PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON l09(2):286-298. 19%
Occurrence of anomuran crabs (Crustacea: Decapoda) in hydro thermal vent and cold-seep communities: a review
Pierre Chevaldonne and Karine Olu (PC & KO) Laboratoire d'Ecologie Abyssale, IFREMER, B.P. 70, 29280 Plouzane, France; (PC) Present address: Institute of Marine and Coastal Sciences. Rutgers University, Dudley Road, Cook College. New Brunswick, New Jersey 08903, U.S.A.
Abstract.—Crabs of the family Lithodidae are frequently encountered in the vicinity of deep-sea hydrothermal vents and cold-seeps. Together with crabs of the families Galatheidae and Chirostylidae, they are the main contributors to the scavenging/predatory fauna of these highly productive areas, and a potential vector for the export of organic carbon to the surrounding deep-sea commu nities. A review of the literature indicates that anomuran crabs have been re ported from such environments since their discovery, and occur virtually any where a reducing habitat is found. These three families are represented by at least eight genera, with at least 14 species occurring in hot vent areas, and eight in cold-seep associated communities.
Two species of lithodid crabs have re Vents and seeps are environments where cently been reported from the south Bar biomass and biological production are high bados accretionary prism cold-seeps at compared to that of the surrounding abyssal depths of 1200-1700 m (Macpherson plains, due to utilization of the expelled re 1994). In addition to the description of a duced compounds by chemoautotrophic mi new species, Paralomis arethusa, Macpher croorganisms that constitute the first step of son mentioned that his new species and Li- a trophic web independent from sea-surface thodes manningi Macpherson, 1988, were production. Although organisms exclusive the first records of the family Lithodidae to ly dependent on fluid emissions usually be found in hydrothermal vent and cold- dominate these communities and live close seep areas. However, a review of the liter to the expelled fluids, other inhabitants of ature showed that occurrence of lithodid the neighboring deep-sea environment are crabs in such environments has been known attracted by the high food supply and enter at least since 1985 (Suess et al. 1985). The these environments. Like anomurans, most taxonomic literature on deep-sea hydrother of these organisms are filter-feeders, scav mal vent and cold-seep organisms often engers, or predators. As top predators/scav lacks ecological data, and ecological sur engers, anomurans are rivalled only in some veys frequently present vague taxonomic cases, in particular by the hydrothermal-en- information. This review is intended to demic brachyuran crab family Bythograei- dae. demonstrate that the Anomura are a partic ularly well represented group in deep-sea Infraorder Anomura chemosynthetically-based ecosystems, and Superfamily Paguroidea (sensu Forest to present a literature survey of the species 1987) and accompanying information relevant to biogeographic studies. This superfamily comprises the Lithodi dae and two families of hermit crabs. Pa-
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• iiHMHaaiiiHi VOLUME 109. NUMBER 2 287 guridae and Parapaguridae. Only one her tionary prism, and was identified by Mac mit crab has been reported to date from pherson (1994). The species is known from cold seeps of the Barbados accretionary depths of 640-777 m in the Caribbean prism, at depths of 1000-2000 m (K. Olu, (Macpherson 1988). pers. obs.), and another one from Monterey Two species of lithodid crabs found at Bay seeps at 600-1000 m (J. P. Barry, pers. seeps of the Monterey Bay, at 600-1000 m, comm.). The Barbados specimen seems to have been assigned tentatively to the genus belong to the genus Parapagurus (M. de Lithodes (J. P. Barry, pers. comm.). These Saint Laurent, pers. comm.). Although the crabs are not considered as seep-endemics. Monterey Bay hermit crab has not been identified, it is possible that it is also a par- Genus Lopholithodes Brandt, 1848 apagurid, which are common at these Lopholithodes foraminatus (Stimpson, depths. 1859) As previously mentioned, Carey et al. Family Lithodidae (1988) observed dense aggregations of Lo The first report of the occurrence of Lith pholithodes foraminatus while exploring odidae in hydrothermal vent or cold-seep apparently extinct seepage areas of the up habitats, is a "large crab," clearly a lith per Oregon subduction zone, at a depth of odid, illustrated in a sketch of the benthic ca. 250 m. There is no mention of this spe community associated with cold-seeps cies at active seeps, but it could be the same along the lower Oregon subduction zone at species as the "large crab" observed at 2037 m depth (Suess et al. 1985, Kulm et 2037 m by Suess et al. (1985). However, al. 1986). Subsequently, Carey etal. (1988) this occurrence would be much deeper than observed Lopholithodes foraminatus from a the known depth range of L. foraminatus, shallower part of this subduction zone. which is known from British Columbia to Whether the "large crab*' and L. forami southern California, at 0-547 m (Hart natus are the same species is not stated. 1982). Tunnicliffe & Jensen (1987) proposed that the deeper water species from the lower Genus Neolithodes Milne Edwards & zone could be the same Paralomis sp. they Bouvier, 1894 found at hydrothermal vents in the Juan de Neolithodes diomedeae (Benedict, 1894) Fuca Ridge. This species represents the first reported In total, four lithodid crabs have been occurrence of Lithodidae in hydrothermal identified to the species level from cold- vents. Grassle (1986) described it as the seep areas, and the status of four others re most common crab found at the active hy mains uncertain. The Lithodidae are repre drothermal vents, at 2000 m, in the Guay- sented at hydrothermal vents by two species mas Basin, Gulf of California. It is known and six occurrences not yet clearly assigned from southern California to South Georgia, to species. Two of the described species at depths of 640-2450 m (Macpherson {Paralomis arethusa and P. jamsteci) are 1988). until now known only from reducing envi ronments. No lithodid species is definitely Genus Paralithodes Brandt, 1848 known from both vents and seeps. Based on submersible observations, Sa- galevich et al. (1992) reported Paralithodes Genus Lithodes Latreille, 1806 sp., at 350-400 m, on the periphery of ac Lithodes manningi Macpherson, 1988 tive hydrothermal vents on the summit of One specimen was collected at 1236 m Piyp Volcano, in the Bering Sea, on dense depth in cold seeps of the Barbados accre populations of actinians. 288 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Genus Paralomis White, 1856 Paralomis multispina (Benedict, 1894) This is the best represented of the lith- Horikoshi & Ishii (1985), Hashimoto et odid genera in chemosynthetically-based al. (1987, 1989), and Ohta (1990b) have de communities. At least four species have scribed cold-seep communities of Sagami been reported, including two exclusively Bay in Japan, at depths of 900-1200 m, from reducing habitats. Considering the nu where large Paralomis multispina and the merous reports of "Paralomis sp." found clams Calyptogena soyoae are the dominant in the literature, more species are likely to species. P. multispina is known from Japan be discovered in the future. to California, at 500-1665 m (Hart 1982). Paralomis verrilli (Benedict, 1894) Paralomis arethusa Macpherson, 1994 Paralomis verrilli is believed to be the Paralomis arethusa is one of two lith- species found at hydrothermal vents of the odid crabs known from cold-seep commu Iheya Ridge, at 1400 m, in the Mid-Oki nities of the Barbados accretionary prism at nawa Trough (S. Ohta, pers. comm.; Ohta 1691 m depth. This species is known only 1990a; Kim & Ohta 1991; Hashimoto et al. from this location (Macpherson 1994). 1995). Although still to be confirmed, this occurrence would not be surprising as P. verrilli is usually found at depths of ca. Paralomis cubensis Chace, 1939 1500-3500 m around Japan and from the Bering Sea to California (S. Ohta, pers. Sassen et al. (1993) produced a photo comm.). Crabs tentatively assigned to this graph of a "crab" (clearly a lithodid) crawl species were also reported from the Sagami ing on tubeworms and mussels at the Green Bay cold-seeps, but in much lower numbers Canyon methane seep, on the upper conti than the dominant P. multispina (J. Hashi nental slope of the Gulf of Mexico, at 620 moto, pers. comm.). m. It was subsequently identified as Para lomis cubensis. It is considered as a "va Additional occurrences of Paralomis spe grant" species of the Gulf of Mexico seeps cies.—Several authors have reported the by Carney (1994), and is not commonly in occurrence of lithodid crabs as belonging to direct contact with the seep community. P. the genus Paralomis. Most reports have cubensis is known from east Florida to Bra been based on collected specimens. Two yet zil, at 329-730 m (Macpherson 1988). unidentified species of Paralomis occur at the Minami-Ensei vents (Mid-Okinawa Trough), living on mytilids or around bac Paralomis jamsteci Takeda & Hashimoto, terial mats, together with the vent endemic 1990 P. jamsteci (Hashimoto et al. 1995). In the hydrothermal areas of the North-Fiji back- In hydrothermal areas of the Okinawa arc basin (2000 m) the "white-coloured Trough, lithodid crabs have been observed lithodid crabs" observed by Jollivet et al. on the Minami-Ensei Knoll, at 700 m, (1989), were later identified as belonging to where a new species was described as Par the genus Paralomis, and have also been alomis jamsteci (Hashimoto et al. 1990, sampled at 1800 m in venting areas of the Takeda & Hashimoto 1990). This species is Lau back-arc basin (Desbruyeres et al. living among mytilid beds, near vent open 1994). On the Juan de Fuca ridge in the ings (Hashimoto et al. 1995). Two other yet northern Pacific, the lithodid crab observed unidentified species of Paralomis also oc and sampled at 1600 m on the Axial Sea- cur at the Minami-Ensei vents (Hashimoto mount hydrothermal vent area by Tunni- et al. 1995). cliffe & Jensen (1987), is a species of Par- VOLUME 109, NUMBER 2 289 alomis that co-occurs with the brachyuran and seeps. As there appears to be no other majid crab Macroregonia macrochira. advantage in getting close to such a toxic These authors suggested that this lithodid milieu than the increased food-supply (par might be the same species found at the Or ticularly near hydrothermal vents with high egon cold-seeps by Suess et al. (1985). This concentrations of hydrogen sulfide and Paralomis sp. and Paralomis verrilli (if heavy metals, low oxygenation and pH), confirmed) would then be the only lithodid they can be considered at least to partially species that occur in both vent and seep en feed on chemosynthetically-derived mate vironments. Galkin & Moskalev (1990b) rial. also observed a crab closely related to Par Cold-seep Lithodidae described to date alomis on the Juan de Fuca ridge. range in depths from 250 to 2037 m. How Remarks on Lithodidae.—The Lithodi- ever, one occurrence is known from cold- dae do not seem to be physiologically de seep communities of the Nankai Trough, Ja pendent on the chemosynthetic production pan subduction zone, at 3800 m (M. Sibuet, of vents or seeps. E. Escobar-Briones (pers. pers. comm.). At hydrothermal vents, the comm.) recently observed that although depth range is 350-2000 m. The geograph Neolithodes diomedeae was seen near bac ical distribution of these observations is terial mats and hydrothermal vent structures worldwide, with some noticeable excep in the Guaymas Basin, it appears to mainly tions: no Lithodidae have ever been ob feed on organic matter of photoautotrophic served near hydrothermal vent areas of the origin. The opportunistic behavior of this East Pacific Rise (EPR), the Galapagos deep-sea scavenger is, however, believed to Rift, nor the Mid-Atlantic Ridge. The crab contribute to the export of hydrothermal mentioned by Guinot & Macpherson (1987) material to the surrounding abyssal environ from the EPR at 12°35'N, although proba ment. In other hydrothermal and cold-seep bly a lithodid, cannot be assigned to a hy settings, direct evidence exists of lithodid drothermal context. It was indeed observed crabs feeding on chemoautotrophic symbi- on an off-axis seamount located 18 km ont-containing fauna. Tunnicliffe & Jensen from the ridge axis and even farther from (1987) reported that pieces of vestimentif- the nearest known vent site in this area (P. eran tubes were found in the stomachs of Chevaldonne, pers. obs.). Paralomis sp. at Axial Seamount vents. Ob servations of P. multispina and P. jamsteci Superfamily Galatheoidea feeding upon live vesicomyid and mytilid bivalves are mentioned by Hashinipto et al. This superfamily includes four families, (1989) and Takeda & Hashimoto (1990) in two of which are represented at deep-sea Sagami Bay cold-seeps and Okinawa vents. hydrothermal vents and cold seeps. Suess et al. (1985) described a large lith odid in the Oregon seeps as actively feeding Family Galatheidae on vesicomyid clams. The Lithodes spp. Galatheid crabs have been found associ from Monterey Bay seep communities are ated with hydrothermal environments since known to have attacked unsuccessfully ves the first submersible expeditions explored icomyid clams (J. P. Barry, pers. comm.). the Galapagos Rift vents (Corliss & Ballard Another type of feeding has been proposed 1977). They have also been known to occur for P. multispina by Horikoshi & Ishii in cold-seep settings since the discovery of (1985), who observed this crab actively this type of community (Paull et al. 1984). scooping up and "feeding" on the black re Galatheids are familiar members of vent duced mud within the clam beds of Sagami and seep communities but often do not at Bay cold-seeps. Most frequently, the Lith tract as much attention as other spectacular odidae occur in greatest densities at vents organisms such as the Vestimentifera and 290 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON large bivalves. Also, because most gala- Genus Munidopsis Whiteaves, I 874 theid crabs appear similar, to the non-spe This is the most commonly encountered cialists, observations on them are numerous galatheid genus in reducing environments, but often vague. Only three of the 11 defi including at least 10 species, of which six nite species known so far have been re are known so far only from hydrothermal ported from cold-seep communities, and vents. Many authors have reported Muni one, Munidopsis crassa, from both hydro- dopsis sp. or Munidopsis-WYe galatheid thermal and seep environments. Reports of crabs, so again, the list of species is likely the latter, however, seem suspect as one is to expand. from the Mid-Atlantic Ridge, and another from the Peru margin. Many occurrences are mentioned only as "galatheid crabs," so Munidopsis alvisca Williams, 1988 much more species are likely to be de This so far exclusively hydrothermal-as- scribed in the future. sociated species was described by Williams (1988) from widely separated venting areas. Munidopsis alvisca occurs at the periphery Genus Munida Leach, 1820 of hydrothermal areas on the Explorer and Munida magniantennulata Baba & Juan de Fuca ridges, at 1545-1800 m. in Tiirkay. 1992 the northeastern Pacific, and at 2000 m in This species was originally described the Guaymas Basin, Gulf of California from three specimens collected at hydro- (Khodkina 1991). Van Dover et al. (1990) thermal vents of the Lau back-arc basin in also reported a galatheid crab which might the southwest Pacific, at a depth of 1750- be M. alvisca, from vents of the Escanaba 2000 m (Baba & de Saint Laurent 1992, Trough on the Gorda Ridge. Baba & Tiirkay 1992). ii was subsequently found in collections from a non-vent envi Munidopsis beringana Benedict. 1902 ronment off Australia (Baba 1994). A galatheid sampled from cold-seeps of the Middle America Trench at 3700-4000 Munida inicrophthalma Milne Edwards, m west of Mexico has been assigned to Munidopsis beringana (E. Macpherson. 1880 pers. comm.). The galatheid crabs observed !n the seepage areas of the Barbados ac- at shallower seen communities (2900 m) cretionary prism, at 1700-2000 m. two dif might belong to this same species (K. Olu. ferent species of Galatheidae have been ob pers. obs ). There is still some doubt 'n he served • K. Olu, per?, obs.). Only one has determination of this species, as the col been collected and identified as M. inicro lected specimens present some variations phthalma (M. de Saint Laurent, pers. from other M. beringana. This occurrence comm.). Jollivet et al. (1990) earlier re is also out of the known geographical and ported Munic lopsis sp. iTom the same area. depth range of this species, usually found They either misidentified M. microphthal- from the Bering Sea to Oregon, at 2800- ma from their towed camera pictures, or ob 3276 m (Ambler !980l. served the second and bigger species, which night be a Munidopsis. Munida mi- Munidopsis crassa Smith. 1885 crophtluibna has been reported from the At As previously mentioned, this species is lantic, south of Iceland to Ascencion Island, believed to be present horn in cold-seep and in the Gulf of Mexico, the Carribbean, and usdrothermal setting'-. Segonzac (1992) re the E.iv of 3iscaye, at uenths of 194-2129 ported it "rom the snake Pit hydrothermal m (Wt-mer I982\ area, al jx'O m. • r. r.e Mid-Atlantic R;d:ie. OLUME 109, NUMBER 2 29! Olu et ai. (1996) recently reported Muni- deep hydrothermal sites of the Mariana dopsis crassa from cold-seeps of the Peru back-arc basin, western Pacific, where it margin, at 3000-3600 m, although uniden lives in 10-25°C waters among dense car tified galatheids were also observed at 5040 pets of actimans (Williams & Baba 1989. m. A comparative examination of Atlantic Hessler & Lonsda.e 1991). and Peru specimens is needed to confirm these findings. M. crassa was previously Munidopsis sonne Baba, 1995 inly known from the Atlantic, at 2679- 5315 m (Wenner 1982). Munidopsis sonne is only known from two specimens collected at active vents of Munidopsis diomedeae (Faxon. 1895) the North Fiji Basin, at 1992 m (Baba 1995). Munidopsis diomedeae was observed near hydrothermal vents of the Guaymas Munidopsis starmer Baba & de Saint Basin, Gulf of California, at 2000 m depth Laurent, 1992 (Khodkina 1991), along with M. alvisca. One of these two species might be the Mun Munidopsis starmer also seems restricted idopsis sp. mentioned by Lutz (1992) from to hydrothermal vents (Baba & de Saint cold seepages in the same area. M. diome Laurent 1992) and occurs at vent sites of deae is common in the Gulf of California, the North-Fiji Basin, at 2750 m. In the same and is known from California to Chile, at area, M. Iauensis, M. sonne, and two spe 768-3790 m (Haig & Wicksten 1975). cies of Chirostylidae are also present. Munidopsis iauensis Baba & de Saint Munidopsis subscpiamosa Henderson, Laurent, 1992 1885 Munidopsis Iauensis occurs with Munida Munidopsis subsquamosa is believed to magniantennuiata at deep-sea vents of the be a complex of deep-sea cosmopolitan Lau back-arc basin, in southwest Pacific, at species (Van Dover 1986), which explains 1750 m. It is also found at 2000 m in hy its considerable geographic range. Although drothermal sites of the nearby North-Fiji this complex clearly needs a complete re back-arc basin (Baba & de Saint Laurent examination and taxonomic work at the 1992). To date, this species is exclusively molecular '.evel. we will simply call it here known from hydrothermal areas. M. subscpiamosa. It was the first galatheid species observed in a chemosynthetically- Munidopsis lentigo Williams & Van based community, and is found at almost Dover. 1983 every vent location in the eastern Pacific, in a depth o\' ca. 2600 m (de Saint Laurent This hydrothermal vent endemic is con 1984, Hessler et al. 1985, Van Dover et ai. sidered a "high temperature" species (Wil 1985). At 21°N on the East Pacific Rise liams & Van Dover 1983) as it seems to be (EPR), it occurs with M. lentigo, but inhab the galatheid that lives closest to high tem its colder waters. It is the only species ob perature fluid venting. It has been found served at 13°N (EPR) and sites of the Ga only at vent sites of the East Pacific Rise at lapagos Rift, and has also been collected 21°N. 2600 m depth. from the EPR at 10-12°N (Van Dover & Hessler 1990) and 9°50'N (Kaartvedt et ai. Munidopsis marianica Williams & Baba, 1994). At the latter location, Lutz (1992) 1989 suggested that there might be two different Munidopsis marianica is another appar species of Munidopsis. ently vent-endemic from the 3600-3700 m Additional occurrences of Munidopsis 292 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON species.—Many reports of "Munich)psis dopsis spp. are much more common than sp/' exist from a great variety of hydro- the chirostylid Eumunida picta that also oc thermal and seepage areas. Hydrothermal curs at the 400-1000 m sites, and they live occurrences include the EPR at I7-20°S in close association with mytilid bivalves (Van Dover & Hessler 1990); three loca and vestimentiferan tubeworms. At 3270 m, tions in the Mid-Okinawa back-arc basin, in cold-seeps of the Florida Escarpment, the Iheya Ridge at 1400 m (Ohta 1990a, Hecker (1985) has reported a large and nu Kim & Ohta 1991), the Izena Caldron at merous white Munidopsis sp. that has not 1400 m (Hashimoto et al. 1995), and the been seen far outside the seep communities. Minami-Ensei Knoll at 700 m (Hashimoto Unconfirmed reports of Munidopsis-Vike et al. 1990, 1995); Manus back-arc basin in galatheids include those from the hydro- the southwest Pacific at 2500 m (Galkin thermal vents of the Piyp volcano, at 800 1992); TAG area on the Mid-Atlantic Ridge m, in the Bering Sea (Sagalevich et al. (Galkin & Moskalev 1990a); and Broken 1992). In the region of the Barbados accre- Spur area farther north on this same ridge tionary prism, Munidopsis-likc galatheids (Murton et al. 1995). Reports from cold- have also been observed associated with seeps include the Nankai Trough in the Jap cold seeps on mud volcanoes, at 4710- anese Trench at 3800 m (Laubier et al. 4980 m (K. OIu. pers. obs.), and galatheids 1986), cabonate seeps of Enshu-nada, Ja tentatively assigned to Munidopsis were re pan, at 1000-1220 m (Ohta et al. 1995), ported by Jollivet et al. (1990), at shallower and the Laurentian Fan at 3850 m (Mayer cold-seep sites (1000-2000 m), in the et al. 1988). southern part of the prism. In Monterey Bay, cold-seep communities occur at two different bathymetric levels, at 3000-3600 m (Embley et al. 1990) and Family Chirostylidae 600-1000 m (J. P. Barry, pers. comm.). At least two different species have been ob Chirostylid crabs are represented in che- served so far, possibly of two different gen mosynthetic communities by two genera era. At deeper sites, a single species has and three species. This family is frequently been found and tentatively assigned to the overlooked by non-specialist observers genus Munidopsis by Embley et al. (1990), when reporting the occurrence of "gala but could also be a species of Manida ac theids." cording to J. P. Barry (pers. comm.). At shallower seeps (600-1000 m), two differ ent species of Galatheidae are known, one Genus Eumunida Smith. 1883 of which might be the same species ob Eumunida picta Smith. 1883 served deeper, and the other apparently is a This species is considered as a rare va seep endemic that might belong to the ge grant in cold-seeps of the Louisiana slope, nus Munidopsis (J. P. Barry, pers. comm.). at 400-1000 m depth (Carney 1994). It has In the Gulf of Mexico seep communities, been reported from Massachusetts to Co MacDonaldet al. (1989) reported a possibly lombia, at 200-600 m (de Saint Laurent & undescribed species of Munidopsis from the Macpherson 1990). Bush Hill site, at 540 m, on the Louisiana Slope. At different seepage depths on the slope (400-1000 m and 2200 m), Carney Genus Uwprychtts Henderson. 1888 (1994) indicated two Munidopsis spp. (in cluding one seep endemic), different from Baba&de Saint Laurent (1992) have de the "background" galatheids. One of these scribed two new species only known from is probably the Bush Hill species. Muni vents of the North-Fiji Basin. VOLUME 109, NUMBER 2 293 Uroptychus bicavus Baba & de Saint at low oxygen levels, where most other in Laurent, 1992 vertebrates of the fjords cannot compete for food and space (Burd & Brinkhurst 1984). Uroptychus bicavus is known from a Stable isotope analyses led Van Dover & depth of 2750 m. Fry (1989) and E. Escobar-Briones (pers. comm.) to characterize Munidopsis sub Uroptychus thermalis Baba & de Saint squamosa and M. alvisca, of the Galapagos Laurent, 1992 and Guaymas hydrothermal vents, as hav Uroptychus thermalis occurs at depth of ing a mixed diet partly including a sulfur- 2000 m. based source of carbon. Similar analyses on Additional occurrences of Galatheo- the Iheya Ridge Munidopsis sp. also show idea.—Several authors have reported the at least a partial nutritional dependence on occurrence of "galatheid crabs" or "squat chemosyntheuc processes (Kim et al. 1989, lobsters" that cannot be assigned with cer Kim & Sakai 1991). At the Monterey Bay tainty to a family. These reports are from cold-seeps, an apparently seep endemic gal hydrothermal areas of the EPR at 21°30'S atheid is found grazing on mats of the fil (Renard et al. 1985), 23°30'S and 26°S amentous bacteria Beggiatoa, and is even (Marchig & Gundlach 1987), the Gulf of covered with the bacterial filaments (J. P. Aden (Juniper et al. 1990) and the Edison Barry, pers. comm.). Seamount in the southwestern Pacific (Her- Galatheids and chirostylids might also zig et al. 1994), and from cold seeps of En- act as vectors for the export of organic mat shu-nada. Japan, at 1000-1220 m (Ohta et ter of hydrothermal origin in the deep sea. al. 1995). They might disperse more efficiently than Remarks on Galatheidae and Chirostyli- lithodids, due to their swimming abilities, dae.—Like Lithodidae, Galatheidae and and can also easily detect new food sources. Chirostylidae are generally not considered Two shark carcasses placed 100 m from a to be dependent on the chemically reduced vent site at 13°N on the EPR (Fig. 1) at fluids emitted at vents and seeps. Their rep tracted hundreds of Munidopsis subsqua resentatives are often restricted to the pe mosa within a day (D. Desbruyeres, pers. riphery of the sites, where they are believed comm.). These characteristics might also to be opportunistic species taking advantage favor dispersal of non-planktonic develop of increased productivity. However, there is mental stages of vent and seep species. Car a significant number of species that seem to ney (1994) reported that Munidopsis sp. of be endemic to hydrothermal vents or cold- the Gulf of Mexico seepage areas are often seeps, and some of them even appear to be covered with juvenile mussels, and may be adapted to this harsh environment, not just an important means of post-settlement dis the periphery. Van Dover (1986) observed persal for seep endemic sessile taxa. Such that gravid females of the hydrothermal phoretic processes might also occur in hy Munidopsis subsquamosa seemed to live drothermal polychaetes (Tunnicliffe & Jen under higher temperature conditions than sen 1987, Zal et al. 1995). males. Although this species is not restrict Galatheidae are found in virtually every ed to vent areas, this behaviour was inter hydrothermal or cold-seep environment, as preted as a possible adaptation to improve well as in other types of reducing habitats reproductive efficiency. Galatheids also such as whale carcasses (Bennett et al. seem to adapt relatively well to hypoxic 1994, Wada et al. 1994) or decomposing conditions, such as those encountered in re wood (Williams & Baba 1989). They do not ducing habitats. In some British Columbia seem to be limited by depth, as their oc fjords, for instance, dense aggregations of currence is reported from as deep as 5040 Munida quadrispina are consistently found m, in the deepest cold-seep communities «.*! 1 Vi m 03 n o Fig. I. Allraclion of Muniilopsis substiiuimoxa, a galalheid crab commonly found in deep-sea hydroihcimal vents of ihe eastern Pacific, to shark carcasses § deposited 100 in from :i vent silo :il 13°N, on tin- I: ml I'ucilic Rise (2600 in " ' '—I GLUME 109, NUMBER 2 295 (Olu et al. 1996). They are also associated Brandt. J. F. 1848. Die Gattung Lithodes Latreille with the shallowest occurrence of vestimen- nebst vier neuen ihr verwandten von Wosnes- senski entdecken, als Typen einer besondern tiferans known to date, at 82 m, in the cold- Unterabtheilung (Tribus Lithodea) der Ed- seep site of Kagoshima Bay, Japan (Hash wards'schen Anomuren.—Bulletin de la Classe imoto et al. 1993;. Physico-Mathematique de l'Academie [mperi- ale des Sciences de Saint-Petersbourg 7:171- Acknowledgments 175. Carey, A. G., Jr., D. L. Stein, G. L. Taghon, & A. E. We thank D. Desbruyeres, M. Sibuet, M. DeBevoise. 1988. Biology and ecology of the Segonzac and anonymous referees for help Oregon continental shelf edge associated with ful comments on this manuscript, and A. the accretionary prism. Pp. 137-149 in M. P. Gebruk, S. Ohta, J. Hashimoto, J. P. Barry, De Luca & I. Babb. eds., Global venting, mid- water, and benthic ecological processes. NOAA, and E. Escobar-Briones for communicating Groton, Connecticut, 442 p. valuable information. R. Lemaitre, T. Com- Carney, R. S. 1994. Consideration of the oasis anal tet, M. de Saint Laurent and D. Guinot pro ogy for chemosynthetic communities at Gulf of vided much appreciated help. Mexico hydrocarbon vents.—Geo-Marine Let ters 14:149-159. Literature Cited Chace. F. A . Jr. 1939. Reports on the scientific results of the first Atlantis Expedition to the West In Ambler. J. W. 1980. Species of MuniJopsis (Crusta dies, under the joint auspices of the University cea, Galatheidae) occurring off Oregon and in of Havanna and Harvard University. Prelimi adjacent waters.—Fishery Bulletin 78:13-34. nary descriptions of one new genus and sev Baba, K. 1994. Deep-sea galatheid crustaceans (An- enteen new species of decapod and stomatopod omura: Galatheidae) collected by the "Cidaris Crustacea.—Memorias de la Sociedad Cubana I" expedition off central Queensland. 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