Systematics, Ecology and Biology of Cirrate Octopods: Workshop Report
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BULLETIN OF MARINE SCIENCE, 71(1): 79–94, 2002 SYSTEMATICS, ECOLOGY AND BIOLOGY OF CIRRATE OCTOPODS: WORKSHOP REPORT Michael Vecchione and Martin A. Collins with an Appendix by Michael J. Sweeney Cirrate octopods are conspicuous members of the benthopelagic and bathypelagic com- munities and include some of the largest invertebrates of the deep-sea. Although they have often been considered to be rare members of deep-sea communities, recent trawling has shown that the relative abundance of some cirrates, especially opisthoteuthids, may be locally or regionally high (e.g., Boyle et al., 1998). Cirrates also are presumed to be primitive, morphologically similar to ancestral octopods (Young et al., 1998), although recent observations indicate unexpected adaptations such as bioluminescence (Johnsen et al., 1999) and possibly diverse feeding modes (Vecchione and Young, 1997). There- fore, knowledge of cirrates may contribute substantially to understanding cephalopod evolution as well as deep-sea biology and ecology. However, because they are fragile, the condition of specimens collected up until the past decade generally has been very poor, and our knowledge of the group is rudimentary. New methods, such as videotapes re- corded in-situ and gentle collection by submersibles, have allowed substantial new ob- servations and renewed interest in the group. WORKSHOP FORMAT Participation in the cirrate workshop was by invitation only and was limited to re- searchers actively working on cirrates. A few members of the world’s rare and widely dispersed community of cirrate specialists were unable to attend, resulting in participa- tion by 20 researchers from 8 countries (Appendix 1). Many of the participants expressed interest in making a presentation to the group. The two days devoted to the workshop were therefore divided into morning and afternoon sessions. The morning sessions fo- cused on brief, informal presentations, freely interspersed with discussion. Nineteen top- ics were thus presented, ranging from evolution based on morphological and molecular systematics, through direct observations of behavior, to translation and discussion of classic cirrate literature. Many of these topics have been formalized as manuscripts and several are published in this volume (e.g., Villanueva et al., 2002). During afternoon sessions the participants separated into working groups to focus their attention on the following top- ics: (1) taxonomy and systematics, (2) behavior and functional anatomy, (3) biology, growth, and feeding, and (4) distribution and abundance. Along with detailed discus- sions, these afternoon breakout sessions allowed workers to examine well-preserved speci- mens of many cirrate taxa and to review videotapes that had been shown all too briefly during the morning presentations. Assembling a ‘critical mass’ of workers enhances the effectiveness of their individual research. This workshop allowed cirrate researchers from around the world to share the recent progress they have made and to discuss possible differences in interpretations. Such observations and interpretations were discussed in light of many recent publica- tions on cirrates. In particular, much discussion focused on stabilizing cirrate systemat- 79 80 BULLETIN OF MARINE SCIENCE, VOL. 71, NO. 1, 2002 Table 1. Recommendations for analysis, fixation and preservation of cirrate octopods. Particularly when the identification of specimens is not known. Submersible observation and capture 1. Video for short period prior to capture attempt. Check for bioluminescence if possible. 2. Capture gently and bring to surface. 3. Recommend use of special gelatinous zooplankton sampler (Contact Jay Hunt) Trawl and submersible caught specimens on the surface 1. Check for bioluminescence. Place in chilled sea-water in dark room. Stimulate mechanically to determine if light is produced. 2. Remove tissue sample for DNA/molecular analysis and place in 80% ethanol. 3. Take basic measurements. (total length, mantle length, head width, arm lengths, fin lengths, fin span, cirri length, weight). 4 . Take photographs of fresh specimens (dorsal, ventral and oral views), covered in seawater with polarised filter. 5. Collect eggs from distal oviduct and incubate in cold sea-water. 6. Obtain kidney smears. 7. Fix in 10% buffered formalin for minimum of 24 hrs. 8. Transfer to Steadmans solution (2.5% formalin, 1% propylene phenoxitol, 10% propylene glycol, made up in sea-water) for storage. ics, which currently is in disarray (Appendix 2). Other questions for which discussions were productive ranged from the very basic (e.g., What are the best methods for fixation and preservation? See Table 1) to complex problems (such as variation in functional morphology of feeding). Although two days was not enough time to resolve this broad range of problems, we feel that substantial progress was made, particularly by facilitation of communications among researchers who are usually widely scattered across the globe. One tool that proved to be especially useful throughout these deliberations was a sum- mary of the current status of cirrate nomenclature, prepared for the workshop by Michael Sweeney of the Smithsonian Institution, who unfortunately was unable to travel to Scot- land. We present an abridged version of the summary (Appendix 2) so that it can continue to be of use to researchers who are interested in these fascinating but difficult cephalo- pods. ACKNOWLEDGMENTS The workshop was funded by the CEC under contract number QLAM-PL1999-30001. LITERATURE CITED Boyle, P. R., M. A. Collins and G. R. Williamson. 1998. The cephalopod by-catch of deep-water trawling on the Hebrides slope. J. Mar. Biol. Ass. UK 78: 1023–1026. Johnsen, S., E. J. Balser and E. A. Widder. 1999. Light-emitting suckers in an octopus. Nature 397: 113–114. Vecchione, M. and R. E. Young. 1997. Aspects of the functional morphology of cirrate octopods: Locomotion and feeding. Vie Milieu 47: 101–110. Villanueva, R., M. A. Collins, P. Sanchez and N. A. Voss. 2002. Systematics and distribution in the Atlantic Ocean of the cirrate octopods of the genus Opisthoteuthis (Mollusca, Cephalopoda), with description of two new species. Bull. Mar. Sci. (this issue). CIRRATE OCTOPODS WORKSHOP REPORT 81 Young, R. E., M. Vecchione and D. T. Donovan. 1998.The evolution of coleoid cephalopods and their present biodiversity and ecology. S. Afr. J. Mar. Sci. 20: 393–420. ADDRESSES: (M.V.) National Marine Fisheries Service Systematics Laboratory, National Museum of Natural History, Washington, DC 20560 USA. E-mail: <[email protected]>. (M.A.C.) Department of Zoology, University of Aberdeen, Tillydrone Avenue, Aberdeen, AB24 2TN, Scotland, UK. E-mail: <[email protected]>. APPENDIX 1 REGISTERED WORKSHOP PARTICIPANTS Slava Bizikov, Russia Peter Boyle, UK Martin Collins, UK Heather Daly, UK Ian Gleadall, Japan Angel Guerra, Spain Eric Hochberg, US Cendrine Hudelot, France Jay Hunt, Japan Tsunemi Kubodera, Japan C.C. Lu, Taiwan Kir Nesis, Russia Chingis Nigmatullin, Russia Martina Roeleveld, South Africa Clyde Roper, US Pilar Sanchez, Spain Mike Vecchione, US Roger Villanueva, Spain Janet Voight, US Dick Young, US 82 BULLETIN OF MARINE SCIENCE, VOL. 71, NO. 1, 2002 APPENDIX 2 NOMINAL TAXA OF CIRRATE OCTOPODS Michael J. Sweeney To assist the individuals participating in the Cirrate Workshop at CIAC 2000 with the daunting task of updating cirrate systematics and biology, the following data are com- piled: (1) Nominal genus-level taxa with taxon, author, date of publication and pagination; current systematics status of the taxon fide the most recent review; type species of the taxon with author, date and designation. (2) Nominal species-level with taxon, author, date of publication, pagination, and illus- trations; current systematics status fide the most recent published review of the taxon; type repository with museum acronym, type designation and catalog number fide pub- lished type catalogs; published type locality for the taxon and water depth (or capture depth); remarks if required. (3) Genera introduced in cirrate families but now known to be non-cirrate (exclusively synonymized with Vampyroteuthis). (4) Acronyms and addresses for museum where cirrate types are deposited; published type catalogs of cephalopods for the respective museums are given in brackets. (5) References for literature cited within. NOMINAL GENUS-LEVEL TAXA Chunioteuthis Grimpe, 1916:355 CURRENT SYSTEMATIC STATUS.—synonym of Stauroteuthis Verrill, 1879 [fide Collins and Henriques (2000:690)] TYPE SPECIES.—Chunioteuthis ebersbachii Grimpe, 1916 by monotypy. Cirroctopus Naef, 1923:675 CURRENT SYSTEMATIC STATUS.— valid genus [fide O’Shea (1999:34)] TYPE SPECIES.— Stauroteuthis mawsoni Berry, 1917 by original designation Cirroteuthis Eschricht, 1836:633 CURRENT SYSTEMATIC STATUS.— valid genus [fide Voss (1988b:266)] TYPE SPECIES.— Cirroteuthis muelleri Eschricht, 1836 by monotypy (Cirroteuthopsis) Grimpe, 1920:233 CURRENT SYSTEMATIC STATUS.— synonym of Opisthoteuthis Verrill, 1883 [fide Villanueva et al. {this issue}] TYPE SPECIES.— Cirroteuthis (Cirroteuthopsis) massyae Grimpe, 1920 by monotypy. Cirrothauma Chun, 1911:5 CURRENT SYSTEMATIC STATUS.— valid genus [fide Voss (1988b:267)] TYPE SPECIES.— Cirrothauma murrayi Chun, 1911 by monotypy CIRRATE OCTOPODS WORKSHOP REPORT 83 Cirrothauma Chun, 1913:22 CURRENT SYSTEMATIC STATUS.— preoccupied in Chun (1911) Enigmatiteuthis O’Shea, 1999:47 CURRENT SYSTEMATIC STATUS.— taxon not yet reviewed TYPE SPECIES.— Enigmatiteuthis innominata