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ISSN 0968-0470 Bulletin of The Natural History ~ Museum PRESENTED fiENERAL LIBRARY | Zoology Series VOLUME 62 NUMBER 2 28 NOVEMBER 1996 The Bulletin ofTheNaturalHistory Museum (formerly: Bulletin ofthe British Museum (NaturalHistory)), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Zoology Series is edited in the Museum's Department ofZoology Keeper ofZoology DrC.R. Curds EditorofBulletin: DrN.R. Merrett Assistant Editor: DrB.T. Clarke Papers in the Bulletin are primarily the results ofresearch carried out on the unique and ever- growing collections ofthe Museum, both by the scientific staffand by specialists from elsewhere who make use ofthe Museum's resources. Many ofthe papers are works ofreference that will remain indispensable foryears to come. All papers submitted for publication are subjected to external peer review for acceptance. A volume contains about 160 pages, made up by two numbers, published in the Spring andAutumn. Subscriptions may be placed for one or more ofthe series on an annual basis. Individual numbers and back numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and enquiries should be sent to: Intercept Ltd. P.O. Box 716 Andover Hampshire SP10 1YG Telephone: (01264) 334748 Fax: (01264) 334058 Claims fornon-receipt ofissues ofthe Bulletin will be met free ofcharge ifreceivedby the Publisher within 6 months forthe UK, and 9 months for the rest ofthe world. WorldListabbreviation: Bull. nat. Hist. Mus. Lond. (Zool.) ©The Natural History Museum, 1996 Zoology Series ISSN 0968-0470 Vol. 62, No. 2, pp. 71-132 The Natural History Museum Cromwell Road London SW7 5BD Issued 28 November 1996 TypesetbyAnnBuchan(Typesetters),Middlesex PrintedinGreatBritainbyHenryLingLtd.,attheDorsetPress,Dorchester,Dorset Bull. nar. Hist. Mus. Lond. (Zool.)62(2): 71-82 Issued28November 1996 Indian Ocean echinoderms collected during the Sindbad Voyage (1980-81): 3. Ophiuroidea and Echinoidea ANDREW R.G. PRICE EcosystemsAnalysisandManagementGroup, DepartmentofBiologicalSciences, UniversityofWarwick, Coventry CV4 7AL, UK THE NATURAL HISTORY MUSEUM FRANCIS W.E. ROWE GoldbrookBoarding Kennels, Nuttery Vale, CrossStreet, Home, Suffolk IP21 5BB, UK 11 L>l.u1996 PRESENTED GENERAL LIBRARY SYNOPSIS. At least 44 ophiuroid and 11 echinoid species are recorded from echinoderm collections made during an internationalexpedition,theSindbadVoyage, fromOmantoChina. Samplinglocalitiesincludethe littleknown Lakshadweep (Laccadive), Islands and Pula We (Sumatra) from which 71% ofthe species were recorded. Following the zoogeographic subdivisionsofClark&Rowe(1971),rangeextensionsarerecordedfortenoftheophiuroids:W.India(Amphioplus(Lymanella) sp.);SriLanka{Ophiactismodesta, Ophiarachnarobillardi, Ophiodyscritainstratus);Maldivesarea{Cryptopeltagranulifera, Ophiochaetahirsuta);andIndonesia/EastIndies(Amphiura(Amphiura)dejectoides,Amphiura(Amphiura)micro,Amphioplus {Amphioplus) sienaspis, Ophiogymna pellicula). In addition to the taxonomic treatment, ecological information for each echinodermspecies(habitattypes,depthrange)isprovidedandbroadly analysed. INTRODUCTION MATERIALSAND METHODS The systematica and distribution of Indian Ocean ophiuroids and Specimenswerecollectedbyoneofus(A.R.G.P.)andotherexpedi- echinoidsaretreatedindetail byClark& Rowe(1971). Regions for tion members from localities at Muscat, Oman; Chetlat, Laks- which limited information is available include the Lakshadweep hadweeps (Laccadives); SW India; SW Sri Lanka; and Pula We, (Laccadive) Islands, Sumatra and other parts of SE Asia. Recent Sumatra. Details ofthe sampling localities are shown in Figure I. studies including the systematica and zoogeography ofophiuroids Samplingwasundertaken principallyoncoralreefsusingscuba.At and echinoids for the Lakshadweeps include Nagabhushanam & each locality detailsofhabitattype and depth range were recorded, Rao(1972)andJames(1989),the latteryielding many newspecies alongwiththenumberofindividualsofeachspecies.Thenumberof records for both echinoderm classes, which fill in gaps in the specimens collected is placed in parenthesis after each station distributionrecordsofClark& Rowe(1971). Recent workhasalso number in the Material lists foreach species. been undertaken in SEAsia, including Indonesia (Aziz, 1981) and Material was fixed and preserved using standard methods (Lin- the westcoastofThailand (Bussarawit & Rowe, 1985; Bussarawit, coln & Shields, 1979). Although several specimens (inadvertently includedwiththeasteroidcollections)hadbeenidentifiedearlierby in prep.). L.M. Marsh andafewpreliminary identifications were made inthe This paper reports on collections of ophiuroids and echinoids field by A.R.G.P., specimens were mostly identified by and all ftrraonmsditshceispelinaarreyasvoaynadgeotahcerrossloctahleitIinesdiadnurOicngeananfrinotmerOnamtaionnalt,o snpoemciicesccoomnmfeinrtmse.d bTyheF.Wo.pEh.iRu.rowidhoanisdaelcsohirneosipdoncsoilblleectfioornstaxaor-e China.Theexpedition,SindbadVoyage,wasundertakenin 1980-81 deposited at the Natural History Museum, London, where the aboard a replica of an ancient Arab sailing vessel, 'Sohar'. In holothurian collection (Price & Reid, 1985) and a representative additiontoasystematicaccount,thezoogeographic significanceof collection of the asteroids (Marsh & Price, 1991) have also been the results and the ecology of each species are broadly assessed. lodged. Details of the holothurian collections (Price & Reid, 1985) and & asteroidcollections(Marsh& Price, 1991)resultingfromtheexpe- sysFtoelmlaotwiicnrgefreerceennctespraarcetkiecpet(teo.ga.mSilnoainm,uCmlabrykcitinTgaymlaojro,r1w9o7r9k)s, dairteiaolnshoabveeianlgrepardeypabreeedn(pMuabrlsihsahleld.&DePtraiicles,oinftphreepc.r),inaoniddsacodlelteacitleedd w1h97e6n;pCohsseirbbloenn(ei.egr.C&larGkui&lleR,ow1e9,781;97R1o;wCelar&k&GatCeosu,rt1m9a9n5)Stforcokm, analysis of the ecology and biogeography of all five echinoderm which the original species descriptions, recent authoritative diag- classes will follow. noses and taxonomic decisions can be traced. In some instances, further references are given (e.g. Sloan et al„ 1979) to provide Addressforcorrespondence:Firstauthor additional systematic orbiological information. ©TheNaturalHistoryMuseum, 1996 72 A.R.G.PRICEANDF.W.E. ROWE O 50 E 60 70 80 90 100 Udjung Ug Bau Lo Me Ug TaPa Ga d a j Fig. 1. (a)MapofnorthernIndianOceanshowingsamplingareas(•)duringSindbadVoyage,withinsert(b)forPulaWeSumatra. INDIAN OCEAN ECHINODERMS 73 RESULTS 4. Amphiura (Amphichilus) ochroleuca (Brock, 1888) See. Clark&Rowe, 1971 78; 100; Rowe&Gates, 1995: 344. : Class Ophiuroidea Material. 810502C/1 (1), 810427B/3 (1). Collection sites. SE Klah, SE Lho Pria Laot, Pula We, Sum- atra. GORGONOCEPHALIDAE Family Habitatand depth. In sponge andon subtidal rock; 0-10m. Remarks. This species is known from the Indo-Malayan region 1. Astroboa nuda (Lyman, 1874) and more orless circumscribestheAustraliancontinental coastline (Rowe & Gates, 1995). Pula We appears to be the most westerly See. Clark&Rowe, 1971:78;92;Clark&CourtmanStock, 1976: pointofitsdistribution known todate. 108; 130; Baker, 1980: 60; Rowe & Gates, 1995: 364. 5. Amphioplus (Amphioplus) stenapsis H.L. Clark, 1938 Material. 810504D/3 (1), 810504D/4 (1). Collection sites. NW Klah, PulaWe, Sumatra. SEE. Clark & Rowe, 1971: 78; 101; Rowe & Gates, 1995: 344. Habitatand depth. Subtidal rock/coral; 13 m. Material. 8I0422B/4 (1). Collection sites. Nr. Klah/Seukundo, PulaWe, Sumatra(disc only). Family AMPHIURIDAE Habitatand depth. Coral reef, 2-8 m. Remarks. Although the single specimen comprises only a com- 2. Amphiura (Amphiura) dejectoides H.L. Clark, 1939 plete disc with the bases of2 arms (6 & 9 segments respectively), thereislittledoubtofitsidentity.Thisrecordextendstherangeofthe See. Clark & Rowe, 1971: 80; 97; Cherbonnier & Guille, 1978: speciesto PulaWe from its type locality, Darwin, NAustralia. The 33. record ofA. stenapsis from Madagascar by Cherbonnier & Guille Material. 810501E/8 (4); 7810428D/5 (2). (th1e97v8e)ryissamlamlolstsiczeertoafinltyhebarsadeidalonshaiemlidssidoefnttihfeiicratsipoenc,ijmuedngsi.ngThbey Collection sites. Ug Tapa Gadja & Ug Seukundo, Pula We, confirmation ofthis species in the western Indian Ocean therefore Sumatra. requires confirmation. Habitatanddepth. Subtidal rock/coral,coralreef; 1 & 15 m. 6. Amphioplus (Lymanella) sp. Remarks. The specimens from stn 8I0501E/8 appear to accord well with the original description (Clark, 1939) ofthe species and Material. 810109A/lb (2). descriptions in Clark & Rowe (1971) and Cherbonnier & Guille Collection sites. Beypore, India (west coast). (1978). The 2 specimens from stn 810428D/5 differ in theirfirmer disc, with coarser ventral scaling and in having 6 arm spines Habitat and depth. Subtidal mud, 9 m. proximallyinsteadoffineventralscalingand5proximalarmspines. Remarks. Only the mouthparts and bases ofthe arms are avail- TClhaeryk&areRoiwdeen,ti1f9ie7d1)assuAs.pedcetjeedcAtmoipdheisurwaitihnhraecseenrvsaitsiBona.liCnlsakrykma(iyn Aabmlpehitoopilduesnt(ifLyymtahnieslltaa)x.onT,hwehidocrhsaclleaarrlmyprleaptreesseanrtestrailsopbeecd,iessugo-f bfeircmoendsbpyecCihfeicrbwointnhieAr.&deGjueicltloeid(e1s97H8)..L.ThCelarrekc,oradscionncclluudseidonhecroeni-n gleaesvtiisng(Leyitmhaenr,sp1e8c7i4e)sinA.the(L.k)eyanpdrroevaideed(LbiyitkCelna,rk1&872R)oowreA(.19(7L1.:) extend the distribution ofthis species eastwards across the Indian 102). Cherbonnier & Guille (1978) indicate that laevis has a wide Ocean from the Red Sea, East Africaand Madagascarto the Indo- rangeintheIndianOceanandIndo-Malayregion, whereasClark& Malayan region. Inthe latterregion itmay prove tobe widespread. Rowe (1971) record andreae only from the Malay region. Without complete specimens it is not possible to determine the species nor indeed whetherandreae and laevis are taxonomically separable. 3. Amphiura {Amphiura) micra H.L. Clark, 1938 SEE. Clark & Rowe, 1971: 80; 97; Cherbonnier & Guille, 1978: Family OPHIACTIDAE 46; Rowe & Gates, 1995: 350. Material. 810423B/2 (1). 7. Ophiactis modesta Brock, 1888 Collection sites. Ug Bau, PulaWe, Sumatra. Habitatand depth. Coral, coral reef; 10-30m. See. Clark&Rowe, 1971: 105; Rowe & Gates, 1995: 379. Remarks. This species is recorded across the tropical coast of Material. 810206A/8 (3). AustraliaandfromMadagascar. ItsdiscoveryatPulaWesuggestsa Collection sites. Negombo, Sri Lanka. wider distribution in the Indo-Malayan region for this very small species. Habitatand depth. Coral/rock; 5 m. 74 A.R.G. PRICEANDF.W.E. ROWE Remarks. The 3 specimens were collected in a batch of6 from 11. Ophiothrix exigua Lyman, 1874 stn 810206A/8 with 3 specimens of O. savignyi. They run unequivocably to the species O. modesta in the key provided by See. Clark&Rowe, 1971: 84; 110; Cherbonnier& Guille, 1978: Clark&Rowe(1971: 105)wheretherelationshipwithotherconge- 140; Rowe &Gates, 1995: 422. nersisdiscussedbyA.M. Clark(notes 25-27, pp. 103-105). Rowe (in Rowe &Gates, 1995) preferred to follow the views ofClark& Material. 810206A/8 (5), 810502C/1 (2), 810502E/2 (3). Rowe (1971) regarding recognising the validity of the species O. COLLECTION sites. Negombo, Sri Lanka; E Klah, Pula We, modesta, a course also followed herein, rather than accept the Sumatra. sweeping synonymy of O. savignyi, which includes O. modesta, & Habitatanddepth. Coral/rock,spongeoncoralreefandsubtidal givenby Cherbonnier Guille (1978). rock; 5-10m. 8. Ophiactis savignyi (Miiller & Troschel, 1842) Remarks. One specimen from stn 810206A/8, both from 810502C/1 andtwo from 810502E/3 are veryjuvenile specimens. See. Clark & Rowe, 1971: 82; 103; Clark & Courtman Stock, & 1976: 164; Cherbonnier & Guille, 1978: 125; Sloan et ai, 1979: 12. Ophiothrix savignyi (Miiller Troschel, 1842) 102; Price, 1983: 61; Rowe &Gates, 1995: 380. See. Clark& Rowe, 1971: 84; 109; Cherbonnier& Guille, 1978: Material. 810206A/3 (5), 810206A/8 (3), 810426A/2 (1), 142; Price, 1983:65. 810501E/1 (1), 810501G/3 (1), 8105011/2 (1), 810502F/4 (2). W Material. 801114B/1 (1). Collection sites. Negombo, Sri Lanka; Rubiah, Ug Seukundo, E Klah, PulaWe, Sumatra. Collection sites. Muscat, Oman (1 specimen). Habitatanddepth. Sponge,coral/rock,subtidalrock,coralreef; Habitatand depth. Coral reef; 2 m. 5-14 m. 13. Ophiothrix trilineata Lutken, 1869 Remarks. The specimens identified here are fissiparous and 6- armedwithtrilobeddorsal armplatesandproximally6arm spines, SEE. Clark & Rowe, 1971: 84; 111; Clark & Courtman Stock, according well with the key characters given in Clark & Rowe 1976: 145; Sloan etal, 1978: 103; Rowe &Gates, 1995: 423. (1971). Material. 810420A/5 (2), 810422B/3 (3), 810426A/2 (1), 810428A/9 (2), 810428C/7 (1), 810428D/3a&b(6&2), 810430A/8 (1), 810430A/18 (1), 810430A/21c (1), 810501D/2(1), 810501E/2 Family OPHIOTRICHIDAE (831)0,580120F5/021E(1/)5,(811),0580110F5/031E(/1)8,(811),08501015G0/11E/(29),(28),108510015G0/13E/l(1I&(1)1, juvenile), 810501G/5 (3), 810502C/2(1 veryjuvenile), 810502D/4 9. Gymnolophus obscura (Ljungman, 1867) (4), 810502E/3 (4juvenile), 810502F/2 (2juvenile), 810502G/3 (2 juvenile). See. Clark& Rowe, 1971:82; 117; Rowe & Gates, 1995:411. COLLECTION sites. Klah /E Klah, Nr. Seukundo, Ug Seukundo, Material. 810124A/11 (1), 810125A/2 (1), 810430A/20d (2). Pula We, Sumatra, W Rubiah, Ug Bau, Ug Tapa Gadja, Pula We, Collection sites. Ala Gala & DeumbaGala, Galle, Sri Lanka; Sumatra. Ug Seukundo, PulaWe, Sumatra. Habitat and depth. Coral reef, soft coral / gorgonian, subtidal Habitatanddepth. Coralreefandepizoiconcrinoidsonsubtidal rock, sponge/subtidal rock, sponge; 2-30 m. rock; 8-15 m. Remarks. The majority of specimens exhibit the characteristic Remarks. This species is commonly epizoic on comasterid armcolourpatternof5 longitudinallinesalternatingwhiteanddark crinoids, thehost speciesforwhichhavenotbeenidentifiedforthe blue. However, afewspecimensbearawidemedianpalelongitudi- specimens recorded herein. nal line along the arms, and at least one specimen (810501E/2) is distinctivelypatternedwithcreamblotches,thelinearpatternbeing discernableonlyneartheendsofthearms(seeClark&Rowe, 1971: 10. Ophiotheladanae Verrill, 1869 111). Nextto O. (A.)purpurea, this is the mostcommon ophiuroid species collected. See. Clark & Rowe, 1971: 84; 116; Clark & Courtman Stock, 1976: 141; Price, 1983: 63; Rowe& Gates, 1995:419. 14. Ophiothrix (Acanthophiothrix) armata Koehler, Material. 810125B/1 (3), 810206A/3/78 (4), 810425F/8 (10+), 1905 810428A/2(7),810428A/14(3),810501A/3(30+),810501A/4(l), 810428B/1 (2), 810428D/3c (3), 810501E/9 (15+). See. Clark& Rowe, 1971: 84; 111; Rowe& Gates, 1995: 423. Collection sites. Ala Galla, Galle & Negombo, Sri Lanka; N. Material. 810422B/4 (15+), 810502F/6 (2). Udjung LoMe (NE Sabang Bay), Ug Bau, Ug Seukundo, UgTapa Gadja, PulaWe, Sumatra Collection sites. Klah/Nr. Seukundo, PulaWe, Sumatra. Habitatanddepth. Coral reef, 2-8 m. Habitatanddepth. Epizoiconmacroalgae,gorgonian,firecoral (Miileporasp.)spongeandonholothurians(Thelenotaananas),all Remarks. This species is recorded from the Indo-Malay region, on rock/coral; 2-30m. tropical Australian coasts and the South Pacific (Clark & Rowe, 1 INDIAN OCEAN ECHINODERMS 75 1971).The presentrecord isthe mostwesterly sofarknownforthe Habitatand depth. Coral rubble, coral reef; 8 m & 20-30 m. species. Remarks. A.M.Clark(1968)transferredthisspeciestothegenus MacrophiothrixreferringOphiothrix(Amphiophiothrix)H.L.Clark, 15. Ophiothrix (Acanthophiothrix)purpurea von Martens, 1867 1946,ofwhichdemessaistype(andonly)speciestothesynonymy ofMacrophiothrixH.L. Clark, 1938. SEE. Clark& Rowe, 1971: 86; 1 12; Cherbonnier&Guille, 1978: 148; Sloan etal., 1978: 103; Rowe &Gates, 1995: 423. 19. Macrophiothrix elongata (H.L. Clark, 1938) Material. 810204A/4 (3), 810421A/2 (1), 810421A/9 (1 juve- See. Clark, 1968:291;Clark&Rowe, 1971:82; 114;Price, 1983: nile), 810421B/1 (1), 810422D/3 (2), 810423A/4 (1), 810423B/2 61; Hoggett, 1992: 125. (1), 810425D/2 (1), 80425D/4b(1), 810425F/7 (7), 810425F/8 (1), 810426A/2 (3), 810427A/2 (2), 810428A/2 (2), 810428A/7 (3), Material. 801111A/5 (1), 801114A/5 (2), 801114B/2 (1). 810428A/8 (4), 810428A/9 (2), 810428D/3f(23), 810430A/3 (1), Collection sites. Muscat harbour, Oman. 810430A/21c (6), 810430A/22b (1), 810430A/26 (5), 810501A/ (1), 810501A/2 (1 juvenile), 810501A/4 (1 juvenile), 810501A/6 Habitat and depth. Coral reef, subtidal rock/coral/sand; 0.5-3 (9), 810501E/7 (2), 810501E/9 (1), 810501E/14 (1), 810501E/15 m. (1), 810504B/2(1). 20. Macrophiothrix longipeda (Lamarck, 1816) Collectionsites. SWKalpitiya,SriLanka;UgBau,Rubiah,Ug Seukundo, UgTapaGadja,NKlah,?N. UdjungLoMe,NESabang SEE. Clark, 1968: 300; Clark & Rowe, 1971: 82; 114; Clark & Bay, PulaWe, Sumatra. Courtman Stock, 1976: 139; Hoggett, 1991: 1103; Hogett, 1992: Habitat and depth. Coral reef, soft coral, fire coral (Millepora 151; Rowe & Gates, 1995:413. sspp.o)n,gseu)b;ti2d-al30romc.k/coral(epizoiconsoftcoral/gorgonian&crinoid, Material. 801212B/3 (1), 810206A/5 (1), 810430A/24a(1). COLLECTION SITES. Chetlat, Lakshadweep (Laccadive) Islands; Remarks. The mostcommon ophiuroid species collected. Negombo, Sri Lanka; Ug Seukundo, PulaWe, Sumatra. Habitatand depth. Coral/rock, coral rubble; 5-10 m. 16. Ophiothrix (Acanthophiothrix) spinosissima Koehler, 1905 21. Macrophiothrix lorioli A.M. Clark, 1968 See. Clark & Rowe, 1971: 86; 112. See. Clark, 1968: 302; Clark & Rowe, 1971: 82; 115; Hoggett, Material. 810422B/4 (4), 810501G/5 (2), 810502F/6 (I ). 1991: 1108; Hoggett, 1992: 161; Rowe& Gates, 1995:414. COLLECTIONsites. UgSeukundo, Klah/UgSeukundo,PulaWe, Material. 810502D/2 (1), 810502E/1 (1). Sumatra. Collection sites. E Klah, PulaWe, Sumatra. Habitat and depth. Coral reef, 2-8 m. Habitat and depth. Coral reef, coral reef/subtidal rock; 5 & Remarks. Thespecimensrundownwell toO. (A.)spinossimain 10m. the key provided by Clark & Rowe (1971). However, 2 specimens fromstn810422B/4haveasingledarklinerunningthelengthofthe 22. Macrophiothrix nereidina (Lamarck, 1816) dorsal side ofthe arms, ratherthan aseries ofdark spots. See. Clark & Rowe, 1971: 86; 107 (as Ophiothrix (Keystonea) 17. Macrophiothrixaspidota (Miiller & Troschel, 1842) nereidina);Hoggett, 1992:228(asMacrophiothrix);Rowe&Gates, 1995: 426 (as O. (Keystonea)nereidina). SEE. Clark, 1968: 285: Clark & Rowe, 1971: 114; Clark & Courtman Stock, 1976: 137; Hoggett, 1992: 91. Material. 810421A/4 (1), 810422B/3 (2), 810430A/20d (1), 810430A/21c (2), 810430A/22b(1), 810501E/6 (1). Material. 810123B/6 (1), 810124A/8 (1), 810206A/6 (1), Collection sites. Ug Seukundo, Pula We, Sumatra. 810206A/7(1). Collectionsites. Negombo,ClosenburgPoint,Galle,AlaGala, Habitatand depth. Coral reef, coral/rock; 2-10 m. Galle, Sri Lanka. REMARKS. The species nereidina (which was placed in the Habitatand depth. Subtidal rock, coral /rock; 5-15 m. subgenusOphiothrix(Keystonea)byA.M.Clark, 1967)isincluded hereininthegenusMacrophiothrix.Hoggett(1991)statedthat 'itis particularlydifficulttodeterminetherespectiveboundariesbetween 18. Macrophiothrix demessa (Lyman, 1861) MacrophiothrixH.L.Clark, 1938andtwosubgeneraofOphiothrix, O. (Placophiothrix) H.L. Clark, 1938 and O. (Keystonea) A.M. See. Clark, 1968: 289; Clark & Rowe, 1971: 82; 114; Hoggett, Clark, 1967.'The differences between these taxahavetraditionally 1991: 1089; 1992: 117; Rowe & Gates, 1995: 412. reliedprincipallyonarmlength,shapeofdorsalarmplates,relative spinular armament ofthe disc plates including cover ofthe radial Material. 801212B/1 (1), 810424B/4 (2). plates(see A.M. Clark, 1967;Clark&Rowe, 1971).Thedifficulty Collection sites. Chetlat, Lakshadweep (Laccadive) Islands; inrecognising the supraspecific limits ofthesetaxa is made all the Seulakoe, PulaWe, Sumatra. more obvious by the treatment of the species Macrophiothrix 76 A.R.G.PRICEANDF.W.E. ROWE propinqua, placedinthesubgenusKeystoneabyA.M.Clark(1967), pointedgranuleswithlarger,conicalspinesinterradiallytowardsthe whilst Devaney (1974) described O. (Placophiothrix) westwardi edge of the disc. Clark & Courtman Stock (1976) include which has beenconsideredconspecific withpropinqua by Hoggett Placophiothrix phrixa H.L. Clark as a synonym of O. fulgens (1991). Later, in a far-reaching and critical treatment of Macro- (Koehler)whichinturnisincludedinthesynonymyofO.pellicula phiothrix, Hoggett (1992: PhD thesis) commits both the taxa byRowe(inRowe&Gates, 1995).Thespeciesthereforeappearsto Placophthiothrix and Keystonea to the synonymy of Macrophio- tobedistributedfromtheGulfofAdentotheIndo-Malayregionand NW thrix,transferringtheincludedspeciesoftheformertwotaxatothe the coastofAustraliain depthsof 10-116 m. lattertaxon andto whom this move is herein credited. 27. Ophiopteron elegans Ludwig, 1888 23. Macrophiothrixpropinqua (Lyman, 1861) See. Clark& Rowe, 1971: 84; 115; Rowe &Gates, 1995: 419. See. Clark & Rowe, 1971: 86; 107 (as Ophiothrix (Keystonea) Material. 810428D/3e (2), 810428D/5 (8). propinqua);Clark, 1980:537;Hoggett, 1991: 1130;Hoggett, 1992: 204; Rowe &Gates, 1995: 415. Collection sites. UgTapa Gadja, PulaWe, Sumatra. MATERIAL. 810428D/5 (5), 810501C/3 (1 juvenile), 810501D/1 Habitatand depth. Coral reef, subtidal rock/coral; 15 m. (1juvenile), 810501E/13 (1), 810501DF/1 (1juvenile), 810501G/1 (1), 810501G/5 (3), 8105011/2 (2), 810502F/3 (1), 810502G/3 (1). Family OPHIOCOMIDAE Collection sites. Ug Tapa Gadja, Ug Seukundo, E Klah, Pula We, Sumatra. Habitatanddepth. Subtidalrock/coral,coralrubble,coralreef, 28. Ophiarthrumpictum Miiller & Troschel, 1842 softcoral; 2-20 m. See. Clark & Rowe, 1971: 86; 121; Rowe& Gates, 1995: 385. Remarks. This species was first transferred to the genus Material. 810502H/1 (1). Macrophiothrix H.L. Clark, 1938 by A.M. Clark (1980) from Ophiothrix(Keystonea)A.M. Clark, 1967. Collection sites. E Klah, PulaWe, Sumatra. Habitatand depth. Subtidal rock, 2m. 24. Macrophiothrix variabilis (Duncan, 1887) 29. Ophiocomella sexradia (Duncan, 1887) SEE. Clark, 1968: 308;Clark&Rowe, 1971: 115; Hoggett, 1991: 1138; Hoggett, 1992: 218; Rowe &Gates, 1995: 416. See. Clark & Rowe, 1971: 86; 118; Devaney, 1974: 162; Material. 810206A/4 (1). Cherbonnier&Guille, 1978: 179; Rowe & Gates, 1995: 389. Collection sites. Negombo, Sri Lanka. Material. 81042D8/3d (1). Habitatanddepth. Coral/rock, 5 m. Collection sites. Ug TapaGadja, PulaWe, Sumatra. Habitatand depth. Coral reef; 15 m. 25. Macrophiothrix virgata (Lyman, 1861) Remarks. The single specimen fromPulaWe measures d.d. = 3 SEE. Clark & Rowe, 1971: 86; 113 (as Ophiothrix (Placophio- mupmp;eram.lo.s=t o1f0+thmem4 a(rbmrokspeinnenseairstdiips).tiWncittlhy tlhoengeexrcetphtainonthtehaltowtehre thrix) virgata); Hoggett, 1992: 236. ones, the character separating Ophiomastix sexradiata A.H. Clark Material. 810423D/2 (1). 1952 (known only from its type locality: BikiniAtoll, Marshall Is, SW Pacific) from Ophiocomella sexradia (Duncan)(identified Collection sites. Ug Bau, PulaWe, Sumatra. throughouttheIndo-WestPacificregionandpossiblytropicopolitan) Habitatand depth. Coral reef; 2-8 m. in Clark & Rowe's (1971) key, all other skeletal characters ofthe presentspecimenaccordwiththosedescribedasfittingO. sexradia REMARKS. See remarks under Macrophiothrixnereidina. (note 65, p. 118) by A.M. Clark. Cherbonnier & Guille (1978), following their study ofMalagasy material, concur with the com- 26. Ophiogymnapellicula (Duncan, 1876) ments expressed by A.M. Clark (in Clark & Rowe, 1971) and tentatively consider Ophiomastixsexradiata tobeconspecific with SEE. Clark & Rowe, 1971: 84; 117; Clark & Courtman Stock, Ophiocomella sexradia simultaneously agreeing with A.M. Clark 1976; 140(as O.fulgens); Rowe&Gates, 1995: 417. that the status ofthe genus Ophiocomella as distinct from Ophio- comaremainsdoubtful.Althoughthepresentspecimendoesnothing Material. 810504B/2 (1). toclarifythegenericstatusofOphiocomella,itdoessupportstrongly Collection sites. Rubiah, PulaWe, Sumatra. tthhee sviyenwontyhamtythperotpwoosendombiynaClhesrpbeocnineisearre&cGounislpleeci(f1i9c78a)n.dconfirm Habitatand depth. Softcoral; 10 m. 30. Ophiocoma dentata Miiller & Troschel, 1842 Remarks. The single specimen has ad.d. = 2.7 mm, a.l. = c. 20 mwimt.h Twihdeedipsicnkisamnodttnlaerdrocwrecarmeaamndbapnidnsk.,Aanndirtrheegualramrsloanrgeibtuadnidneadl See. Devaney, 1970: 13; Clark& Rowe, 1971: 86; 119; Rowe & Gates, 1995: 386. lineofcream spots isevidentalongthedorsal midlineofthe arms. The disc is covered (except for the radial shields) with minute, Material. 810502H/2b (1). INDIAN OCEAN ECHINODERMS 77 Collection sites. E Klah, PulaWe, Sumatra. Habitatand depth. Coral reef; 20 m. Habitatand depth. Subtidal rock, 2m. 33. Ophiocomapusilla (Brock, 888) 1 Remarks. This single specimen (d.d. c. 11.3 mm, distorted) exhibits two of the described colour forms for the species. At the SEE. Devaney, 1970: 25; Clark & Rowe, 1971: 86; 118; Clark & centre ofthe dorsal surface ofthe disc is a cream spot (c. 1.1 mm CourtmanStock, 1976: 174; Sloanetal, 1979: 106;Rowe&Gates, mm diameter).Thisis surroundedby anirregularring(c. 2.2 wide) 1995: 388. twhheicdihscisduonrisaflolrymlayndduvseknyt/rgalrleyyiwsirtehtidcaurlkaetredsdpoutssk.yT/ghreeyreomnaiancderreaomf Material. 810422E/4 (1), 810425C/2 (1), 810430A/21c (1), 810501E/3(1),810501K/4(1). background. COLLECTION sites. N Klah island, Sabang Bay, Ug Seukundo, 31. Ophiocoma erinaceus Muller & Troschel, 1842 Pula We, Sumatra. SEE. Clark & Rowe, 1971: 114;119; Clark & Courtman Stock, Habitat and depth. Coral/sand, coral conglomerate, coral reef; 1976: 173; Sloansa/, 1979: 106; Bussarawit&Rowe, 1985: 1 (as 2-10m. O. similanensis n. sp.); Rowe& Gates, 1995: 387. Remarks. The 5 specimens range in size from d.d. = 3-7.5 mm. Thecharacteristic,enlarged,tissue-coveredarmspines(seeClark& M8a1t0e4r2i7aDl/.5 (1)8,018211024A28/C3/6(1)(,2j8u1ve0n4i2le2sB)/,381(0104)2,8E8/1504(127jBu/ve1nil(e1)),, Rowe, 1971) appearon specimensfrom d.d. > 5 mm. 810430A/3 (2), 810501F/1 (1), 810501G/5 (3), 810501G/6 (2), 34. Ophiomastix annulosa (Lamarck, 1816) 810501K/1 (2). Collection sites. Chetlat, Lakshadweep (Laccadive) Islands; See. Clark & Rowe, 1971: 86; Rowe & Gates, 1995: 390. UgSeukundo, Ug Bau,LhoPriaLaot, UgMurung, UgTapaGadja, Pula We, Sumatra. Material. 810123A/3 (1), 810212A/3 (2), 810213A/4 (1). Habitatanddepth. Coralreef,coralconglomerate,coralrubble, COLLECTION SITES. Kakoni rocks. Pigeon Island & Unawatuna, Galle,Tangalla, Sri Lanka. subtidal rock; 2-25 m. Remarks. This is the commonest species of Ophiocoma col- rHoacbki;t3a-t10anm.d DEPTH. Coral reef, subtidal rock/coral, subtidal lected.Thecollectioncomprisessome26specimens,ranginginsize fromd.d. = 3.6-22.2 mm, which showclearly both colourchanges and development ofdisc granulation with growth. Juveniles up to 35. Ophiomastix caryophyllata Lutken, 1869 d.d. = c. 5 mm bear no granules and are usually marked radially See. Clark&Rowe, 1971:86; 120;Cherbonnier&Guille, 1978: a&crRoossweeac(h19r8ad5i)alfosrhitehledirwintehwascpreecaimesliOn.e,saismidleasncernisbiesd.bByyBuds.ds.ar=aw5i.t7 185; Rowe& Gates, 1995: 390. mm radiagtriangnulliensesarteodtehveeleodpgeedoafttthhee cdeisnctrewhoefretheandiisrcreagnudlaarlloinnge 1o0f Material. 810425C/1 (2), 810428C/7 (72), 810430A/21b (2). granules is developed around the peripheryjoining these radiating Collection sites. E Sabang Bay, Ug Bau, Ug Seukundo, Pula We, Sumatra linesandthus leavingbarethedorsal interradial andradial portions ofthe disc. By d.d. = c. 11 mm granules are developed over the Habitatand depth. Coral reef, coral/rock; 3-10 m. interradialbutnotradialregionsofthedisc,butgranulesarestillnot developed ventrally. By d.d. = 12.5 mm granules cover the whole surface ofthe disc except forthe radial shields which remain bare, Family OPHIONEREIDAE whilegranulesbegintoextendinawedgeshape,ontheventral side ofthedisc.Thisarrangementmayremain inspecimensuptod.d.= 14 mm but generally from about d.d. = > 13 mm the radial shields & become covered by granules. In specimens up to d.d. = 12.5 mm 36. Ophionereis dubia (Muller Troschel, 1842) some central and peripheral granules may be more prominent by tahreeirmoslrieghetvlyenmloyrreoeulnodnegdataendshmapoer,ebourtliensslaervgeernlsypesciziemde.nTshgeracnruelaems PSreiec.e, 1C9l8a3r:k6&7;RRoowwee,&197G1a:tes1,221;99C5l:ar4k08&. Courtman, 1976: 179; colourpatterndisappearswithincreasedd.d. and isabsentinspeci- Material. 810502D/5 (1). mens with d.d. > 11 mm. ItisveryclearthatOphiocomasimilanensisBussarawit&Rowe, Collection sites. E Klah, PulaWe, Sumatra. 1985 is based onjuvenile specimens of O. erinaceus, to the syn- Habitatand depth. Subtidal rock/sand; 10 m. onymy ofwhich O. similanensis is herein committed. 32. Ophiocomapica Muller & Troschel 37. Ophionereisfusca Brock, 1888 SCeoeu.rtmDaenvaSnteocyk,,11997706::2157;3;ClSalrokan&etRaolw,e1,97199:711:068;6;Ro1w18e;&ClGaartkes&, SReoew.e &ClaGrakt,esA,.1M9.951:945038:.69; 78; Clark & Rowe, 1971: 88; 122; 1995: 387. Material. 810421B/2 (1). Material. 801212A/3 (1). Collection sites. Nr. Seukundo, PulaWe, Sumatra. Collection sites. Chetlat, Lakshadweep (Laccadive) islands. Habitatand depth. Subtidal sand; 10-20m. 78 A.R.G.PRICEANDF.W.E. ROWE Family OPHIODERMATIDAE occurrence ofspinelets on the discs of4 specimens from Aldabra, western Indian Ocean and 2 specimens from Palau in the western Pacific Ocean which they examined suggested that Ophiochaeta 38. Cryptopeltagranulifera H.L. Clark, 1909 boschmaiA.H. Clark, 1964 is a synonym of Ophiochaeta hirsuta Liitken, 1869. They pointed out that Cherbonnier & Guille,1978, See. Clark& Rowe, 1971: 88; 128; Rowe& Gates, 1995: 394. haddescribedanewspecies,O.crinita,basedonasinglespecimen, from Madagascar, but did not comment further. In the present Material. 801212B/1 (1). collectionthespecimenfromtheLakshadweepIslandsaccordswith Collection sites. Chetlat, Lakshadweep (Laccadive) Islands. the description of boschmai in that the disc is granule-covered dorsally but bears spinelets on its ventral surface. The specimen Habitat and depth. Coral rubble; 8 m. from Pula We, on the other hand, accords with the description of Remarks. Originally described from Mauritius. Rowe & Gates crinitainthatthedisc iscovereddorsallyandventrallybyelongate (1995) describe the distribution as including tropical Australia, the spines. Considering the comments by Sloan etal. (1979), the two Indo-MalayanregionandPhilippineIslands.Thespecimenisiden- specimens reported herein are referrred to O. hirsuta, with the tified here fromthe Laccadive Islands forthe firsttime. implication that O.crinita Cherbonnier & Guille should also be referred to the synonymy on the basis that it exhibits the extreme 39. Ophiarachna affinis Liitken, 1869 spiny formofO. hirsuta. See. Clark&Rowe. 1971: 88; 123; Sloanetal., 1979: 1 1 1;Rowe 42. Ophiodyscrita instratus (Murakami, 1944) n. comb. & Gates, 1995:395. SEE. Murakami, 1944: 272 (as Ophiostegastus instratus); Material. 810425C/la,b (1). A.M.Clark, 1968: 320(as Ophiostegastus instratus; discussion) Collection sites. E. Sabang Bay, PulaWe, Sumatra. Material. 810124A/4 (1). Habitat and depth. Coral conglomerate; 3-6 m. Collection sites. AlaGala, Galle, Sri Lanka. Remarks. A.M. Clark(inClark&Rowe, 1971: Note 83, p. 123) Habitatand depth. Subtidal rock, 10-15 m. describes in detail colour variation in Ophiarachna affinis and O. mauritiensisdeLoriol,concludingthatspecificdistinctionbetween Remarks. This specimen, apart from its smaller size and fewer thetwoisdifficulttomake. Inthepresentspecimens withd.d. =22 nakeddiscplates,accordssowell withMurakami's(1944)descrip- mm, thedisc is uniformly brownish-grey andthedorsal side ofthe tionofOphiostegastusinstratusthatits identity isin nodoubt.The arms has a broad longitudinal central dusky band either side of species is, however, transferred to the genus Ophiodyscrita H.L. which is anarrowerpale band, the 3 bands being demarcated by 4 Clark, 1938 (type-species O. acosmeta H.L. Clark), with irregular, very narrowlongitudinal darkbands.Thiscorrespondsto Ophiostegatus Murakami, 1944 (of which instratus is the type- A.M. Clark's form C colour pattern. If O. affinis and O. mauriti- species) reduced to a junior synonym of Ophiodyscrita. The ensisareconspecific the species isclearly widely distributedin the distinctness of two genera has been questioned by A.M. Clark Indo-WestPacific region. (1968) and Guille & Vadon (1985) on the grounds of variation of granulation with increased specimen size. Tabulation of measure- 40. Ophiarachna robillardi de Loriol, 1893 ments(Table 1)takenfromoriginal speciesdescriptionsandsimilar details of the specimens from Sri Lanka, shows an interesting See. Clark&Rowe, 1971: 88; 123. picture. From this table it becomes apparent that only two species canberecognised:a)Ophiodyscritainstratus(Murakami, 1943)(d.d. Material. 810126B/4 (1), 810213A/3 (2). =7mm)ofwhichthelargerOphiostegastusnovaecaledoniaeGuille &Vadon (d.d.=9-11 mm)isasynonym,beinganextremeformof Collection sites. Galle, Tangalla, Sri Lanka. instratus in which many disc plates, including the radial shields, Habitat and depth. Coral reef, 3-5 m. havebecomeprominent(convex)andnakedofgranules;thisspecies possesses supplementary oral shields (granule covered in small Remarks. Thisisasignificantextensionofrangeforthisspecies specimens < 7 mm d.d.) and b) O. acosmeta H.L. Clark (d.d. = 5 described from Mauritius. The species is recorded as having 5 arm mm) with which O. pacifica (Murakami, 1943)(d.d. 4 mm) and spines (H.L. Clark, 1909; size not recorded) but the present 3 Ophiostegastus compsus A.M. Clark (1968) type locality Bahrain satped.cdi.me31n.s5hmavme7a-n9da1r0m-1sp1ianresmastpdi.nde.s=at21d..d5.m=m3;6.95-1m0m.armspines e(dv.edn.c=ov8e-r1i0n.g5ofmgmr)anaulpepseaorvetrotbheedciosncspwehciifcihc.areO.graacdouaslmleytaloshtaosnalyn from the oral shields (as in thetype ofacosmeta; d.d. = 5 mm)and 41. Ophiochaeta hirsuta Liitken, 1869 adoral plates (as in the type series ofcompsus; d.d. = 8-10.5 mm; though A.M. Clark (1968) does note that the largest paratype of SEE. Clark&Rowe, 1971:88; 127; Sloanetal, 1979: 115; Rowe compsus(d.d.= 10.5mm)hasasmallbarepatchdorsallyatthebase & Gates, 1995 398. of 4 of the arms). Supplementary oral shields are absent from compsus, according to A.M. Clark, and are not recorded foreither Material. 801212B/1 (1), 810425C/2 (1). acosmetaoxpacifica. Inbothinstratusandacosmeta. asrecognised Collectionsites. Chetlat,Lakshadweep(Laccadive)Islands;E. herein, it is clearthatarm spine numberincreases with size. Sabang Bay, PulaWe, Sumatra. There is clearly insufficient justification for recognising Ophiodyscritaand Ophiostegastusasseparategeneraonthebasisof Habitatand depth. Coral rubble, 3-8 m. the extent of disc granulation, for it is clear (see Clark & Rowe, Remarks. Sloan et al. (1979) concluded that variation in the 1971) that such differences occurbetween species included within

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