The Carcinological Society of Japan

CRUSTACEAN RESEARCH, NO. 30: 160-171, 2001

A new of the genus Periclimenes (: : ) collected from hydrothermal vent fields in Kagoshima Bay, Japan

Ken-Ichi Hayashi and Jun Ohtomi

Abstract.—Based on material col­ Mirocarididae Vereshchaka, 1997 are the lected from shallow hydrothermal vent typical vent carideans (Gebruk et al., fields in Kagoshima Bay, southern Japan, 1997). No species of the family a neAV species of the subfamily Palaemonidae has been known from these Pontoniinae, Periclimenes environments so far, while a few species thertnohydrophiluSy is described and il­ of Hippolytidae, Crangonidae and lustrated. A combination of several char­ have been reported acters distinguishes the present new^ spe­ (Desbruyeres & Segonzac, 1997; cies from all other members of this genus. Tunnicliffe et al, 1998). A short and narrow rostrum w^ithout Periclimenes is a large genus contain­ basal crest, an isolated epigastric spine, a ing more than 170 species and most spe­ submarginal antennal spine and a long cies are commensal with scleractinean carpus of the second pereopod are char­ corals or their associated invertebrates in acteristics of the new^ species. The rela­ tropical and subtropical waters (Chace & tionship betw^een this shrimp and host Bruce, 1993; Bruce, 1994). Recently sev­ tube w^orms is briefly discussed. eral species of this genus have been de­ scribed from deep waters, more than 100 m deep (Bruce, 1990, 1991b, 1996). Al­ Introduction though collected from a curious and com­ Recently shallow hydrothermal vent paratively deep habitat, the present spe­ fields were explored in the eastern end of cies has the general form of the genus. Kagoshima Bay and an interesting associ­ Careful comparison with the related ated fauna was discovered (Hashimoto et deep-water congeners, however, reveals al, 1993; Miura et al, 1997). The junior that the present species is distinguished author and his colleagues carried out from them in having a short and narrow many cruises while researching the deca­ rostrum, an isolated epigastric spine, a pod fauna of Kagoshima Bay including submarginal antennal spine, and un­ these vent fields. Considerable numbers armed long carpus of the second pereo- of unusual shrimps were found in the pods. tube worm colonies at depths of about 100 All specimens examined are deposited m. They constitute a single species of the in the collection of the National Fisheries genus Periclimenes Costa, 1844. This is University (NFU). The carapace length the first species of the family Palae­ (CL) represents the specimen size, mea­ monidae collected from the hydrothermal suring from the posterior margin of the vent fields. Species of the families orbit to the posterior margin of the cara­ Alvinocarididae Christoffersen, 1986 and pace.

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Systematic account ginal, situated just inside anterior margin of carapace, apex reaching rounded sub­ Periclimenes thermohydrophilus orbital angle (Fig. 3D). Anterolateral new species margin from suborbital angle to Figs. 1-4 pterygostomial angle gently convex. Pterygostomial angle largely rounded. Material examined,—Holotype, ovig. Abdominal somites smooth, glabrous. $ (NFU 530-2-2367, 6.1 mm CL), Few simple setae on terga of fifth and Kagoshima Bay, "Shinkai 2000," dive sixth somites only. Third somite not pro­ 980, depth 90-100 m, associated with duced posteriorly. Pleura of first three Lamellibrachia satsuma, 30 September somites broadly rounded; those of fourth 1997, coll, T. Miura. and fifth somites produced posteriorly, Paratypes: 2 ovig. 9 (NFU 530-2-2369, not sharply pointed, fifth more pointed CL 6,0, 7.0 mm), Kagoshima Bay, "Dol­ than fourth (Fig. 3E). Sixth somite 1.8 phin 3K;'d\ye 340, 31°39.5'N, 130°48.2'E, times as long as fifth somite and 1.4 times depth 97 m, associated with as long as deep, posterolateral and Lamellibrachia satsuma, 1 September posteroventral angles both acute (Fig. 1997, coll. T. Miura; — 11 ovig. 9 (NFU 3E). Telson 1.4 times as long as sixth 530-2-2368, CL 4,4-6.6 mm), 1 9 (CL 5,3 somite, moderately slender, lateral mar­ mm), collected together with holotype; — gin nearly straight. Dorsolateral spines in 4 S (NFU 530-2-2370, CL 3.4-4.4 mm), 1 2 pairs, but small additional spine distal $ (CL 6.8 mm), northern part of to distal pair on left side, proximal pair at Kagoshima Bay, depth 105 m, hydrother- midlength of telson, distal pair at mal vent field, 20 April, 2001, coll. J. midlength between proximal pair and dis­ Ohtomi. tal margin. Distal margin with 1 central Description of holotype.—Moderate spine and 3 pairs of unequal spines (Fig. sized species (Fig. 1). Rostrum com­ 3F). pressed, slender and gradually tapered; Eyes moderate, with well-pigmented directed straight forward, not reaching cornea, with small ocellus, eyestalk distal end of antennular peduncle (Fig. slightly longer than cornea, swollen me- 3A), Lateral carina not developed, dorsal sially (Fig. 3G). Antennular peduncle margin with 8 teeth, posterior 4 teeth moderate, reaching distal margin of an­ closely set, of which posterior 2 on cara­ tennal scale (Fig. 3B). First segment of pace, third tooth just above orbital mar­ peduncle twice as long as wide, with me­ gin; 3 equidistant teeth on distal half of sial margin straight, entirely setose, with ventral margin; distal tooth small (Fig. small ventral spine; lateral margin 3A). Carapace glabrous, smooth, with slightly convex mesially ending in acute shallow hepatic depression below hepatic spine reaching nearly midlength of sec­ spine posterior to anterior half of cara­ ond segment; distolateral lobe developed pace. Epigastric spine situated at anterior (Fig. 3B, H). Stylocerite acute, slender, third of carapace, considerably separated overreaching midlength of first antennu­ from last tooth of rostral series (Fig. 3A, lar segment. Second and third antennular B); dorsal margin between epigastric segments cylindrical, equal in length, spine and last rostral tooth convex in lat­ much slenderer than first segment; lat­ eral view; indistinct tubercle present just eral and mesial margins of second seg­ posterior to epigastric spine (Fig. 3A). ment setose. Upper antennular flagellum Middorsal carina vanishing behind tu­ biramous, with 7 or 8 proximal segments bercle. Antennal spine slender, submar- fused and unarmed. Shorter free ramus

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162 K. HAYASHI & J. OHTOMI

Fig. 1. Periclimenes thermohydrophilus new species. Holotype, ovigerous female, CL 6.1 mm (NFU 530-2-2367) from Kagoshima Bay. Scale = 1.0 mm.

with about 10 segments, as long as fused both setose distally; coxal endite obsolete; portion, each segment with few scaphognathite well developed (Fig. 4C). aesthetascs; longer ramus not complete, First maxilliped with slender tapering more than twice as long as shorter ramus. palp, reaching distal margin of basal Lower flagellum slender, filiform, not endite, with subterminal seta; basal complete, as long as upper flagellum. An­ endite large, broadly rounded distally, tenna with basicerite bearing small densely setose; coxal endite small, thick­ distolateral tooth; dorsodistal corner pro­ ened, sparsely setose; exopod with flagel­ duced, but blunt. Carpocerite cylindrical, lum well developed, broad, with several reaching proximal third of scaphocerite. long plumose setae, caridean lobe large, Scaphocerite broad, 0.6 times as long as epipod large, weakly bilobed (Fig. 4D). carapace, 2.4 times as long as wide; lat­ Second maxilliped with normal endopod eral margin nearly straight; distolateral with distal segment short and wide setose tooth stout, slightly falling short of distal distally; exopod long, with several long margin of rounded blade. Flagella not setae distally and short setae on mesial complete, extending forward to distal end margin; epipod large, rectangular (Fig. of second chela. 4E). Third maxilliped normal, moderately Mouthparts (Fig. 4A-F) not different elongate and slender, reaching rostral from typical form of genus. Mandible apex (Fig. 4F). Distal segment little without palp; incisor process well devel­ shorter than half of third segment, taper­ oped, with 4 acute teeth, molar process ing distally, mesial margin with several stout, with several blunt teeth (Fig. 4A). groups of strong setae; second segment First maxilla with bilobed palp, upper 1.2 times longer than distal segment, me­ and lower laciniae densely setose distally sial margin sparsely setose; third seg­ (Fig. 4B). Second maxilla with simple na­ ment faintly articulated with basis, me­ ked palp; basal endite deeply bilobed, up­ sial margin with sparse long setae, per lobe slightly larger than lower lobe. distolateral part with 2 (right) or 4 (left)

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NEW PERICLIMENES FROM HYDROTHERMAL VENT FIELDS 163

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,;<^ •-•"•• •^: "^miil^} .•,.-»- -J ..-.;,,• -•-;'--*.-;•-: S.;---!?;-'-.-",' >>l - ••;, ;—;„'.^,-;-'-,' ,j' '.'. °. '.'"-., V" r " * *-^ , .. ..-.,•.:-!-•.'•....-.-•<:'->• ::*^zy- .,•„ . •_- c. -%^ rvi.r . ,, -•.•.-r,-.-r'--T •:;.-4

Fig. 2. Periclimenes thermohydrophilus new species. Paratype, female, CL 6.8 mm (NFU 530-2- 2370) from Kagoshima Bay. Lateral (upper) and dorsal (lower). Scale = 0.5 mm.

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164 K. HAYASHI & J. OHTOMI

spinules; basis sparsely setose; exopod cessory tooth overreaching midpoint of long, with distal setae only; coxa feebly unguis (Fig. 4L). Propodus 4.8 times as produced mesially, sparsely setose, with long as dactylus; ventral margin with 2 small oval plate laterally, without rather long distoventral spines, preceded arthrobranch. by 5 separate spines, all spines with few Epistome with pair of low rounded pro­ setae; dorsal margin sparsely setose (Fig. cesses. Fourth thoracic sternite without 4L). Carpus about half as long as median process but with posterior plate propodus, unarmed; merus about 0.9 as on fifth sternite; these posterior plates times as long as propodus, unarmed; is­ with median deep incision. chium about 0.6 times as long as merus, First pereopod slender, exceeding dis­ unarmed (Fig. 4K). Fourth and fifth tal end of antennular peduncle by chela pereopods similar in shape and propor­ (Figs. 1, 4G); palm of chela cylindrical, tion (Fig. 4M, N); propodus of fifth pereo­ with 5 rows of short setae on ventral sur­ pod with row of 4 or 5 separate spines on face near carpal articulation; fingers 1.8 ventral margin, and one pair of spines on times as long as palm, bearing some distal end, all obscured by 5 rows of setae groups of setae, cutting edge smooth (Fig. (Fig. 4P). 4H). Carpus 1.6 times as long as chela, cy­ First pleopods (female) unequally bi- lindrical, slightly tapered proximally, ramous. Endopod short, about one-fourth with tuft of long setae distoventrally. length of exopod, both densely setose. Sec­ Merus 0.8 times as long as carpus and 1.4 ond to fifth pleopods biramous in nearly times as long as chela, slightly swollen equal length, densely setose, endopod centrally. Ischium as long as chela, ob­ with long slender appendix interna. Uro- liquely articulated with basis (Fig. 4G). pods longer than telson, protopod with Second pereopods similar in shape and acute posterolateral process; endopod very slightly unequal in length; both legs slightly shorter than exopod, about 4 exceeding distal end of antennular pe­ times as long as broad; exopod ending in duncle by carpus and chela (Fig. 1); fin­ small spine on outer distal end, accompa­ gers, carpus and merus 1.1 times and nying with longer movable spine just in­ palm 1.3 times longer in left than in right; side; diaeresis well marked, smooth (Fig. chela smooth, glabrous, cylindrical, 1.3 3F). (right) or 1.5 (left) times longer than cara­ Variation of paratypes.—Rostrum not pace. Fingers 0.7 (right) or 0.6 (left) times extending beyond distal end of antennu­ as long as palm, with strongly hooked lar peduncle. Of 19 paratypes 18 with in­ tips, sparsely setose; cutting edges sharp, tact rostrum, comparative length var3dng entire on distal three-fifths, and with sev­ from 0.50 to 0.68 times as long as cara­ eral small teeth on remaining proximal pace. Armature also variable, upper mar­ part (Fig. 4J). Carpus 0.9 (right) or 0.8 gin with 7 teeth in 2 specimens, 8 teeth in (left) times as long as palm, unarmed, ex­ 9 specimens, 9 teeth in 5 specimens, and panded distally. Merus 1.0 (right) or 0.9 10 teeth in 2 specimens, 2 (in 5 speci­ (left) times as long as palm, cylindrical, mens) or 3 teeth (in 13 specimens) of these unarmed; ischium slightly shorter than on carapace. Lower margin with 1 tooth in merus, cylindrical, unarmed (Fig. 41). 3 specimens, 2 teeth in 13 specimens, and Ambulatory pereopods slender, simi­ 3 teeth in 2 specimens. One male (3.4 mm lar in shape and size, all overreaching tip CL) entirely lacking epigastric spine or of first pereopod by dactylus (Fig. 4K, M, small tubercle; large specimens usually N). Dactylus of third pereopod gently with both but remaining males with epi­ curved, simply biunguiculate, lower ac­ gastric spine only.

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Fig. 3. Periclimenes thermohydrophilus new species. A, B, D-F, holotype, ovigerous female, CL 6.1 mm (NFU 530-2-2367); C, J-K, paratype, male, CL 4.4 mm (NFU 530-2-2370); G-I, paratype ovigerous female, CL 6.8 mm (NFU 530-2-2368); all from Kagoshima Bay: A, anterior part of carapace, with cephalic appendages, lateral; B, same, dorsal; C, carapace, lateral; D, anterior margin of carapace, lateral; E, posterior part of abdomen, lateral; F, tail fan, dorsal; G, eye, dorsal; H, first antenna, ventral; I, second antenna, ventral; J, endopod of left first pleopod; K, appendix interna and appendix masculina of left second pleopod. Scales for A-I = 1.0 mm; scale for J, K = 0.2 mm.

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Fig. 4. Periclimenes thermohydrophilus new species. A-F, paratype, ovigerous female, CL 6.8 mm (NFU 530-2-2368); G-N, P, holotype, ovigerous female, CL 6.1 mm (NFU 530-2-2367); O, paratype, male, CL 4.4 mm (NFU 530-2-2370); all from Kagoshima Bay; A- N, left side; O, P, right side: A, mandible; B, first maxilla; C, second maxilla; D, first maxilliped; E, second maxil- liped; F, third maxilliped; G, first pereopod, lateral; H, same, chela, mesial; I, second pereopod, lateral; J, same, chela, lateral; K, third pereopod, lateral; L, same, distal two segments, mesial; M, fourth pereopod, lateral; N, fifth pereopod, lateral; O, same, distal two segments, lateral; P, same, dactylus and distal part of propodus, lateral. Scales = 1.0 mm.

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Sixth abdominal somite 1.7-2.1 times Males smaller and more slender than as long as fifth somite and 1.4-1.8 times females, especially shorter and slenderer as long as deep. Telson 1.3-1.6 times as in pereopods but not clearly different long as sixth somite. Dorsolateral spines from females (Figs. 3C, 40). Endopod of usually in 2 pairs, one specimen with only male first pleopod leaf-shaped, broadened 1 pair, two specimens with unequally 1 distally, outer distal margin convex, with spine or 3 spines on either side. Upper 10 plumose setae; inner margin slightly antennular flagellum with 5—7 basal concave, with a rather long plumose seta joints and biramous distal part; shorter near proximal end and 4 short simple se­ ramus composed of 10-14 free joints, as tae on proximal half (Fig. 3J). Male sec­ long as or slightly longer than carapace. ond pleopod equally biramous. Endopod Longer ramus of upper flagellum and with appendix interna and appendix lower flagellum, distal part missing, more masculina, appendix interna longer than than 3.5 times as long as carapace. corpus of appendix masculina, with 5 Scaphocerite 1.9-2.7 times as long as simple setae on distal half of mesial mar­ wide; outer distal spine shorter or longer gin and 5 unequal apical setae (Fig. 3K). than distal margin of lamella. Distal part Size.—The holotype is an ovigerous fe­ of antennal flagellum usually missing, male, 6.1 mm CL and about 22.5 mm in but more than 5 times as long as carapace total length. The paratypes are 4.6—7.0 length. mm CL in ovigerous females and 3.4—4.4 Two or more spinules on third segment mm CL in males. Eggs small, 0.43—0.45 x of third maxilliped present; of 18 0.46-0.50 mm in uneyed eggs and 0.50— paratypes with third maxillipeds on both 0.53 X 0.63-0.66 mm in eyed eggs. The sides, 8 specimens with 3 spinules, and 2 holotype carries 260 uneyed eggs, the specimens with 2 or 4 spinules on both smallest ovigerous female, 4.4 mm CL, sides, 3 specimens unequally with 2 and 3 has 140 eyed eggs and the largest one, 7.0 spinules, one specimen with 3 and 4 mm CL, has 510 uneyed eggs. spinules and 2 specimens with unequally Color.—Body entirely crimson red 5 and 6 spinules on one side; number of (Fig. 2). Rostrum and uropod semitrans- spinules related neither to sex nor speci­ parent, with many red chromatophores. men size; these spinules usually situated Cornea black, eyestalk and antennular at distal third of lateral surface in case of peduncle crimson red. Antennular fla- less than 3 spinules, but posterior 1 or 2 gella, antennal flagellum and thoracic ap­ spinules at midlength of segment in case pendages including second pereopod red­ of 4 or more spinules. dish orange. Seven specimens bearing pair of sec­ Etymology.—The species name, ond pereopods, 4 specimens with second thermohydrophilus, is the combination of pereopod on only one side and 8 speci­ the Greek, "thermos" (= hot), "hydro" (= mens lacking those from both sides. One water), and "philos" (= loving, fond of), male (4.4 mm CL) with nearly equal pair, with reference to the habitat preference of but other females with slightly unequal the new species. All materials examined chelipeds, all reaching beyond were collected from submarine hot spring scaphocerite by carpus and chela or chela areas of Kagoshima Bay. only. Major chela 1.05-1.60 times as long Distribution.—Known only from the as minor chela; each segment of major eastern end of Kagoshima Bay at depths and minor chelipeds with nearly same of 90-115m. The shrimps were always proportion as chela. Carpus 0.8-1.0 times collected from in or around colonies of as long as palm; merus as long as or Lamellibrachia satsuma Miura, slightly longer than carpus. Tsukahara & Hashimoto, 1997 and an-

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other unidentified polychaete tube worm, Periclimenes ordinarius differs from the growing at the hydrothermal vent fields. present new species in the large hepatic spine on the carapace and the ogival Discussion shaped cornea. Periclimenes richeri and P. ordinarius bear a tubercle instead of an Morphological comparison with congeners epigastric spine. In the new species the The present new species differs from hepatic spine and the cornea are of nor­ more than 130 shallow-water species of mal shape and size. The epigastric spine Periclimenes in having the combination of usually is present far from the most poste­ the following characters. The carapace is rior tooth of the rostral series and the dor­ provided with an antennal spine and an sal margin between them is concave in epigastric spine or tubercle, but without a lateral view. A tubercle is also discernable supraorbital spine. The rostrum is slen­ just behind the epigastric spine in large- der, without dorsal crest. The carpus, sized specimens. These features of the merus and ischium of the second pereo- epigastric region are characteristic of the pod are unarmed. The ambulatory pereo­ new species. pods have biunguiculate dactyli. Two In P. latipollex, P. ordinarius, and P. pairs of spines are present on the dorso­ richeri, the rostrum is relatively long, lateral surface of the telson. These char­ reaching or overreaching the distal end of acters, on the other hand, are shared by the antennular peduncle, but falls short the following seven deep-water conge­ of it in the new species. In P. involens the ners: P. involens Bruce, 1996, P. rostrum does not reach the end of the an­ laccadivensis (Alcock & Anderson, 1894), tennular peduncle, but it is actually long, P. latipollex Kemp, 1922, P. ordinarius as long as the carapace length, while in Bruce, 1991b, P. richeri Bruce, 1990, P. the new species it is only 0.5-0.7 times as tenuirostris Bruce, 1991a, and P. vaubani long. Bruce, 1990. In P. latipollex, P. ordinarius, P. The new species, however, differs from richeri and P. vaubani, the accessory these deep-water species in having the tooth on the ventral margin of the corpus submarginal antennal spine, the base of of the ambulatory dactyli is much smaller which is located slightly inside the ante­ than the unguis, while in the remaining rolateral margin of the carapace and no four species including the new species it is part of the base of the spine is on the mar­ a considerable size. gin. The antennal spine is marginal in all other species. The second pereopods in Consideration to the distribution the new species are similar in shape and Periclimenes thermohydrophilus is the usually unequal in length, and the carpus only member of the family Palaemonidae is 0.8 or 1.0 times of the palm length. In­ represented at hydrothermal vent fields cluding P. laccadivensis and P. vaubani, (Desbruyeres & Segonzac, 1997; which have the unequal second pair of Tunnicliffe et al, 1998). The shrimp was pereopods, all other species have a short always found from the samples sucked carpus of the second pereopod, less than from colonies of hard tubes of the half of the palm length, although in P. or­ vestimentiferan worm, Lamellibrachia dinarius and P. richeri, only one side is satsuma, as well as soft tubes of unidenti­ known. fied polychaete tube worms. These worms The new species, moreover, differs are the typical associates of the so-called from P. tenuirostris in having smooth ab­ "tagiri," hydrothermal vent fields, around dominal somites, not strongly produced the eastern end of Kagoshima Bay at into a carinate lobe on the third somite. depths of about 100 m (Miura et al., 1997).

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Judging from the collecting data, the was originally collected from an echinoid shrimp is restricted to these vent fields host, PhorTTiosoma sp. (Balss, 1913). The only and has not been obtained from any subsequent collections, moreover, were other parts of that bay, where more than from the echinoids, Areosoma thetidis (H. 300 hauls were taken by the junior author L. Clark) or A. owstoni Mortensen (Bruce, from 1993. This is probably the first 1975; Okuno & Minemizu, 1998). Bruce pontoniine shrimp in association with the (1990) also suggested echinoid hosts for P. vestimentiferan or polychaete tube curvirostris^ worms (Bruce, 1976; Chace & Bruce, The body color of the new species is 1993; Monod & Laubier, 1996), though entirely crimson red and clearly differs the relationship between this shrimp and from cryptic, disruptive or transparent the host invertebrate has not clearly been patterns often seen in the shallow-water established. congeners (Bruce, 1976). The body does The genus Periclimenes is mostly com­ not show any unusual parts and the eggs posed of shallow-water inhabitants are small and numerous in number, (Chace & Bruce, 1993), whereas most of which means that the shrimp probably the specimens we examined were col­ has long-lived planktotrophic larvae and lected at about 100 m deep, which is con­ is highly mobile adults. This shrimp, sidered to be deep for most species of this therefore, should be found from other genus. This is not an extreme depth, be­ shallow water hydrothermal vent fields. cause about 35 species have been reported from depth of more than 100 m (Bruce, Acknowledgments 1991b, 1996). We would like to thank Dr. Tomoynki The following five species have been Miura of Miyazaki University for collect­ collected from deep waters around Japan; ing and donating this interesting mate­ P. alcocki Kemp, 1922 from Tosa Bay, and rial for us and Dr. Mary K. Wicksten of Kumanonada, in 311-400m (Kubo, 1940; Texas A&M University and Dr. Junji Hayashi, 1986), P. curvirostris Kubo, Okuno of Coastal Branch of Natural His­ 1940 from off Owase, Mie Prefecture, in tory Museum and Institute, Chiba for 311 m (Kubo, 1940), P. granuloides critical reading of the manuscript. The Hayashi, 1986 from Tosa Bay in 130 m junior author thanks Dr. Jun Hashimoto (Hayashi, 1986), P. hertwigi Balss, 1913 of Japan Marine Science and Technology from Suruga Bay to East China Sea off Center, technical operators and crew of Kushikino, Kagoshima Prefecture, in 45- "Shinkai 2000," "Dolphin 3K" and the 311 m (Balss, 1913, 1914; Kubo, 1940; mother ship "Natsushima" for capable Fujino & Miyake, 1970; Okuno & support at sea. Minemizu, 1998) and P. macrophtalmus Fujino & Miyake, 1970 from East China Literature Cited Sea, off Goto Islands in 145 m (Fujino & Alcock, A., & Anderson, A. R., 1894. An ac­ Miyake, 1970). P. curvirostris, P. count of a recent collection of deep-sea granuloides and P. macrophtalmus have Crustacea from the Bay of Bengal and Laccadive Sea. Natural History Notes from only been known from the type locality, H. M. Indian Marine Survey Steamer "In­ while P. alcocki and P. hertwigi have also vestigator", Commander C. F. Oldham, R. been recorded from other localities of the N., commanding. Ser. II, No. 14, Journal of Indo-West Pacific region at depths down Asiatic Society of Bengal, 63: 141-185, pi. to more than 600 m (Chace & Bruce, 9. 1993). Balss, H., 1913. Diagnosen neuer Ostasiatischer Macruren. Zoologischer The hosts of these deep-water shrimps Anzeiger, 42: 234-239. are not known, except for P. hertwigi. It , 1914. Ostasiatsche Decapoden 11. Die

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vances in Marine Biology, 34: 355-442. Aquabiology, National Fisheries University, 2- Vereshchaka, A. L., 1997. A new family for a 7-1 Nagata-honmachi, Shimonoseki 759-6595, deep-sea caridean shrimp from North At­ Japan; (JO) Faculty of Fisheries, Kagoshima lantic hydro thermal vents. Journal of the University, 4-50-20 Shimoarata, Kagoshima Marine Biological Association of the 890-0056, Japan United Kingdom, 77: 425-538. E-mails: (KH) [email protected]; (JO) ohtomi@fish .kagoshima-u .ac.jp

Addresses: (KH) Department of Applied

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