus and algae.Orthonectida: A very small phylum (about 20 species) of very small (no larger than 1 mm) internal parasites of other invertebrates such as bivalves, polychaetes, tunicates, turbellarians, and nemerteans.Dicyemida: This phylum contains about 75 species of very thin renal parasites of cephalopods.Nemertea: This diverse phylum contains approximately 1150 species of ribbon worms, which tend to be much larger and longer than flatworms. Unlike flatworms, nemerteans have a true coelomic circulatory system. Most are marine, but there are a few freshwater and terrestrial forms. Nemerteans are predators and use a long, eversible proboscis to capture and retain prey.Mollusk groupsAplacophora: This class consists of about 300 species of small, vermiform, marine animals that live at depths of between 200 and 7000 m.Polyplacophora: Commonly known as the chitons, these interesting mollusks are mobile but spend most of their time tightly adhered to rocky substrates. There are approximately 800 exclusively marine species described. All have eight valves or plates (hence the name of the class) that overlap and are connected by soft tissue and surrounded by a muscular “girdle.” Most species could rest in your palm, but one, Cryptochiton sp., the stocky gumshoe chiton, can reach a length of about 40 cm.Scaphopoda: Known as the “tusk” or “tooth” mollusks because of their shell shape. The approximately 500 species are all marine, and most are burrowers with the head facing down within the substrate.Echiura: Commonly known as the spoon worms, most of the 150 species either live in U-shaped burrows or between rocks closely associated with the marine environment. Most are deposit feeders, and some are an important food source for fishes. The name comes from the large and flared prostomium that resembles a spoon or small scoop.Sipuncula: The sipunculids, or peanut worms, are a group of about 150 marine burrowing species. Most are smaller than 10 cm, but some can reach 70 cm in length. They possess an interesting feeding structure termed the introvert that can be expelled from or retracted into the main body or trunk.Onychophora: This group of tropical, terrestrial animals (110 known species) is commonly referred to as velvet worms or walking worms. They are segmented and aligned with arthropods. In fact, some workers include the phyla Onychophora, Tardigrada, and Arthropoda in the superphylum Panarthropoda. Velvet worms prey on smaller arthropods by capturing them with slime ejected from paired glands near the mouth.Tardigrada: If the water bears, as they are commonly known, grew larger (most are less than 1 mm long), they would surely be common and popular pets and display animals. There are marine, freshwater, and terrestrial representatives among the 800-plus species in this group of taxonomically mysterious animals. They have features in common with the arthropods but are different enough to warrant their own phylum. Perhaps their most interesting attribute is their ability to undergo cryptobiosis and form desiccated tuns, which can withstand adverse environmental conditions. In fact, some tardigrades may live as long as 100 years with the aid of cryptobiosis.Gastrotricha: Many of the 500 species belonging to this microscopic phylum are interstitial. Most look like miniature bowling pins atop two small pegs. There are freshwater and marine forms.Nematomorpha: The horsehair worms superficially resemble nematodes but are very long and free-living as adults. The larvae usually parasitize either crustaceans or insects. Approximately 325 species have been described.Priapulida: This small phylum containing just 18 species is all marine and benthic. They are cylindrical and resemble a small cactus.Loricifera: This interesting and microscopic marine phylum (all appear to be interstitial) was not known to science until 1983. Many of the 100 or so known species have not yet been described due to the difficulty of examining fresh, living specimens. These little creatures are so dogged in their attachment to sand grains that only freshwater will dislodge them, causing osmotic damage and distortion of their anatomy.Kinorhyncha: The mud dragons somewhat resemble the Gastrotricha in general shape but have an oral feeding structure called the oral styles at the end of a movable introvert. Most are microscopic and are either interstitial or benthic on mud and sand. There are approximately 150 species and all are marine.Gnathostomulida: Virtually all 80 known species are marine, interstitial, and less than 1 mm long. They are vermiform and were not known to science until 1956.Rotifera: Most occur in freshwater, but there are marine and terrestrial (primarily in water films) species. They are defined and frequently identified by the ciliated corona or wheel organ near the head. Some rotifers are extremely important in freshwater and marine food chains (in some cases, hundreds may be found in a liter of water) and are also commonly reared to support invertebrate and finfish aquaculture. There are approximately 2000 described species.Acanthocephala: A totally parasitic group containing 1150 species. They are commonly known as thorny-headed worms, and some are important parasites of wild and domestic vertebrates. Most use other invertebrates as intermediate hosts.Kamptozoa: Also known as Entoprocta, the 150 species are nearly all marine. Most are stalked, and some people refer to them as nodders because of the zooid’s tendency to nod or rock at the end of the stalk. Although some zoologists still classify them as bryozoans, these animals differ in their complete lack of a coelomic cavity. Some zoologists feel the morphological similarities between the groups are convergent.Cycliophora: This small (in size and species number) phylum was not introduced to science until 1995. The single described species, Symbion pandora, exhibits a commensal lifestyle with a lobster (Nephrops sp.). Other as yet undescribed species are commensal with other crustaceans, including the American lobster, Homarus americanus. They are suspension feeders and have a complex reproductive cycle with both asexual and sexual life stages. None of the life stages are over 0.5 mm long.Phoronida: There are just 14 species in two genera of these sessile marine creatures. These vermiform animals live in chitinous tubes that they secrete. Although externally they are bilaterally symmetrical, internally the left side is dominant. They feed by means of a lophophore and are grouped into the superphylum Lophophorata along with the bryozoans and brachiopods.Brachiopoda: The brachiopods, or lamp shells, are an interesting group of 350 extant marine species that grossly resemble bivalve mollusks. Thousands of species are known from the fossil record, in part due to their mineralized valves that are preserved well. They are not related to mollusks, and the hard valves that protect the soft body are oriented opposite that of the bivalve’s. They feed with the aid of a lophophore, placing them in the superphylum Lophophorata. Most are the size of small cherrystone clams and frequently turn up in shops specializing in fossils. Most species occur in colder waters.Bryozoa: Known as the moss animals, these are common animals that can be found on many marine substrates (there are a few freshwater species), including rocks, algae, pilings, and even living animals such as sea turtles. With nearly 5000 species, this phylum is the best known of the Lophophorata and is studied as part of nearly all basic invertebrate zoology courses. The vast majority are colonial, although there is one solitary genus. From a distance, they may look more like plants than animals to casual observers. Some colonies are polymorphic, whereas other species are monomorphic. They are filter feeders, using the lophophore to trap and retain small food items.ArthropodaPycnogonida: Known commonly as the sea spiders, this class of arthropods contains about 1000 known species. They are all marine and widely distributed, with most occurring in benthic habitats. Very few species are larger than 1 cm, and although they resemble a true spider, they are not close relatives.Xenoturbella: This flatworm-like group of benthic, ciliated, marine organisms was first discovered in the early years of the 21st century but initially and incorrectly considered a free living turbellarian. Later, it was erroneously linked to other taxonomic groups, including hemichordates and echinoderms. In the 1990s, based on Xenoturbella bocki samples that were “contaminated” by food the animal was consuming, the group was reclassified as a bivalve mollusk. Recent molecular work has elevated this cryptic and small group of animals into their own phylum of deuterostomes (bilaterally symmetrical animals like the echinoderms, hemichordates, and chordates, which possess radial cleavage, an ectodermal mouth, and anal opening originating from the blastopore). Unique and unusual traits include lack of a gut or defined coelomic cavity, absence of gonads or excretory organs, and a nerve net system without a cephalic or central ganglionic region.

Table 1.2. Habitats and approximate metazoan species numbers

Modified from Pearse et al. (1987), p. 7; with taxonomic and number updates from Ruppert et al. (2004).

#, 1–100;

##, 100–1000;

###, 1000–10,000;

####, 10,000–100,000; and

#####, over 100,000.

References

Bourlat SJ, Nielsen C, Lockyer AE, Littlewood DTJ, and Telford MJ. 2003. Xenoturbella is a deuterostome that eats molluscs. Nature 424:925–928.

Bourlat SJ, Juliusdottir T, Lowe CJ, Freeman R, Aronowicz J, Kirschner M, Lander ES, Thorndyke M, Nakano H, Kohn AB, Heyland A, Moroz LL, Copley RR, and Telford MJ. 2006. Deuterostome phylogeny reveals monophyletic chordates and the new phylum Xenoturbellida. Nature 444:85–88.

Federov A, and Federova L. 2006. Where is the difference between the genomes of humans and annelids? Genome Biol 7(1):203.

Pearse V, Pearse J, Buchsbaum M, and Buchsbaum R. 1987. Living Invertebrates. Blackwell Scientific, Palo Alto, CA.

Raible F, Tessmar-Raible K, Osoegawa K, Wincker P, Jubin C Balavoine G, Ferrier D, Benes V, de Jong P, Weissenbach J, Bork P, and Arendt D. 2005. Vertebrate-type intron-rich genes in the marine annelid Platynereis dumerilii. Science 310:1325–1326.

Ruppert EE, and Barnes RD. 1994. Invertebrate Zoology, 6th ed. Saunders College, Philadelphia, pp. 499–595.

Ruppert EE, Fox RS, and Barnes RD. 2004. Invertebrate Zoology: A Functional Evolutionary Approach, 7th ed. Brooks/Cole—Thomson Learning, Belmont, CA.

Telford MJ. 2004. Animal phylogeny: Back to the Coelomata? Curr Biol 14:R274–R276.

Wolf YI, Rogozin IB, and Koonin EV. 2004. Coelomata and not Ecdysozoa: Evidence from genome-wide phylogenetic analysis. Genome Res 14:29–36.

General Invertebrate Zoology References

Barnes RSK, Calow P, Olive PJW, Golding DW, and Spicer JI. 2001. The Invertebrates: A Synthesis, 3rd ed. Blackwell Science Ltd., Oxford.

Barrington EJW. 1979. Invertebrate Structure and Function, 2nd ed. John Wiley & Sons Inc., New York.

Brusca RC, and Brusca GJ. 2003. Invertebrates, 2nd ed. Sinauer Associates, Inc., Sunderland, MA.

Cohen WD, ed. 1985. Blood Cells of Marine Invertebrates: Experimental Systems in Cell Biology and Comparative Physiology. Alan R. Liss, Inc., New York.

Conn DB. 1991. Atlas of Invertebrate Reproduction and Development. John Wiley & Sons, Inc., New York.

Fretter V, and Graham A. 1976. A Functional Anatomy of Invertebrates. Academic Press, London.

Harrison FW. 1991–1999. Microscopic Anatomy of Invertebrates, 15 Volumes. Wiley-Liss, New York.

Hyman LH. 1940–1967. The Invertebrates, Volumes 1–6. McGraw-Hill, New York.

Kozloff EN. 1990. Invertebrates. Saunders, Philadelphia.

Meglitsch PA, and Schram FR. 1991. Invertebrate Zoology, 3rd ed. Oxford University Press, New York.

New TR. 1995. Introduction to Invertebrate Conservation Biology. Oxford University Press, Oxford.

Pearse V, Pearse J, Buchsbaum M, and Buchsbaum R. 1987. Living Invertebrates. Blackwell Scientific, Palo Alto, CA.

Pechenik JA. 2005. Biology of the Invertebrates, 5th ed. McGraw-Hill, Higher Education, Boston.

Ruppert EE, Fox RS, and Barnes RD. 2004. Invertebrate Zoology: A Functional Evolutionary Approach, 7th ed. Brooks/Cole—Thomson Learning, Belmont, CA.

Sherman IW, and Sherman VG. 1976. The Invertebrates: Function and Form, 2nd ed. Macmillan, New York.

Stachowitsch M. 1992. The Invertebrates: An Illustrated Glossary. Wiley-Liss, New York.

Young CM, Sewell MA, and Rice ME. 2002. Atlas of Marine Invertebrate Larvae. Academic Press, London.

General References for Invertebrate Medicine, Husbandry, Culture, and Pathology

Frye FL. 1992. Captive Invertebrates: A Guide to Their Biology and Husbandry. Krieger Publishing, Malabar, FL.

Kinne O, ed. 1980–1990. Diseases of Marine Animals, Volumes 1–3. John Wiley & Sons, New York. All volumes are now available online by the Inter-Research Science Center http://www.int-res.com (accessed May 24, 2011).

Lewbart GA. 2012. Invertebrates. In: Carpenter J, ed. Exotic Animal Formulary, 4th ed. Elsevier Publishing, Philadelphia, in press.

Mitsuhashi J. 2002. Invertebrate Tissue Culture Methods. Springer-Verlag, Tokyo.

Mothersill C, and Austin B. 2000. Aquatic Invertebrate Cell Culture. Springer Praxis Publishing, Chichester, UK.

Stolen JS, Fletcher TC, Smith SA, Zelikoff JT, Kaattari SL, Anderson RS, Soderhall K, and Weeks-Perkins BA. 1995. Techniques in Fish Immunology, Fish Immunlogy Communications 4 (FITC 4). SOS Publications, Fair Haven, NJ.

Chapter 2

SPONGES

Gregory A. Lewbart

Natural History and Taxonomy

The phylum Porifera is a diverse group of primitive animals commonly referred to as the sponges. Until the middle of the 18th century, sponges were actually classified as plants (Ruppert and Barnes, 1994). Sponges occur in the fossil record back to the Precambrian era (over 600 million years ago) and were the most important contributors to reefs during the Paleozoic and Mesozoic eras (Hooper and Van Soest, 2002). All members lack defined organs; differentiated cells within connective tissue perform necessary biologic functions. A unique system of water canals facilitates transport of food, waste products, and gametes. Nearly all species are sessile and most are marine. Of the approximately 8300 species belonging to over 680 genera, only about 3% occur in freshwater (Hooper and Van Soest, 2002; Ruppert et al., 2004). Sponges are normally found on firm substrates in shallow water, although some occur on soft bottoms.