V.L. KflSVflNOV>G.fl. KRVUCHKOVfl VAKUUKOVfl-LAIVICDVCDevn Scientific Cditor Dovid L pQiuson LARVAE OF MARINE BIVALVES AND ECHINODERMS V.L. KASYANOV, G.A. KRYUCHKOVA, V.A. KULIKOVA AND L. A. MEDVEDEVA Scientific Editor David L. Pawson SMITHSONIAN INSTITUTION LIBRARIES Washington, D.C. 1998 Smin B87-101 Lichinki morskikh dvustvorchatykh moUyuskov i iglokozhikh Akademiya Nauk SSSR Dal'nevostochnyi Nauchnyi Tsentr Institut Biologii Morya Nauka Publishers, Moscow, 1983 (Revised 1990) Translated from the Russian © 1998, Oxonian Press Pvt. Ltd., New Delhi Library of Congress Cataloging-in-Publication Data Lichinki morskikh dvustvorchatykh moUiuskov i iglokozhikh. English Larvae of marine bivalves and echinodermsA^.L. Kasyanov . [et al.]; scientific editor David L. Pawson. p. cm. Includes bibliographical references. 1. Bivalvia — Larvae — Classification. 2. Echinodermata — Larvae — Classification. 3. Mollusks — Larvae — Classification. 4. Bivalvia — Lar- vae. 5. Echinodermata — Larvae. 6. Mollusks — Larvae. I. Kas'ianov, V.L. II. Pawson, David L. (David Leo), 1938-III. Title. QL430.6.L5313 1997 96-49571 594'.4139'0916454 — dc21 CIP Translated and published under an agreement, for the Smithsonian Institution Libraries, Washington, D.C., by Amerind Publishing Co. Pvt. Ltd., 66 Janpath, New Delhi 110001 Printed at Baba Barkha Nath Printers, 26/7, Najafgarh Road Industrial Area, NewDellii-110 015. UDC 591.3 This book describes larvae of bivalves and echinoderms, living in the Sea of Japan, which are or may be economically important, and where adult forms are dominant in benthic communities. Descriptions of 18 species of bivalves and 10 species of echinoderms are given, and keys are provided for the iden- tification of planktotrophic larvae of bivalves and echinoderms to the family level. Information on identified species is preceded by a description of the morphology, physiology, and behavior of larvae of the given classes. The book is addressed to marine biologists, embryologists, zoologists, and specialists engaged in fisheries. FOREWORD TO THE ENGLISH EDITION The Smithsonian Institution Libraries, in cooperation with the National Science Foundation, has sponsored the translation into English of this and hundreds of other scientific and scholarly studies since 1960. The program, funded with Special Foreign Currency under the provisions of Public Law 480, represents an investment in the dissemination of knowledge to which the Smithsonian Institution is dedicated. In this volume, the authors review and summarize many aspects of the early life history of the marine bivalve mollusks and echinoderms that are common in the Sea of Japan. Several of the species covered are of commercial value, and the scientific information given here will be very useful in planning successful exploitation and management. Many of these species have a wide distribution in the Northern Hemisphere, and thus the book is of broad regional interest. For many readers, this English- language translation presents for the first time a review of Russian research on mollusks and echinoderms that would be otherwise inaccessible. It updates a 1980 book (in Russian) by V.L. Kasyanov, L.A. Medvedeva, Y.M. Yakovlev and S.N. Yakovlev, and it includes relevant literature references up to, and including, the year 1990. The editors have taken great care to ensure that this translation accurately reflects the original text. However, some scientific names and parts of the classification, especially in the Echinodermata section, have been changed to reflect current thinking, and to make the data more readily accessible. This work is a significant addition to the literature on embryology and larval development of mollusks and echinoderms, and it should prove to be important and useful to fishery biologists, embryologists and marine biologists. DAVID L. PAWSON National Museum of Natural History Smithsonian Institution Washington, D.C. CONTENTS FOREWORD TO THE ENGLISH EDITION v INTRODUCTION 1 CHAPTER I. LARVAE OF MARINE BIVALVES (MORPHOLOGY, 3 PHYSIOLOGY AND BEHAVIOR) Identification of Pelagic Larvae of Bivalves 41 Pacific Mussel, Mytilus trossulus Gould 74 Crenomytilus grayanus (Diinker) 75 Long-bristled Modiolus, Modiolus difficilis 11 (Kuroda and Habe) Musculista senhousia (Benson) 77 Crescent-shaped Adula, Adula falcatoides Habe 78 Boucard's Area, Area boucardi Jousseaume 80 Giant Oyster, Crassostrea gigas Thiinberg 80 Chlamys farreri nipponensis Kuroda 81 Swiftopecten swifti (Bernard) 84 Mizuhopecten yessoensis (Jay) 84 Hiatella arctica Linne 88 Kellia japonica Pilsbry 90 Keenocardiitm californiense (Deshayes) 90 Baltic Macoma, Macoma balthica (Linne) 91 Tall Siliqua, Siliqua alta (Broderip and Sowerby) 91 Razor Clam, Solen krusenstemi Schrenck 92 Chinese Mactra, Mactra chinensis Philippi 92 Spisula sachalinensis (Schrenk) 93 Philippine Ruditapes, Ruditapes philippinarum (Adams 97 and Reeve) 1 Vlll Mya japonica Jay 97 Truncate Mya, Mya tnincata (Linnaeus) 98 Shipworm. Teredo navalis Linne 99 Bankia setacea (Tyron) 101 Crisj^ate Zirphaea, Zirfaea crispata (Linne) 102 Japanese Bamea, Barnea japonica (Yokoyama) 103 CHAPTER II. LARVAE OF SEA STARS (MORPHOLOGY, 104 PHYSIOLOGY, BEHAVIOR) Identification of Pelagic Larvae of Sea Stars 127 Patiria pectinifera (Muller and Troschel) 132 Asterias amurensis Liitken 134 CHAPTER III. LARVAE OF SEA URCHINS (MORPHOLOGY, 138 PHYSIOLOGY, AND BEHAVIOR) Identification of Pelagic Larvae of Sea Urchins 170 Strongylocentrotus nudus Agassiz 177 Strongylocentrotus intermedins (Agassiz) - 179 Echinarachinus parma (Lamarck) 182 Scaphechinus mirabilis (Agassiz) 185 Scaphechinus griseus (Mortensen) 187 Echinocardium cordatum (Pennant) 190 CHAPTER IV. LARVAE OF BRITTLE STARS (MORPHOLOGY, 195 PHYSIOLOGY, AND BEHAVIOR) Identification of Pelagic Larvae of Brittle Stars 216 Ophiura sarsi Liitken 221 Amphipholis kochii Liitken 223 CHAPTER V. LARVAE OF SEA CUCUMBERS (MORPHOLOGY, 225 PHYSIOLOGY, AND BEHAVIOR) Identification of Pelagic Larvae of Sea Cucumbers 244 Far Eastern Trepang, Stichopus japonicus Selenka 246 Japanese Cucumaria, Cucumaria japonica Semper 25 Fraudulent Eupentacta, Eupentacta fraudatrix 253 (Djakonov and Baranova) CONCLUSION 255 REFERENCES 258 8 INTRODUCTION The life cycles of commercially exploited and cultured mollusks and echino- derms comprise, in most cases, a prolonged benthic period and a brief pelagic period. In the benthic period the animal feeds, grows, attains sexual maturity, produces gametes, and participates in reproduction. The prodijction of gametes (gametogenesis) and their subsequent release and fertilization by mollusks and echinoderms inhabiting temperate waters usually occurs every year. The an- nual reproductive processes in bivalves and echinoderms inhabiting the Sea of Japan were described earlier by researchers in the Laboratory of Embryology, Institute of Marine Biology [Razmnozhenie iglokozhikh i dvustvorchatykh mollyuskov (Reproduction of Echinoderms and Bivalves), Kasyanov et al., 1980]. The present book is a continuation of the 1980 publication. Here we describe the larvae of bivalves, sea stars, sea urchins, brittle stars, and sea cucumbers of the Sea of Japan, which are economically important or abundant forms in benthic communities. The larvae of these species are planktonic, feed on phytoplankton, grow, and are carried out to sea by currents. Then they migrate froni the surface to the bottom, settle there, and metamorphose, thus completing the pelagic period of their life cycle. This period is less known and understood than the benthic, even though many papers and books deal with the subject. Among such works, mention should be made of those by Thorson (1936, 1946, 1950), Mortensen( 1921, 1937, 1938), Mileikovskii (1977, 1981), and Jagersten (1972). The structure of the larvae discussed here have been examined by O.M. Ivanova-Kazas in her two-volume work on the embryology of mollusks and echinoderms (Ivanova-Kazas, 1977, 1978). Particular attention is paid in the present book to those characters which make possible the determination of the species affinities of larvae. Thus 1 species of bivalves and 10 species of echinoderms are described in detail, and keys are provided to enable identification to family level of planktotrophic larvae of bivalves and echinoderms from Peter the Great Bay. The descriptions of the larvae are preceded by descriptions of the morphology, physiology, and — behavior of the larvae of individual classes, based on published and original material. Larvae are examined not so much from an embryological point of view (as a developmental stage of a definitive animal), but from a zoological viewpoint (as an organism endowed with all the systems required for a viable existence in the plankton). The general description of the larvae studied begins with a brief report on early development (up to the stage of trochophore or dipleurula). Cytoembryological aspects of early development are not described because extensive literature is available on this topic. The description con- cludes with a summary of the process of metamorphosis and a brief charac- terization of lecithotrophic larvae. Ecological aspects of the pelagic life of larvae and their settling to the bottom are not described due to space con- straints of this book. Plankton samples were collected mainly in Vostok Bay, which is situated in the eastern part of Peter the Great Bay of the Sea of Japan. To obtain the required stage of larval development, artificial fertilization with subsequent culturing of larvae was done, in addition to