JOURNALOF CRUSTACEANBIOLOGY, 9(3): 445-452, 1989

KORORMISTICIUS, NEW GENUS, NEW SPECIES (:HIPPOLYTIDAE), A CAVE SHRIMP FROM PALAU

Janice Clark

ABSTRACT A new genus and species, Koror misticius,a shrimp belongingto a group of 6 monotypic generawithin the Hippolytidaeis describedfrom a marinecave in Palau.It differsfrom Barbouria and Janicea in having arthrobranchson the 4 anterior pairs of pereiopods and epipods on maxilliped 1, from Ligur in differentplacement of the "antennal spine" and in having the propodusof the 3 posteriorpereiopods subdivided, and from Parhippolyteand Somersiellain havinga slenderrostrum and the appendixmasculina distinctly longer than the appendixinterna on the second male pleopod.

Onlytwo speciesof carideanshrimps have of all pereiopods.Mandible bearing 3-joint- been recordedfrom the inland saline ponds ed palp. Pereiopods 1 and 2 chelate; pe- and wells and the anchialine and marine reiopod 2 merus and carpussubdivided; is- caves of Palau: Halocaridinidestrigonoph- chium subdivided in distal one-third, thalma (Fujino and Shokita, 1975) (Hart, although less distinctly than carpus and 1980; Holthuis, 1982) and Caridina typus merus; pereiopods 3, 4, and 5 with propo- Kubo, 1941. The species here described is dus subdivided and merus bearing strong allied to BarbouriaRathbun, 1912, Janicea lateral spines. Endopod of first pleopod of Manning and Hart, 1984, Ligur Sarato, male without appendix interna, that of sec- 1885, ParhippolyteBorradaile, 1899, and ond with slenderappendix masculina longer Somersiella Hart and Manning, 1981. Un- than appendixintera and bearing2 clusters like the previously described shrimps be- of setae. with- longingto these five monotypicgenera TypeSpecies. -Koror misticius,new species. in the Hippolytidae,the new species has no -Named afterthe unique characterthat may be relied upon Etymology. type locality, Koror Palau. to distinguish it from them. However, cer- Island, tain characteristicsthat occur in members Gender.-Masculine. combined of the other genera are uniquely Relationships.--The features shared by the be in this new shrimp. Because it cannot 6 genera are: presence of "antennal" and as assigned to any one of these genera they branchiostegalspines, pigmented eyes, shape are currentlydefined, the new genus Koror and armament of pleura 6, two arthro- is erected to receive it. branchs on maxilliped 3, pleurobranchs above the Hippolytidae pereiopods, epipods on maxil- liped 3 and anterior 4 pereiopods, lack of new Koror, genus incisor process on mandible, 3-jointed Diagnosis. -Carapace smooth, bearing mandibular palp with last segment being "antennal"and branchiostegalspines; ros- long, chelate pereiopods 1 and 2, multiar- trum dentate, short. Eyes well developed ticulatepropodus, carpus, and merus on pe- and pigmented. First 4 abdominal pleura reiopod 2, merus of pereiopods 3-5 undi- broadly rounded and unarmed; fifth and vided and armed with spines, 2 pairs of sixth producedin sharptooth. Telson bear- dorsalspines on telson, and lack of appendix ing 2 pairs of dorsal spines, 3 pairs of mov- intema on first pleopod of male. able spines on posterior margin. Exopods The differencesamong the generaare not- presentonly on maxillipeds. Epipods borne ed in the key. The charactersin the key are on all maxillipeds and first 4 pereiopods. derived from data in: Chace, 1972; Crosnier Pleurobranchs present above all pereio- and Forest, 1973; Hart and Manning, 1981; pods, arthrobranchson third maxillipedand Gordon, 1936; Hobbs et al., 1977; Hobbs first4 pereiopods,and setobranchson coxae III, 1978; Holthuis, 1963; Lemaitre, 1984;

445 446 JOURNAL OF BIOLOGY, VOL. 9, NO. 3, 1989

Fig. 1. Koror misticius, new genus, new species. Lateral view of male paratype.

Manning and Hart, 1984; Monod, 1968; Telson 3.7 times as long as broad, taper- Rathbun, 1912; Senna, 1903; Wear and ing in width posteriorly; dorsal spines of Holthuis, 1977. telson small, anterior pair situated in pos- terior half, posterior pair lying midway be- Koror new misticius, species tween anterior pair and posterior margin. 1-4 Figs. Posterior margin tapering to sharp median Material. -Male holotype (USNM 235027), and 4 male point and armed with 3 pairs of movable paratypes (USNM 235028) collected from South Point posterior spines (mesial pair one-third length 7?18'32"N Cave, Koror Island, Ngermeuangel, Palau, of middle ones, lateral pair one-half length 134?30'05"E, on 18 April 1985 by Dennis Williams and JeffBozanic. One male (USNM 235029) collected of mesial ones). in limestone cavern on Ngargol Island, Palau, at depth Angle on outer margin of outer ramus of of 20 feet (6.1 m) in nearly complete darkness, 9 Jan- uropod acute, bearing single curved artic- uary 1972, by W. A. Starck and G. R. Allen. ulated spine mesial to angle. Description of Holotype.-Rostrum later- Eyes prominent; cornea pigmented, ally compressed, slender, about 5 times slightly broader than stalk. longer than high, extending almost to distal Antennular peduncle with acute styloce- end of basal segment of antennular pedun- rite not reaching distal end of basal segment. cle, bearing 2 or 3 teeth on dorsal margin Second segment equal in length to third. and 1 or 2 on ventral margin; 1 tooth dis- Antennal scale 4.5 times as long as wide; tinctly postorbital. Antennal angle pro- outer margin straight throughout most of duced anteriorly beyond tip of antennal length, distal tooth overreaching narrowly spine; antennal spine placed behind anterior rounded distal margin of blade, scale over- margin of carapace; branchiostegal spine reaching antennular peduncle by about half placed behind margin, its tip surpassing its length. margin; both antennal and branchiostegal Mandible without incisor process, bear- spines continuing posteriorly in short hor- ing slender 3-segmented palp, distal seg- izontal carina; pterygostomian angle round- ment nearly as long as combined length of ed, not produced. proximal 2 segments. First maxilla with dis- CLARK: NEW HIPPOLYTID SHRIMP FROM PALAU 447

C

F 3 4

Fig.3, n g n , 4 6 I - _ Fig. 2. Koror misticius, new genus, new species, holotypic male. A, cephalic region in lateral aspect; B, same, in dorsal aspect; C, lateral view of branchial region showing pleurobranchs, arthrobranchs, mastigobranchs, and epipods; D, telson and uropods; E, detail of terminal part of telson; F, detail of distal part of lateral ramus of uropod; G, antennule in dorsal aspect; H, antenna in ventral aspect; I, pleura 3-6. tal lacinia spatulate, supporting several rows tal one-third, bearing terminal and subter- of short spines on mesial face; proximal la- minal setae. Second maxilla with scaphog- cinia digitiform, bearing many terminal and nathite broadly rounded proximally, not as subterminal setae; palp slightly rolled in dis- broad distally; palp with single subterminal 448 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 9, NO. 3, 1989

Fig. 3. Koror misticius, new genus, new species, holotypic male. A, third pereiopod; B, same, dactyl; C, fourth pereiopod; D, same, dactyl left member; E, fifth pereiopod; F, same, dactyl; G, second pereiopod; H, same, propodus and dactyl; I, first pereiopod; J, same, propodus and dactyl. CLARK: NEW HIPPOLYTID SHRIMP FROM PALAU 449

A

4 ^^/^^ ~s G G A

Fig.4. Korormisticius, new genus,new species,holotypic male. A, mandible;B, firstmaxilla; C, secondmaxilla; D, first maxilliped; E, second maxilliped; F, third maxilliped; G, same, terminal spines; H, first pleopod; I, same, distal part of endopod; J, second pleopod; K, same, distal part of appendix intema; L, same, distal part of appendix masculina.

spine and several mesial setae; distal, mid- dopod. Third maxilliped overreaching dle, and proximal endites unremarkable. antennal scale, exopod reaching slightly be- First maxilliped with exopod bearing ca- yond midlength of ischium; with 1 epipod ridean lobe on lateral face; epipod bilobed; and 2 arthrobranchs, one small and ob- palp with 2 articles, bearing single subter- scured by larger, more ventrally situated one; minal spine and several setae on mesial face; terminal segment bearing transverse rows proximal endites subtruncate, distal one of weak spines and 1 terminal and 5 sub- broadly rounded. Second maxilliped bear- terminal heavy corneous denticles. ing single epipod, podobranch, and well de- Branchiae and endites as illustrated and veloped exopod reaching much beyond en- as follows (P = present): 450 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 9, NO. 3, 1989

Maxillipeds Pereiopods 1 2 3 1 2 3 4 5 Pleurobranchs 1 1 1 1 1 Arthrobranchs _ -- 2 1 1 1 1 Podobranchs - 1 - Setobranchs P P P P Epipods 1 1 1 1 1 1 - Exopods 1 1 1

Pereiopods as illustrated. Pereiopods 1 onymy of Ligur, assigned Barbouria anti- and 2 chelate, fingers of both shorter than gensis to the new genus Janicea, and as- palms. Pereiopod 2 longer than 1; propo- signeda fifth species to the monotypic genus dus, carpus,and merus subdivided,ischium Somersiella. Koror misticius belongs to a with several rudimentary subdivisions in sixth monotypic genus. Although it has no distal one-third. Pereiopods 3-5 with 3 charactersthat are unique, it does not have spines on posterior surfaceof dactyl, prop- charactersthat allow it to be placed into any odus subdivided, carpus and merus undi- of the existing genera. vided, merus armed with conspicuous Janicea, Barbouria, Ligur, and Somer- spines. siella have unique charactersthat separate Endopod of first pleopod of male half them. Janicea and Barbouriaare the only length of exopod; with cluster of coupling generain this group that lack arthrobranchs hooks at distal end; without appendix in- on the four anteriorpairs of pereiopodsand tera. Endopod of second pleopod of male epipods on the first maxilliped. with thin appendix masculina longer than Ligur is the only genus with a long ros- appendix intera, armed with distal and trum, distinctly overreachingthe antennu- mesial clustersof spines;longest spine about lar peduncle. It is also the only genus with one-half length of appendix masculina. the "antennal spine" conventionally situ- -Rostrum with 1 or 2 ventral ated ventral to the indistinct orbital angle. Variability. In spines.One male with 2 dorsalrostral spines. Barbouria,Janicea, Koror,Parhippolyte, and Somersiellathis "antennal is un- Length of rostrum varying from barely spine" situated to the dis- reaching end of basal segment of antennal usually posterodorsad tinct orbital Chace com- peduncle to slightly overreaching it. Sub- edge. (personal believes that "the 'antennal division of ischium of pereiopod 2 varying munication) in is not from distinct to vague. spine' Ligur probably homologous with the nearly postorbitalspines that have Size. -Male holotype, postorbitalcarapace been masqueradingunder that name in the mm mm length 8.5 (10.1 including ros- other 5 genera."He "suspectsthis character trum). Illustratedmale paratype,postorbit- may be more importantto an understanding mm in- al carapace length 7.8 (10.0 mm of the relationships of the 6 'genera' in- cluding rostrum).Size range of other males volved here than either arthrobranchsor postorbitalcarapace length 7.4-9 mm (10.1- subdivided propodi of the posteriorpereio- 12.1 mm including rostrum). Females un- pods." known. Somersiella is the only genus with ar- Distribution. -Palau. throbranchs instead of podobranchs on maxilliped 2. -From Latin, misticius, of Etymology. Parhippolyte and Koror can be distin- mixed race, hybrid, referringto the com- charactersthat are not bination of charactersshared with the re- guished by unique to them, but are differentin the two genera. latedgenera and the lackof any singleunique In Kororthe rostrum is about five character. slender, times longer than high, whereas in Parhip- Remarks.-The relationshipsof the genera polyte it is much deeper,about two and one- Barbouria and Ligur were reevaluated by half times longer than high. In Koror the Manning and Hart in 1984 at which time appendixmasculina is distinctlylonger than they removed Parhippolytefrom the syn- the appendix interna on the second male CLARK:NEW HIPPOLYTIDSHRIMP FROM PALAU 451

pleopod, whereas in Parhippolyteit is sub- Islandsand elsewhere, collected during the years1895, equal in length to the appendixintera. The 1896, and 1897 4: 395-428. of Chace, F. A., Jr. 1972. The shrimps of the Smith- pleuron the fourth abdominal somite is sonian Bredin CaribbeanExpeditions with a sum- roundedposteroventrally in Koror,whereas mary of the West Indian shallow-water species in Parhippolyteit is acute posteroventrally. (Crustacea: Decapoda: Natantia).-Smithsonian From the distributionalviewpoint Kororis Contributionsto Zoology 98: 1-179. included in the same as Crosnier,A., and J. Forest. 1973. Les crevettespro- geographicalregion fondes de l'Atlantique oriental tropical.-Faune Parhippolyte,which is the only other genus Tropicale(O.R.S.T.O.M.) 19: 1-409. of the group known from the Pacific. Fujino, T., and S. Shokita. 1975. Report on some new atyid shrimp (Crustacea,Decapoda, ) KEY TO SIX RELATEDMONOTYPIC from the Ryukyu Islands.-Bulletin of Science & GENERA WITHINTHE HIPPOLYTIDAE EngineeringDivision, University of the Ryukyus (Mathematics& Natural Sciences) 18: 93-113. 1. Pereiopodswithout arthrobranchs 2 Gordon, I. 1936. On the hippolytid prawns of the - Four anteriorpairs of pereiopodswith arthro- genus Ligur Sarato.-Proceedings of the Zoological branchs 3 Society of London 1935-1936: 102-108. 2. Corneanarrower than eyestalk;3 posteriorpairs Hart,C. W., Jr. 1980. A new atyid shrimp,Palauatya of pereiopodswith propodusentire, not subdi- dasyomma,from Palau, Caroline Islands. -Proceed- vided; appendixmasculina shorter than appen- ingsof the BiologicalSociety of Washington93:481- dix internaon second male pleopod...... Barbouria 489. - Corneabroader than eyestalk;3 posteriorpairs ,and R. B. Manning. 1981. The cavericolous of pereiopods with propodus subdivided; ap- carideanshrimps of (Alpheidae, Hippol- pendix masculinalonger than appendixinterna ytidae, and Atyidae).-Journal of CrustaceanBiol- on second male pleopod Janicea ogy 1: 441-456. 3. Rostrumdistinctly overreaching antennular pe- Hobbs, H. H., III. 1978. The female of Barbouria duncle; antennal spine conventionally situated cubensis (von Martens) (Decapoda, Hippolytidae) ventral to indistinct orbital angle; 3 posterior with notes on a populationin the Bahamas.- Crus- pairs of pereiopods with propodus entire, not taceana35: 99-102. subdivided Ligur Hobbs, H. H., Jr., H. H. Hobbs, III, and M. A. Daniel. - Rostrumreaching little if at all beyond level of 1977. A review of the troglobiticdecapod crusta- distal margin of basal antennularsegment; an- ceans of the Americas.- SmithsonianContributions tennal spine situated posterodorsadto distinct to Zoology 244: 1-176. orbitalangle; 3 posteriorpairs of pereiopodswith Holthuis, L. B. 1963. On red colouredshrimps (De- propodussubdivided 4 capoda,Caridea) from tropicalland-locked saltwater 4. Pleuronof fourthabdominal somite acute pos- pools.-Zoologische Mededelingen38: 261-279. teroventrally Parhippolyte 1982. Notes on Indo-WestPacific Crustacea - Pleuron of fourth abdominal somite rounded Decapoda I and II.-Crustaceana 42: 26-36. posteroventrally 5 Kubo, I. 1941. On some fresh-watershrimps from 5. Rostrumdeep, 2.5 times as long as high, with 4 the Ryukyu Islands.-Transactions of the Biogeo- or 5 ventralteeth; appendix masculina subequal graphicalSociety of Japan 3: 303-318. in length to appendix intera on second male Lemaitre,R. 1984. Decapod crustaceansfrom Cay pleopod Somersiella Sal Bank, Bahamas, with notes on their zoogeo- - Rostrumshallow, 5 times as long as high, with graphicaffinities.-Journal of CrustaceanBiology 4: 1 or 2 ventral teeth; appendix masculina dis- 425-447. tinctly longer than appendixintera on second Manning,R. B., and C. W. Hart,Jr. 1984. The status male pleopod Koror of the hippolytidshrimp genera Barbouria and Ligur (Crustacea:Decapoda): a reevaluation.- Proceed- ACKNOWLEDGEMENTS ingsof the BiologicalSociety of Washington97:655- 665. I am gratefulto C. W. Hart, Jr., Departmentof In- Monod, Th. 1968. Nouvelle capturede Ligur uveae vertebrateZoology, National Museumof NaturalHis- (Borradaile)aus Iles Loyalty (Crustacea,Decapo- tory, for advice. My thanks go also to Dr. Brian F. da).-Bulletin du Museum National d'Histoire Na- Kensley, Dr. Horton H. Hobbs, and Dr. Fenner A. turelle,Paris (2)40: 772-778. Chace, Jr. for their constructivecomments and sug- Rathbun,M. J. 1912. Some Cuban Crustacea,with gestions on the manuscript.I am also gratefulto Mr. notes on the Astacidae,by WalterFaxon, and a list Dennis Williams, Mr. Jeff Bozanic, Mr. W. A. Starck, of the Isopoda,by HarrietRichardson. -Bulletin of and Mr. G. R. Allen who collected the shrimp and the Museum of ComparativeZoology 54: 449-460. made them available for study. Sarato,C. 1885. Etude sur les Crustacesde Nice.- Le Moniteurdes Etrangersa Nice 222: 2. LITERATURECITED Senna, A. 1903. Nota sui Crostacei Decapodi. Le esplorazioniabissali nel Mediterraneodel R. Piros- Borradaile,L. A. 1899. On the Stomatopoda and cafo Washingtonnel 1881, II.-Bolletino della So- Macrura brought by Dr. Willey from the South cieta EntomologicaItaliana 34: 235-367. Seas.-In: A. Willey, Zoological results based on Wear,R. G., and L. B. Holthuis. 1977. A new record material from New Britain, New Guinea, Loyalty for the anchialine shrimp Ligur uveae (Borradaile, 452 JOURNALOF CRUSTACEANBIOLOGY, VOL. 9, NO. 3, 1989

1899) (Decapoda, Hippolytidae) in the Philippines Address: Department of Invertebrate Zoology, Na- with notes on its morphology, behavior and ecolo- tional Museum of Natural History, Smithsonian In- gy.-Zoologische Mededelingen 51: 125-140. stitution, Washington, D.C. 20560. RECEIVED:26 April 1988. ACCEPTED:20 March 1989.

ANNOUNCEMENT

Under the auspices of the Commission on Systematic and EvolutionaryBiology of the InternationalUnion of BiologicalSciences (IUBS), The InternationalCongress of System- atic and EvolutionaryBiology IV (ICSEB-IV)will be held at the University of Maryland, College Park, Maryland, 1-7 July 1990. The meeting is cosponsored by the Smithsonian Institution and the University of Maryland. The overall theme for the meeting is "The Unity of EvolutionaryBiology." The program will featurea number of major symposia aimed at illuminatingsome of the most exciting new areas of evolutionary biology as well as providing a synthesis of historical and mechanistic approachesto evolution in , plants, and microorganisms.Congres- sional symposia will focus on three major ideas: Evolutionin perspective:biodiversity, conservation, biotechnology, and globalchange; Tempo and pattern of evolution: micro- and macroevolutionaryprocesses; and Systematicsand phylogenetic reconstruction. The goal of the Congress will be to foster a resynthesis of the theory of evolution, incorporatingnew and traditionalapproaches. Keynote speakerswill include Douglas Futuyma, Stephen J. Gould, Richard Leakey, Robert May, Peter Raven, John MaynardSmith, Geerat Vermeij,and Edward0. Wilson. Individuals may present contributed papers in either oral or poster format, although posters are stronglyencouraged. For furtherinformation on travel assistance contact Dr. ArthurPopper, Department of Zoology, University of Maryland,College Park, Maryland 20742. For furtherinformation, write: CongressSecretary, ICSEB-IV Department of Microbiology University of Maryland College Park, Maryland20742