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United States National Museum Bulletin 291 SYSTEMATICS AND ZOOGEOGRAPHY OF THE WORLDWIDE BATHYPELAGIC SQUID BATHYTEUTHIS (CEPHALOPODA: OEGOPSIDA) For sale by the Superintendent of Documents, U.S. Government Printing Office Washington, D.C. 20402 - Price $1.50 (paper covers) UNITED STATES NATIONAL MUSEUM BULLETIN 291 Systematics and Zoogeography of the Worldwide Bathypelagic Squid Bathyteuthis (Cephalopoda: Oegopsida) CLYDE F. E. ROPER Division of Mollusks Smithsonian Institution SMITHSONIAN INSTITUTION PRESS CITY OF WASHINGTON • 1969 Publications of the United States National Museum The scientific publications of the United States National Museum include two series, Proceedings of the United States Nationcd Museum and United States National Museum Bulletin. In these series are published original articles and monographs deal- ing with the collections and work of the Museum and setting forth newly acquired facts in the field of anthropology, biology, geology, history, and technology. Copies of each publication are distributed to libraries and scientific organizations and to specialists and others in- terested in the various subjects. The Prooeedings^f begun in 1878, are intended for the publication, in separate form, of shorter papers. These are gathered in volumes, oc- tavo in size, with the publication date of each paper recorded in the table of contents of the volume. In the Bulletin series, the first of which was issued in 1875, appear longer, separate publications consisting of monographs (occasionally in several parts) and volumes in w^iich are collected works on related subjects. Bulletins are either octavo or quarto in size, depending on the needs of the presentation. Since 1902, papers relating to the bo- tanical collections of the Museum have been published in the Bulletin series under the heading Contributions from the. United States Na- tional Herbarium. This work forms number 291 of the Bulletin series. Frank A. Taylor Director^ United States National Museum IV Contents Page Introduction 1 Material and Methods 5 Part I: SYSTEMATICS 7 Historical R6sum6 8 Bathyteuthis abyssicola Hoyle, 1885 25 Bathyteuthis bacidifera Roper, 1968 43 Bathyteuthis berryi Roper, 1968 53 Comparison of B. abyssicola and B. bacidifera 59 Geographical Variation in Bathyteuthis 68 Buccal Suckers 68 Number of Arm Suckers 75 Gill Size 78 Relationship of Ctenopteryx to Bathyteuthis 85 Familial Relationships of the Bathyteuthidae 89 Part II: ZOOGEOGRAPHY 95 Review of Antarctic Ocean Oceanography 96 Analysis of Environmental Parameters and Distribution 102 Geographic Distribution in Relation to Water Masses 140 Biological Factors Governing Distribution 157 Vertical and Regional Distribution 165 Calculation of Maximum Depth of Capture and Vertical Range . 165 Vertical Distribution by Size 171 Depth of Capture in Relation to Depth of Ocean 175 Aspects of Regional Distribution and Relative Abundance ... 176 Summary and Conclusions 197 References 199 Appendix 209 V Systematics and Zoogeography of the Worldwide Bathypelagic Squid Bathyteuthis (Cephalopoda : Oegopsida)' Introduction Study the deep-sea squid and see how he does only what he has to. Carl Sandburg Bathyteuthis ahyssicola Hoyle has been recorded as an uncommon deep-sea species from widely scattered localities throughout the major oceanic regions of the world. In recent years the relatively large quan- tity of material that has accumulated allows a more thorough analysis of the systematics and distribution of Bathyteuthis. Newly described species and range extensions are represented in small collections from lower latitudes ; however, the bulk of the material, useful in determin- ing details of horizontal and vertical distribution, comes from the Antarctic Ocean. The cephalopods of the open waters of the Southern or Antarctic Ocean and the shallower waters along the shores of the Antarctic Continent are very poorly known despite an impressive list of vessels and expeditions that have collected in these regions. Most of the species known in the Antarctic cephalopod fauna were described from collec- tions made during the great era of exploration prior to World War I. Berry (1917) reviewed the literature and listed the species that occur south of 60° south latitude; with the addition of Berry's 5 new species the list at that time contained 17 species—11 species of the order Octo- poda and 6 species of the suborder Oegopsida. Thiele (1921) and Odhner (1923), reporting on the collections of the German and Swedish Expeditions (both 1901-1903), added to the knowledge of the Antarctic fauna. Robson (1930, 1932) struggled with the complexi- ties of the octopod fauna represented in the early collections of the Discovery. Dell (1959) reported on the cephalopod material col- lected during the British Australian New Zealand Antarctic Research Expedition (B.A.N.Z.A.R.E.) and listed the species that occur in the ^ Contribution no. 982 from the Institute of Marine Sciences, University of Miami and paper no. 65 from tlie Bana oceanographical collections. This paper was submitted in partial fulfillment of the requirements of the Degree of Doctor of Philosophy, University of Miami. 1 . 2 U.S. NATIONAL MUSEUM BULLETIN 291 zoogeographical regions defined by Powell (1951) for the Southern Ocean. Dell listed 20 species of c«phalopods in the Antarctic Province that is bounded in the north by the mean location of the Antarctic Convergence. The list included 12 species of octopods and 8 species of oegopsids (4 of them cranchiids). Every species that Dell listed, endemic or cosmopolitan, presents a systematic problem, and the dis- tribution and biology of these forms have been virtually unknown. Some other groups of the Antarctic marine fauna, however, have been studied more thoroughly, primarily because of the extensive pro- gram carried on by the Discmiery office. More than 30 volumes of Discovery Reports have been prepared on the biology of Southern Ocean organisms. The systematics and distribution of planktonic and nektonic groups have been presented by Mackintosh (1934, 1937), Hardy and Gunther (1935), Fraser (1936), Baker (1954), Tebble ( 1960) , Marr ( 1962) , and many others. In 1962 the Office of Antarctic Programs of the National Science Foundation initiated its program in oceanography by the deployment of the USNS Eltanin to the Southern Ocean. A biological collecting program of broad scope provided the opportunity to conduct detailed studies on the marine fauna of Antarctica. A grant to study the sys- tematics and distribution of Antarctic cephalopods was awarded to G. L. Voss of the Institute of Marine Sciences, University of Miami. As the large collections were sorted and identified, it became increas- ingly clear that the cephalopod fauna of Antarctic waters was con- siderably more extensive and more complex than had been indicated by all previous surveys. Preliminary sorting and identification of the collections by the writer in the winter of 1965 revealed that approxi- mately 30 species of the suborder Oegopsida occur in the Southern Ocean. Some of these species are relatively well known ; some represent long extensions in range; several are undescribed. Nearly the same situation holds for the dozen or so nominal species of benthic octopods that are being studied by G. L. Voss. The finned octopods, a perpetual problem group, are represented by about a half-dozen species. Recent additions to the cephalopod fauna of the Antarctic include the Batoteuthidae, a new family of oegopsids (Young and Roper, 1968), the second and third specimens of the curious Prornachoteuthis Hoyle (Roper and Young, 1968), the second known specimen of Cir- rothauma rwuTrafyl Chun, and new species in several oegopsid families (e.g., Cranchiidae, Histioteuthidae) While new or rare species were being added to the fauna with the increasing collections, one species, Batliyteuthlfi ahysshoJn. Hoyle, 1885, emerged as the overwhelmingly dominant species of pelagic cepha- lopod. It seemed to be common everywhere in the Antarctic Ocean, BATITi'PELAGIC SQUID BATHYTEUTHIS 3 where it was captured over a great range of depths ; often it was taken in what is considered great abundance for a deep-sea squid, with as many as two-dozen specimens in a single midwater tow. Although B. abyssicoki has been recorded in the literature from several different localities in the Atlantic, Pacific, Indian, and Antarctic Oceans, it has been regarded as a rather rare bathypelagic squid that exhibits a world- wide distribution. The overall objectives of the Antarctic Celphalopod Project are to delineate the Antarctic cephalopod fauna, to define the horizontal and vertical distributions of its components, and to determine its zoogeo- graphic relationships with the faunas of adjoining regions. The sys- tematics of the fauna is currently being worked out group by group, but it seemed that the ultimate objectives of the program could be achieved by first working out the distribution and zoogeographic rela- tionships of a single species as a model for comparison in future studies on other species. Bathyteuthis ahyssicola was the obvious choice for the initial study because of its numbers and because it occurs throughout the area of operations of the Eltanin. In addition to the material from the Antarctic, a number of speci- mens of Bathyteuthis from the Atlantic, East Pacific, and Indian Oceans were available from the collections of the Dana, PiUsbury^ Velero, Anton Bruun, Chain, and Bureau of Commercial Fisheries vessels. The work that is presented here is divided into two major parts. The part on
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