23 April 2003

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1I6(1):I58-I67. 2003.

A new caridean shrimp 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: tshank@whoi.edu;
(JWM) Natural History Museum of Los Angeles County, 900 Exposition Boulevard,
Los Angeles, California 90007, U.S.A., e-mail: jmartin@nhm.org

Abstract.—A new species of alvinocaridid shrimp, Alvinocaris williamsi, is

described from the Menez Gwen hydrothermal vent 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
genus recently described from the M i d - O k i n a w a Trough, Japan. Molecular phy-
logenetic studies of A. williamsi, the eighth described m e m b e r 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.

T h e caridean shrimp family Alvinocari- T h e 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. T h e family was further diagnosed by Ensei Knoll in the M i d - O k i n a w a 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; W i l l i a m s , 1 9 8 8 , from the M i d - A t l a n t i c
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. T h e 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 N e w 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 crabs, 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 taxonomy,
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 crab 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-
cies of the shrimp genus Chorocaris Martin &
ported in part by the Postdoctoral Scholar
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
Museum of Los Angeles County, Science Series
tematic Biology program), and DEB No. 39:1-124.
0120635. , & R. R. Hessler. 1990. Chorocaris vandov-
erae, a new genus and species of hydrothermal
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