The Mollusks
A Guide to Their Study, Collection, and Preservation
Edited by Charles F. Sturm Timothy A. Pearce Ángel Valdés
A Publication of the American Malacological Society Los Angeles and Pittsburgh
Universal Publishers Boca Raton, Florida USA • 2006
The Mollusks: A Guide to Their Study, Collection, and Preservation
Copyright 2006 American Malacological Society
All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or other- wise without the prior permission of the copyright owner.
Acknowledgment of permission to use copyrighted material: Carl Zeiss, Germany: Figure 7.1 Eastman Kodak Company: Figure 7.4 A. & C. Black, London: Figure 19.2 Daniel Geiger: Figures 20.1 and 20.3 Santa Barbara Museum of Natural History: Figures 27.1-27.9
Acknowledgment of permission to use non-copyrighted material: David Mulliner: Figure 3.1 Illinois Natural History Survey: Figure 25.1 The Festivus, San Diego Shell Club: Figure 24.4
This volume should be cited as follows: Sturm, C. F., T. A. Pearce, and A. Valdés. 2006. The Mollusks: A Guide to Their Study, Collection, and Preservation. American Malacological Society, Pittsburgh, PA, U.S.A. Pp. xii + 445.
Chapters should be cited as based on the following example: Sturm, C. F., T. A. Pearce, and A. Valdés. 2006. Chapter 1. The Mollusks: Introductory Comments. In: C. F. Sturm, T. A. Pearce, and A. Valdés, eds., The Mollusks: A Guide to Their Study, Collection, and Preservation. American Malacological Society, Pittsburgh, PA., U.S.A. Pp. 1-7.
Universal Publishers Boca Raton, Florida USA • 2006 ISBN: 1-58112-930-0 (paperback) ISBN: 1-58112-931-9 (ebook)
Universal-Publishers.com
Library of Congress Cataloging-in-Publication Data
The mollusks : a guide to their study, collection, and preservation / edited by Charles F. Sturm, Timothy A. Pearce, Ángel Valdés. p. cm. ISBN 1-58112-930-0 (pbk. : alk. paper) -- ISBN 1-58112-931-9 (ebook) 1. Mollusks--Collection and preservation. 2. Mollusks--Study and teaching. I. Sturm, Charles, F. 1953- II. Pearce, Timothy A., 1954- III. Valdés, Ángel. QL406.5.M65 2006 594.075--dc22 2006016238 Preface
This volume promotes the educational mission of the American Malacological Society. The editors and contributors have brought together a broad range of topics within the field of malacology. It is our expectation that these topics will be of interest and use to amateur and professional malacologists.
The chapters in this publication have been peer-reviewed. Each chapter was reviewed by at least one of the editors and by a minimum of two outside reviewers, including at least one amateur and one professional malacologist. Chapters were then revised by authors and again reviewed by the editors, and occasionally, when warranted, by another outside reviewer. Then authors made a second round of changes and editors conducted a final review. While this review process has eliminated some errors and inconsistencies, some may remain.
If anyone does uncover any error or inconsistency in formatting, grammar, or style, the editors would appreciate it being brought to their attention. Constructive suggestions for improving this volume are welcome. Comments can be sent to the editors at
C. F. Sturm T. A. Pearce A. Valdés
vii
Table of Contents
Frontispiece ...... ii Preface ...... vii Table of Contents ...... ix 1 The Mollusks: Introductory Comments ...... 1 2 Field and Laboratory Methods in Malacology ...... 9 3 Remote Bottom Collecting ...... 33 4 Snorkeling and SCUBA Diving ...... 41 5 Archival and Curatorial Methods ...... 45 6 Digital Imaging: Flatbed Scanners and Digital Cameras ...... 59 7 Applied Film Photography in Systematic Malacology ...... 73 8 Computerizing Shell Collections ...... 101 9 The Molluscan Literature: Geographic and Taxonomic Works ...... 111 10 Taxonomy and Taxonomic Writing: A Primer ...... 147 11 Cladistics and Molecular Techniques: A Primer ...... 161 12 Organizations, Meetings, and Malacology ...... 173 13 Museums and Malacology ...... 181 14 Donating Amateur Collections to Museums ...... 189 15 Fossil Mollusks ...... 197 16 Aplacophora ...... 207 17 Monoplacophora ...... 211 18 Polyplacophora ...... 217 19 Scaphopoda: The Tusk Shells ...... 229 20 Cephalopoda ...... 239 21 Freshwater Gastropoda ...... 251 22 Terrestrial Gastropoda ...... 261 23 Rearing Terrestrial Gastropoda ...... 287 24 Marine Gastropoda ...... 295 25 Unionoida: Freshwater Mussels ...... 313 26 Non-Unionoid Freshwater Bivalvia ...... 327 27 Marine Bivalvia ...... 339 28 The Marine Aquarium: A Research Tool ...... 349 29 An Introduction to Shell-forming Marine Organisms ...... 359 30 Conservation and Extinction of the Freshwater Molluscan Fauna of North America ...... 373 31 Issues in Marine Conservation ...... 385 Appendices ...... 417 Appendix 1: Morphological Features of Gastropod and Bivalve (Pelecypod) Shells ...... 418 Appendix 2: Expanded Table of Contents ...... 427 Glossary ...... 441
ix
Contributors
When two addresses are listed for contributors, the first is their current address while the second is the institution where they were when they submitted the first draft of their chapter.
Frank E. Anderson Jim Cordy Chapter 20 Chapter 4 Department of Zoology Merritt Island, FL Southern Illinois University Carbondale, IL Clement L. Counts, III Chapter 17 Patrick Baker Department of Natural Sciences Chapter 31 University of Maryland Eastern Shore Department of Fisheries and Aquatic Sciences Princess Anne, MD University of Florida Gainesville, FL Kevin S. Cummings Chapters 9, 25 B. R. Bales (1876-1946) Curator of Mollusks Chapter 2 Illinois Natural History Survey Center for Biodiversity Arthur E. Bogan Champaign, IL Chapters 25, 30 Curator of Aquatic Invertebrates Robert T. Dillon, Jr. North Carolina Museum of Natural Sciences Chapter 21 Raleigh, NC Department of Biology College of Charleston Thomas A. Burch, MD Charleston, SC Chapter 3 Bremerton, WA Daniel L. Geiger Chapters 7, 10, 24 David Campbell Research Associate Chapters 11, 15 Santa Barbara Museum of Natural History University of Alabama Santa Barbara, CA Tuscaloosa, AL Lucía M. Gutiérrez Department of Geology Chapter 29 University of North Carolina Bióloga Marina y Científica del Medio Ambiente Chapel Hill, NC Guatemala City, Guatemala
Eugene V. Coan Environmental Protection Commission of Hills- Chapter 27 borough County Santa Barbara Museum of Natural History Tampa, FL Santa Barbara, CA Alexei V. Korniushin (1962-2004) Bobbi Cordy Chapter 26 Chapter 4 Institute of Zoology and Zoological Museum Merritt Island, FL The National Academy of Sciences of Ukraine Kiev, Ukraine
xi Ross Mayhew Amélie H. Scheltema Chapter 2 Chapter 16 Schooner Specimen Shells Woods Hole Oceanographic Institution Halifax, Nova Scotia, Canada Woods Hole, MA
Fabio Moretzsohn Enrico Schwabe Chapter 6 Chapter 9, 18 Center for Coastal Studies Department Mollusca Texas A&M University - Corpus Christi Zoologische Staatssammlung Muenchen Corpus Christi, TX Munich, Germany
Department of Zoology Charles F. Sturm University of Hawaii Chapters 1, 2, 3, 5, 9, 12, 13, 28, Glossary Honolulu, HI Research Associate - Section of Mollusks Carnegie Museum of Natural History Aydin Örstan Pittsburgh, PA Chapters 22, 23 Research Associate - Section of Mollusks Ángel Valdés Carnegie Museum of Natural History Chapter 1 Pittsburgh, PA Associate Curator of Malacology Natural History Museum of Los Angeles County Timothy A. Pearce Los Angeles, CA Chapters 1, 9, 14, 22, Glossary Curator - Section of Mollusks Paul Valentich-Scott Carnegie Museum of Natural History Chapter 27 Pittsburgh, PA Curator of Mollusks Santa Barbara Museum of Natural History Delaware Museum of Natural History Santa Barbara, CA Wilmington, DE Andreas Wanninger Richard Petit Chapters 9, 18 Chapter 9 Department of Zoomorphology North Myrtle Beach, SC University of Copenhagen Copenhagen, Denmark Gary Rosenberg Chapter 8 Beatrice Winner Chairman - Department of Malacology Chapter 28 Academy of Natural Sciences North Palm Beach, FL Philadelphia, PA
Patrick D. Reynolds Chapter 19 Biology Department Hamilton College Clinton, NY
xii C. F. Sturm, T. A. Pearce, and A. Valdés. (Eds.) 2006. The Mollusks: A Guide to Their Study, Collection, and Preservation. American Malacological Society.
CHAPTER 1 THE MOLLUSKS: INTRODUCTORY COMMENTS
CHARLES F. STURM TIMOTHY A. PEARCE ÁNGEL VALDÉS
1.1 INTRODUCTION subjected to a good revision in many decades. An advanced amateur could review the world literature Mollusks have been important to humans since our and summarize it into a well-researched revision- earliest days. Initially, when humans were primarily ary work. Though this can be a daunting task, it interested in what they could eat or use, mollusks could be the culmination of years of demanding were important as food, ornaments, and materials and painstaking work. While such a project may for tools. Over the centuries, as human knowledge take years or decades, if done correctly, it will be a branched out and individuals started to study the valuable contribution to practicing malacologists. world around them, mollusks were important sub- jects for learning how things worked. This book is intended for three groups of people. The first and foremost is the amateur community. Initially, nobility and the wealthy (or scientists Amateur malacologists are those who study mol- with wealthy patrons) carried out such studies on lusks out of an avocational desire, in contrast to the the natural world. Later, in the 19th and 20th Cen- professional who is employed or was trained as a turies, a professional class of scientists developed. malacologist or biologist. Some amateurs may be Governments and industry also started supporting classified as paraprofessionals meaning that their scientists. Although in some fields, as professional- depth of knowledge on a particular subject may be at ism developed, non-professionals took a back seat; a professional level, however, they may not have the in contrast, non-professionals studying mollusks breadth of malacological knowledge that a profes- have consistently made important contributions. sional has. We hope that this book will give amateurs the guidance and skills to deepen their interest in Just as bird watchers contribute important ob- malacology and do so in a professional manner. servations that allow professionals to study bird migration and changes in populations, so amateurs Another group of people for whom this book is in- play important roles to professional malacologists. tended is professional biologists, those individuals Amateurs can study the mollusks of a given region whose work, at least in part, relates to mollusks. This over time and see what changes occur in response group would include, among others, malacologists to interactions with humans and natural forces. The and ecologists. The techniques outlined in this book molluscan fauna is inadequately studied in many will be of use to them in their work. If you are a bi- areas; amateurs can study such areas. You may have ologist, and have a need for an up-to-date reference the opportunity to describe a new species. While on mollusks, this book is for you. It will be of use to amateurs may not have the resources for undertak- biologists studying both fossil and recent mollusks. ing molecular studies, they might collaborate with professionals by providing samples. In addition, The final group of people is biology students begin- many molluscan families and genera have not been ning their studies of mollusks. As with the above 2 Introductory comments groups of people, this book will help open up the field • mesodermal origin of pericardioducts of malacology to them. It will provide students with • rhogocytes (pore cells) associated with the ne- a sourcebook that they can use in their studies. phridia (kidneys) • tetraneury (two pairs of main longitudinal nerve The terms conchology and malacology need clari- bundles) fication. Conchology has traditionally been thought • intercrossing dorsoventral muscles of as the study of molluscan shells. Malacology is a • crystalline style and associated ciliated midgut broader term that includes the study of the animal digestive organs that made the shell in addition to the molluscan • esophageal pouches shell. In this book, we use the term malacology to • broad creeping sole or narrow hydrostatic foot mean the study of mollusks and malacologist to • large ventral pedal glands that secrete mucus describe anyone interested in mollusks, whether their interest is only the shell or the molluscan fauna Mollusks will share some or all of these charac- in the broadest sense. teristics. You may be unfamiliar with some of the terms used; many are defined in the glossary found What is a mollusk? Among the animals, Mollusca at the end of this book. are in a group called Spiralia or Protostomia, which also includes Annelids, Arthropods, and other small Mollusks first appeared at the end of the Pre- phyla. The Spiralia, which make up more than 90% Cambrian. Many lineages of mollusks have died of all living multicellular animals, share several out before Recent times. The living mollusks developmental features including spiral cleavage comprise the following classes: Solenogastres of blastomeres, formation of the mouth from the and Caudofoveata (together the Aplacophora), blastopore, and predictable cell fates such as all Monoplacophora, Polyplacophora, Scaphopoda, mesoderm being formed from the single cell 4d. Cephalopoda, Gastropoda, and Bivalvia. Mollusks The circumpharyngeal nerve ring is present in many can be found in terrestrial, freshwater, and marine Spiralia. Within the Spiralia, the group Eutrochozoa environments. includes the Mollusca, Annelida, and other small phyla, which all share a trochophore larva (top As indicated in the title, this book focuses on shaped larva with an equatorial ciliated band and studying, collecting, and preserving mollusks. The a dorsal cilia tuft), schizocoely (formation of body study of mollusks can take place in the field, in an cavities from multiple bilaterally paired masses of aquarium, or in a collection in your home or at a mesoderm), and paired excretory organs and ducts museum. Collection can refer to the collection of that open externally. Regarding the Mollusca them- field observations or specimens. Preservation can selves, despite the wide diversity of body forms, refer to the preservation of specimens in collec- the groups of animals we classify as Mollusca share tions, or to conserving living mollusks and their the following unique features (Eernisse et al. 1992, habitats. Haszprunar 2000): It is also important to know what this volume • radula (absent in Bivalvia) does not cover. It provides only basic information • mantle capable of secreting a calcium carbonate- on anatomy. It does not cover the biochemistry, based shell or spicules physiology, or genetics of mollusks. Though raising • mantle cavity and maintaining mollusks in aquaria or terraria is • ctenidia (specialized gills having countercurrent mentioned in a few chapters, except for Chapter oxygen exchange) 31, there is very little information on aquaculture • osphradia (chemosensory epithelial organs) of mollusks. (absent in Monoplacophora and Scaphopoda) • pericardium around the heart (not around heart Now we give a brief overview of the remainder of in Scaphopoda) this book. This overview should give you an idea Sturm, Pearce, and Valdés 3
of the information to be found in each chapter and Moretzsohn, in Chapter 6, discusses digital imag- help you use this book effectively. ing with flatbed scanners and digital cameras. He begins with a basic discussion of the theory behind 1.2 CHAPTER REVIEWS digital imaging. He goes on to compare digital and film imaging technologies. Digital cameras and flat- The book is composed of 31 chapters. The first bed scanners are discussed. Finally, the printing and 14 chapters cover basic topics in malacology. The editing of digital images are reviewed. Moretzsohn next seventeen chapters cover malacological issues also provides an extensive list of Internet sites related to specific groups of mollusks. where one can go for further information.
Chapter 2 presents many techniques for the collec- Geiger completes the topic of imaging with a re- tion, cleaning, and preservation of mollusks. There view of traditional film photography. In Chapter 7, are also sections about tagging and narcotizing mol- he begins with a discussion of the nature of light lusks. In this chapter, Sturm, Mayhew, and Bales and basic photographic theory. He discusses camera have tried to include not only the latest techniques, equipment and accessories. There is a discussion but also older and still useful methods. The Walker regarding film. Special topics include infrared and Dipper, first described in 1904, is illustrated here ultraviolet photography, underwater photography, for the first time. photography of mollusks in aquaria, and storage of images. Chapter 3, primarily written by Burch, is on remote bottom sampling. It updates his paper presented at In Chapter 8, Rosenberg discusses the use of the annual American Malacological Union meet- computer databases in managing collections. In ing in 1941. Burch relates stories about a lifetime his discussion he includes choosing a software of dredging and provides useful insights on this package, the suggested fields to be included, and activity. In the second part of this chapter, Sturm constructing the database. The discussion is general discusses several other methods of sampling un- enough that it will apply to many different operat- derwater sediments including grabs, box corers, ing systems. tangle nets, and the like. Chapter 9 by Sturm, Petit, Pearce, Cummings, Snorkeling and SCUBA diving are covered by Schwabe, and Wanniger provides an introduction Cordy and Cordy in Chapter 4. This chapter will to the malacological literature. The first part of expose you to new ways of coming face to face with the chapter discusses the various types of malaco- mollusks. While this chapter will not teach you how logical publications: monographs, iconographies, to dive, the authors have covered the equipment that journals, and separates, just to name a few. The is needed for snorkeling and SCUBA diving. The rest of the chapter categorizes over 700 works by inherent risks of these activities are also covered. biogeographic zones and molluscan families. Thus, if you have an unknown shell from Ecuador, under Chapter 5 moves us out of the field and into our col- the Panamic Biogeographic Zone listing, you will lections. If you want to learn how to help preserve find a list of references to help in identifying it. Also your collection for the next millennium, this chapter listed are books of general interest and others for will interest you. This chapter discusses curatorial those interested in taxonomic research. practices and methods for archival protection of your collection. The chapter starts with a discussion How to write a taxonomic paper is an art that is of the risks to stored collections. It continues with a handled by Geiger in Chapter 10. Geiger shows discussion of materials that are used in maintaining the components of scientific papers dealing with collections, stressing archival materials and prac- both the new description of a genus or species and tices. This chapter concludes with a list of sources of a revisionary paper. Geiger also gives a brief for archival materials. overview of the International Code of Zoological 4 Introductory comments
Nomenclature, a set of rules governing the naming with them. Data can be intrinsic (for example color, of species. Familiarity with the code is essential for size, and weight) or extrinsic (for example collected anyone describing a new genus or species; for those where, when, and by whom). The importance of the just interested in cataloging shells, it is useful know- extrinsic data is emphasized, as extrinsic data are ing how names are determined and why a familiar often what makes specimens of vital importance to name is sometimes replaced by a less familiar one. a museum. Also covered in this chapter are ways to preserve soft tissue from molluscan specimens If you are interested in finding out how researchers and how these may be of importance to a museum. determine the tree of life, or how closely related dif- Finally, if you are contemplating donating your ferent species are, Chapter 11, on cladistic analyses collection to a museum, Pearce discusses the way and phylogenetic trees, will interest you. Campbell to do this so that the collection will be of greatest explains cladistics and then covers some of the use to the museum and other researchers. major techniques used in a cladistic analysis. He then concludes with a discussion of molecular With Chapter 15, we will begin exploring differ- biology and how it relates to modern phylogenetic ent molluscan groups, as well as techniques for analyses. rearing mollusks. These chapters are followed by a chapter on the marine biology of non-molluscan If you have ever wondered whether or not to join organisms. a malacological organization or go to a meeting, Chapter 12 will be of use to you. Here Sturm dis- Chapter 15 by Campbell is a discussion of paleo- cusses the basic functions that both amateur and malacology. Here you will learn about some mol- professional malacological associations provide. lusks that exist only as fossils. Campbell provides He describes the functions of such major organi- a general overview of fossils and ways to collect zations as the American Malacological Society, fossil mollusks. Conchologists of America, and others. He discusses the benefits of meetings, both local and national. Chapter 16, by Scheltema, is a review of the Six Internet-based discussion groups are listed with Aplacophora (worm mollusks). These organisms directions on how to join them. comprise a fascinating group of mollusks that, while often under our feet, are rarely collected Museums are one of the places where we can go to by professionals or non-professionals. Scheltema see fossil and Recent mollusks and to learn more discusses the basic biology of these mollusks and about them. In Chapter 13 by Sturm, you will their ecologic context. She goes on to discuss ways learn about the basic roles of museums in keeping of collecting aplacophorans and techniques for collections and conducting research: they are not studying them. just places that exhibit natural history artifacts! In addition, you will take a quick tour of 22 museums Chapter 17 deals with the smallest class of living in the United States and Canada. These museums mollusks, the Monoplacophora. Counts discusses all have major malacological collections. So you this group, which was first known from the fos- can learn more about these institutions and what sil record; the first living monoplacophoran was they have to offer, the addresses for their Internet identified only in 1957. These mollusks are found sites are included. in marine environments and at great depths. Counts covers the biology and zoogeography of these If you ever wondered about museum collections and organisms. If you are fortunate enough to find a what museums do with donated specimens, Chapter monoplacophoran, directions for their preservation 14 by Pearce will inform you. Pearce begins by are covered. discussing what makes a specimen valuable to a museum. Most specimens will have some value if Schwabe and Wanniger write about the Polypla- they have clear and accurate locality data associated cophora (chitons) in Chapter 18. These mollusks Sturm, Pearce, and Valdés 5
have a shell composed of 8 plates and attach tena- how to collect snails with a leaf blower! They con- ciously to the rocks on which they live. Schwabe clude with an exhaustive description of processing and Wanniger discuss the external and internal and storing land snails as well as a discussion of anatomy of these organisms as well as their mode record keeping. of reproduction and where they live. They include techniques for collecting and preserving the soft The discussion of land snails is continued in Chap- tissues and shells of chitons. ter 23. Örstan begins with a review of the literature on maintaining land snails in a terrarium. He goes The Scaphopoda (tusk shells) are addressed in on to discuss short-term maintenance of a terrarium Chapter 19. Here Reynolds begins with a discus- and rearing some North American woodland snails. sion of the biology of the tusk shells and goes on to He concludes with a discussion of factors that can review them in an ecological context. He discusses affect the health of land snails. Hans (see the fron- ways to collect, preserve, and maintain collections tispiece) has been raised in captivity for the past 4 of tusk shells. years following the advice in this chapter.
If you ever wanted to know how to preserve a giant The treatment of the Gastropoda concludes in squid, this next chapter is for you. In Chapter 20, Chapter 24. Here Geiger presents information on Anderson discusses the Cephalopoda (i.e., squids the marine Gastropoda. He begins with an extensive and octopuses). He begins with a review of the discussion of the shell and soft tissue anatomy of biology and behavior of these mobile organisms. the marine gastropods. He continues with a discus- This review is followed by a taxonomic review of sion of the major groups of marine gastropods. He the class. He then goes on to discuss collecting tech- then discusses the various habitats where marine niques and protocols for preserving cephalopods, mollusks can be found and the ecological aspects including Architeuthis, the giant squid. Although of the marine gastropods. He concludes with a you will need hundreds of liters of preservative, an discussion of techniques for collecting and preserv- enormous vat, and weeks to months, this chapter ing marine gastropods, including the preparation will prepare you for the undertaking. Anderson of radulae. concludes the chapter with information on identify- ing cephalopods and a section on the difficulties of The next three chapters cover the Bivalvia. Chapter maintaining cephalopods in an aquarium. 25 by Cummings and Bogan reviews the Unionoida or freshwater mussels. The larvae of these mollusks Chapter 21 reviews the freshwater Gastropoda. spend part of their lives as parasites of vertebrates. While some of these gastropods are vectors of Cummings and Bogan begin with a description of disease, others are common aquarium inhabitants. the biology and ecology of the freshwater mussels. Dillon starts off with a discussion of the biology This section is followed by a taxonomic treatment and ecology of the freshwater gastropods. He also of this group. They briefly touch on issues relating touches on some conservation issues relating to to conservation, a topic more fully developed in this fauna. He describes techniques for collect- Chapter 30. They then provide an extensive descrip- ing, preparing, and storing freshwater gastropods. tion of field collecting equipment and techniques, Dillon concludes with techniques for maintaining information on identifying freshwater mussels, freshwater gastropods in an aquarium. and methods of curating wet and dry collections of these organisms. Pearce and Örstan discuss the terrestrial Gastropoda in Chapter 22. They begin by discussing the biology Korniushin, in Chapter 26, provides a similar treat- of this group. There is an extensive discussion of the ment for the non-unionoid freshwater bivalves. habitats of the land snails and how to locate them. He begins with a treatment of the Sphaeriidae, They continue with a discussion of field methods commonly known as the fingernail, pea, or pill used in collecting land snails; you will even learn clams because of their small size. This includes 6 Introductory comments the biology and ecology of and methods for col- he also discusses the fisheries for abalones, Queen lecting and preserving these organisms. He then Conchs, scallops, clams, mussels, and cephalopods. briefly discusses the Corbiculidae (Asian clams) He includes tables listing gastropods and bivalves and Dreissenidae (zebra mussels). that are covered by state or federal regulations, common edible or commercial mollusks that are The marine Bivalvia are discussed in Chapter 27. not covered by specific regulations, and information Coan and Scott begin with a discussion of the biol- for states with marine fishery agencies. ogy of these organisms. They then give an overview of the five groups (orders) of marine bivalves. They Following Chapter 31 are two appendices and a then discuss techniques for the collection, preserva- glossary. Appendix 1 consists of four plates il- tion, and study of these animals. lustrating morphological and anatomic features of gastropods and bivalves. Appendix 2 is an Chapter 28 deals with establishing and maintaining expanded table of contents. It will give you a de- a marine aquarium. Winner begins the chapter with tailed overview of each chapter and help locating a discussion of the uses of and observations that can information in the book. In the Glossary, you will be made in an aquarium. The physical components find definitions for many of the technical terms used of the aquarium are then discussed: water, plants, throughout this volume. rocks, and food. She then describes fourteen marine organisms that she maintained in her aquaria and This book is meant to be sampled piecemeal and the observations that she made. not necessarily read from cover to cover, though one could do this. The book is meant to be sampled In Chapter 29, Gutierrez gives a brief overview of in portions relating to one’s interests. If you are marine biology. The aim of her chapter is to intro- interested in bivalves, you will concentrate on duce marine organisms that are sometimes mistaken Chapters 25-27. If you wish to delve into freshwater for mollusks. Some examples are calcareous algae, malacology, Chapters 21, 25, and 26 would be on annelids, brachiopods, and echinoderms. your reading list. If you are interested in field tech- niques, read Chapters 2-4, 6-7, and the chapters for The last two chapters cover issues of conservation. the organisms in which you have an interest. One covers freshwater mollusks while the other covers marine mollusks. We hope that you will find the information and dis- cussions in this book helpful. As with all books of In Chapter 30, Bogan reviews the sad conservation this type, compromises had to be made. The authors history of freshwater Gastropoda and Bivalvia in and editors sought to present a balanced handbook North American streams, rivers, and lakes. He of techniques and background information. In explores the patterns of extinction that have oc- preparing each chapter, the authors attempted to be curred and the reasons behind them. On an upbeat comprehensive in approach while presenting only note, he does discuss some reversals in these what they felt would be useful. If there are areas trends. Bogan includes tables listing the threatened, or topics that you feel were slighted or overlooked, endangered, and extinct freshwater mollusks of let us know. North America. 1.3 ACKNOWLEDGMENTS In the final chapter of this book, Chapter 31, Baker discusses marine molluscan fisheries. He explores The senior editor (CFS) owes a debt of gratitude the attitudes towards marine mollusks and marine to many individuals for their involvement with this conservation in the 20th and 21st Centuries. He goes book. The book is partially an outgrowth of a work- on to examine specific examples involving mol- shop at the 1999 American Malacological Society lusks. He extensively reviews the oyster fisheries on (AMS) meeting held in Pittsburgh, Pennsylvania. both coasts of the United States. Less extensively, Opinions as to what topics should be addressed in Sturm, Pearce, and Valdés 7
the workshop were offered by Paul Drez (1947- The American Malacological Society sponsored 2004), William Frank, Ross Mayhew, Timothy this project and provided financial support. Two Pearce, Michael Penziner, Richard Petit, Robert chairs of the AMS Publications Committee have Prezant, Gary Rosenberg, and Evangelos Tzimas. helped us with this book. The project was started Presenters at that workshop included Kevin Cum- under the guidance of Ronald Toll and came to mings, José Leal, Timothy Pearce, Richard Petit, completion under Janice Voltzow’s tenure. Paul and Gary Rosenberg. Robert Prezant deserves spe- Callomon offered valued insight into the intricacies cial thanks, for presenting CFS with the opportunity of publishing books. to organize this workshop. The quality of this book has been greatly enhanced We would like to extend our thanks to all the con- by the assistance of three librarians who helped track tributors to this book: Frank E. Anderson, Patrick down cited literature and provided help in obtaining Baker, B. R. Bales (1876-1946), Arthur E. Bogan, interlibrary loans. This work has benefited greatly from Thomas A. Burch, David Campbell, Eugene V. the assistance of Bernadette Callery, Sun Xianghu, and Coan, Bobbi Cordy, Jim Cordy, Clement L. Counts, Marie Corrado of the Carnegie Museum of Natural III, Kevin S. Cummings, Robert T. Dillon, Jr., History Library. Daniel L. Geiger, Lucia M. Gutierrez, Alexei V. Korniushin (1962-2004), Ross Mayhew, Fabio We would like to thank Emily C. Ullo and Amanda Moretzsohn, Aydin Örstan, Richard Petit, Patrick D. E. Zimmerman for commissioned artwork request- Reynolds, Gary Rosenberg, Amélie H. Scheltema, ed by CFS. Ms. Ullo painted the cover and drew the Enrico Schwabe, Paul Valentich-Scott, Andreas illustrations in Chapter 21. Ms. Zimmerman drew Wanninger, and Beatrice Winner. illustrations in Chapters 2, 3, 9, 11, and 29.
We are thankful for all those individuals who agreed CFS is extremely appreciative of the help given by to review submissions. They are: James Albarano, the two associate editors, Timothy Pearce and Ángel Amir Amiri, Kurt Auffenberg, Glenn Burghardt, Valdés. They joined the project in an editorial capac- Laura Burghardt, Henry Chaney, Eugene Coan, ity in 2004. Their assistance in copy-editing and Louise Corpora, Robert Dillon, Douglas Eernisse, layout helped bring this project to its conclusion. James Fetzner, Daniel Geiger, Jose Juves, Al- bert Koller, Harry Lee, James Lee (1922-2005), Lastly, CFS thanks his wife Pat for her patience Richard Lee, James McLean, Paula Mikkelsen, with and encouragement for this project. Paul Monfils, Mohan Paranjpe, Kristin Petersen, Robert Prezant, John Rawlins, Lutfried von Salvini- 1.4 LITERATURE CITED Plawen, Gerhard Steiner, Patricia Sturm, Michael Vecchione, Janice Voltzow, and Amy Wethington. Eernisse, D. J., J. S. Albert, and F. E. Anderson. 1992. Annelida and Arthropoda are not sister taxa: a phy- In addition to individuals already mentioned, we logenetic analysis of spiralian metazoan morphol- would also like to thank James T. Carlton, Harry ogy. Systematic Biology 41: 305-330. A. ten Hove, and Peter J. Wagner for helping to Haszprunar, G. 2000. Is the Aplacophora monophyletic? resolve specific questions that arose during the A cladistic point of view. American Malacological editing process. Bulletin 15: 115-130.
C. F. Sturm, T. A. Pearce, and A. Valdés. (Eds.) 2006. The Mollusks: A Guide to Their Study, Collection, and Preservation. American Malacological Society.
CHAPTER 2 FIELD AND LABORATORY METHODS IN MALACOLOGY
CHARLES F. STURM ROSS MAYHEW B. R. BALES (1876-1946)
2.1 INTRODUCTION 2.2 COLLECTING BASICS
Your first exposure to the world of mollusks may Other than being given a collection, there are three have been picking up a seashell on a beach while basic ways to build a collection. You can buy speci- you were on vacation. It may have been finding mens, trade for them, or collect them yourself. a snail or slug in your garden. It may have taken many years before you decided to pursue a more 2.2.1 Purchasing. If you chose to purchase shells, in-depth interest in these organisms. This chapter you first have to find a dealer. They can be found will introduce you to ways to collect, clean and advertising in publications such as American Con- prepare mollusks. chologist, La Conchliglia, and Of Sea and Shore. A comprehensive worldwide listing of dealers can Part of this chapter was a paper written by B. R. be found in Tom Rice’s A Sheller’s Directory of Bales and presented for him by R. Tucker Abbott Clubs, Books, Periodicals and Dealers compiled by at the Eleventh Annual Meeting of the American Tom Rice (2003) and at several good Internet sites Malacological Union in 1941 (Bales 1942). This such as
publications. The benefits (and hazards) are the same as with purchasing from dealers. In addition, if you are not trading for a specific shell, you will have the excitement that comes with receiving a shipment of unknown material. As with purchasing from dealers, when you trade with someone, make your first trades modest ones of material that you can afford to lose. If all goes well, mutual trust will develop and larger trades will ensue with confidence. Often, but not always, the material you trade for has been self-col- lected and therefore the collection data may be more complete and reliable than with purchased material.
2.2.3 Self-collecting. In spite of building a collec- tion by purchasing or trading shells, eventually you may desire to start field collecting yourself. You may decide to collect shells in which the animals have died (dead collecting) or living material (live collecting). Section 2.4 deals with some techniques that you can use to collect mollusks.
The simplest way to self-collect is to look for dead material. You can walk along the beach picking Figure 2.1 Collecting devices and aids. up seashells, an activity known as beachcombing. A. Allison Scoop, B. Ferriss Hoe, C. Davis Rake Drag, A particularly good time to try beachcombing is D. Walker Dipper, E. van Eeden Scoop, F. Clam tube or clam gun, G. Bales Hook. after a storm. Sometimes you will find material from deeper water thrown up on the beach. Also, eye, “F++ or Gem-”, which some dealers call “F+++” pelagic material may be blown onto the beach by specimens, are so close to perfect that any flaws have the storm’s winds. to be searched for, “F+” indicates a specimen with minor flaws that do not detract much from its aesthetic You might walk along the shores of a lake, pond, or appeal, “F” (stands for “Fine”) specimens have sig- river and search for freshwater material. Sometimes nificant flaws, Good specimens are not good at all, and you may be lucky and find a shell midden. This is Fair and Poor are recognizable as that species, but just a pile of shells left behind by an animal such as a barely. That said, grading is subjective, and there is an raccoon or muskrat after it has eaten a meal of mus- understandable temptation to over-grade. If you see sels. Section 2.4 describes some of the techniques a dealer’s list with mostly gem designations, beware. of field collecting. Caveat emptor is a useful watch-phrase. When plac- ing an order from a dealer you are unfamiliar with, it You might also be interested in collecting terrestrial is best to start small. Make a few modest purchases, gastropods. These can be found under rocks or leaf and if you agree with the dealer’s grading of the shells litter. Techniques for collecting these mollusks are and the quality of the associated data, you can make covered below and in Chapter 22, Sections 22.4 larger purchases with confidence. Often dealers will and 22.6. learn your interests and notify you of material in which you might be interested. 2.3 COLLECTING EQUIPMENT
2.2.2 Trading. Trading has several rewards. One can Many different tools or implements may come trade shells and/or publications for other shells and/or in handy from time to time, and, while you may Sturm, Mayhew, and Bales 11
achieve a measure of success with very little equip- strainers, these may be constructed or purchased in ment (such as a spade, a kitchen strainer, some vials many sizes and forms. The size of the screen and and collecting jars, rubber boots, snorkeling equip- height of the sides depend upon the individual user. ment, sharp eyes, and plenty of patience), you will It should not be so large that it taxes the strength, eventually find that some specialized tools will be for it should be remembered that to be successful in necessary as you progress in the study of malacol- screening, the collector must be persistent and many ogy (i.e., the study of mollusks as living animals), hours are usually spent in this manner. It is a fascinat- or conchology (i.e., the study of the calcareous ing form of collecting and the time flies all too soon. exoskeletons of mollusks, otherwise known as Seldom will the collector stop sifting without trying “shells”). What follows now are descriptions of “just one more screenful” of material. some such tools. Some collectors prefer several graduated sizes of 2.3.1 Allison scoop. Allison (1942) described a screen (brass, aluminum, stainless steel, or zinc- scoop for collecting Campeloma (a freshwater coated mesh are best, since they are corrosion-re- gastropod) from stream bottoms. The scoop was sistant), but most prefer just two; the inside screen 1 triangular in shape, had a wire basket with a rein- to be 6-8 mm mesh ( /4 inch) and the outer one of 1 forced leading edge and it was attached to a pole 3-4 mm mesh ( /8 inch). If you are interested in (Figure 2.1A). By varying the size of the scoop microshells at all, a third one, of 1 mm or smaller and the mesh size of the basket, this scoop can be mesh (0.04 in or less) will be necessary. This one modified to collect a wide variety of mollusks. will catch most of the smallest shells. The inner screen with the larger mesh should fit snugly into 2.3.2 Ferriss hoe. Walker (1904) described an the outer one with the smaller mesh, but not too implement called the Ferriss hoe. This is a garden snugly. Allowances should be made for the natural hoe with its blade trimmed to 75 mm (3 in) at the swelling of the frames, although much swelling will top and tapering to a sharp point. The handle is be avoided by painting the frames. Some collectors trimmed to the length of a walking cane (Figure nail a small cleat to the ends of the outer screen 2.1B). This device is a good tool for turning over to obtain a firmer grasp. The same results may be logs and rocks, breaking up rotting logs, and dig- obtained by sawing a narrow horizontal slit in the ging through rotting leaves and around stumps. It is end of the frame. The size of frame and mesh being also long enough to pull down tree branches. of individual preference, it is sometimes advisable to try out several before the ideal one is found, and 2.3.3 Davis rake drag. Davis (1964) described a even then, many collectors change from time to type of dredge to be used from the shoreline. He time as the occasion demands. You might also find took a garden rake and to the crossbar of the rake soil sieves useful. These are nested sieves about 1 he attached a 6-8 mm ( /4 in) wire mesh that looped 20 cm (8 inches) in diameter. They are generally around and was attached to the ends of the tines. He made of brass. They can be purchased from general then attached a pipe, filled with sand, to the crossbar scientific and forestry supply companies (see the ap- to give the device added weight (Figure 2.1C). A rope pendix in Chapter 5 for a list of such companies). was attached to the handle of the rake. The device was then thrown out into the water and pulled back One form of screen that is sometimes used has no so that the wire basket and tines would bite into the upright frame on one side and is held in place on the substratum. The rope was pulled drawing the device bottom of a body of water by the collector’s foot. back to shore. The device was then tilted, emptied, The sand, mud, marl, or other material is raked or and the contents examined for specimens. drawn into the screen by the use of a hoe, rake, or other utensil. This type of screen avoids having to 2.3.4 Screens, dippers, and nets. To collect the lift the material and to deposit it in the screen for smaller specimens from shallow water, a screen those who like to adhere to a less energetic regimen comes in handy. Once you progress beyond kitchen while collecting. 12 Field and laboratory methods in Malacology
A handy device often used consists of a small round (3-6 inches) and 1 to 1.5 meters long (3-4 feet). It is sieve that has been attached to a long handle. It is closed on one end with a cap that has a handle and easily made from a 12-15 cm (5 or 6 inches) gravy small vent hole (Figure 2.1F). When you locate a strainer to be had at most hardware or discount clam burrow, place the tube over the burrow. Rock stores. They have two bent prongs in front that must the tube back and forth and twist it so that it drills be bent backward so as not to interfere with the use of into the sediment. When you believe that the shell the net. No device equals this when working in waist is within the tube, cover the vent hole. Doing so or chest deep water. You might also want to make creates a suction effect when you withdraw the and use a Walker Dipper (see Figure 2.1D, Chapter tube from the ground. You will pull up a plug of 21, Freshwater Gastropoda, and Walker 1904). sediment that should include the clam. Empty the contents onto a screen, wash them, and remove When collecting the specimens on various types the shell. This device works best in muddy and of aquatic grasses (eel, turtle, etc.), a net made of sandy substrates; it does not work well in bottoms mosquito or other fine netting sewn around a but- composed of gravel and rock. terfly net hoop is extremely useful, especially if you wish to be as ecologically sensitive as possible, 2.3.8 Hammer. A geologist’s or bricklayer’s ham- leaving the grasses in place instead of collecting mer with a chisel end can be quite useful in several them and washing the mollusks into fine (the 24- respects. You can scrape through leaf litter, dig into mesh) screens using fresh water. the upper layers of soil, turn over rocks (which of course should always be replaced as they were, 2.3.5 van Eeden scoop. Another variation of scoop before leaving the scene), use it in the pursuit of was described in van Eeden (1960) (Figure 2.1E). paleomalacology (fossil collecting), chip away The scoop was designed for collecting freshwater the soft rocks in which burrowing species such gastropods. It is a square frame made from a 5 mm as many of the family Pholadidae (e.g., Barnea iron rod. The square is 25 cm on a side. The frame and Zirfaea spp.) live, and the chisel end comes in is angled upwards about 30 degrees two thirds of handy when taking apart pieces of sunken wood the way back from the leading edge. The leading that may harbor ship worms (Families Teredidae edge is reinforced with iron or tin sheeting. A wire and Xylophagidae). It is useful to paint part of screen of appropriate mesh size is attached to the these tools a bright yellow or orange. Otherwise, frame. The frame in turn is attached to a handle of you will learn the hard way how easily such tools appropriate length (2-3 m). can be overlooked when put down in the field and searched for at a later time. 2.3.6 Shovels. Some collectors use a shovel to dig up clams. Commonly, these shovels have a blade 2.3.9 Bales hook. According to Bales, one of the 75-100 mm wide (3-4 inches). The blade is longer most important tools is a device made from a 15 5 than it is wide, and it is slightly curved. These mm ( /8 inch) metal rod. One end is looped to make shovels are sometimes called clam guns. Insert a handle and the other is formed into a point. About the shovel so that it curves away from where you 75 or 100 mm (3 or 4 inches) above the point, a believe the clam to be and start digging. Dig and curved hook is welded onto the rod giving a form scoop the sediment away from the clam burrow; similar to an elephant hook (Figure 2.1G). With keeping the shovel blade parallel to the clam bur- such a device, you have an implement that can be row. When you see part of the clam reach down used as a walking stick, to turn over rocks, to pull and grab it. It is important not to dig towards the things closer to you, and to pull down branches clam; doing so may damage it. when looking for arboreal snails. Estwing makes a similar tool called the Gem Scoop®. Instead of a 2.3.7 Clam tube. There is another device also hook, it has a small basket. Of course, many collec- called a clam gun or sometimes a clam tube. This tors make their own tools as best as they can, given is a metal or plastic tube, 75-150 mm in diameter their budget and circumstances, so the implements Sturm, Mayhew, and Bales 13
described in this chapter can be taken as a starting to mix things up and difficult to take detailed notes, point - an assemblage of ideas and advice, for you but care nevertheless should be taken to keep facts to adopt or adapt as you see fit. and specimens straight. Any details that you note regarding the habitat can be jotted down afterwards 2.3.10 Water pumps. When the ocean floor is of while they are fresh in the memory. rock that is more or less honeycombed with small potholes, it is surprising what fine specimens of the Freshly taken specimens should never be placed, smaller varieties may be obtained by the use of a even temporarily, in a rusty metal container or in common bilge pump, which is standard equipment contact with rusty chains or other rusty objects for on small boats. The end of the pump is placed in it is remarkable how soon they will become rust a sand-filled pocket in the rock. The sand as well stained. It is almost impossible to remove these as the mollusks that have taken refuge in the hole stains without damaging the shell in the process. are pumped with the water onto a screen; the sand Sturdy plastic buckets serve the same purpose at and water flow through, leaving the specimens all a very reasonable cost, and are easier on fragile ready for the collector. species than metal ones.
2.3.11 Bags and collecting containers. Very impor- 2.3.12 Glass-bottomed bucket. A useful adjunct tant to the collector’s outfit is your collecting bag, to shallow water collecting is the glass-bottomed and this may consist of almost anything from an old bucket (first used by sponge collectors), or an tin can, a burlap or nylon sack, a pocket handkerchief equivalent device - sometimes called a water glass or or some such makeshift affair, to a real game bag or water bucket. Collectors often use one that is square collecting bag. They are usually made from light- or oblong. An easy way to construct one is to make weight canvas and carried in some cases by a strap the frame or box of not too heavy wood and fasten over the shoulder. Such bags are useful when extra the glass or plexiglas to the open bottom by means heavy specimens are anticipated. A bag that may of quarter rounds available at any lumberyard. A be tied or secured about the waist is much handier bead of silicone caulking is placed around the open- and has the advantage of always being in place and ing between the wood and plexiglas. This ensures does not drop in front of the collector when he or a watertight seal. Visualization will be improved if she stoops to secure a specimen. Many collectors the inside is painted dull black. You can also take a favor a bag containing partitions: one compartment plastic or metal bucket and cut out the bottom leaving 1 for tools, vials and other equipment apart from the a 10 mm ( /2 inch) rim. To this rim, a circular piece shells that are collected, and others for samples of of Plexiglas can be attached with silicone caulking. various sorts. You might even find that a carpenter’s When in use, frequent wetting of the inside of the apron makes a useful collecting bag. glass makes vision clearer and eliminates fogging.
When collecting small mollusks, a good supply of 2.3.13 Lights for night collecting. If you have glass or plastic vials of various sizes, will prove never collected at night, hunting by the aid of ar- most useful. These should be cylindrical and of tificial light is a revelation. Many mollusks (such the screw top sort without a narrow neck. Nylon as Conus) hide wherever they can in the daytime mesh bags are often useful when collecting larger and mainly venture out to feed at night. This is also shells such as bivalves. You should be careful to true of Marginella, Cypraea, Hydatina, and the like. keep shells from different habitats separate, label- While specimens may be taken by the combined ing the vials and containers in the field using pencil use of water glass (see above) and flashlight, you and strong paper, and/or a grease pencil applied might also try snorkeling or SCUBA (see below to the container. Basic information will include and Chapter 4, Snorkeling and SCUBA Diving). date, locality, collector, and as a good and brief a Use a bright waterproof flashlight and make cer- description of the habitat as possible. In the heat of tain that you have spare batteries. Extremely good collecting, especially when racing the tide, it is easy underwater lights can be purchased for less than