sign in sign up
<<
Home , Bathypolypus, Cirroteuthidae, Cirroteuthis, Cirrothauma, Cirrothauma murrayi, Eledone

MALACOLOGIA, 1988, 29(1): 295-307

THE BIOGEOGRAPHY OF THE DEEP-SEA OCTOPODA

Gilbert L. Voss

Rosenstiel School of Marine and Atmospheric Science, University of Miami, Miami, Florida 33149, U.S.A.

ABSTRACT

The deep-sea octopods are in both octopod suborders, the Cirrata with three families, eight

genera and 29 , and the with three octopodid subfamilies, 1 1 genera and 48 species. In addition, some families in the Incirrata contain deep-sea pelagic species. Only benthic or "near benthic" species are dealt with m this paper. Among the Cirrata, only the contain species with a multi-ocean distribution. murrayi is possibly cosmopolitan, occurring from polar seas to the depths of the tropics. muellen has an amphiboreal distribution in the North Atlantic and North Pacific. In the monotypic Stauroteuthidae, Stauroteulhis syrtensis is known only from the temperate western North Atlantic. The Opisthoteuthidae are represented by two genera.

Opisthoteuthis with 10 species is the shallowest-dwelling of the cirrates, with all but three species living between 100 and 1000 m. The is distributed throughout the oceans except

in high latitudes. Gnmpoteuthis, with 13 species, occurs in all oceans but no species is recorded from more than one ocean. A specimen has been trawled from 7279 m. The deep-sea Incirrata are found among three subfamilies of the . In the Bathypolypodmae. has five species in the Atlantic, one off Japan, one in the Indian Ocean; Teretoclopus has two species in the Indian Ocean, and Benthoctopus has approximately 15 species distributed worldwide from the Antarctic to the Arctic, with individual species restricted to single oceans. Among the Pareledonmae, Pareledone, Vosseledone and Velodona are found only in the southern hemisphere, and Tetracheledone only in the western North Atlantic. Pareledone is the most speciose and has a circum-Antarctic distribution; the monotypic Vosseledone is known only from Brazil; the two species of Velodona occur off East Africa. Among the Graneledoninae, Graneledone has two species in the Southern Ocean, one in the North Atlantic, one in the North Pacific and one in the Panamic region. Thaumeledone and Bentheledone are confined to the Southern Ocean. Key words; biogeography, deep sea; Octopoda; Cirrata; Incirrata.

INTRODUCTION have a moderate to deep web and vestigial paired shell-remnants or stylets. They crawl The deep-sea octopods (defined as those about the bottom or swim by jet propulsion for octopods that live below the edge of the short distances. While there are several fam- continental shelf) are in both suborders, the ilies that have deep-sea pelagic octopods Cirrata and the Incirrata. The cirrates, or the (see Thore, 1949), only the benthic species finned octopods, represent the oldest evolu- are considered here. tionary lineage of the octopods (Voss, 1988). The cirrates are soft-bodied, semigelatinous In both orders, the benthic and "near benthic" females large result- with a pair of small to large fins, a produce eggs deep and sometimes complicated web and a ing in young that, we suppose, take up the relatively large, single internal shell. They adult habitat and mode of life immediately after hatching from the swim by medusoid action of the web, by jet egg. propulsion, by beating the fins or by a combi- One might expect that the genera and nation of all three (Pereyra, 1965; Roper & species of the deep-sea cirrates would have a Brundage, 1972). Although swimmers, they much broader distribution than would the live just above the bottom and are rarely slower moving, bottom crawling deep-sea captured in the mid-depths. incirrates. Such, however, is not the case The deep-sea benthic incirrates resemble, although our studies are limited by a paucity in general appearance, the typical shallow- of specimens and much confusion concerning water octopods. They are soft to firm bodied. identifications.

(295) 296 G. L. VOSS

TABLE 1 . List of cirrate octopods arranged accord- Cirroteuthis muelleri is known from the ing to Voss (1988) with depth of capture and northwestern Atlantic (Voss, unpublished), nomenclatural status. Baffin Bay to Iceland (Eschricht, 1836; Voss, unpublished), off Jan Mayen (Appellöf, Cirroteuthidae 1893) and from off Oregon Pearcy, in Cirroteulhis muellen Eschncht, 1836 0-2342 m (Voss &

Cirrothauma murrayi Chun. 191 1 1500—1500 m preparation). These data suggest an amphib- Stauroteuthidae oreal distribution (Fig. 1). Although a number Stauroleuthis syrtensis Verrill, 1879. 457-2463 m of species of fish show this amphiboreal dis- Opisthoteuthidae tribution, most are separated into Atlantic Gnmpoteuthis megaptera (Verrill, 1885) 1929-4710 m and

Grimpoleulhis umbellata (Fischer, 1883). 1 140-5274 m. Pacific subspecies (Ekman, 1953). The ap- Gnmpoteuthis pacifica (Hoyle, 1885) 4465 m parent absence of a median palliai adductor Gnmpoteuthis meangensis (Hoyle. 1885) 915-1098 m in Atlantic specimens (Robson, 1932) is erro- Gnmpoteuthis caudani (Joubin. 1896) 650 m Nomen dubium neous since examination of well-preserved Gnmpoteuthis plena (Verrill, 1885) 1964 m. WALTHER HERWIG specimens from the Gnmpoteuthis hippocrepium (Hoyle, 1904) 3336 m North Atlantic proved that this structure was Nomen dubium present. No other characters have been Gnmpoteuthis mawsoni (Berry, 1917). 527-549 m. found Nomen dubium. by which to separate the two popula- Gnmpoteuthis gnmaldi (Joubin. 1903) 1804-1901 m tions. Because of the high northern occur- = ?) ( rence in the Atlantic, the distribution could be Gnmpoteuthis wuelken (Grimpe, 1920). 2057 m. continuous through the Arctic Ocean even Nomen dubium. though no high Arctic records are Gnmpoteuthis albatross/ (Sasaki, 1920). 487-1680 m known. The Gnmpoteuthis glacialis (Robson, 1930). 500 m. species has been taken from the surface (off Gnmpoteuthis bruuni Voss, 1982 250-360 m Greenland) to 2342 m. Opisthoteuthis agassizi Verrill. 1883. 922-2250 m has been mono- Opisthoteuthis depressa Ipma & Ikeda, 1895 128-1074 m graphed by Aldred et al. (1983) who gave all Opisthoteuthis pluto Berry, 1918 833 m. known records. Additional records in Fig. 1 Opisthoteuthis persephone Berry. 1918 549 m have been added from specimens in the Opisthoteuthis medusoides Thiele, 1915. 399 m University of Miami collections. This species Opisthoteuthis extensa Thiele, 1915 769 m. is known from the coasts of Opisthoteuthis californiana Berry, 1949 125-1 100 m Europe and Opisthoteuthis ¡aponica Taki, 1962 150 m northwestern Africa to off northeastern South Opisthoteuthis phillippi Oomen, 1976. 275-365 m America and along the eastern coast of North Uncertain status America from about New York to the Baha- Chunioteuthis ebersbachi (Grimpe, 1916) 1100 m and questionably from the Indian Chunioteuthis gilchnsti (Robson, 1924). 2562 m. mas Ocean. Froekenia clara Hoyle. 1904. 1016 m. Its normal depth range is 1500 to 3000 m. Cirroteuthopsis massyae Grimpe, 1920 1409 m One specimen was taken from an ice hole in

the Arctic Ocean. If all specimens prove to belong to a single species, this species will be CIRRATA the only widely distributed, nearly cosmo- politan, species among the deep-water octo- The Cirrata contains 29 species in 8 genera pods.

(Table 1). These are divided into three fami- lies based upon web and shell types: Cir- Stauroteuthidae roteuthidae, Stauroteuthidae and Opistho- teuthidae. Like the cirroteuthids, stauroteuthids are

fragile, soft-bodied animals. This family is

Cirroteuthidae represented only by syrtensis. It occurs from Nova Scotia (Verrill, 1879) along The cirroteuthids contain two monotypic the Atlantic coast of North America to north of genera represented by Cirroteuthis muelleh the Bahamas (Voss, unpublished) (Fig. 1). and Cirrothauma murrayi. All of the cir- Except for one questionable record, this spe- roteuthids are fragile, soft-bodied animals, cies occurs only in the western North Atlantic. especially Cirroteuthis. Meaningful measure- Its depth range is 457 to 2463 m. ments are exceedingly difficult to obtain from the specimens, and internal organs are rela- Opisthoteuthidae tively unstudied. Perhaps these genera will be found to be more speciose when sufficient As now understood, this family includes the specimens are available. majority of the cirrates: 22 species belong to DEEP-SEA OCTOPOD BIOGEOGRAPHY 297

FIG. 1 . Cirroteuthidae. Cirroteuthis muellen, • Cirrothauma murrayi. Stauroteuthidae. a .

the two genera and Opis- off northern Chile and Peru (Voss, 1982). G. thoteuthis. umbellata, the type species, is still relatively The genus Grimpoteuthis occurs through- unknown but occurs off Mauritania and the out the Atlantic and Pacific oceans and the Azores (Fischer, 1883; Robson, 1932). Sev- Antarctic (Fig. 2), living in depths from about eral specimens identified as this species 200 m to below 7000 m (Robson, 1932). The (Robson, 1932; personal observations) are identity of the majority of species is uncertain questionable. and many species are represented by single Verrill (1885) described G megaptera and specimens. As a result, the geographical G. plena from off New England. The latter is a range of many species is not yet established. nomen dubium, but G. megaptera, with many At present no species is known to be wide- misidentified specimens, may be widespread spread. in the North Atlantic. Nine species have been described and In the same year Hoyle (1885) described three are in manuscript (Voss & Pearcy, in two species taken by the CHALLENGER: G. preparation; Voss, in preparation). Brief in- pacifica from near the Coral Sea and G. spection of the GALATHEA collections sug- meangensis from north of the Celebes. No gests that this number may be doubled when subsequent specimens identifiable to either these specimens are reported upon. of these two species have been reported. Grimpoteuthis glacialis is known from south In 1904 Hoyle also described G. hip- of the Antarctic Convergence from South pocrepium from off the Gulf of Panama. The Georgia, the Palmer Peninsula and the Ross validity of all three species is questionable Sea (Robson, 1930; unpublished ELTANIN because of the brief descriptions and poor records, University of Miami). It probably oc- illustrations. curs throughout the Southern Ocean south of Grimpoteuthis albatrossi was described by the convergence. A second undescribed Sasaki (1920) from south of the Aleutians Southern Ocean species occurs in the Scotia from specimens taken by the United States

Sea (WALTHER HERWIG specimens, Voss, Fish Commission steamer ALBATROSS. It in preparation). Grimpoteuthis bruuni occurs also occurs off Japan (Sasaki, 1929). 298 G. L. VOSS

FIG. 2. Opisthoteuthidae Grimpoteuthis bruuni, • G glacialis, a G. sp. A, a G. sp. , G. sp OG meangensis, • G. pacifica. A G. umbellata, Lj G. megaptera, G p/ena. G hippocrepium, t> G. grimaldi, ® G. albatrossi

The beautifully illustrated and well de- Opisthoteuthis califomiana occurs in north- scribed G. grimaldi Joubin, 1903. is a prob- ern California (Berry, 1949) and Washington lem. Since its original description from off the (Pereyra, 1965) waters and has since been Azores, additional specimens have been as- reported by Taki (1963) from off Honshu. Two signed to this species in the literature (Joubin, other species, O. depressa and O. japónica,

1 920; Bruun, 1 945). My study of the optic lobe also occur in Japanese waters. of cirrates (unpublished) indicates, however, Thiele (1915a) described O. extensa from a that G grimaldi is probably an Opisthoteuthis VALDIVIA station southwest of Sumatra and and may even be O. agassizi. 0. medusoides from off Zanzibar. In 1918 The genus Opisthoteuthis occurs in moder- Berry described O. persephone and pluto ate depths, mostly between 100 and 1000 m from captures by the ENDEAVOUR in the in tropical and temperate seas (Fig. 3). It Great Australian Bight. Many more speci- ranges from about 50 N to 45 S. Only three mens of the latter two species have since species have been taken in depths greater been taken. Both apparently are limited to than 1000 m; the deepest record is 2250 m Australian waters. (Verrill, 1885). The last species described is Opistho-

If identifications are correct, Opisthoteuthis teuthis philippi from off southwest India agassizi occurs in the eastern Atlantic from (Oomen, 1976). southwest of Ireland (Chun, 1 91 3) to off Spain, The paucity of records prevents generaliza- and the western Mediterranean (Morales, tions on the biogeography of the individual 1959), the Gulf of Guinea (Voss, in prepara- species. tion) and south to off southwest Africa (Adam,

1 962). Its distribution in the W Atlantic appears to be more limited, ranging from northeast of INCIRRATA the Bahamas (Univ. of Miami collections) and Gulf of Mexico (Voss, 1956) to northeastern The incirrate deep-sea octopods contain 48

Brazil (Univ. of Miami collections). species distributed unevenly in 1 1 genera (Ta- —

DEEP-SEA OCTOPOD BIOGEOGRAPHY 299

,/^ • ..»**~ £ ,

FIG. 3. Opisthoteuthidae. (J Opislholeulhis agassizi, @ depressa, O O. extensa. O. medusoides, < . japónica, • . californiens, Ü . philippi, . p/ufo. . persephone.

ble 2). I have recently (Voss, 1988) grouped Denmark Straits, Iceland, northern Great these in three subfamilies: Bathypolypodinae, Britain and the North Sea to Spitzbergen and Pareledonmae and Graneledoninae. In gen- the Arctic Ocean north of European Russia eral, the deep-sea incirrates do not occupy as (Nesis, 1982; Kumpf, 1958). It appears to be great depths as do the cirrates, ranging from an Atlantic Boreal and Arctic species. B. about 200 m to nearly 4000 m but with many arcticus is also confused with B. faeroensis, living to about 500 m. All, however, show mod- recently reinstated by Toll (1985), and S. ifications to reflect a deep-sea habitat. proschi Muus, 1962. Further work may show Robson's (1932) classification of the incir- that true B. arcticus may have a more limited rates. in my opinion, cannot be maintained. distribution. Some major shifts are made here, based B. sponsalis occurs along the Atlantic partly on the number of longitudinal sucker Spanish coast southward to Senegal (Adam, rows and the presence or absence of an ink 1960) and the Cape Verde Islands (Fischer & sac and crop. Fischer, 1892) and the western Mediterra- nean Basin (Wirz, 1955; Morales, 1958) in Bathypolypodinae depths from 400 to 930 m. Its distribution reflects the typical Mauritanian— Lusitanian This subfamily contains three genera, Mediterranean pattern (Ekman, 1953: 80). Bathypolypus. Teretoctopus and Benthoc- B. salebrosus is found off northern Honshu, topus. All have biserial arm suckers and no the Kurile Islands and in the Okhotsk Sea ink sac. (Sasaki, 1929). It lives between about 450 Bathypolypus contains five species of and 800 m. which three are found in the North Atlantic B. valdiviae is distributed from near (Mangold-Wirz, 1963; Toll, 1985), one off Durban, South Africa, to off Walfisch Bay, Japan (Sasaki, 1929) and one in the Indian Namibia, in depths from 400 to 800 m (Thiele, Ocean (Thiele, 1915b) (Fig. 4). B. arcticus is 1915b; Sanchez & Moll, 1984). Its range found in the North Atlantic from the Straits of encompasses the Cape Province and the Florida (in deep water) north to Baffin Bay, Namaquan Province, both dominated by the 300 G. L VOSS

TABLE 2 List of mcirrate octopods arranged ac- the Andaman Sea to the upper Arabian Sea cording to Voss (1988) with depths of capture and (Robson, 1932) between 500 and 1281 m. nomenclatural status. Little is known of these two Indian Ocean species other than Robson's (1932) com- Bathypolypodinae ments. Bathypolypus ardicus (Prosch, 1849) To 1543 m. (P & H Fischer. 1892) 930-1250 m The genus Benthoctopus occurs from the Bathypolypus faeroensis (Russell, 1909) 480-1030 m Arctic Ocean to the Southern Ocean (Fig. 5). Bathypolypus valdiviae (Thiele, 1915) 403-860 m Specific identifications are difficult; there are Bathypolypus salebrosus (Sasaki. 1920) 487-805 m numerous is Bathypolypus proschi Muus, 1962 Nomen dubium synonyms and the literature filled Teretoctopus indicus Robson, 1932 996 m with mtsidentifications. All of the 15 or so Teretoctopus alcocki Robson, 1932 353-1281 m species have somewhat limited ranges. They Benthoctopus piscatorum Verrill, 1879 220-2492 m live from about 200 m to in excess of 3000 m. Benthoctopus levis (Hoyle. 1885) 13-137 m Benthoctopus piscatorum is Benthoctopus ¡anuari (Hoyle. 1885) 350-732 m an Atlantic Bo- Benthoctopus ergasticus (P &H Fischer. 1892) 470-915 m real species found from New England to the Benthoctopus lothei (Chun, 1914) 1365 m British Isles, North Sea and off Norway Benthoctopus hokkaidensis (Berry, 1921) 487-920 m (Robson, 1932). It may also be found in the Benthoctopus abruptus (Sasaki, 1920) 1074 m High Arctic and Spitsbergen Benthoctopus pseudonymus (Grimpe. 1922) 1599 m. around (Robson, Benthoctopus berryi (Robson. 1924) 2196 m 1932) but its northern and eastern ranges are Benthoctopus eureka (Robson. 1929) Depth unknown not known. B. ergasticus is in the Eastern Benthoctopus sibincus Lonmg. 1930 Stomach of walrus Atlantic southwest of Ireland (Massy, 1909) Nomen dubium and the Mauritanian upwelling area Benthoctopus thielei Robson. 1932 On shore (Ficsher Benthoctopus prolundorum Robson, 1932 741-3431 m & Fischer, 1892). B. ¡anuari is strictly Atlantic, Benthoctopus magellamcus Robson, 1932 145 m distributed from the Gulf of Mexico (Voss, Benthoctopus oregonae Toll. 1981 640-1080 m 1956) to middle Brazil (Hoyle, 1886). Toll Pareledonmae described B. the Pareledone charcoti (Joubin. 1905) 18-595 m (1981) oregonae from mid- Pareledone turqueti (Joubin. 1905) 25-1116m dle of the former's range. Pareledone harnsoni (Berry. 1917) 494-655 m The only other species with sufficient infor- Nomen dubium mation to permit speculation on its range is B. Pareledone adeliana (Berry, 1917) 527-549 m Nomen dubium magellanicus from the Valdés Peninsula, Ar- Pareledone polymorpha (Robson. 1930) 15-1 116 m gentina (Ré, personal communication) to Velodona togata togata Chun. 1915 748 m. Tierra del Fuego (Robson, 1932) and appar- Velodona togata capensis Robson, 1929 402-457 m ently N to Chiloe Island (Voss, unpublished). Vosseledone charrúa Palacio. 1978 10-200 m Greater deep-trawling efforts will certainly Tetracheledone spinicirrus Voss, 1955 183-544 m. moschala (Lamarck, 1798) Above 300 m reveal more species and wider distributions. Eledone cirrhosa (Lamarck. 1798) 10-800 m There are four species off Japan and Voss & Eledone caparti Adam, 1950 60-170 m Pearcy (in preparation) have four new spe- Eledone thysanophora Voss, 1962 Tide pool cies from off are Voss, 1964 60-170 m Oregon. But records few and Graneledoninae scattered widely. Surprisingly, few are known Craneledone verrucosa (Verrill, 1881) 852-2297 m from the Southern Ocean. Graneledone challenger: (Berry, 1961) 1153-1867 m Graneledone verrucosa media (Joubin, 1918) 1458 m Nomen dubium Pareledonmae Graneledone antárctica Voss, 1976 2341 m Graneledone macrotyla Voss, 1976 1647-2044 m This subfamily contains the deep-sea gen- Graneledone boreopacitica Nesis 1982 1165-1500 m era Pareledone, Velodona, Tetracheledone Thaumeledone brevis (Hoyle, 1885) 800-3931 m and the closely related Thaumeledone gunten Robson. 1930 410 m. shallow-water genus Bentheledone rotunda (Hoyle. 1885) 3596 m Vosseledone and the more distantly related Bentheledone albida (Berry. 1917) 3111 m genus Eledone. Eledone is retained only pro-

visionally in this subfamily as it shares the common characters of uniserial arm suckers, Agulhas Current, which probably is responsi- an ink sac and a crop. However, Robson ble for this distribution. (1932) and others have held that it is not truly The genus Teretoctopus contains only two related to the other members of the subfamily. species, T. indicus and T. alcocki. T indicus The species of Pareledone are all from the has been reported from the pearl grounds off Southern Ocean (Fig. 6), living primarily south Ceylon (Robson, 1932) and the upper Ara- of the Antarctic Convergence (Robson, 1932; bian Sea in 1000 m (Massy, 1916b). T. Dell, 1959; Taki, 1961; Voss, in preparation) alcocki is known from the Bay of Bengal and although P. turqueti was recorded off Para, DEEP-SEA OCTOPOD BIOGEOGRAPHY 301

FIG. 4. Bathypolypodinae. = . sponsalis. ill B. salebrosus, B. valdiviae, • laeroensis. Teretoctopus indicus, A T. alcocki.

Brazil, by Massy (1916a) from deep trawling Eledone was originally thought to be Med- of the TERRA NOVA expedition. It was also iterranean with only two species, cirrosa and taken in Antarctic Deep Water. All of the moschata. Later a West African species, E. species are circumpolar, living from tide pools capartl, was described (Adam, 1950) from the in the Ross Sea to below 2000 m, with Guiñean Province. More recently another temperature apparently the controlling factor species, E massyae, was added from off (Voss, in preparation). Brazil (Voss, 1964) where an additional, Velodona togata has been taken twice from undescribed species occurs (Haimovici, in off Zanzibar (Chun, 1915; Voss, unpublished). preparation), and finally one from southwest A subspecies, capensis, occurs in the Cape Africa, E. thysanophora, was described Province (Robson, 1932). This species is (Voss, 1962). A new species from Australia probably widely distributed along the East Af- has recently been found (Lu, personal com- rican coast in depths between 400 m and 800 munication). Apparently the genus is circum- m (British Museum collections). Vosseledone tropical and warm temperate in distribution charrúa has a similar distribution along the (Fig. 7). It is found in moderate depths, mostly eastern coast of South America in shallow wa- above 300 m with a single record to 800 m. Its ter, only 10 m to 200 m (Palacio, 1978), but it true position and relationship with both the is closely related to the deep-sea Pareledone deep-water and shallow-water genera has yet in many of its features. Tetracheledone to be solved. splnicirrus is known only from north Florida to the Gulf of Mexico and the northern Caribbean Graneledoninae in 183 to 544 m (Voss, 1956). Vosseledone, Velodona and Tetracheledone, while occur- ring in shallow to moderate depths, in all other This subfamily contains the genera respects are deep-water animals closely re- Graneledone, Thaumeledone and Benthele- sembling the genus Graneledone to which, on done (Fig. 8). They are all deep dwellers, superficial examination, many of them would living between 1 000 m to in excess of 3500 m. seem to belong. The genus Graneledone shows a somewhat 302 G. L. VOSS

FIG. 5. Bathypolypodinae. ¡s Benthoctopus piscatorum, . ergasticus. Si B. magellamcus. B. ¡anuan, t> ® oregonae. eureka, I J . berryi. . pseudonymus, . fuscus, . hokkaidensis. S. abruptus, • . profundorum, . levis, S thielei. ir , sp. , ® . spp. , , , D.

FIG. 6. Pareledoninae. Pareledone polymorphe, • P. turqueti, "7 P. senoi. A P hamsoni, ü P. charcoti. DEEP-SEA OCTOPOD BIOGEOGRAPHY 303

FIG. 7. Pareledoninae. Velodona lógala. • Vosseledone charrúa, Tetracheledone spinicirrus. Eledone massyae, E. nigra, U A E thysanophora. E. arrosa. . E. moschata. . caparti.

bipolardistributionwithG. verrucosa. G.boreo- DISCUSSION pacifica and G. sp. A (the latter two perhaps identical), in N waters (Voss & Pearcy, in Unfortunately the systematics of the deep- preparation), and G. antárctica, G. macrotyla sea octopods, and in particular the cirrates, and G. challengeri (Voss, unpublished) in the have been sadly neglected. Robson's (1932) Southern Ocean and north of New Zealand. monographic study of the cirrates was the last

An unidentified specimen, examined by me attempt to make order out of the group. It did from off the Gulf of Panama, may represent not succeed, partly because of Robson's an additional species (sp. B). mental decline and partly because of the poor The genus Thaumeledone contains two quality and quantity of available collections. nominal species, 7". brews and T. gunteri. Un- The situation has not altered appreciably published records of T. brevis show that it is since then because, while considerable addi- distributed from the Palmer Peninsula to the tional material has accumulated, the existing Ross Sea and is probably circumpolar south of types have deteriorated to such an extent that the Antarctic Convergence. T. gunteri is prob- reliance must be given to the original, often ably a synonym and was reported from off inadequate, descriptions. The next few years,

South Georgia. It was taken in 41 m (Robson, however, should show progress in cirrate

1 930) while T. brevis lives between 800 m and systematics and biogeography as a result of 3931 m (Robson, 1932; Voss, unpublished). study of the collections made by the USNS Bentheledone consists of two described ELTANIN in the Antarctic and the Danish species, B. rotunda, and B. albida, with two GALATHEA on her round-the-world, deep- undescribed species (Voss, in preparation). trawling expedition, all now at Miami. All occur south of the Antarctic Convergence Among the cirrates, several distributional in depths from 3000 m to 3596 m. They are patterns are evident, even at this time. Cir- probably circumpolar. Despite the extensive rothauma is found in both the Atlantic and trawling effort in the Southern Ocean, no Pacific Oceans. Stauroteuthis is known defi- specimens of Bentheledone have been taken nitely only from the western North Atlantic, from moderate depths. including the Caribbean Sea. 304 G. L VOSS

FIG 8 Graneledoninae Graneledone verrucosa. • G sp. A, A G. challengeri, A G. sp. B, Ü G. antárctica, QG macrotyla, iiThaumeledone brevis, 7". gunteri, Bentheledone rotunda, à S, albida.

Grimpoteuthis is widely distributed but does The sole southern exception is B. valdiviae, not occur in the Arctic, living no further north which is found off South Africa in the Agulhas than about 50N. It is the only genus of Current or the southern end of the cold cirrates that has representatives in the South- Benguela Current. Teretoctopus is limited to ern Ocean south of the Antarctic Conver- the deep underlying cold water of the tropical gence. Indian Ocean. Opisthoteuthis is known only from 45 N to Benthoctopus is the most widely distributed 45°S. Living in shallower depths, its distribu- of the Bathypolypodinae but only four of the tion reflects the shallow-water hydrographie nineteen species are found in the southern regimes. hemisphere with two south of the Antarctic

The Incirrata have fared little better. Again Convergence. Of the remaining 15 species, Robson (1932) brought together all of the one is widely distributed in the North Atlantic known information on the group but it is a Boreal to the Subarctic and perhaps to the hodge-podge of misidentification by previous High Arctic. Two are tropical and nine are authors and reflects a lack of comparative northern temperate. morphological data. Many of the species de- The Pareledoninae comprises two groups scribed both before and after Robson were showing distinctly different distributional pat- done without referring to the types, and since terns. Pareledone is strictly Southern Ocean;

1932 are based upon Robson's confused all of the known species have their main decisions. Again, many of the types are in concentrations south of the Antarctic Conver- poor condition or are immature specimens gence and are circumpolar. The few catches lacking important specific characters (Voss, in north of the convergence, one off Montevideo preparation). and the others near the Falkland Islands, The Bathypolypodinae contains primarily were in cold Antarctic Deep Water. northern hemisphere genera and species. The remaining genera of the Pareledoninae Five of the six species of Bathypolypus occur are all Atlantic and Indian Ocean in occur- from about 20 N to the Atlantic and Pacific rence. Their distribution reflects the general Subarctic with one extending to Spitsbergen. surlace hydrographie regimes and all have .

DEEP-SEA OCTOPOD BIOGEOGRAPHY 305 restricted distributions, reflecting their moder- contribution from the Rosenstiel School of ate-depth, continental slope habitats. Marine and Atmospheric Science, University The Graneledoninae are in the western of Miami. North and South Atlantic, Pacific and South- ern Ocean. Graneledone has a somewhat bipolar distribution with two species in the LITERATURE CITED colder North Atlantic and North Pacific Subarctic Water and three in Southern ADAM, W., 1950. Notes sur les Céphalopodes. Ocean. However, one species occurs north of XXII. -Deux nouvelles espèces de la côte New Zealand and one off the Gulf of Panama, africaine occidentale. Bulletin de l'Institut royal des Sciences naturelles de Belgique. 26(45): both in deep water. 1-9. The genera Thaumeledone and Benthele- ADAM, W., 1960. Les Céphalopodes de l'Institut done contain the deepest dwelling of all the Français de l'Afrique Noire. II. Bulletin de l'Institut Incirrata and are found in the Southern Ocean Français d'Afrique Noire. 22(2): 465-51 1 south of the Antarctic Convergence with only ADAM, W., 1962, Céphalopodes de l'Archipel du one record in Antarctic Deep Water off Argen- Cap-Vert, de l'Angola et du Mozambique. tina. Memorias do Junta de Investigaçôes do Ultra- Of the 77 species of deep-water octopods, mar. Lisboa, ser. 2, 33: 9-64, 2 pi. R. G., 25 percent are confined to the Southern ALDRED, NIXON, M. & YOUNG, J. Z , 1983, Cirrothauma murrayi Chun, a fmned Ocean in Antarctic Water; 22 percent are octopod. Philosophical Transactions of the Royal limited to Atlantic and Pacific Subarctic Wa- Society of London, ser. , 301: 1-54. ter. Ten percent may be considered shallow APPELLÖF, ., 1893. Teuthologische Beiträge III inhabitants to moderate depth of the warm Bemerkungen über die auf norwegischen water regions of the world. The remaining 43 Nordmeer-Expedition (1876-78) gesammelten percent occupy the great depths below the Cephalopoden. Bergens Museum Aarbog. 1892: tropical and warm temperate regions of the 1-13. world, living in cold deep water derived from BERRY, S. S., 1918, Report on the Cephalopoda. either the Antarctic or the Arctic Deep or Biological Results of the Fishing Experiments Intermediate Water. carried on by the F.I.S. "Endeavour,'' 1909-14, 4: 203-298. The high percentages of species occurring BERRY, S. S., 1949, A new Opisthoteuthis from in the Southern Ocean is probably due to the

the eastern Pacific. Leaflets in Malacology, 1 : high productivity of waters the the between 23-26. Antarctic Convergence and the Antarctic con- BRUUN, A.F.. 1945, Cephalopoda. Zoology of Ice- tinent, with somewhat lower levels northward land. 4(64): 1-15. to the Subtropical Convergence. The same CHUN, O, 1913, Cephalopoda from the "Michael situation is found with regard to the species in Sars" North Atlantic Deep Sea Expedition 1910. the North Atlantic and North Pacific in Repon" of the Scientific Results of the "Michael Subarctic Water. The dearth of species in the Sars'' North Atlantic Deep Sea Expedition 1910. Arctic Ocean may well correlate with the Zoology. 3(1): 1-28, 2 pis. CHUN, C, 1915. Octopoda Leach, 1818 In: lower productivity of the Arctic Ocean in gen- CHUN, C, The Cephalopoda. Part II; Myopsida. eral. If one compares the distributions of the Octopoda. Scientific Results of the German deep-water octopods in the world oceans with Deep-Sea Expedition on the Steamer "Valdivia'' the of ocean productivity map as depicted by 1898-1899. 18: 373-378. Ebeling (1962), one finds a high correlation DELL, R. K., 1959, Cephalopoda. BANZ. Ant- with species richness and high productivity arctic Research Expedition 1929-1931. 2 (above 100-200 gC/m /year) with few genera B.A.N.Z.A.R.E. Scientific Reports, ser. (Zool- and species in less productive waters. ogy and Botany). 8(4): 89-106.

EBELING, A. W., 1962, Melamphidae I. Systemat- ics and zoogeography of the species in the ACKNOWLEDGEMENTS bathypelagic fish genus Melamphaes Günther. Dana-Report 58: 1-64. EKMAN, S., 1953, Zoogeography of the Sea. N. A. Voss read the manuscript critically Sidgwick & Jackson, London. and offered many suggestions. I have bene- ESCHRICHT, D. F., 1836, Cirroteuthis Müllen. fited from the study of numerous specimens eine neue Gattung der Cephalopoden Bilden. in many museums in this country and Europe, Nova Acta Academiae Caesarea Leopoldino too numerous to mention here but to whom Carolinae Germanicae Naturae Cunosorum. thanks are extended. This paper is a scientific Halle. 18(2): 627-634. .

306 G. L VOSS

FISCHER, P., 1883, Note préliminaire sur une of the Marine Biological Association of India. 18 nouvelle espèce du genre Cirroteuthis. Journal 368-374. de Conchyliologie. 31: 402-404. PALACIO, F. J., 1978, Vosseledone charma: a FISCHER, P. & FISCHER H., 1892, Diagnoses new Patagonian (Octopodidae) with d'espèces nouvelles de Mollusques Céphalo- notes on related genera. Bulletin of Marine Sci- podes. Journal de Conchyliologie, 75: 319. ence. 28: 282-296 PEREYRA. T., HOYLE, W. E , 1885, Diagnoses of new species of W 1965, New records and obser-

Cephalopoda ... Pari I. The Octopoda. Annals vations on the flapiack devilfish, Opisthoteuthis and Magazine of Natural History, ser. 5,: californiana Berry. Pacific Science. 19: 427-441

222-236. ROBSON, G. C, 1930, Cephalopoda, I. Octopoda.

HOYLE, W. E., 1886, Report on col- Discovery Reports. 2: 371-402, 2 pl.

lected by H.M.S. CHALLENGER during the ROBSON, G. , 1932, A monograph of the recent

years 1 873-76. Report on the Scientific Results Cephalopoda ... Part II The Octopoda (exclud- of the Voyage of H.M.S. CHALLENGER ing the Octopodinae). Bulletin of the British Mu- 1873-76. Zoology. 16(44): 1-245. seum (Natural History), pp. i-xi, 1-359, 6 pl. HOYLE, W. E., 1904, Reports on the ROPER, F E., & BRUNDAGE, JR.. W L, Cephalopoda. In: Reports on the scientific re- 1972, Cirrate octopods with associated deep-sea organisms, new biological data based on deep sults of the expedition to the eastern Pacific . . . ALBATROSS. Bulletin of the Museum of Com- benthic photographs (Cephalopoda). Smithson- parative Zoology, Harvard, 43: 1-71. ian Contributions to Zoology. 121: 1-46. JOUBIN, L. 1903, Sur quelques Céphalopodes SANCHEZ. P & MOLI, B. 1984, Cefalópodos de recueillis pendant les dernières campagnes de las costas de Namibia (Atlántico sudoriental). S A. S. le Prince de Monaco (1901-1902). Resultados Expediciones Científicas. Barcelona. Comptes Rendus des Seances de l'Académie 12: 3-22.

des Sciences. Paris. 136: 100-102. SASAKI, M , 1920, Report of Cephalopoda col- lected during 1906 by the U.S.B.F. Steamer JOUBIN, L , 1920, Céphalopodes provenant des Albatross' in the N Pacific. Proceedings of campagnes de la PRINCESS-ALICE (1898- W. the United 1910) Resultats des Campagnes Scientifiques States National Museum. 57: e 163-203. acomplies sur son yacht par Albert I ' Prince

SASAKI. M , 1929, A monograph of the dibran- Souverain de Monaco, fase, 54:1-95, 16 pi chiate cephalopods of the Japanese and adja- KUMPF, H., 1958, A study of the Bathypolypus cent waters. Journal of the College of Agricul- arcticus-bairdii-lentus-obesus complex of the At- ture. Hokkaido University, vol. 20, suppl., 357 lantic (Cephalopoda. Octopoda) Master's thesis, p., 30 pl. University of Miami, 135 p. TAKI, I., 1961, On two new eledonid octopods from MANGOLD-WIRZ, K., 1963, Biologie des Cépha- the Antarctic Sea. Journal of the Faculty of lopodes benthiques et nectoniques de la Mer Fisheries and Husbandry. Hiroshima Uni- Catalane. Vie et Milieu, suppl. 13: 1-283, 4 pi versity. 3(2): 297-316, 3 pl. MASSY. A. L., 1909. The Cephalopoda Dibran- TAKI, I., 1963. On four newly known species of chiata of the coasts of Ireland. Fisheries of octopods from Japan. Journal of the Faculty of Ireland. Scientific Investigations. 1907: 3-39, 3 Fisheries and Animal Husbandry. Hiroshima Uni- pl- versity. 5: 57-93.

MASSY, A. L ., 1916a, Part II Cepha- THIELE, J., 1915a, Cirroteuthidae In: CHUN, lopoda. British Antarctic ( "Terra Nova") Expedi- ed.. The Cephalopoda. Part II Myopsida. tion. 1910. Zoology. 2: 141-176. Octopoda. Scientific Results of the German

MASSY, A. L , 1916b, Cephalopoda of the Indian Deep-Sea Expedition on the Steamer "Valdivia" Museum. Records of the Indian Museum. 1898-1899, 18: 421^*26. Calcutta. 12(5): 185-247, 2 pl.

THIELE, J , 1915b, Polypus Lamarck, 1799. In:

MORALES, E . 1958, Cefalópodos de Cataluña. I. CHUN, C. ed. The Cephalopoda, Part II.

Investigación Pesquera, Barcelona, 1 1 : 3-32, Myopsida. Octopoda. Scientific Results of the MORALES, E., 1959, Sobre la presencia de German Deep-Sea Expedition on the Steamer Opisthoteuthis agassizi Vernll, en el Mediter- "Validivia" 1898-1899, 18: 378-379. ráneo. Investigación Pesquera, Barcelona, 15: THORE. S.. 1949. Investigations on the DANA 113-123. Octopoda. Dana-Report, 33: 1-85. MUUS. B. J., 1962, Cephalopoda, The Godthaab TOLL, R ., 1981 , Benthoctopus oregonae. a new Expedition 1928. Meddelelser Grönland. 81: species of octopod (Mollusca: Cephalopoda) 4-23. from the southern Caribbean with a redescription

NESIS, . N.. 1 982, Brief diagnoses of the cepha- of Benthoctopus januaru (Hoyle, 1885). Bulletin

lopod mollusks of the world ocean Light and of Marine Science. 31 : 83-95. Food Industries, Moscow, 358 p. (In Russian). TOLL, R ., 1985, The reinstatement of Bathy- OOMEN, V. P., 1976, A new species of the genus polypus faeroensis (Russell, 1909) (Octopoda: Opisthoteuthis Verrill, 1883 (Cephalopoda: Mol- Bathypolypodmae) Proceedings of the Biologi- lusca) from the southwest coast of India. Journal cal Society of Washington. 98: 598-603. DEEP-SEA OCTOPOD BIOGEOGRAPHY 307

VERRILL, A. E-, 1 879, Notice of recent additions to octopod, Eledone massyae. Bulletin of Marine the marine fauna of the eastern coast of North Science of the Gulf and Caribbean, 14:511-516. America. No. 7. Brief contributions to zoology VOSS, G. L., 1982, Gnmpoleuthis bruuni. a new from the museum of Yale College No. 44. Amer- species of finned octopod (Octopoda: Cirrata) ican Journal of Science and Arts, ser. 3, 18 from the southeastern Pacific. Bulletin of Marine (108): 468-470. Science. 31: 426-433. VERRILL, A. E., 1885, Third catalogue of Mollusca VOSS, L, Evolution recently added to the fauna of the New England G 1988, and phylogenetic relationships of deep-sea octopods (Cirrata and coast. . .. Transactions of the Connecticut Incirrata). In: R. Academy. 6: 395-452. TRUEMAN, E & CLARKE, M. R-, eds.. The Mollusca: VOSS, G. L„ 1956, A review of the cephalopods of Paleontology and the Gulf of Mexico. Bulletin of Marine Science of Neontology of Cephalopods, Academic Press. vol 12: 253-291 the Gulf and Caribbean, 6: 85-178

VOSS, G. L, 1962, South African cephalopods. WIRZ, , 1955, Contribution a l'étude des Octo-

Transactions of the Royal Society of South Af- podes de profondeur. I, Bathypolypus sponsalis rica. 36: 245-272. (P. et H. Fischer) espèce commune dans la VOSS, G. L., 1964, A note on some cephalopods partie ouest de la Méditerranée. Vie et Milieu, 6: from Brazil with a description of a new species of 129-147.