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Vertical Distribution of Pelagic Cephalopods * * Vertical Distribution of Pelagic Cephalopods CLYDE F. E. ROPER and RICHARD E. YOUNG SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 209 SERIAL PUBLICATIONS OF THE SMITHSONIAN INSTITUTION The emphasis upon publications as a means of diffusing knowledge was expressed by the first Secretary of the Smithsonian Institution. In his formal plan for the Insti- tution, Joseph Henry articulated 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 keynote of basic research has been adhered to over the years in the issuance of thousands of titles in serial publications under the Smithsonian imprint, com- mencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Annals of Flight Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Studies in History and Technology In these series, the Institution publishes original articles and monographs dealing with the research and collections of its several museums and offices and of professional colleagues at other institutions of learning. These papers report newly acquired facts, synoptic interpretations of data, or original theory in specialized fields. These pub- lications are distributed by mailing lists to libraries, laboratories, and other interested institutions and specialists throughout the world. Individual copies may be obtained from the Smithsonian Institution Press as long as stocks are available. S. DILLON RIPLEY Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 209 Vertical Distribution of Pelagic Cephalopds Clyde F. E. Roper and Richard E. Young SMITHSONIAN INSTITUTION PRESS City of Washington 1975 ABSTRACT Roper, Clyde F. E., and Richard E. Young. Vertical Distribution of Pelagic Cephalopods. Smithsonian Contributions to Zoology, number 209, 51 pages, 31 figures, 1975.—The vertical distributions of pelagic cephalopods are analyzed, based primarily on studies undertaken in the waters off California, Bermuda, and Hawaii. Much of the information is derived from midwater trawl nets equipped with closing apparatus. Each family that contains pelagic or quasi- pelagic species is discussed; selected records from the literature are included where necessary. A wide variety of patterns of vertical distribution exists among pelagic cephalo- pods; patterns may be associated with stage of maturity, diel rhythms, or seasonal behavior. The study revealed that several basic types of vertical distributional patterns occur: near-surface dwellers, first order diel vertical migrators, second order diel vertical migrators, diel vertical shifters, diel vertical spreaders, non- migrators, vertical wanderers, species associated with the ocean bottom, species exhibiting ontogenetic descent. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 5389. SERIES COVER DESIGN: The coral Montastrea cavemosa (Linnaeus). Library of Congress Cataloging in Publication Data Roper, Clyde F. E. 1937- Vertical distribution of pelagic Cephalopods. (Smithsonian contributions to zoology; no. 209) Bibliography: p. Includes index. 1. Cephalopoda—North Pacific Ocean. 2. Cephalopoda—Bermuda Islands. I. Young, Richard E., joint author. II. Title. III. Series: Smithsonian Institution. Smithsonian contributions to zoology; no. 209. QL1.S54 no. 209 [QL430.2] 591'.08s [594'.5'092] 75-6936 Contents Page Introduction 1 Materials and Methods 3 California Study 3 Bermuda Study 5 Hawaii Study 6 Subclass Nautiloidea 6 Nautilidae 6 Subclass Coleoidea 6 Order Sepioidea 6 Spirulidae 6 Sepiidae 7 Sepiadariidae, Sepiolidae, Idiosepiidae 7 Order Teuthoidea 8 Suborder Myopsida 8 Loliginidae 8 Pickfordiateuthidae 9 Suborder Oegopsida 9 Lycoteuthidae 9 Enoploteuthidae 9 Ommastrephidae 13 Histioteuthidae 14 Psychroteuthidae 15 Neoteuthidae 15 Architeuthidae 16 Bathyteuthidae 16 Gonatidae 17 Octopoteuthidae 21 Onychoteuthidae 22 Thysanoteuthidae 23 Cycloteuthidae 23 Ctenopterygidae 23 Brachioteuthidae 23 Lepidoteuthidae 24 Joubiniteuthidae 24 Chiroteuthidae 24 Mastigoteuthidae 27 Promachoteuthidae 28 Batoteuthidae 28 Grimalditeuthidae 28 Cranchiidae 28 iv SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Page Order Vampyromorpha 32 Vampyroteuthidae 32 Order Octopoda 33 Suborder Cirrata 33 Cirroteuthidae and Stauroteuthidae 33 Opisthoteuthidae 34 Suborder Incirrata 35 Bolitaenidae 35 Vitreledonellidae 35 Amphitretidae 36 Allopsidae 36 Tremoctopodidae 36 Ocythoidae 36 Argonautidae 36 Discussion 37 Literature Cited . 45 Index 49 Vertical Distribution of Pelagic Cephalopds Clyde F. E. Roper and Richard E. Young Introduction zones that are uninhabited by adults. Therefore, because distributional patterns often differ Information concerning the vertical distribution markedly between growth stages, it is necessary to of pelagic cephalopods is relatively sparse. Several separate and distinguish between depth records of factors are primarily responsible: (1) Oceanic larval and juvenile-adult populations. Seasonal or cephalopods are often large, fast-swimming ani- daily differences in depth distributions also may mals that are able to avoid being captured by occur; for example, some species migrate to near- conventional sampling gear. (2) Until recently, shore waters in the summer to feed and/or spawn, most specimens were taken in open nets that, in then migrate into deeper, offshore waters during addition to the desired sampling depth, sample the winter; many species in the day inhabit depths that entire water column during setting and retrieval; differ from their nighttime depths. Because of as a result the true depth of capture is difficult to these factors, it is extremely difficult to draw con- define precisely. (3) In many groups only larvae clusions from most of the literature. and juveniles are captured and adults are rare or A factor that has led to considerable confusion unknown. (4) Most sampling has been conducted in the literature is the inability of many workers along geographical transects where a given local- to recognize the phenomenon of diel vertical mi- ity is generally sampled only once, at one time of gration in which animals ascend toward the surface day and at one season of the year. Compounding zone at night and descend to deeper water during these sampling problems are the complexities of the day. Although vertical migrations are well the distributional patterns of the cephalopods. documented in several groups of pelagic animals, To understand the vertical distributions of evidence of this phenomenon in cephalopods has cephalopods (as indeed of many oceanic organ- been inconclusive until recently. The first indica- isms) a knowledge is required of more than tion of vertical movements of cephalopods was merely the bathymetric range, with carefully de- given in 1841 by d'Orbigny who noted that cepha- fined upper and lower limits, over which speci- lopods occurred at the surface at night but were mens of a species are known to occur. Often the absent during the day (vide G. L. Voss, 1967:511). bathymetric range will extend over a great vertical Similar observations have been recorded since that distance, but the vast majority of a population will time. For instance, Sasaki (1914:92) stated that occupy only a restricted depth zone. In addition, Watasenia scintillans comes to the surface at night, larvae of a species may inhabit one or more depth where it may be captured by fishermen, and dis- appears to greater depths during the day. More Clyde F. E. Roper, Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smithsonian Institution, recently Pearcy (1965) presented trawling data to Washington, D.C. 20560. Richard E. Young, Department of suggest that some oceanic species show greater fre- Oceanography, University of Hawaii, Honolulu, Hawaii 96822. quencies of occurrence in the surface layer SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY (0-200 m) during night than during the day, while about which we have no knowledge of the circum- in the layer of 0-500 m abundance of these species stances related to the capture. is greater during the day than during the night. The vertical distribution of cephalopods is sur- Clarke (1969) compared day-night captures using veyed by discussing representative species in each several different types of sampling gear, among family. Knowledge is insufficient to allow a presen- them a closing-net, during the SOND cruises in the tation based upon recognized distributional pat- eastern tropical Atlantic. His results indicate more terns or vertical faunal zones in the open ocean. clearly the existence of vertical migration. The systematic approach allows us to present data Our studies presented here confirm the occur- wherever they exist without having to force them rence of vertical migration in cephalopods and into distributional concepts. Hopefully, within the indicate that this phenomenon is widespread in next few years it will be possible to discuss vertical oceanic squid. These conclusions have relied most distribution of cephalopods in relation to the phys- heavily on sampling programs conducted in three ical and biological parameters
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