Systematics, Ecology and Biology of Cirrate Octopods: Workshop Report

Home , Cirroteuthis, Cirrothauma murrayi, Opisthoteuthis, Stauroteuthis, Stauroteuthis syrtensis

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 communities 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 recorded in-situ and gentle collection by submersibles, have allowed substantial new observations and renewed interest in the group.

WORKSHOP FORMAT

Participation in the cirrate workshop was by invitation only and was limited to researchers 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 participation 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 focused on brief, informal presentations, freely interspersed with discussion. Nineteen topics 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 topics: (1) taxonomy and systematics, (2) behavior and functional anatomy, (3) biology, growth, and feeding, and (4) distribution and abundance. Along with detailed discussions, these afternoon breakout sessions allowed workers to examine well-preserved specimens 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 publications 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 summary 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 cephalopods.

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: . (M.A.C.) Department of Zoology, University of Aberdeen, Tillydrone Avenue, Aberdeen, AB24 2TN, Scotland, UK. E-mail: .

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 published 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 O’Shea, 1999 by original designation

Froekenia Hoyle, 1904:7 CURRENT SYSTEMATIC STATUS.— valid genus [fide Guerra et al. (1998:75)] TYPE SPECIES.— Froekenia clara Hoyle, 1904 by monotypy

Grimpoteuthis Robson, 1932:136 CURRENT SYSTEMATIC STATUS.— valid genus [fide Voss (1988b:267)] TYPE SPECIES.— Cirroteuthis umbellata Fischer, 1883 by original designation

(Heterohistion) Grimpe, 1920:232 CURRENT SYSTEMATIC STATUS.— unresolved TYPE SPECIES.— [?]Cirroteuthis magna Hoyle, 1885

Luteuthis O’Shea, 1999:56 CURRENT SYSTEMATIC STATUS.— taxon not yet reviewed TYPE SPECIES.— Luteuthis dentatus O’Shea, 1999 by original designation

Opisthoteuthis Verrill, 1883:113 CURRENT SYSTEMATIC STATUS.— valid genus [fide Voss (1988b:267)] TYPE SPECIES.— Opisthoteuthis agassizii Verrill, 1883 by monotypy

Sciadephorus Reinhardt and Prosch, 1846:189 CURRENT SYSTEMATIC STATUS.— synonym of Cirroteuthis Eschricht, 1836 [fide Hoyle (1910:412)] TYPE SPECIES.— Cirroteuthis muelleri Eschricht, 1836 by monotypy

Stauroteuthis Verrill, 1879:468 CURRENT SYSTEMATIC STATUS.— valid genus [fide Voss (1988b:267)] TYPE SPECIES.— Stauroteuthis syrtensis Verrill, 1879 by monotypy

(Teuthidiscus) Berry, 1918:284 CURRENT SYSTEMATIC STATUS.— synonym of Opisthoteuthis Verrill, 1883 [fide Voss (1988b:265)] TYPE SPECIES.— Opisthoteuthis (Teuthidiscus) pluto Berry, 1918 by original designation

NOMINAL SPECIES-LEVEL TAXA (AS INTRODUCED BINOMIAL)

Grimpoteuthis abyssicola O’Shea, 1999:46, figs 28–29 CURRENT SYSTEMATIC STATUS.— taxon not yet reviewed TYPE REPOSITORY.— NZOI Holotype H-661 TYPE LOCALITY.— New Zealand (35o35.1'S, 160o57.1'E), in 3180–3154 m. 84 BULLETIN OF MARINE SCIENCE, VOL. 71, NO. 1, 2002

Opisthoteuthis agassizii Verrill, 1883:113, pl 1 fig 1, pl 2 fig 1 CURRENT SYSTEMATIC STATUS.— valid species [fide Nesis (1987:288)] TYPE REPOSITORY.— MCZ? Holotype (not traced) TYPE LOCALITY.— off Grenada [12o03'30"N, 61o47'10"W], in 291 fm [532 m]. REMARKS.— Type species of Opisthoteuthis Verrill, 1883.

Stauroteuthis albatrossi Sasaki, 1920:169, pl 23 figs 2–3 CURRENT SYSTEMATIC STATUS.— valid species; Opisthoteuthis albatrossi [fide O’Shea (1999:24)] TYPE REPOSITORY.— NMNH Holotype 332949 [fide Roper and Sweeney (1978:3)] TYPE LOCALITY.— off Kinka-san, Rikuzen Province [38oN, 142oE], Japan, in about 250 fm [457 m]. REMARKS.— See Sasaki (1929:7) for additional figures of type.

Grimpoteuthis antarctica Kubodera and Okutani, 1986:129, text-fig 1, pl 1 CURRENT SYSTEMATIC STATUS.— valid species; Cirroctopus antarctica [fide O’Shea (1999:35)] TYPE REPOSITORY.— NSMT Holotype 63958 [fide Kubodera and Tsuchiya (1993:unpaginated)] TYPE LOCALITY.— 62o59'S, 62o09'W (Southern Ocean), in 803–804 m.

Grimpoteuthis bathynectes Voss and Pearcy, 1990:57, figs 4–5 CURRENT SYSTEMATIC STATUS.— valid species; taxon not yet reviewed TYPE REPOSITORY.— NMNH Holotype 730715 [fide Sweeney, pers. comm.] TYPE LOCALITY.— 45o01.1'N, 135o12.0'W (NE Pacific Ocean), in 3932 m.

Grimpoteuthis bruuni Voss, 1982:426, figs 1–2 CURRENT SYSTEMATIC STATUS.— valid species [fide Nesis (1987:286)] TYPE REPOSITORY.— NMNH Holotype 730616 [fide Sweeney, pers. comm.] TYPE LOCALITY.— off Antofagasta, Chile (23o41'S, 70o34'W), in 250–360 m.

Opisthoteuthis californiana Berry, 1949:23 CURRENT SYSTEMATIC STATUS.— valid species [fide Nesis (1987:288)] TYPE REPOSITORY.— CASIZ Holotype 021666 [fide Sweeney et al. (1988:12)] TYPE LOCALITY.— northwest by west of Eureka Bar [40oN, 124oW], California, U.S.A., in 188 fm [343 m].

Opisthoteuthis calypso Villanueva et al., {this issue} CURRENT SYSTEMATIC STATUS.— valid species, taxon not yet reviewed TYPE REPOSITORY.— {not stated in ms} TYPE LOCALITY.— Cantabric Sea, 43o34.64'N, 02o16.47'W.

Opisthoteuthis chathamensis O’Shea, 1999:25, figs 13–14 CURRENT SYSTEMATIC STATUS.— valid species, taxon not yet reviewed TYPE REPOSITORY.— NZOI Holotype H-664 TYPE LOCALITY.— New Zealand (approx. 44o43'S, 177o15'W), in 1174–1195 m.

Froekenia clara Hoyle, 1904:7, pl 2 fig 2, pl 3 fig 5 Nomen dubium [fide Voss (1988b:265)]; Valid species [fide Nesis (1987:282)] TYPE REPOSITORY.— NMNH Holotype 382468; specimen no longer extant [fide Roper and Sweeney (1978:5)] TYPE LOCALITY.— 06o30'N, 81o44'W [Pacific Ocean, off Panama], in 555 fm [1015 m]. REMARKS.— Type species of Froekenia Hoyle, 1904 CIRRATE OCTOPODS WORKSHOP REPORT 85

Cirroteuthis caudani Joubin, 1896:247 CURRENT SYSTEMATIC STATUS.— synonym of Grimpoteuthis grimaldii (Joubin, 1903)? [fide Nesis (1987:285)] TYPE REPOSITORY.— “Zool. Dept., Univ. Lyon”? [fide Robson (1932:145)] (not traced) TYPE LOCALITY.— Bay of Biscay (44o36'N, 04o25'W), in 650 m.

Luteuthis dentatus O’Shea, 1999:57, figs 36–38 CURRENT SYSTEMATIC STATUS.— valid species, taxon not yet reviewed TYPE REPOSITORY.— NMNZ Holotype M.131564 TYPE LOCALITY.— New Zealand (40o01.3'S, 167o49.9'E), in 991 m. REMARKS.— Type species of Luteuthis O’Shea, 1999

Opisthoteuthis depressa Ijima and Ikeda, 1895:323, pl 33 CURRENT SYSTEMATIC STATUS.— valid species [fide Okutani et al. (1987:155)] TYPE REPOSITORY.— MSUT Holotype; specimen no longer extant (not traced) TYPE LOCALITY.— south of Misaki [35oN, 139oE], Japan, in about 250 fm [457 m].

Chunioteuthis ebersbachii Grimpe, 1916:357, text-figs 1–3 CURRENT SYSTEMATIC STATUS.— synonym of Stauroteuthis syrtensis Verrill, 1879 [fide Collins and Henriques (2000:690)] TYPE REPOSITORY.— ZMUB Holotype 36371 [fide Collins and Henriques (2000:688)] TYPE LOCALITY.— north of Sargasso Sea, 1100 m [42o59'N, 51o51'W fide Robson (1932:157)] REMARKS.— Type species of Chunioteuthis Grimpe, 1916

Opisthoteuthis extensa Thiele, in Chun, 1915:537, pl 94 fig 3, pl 95 fig 2 CURRENT SYSTEMATIC STATUS.— unresolved; synonym of Opisthoteuthis meangenis (Hoyle, 1885) [fide Hochberg & Norman, pers. comm.] TYPE REPOSITORY.— ZMB Holotype Moll.-110010 [fide Glaubrecht and Salcedo-Vargas (2000:274)] TYPE LOCALITY.— 0o57.5'S, 99o51.1'E, Mentawei Basin (Indian Ocean), in 768 m.

Cirrhoteuthis gilchristi Robson, 1924a:204 CURRENT SYSTEMATIC STATUS.— valid species; Stauroteuthis gilchristi [fide Collins and Henriques (2000:694)] TYPE REPOSITORY.— BMNH Holotype 1924.9.9.7 [fide Lipinski et al. (2000:111)] TYPE LOCALITY.— off Cape Town [33oS, 18oE], South Africa, in about 1400 fm [2561 m]. REMARKS.— See also Robson (1924b:677, text-fig 46) for illustration of type.

Cirroteuthis glacialis Robson, 1930:375, figs 1–5 CURRENT SYSTEMATIC STATUS.— valid species; Cirroctopus glacialis [fide O’Shea (1999:35)] TYPE REPOSITORY.— BMNH Holotype 1951.4.26.1 [fide Lipinski et al. (2000:111)] TYPE LOCALITY.— Palmer Archipeligo, 64o21'S, 62o58'W (Southern Ocean), in 278–500 m.

Cirroteuthis grimaldii Joubin, 1903:100 CURRENT SYSTEMATIC STATUS.— valid species; Opisthoteuthis grimaldii [fide Villanueva et al. {this issue}] TYPE REPOSITORY.— MOM Holotype [fide Belloc (1950:2)] TYPE LOCALITY.— NNW of Fayal, Azores [39oN, 29oW], in 1900 m [fide Joubin (1912:1)] REMARKS.— See Joubin (1912, 1920) for additional description and illustration of type. 86 BULLETIN OF MARINE SCIENCE, VOL. 71, NO. 1, 2002

Cirroteuthis grimaldii Joubin, 1912:1 CURRENT SYSTEMATIC STATUS.— preoccupied in Joubin (1903)

Opisthoteuthis hardyi Villanueva et al. {this issue} CURRENT SYSTEMATIC STATUS.— valid species, taxon not yet reviewed TYPE REPOSITORY.— NMS Holotype 199958.088 TYPE LOCALITY.— South Georgia (53o18'S, 42o12'W), in 1000 m.

Stauroteuthis hippocrepium Hoyle, 1904:6, pls 1–3 CURRENT SYSTEMATIC STATUS.— valid species; Grimpoteuthis hippocrepium [fide O’Shea (1999:46)] TYPE REPOSITORY.— NMNH Holotype 382467 [fide Roper and Sweeney (1978:8)] TYPE LOCALITY.— SW of Malpelo Island, off Colombia, Pacific Ocean (02o35'N, 83o53'W), in 1823 fm [3334 m].

Cirroctopus hochbergi O’Shea, 1999:35, figs 23–25 CURRENT SYSTEMATIC STATUS.— valid species, taxon not yet reviewed TYPE REPOSITORY.— NZOI Holotype H-674 TYPE LOCALITY.— New Zealand (39o58–54'S, 178o14–16'E), in 800–1070 m.

Cirroteuthis(?) hoylei Robson, 1932:161 CURRENT SYSTEMATIC STATUS.— questionable species; Cirroteuthis sp. [fide Guerra et al. (1998:77)] TYPE REPOSITORY.— BMNH Holotype 1890.1.24.2 [fide Lipinski et al. (1998:77)] TYPE LOCALITY.— 34o07'S, 73o56'W [off Valparaiso, Chile] (SE Pacific Ocean), in 2225 fm [4071 m].

Enigmatiteuthis innominata O’Shea, 1999:48, figs 32–33 CURRENT SYSTEMATIC STATUS.— valid species, taxon not yet reviewed TYPE REPOSITORY.— NMNZ Holotype M.109381 TYPE LOCALITY.— off New Zealand, 42o36.7'S, 176o09.8'E, in 1999–2002 m. REMARKS.— Type species of Enigmatiteuthis O’Shea, 1999

Opisthoteuthis (Teuthidiscus) japonica Taki, 1962:397 CURRENT SYSTEMATIC STATUS.— valid species; Opisthoteuthis japonica [fide Nesis (1987:288)] TYPE REPOSITORY.— YTKC? Holotype [not extant?] (not traced) TYPE LOCALITY.— off Minabe [33oN, 135oE], Wakayama Prefecture, Japan, in 152 m [fide Taki (1963:77)]. REMARKS.— See also Taki (1963:74) for additional description and illustration of type.

Cirroteuthis magna Hoyle, 1885:233 CURRENT SYSTEMATIC STATUS.— valid species [fide Guerra et al. (1998:55)] TYPE REPOSITORY.— BMNH Holotype 1890.1.24.1 [fide Lipinski et al. (2000:112)] TYPE LOCALITY.— 46o46'S, 45o31'E [between Prince Edward Island and Crozet Islands] (S Indian Ocean), in 1375 fm [2516 m]. REMARKS.— Type species of (Heterohistion) Grimpe, 1920[?]. See also Hoyle (1886:56) for additional description and illustration of type. See also Guerra et al. (1998:51) for redescription of species. CIRRATE OCTOPODS WORKSHOP REPORT 87

Cirroteuthis (Cirroteuthopsis) massyae Grimpe, 1920:230 CURRENT SYSTEMATIC STATUS.— valid species; Opisthoteuthis massyae [fide Villanueva et al. {this issue}] TYPE REPOSITORY.— NMI Holotype (not traced) [fide Massy (1924:128)] TYPE LOCALITY.— [off Ireland] 50o31'N, 11o31'W (Atlantic Ocean), in 670–770 fm [1226–1409 m] [fide Robson (1932:160)]. REMARKS.— Type species of (Cirroteuthopsis) Grimpe, 1920. See also Massy (1909:4) for measurements of type.

Stauroteuthis (?) mawsoni Berry, 1917:8, text-figs 1–4, pl 10 fig 1 CURRENT SYSTEMATIC STATUS.— valid species; Cirroctopus mawsoni [fide O’Shea (1999:35)] TYPE REPOSITORY.— AMS Holotype C40886 [fide Sweeney et al. (1988:15)] TYPE LOCALITY.— off Mertz Glacier, 66o55'S, 145o21'E (Antarctic Ocean), in 288–300 fm [527– 549 m]. REMARKS.— Type species of Cirroctopus Naef, 1923

Cirroteuthis meangensis Hoyle, 1885:234 CURRENT SYSTEMATIC STATUS.— valid species; Opisthoteuthis meangenis (Hoyle, 1885) [fide Hochberg & Norman, pers. comm.] TYPE REPOSITORY.— BMNH Holotype 1890.1.24.4 [fide Lipinski et al. (2000:112)] TYPE LOCALITY.— [S. Philippines], 04o33'N, 127o06'E (Pacific Ocean) [fide Robson (1932:143)], in 500 fm [915 m]. REMARKS.— See also Hoyle (1886:63) for additional description and illustration of type.

Opisthoteuthis medusoides Thiele In Chun, 1915:538, pl 94 figs 1–2, pl 95 fig 1 CURRENT SYSTEMATIC STATUS.— unresolved; synonym of Opisthoteuthis meangenis (Hoyle, 1885) [fide Hochberg & Norman, pers. comm.] TYPE REPOSITORY.— ZMB Lectotype Moll.-63854a [fide Glaubrecht and Salcedo-Vargas (2000:277)] TYPE LOCALITY.— [off Tanzania], 06o39.1'S, 39o30.8'E (W Indian Ocean), in 400 m.

Cirrhoteuthis megaptera Verrill, 1885:405, pl 43 figs 1–2 CURRENT SYSTEMATIC STATUS.— valid species; Grimpoteuthis megaptera [fide Robson (1932:140)] TYPE REPOSITORY.— NMNH Syntypes: [unresolved, types not recently systematically reviewed] 39916 one specimen not extant (sta. 2220); 39963 one specimen not extant (sta. 2225); 40127 one specimen plus one specimen not extant (sta. 2224); 40131 one macerated specimen (sta. 2225) [fide Roper and Sweeney (1978:9)] TYPE LOCALITIES: 39o43'30"N, 69o23'00"W, in 1054 fm [1929 m] (sta. 2220); 36o05'30"N, 69o51'45"W, in 2512 fm [4597 m] (sta. 2225); and 36o16'30"N, 68o21'W, in 2574 fm [4710 m] (sta. 2224) (all Atlantic Ocean)

Opisthoteuthis mero O’Shea, 1999:17, figs 9–10 CURRENT SYSTEMATIC STATUS.— taxon not yet reviewed TYPE REPOSITORY.— NZOI Holotype H-673 TYPE LOCALITY.— New Zealand (approx. 36o51'S, 176o18'E), in 509–514 m.

Cirroteuthis mulleri Eschricht, 1836:633, figs 1–3 CURRENT SYSTEMATIC STATUS.— valid species [fide Nesis (1987:283)] TYPE REPOSITORY.— ZMUC Syntype? [fide Kristensen and Knudsen (1983:223)] TYPE LOCALITY.— Jakobshavn [69oN, 51oW], West Greenland, [depth not given] REMARKS.— Type species of Cirroteuthis Eschricht, 1836. Type species of Sciadephorus Reinhardt and Prosch, 1846. 88 BULLETIN OF MARINE SCIENCE, VOL. 71, NO. 1, 2002

Cirrothauma murrayi Chun, 1911:5, figs 2–6 CURRENT SYSTEMATIC STATUS.— valid species [fide Nesis (1987:279)] TYPE REPOSITORY.— ZMUB? Holotype (not traced) TYPE LOCALITY.— 48o24'N, 36o53'W (Atlantic Ocean), with 3000 m wire out [fide Robson (1932:165)] REMARKS.— Type species of Cirrothauma Chun, 1911. See also Chun (1913:22, figs 7–9) for En- glish language description of type.

Cirrothauma murrayi Chun, 1913:22. CURRENT SYSTEMATIC STATUS.— preoccupied in Chun (1911)

Cirroteuthis pacifica Hoyle, 1885:235 CURRENT SYSTEMATIC STATUS.— valid species; Enigmatiteuthis pacifica [fide O’Shea (1999:50)] TYPE REPOSITORY.— BMNH Holotype 1890.1.24.3 [fide Lipinski et al. (2000:112)] TYPE LOCALITY.— Pacific Ocean between New Guinea and Australia, at surface, station 181 [fide Hoyle (1885:235; 1885b:112)]; off SE extremity of New Guinea, 13o50'S, 151o49'E (SW Pa- cific Ocean), in 2440 fm [4465 m] [fide Hoyle (1886:61)]. REMARKS.— See also Hoyle (1886:61) for additional description and illustration of type.

Opisthoteuthis persephone Berry, 1918:290, text-figs 66–67, pls 81–82, 85–88 CURRENT SYSTEMATIC STATUS.— valid species [fide Nesis (1987:288)] TYPE REPOSITORY.— AMS Holotype C148253 [fide Sweeney et al. (1988:16)] TYPE LOCALITY.— 42 miles SE of Genoa Peak, Victoria [Bass Straits, Australia], in 232–260 fm [424–475 m].

Opisthoteuthis philipii Oommen, 1976:368, fig 1 CURRENT SYSTEMATIC STATUS.— valid species; taxon not yet reviewed TYPE REPOSITORY.— IFLC Holotype [fide Oommen (1976:371)] TYPE LOCALITY.— Off Alleppey, SW India, 09o32'N, 75o45'E (Indian Ocean), in 275 m.

Cirrhoteuthis plena Verrill, 1885:404, pl 42 fig 3 CURRENT SYSTEMATIC STATUS.— [unresolved, types not recently systematically reviewed]; valid species; Grimpoteuthis plena [fide Robson (1932:145)] TYPE REPOSITORY.— NMNH Holotype 39908 [fide Roper and Sweeney (1978:11)] TYPE LOCALITY.— [off New Jersey, U.S.A.] 39o35'00"N, 71o18'45"W (Atlantic Ocean), in 1073 fm [1963 m]. REMARKS.— Latitude originally published incorrectly by Verrill as 37o35’N.

Opisthoteuthis (Teuthidiscus) pluto Berry, 1918:284, text-figs 64–65, pls 81–84 CURRENT SYSTEMATIC STATUS.— valid species; Opisthoteuthis pluto [fide Nesis (1987:288)] TYPE REPOSITORY.— AMS Holotype; specimen not extant [fide Rudman (1983:67)] TYPE LOCALITY.— Great Australian Bight, in 350–450 fm [640–823 m]. REMARKS.— Type species of (Teuthodiscus) Berry, 1918

Cirroteuthis richardi Joubin, 1903:101 CURRENT SYSTEMATIC STATUS.— [unresolved, type not recently systematically reviewed]; synonym of Grimpoteuthis megaptera (Verrill, 1885) [fide Joubin (1920:15)] TYPE REPOSITORY.— MOM Holotype [fide Robson (1932:141)] TYPE LOCALITY.— off Cape Verde Island, in 3890 m. CIRRATE OCTOPODS WORKSHOP REPORT 89

Opisthoteuthis robsoni O’Shea, 1999:28, figs 17–18 CURRENT SYSTEMATIC STATUS.— taxon not yet reviewed TYPE REPOSITORY.— NMNZ Holotype M.95255 TYPE LOCALITY.— New Zealand (42º41.7'S, 174º28.0'E), in 1723–1549 m.

Stauroteuthis syrtensis Verrill, 1879:469 CURRENT SYSTEMATIC STATUS.— valid species [fide Collins and Henriques (2000:690)] TYPE REPOSITORY.— NMNH Holotype 382471 [fide Roper and Sweeney (1978:13)] TYPE LOCALITY.— off Nova Scotia, 43º54'N, 58º44'W (Atlantic Ocean), in 250 fm [457 m]. REMARKS.— Type species of Stauroteuthis Verrill, 1879. See also Verrill (1880–1881:382) and Collins and Henriques (2000:690) for additional description and illustration of type.

Grimpoteuthis tuftsi Voss and Pearcy, 1990:63, figs 6–8 CURRENT SYSTEMATIC STATUS.— valid species; Luteuthis tuftsi [fide O’Shea (1999:56)] TYPE REPOSITORY.— NMNH Holotype 730714 [fide Sweeney, pers. comm.] TYPE LOCALITY.— off Oregon, U.S.A., 45º05.2'N, 134º43.4'W (Pacific Ocean), in 3900 m.

Cirroteuthis umbellata Fischer, 1883:404 CURRENT SYSTEMATIC STATUS.— [unresolved, types not recently systematically reviewed]; valid species; Grimpoteuthis umbellata [fide Robson (1932:137)] TYPE REPOSITORY.— MNHN Syntypes (originally 3) 3-6-698 (arms only, very poor condition) and 3-6-699 (poor condition) [fide Lu et al. (1995:326); remaining syntype (sta. 80) not recorded.] TYPE LOCALITY.— off Azores at 37º55'N, 29º22'W [“Talisman” sta. 130, MNHN 3-6-698], in 2235 m; off Sahara, near Cape Garnett at 23º50'N, 19º37'W [sta. 80], in 1139 m; and between Cape Nounn and Canary Islands, in 1235 m [sta. 45, MNHN 3-6-699] [fide Joubin (1900:22) and Lu et al. (1995:326)] REMARKS.— Type species of Grimpoteuthis Robson, 1932. See English version of Joubin (1900:22) for translation of syntype description. See Fischer and Joubin (1907:318) for additional description and illustration of syntypes.

Opisthoteuthis vossi Sanchez and Guerra, 1989:1159, figs 1–2 CURRENT SYSTEMATIC STATUS.— synonym of Opisthoteuthis massyae (Grimpe, 1920) [fide Villanueva et al., {this issue}] TYPE REPOSITORY.— NMNH Holotype 816660 [fide Sweeney, pers. comm.] TYPE LOCALITY.— Valdivia Bank, 22º29'S, 06º07'E (Atlantic Ocean), in 900 m.

Stauroteuthis wulkeri Grimpe, 1920:235, figs 1–6 CURRENT SYSTEMATIC STATUS.— valid species; Enigmatiteuthis wulkeri [fide O’Shea (1999:50)] TYPE REPOSITORY.— ZMUB Holotype [fide Robson (1932:152)] TYPE LOCALITY.— off Morocco, 35º46'N, 08º16'W (NE Atlantic Ocean), in 2055 m. REMARKS.— See Robson (1932:139, 152) for discussion of type material.

GENERA INTRODUCED IN CIRRATE FAMILIES, NOW SYNONYMIZED AS NON-CIRRATE.

Danateuthis Joubin, 1929:375 CURRENT SYSTEMATIC STATUS.— Vampyroteuthis Chun, 1903 [fide Pickford (1946:7)] TYPE SPECIES.— Danateuthis schmidti Joubin, 1929 by monotypy REMARKS.— See also Pickford (1949:122) 90 BULLETIN OF MARINE SCIENCE, VOL. 71, NO. 1, 2002

Hansenoteuthis Joubin, 1929:388 CURRENT SYSTEMATIC STATUS.— Vampyroteuthis Chun, 1903 [fide Nesis (1987a:276)] TYPE SPECIES.— Hansenoteuthis lucens Joubin, 1929 by monotypy REMARKS.— See also Pickford (1949:123)

Hymenoteuthis Thiele, 1916:4 CURRENT SYSTEMATIC STATUS.— Vampyroteuthis Chun, 1903 [fide Pickford (1946:7)] TYPE SPECIES.— Cirroteuthis macrope Berry, 1911 by monotypy REMARKS.— See also Pickford (1949:127)

Melanoteuthis Joubin, 1912:1, figs 1–12 CURRENT SYSTEMATIC STATUS.— Vampyroteuthis Chun, 1910 [fide Pickford (1939:1) TYPE SPECIES.— Melanoteuthis lucens Joubin, 1912 by monotypy REMARKS.— See also Pickford (1949:125)

Retroteuthis Joubin, 1929:383 CURRENT SYSTEMATIC STATUS.— Vampyroteuthis Chun, 1903 [fide Pickford (1946:7)] TYPE SPECIES.— Retroteuthis pacifica Joubin, 1929 by monotypy REMARKS.— See also Pickford (1949:126)

Watasella Sasaki, 1920:168 CURRENT SYSTEMATIC STATUS.— Vampyroteuthis Chun, 1903 [fide Nesis (1987:276)] TYPE SPECIES.— Watasella nigra Sasaki, 1920 by monotypy REMARKS.— See also Pickford (1949:128)

TYPE REPOSITORIES

AMS— Australian Museum, P.O. Box A285, 6-8 College Street, Sydney South, New South Wales 2000, Australia. [Rudman (1983); Sweeney, Roper and Hochberg (1988)] BMNH— The Natural History Museum, Cromwell Road, London SW7 5BD, England (formerly British Museum (Natural History)). [Lipinski et al. (2000)] CASIZ— California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118, U.S.A. [Smith (1974); Sweeney, Roper and Hochberg (1988)] IFLC— Integrated Fisheries Project Marine Research Laboratory, P.O. Box 1801, Cochin 682016, India. MCZ— Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138, U.S.A. MOM— Musee Oceanographique, Avenue Saint-Martin, Monaco-Ville, MC 98000, Monaco. [Belloc (1950)] MSUT— University Museum, University of Tokyo, Hongo 7-3-1, Bunkyoku, Tokyo 113, Japan (includes former Tokyo Imperial University (TIU) collection and University of Tokyo Depart- ment of Fisheries (FUMT) collection). NMI— National Museum of Ireland, Kildare Street, Dublin 2, Ireland. NMNH— National Museum of Natural History, 10th & Constitution Avenue, Smithsonian Institu- tion, Washington, DC 20560, U.S.A. [Roper and Sweeney (1978); Sweeney, Roper and Hochberg (1988)] NMS— National Museums of Scotland, Chambers Street, Edinburgh, EH1 1JF, Scotland. NMNZ— National Museum of New Zealand - Te Papa Tongarewa, Cable Street, Wellington, New Zealand (formerly Colonial Museum; Dominion Museum). CIRRATE OCTOPODS WORKSHOP REPORT 91

NSMT— National Science Museum, 3-23-1 Hyakunin-cho, Shinjuku-ku, Tokyo 160, Japan. [Kubodera and Tsuchiya (1993)] NZOI— National Institute of Water and Atmospheric Research, P.O. Box 14-901, Kilbirnie, Wellington, New Zealand. [O’Shea (1999)] YTKC— Private collection of the late Dr. Iwao Taki stored at the home of Dr. Yoshio Taki, Kyoto City, Japan. [fide T. Okutani, pers. comm.]. ZMB— Zoologisches Museum, Museum für Naturkunde der Humboldt-Universitaet, Invalidenstrasse 43, D-1040 Berlin, Germany. [Glaubrecht and Salcedo-Vargas (2000)] ZMUB— Zoological Museum, Universitetet i Bergen, Museplass 3, 5000 Bergen, Norway. ZMUC— Kobenhavns Universitet, Zoologisk Museum, Universitetsparken15, DK 2100 Copenhagen, Denmark. [Kristensen and Knudsen (1983); Also see Volsoe et al. (1962) for Steenstrup papers]

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ADDRESS: (M.J.S.) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. E-mail: CURRENT ADDRESS: Biology Department, New Mexico State University, Las Cruces, New Mexico 88003.