23 April 2003 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1I6(1):I58-I67. 2003. A new caridean of the family Alvinocarididae from thermal vents at Menez Gwen on the Mid-Atlantic Ridge

Timothy M. Shank and Joel W. Martin (TMS) Biology Department, Woods Hole Oceanographic Institution, Woods Hole, Massachusetts 02543, U.S.A., e-mail: [email protected]; (JWM) Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007, U.S.A., e-mail: [email protected]

Abstract.—A new species of alvinocaridid shrimp, Alvinocaris williamsi, is described from the Menez Gwen field on the Mid-Atlantic Ridge. The new species is most similar to another relatively shallow water alvinocaridid, A. stactophila Williams, known only from a cold seep area off the coast of Louisiana, but differs in details of the rostrum and appendages. Alvinocaris williamsi is also morphologically similar to two species of the recently described from the Mid-Okinawa Trough, Japan. Molecular phy- logenetic studies of A. williamsi, the eighth described member of the genus, may provide significant insights into the role that deep-sea hydrothermal vents at mid-ocean ridges and hydrocarbon seeps on continental margins have played in the evolution of fauna endemic to these chemosynthetic habitats.

The caridean shrimp family Alvinocari­ The type genus of the Alvinocarididae, didae was originally proposed by Christof- Alvinocaris, is the most diverse genus of fersen (1986) to accommodate a number of the family and contains five hydrothermal morphologically similar shrimp known vent-endemic species: A. brevitelsonis Kik- from hydrothermal vents and hydrocarbon uchi & Hashimoto, 2000, from the Minami- seeps. The family was further diagnosed by Ensei Knoll in the Mid-Okinawa Trough off Segonzac et al. (1993) in a footnote, and Japan; A. leurokolos Kikuchi & Hashimoto, also by Vereshchaka (1996a, 1996b, 1997) 2000, also from the Minami-Ensei Knoll; A. (see Martin & Davis 2001). To date, five longirostris Kikuchi & Ohta, 1995, from genera have been proposed for the family. the Iheya Ridge off Japan; A. lusca Wil­ In the order in which they were proposed, liams & Chace, 1982, from the Galapagos they are: Alvinocaris Williams & Chace, Rift in the eastern Pacific; and A. markensis 1982; Rimicaris Williams & Rona, 1986; Williams, 1988, from the Mid-Atlantic Chorocaris Martin & Hessler, 1990; Opae- Ridge. Another two species of Alvinocaris pele Williams & Dobbs, 1995; and lorania are endemic to chemosynthetic habitats as­ Vereshchaka, 1996b (see Shank et al. 1998 sociated with hydrocarbon and brine seeps: for the synonymy of lorania and Rimicar­ A. muricola Williams, 1988, from the West is). A sixth genus of vent shrimp, Mirocaris Florida Escarpment; and A. stactophila Wil­ Vereshchaka, 1997, was proposed by Ver­ liams, 1988, from the Louisiana Slope in eshchaka (1997) for Chorocaris fortunata the northern Gulf of Mexico. The study of Martin & Christiansen (1995) and for a new additional collections, including unde- species (M. keldyshi Vereshchaka, 1997); scribed hydrothermal-associated shrimp however, this genus was transferred to the specimens from the Edison Seamount of the newly created family Mirocarididae by Ver­ Bismark Archipelago in Papua New Guinea eshchaka (1997). (Shank et al. 1999; K. Baba, pers. comm.). VOLUME 116, NUMBER I 159 from the Logatchev vent field of the Mid- portion of a hydrothermally-active mussel Atlantic Ridge (A. Vereshchaka, pers. bed community that included a more abun­ comm.), and several recently recovered dant alvinocaridid species, Mirocaris keld- from active seamounts north of the Bay of yshi, amphipods, Luckia striki Bellan-San- Plenty (northern North Island, New Zea­ tini & Thurston, gastropods, e.g., Lepeto- land) (Rick Webber and Neil Bruce, pers. drilus spp. and Protolira valvatoides Waren comm., http;//www.niwa.co.nz/pubs/bu/05/ & Bouchet, polychaetes, e.g., Branchipo- blind) may yield additional new species of lynoe seepensis Fettibone, and , S«?- Alvinocaris. The pan-equatorial biogeo­ gonzacia mesatlantica (Williams, 1988) graphic distribution of the genus is note­ and Chaceon affinis (A. Milne-Edwards & worthy. Alvinocaris species occur in almost Bouvier, 1894). Specimens were brought to every biogeographic vent province (except the surface in an insulated container and the Northeast Pacific vents; Shank et al. placed in chilled water on-board ship. 1999). However, recent initial investiga­ Whole shrimp were sorted by morphotype tions of hydrothermal vents in the Indian and either preserved in 4% buffered for­ Ocean did not reveal the presence of Alvi­ malin in seawater and subsequently trans­ nocaris shrimp despite the common ap­ ferred to 70% ethanol or frozen at — 70°C pearance of the other dominant Atlantic and and subsequendy transported on dry ice and Pacific genera Rimicaris and Chorocaris stored at -80°C. Carapace length (CL) of (Hashimoto et al. 2001, Van Dover et al. each individual was measured in millime­ 2001). The Alvinocaris species described ters (mm) from the orbital margin to the herein occupies the northernmost extent of posteriomedial margin of the carapace. We the generic range, as it inhabits the Menez recognize that the description of new spe­ Gwen vent site, the most shallow vent field cies within this family from relatively few (850 m) and the most northern known ac­ individuals can be risky (see Shank et al. tive deep-sea hydrothermal site along the 1998); however, preliminary results of mi­ Mid-Atlantic Ridge {37°50.5'N, 3l°3l3'W) tochondrial DNA sequence data (Shank, (Gebruk et al. 1997, Colago et al. 1998, pers. obs.) complement morphological evi­ Comtet & Desbruyeres 1998, Desbruyeres dence that distinction of this species from et al. 2001). Below, we describe a new spe­ other Alvinocaris species is warranted. The cies of Alvinocaris from the Menez Gwen holotype and paratypes are deposited in the hydrothermal vent field, and suggest evo­ collections of the National Museum of Nat­ lutionary relationships with other alvino- ural History, Smithsonian Institution, carid species. Washington, D.C. (USNM). The remaining specimens are cryo-preserved at the Woods Hole Oceanographic Institution for ongoing Materials and Methods molecular phylogenetic investigations by All specimens were collected using the TMS. human occupied submersible DSV Alvin at the Menez Gwen hydrothermal vent field, northern Atlantic Ocean. Shrimp were col­ Alvinocaris williamsi, new species lected from a cluster of active individuals Figs. 1-3 among clumps of mussels, Bathymodiolus azoricus Von Cosel, Comtet & Krylova, Material examined.—14 females CL: 8.2, within vent flow with a 30 cm X 30 cm 8.2, 8.5, 8.7, 8.9, 8.9, 9.3, 9.5, 9.5, 9.5, 9.6, square "black net" (BN) operated using the 9.6, 9.7, 10. L DSV Alvin, Dive 3117, Menez manipulator arm by the pilots oi DSV Alvin. Gwen hydrothermal vent field, North Atlan­ A total of 14 specimens were collected tic Ocean, 37°50.5'N, 3r31.3'W, 850 m, 7 (from a single net sample) from areas of Jul 1997. most intense diffuse fluid within the central Types.—All from DSV Alvin Dive 3117. 160 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. L Alvinocaris williamsi, new species, iiolotype ovigerous female, USNM 1009651. a, body, cephalic appendages, and abdomen, lateral view; b, anterior region of carapace, including eyes and antennae, dorsal view; c, telson and uropods, dorsal view. Scale bars equal 1.0 mm (bar 1 for a, bar 2 for b, bar 3 for c).

Holotype: adult female CL 9.5 mm, ovig­ midlength of second segment; dorsal mar­ erous (specimen ID: BN 70), USNM gin raised into thin serrate crest containing 1009651. Paratypes: 2 adult females CL 13 or 14 teeth of nearly equal strength in 9.3, 9.7 (specimen ID: BN 68 and BN 51, central sector of row, about 0.5 length of respectively), abdomen torn from body, tail crest continued onto carapace, deflecting fan missing, USNM 1009652, USNM from dorsal line of carapace at approximate 1009653. 45° angle; ventral margin less prominent, Description.—Integument thin, shining, straight, and unarmed or with 1 minute sub- minutely punctuate. Rostrum (Fig. la, b) al­ terminal tooth; lateral carina (not visible in most straight, imperceptibly elevated above Fig. la) broadened proximally and conflu­ horizontal in distal half; sharply pointed tip ent with orbital margin. Carapace (Fig. la) (broken or slightly blunted in all speci­ with buttressed and distinct antennal spine; mens) reaching at least to level of articu­ pterygostomian spine prominent, acumi­ lation between first and second peduncular nate. Anterior antennal carina curving pos- segment of second antenna or sometimes to teroventrally from near base of antennal VOLUME 116. NUMBER 1 161 spine to intersect at about midlength of car­ with stronger mesiodistal spine. Antennal apace with carina extending posteriorly scale (scaphocerite) broadly rounded distal- from pterygostomial spine, its associated ly, with broad triangular tooth on distola- groove (not visible in Fig. lb) continuing teral border extending to same level as indistinctly posteriorly. rounded distal border. Abdomen of female broadly arched dor- Mouthparts (Fig. 2) fairly typical for the sally, gradually tapering posteriorly, height genus. Mandible (Fig. 2a, b) with 6 blunt, of sixth somite about half that of first so­ uneven, terminal teeth plus one slightly mite. Pleura of third somite broadly round­ sharper dorsal subterminal tooth along cut­ ed, that of fourth somite irregularly round­ ting border, and with long, blunt, posterior ed, flanked dorsally by single remote ob­ tooth ("molar process" of some authors) solescent spine; fifth pleuron with strong separated from cutting edge by a wide gap; acute triangular posteroventral tooth mandible deeply excavate on internal (pos­ flanked dorsally by cluster of 2 or 3 remote terior) surface just above this blunt poste­ small spines on margin; sixth somite with rior tooth; mandibular palp 2-segmented, middorsal length about 1.4-1.9 that of fifth distal article bearing numerous plumose se­ somite, broad-based posterolateral tooth tae; basal article with 2 plumose setae on overlapping base of telson. smaller postero­ distal border. First maxilla (Fig. 2c) with 2 lateral spine acute. Telson (Fig. Ic) elon­ endites; coxal (di.stalmost) endite with row gate, subrectangular, length (not including of evenly spaced and evenly sized blunt posterior teeth) about 2.0-2.7 anterior spines (arrow) in addition to dense plumose width, 3.2-3.7 posterior width, about 1.3- and simple setae; basial endite (proximal 1.5 length of sixth somite; armed with 5-7 endite) with longer, curved setae but lack­ dorsolateral spines of nearly uniform size, ing spine row; palp unsegmented, with 2 occasionally unequal in number on either short and 1 long terminal setae as illustrat­ side; posterior margin slightly convex, ed. Second maxilla (Fig. 2d) scaphognathite armed with 1 spine at each corner and 12 with dorsal lobe broad, distally almost trun­ feathered strong setae on posterior margin cate, and lined with evenly spaced plumose between them. setae; posterior lobe of scaphognathite ta­ pering posteriorly, almost triangular, with Eyes (Fig. la, b) with cornea imperfectly setae increasing in length posteriorly, each developed; unfaceted though diffusely pig­ seta angled and minutely serrate (for mented; ovate in outline though fused to grooming); coxal lobe simple, blunt, setose; each other mesially beneath rostrum, each basial endite bilobed, each lobe fringed with upturned spine on anterodorsal sur­ with plumose setae; palp thin, nantjw, face. strap-like. First maxilliped (Fig. 2e) phyl- Antennular peduncle (Fig. lb) extending lopodous; coxal endite distally blunt and slightly short of end of antenna! scale; basal slightly recurved; basial endite bilobed, segment 2.1 length of second segment and with distal lobe approximately twice length about 2.2-2.5 length of third, all measured of proximal lobe; palp thin, tapering to on dorsal margin; stylocerite well separated acute distal tip bearing short setae and vis­ from peduncle, tapering to slender elongate ible only in posterior view; exopod broad, tip variably reaching as far as midlength of expanded and rounded distally, lined on ei­ second segment; basal segment with disto- ther side with evenly spaced plumose setae; dorsal margin exceeded by rostral tip epipod bilobed at base, fused distally, with though extended laterally into strong lateral dorsolateral lobe bearing weak posterior spine reaching level nearly equal to that of projection. Second maxilliped (Fig. 2f) ped- stylocerite and closely appressed to second iform, 6-segmented; coxa expanded and segment, much smaller distomesial spine rounded on medial surface, which bears nu- slightly divergent; shorter second segment 162 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Alvinocaris williamsi, new species, paratype female, USNM 1009653. Mouthparts, right side, a, mandible, outer (external) view; b, mandible and palp, inner view; c, first maxilla, with row of evenly spaced spines magnified and indicated by arrow; d, second maxilla; e, first maxilliped; / second maxilliped; g, third maxilliped, with tip enlarged (upper arrow) and inner surface with rows of setae displayed (lower arrow). Scale bar 1 equals 0.5 mm (a, h, c), and 1.0 mm {d, e, f, g). merous plumose setae; fused basi-ischium tae on proximal region. Epipod nearly tri­ with evenly spaced plumose setae along angular, with weak lobe coming off poste­ medial border; merus and carpus short; pro- rior surface. Third maxilliped (Fig. 2g) ped- podus trapezoidal; dactylus short, distally iform, elongate, 4-segmented; coxal seg­ rounded, with brush like patch of dense se­ ment short, with minute epipod bearing 1 VOLUME 116, NUMBER 1 163 or 2 setae; distal 3 segments elongate and First pereopods (Fig. 3a, b) chelate, sub- pediform; basal of these with plumose setae equal; fingers curved ventrally and slightly on proximal medial border and row of plu­ laterally; dactylus more slender than fixed mose setae along dorsolateral border; dac- finger, tips varying slightly in relative tylus tipped with acute sclerotized spines length, mesial surface of each finger con­ surrounding central claw-like tip (arrow), cave; cutting margins uniformly offset, and with evenly spaced densely packed closing without gape, each armed with row rows of stiff, minutely serrulate setae on of almost uniform teeth so closely set as to ventro-medial surface (curved arrow). be almost contiguous, line of sensory setae

Fig. 3. Alvinocaris wilUamsi, new species, paratype female, USNM 1009652. a, right first pereopod (che- liped), inner view; b, right first pereopod, outer view; c, left second pereopod, tip of chela; d, left second pereopod; e, left third pereopod; / left fourth pereopod; g, left fifth pereopod. Scale bars 1 and 2 equal 1.0 mm; scale bar 3 equals 0.5 mm (bar 1 for d, e, f, g; bar 2 for a, b; bar 3 for c). 164 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON mesial to cutting edges, acute tip of dacty- those on merus; fourth leg with similar lus slightly spooned by elongate teeth slant­ spines on merus but ischium not always ed posteriorly and curving around external bearing spines; fifth leg without spines on edge; entire leg slightly shorter than third merus or ischium but with ventral row of maxilliped. Palm inflated but not elongate, small spines on propodus preceding base of approximately equal in length to fingers. dactylus. Carpus measured along dorsal border Pleopods (not illustrated) well developed, slightly longer than palm, bearing oblique first pair with endopods about half length ventral crest, ending in strong distolateral of exopods; endopods tapering to acute tip. spine (Fig. 3b) and flanked mesially by Uropods (Fig. Ic) with rami subequal in patch of setae on polygonal raised area; length, slightly exceeding posterior margin notch above spine smoothly concave, op­ of telson; lateral ramus of exopod with posing low ridge ending in small rounded movable spine mesial to larger distolateral spine on heel of palm; shallowly concave tooth, diaeresis sinuous. anteromesial margin of carpus leading dor- Etymology.—The species name com­ sally to 2 low rounded lobes. Merus swol­ memorates the late Austin B. Williams, Re­ len in distal half, distinct from ischium but search Scientist of the Systematics Labo­ fused to it, neither armed. ratory and National Marine Fisheries Ser­ Second pereopod (Fig. 3c, d) shorter and vice at the National Museum of Natural more slender than first, reaching to between History, Smithsonian Institution (see Le- midlength and end of antennal peduncle; maitre & Collette 2000). We are all bene­ finger slightly longer than palm, similar in ficiaries of his ceaseless pursuit of excel­ size and shape, opposed margins without lence in his contribution to the , gape, each pectinate with single row of systematics, biogeography, and evolution of teeth in distal half directed obliquely dis- various decapod groups, including vent tally and increasing slightly in size to end shrimp and species. in noticeably stronger tooth crossing op­ Remarks.—Alvinocaris wilUamsi appears posite member when closed, but spineless morphologically most similar to A. stacto- proximally; carpus slender, about 1.2 times phila, another species from a relatively longer than chela; merus unarmed, ischium shallow site (530 m). However, A. stacto- with stout spine at approximately 0.75 phila is known only from a distant hydro­ length. carbon seep site (as opposed to a hot vent) Third to fifth pereopods (Fig. 3e, f, g) off the coast of Louisiana, northern Gulf of similar in length and structure, third reach­ Mexico. Characters shared by the two spe­ ing beyond antenna! scale by about 0.3 the cies include a rostrum that is unarmed (or length of the propodus. Segments of these occasionally armed with a single tooth) on pereopods composed of: short dactylus the ventral margin (a feature also shared armed with about 6 corneous spines on with A. hrevitelsonis and A. leurokolos flexor surface, grading from small proxi­ from the western Pacific). The dorsal and mally to longest and strongest distally; ven­ ventral margins of the rostrum are heavily tral row of spines on propodus leading to toothed in all other described Alvinocaris base of dactylus; carpus of each leg with species. Additionally, both A. williamsi and distodorsal extension variously projecting A. stactophila have a relatively short ros­ as a stop along proximal part of propodal trum that continues posteriorly as a toothed extensor surface; third leg with ischium and carina that meets the dorsum of the cara­ merus stronger than on fourth and fifth leg, pace at an angle; this angle is sharper in A. merus of third with ventral spine at 0.3 and williamsi and A. stactophila than in any 0.6 length, distal spine tending to be stron­ other species except for A. longirostris, the gest, and ischium with 2 spines in line with distinctive rostrum of which, because of its VOLUME 1J6. NUMBER 1 165

length and armature, could not be confused scribed Edison Seamount specimens) were with that of either A. stactophila or A. wil- derived from the A. stactophila (seep) lin­ liamsi. Hovt'ever, A. williamsi can be easily eage. This suggests that the extant Alvino­ separated from A. stactophila by the sharper caris lineages share a common seep ances­ angle of the rostrum and carina where they tor. However, other vent-endemic lineages meet the dorsal line of the carapace (ap­ are basal to Alvinocaris, suggesting that proximately 45° in A. williamsi vs. approx­ vent lineages gave rise to a seep lineage that imately 30° in A. stactophila), by the rela­ in turn gave rise to other extant vent line­ tive length and width of the telson (which ages. Based on morphology, A. williamsi is considerably longer and narrower in A. from the relatively shallow Menez Gwen stactophila), and by the shorter rostrum rel­ site more closely resembles a seep-endemic ative to the carapace in A. williamsi. species (i.e., A. stactophila) than other hot Based on the absence of ventral teeth on vent species, and therefore molecular ge­ the rostrum and the angle of the rostral ca­ netic comparisons of A. williamsi with con­ rina and carapace, A. williamsi is also mark­ geners from deep-sea hydrothermal vents, edly similar to two species, A. hrevitelsonis seamounts, back-arc basins, and hydrocar­ and A. leurokolos Kikuchi & Hashimoto bon seeps would markedly improve our un­ (2000), recendy described from the rela­ derstanding of the evolution and radiation tively shallow (~700 m) Mid-Okinawa of these shrimp among diverse chemosyn- Trough. However, Kikuchi & Hashimoto thetic environments throughout the world's (2000) point out that there is significant var­ oceans. iability in rostral characters, especially in A. hrevitelsonis where the ventral rostral mar­ Acknowledgments gin is usually toothed (there are seven spines on the ventral rostral margin of the We owe an enormous debt of gratitude holotype) but may be unarmed, especially to Austin B. Williams, whose early work on in young individuals. Ahnnocaris leuroko­ shrimp from hydrothermal vents established los lacks teeth on the ventral rostral margin, the general framework within which all and is thus more similar to the new species subsequent work on these shrimp is now A. williamsi. Alvinocaris williamsi is read­ viewed, and who unfortunately did not live ily distinguishable from both of those spe­ to see the completion of this species de­ cies in having a lightly pigmented eye; the scription. Special thanks to Ruth Gibbons eye is unpigmented in both A. hrevitelsonis for persevering to make sure that several and A. leurokolos. manuscripts, including this one, begun by The distribution of Alvinocaris species A. B. Williams before his death were within the global biogeographic vent prov­ brought to fruition, and to M. Oremland inces is centered in lower-latitude regions and K. Hiratsuka Moore for providing il­ (Shank et al. 1999), and the strong morpho­ lustrations. We thank the crews and pilots logical similarities among cxiani Alvinocar­ of the R/V Atlantis and the DSV Alvin for is species living in relatively shallow water their skillful collecting efforts, A. Gebruk vent and seep environments (<800 m) in for shipboard expertise, and the co-chief the Atlantic and Gulf of Mexico is striking. scientists of the expedition, R. C. Vrijen- Gebruk (1997) and others have hypothe­ hoek and R. A. Lutz; some of the shared sized that vent-endemic shrimp species are funding was via the National Science Foun­ derived from shallow-water seep ancestors. dation (OCE-96-33131). Finally, we thank A molecular phylogenetic approach to this T. Komai and A. L. Vereshchaka, and two hypothesis by Shank et al. (1999) suggested anonymous reviewers, for suggestions that that the analyzed vent-endemic Alvinocaris greatly improved the manuscript. This is (i.e., A. markensis, A. lusca, and unde- contribution No. 10700 of the Woods Hole 166 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Oceanographic Institution. TMS was sup­ Martin, J. W., & J. C. Christiansen. 1995. A new spe­ ported in part by the Postdoctoral Scholar cies of the shrimp genus Chorocaris Martin & Hessler, 1990 (Crustacea; Decapoda; Bresili­ Program at the Woods Hole Oceanographic idae) from hydrothermal vent fields along the Institution, with funding provided by the Mid-Atlantic Ridge,—Proceedings of the Bio­ Devonshire Foundation. JWM was support­ logical Society of Washington 108:220-227. ed in part by NSF grants DEB 9972100, , & G. E. Davis. 2001. An updated classifica­ DEB 9978193 (a FEET grant from the Sys­ tion of the Recent Crustacea.—Natural History tematic Biology program), and DEB Museum of Los Angeles County, Science Series No. 39:1-124. 0120635. , & R. R. Hessler. 1990. Chorocaris vandov- erae, a new genus and species of hydrothermal Literature Cited vent shrimp from the Western Pacific.—Contri­ butions in Science, Natural History Museum of Christoffersen, M. L. 1986. Phylogenetic relationships Los Angeles County 417:1-11. between Oplophoridae, Atyidae. Pasiphaeidae, Milne-Edwards, A., & E.-L. Bouvier 1894. Brachy- Alvinocarididae fam. n., Bresiliidae, Psalido- oures et anomures, Crustaces decapodes prov- podidae and Disciadidae (Cru.stacea Caridea enant des campagnes du yacht VHirondelle Atyoidea).—BoUetim de Zoologia 10:273-281. (1886. 1887, 1888). Premiere partie.—^Resultats Cola^:o, A., D. Desbruyeres, T. Comtet, & A.-M. des campagnes scientifiques accomplies sur son Alayse. 1998. Ecology of the Menez Gwen hy­ yacht par Albert ler, Prince Souverain de Mon­ drothermal vent field (Mid-Atlantic Ridge/ aco 7:1-112. Azores Triple Junction).—Cahiers de Biologic Segonzac, M., M. de Saint Laurent, & B. Casanova. Marine 39:237-^240. 1993. L'enigme du comportement trophique des Comtet, T, & D. Desbruyeres. 1998. Population struc­ crevettes Alvinocarididae des sites hydrother- ture and recruitment in mytilid bivalves from maux de la dorsale medio-atlantique.—Cahiers the Lucky Strike and Menez Gwen hydrother­ de Biologic Marine 34:535-571. mal vent fields (37°17'N and 37°50'N on the Shank, T. M.. R. A. Lutz, & R. C. Vrijenhoek. 1998. Mid-Atlantic Ridge).—Marine Ecology Pro­ Molecular Systematics of shrimp (Decapoda: gress Series 163:16.5-177. Bresiliidae) from Deep-Sea Hydrothermal Desbruyeres, D., et al. 2001. Variations in deep-sea Vents: enigmatic "Small Orange" Shrimp from hydrothermal vent communities on the Mid-At­ the Mid-Atlantic Ridge are juvenile Riinicaris lantic Ridge near the Azores plateau.—Deep- exoculata.—Molecular Marine Biology and Sea Research 48(5): 1323-1346. Biotechnology 7(2):88-96. Gebruk, A. V., S. V. Galkin, A. L. Vereshchaka, L. I. , M. B, Black, K. Halanych, R. A. Lutz, & R. Moskalev. & A. J. Southward. 1997. Ecology and biogeography of the hydrothermal vent fau­ C. Vrijenhoek. 1999. Miocene radiation of na of the Mid-Atlantic Ridge.—Advances in deep-sea hydrothermal vent shrimp (Caridea: Marine Biology 32:93-144. Bresiliidae): evidence from mitochondrial cy­ Hashimoto, J., O Suguru, T. Gamo, H. Chiba, T. Ya- tochrome oxidase subunit 1.—Molecular Phy- maguchi, S. Tsuchida, T. Okudaira, H. Watabe, logenetics and Evolution 13(2):244-254. T. Yamanaka, & M. Kitazawa. 2001. First hy­ Van Dover, C. L., S. B. Humphris, D. Fornari, C. M. drothermal vent communities from the Indian Cavanaugh, R. Collier, S. K, Goffredi, J. Hash­ Ocean discovered.—Zoological Science 18(5); imoto, M. D. Lilley, A. L. Reysenbach, T. M. 717-721. Shank, K. L. Von Damm, A. Banta, R. M. Gal­ Kikuchi, T, & J. Hashimoto. 2000. Two new caridean lant. D. Gotz, D. Green, J. Hall, T. L. Harmer, of the family Alvinocarididae (Crusta­ L. A. Hurtado, P. Johnson, Z. P. McKiness, C. cea, Decapoda) from a hydrothermal field at the Meredith, E. Okson, 1. L. Pan, M. Turnipseed, Minami-Ensei Knoll in the Mid-Okinawa Y. Won, C. R. Young, 01, & R. C. Vrijenhoek. Trough, Japan.—Species Diversity 5:135-148. 2001. Biogeography and ecological setting of -, & S. Ohta. 1995. Two caridean shrimps of the Indian Ocean hydrothermal vents.—Science families Bresiliidae and Hippolytidae from a 294:818-823, hydrothermal field on the Iheya Ridge, off the Vereshchaka, A. L. 1996a. Comparative analysis of Ryukyu Islands, Japan.—Journal of taxonomic composition of shrimps as edifica- Biology 15(4):771-785. tors of hydrothermal communities in the Mid- Lemaitre, R., & B. B. Collette. 2000. Austin Beatty Atlantic Ridge.—Doklady Biological Sciences Willliams (17 October 1919-27 October 1999). 351(l):576-578. Biographical summary. Proceedings of the Bi­ . 1996b. A new genus and species of caridean ological Society of Washington 113:1-12. shrimp (Crustacea: Decapoda: Alvinocarididae) VOLUME 116. NUMBER 1 167

from Nonh Atlantic hydrothermal vents.—Jour­ shrimp of the family Bresiliidae from thermal nal of the Marine Biological Association of the vents of the GaMpagos Rift.—Journal of Crus­ United Kingdom 76(4):951^961. tacean Biology 2:136-147. . 1997. A new family for a deep-sea caridean ~, & R C. Dobbs. 1995. A new genus and spe­ shrimp from North Atlantic hydrothermal cies of caridean shrimp (Crustacea: Decapoda: vents.—Journal of the Marine Biological As­ Bresiliidae) from hydrothermal vents on Loihi sociation of the United Kingdom 77:425-438. Seamount, Hawaii.—Proceedings of the Biolog­ Williams, A. B. 1988. New marine decapod crusta­ ical Society of Washington 108:228-237. ceans from waters influenced by hydrothermal —, & R A. Rona. 1986. Two new caridean discharge, brine, and hydrocarbon seepage.— shrimps (Bresiliidae) from a hydrothermal field Fishery Bulletin 86:263-287. on the Mid-Atlantic Ridge.—Journal of Crus­ , & F. A. Chace, Jr 1982. A new caridean tacean Biology 6:446-462.