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Larval" and Juvenile Cephalopods: a Manual for Their Identification

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Larval * Larval" and Juvenile Cephalopods: A Manual for Their Identification MICHAEL J. SWEENEY, CLYDE F.E. ROPER, KATHARINA M. MANGOLD, MALCOLM R. CLARKE, and P^ MfcSIGURD V. BOLETZKY 3J EDITORS i SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 513 SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of "diffusing knowledge" was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to the Marine Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Folklife Studies Smithsonian Studies in Air and Space Smithsonian Studies in History and Technology In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world of science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. Robert McC. Adams Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 513 'Larval" and Juvenile Cephalopods: A Manual for Their Identification Michael J. Sweeney, Clyde F.E. Roper, Katharina M. Mangold, Malcolm R. Clarke, and Sigurd v. Boletzky EDITORS ISSUED FEB 1 0 1992 SMITHSUMAN INSriUJTION SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1992 ABSTRACT Sweeney, Michael J., Clyde F.E. Roper, Katharina M. Mangold, Malcolm R. Clarke, and Sigurd v. Boletzky, editors. "Larval" and Juvenile Cephalopods: A Manual for Their Identification. Smithsonian Contributions to Zoology, number 513,282 pages, 277 figures, 6 tables, 1992. This manual provides a practical guide to the identification of "larval" and juvenile cephalopods. A comprehensive glossary of morphological terms is presented, as is a general pictorial key to the young stages of cephalopods, primarily to the family level. Systematically organized by order and family, the manual presents characters for adult and young forms, keys, illustrations, distributional and biological information, and references to pertinent literature. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus). Library of Congress Cataloging-in-Publication Data "Larval" and juvenile cephalopods : a manual for their identification / edited by Michael J. Sweeney ... [et al.]. p. cm.—(Smithsonian contributions to zoology : no. 513) "This manual represents the product of a workshop organized by the Cephalopod International Advisory Council (CIAC) held at Laboratoire Arago, Banyuls-sur-Mer, France, 17-28 June 1985"—Introd. Includes bibliographical references. 1. Cephalopoda—Identification. 2. Cephalopoda—Larvae—Identification. I. Sweeney, Michael J., 1945- . II. Cephalopod International Advisory Council. III. Series. QL1.S54 no. 513 [QL430.2] 591 s-dc20 [594'.5] 91-22098 ® The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48—1984. Contents Page Preface vii Introduction 1 Glossary of Terms 3 Provisional Key to Young Stages of Cephalopods 11 Family Accounts 21 SEPIOIDEA (by C.C. Lu, A. Guerra, F. Palumbo, and W.C. Summers) 21 Key to Families of SEPTOIDEA 21 SPIRULIDAE 21 SEPIIDAE 21 Key to Genera of SEPIIDAE 21 SEPIADARIIDAE 27 Key to Genera of SEPIADARIIDAE 27 SEPIOLIDAE 27 Key to Subfamilies of SEPIOLIDAE 27 HETEROTEUTHINAE 27 Key to Genera of HETEROTEUTHINAE 27 ROSSIINAE 27 Key to Genera of RossnNAE 27 SEPIOLINAE 27 Key to Genera of SEPIOLINAE 31 IDIOSEPIIDAE 31 TEUTHOIDEA 37 MYOPSIDA (by R.T. Hanlon, S. v. Boletzky, T. Okutani, G. Perez-Gandaras, P. Sanchez, C. Sousa-Reis, and M. Vecchione) 37 PlCKFORDIATEUTHIDAE 37 LOLIGINIDAE 37 Key to Genera of LOLIGINIDAE (Adults and Subadults) 37 Western North Atlantic LOLIGINIDAE 40 Eastern North Atlantic and Mediterranean LOLIGINIDAE 40 Eastern North Pacific and Hawaiian LOLIGINIDAE 47 Western Pacific and Japanese LOLIGINIDAE 47 OEGOPSIDA 55 The Enoploteuthid Group of Families (by R.E. Young, K.M. Mangold, and M. Vecchione) 55 Key to Families and Genera of the Enoploteuthid Group (Young) 55 ENOPLOTEUTHIDAE 55 ANCISTROCHEIRIDAE 57 PYROTEUTHIDAE 57 LYCOTEUTHIDAE (by GJL Voss and S.J. Stephen) 67 Key to Subfamilies, Genera, and Species of LYCOTEUTHIDAE (Adults) 67 LAMPADIOTEUTHINAE 67 LYCOTEUTHINAE 67 Key to Species of LYCOTEUTHINAE 67 HISTIOTEUTHIDAE (by N.A. Voss, S.J. Stephen, and Zh. Dong) 73 in iv SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Key to Species and Subspecies of HISTIOTEUTHIDAE (Adults, Subadults, and Juveniles) 73 BATHYTEUTHIDAE (by C.F.E. Roper and MJ. Sweeney) 93 Key to Species of Bathyteuthis (Adults and Juveniles) 94 PSYCHROTEUTHIDAE (by CJ\E. Roper and MJ. Sweeney 95 ARCHITEUTHIDAE (by C.F.E. Roper) 99 NEOTEUTHIDAE (by C.F.E. Roper) 101 Key to Genera of NEOTEUTHIDAE (Adults and Juveniles) 101 OMMASTREPHIDAE (by J.H. Wormuth, R.K. O'Dor, N. Balch, M.C. Dunning, E.C. Forch, RP. Harman, and T.W. Rowell) 105 Key to Subfamilies and Genera of OMMASTREPHIDAE (Adults and Juveniles) 105 ILLJCINAE 109 TODARODINAE 109 OMMASTREPHINAE 112 THYSANOTEUTHIDAE (by SJ. Stephen) 121 CTENOPTERYGIDAE (by K. Jefferts) 125 ONYCHOTEUTHIDAE (by M.R. Clarke) 127 Key to the Genera and Most Species of ONYCHOTEUTHIDAE 127 GONATIDAE (by T. Okutani and M.R. Clarke) 139 Key to Genera of GONATIDAE 139 Key to the North Pacific Species of GONATIDAE 139 Key I (Stage with no tentacular club developed) 139 Key II (Stage with tentacular club with minute sucker buds).... 142 Key III (Stage with developed tentacular club) 142 BRACHIOTEUTHIDAE (by C.F.E. Roper and MJ. Sweeney) 157 CYCLOTEUTHIDAE (by K. Jefferts and C.F.E. Roper) 161 Key to Genera of CYCLOTEUTHIDAE (Young) 161 OCTOPOTEUTHIDAE (by S J. Stephen and K. Jefferts) 165 Key to Genera of OCTOPOTEUTHIDAE (Juveniles) 166 LEPIDOTEUTHIDAE (by MJ*. Clarke) 167 PHOLIDOTEUTHIDAE (by M.R. Clarke) 168 BATOTEUTHIDAE (by Ci\E. Roper and MJ. Sweeney) 171 CHIROTEUTHIDAE (by CJ\E. Roper and MJ. Sweeney) 171 Key to Genera of CHIROTEUTHIDAE (Adults, Subadults, and Late Juveniles) 171 Key to Genera of CHIROTEUTHIDAE ("Larvae") 175 MASTIGOTEUTHIDAE (by C.F.E. Roper and MJ. Sweeney) 175 Key to Genera of MASTIGOTEUTHIDAE (Adults, Subadults, and Juveniles) 175 JOUBINTTEUTHIDAE (by C.F.E. Roper and M.R. Clarke) 179 GRIMALDITEUTHIDAE (by M.R. Clarke) 181 PROMACHOTEUTHIDAE (by N.A. Voss) 183 Key to Species of PROMACHOTEUTHIDAE (Juveniles -10-17 mm ML) 183 CRANCHIIDAE (by N.A. Voss, S J. Stephen, and Zh. Dong) 187 Key to Genera of CRANCHIIDAE ("Larvae" and Early Juveniles)... 187 CRANCHIINAE 188 TAONIINAE 192 VAMPYROMORPHA (by F.G. Hochberg and M. Nixon) 211 VAMPYROTEUTHIDAE 211 OCTOPODA (by F.G. Hochberg, M. Nixon, and R.B. Toll) 213 ClRRATA 213 ClRROTEUTHIDAE 213 NUMBER 513 STAUROTEUTHIDAE 217 OPISTHOTEUTHIDAE 217 INCIRRATA 219 AMPHITRETIDAE 219 BOLITAENIDAE 222 VLTRELEDONELLIDAE 225 ALLOPOSIDAE 227 TREMOCTOPODIDAE 228 OCYTHOIDAE 230 ARGONAUTTDAE 232 OCTOPODIDAE 237 Unidentified or Unverified Material 268 Appendix 1: Octopod Worksheet 280 Appendix 2: Workshop Participants 281 This volume is dedicated with gratitude and honor to Professor Gilbert L. Voss (1918-1989), a leading figure in cephalopod systematics and biology for nearly 40 years. In the early 1950s Dr. Voss intended to conduct research on larval and juvenile cephalopods of the Florida Current, but he was forced to abandon the project because even the adults from the Gulf of Mexico and the Caribbean Sea were mostly unknown. This was a world-wide problem and only by the mid 1980s was faunal knowledge sufficiently improved to attempt a compilation about larval cephalopods on a world-wide basis. This volume, in no small part, represents the tremendous contributions Gil Voss has made to cephalopod systematics and distribution. Preface This manual results from an intensive international workshop and subsequent efforts by most participants. The following summary (adapted from Roper, 1985:94, 95) provides readers with the historical perspective and organizational context from which this collaborative volume emerged. The concept of establishing an international body of cephalopod specialists to assess and encourage the direction and needs of cephalopod research was introduced and accepted at the International Workshop on the Biology and Resource Potential of Cephalopods,
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