JOURNAL OF BIOLOGY, 1 l(4): 583-593, 199 1

HAMALPHEUS ACANTHOPS, NEW , NEW SPECIES, A STYGIOPHILIC ALPHEID SHRIMP FROM A SAMOAN LAVA TUBE

A. J. Bruce and Thomas M. Ilzfle

ABSTRACT A new stygiophilic alpheid shrimp from Upolu, Samoa Islands, is described and illustrated. Hamalpheus acanthops, new genus, new species, is the only alpheid shrimp definitely known from a marine lava tube. The new genus is characterized by a conspicuous spinifom process of the anterior eyestalk and a modified caudal fan with unique holdfast adaptations. The new genus is most closely related to Betaeopsis Yaldwyn, from which it is distinguished by the presence of the above-mentioned features. A key is also provided to the 29 alpheid genera currently known from the Indo-West Pacific Region.

Alpheid shrimps are abundant in marine Second pereiopods slender, carpus 5-seg- habitats, particularly in the tropics and on mented. Ambulatory pereiopods robust, coral reefs. Few occur in brackish or fresh dactyls biunguiculate; first and second per- waters and their occurrence in troglobitic eiopods with epipods. Uropod with proto- situations is particularly rare. Potamal- pod distolaterally acute, exopod with well- pheops stygicola (Hobbs, 1973) has been re- developed diaeresis, proximal margin with ported from a fresh-water cave in Mexico single lateral spine only; endopod with pair and two species of Metabetaeus, M. lohena of stout spines distally. Banner and Banner, 1960, and M. minutus (Whitelegge, 1897) occur in anchialine lava Type Species. -Hamalpheus acanthops, new and coral pools (Holthuis, 1986). The pres- species. ent shrimp appears to be the first alpheid Systematic Position. -Hamalpheus is most shrimp to be found in a marine lava tube. closely related to. the genera Betaeus Dana, SYSTEMATICACCOUNT 1852, and BetaeopsisYaldwyn, 197 1. These three genesa share the following features: Hamalpheus, new genus rostrum absent; eyes completely covered by DeJinition.-Body subcylindrical, feebly anterior carapace; carapace spines com- compressed, smooth, glabrous. Rostrum pletely lacking; first pereiopods with chelae absent; carapace without supracorneal, su- camed in extended position, with dactyl praorbital, extracorneal, or infracorneal ventrally, fingers without molar process and spines; sixth abdominal segment with pos- fossa; second pereiopods with 5-segmented teroventral angle nonarticulate; pleura of carpus; ambulatory pereiopods robust, only first 4 abdominal segments rounded, fifth first and second with epipods; uropod with posteroventrally acute; telson without dor- diaeresis of exopod nondenticulate. The sal spines, anal tubercles absent. Antenna1 closely related genera Betaeus and Betaeop- peduncles robust; stylocerite well devel- sis are distinguished primarily by the pres- oped; basicerite with large lateral tooth, ence of a moveable articulated plate at the scaphocerite normal; antenna1 flagellum ro- posteroventral angle of the sixth abdominal bust, semirigid proximally; eyes with cor- segment in the former genus and its absence nea completely covered in dorsal view, stalk in the latter. In this feature, Hamalpheus with stout, acute anterior tooth. Mouthparts resembles Betaeopsis. Hamalphas may be normal; mandible with 2-jointed palp; mo- distinguished from Betaeopsis by (1) the ab- lar process well developed, incisor process sence of dorsal telson spines, associated with dentate; third maxilliped with slender en- (2) the recurved inner posterior telson spines dopod, not suboperculate. First pereiopods and (3) the strong spines at tip of the exopod small, robust, subequal, similar; chelae car- of the uropod, and (4) the acute spinous ried extended (?), palm tuberculate medi- processes on the anterior aspect of the eye ally, fingers without molar tooth and fossa. stalk. The mouthparts in the two genera are 584 JOURNAL OF CRUSTACEAN BIOLOGY, VOL I I, NO 4, 1991 essentially similar. In Hamalpheus the epi- locerite well developed, slender, acute, pod of the first maxilliped is more triangular reaching to about level of distal margin of and bilobed than in Betaeopsis and that on intermediate segment; statocyst normal; in- the second maxilliped more triangular than termediate segment subcylindrical, with oval. On the third maxilliped, the lateral feeble ventromedial lamina, about as long plate of the coxa is distinctly acute in Ham- as wide; distal segment subcylindrical, alpheus, but blunt in Betaeopsis. slightly compressed, about 1.6 times longer Etymology.-From hamus, Latin, a hook, than wide; upper flagellum feebly biramous, refemng to the spines on the caudal fan, rami fused for proximal 11 segments, com- and Alpheus, a generic name first used by pressed, each segment with paired groups Fabricius (1 798). of aesthetascs ventrally; shorter free ramus with single incompletely free segment; lon- Hamalpheus acanthops, new species ger free ramus slender, about 0.6 of carapace Figs. 1-5 length; lower flagellum slender, filiform, subequal to carapace length. Material Examined. - 1 P, Station 83--034, Tosua-To- Antenna with stout basicerite, with con- lesua lava tube cave located near Lotofaga village, south spicuous triangular laminar tooth antero- coast of Upolu Island, Western Samoa, 17 April 1988, laterally; carpocerite stout, far exceeding collected by T. M. Iliffe and S. Sarbu. scaphocerite and antennular peduncle, sub- Description. -Body form subcylindrical, cylindrical, about 2.6 times longer than dis- slightly compressed (particularly carapace), tal width; merocerite and ischiocerite nor- smooth, glabrous. mal; flagellum about 2.0 times carapace Carapace with rostrum completely lack- length, proximal segments fully fused, prox- ing, frontal region broad, feebly concave, imal half of flagellum semirigid, distal half completely covering eyes in dorsal view, more flexible; scaphocerite well developed, without dorsal canna; supracorneal, extra- reaching to about middle of distal segment corneal, and infracorneal spines or teeth ab- of antennular peduncle, broad, about 1.75 sent; anterior margin of branchiostegite times longer than wide, lateral margin fee- slightly produced, broadly rounded; cardiac bly convex, with strong distolateral tooth, notch distinct. reaching about to distal margin of lamella. Abdomen with pleuron of first 3 segments Eye completely concealed dorsally by car- broadly rounded, fourth slightly produced apace, cornea hemispherical, well pigment- posteriorly, rounded, fifth strongly pro- ed, obliquely orientated on stalk; stalk ovoid, duced posteriorly, posteroventrally acute; with large acute tooth projecting anteriorly sixth segment about 1.3 times length of fifth, from anterior margin adjacent to cornea, 1.3 times longer than deep; posterolateral without setae. angle slender, acute; posteroventral angle Mandible (right) robust, with 2-segment- blunt, nonarticulate. Telson subequal to ed palp, proximal segment irregularly sub- sixth segment length, strongly convex dor- cylindrical, slightly expanded distally, non- sally, concave ventrally, without anal tu- setose, distal segment laminar, oval, with bercles, about 1.6 times longer than anterior feebly plumose marginal setae; molar pro- width, lateral margins feebly sinuous, con- cess well developed, subcylindrical, vergent, without dorsal spines, posterior obliquely truncate distally, with dense tuft margin about 0.5 of anterior width, with of longer setae proximoanteriorly; incisor small acute lateral tooth, convex, with small process very stout, deeply concave medi- inner and outer spines laterally, inner spine ally, with 5 stout acute teeth distally. Max- smaller, curved dorsomedially, outer spine illula with bilobed palp, upper lobe with 2 longer, straight, projecting posteriorly, low- setae, lower lobe with 1 longer stouter seta; er convex margin with 13 long slender seg- upper lacinia broad, with double row of 14 mented plumose setae projecting posteri- short stout spines distally, with numerous orly, upper margin with numerous shorter feebly setulose setae distodorsally, coarsely simple setae projecting dorsally. setulose setae distoventrally; lower lacinia Antennular peduncle, short, robust; prox- slender, with 4 stout spines distally, nu- imal segment about 1.9 times longer than merous serrulate setae. Maxilla with short, wide, with large ventromedial tooth, sty- tapering, angulate palp, with short simple BRUCE AND ILIFFE: NEW STYGIOPHILlC ALPHEID SHRIMP FROM SAMOA

Fig. 1. Hamalpheus acanthops, new genus, new species, holotype female, Upolu, Samoa. Scale bar in mm. distal setae, several short weakly plumose without medial protuberance, sparsely se- setae proximolaterally; basal endite large, tose, with large triangular simple epipod bilobed, lobes subequal, broad, densely laterally, without podobranch. Third max- fringed with short setae medially; coxal en- illiped with endopod slender, not suboper- dite reduced, angular, with few long setae; culate, ischiomerus and basis completely scaphognathite well developed, narrow, fused, about 5.6 times longer than central about 3.5 times longer than wide; anterior width, subuniform, with few short setae dis- lobe 1.75 times longer than wide, distal half tally on lateral margin, medial margin gen- narrow; posterior lobe well developed, about erally sparsely setose; carpal segment short, 0.3 of scaphognathite length. First maxilli- 2.0 times longer than central width, about ped with slender tapering simple palp, with 0.23 of antepenultimate segment length, slender feebly plumose setae along medial centrally slightly swollen, tapering distally, margin and distally, basal endite broad, with sparsely setose laterally, small groups of se- feebly plumose setae along anterodistal tae medially; terminal segment slender, margin, dense fringe of short feebly setulose about 0.5 of antepenultimate segment setae along medial margin; coxal endite length, 6.0 times longer than wide, tapering weakly bilobed, sparsely setose; exopod well distally, with subdivision into larger prox- developed, flagellum slender, with numer- imal and smaller distal segments, with sin- ous plumose setae distally, with small, nar- gle spine distally, with numerous groups of row caridean lobe; epipod large, triangular, long slender finely serrulate setae on ven- weakly bilobed. Second maxilliped of nor- tromedial margins, sparsely setose laterally, mal form, endopod with dactylar segment exopod well developed, not reaching to car- narrow, densely spinulate medially; pro- pomeral articulation, with numerous plu- podal segment expanded anteromedially, mose setae distally; coxa not medially pro- anterior margin with feebly setulose setae duced, sparsely setose, with well-developed and spiniform setae, medial margin with lateral plate, acutely produced anterolater- spines; ischiomerus with short, feebly setu- ally, lateral margin with sparsely setulose lose setae laterally; basis medially excavate; setae, with epipod ventrolaterally; with small exopod slender, with numerous plumose se- arthrobranch. tae distally, short, feebly setulose setae me- Thoracic sternites narrow, unarmed. dially and laterally on proximal hale coxa Branchial formula: JOURNAL OF CRUSTACEAN BIOLOGY. VOL. 1 1, NO. 4, 1991

Maxilli- la length, 5.0 times longer than central width; peds Pereiopods ischium subequal to meral length, about 5.0 123 12345 times longer than distal width, tapering Pleurobranchs - - - +++++ proximally; basis short, obliquely articulat- Arthrobranchs - - + ----- ed with ischium, sparsely setose ventrally; Podobranchs ------coxa stout, sparsely setose ventrally, with Mastigobranchs - - - +++-- trisetose setobranch and epipod dorsolater- Epipods +++ ++--- ally. Exopods +++ ----- Ambulatory pereiopods robust; third pe- reiopod exceeding carpocerite by dactyl and First pereiopods small, robust, subequal, 0.2 of propod; dactyl robust, strongly com- similar, reaching to about level of distal end pressed, 3.0 times longer than proximal of carpocerite, chela camed in extended po- depth, distally acute, biunguiculate, with sition (?); chela stout, about 0.7 of carapace stout, blunt accessory tooth at 0.6 of ventral length, palm subcylindrical, slightly com- margin length, about 0.5 of distal tooth pressed, about 1.8 times longer than central length, ventral border mainly convex, blunt, width, slightly swollen centrally, generally corpus with groups of setae distolaterally smooth, with blunt tubercles ventromedi- and medially; propod about 0.38 of cara- ally, with long simple setae; fingers robust, pace length, 3.5 times dactyl length, subuni- dactylus curved, about 0.7 of palm length, form, 7.0 times longer than deep, with pair 4.0 times longer than proximal depth, with of short, stout, feebly hooked distoventral stout hooked tip, cutting edge slightly lat- spines, ventral border with medial and lat- erally situated, with 4 low teeth on proximal eral rows of sparse, short slender spines; car- half, smaller, irregular, more acute teeth pus about 0.6 of propod length, 3.5 times (damaged?) over distal halt fixed finger sim- longer than distal width, with strong dis- ilar, with 4 low teeth over proximal 0.6 of todorsal lobe, unarmed; merus about 1.2 cutting edge, with several small teeth and times propod length, 3.7 times longer than short sharp cutting edge distally, with stout wide, unarmed; ischium about 0.3 of merus acute hooked tip; carpus short, stout, about length, 1.5 times longer than distal width, 0.4 of palm length, smooth, unarmed, deep- tapering proximally, unarmed; basis normal; ly excavate distally; merus subequal to palm coxa stout, with bisetose setobranch, with- length, robust, about 2.5 times longer than out epipod. Fourth and fifth pereiopods distal width, slightly tapered proximally, similar; fifth pereiopod with propod about deeply excavate distoventrally, lateral mar- 0.9 of third propod length, with transverse gin entire, medial margin with large blunt rows of short setae distomedially. tubercles and setae; ischium short, stout, Pleopods without special features; second unarmed, about 0.4 of merus length, ventral pleopod with basipodite 2.0 times longer margin feebly carinate, setose; ischium about than distal width, with pair of short, rudi- 0.5 of merus length, 2.0 times longer than mentary ovigerous setae (?)at 0.3 of medial distal width, tapering proximally, unarmed; margin length. coxa robust, with small setose distoventral Uropod with protopodite robust, with process, epipod, and trisetose setobranch. stout acute distolateral lobe, smaller acute Second pereiopods small, slender, sub- distomedial lobe; exopod broad, far exceed- equal, similar, extending to slightly beyond ing telson, 1.2 times sixth abdominal seg- carpocerite; chela small, about 0.33 of first ment length, 2.0 times longer than wide, pereiopod chela length, palm subcylindri- ovoid, with very well-marked diaeresis at cal, slightly compressed, feebly tapering dis- about 0.6 of length, proximal margin non- tally, 1.6 times longer than deep, fingers spinulate, distal portion of lamella highly about 0.85 of palm length, slender, simple, flexible; lateral margin feebly convex, with with small, acute hooked tips, slightly lat- dense row of short stiff plumose submar- erally situated, entire cutting edges along ginal ventral setae, with small acute distal distal 0.6 of opposing margins, numerous ventral tooth, with single well-developed groups of setae; carpus about 2.0 times chela mobile spine medially, separated by dense length; 5-segmented, segment lengths in ra- tuft of long simple setae projecting dorso- tio4:1:1:1:2.5; merusabout 1.75 timesche- laterally, distolateral and medial margins BRUCE AND ILIFFE. NEW STYGIOPHILIC ALPHEID SHRIMP FROM SAMOA 587

Fig. 2. Hamalpheus acanthops, new genus, new species, holotype female. A, anterior carapace and antennal peduncles, dorsal aspect; B, same, lateral aspect; C, antennule; D, antennular peduncle, medial aspect; E, antennal peduncle, ventral; F, scaphocerite, dorsal; G, eye, dorsal; H, telson; I, uropod. densely fringed with long robust annulate of long simple setae medially, medial spine plumose setae; endopod about 1.1 times ex- small, subconical, intermediate and lateral opod length, 2.7 times longer than wide, spines robust, about 0.12 of endopod length, proximal half uniform, distal half tapering divergent, strongly curved ventrally; medial medially, posterior margin about 0.33 of and lateral margins fringed with long, robust width with 3 spines laterally and dense tuft annulate plumose setae. 588 JOURNAL OF CRUSTACEAN BIOLOGY. VOL. 11. NO. 4, 1991

Fig. 3. Harnalpheus acanthops, new genus, new species, holotype female. A, right mandible, dorsal; B, maxillula, ventral; C, maxilla; D, first maxilliped; E, second maxilliped; F, third maxilliped.

Type.-The single available specimen is the Coloration.-No data. holotype and is deposited in the collection of the Northern Territory Museum, Dar- Habitat. -The Tosua-To~~su~ lava tube win, catalogue number NTM Cr.007421. consists of two large collapsed sinkholes, interconnected by a lava tube, but also di- Measurements (mm).-Total body length rectly connected with the sea. Salt water from (approximate), 27.3; carapace length, 7.7; the ocean extends most of the way back into second pereiopod chela, 4.5. the system. Beginning at the base of 15-m BRUCE AND ILIF'FE NEW STYGIOPHILIC ALPHEID SHRIMP FROM SAM04

Fig. 4. Hamalpheus acanthops, new genus, new species, holotype female. A, first pereiopod, right, lateral aspect: B, same, chela; C, same, fingers; D, second pereiopod, right; E, same, chela and distal carpus; F, third pereiopod; G, same, propod and dactyl; H, fourth pereiopod, propod and dactyl; I, fifth pereiopod, propod and dactyl; J, second pleopod. high coastal sea cliffs, an underwater pas- deep). A further 30-m section of lake ends sage continues inland for 20 m to a large at a breakdown slope ascending to the To- pool situated directly beneath the Tosua en- lesua entrance (15-m diameter, 8-m depth). trance (30 m long by 20 m wide and 26 m Brackish water emerging out of this break- JOURNAL OF CRUSTACEAN BIOLOGY, VOL 11, NO 4,199 1

Fig. 5. EIamalpheus acanthops, new genus, new species, holotype female. A, mandible, molar and incisor processes, palp removed; B, same, palp; C, maxillula, distal palp; D, third maxilliped, distal endopod; E, third pereiopod, distal propod and dactyl; F, uropod, distolateral angle of exopod; G, same, distal endopod; H, telson, posterior margin; I, same, lateral aspect. down suggests the presence of an inacces- of strong currents within the cave. Total sible inland continuation of the cave under length of the cave is 181 m, with a maxi- the collapse. The floor of most of the pool mum depth of 26 m. is coarse sand and rubble with very large The single specimen of Hamalpheus sand ripples indicating the existence at times acanthops was collected by hand from a BRUCE AND ILIFFE: NEW STYGIOPHILIC ALPHEID SHRIMP FROM SAMOA 591 shallow intertidal pool in the rear portion Etymology. -From akantha, Greek, a thorn of a small side gallery of the cave, close to or spine, and ops, Greek, an eye. the sea.

Remarks. -The unusual acute process in the Hamalpheops acanthops is the first alpheid anterior aspect of the eye stalk in H. acan- definitely recorded from a lava tube. How- thops is unique and its functions are not ever, Metabetaeus lohena has also been re- immediately obvious. Other alpheid corded from a pool in a marine cave on shrimps may also have small acute pro- Hawaii that may possibly also be the re- cesses in this situation, among them Alphe- mains of a lava tube (Banner and Banner, us brucei, but in these the processes are com- 1960b). paratively small (Banner and Banner, A key to the then-known genera of the 198 lb). The caudal fan is also unique, with family was provided by Holthuis the lack of dorsal telson spines, of which in his invaluable volume published in 19 55. two pairs are almost invariably present in This included 19 genera, of which 16 were the rest of the Alpheidae, and the dorsally represented in the Indo-West Pacific region. recurved inner posterior marginal spines, Since then, Arete Stimpson has been syn- together with the group of ventrally curved onymized with Athanas Leach (Banner and spines at the tip ofthe exopod ofthe uropod, Banner, 1960a). The genus Metalpheus a position devoid of spines in most other Coutibre has also been resurrected (Chace, alpheid shrimps, although some species of 1988). A further 9 genera have now been Alphew, such as A. bucephalus Coutibre, described which contain representatives in may have numerous small spinules around the Indo-West Pacific fauna, including the the margin of the endopod of the uropod present genus Hamalpheus. There are 25 (Banner and Banner, 1981a). The mor- genera now reported from the Indo-West phology of the caudal fan suggests an ad- Pacific region. Johnson (1961) provided a aptation to life in a rapidly fluctuating water key to eight genera known from the Indo- current, possibly beneath stones, slabs, or Australian region. Subsequently Banner and rocks. Most shrimps actively face into a wa- Banner (1973) published a key to nine gen- ter current and retain their position by their era from Australian waters. Recently, Chace walking legs. Hamalpheus acanthops may (1988) provided a key to the 15 genera use its caudal fan as an anchor, in a reversing known from the Philippine Islands. To fa- current, with the uropod spines attaching to cilitate the identification of all Indo-West the substrate and the inner telson spines Pacific genera, the following key is provid- possibly to the roof of the cavity occupied. ed. The mode of carriage of the chelae of the first pereiopods is uncertain. The ventral merus is deeply excavate and can readily accommodate the flexed carpus and chela, but these appear to be naturally held in an 1. Body very strongly flattened, first pleuron extended position, with the dactyl ventrally, covering much of posterior carapace; eyes in the preserved specimen. The presence of dorsally exposed; first pereiopods very feebly biunguiculate dactyls and robust ambula- developed, smaller than second pereiopods terocaris Heller, 1862 tory pereiopods in alpheid and some other - ed, first pleuron shrimps also generally suggests a commen- covering only posterior branchiostegte; first sal lifestyle, often in association with mod- pereiopods larger than second pereiopods ...... 2 ifications of the caudal fan, all adaptations 2. Fingers of major first pereiopod with sound- to retaining an association with the host an- P nlf 3 imal. No such associations appear to have - Fi been possible in the case of H. acanthops, producing molar process and fossa mecha- where the habitat was generally devoid of nism 6 potential hosts. An alternative explanation 3. Body pace and abdomen with strong median dorsal is that the dactyls and pereiopods may also canna; chelae of first pereiopods strongly be developed as an independent adaptation compressed ...... Racilius Paulson, 1875 to strong currents. - Body not strongly compressed; carapace with 592 JOURNAL OF CRUSTACEAN BIOLOGY, VOL 11. NO. 4, 1991

at most feeble median carina, abdomen dor- extended position; free-living or associated sally noncarinate; chelae of first pereiopods with gastropods ...... Betaeus Dana, 1852 not strongly compressed ...... 4 - First pereiopods markedly unequal, dissimi- 4. Antepenultimate segment of endopod of third lar, camed in flexed position; associated with maxilliped markedly broadened, flattened, thalassinids ...... Leptalpheus Williams, 1965 suboperculate ...... Metalpheus Coutikre, 1908 18. Posterior margin of telson acutely produced - Antepenultimate segment of endopod of third Neoalpheopsis Banner, 1953* maxilliped subcylindrical, not suboperculate f telson not acutely pro- ...... 5 19 ...... 5. Pereio~odswithout e~i~ods- A Synalpheus Bate, 1888 20 - Exopod of uropod lpheus Fabricius, 1798 tolateral tooth and spine only 22

to dactylar hinge; carpus subrectangular ...... cisor process greatly enlarged, without palp ...... Nennalpheus Banner and Banner, 198 1 ...... Prionalpheus Banner and Banner 1960 - Mandible normal, with not subrectangular 7

8

ment with articulated triangular plate ...... 14 ...... Alpheopsis Coutikre, 1896 - At most, first 2 pairs of pereiopods with epi- pods (sometimes rudimentary) ...... 23 23. Eyes fully concealed - 24. orbital teeth; major chela with palm strongly ...... compressed, dorsally and ventrally carinate; Telson without dorsal spines; endopod with mandible without palp; associated with pa- strong terminal spines; eyestalk with conspic- gurid crabs ...... Aretopsis de Man, 19 10 uous acute anterior process - Rostrum long, acute, with periorbital teeth; chelae oval in section, noncarinate; mandible with palp; free-living or associated with echi- noderm Athanas Leach, 18 14 Bannereus Bruce, 1988 ct rostrum and su- * Neoalpheopsis is possibly a junior synonym of Para- nondentate ...... 12 betaeus (see Banner and Banner, 1985). Pereiopods without epipods; exopod of uro- pod without distinct diaeresis Bat 955 s with epipods; ex- opod of uropod with distinct diaeresis ...... This research was conducted as part of a 12-month Vexillipar Chace, 1988 expedition investigating the biology of anchialine caves Rostrum well developed, acute; eyes dorsally in the South Pacific region. Support for this work was provided by grants (TMI) from the United States Na- tional Science Foundation (BSR-8700079) and the Na- 955 tional Geographic Society (#3412-86). Invaluable as- sistance for the study was generously supplied by Mr. Dan Sua and the Western Samoa Department of Fish- eries. Yolanda Iliffe also assisted with cave collections. 888 Dr. Yasuhiro Miya kindly provided much helpful ad- West Pacific s~ecies) vice on the key to alpheid genera. - 15.

- ...... 16 Banner, A. H., and D. M. Banner. 1960a. Contri- 16. Posterior margin of telson acutely produced butions to the knowledge of the alpheid shrimp of the Pacific Ocean. Part V. The Indo-Pacific members of the genus Athanas. -Pacific Science 14: 129-1 55. - ...... 17 -, and -. 1960b. Contributions to the 17. First pereiopods subequal, similar, camed in knowledge of the alpheid shrimp of the Pacific Ocean. BRUCE AND ILIFFE: NEW STYGIOPHILIC ALPHElD SHRIMP FROM SAMOA 593

Part VII. On Metabetam Borradaile, with a new Holthius, L. B. 1955. The recent genera of the caride- species from Hawaii.-Pacific Science 14: 299-303. an and stenopodidean shrimps (Class Crustacea: Or- , and -. 1973. The alpheid shrimp of der : Supersection Natantia) with keys for Australia. Part I. The lower genera.-Records of the their determination. -Zoologische Verhandelingen Australian Museum 28: 29 1-382. 26: 1-157. Banner, D. M., and A. H. Banner. 198 1a. The alpheid -. 1986. Decapoda.-In: L. Botosaneanu, ed. shrimp of Australia. Part 111. The remaining al- Stygiofauna mundi. A faunistic, distributional, and pheids, principally the genus Alpheus, and the family ecological synthesis of the world fauna inhabiting Ogyrididae. -Records of the Australian Museum 34: subterranean waters (including the marine intersti- 1-357. tial). Pp. 589-6 15. E. J. Brill, Leiden, The Nether- , and -. 198 1b. The alpheid shrimp of lands. Pp. 1-740. Australia. Supp. I.-Records of the Australian Mu- Johnson, D. S. 196 1. A synopsis of the Decapoda seum 34: 359-362. and Stenopodidea of Singapore with notes , and -. 1985. The alpheid shrimp of on their distribution and a key to the genera of Ca- Indonesia, based upon J. G. de Man's "The Decap- ridea occumng in Malayan waters.-Bulletin of the oda of the Siboga Expedition, Part 11. Family Al- National Museum, Singapore 30: 44-79. pheidae." (1 9 l l). -Marine Research in Indonesia Yaldwyn, J. C. 1971. Preliminary descriptions of a 25: 1-79. new genus and twelve new species of natant decapod Chace, F. A., Jr. 1988. The caridean shrimps (Crus- Crustacea from New Zealand. -Records of the Do- tacea Decapoda) of the Albatross Philippine Expe- minion Museum 7: 85-94. dition, 1907- 19 10, Part 5: Family Alpheidae. - Smithsonian Contributions to Zoology 466: i-vi, 1- RECEIVED:28 September 1990. 99. ACCEFTED:I5 May 199 1. Dana, J. D. 1852. Conspectus crustaceorum quae in orbis terrarum circumnavigatione, Carolo Wilkes e Addresses: (AJB) Division of Natural Sciences, Classe, Reipublicae Foederatae Duce, lexit et descrip- Northern Temtory Museum, P.O. Box 4646, Darwin, sit.-Proceedings of the Academy of Natural Sci- Australia 080 1; (TMI) Department of Marine Biology, ences, Philadelphia. 1852: 6-28. Texas A&M University Mitchell Campus, P.O. Box Fabricius, J. C. 1798. Supplementum entomologiae 1675, Galveston, Texas 77553. systematicae.-Hafniae, Copenhagen. Pp. 1-572.