(Echinodermata: Holothuroidea) from Papua New Guinea

'' BULLETIN DE L'INSTITUT ROYAL DES SCIENCES NATURELLES DE BELG IQUE, BIOLOGIE, 75: 6I- 80, 2005 BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN, BIOLOGIE, 75: 6I-80, 2005

New records of Dendrochirotida (Echinodermata: Holothuroidea) from Papua New Guinea

by Claude MASSIN

Summary: Thi s paper describes 12 Dendrochirotida from Papua petmanent storage. Dredged specimens were immediately New Guinea of which II are new local records. Five of them have preserved in I 0% formalin and later transferred to 75 % buff­ been collected by dredging, the remaining 7 by hand-picking (div­ ered alcohol for permanent storage. ing and shore co ll ecting). The present work also provides the first Ossicles were prepared for light microscopy and scanning accou nt of brooding behaviour in a tropical Dendrochirotid a. Num­ electron microscopy (SEM). Juveniles were dehydrated in bers of dendrochirotids to numbers of other species of holothuroids from Papua New Guinea is compared with different areas of the two baths of Hexamethyl-Disilazane (Sigma) for more or Indo-West-Pacific Ocean. less two hours. SEM observations of ossicles and juveniles were carried out usi ng a JEOL JSM 5400 LV. Key words: Echinodermata, Holothuroidea, Dendrochirotida, brooding, Papua New Guinea. Results

Introduction Cucumariidae LUDWIG, 1894 Colochirinae PANNING, 1949 Only few studies exist on the holothuroid fauna of Papua Leptopentacta H.L. CLARK, I 938 New Guinea. THEEL (1886), LAM ~ERT (1885, 1889), H.L. Leptopentacta imbricata (SEMPER, 1868) CLARK ( 1921), LEVIN (1979) and MASS IN et al. (2002) re­ Fig. lA-G ported several aspodichirotids from Papua New Guinea, but no dendrochirotids. BROUNS & HEIJS (1985) listed Ocnus imbricatus SEMPER, 1868: 54, pl. I 1 fig . 2, pl. I 3, fig. holothuroids living in a seagrass bed close to Port Moresby, 12-13, pl. 14, fig 12-13; LAMPERT, 1885: 130; THEEL, 1886: where onl y a single species in a total of 15 belonged to the 76. Dendrochirotida. The bulk of the works published on Cucumaria imbricata; LUDWIG 1889-1892: 344, pl. 16, fig. holothuroids from Papua New Guinea deals with trepang 3; KOEHLER, 1895: 376; SLUITER, 1901 : 79; PEARSON, 1903: fisheries and take into account only the Aspidochirotida (see 190; KOEHLER & YANEY, 1908: 29. among others LONG & SKEWES , 1997: 12 species; KINCH , Trachythyone imbricata; PANNING, 1949: 42; A.M. CLARK 2003 : 15 species ; DESURMONT, 2003: 20 species). & ROWE, 1971: 182. This work li sts the Dendrochirotida from Papua New Guinea Parocnus im.bricatus; DEICHMANN 1941 : 93. (mainly from the north coast, Madang Province) and dis­ Leptopentacta imb1:icata; H.L. CLARK, 1938: 453; cusses their importance in holothuroid diversity. DEICHMANN , 1941 : 94; PANN ING, 1966: 57, fig. 4-5; A.M. CLARK , 1982: 489; ROWE, 1983: 159; L!AO &A.M. CLARK, 1995 : 476, pl. 23, fig. 16; LIAO, 1997: 166, fig. 96a-c; LANE Material and methods et al. , 2000: 490; LANE & VANDENSPEGEL, 2003: 153. Ocnus javanicus SLUITER, 1881: 342, pl. 4, fig. 1. The material examined has been co ll ected by shore collect­ Leptopentacta javanicus; PRICE, 1982: 11 . in g, by diving (between zero and 40 m depth) at Madang's Ocnus typicus THEEL, 1886: 75 pl. 6, fig . 10, pl. 14, fig. I I. Reefs and in Hansa Bay (Madang Province, Papua New Guinea) in 1983, 1987, 1989, 1990 and 1996, and by soft TYPE LOCALITY bottom dredging between 20-60 m depth in Hansa Bay in 1978. For the letterin g of the collecting points around Laing Bohol (Philippines). Island (Hansa Bay) see the map in CLAEREBOUDT et al. (1990). For a detailed map of the collecting points in the MATERIAL EXAMINED Madang reefs see JEBB & LOWRY ( 1995). Specimens coll ected by divers were anaestheti sed in 3.5 % IRSNB IG 29848/11; Hansa Bay (Madang Province, Papua MgCI, for a few hours, preserved in I 0% buffered formalin New Guinea); coli. B. TURSH & G. SEGHERS , 1l .x.1978, (pH 8~2) and later transfetTed to 75 % buffered alcohol for dredging at 2 1 m on sandy-muddy bottom; 7 specimens. I I 62 CLAUDE MAS SIN

DESCRIPTION forked processes but with a posterior notch; retractor mus­ cles of the calcareous ring short (Ill 0 of body length); longi­ Specimens small, ranging from 23x3 mm to 53x5 mm. Body tudinal muscles inconspicuous. One short Polian vesicle. pentagonal in cross section; colour in alcohol whitish-grey. Gonad not observed. Body tapering at both ends, particularly at the anal end. Pos­ Ossicles of dorsal and ventral body wall , large (up to 860 lli11 terior part curved. Body wall thin, firm, gritty to the touch, across), squarish or rounded (fig. lB) densely crowded translucent (intestinal content visible against light). Mouth scales. Between the large scales, nodulous buttons (fig I C), and anus terminal; 5 valves closing the mouth opening; anus 25-60 lli11 across, and small basket-like ossicles (fig. I D), 17- surrounded by numerous small overlapping calcareous 24 lli11 across. Tube feet with perforated plates (fig. I E), 70- plates. Tube feet large, conical, rigid, located in a single row 100 )lm across, nodulous buttons (fig. I F) with some bearing along each ambulacrum; tube feet largely separated from a long spine (fig. I F), and a few irregular rod-like ossicles each other, their number varying with body length: from 14 (fig. I G); no end plate. tube feet/row for a 23 mm long specimen to 37-38 tube feet! row for a 53 mm long specimen. Tentacles fully retracted, REM ARKS difficult to count; at least seven large bushy tentacles. Calcareous ring narrow with radial and interradial plates The fact that the species imbricata has been allocated to five nearly the same size (fig. I A); radial pieces without posterior different genera (Genus, Cucumaria, Trachythyone,

R 8 GS-W-® C I

0 0 0 0

E

1 mm A 500 j..tm 8 50 j..tm 20 j..tm C,E-G D

Fig. I. Leplopentacta imbricala (SEMPER, 1868). A: calcareous ring (R: radi al; IR: interradi al); 8 : body wall scale; C: nodulous buttons from body wall ; D: baskets from body wall ; E: perforated plates from tube fee t; F: nodulous buttons from tube feet; G: irregular rods from tube feet. New record s of Dendrochi rotid a (Echinodermata: Holothuroidea) fro m Papua New Guin e~ ' 63

Parocnus and Leptopentacta) illustrates that the diagnoses DESCRI PTIO N of many dendrochirotid taxa need revision. Here, the speci­ mens studied are all ocated to the genus Leptopentacta be­ Specim ens small , JJ.Ox3.5 mm, 11.5x4.6 mm and 16.0x2.0 mm, quadrangul ar in cross secti on. Colour in alcohol beicre · cause of their general aspect, positi on of their tube fee t, pres­ w hit~ bl a~k ~ ence of 5 oral valves and knobbed pl ates and irregul ar bas­ tentacles with base and main stem speckl ed with kets in the body wall. However, the aspect of the calcareous brown. M outh and anus terminal; mouth closed by 5 buccal ring of the material from Papua New Guinea is closer to the valves; tentacles I 0 (8 large + 2 ventral small); anus guarded genus Trachythyone than Leptopentacta. Leptopentacta by 5 anal teeth. Body wall gritty to the touch, thick, ri gid grisea CLAR K, 1938, the type species of the genus because of the presence of numerous large scales. Tube feet Leptopentacta, has the radial plates of the calcareous ring onl y along ambulacra, in a single row along the 2 dorsal indented by a deep posteri or notch whereas Trachythyone ambul acra and in a double row along the 3 ventral ambulacra muricata STUDER, 1876, the type species of the genus (fig. 2A); in the slender specimen (2 mm across) tube feet in Trachythyone, presents radial pl ates of the calcareous ring a sin gle row in all the ambulacra. Dorsal tube feet few, small , with a shall ow posteri or notch. Several species currently sometimes di fficult to see because of the body wall scales. classified in the genus Leptopentacta have their radial pl ates with onl y a shall ow posteri or notch, (e.g. Leptopentacta Calcareous ring without posterior processes but with poste­ ignava (LUDW IG, 1875); L. nova DEICHMANN, 1941). It thus ri or margin strongly undul ating (fig. 2B) due to deepl y seems that the vari ati on of the depth of the notch has value in notched interradial plates; radi al and intem adial plates of separatin g species but not genera. In '~ Y opinion the pres­ similar size; anteri or prolongati on of the radial plates promi­ ence of oral valves and small-knobbed perforated plates in nent; one short, pyrifonTI Pol ian vesicle; one stone canal end­ the body wall are characteristi cs of the genus Leptopentacta. ing in a spheri cal madrepori c plate located close to the cal­ T he specimens from Papua New Guinea are very close to careous ri ng; gonad made up of a few unbranched tubule.s; specimens fro m Hong-Kong illustrated by PANN ING (1966, retractor muscles of the calcareous ring attached at mid­ fig. 5). L. imbricata is a common species with a wide di stri­ body. bution from south-east Arabia to Indonesia and from the Dorsal and ventral body wall with large multi-layered, roun d, Philippin es to China. The species is new to the fa un a of oval or tri angul ar scales, 230-670 llJTI long (fig. 2C, D), nodu­ Papua New Guinea. lous buttons with 4-6 holes, 60-85 I-UTI across (fig. 2E, F) and Type of bottom and depth records are in accordance with nod ul ous-spiny baskets with (fig. 2G, H) or without (fig. 2J) published data on the species (SLUITER, 190 I; LANE et al., a transverse bar, 45-65 llJTI across. Tube feet with perforated 2000). rods, 11 5-220 1-1m long (fig. 2K) and perforated plates, 70- 125 I-UTI long (fig. 2L); the latter located close to end plate which is 140-200 !lll1 across (fig. 2M). Ossicles of tentacles highl y variable according to the specimen; main ones large, Plesiocolochirus CHERBONN IER , 1946 irregul ar, perforated rods 125-260 llJn long, located at the Plesiocolochirus australis (LUDW IG, 1875) base of the tentacles (fig. 3A ), small straight or curved perfo­ Fi gs 2A-M, 3A-D rated rods, 55-II 0 llJn long (fig. 3B ), small , branched ro­ settes, 20-40 I-UTI long (fi g. 3C), and rounded or elliptical ro­ Colochirus australis L UDW IG, 1875: 88, pl. 6, fig. !Sa-c. settes, 25-50 !lll11ong (fig. 3D); vari ous perforated plates also Plesiocolochirus australis; ROWE & GATES , 1995: 278 (syn­ present (fig. 3E). onymy); M ASS IN, 1999:83, figs 68a-k, 69a-j , 70, 11 2 h (syn­ onymy and list of records before 1999). REMAR KS C ucumari d sp. 3; LANE & V ANDENSPEGEL, 2003: 158. T he specimens of Plesiocolochirus australis from Papua TYPE LOCA LITY New Guinea are very similar to the ones described by M ASS IN ( 1999: figs 68-69) from Sul awesi. Ossicles of the Bowen (Australi a). tentacles present the same hi gh variability. T he species was previously known from Australi a, In donesia MATERI AL EXAM INED and New Caledoni a. It is new to the fa una of Papua New G uinea. IRSNB IG 27754/77, patch reef between Tabat and Woncrat Islands (Madang's Reef, Madang Province, Papua N:w Guinea), coli. C. MASS IN, I O.i x. l 990, by divin g, un der an Phyllophoridae O STERGREN, 1907 Echinopora sp. at 3 m depth, one specimen; IRSNB IG Thyoninae P ANNING, 1949 27754/102, patch reef in front of Ruo Island (Madang's Stolus SELENKA , 1867 Reef, Madang Province, Papua New G uinea), coli. C. Stolus rapax (KOE HLER & V ANEY, 1908) M ASS IN, 14.ix. J990, by di ving, under a block of Favites sp. Fig. 4A-H at 3 m depth , one specimen; IRSNB IG 28455/39, Durangit Reef (Hansa Bay, Madang Provin ce, Papua New Guinea), Cucumaria raJ?ax KOE HLER & YANEY , 1908.· 39 , pI. 2 ' f.,. g. coli . J.-C. BUSSERS 13.x. l996, by di vin g, among Halimeda 17- 19; DAN IEL & HALDE R,1974: 4 19. sp., Sm depth, one specimen. Stolus rapax; PANN ING , 1949: 462; A.M . CLARK & R OWE, I I

64 CLAUDE MASS IN

-0I ~0

Smm

1 mm

500 J..lm C,D 100 J..lm K 50 J..lm E-J,L 200 J..lm M

Fig. 2. Plesiocolochims aus1ralis (LUDWIG, 1875). A: ventral view of a specimen; 8 : calcareous ring (R: radial; IR: interradi al); C: large scales fro m dorsal body wall ; 0 : large scales from ventral body wall ; E: nodulous buttons from dorsal body wall ; F: nodulous buttons fro m ventral body wall ; G: baskets fro m dorsal body wall ; H: baskets from ventral body wall; J: basket without transverse bar; K: perforated rods from tube feet; L: perforated plates fro m tube feet; M: tube feet end plate. I I

New records of Dendrochirotida (Echin odermata: Holothuroidea) from Papua New Guinea 65

100 ~m A 50 ~m B,D,E 20 ~m c

Fig. 3. Plesiocolochirus australis (L UDWIG, 1875). Ossicles from the tent acles. A: large perforated rods; 8: small perfo rated rods; C: small rosettes; D: large rosettes; E: perforated plates.

1971 : 182; RAJP AL & TH ANDA R, 1999: 131 ; TH AN DAR (in narrow posteri or processes (fi g. 4A) and an anteri or groove press). fo r the inserti on of retractor muscl e; interradi al pl ates in a mosaic of pieces and with an anteri or tooth (fi g. 4A). One TYPE LOCALITY Pol ian vesicl e, I 110 of body length . Stone canal not ob­ served. Longitudinal muscl es inconspicuous. Gonad consist­ Bay of Bengal in g of a fe w unbranched tubules. Ossicles of body wall , both dorsall y and ventrall y, numerous MATERI AL EXAMI NED large, multi-layered pl ates, 150-500 ]ll11 across (fi g. 4B-C), and few small , simple perfo rated pl ates, 80-100 ]..lm across IRSNB IG 29848/10; Hansa Bay (Madang Province, Papua (fig. 40 ). In the tube feet, multi-layered plates at the base, New Guinea); coli. B. TURS H & G. SEG HERS, 17.x. l978 , 125-1 60 !lJn across, similar to those of body wall (fi g. 4 E); at dredging at 40-60 m on sand y-muddy bottom; 2 specimens. the tip , small perforated pl ates, some with 2 short pillars (fi g. 4F), and curved tabl es with 2 central pillars (fig. 4G); the DESCRIPTION latter arranged in concentric circles around end plate (fi g. 4H). Specimens small , 32x6.0 mm and 38x6.5 mm, width meas­ ured at mid-body. Body U-shaped, tapering stro ngly toward REMARKS both end s; near to mouth, the body is onl y 2.5 to 3 mm across. Mouth and anus terminal. Body wall very thin, soft, The specimens fit very well with description of Stolus rapax translucent , gritty to the touch because of the numerous large (KOE HLER & YANEY, 1908), a species known up to now onl y scales clearl y visi bl e to the naked eye. Colour in alcohol from the Bay of Bengal (at mouth of the Hughli Ri ver). S. whiti sh. Tube feet restri cted to ambulacra; close to the mouth rapax can be easil y separated fro m nearl y all the other spe­ and the anus they are short, in a zigzag row; mi d-ventra ll y cies at present included in the genus Stolus [S. albescens they are longer and more numerous (in 2-3 rows). Tentacles LI AO in LlAO & CLARK , 1995; 5. buccalis (STIMPSON, 1856); not observed because aquapharyngeal bul b full y retracted. S. canescens (SEMPER, 1868); S. cognatus (LAMPERT, 1885); Radi al plates of the calcareous ri ng with subdivided, long, 5. conjungens (SEMPER, 1868); 5. crassus LIAO & PAWSON , I I 66 CLAUDE MASS IN

3cm A 200 J.lm B,E 100 J.lm c 50 J.lm H D,F-H

Fig. 4. Sto/us rapax (KOEHLER & Y ANEY, 1908). A: calcareous ring (R: radial; IR: interradial); B: large plate from dorsal body wall ; C: large plate from ventral body wall ; D: small perforated plate from body wall; E: multi-layered plate from tube feet; F: small perforated plates from tube feet; G: curved tables from tube feet; H: end plates from tube feet.

200 I ; S. dentatus THANDAR (in press); S. micronodosus LlAO the presence of divided or undivided plates of the calcareous & PAWSON, 200 I ; S. minutus H.L. CLARK, 1938; S. ring, respectively. This is an example of two species that papillatus (SLUITER, 1887); S. pseudoalbescens THANDAR have a remarkably similar ossicle assemblage but at the same (in press); S. punctata (OHSHIM A, 19 15) ] due to the ahsence time different calcareous rings. By their calcareous ring tliey of nodulous buttons and the presence of thick multi-layered have been placed in separate famil ies (Phyllophoridae and plates in the body wall and the tube fee t. Onl y S. kilberti Sclerodactylidae) ! This questions again (cf supra RAJPAL & THANDAR, 1999 is also characterised by large Leptopentacta imbricata) the importance of the calcareous multi-layered plates and the absence of nodulous buttons. ring in dendrochi rote taxonomy, and the definition of several However, S. kilberti is readily separated from S. rapax by the dendrochirote genera which are in need of revision. In the size and the shape of the large plates (RAJPAL & THANDAR, meantime, I wi ll adopt the point of view of RAJPAL & 1999; THANDAR in press). As already noted by KOEHLER & THANDAR ( 1999) and refer the species rapax to the genus V ANEY ( 1910) and RAJP AL & THANDAR ( 1999) S. rapax is Srolus. also very close to Pseudothyone mosaica (KOEHLER & The species is new to the fa una of Papua New Guinea. V ANEY, 1910). The main differences between both species is I I New records of Dendrochirotida (Echinodermata: Holothuroidea) from Papua New Guinea 67

Phyllophorinae OSTERGREN, 1907 (fig. SG). Tentacles with numerous very narrow rods, 50-95 Phyllophorus GRUBE, 1840 llJll long (fig. SH). Phyllothuria HEDING & PANNING , 1954 Phyllophorus ( Phyllothuria) hypsipyrga (MARENZELLER, REMARKS 188 1) Fig. SA-H The calcareous ring and the tables with a hi gh spire are char­ acteris ti c of the genus Phyllophorus and the subgenus Orcu!a hypsipyrga MARE NZELLER , 1881: 135, pl. V, fig. I 0. Phyllothuria (see HEDING & PANN.ING , 1954). The speci­ Phyllophorus hypsipyrgus ; CH ANG & LIAO, 1964: 30, text mens are very close to the Phyllophorus (Phyllothuria) fig. hypsipyrga (MARENZELLER, 1881) reported by 0HSHIMA Phyllophorus ( Phyllothuria) hypsipyrga; HEDING & PAN­ ( 1912). They also present some affi nities with Phyllophorus NING , 1954: 151 , fig. 64a-c, fig. 65a-e (sy nonymy and ( Phyllothuria) discrepans (SLUITER , 1901) but are distinct by records before 1954); CLARK & ROWE, 1971: 184, pl. 29, fig . the presence of more numerous and narrower subdivisions of 19; RHO & WON , 1993: 127, pl. 3, fig s 1-9; LIAO, 1997: 211 , the posterior processes of the radial plates, by the regu lar ta­ fig. 126a-j; LANE eta!. 2000: 491 . bles, and by the very tall spire of the tables. The species was previously known from Korea, Japan, China and Indonesia and is new to the faun a of Papua New Guinea. MATERIAL EXAMINED

IRSNB IG 29848/3 , Hansa Bay (Madang Province, Papua Semperiellinae HEDING & PANNING, 1954 New Guinea); coli. B. TURSH & G. SEGHERS , 12.x. l978, Massinium SAMYN & TH AN DAR , 2003 dredging at 46 m on sandy-muddy bottom; 4 specimens; Massinium magnum. (LUDW IG , 1882) IRSNB IG 29848/8, Han sa Bay (Madang Province, Papua Fig. 6A-K New Guinea) ; coli. B. TURSH & G. SEGHERS , 17. x. l978, dredging at 40-60 m on sandy-muddy bottom; 2 specimens. Th yonidiwn magnum LUDWIG , 1882: 132. Neothyonidium magnum; HEDING & PANNING , 1954:197, TYPE LOCALITY fig. 98A-M; MASSIN , 1999: 88 , fig s. 72a-k, 73 , 113a (syn­ onymy). Japan. Massiniummagnum; SAMYN & THANDAR: 2003: 136.

DESCRIPTION TYPE LOCALITY Specimens small , ranging from 18x5 mm to I 04x20 mm . Ambon (Indonesia). Body cylindrical, with a short narrow tail distally. Mouth and anus terminal, an us surrounded by 5 anal papillae. Body wall MATERIAL EXAMINED gritty to the touch. Colour in alcohol from white to grey­ beige with brown areas. Calcareous ring and ossicles visible IRSNB IG 28455/51 , Laing Island HI H2 (Hansa Bay, through transparent body wall. Tube feet large, sparsely dis­ Madang Province, Papua New Guinea), coli . C. MASSIN , tributed over the whole body surface, more numerous 16.x. l996, by diving at 5 m depth in coarse sand , one speci­ ventrally. men (tentacles + introvert); IRSNB IG 28455/54, Laing Is­ Calcareous ring 1/7 to 1/6 of body length ; radial plates with land HIH2 (Hansa Bay, Madang Province, Papua New posterior processes, each made up of 5-7 pieces, the last ones Guinea), col i. C. MASS IN, 17 .x.1996, hand picking 6 m depth ( 4-7) very narrow (fig. SA) ; anterior part of radial plate with in coarse sand , one SP.ecimen (tentacles + introvert+ a piece a deep groove for the attachment of prominent retractor mus­ of body wall ). cles; interradial plates each with a strong anteri or tooth (fig. SA); retractor muscles attached at I /4 to 113 of body length. DESCRIPTION Polian vesicle, single, short; stone canal si ngle. In the body wa ll and tube feet , tables on ly. Table bases with Large species with tentacle crown up to 30 em across when one large central hole and 8-24 peripheral ones (fig. SB-E); fully extended Colour of large tentacles of li vin g specimen: majority with 8 large, regular peripheral holes (fig. SB , D). In at base, alternation of beige and black-brown bands, at tip the largest specimen the 8 peripheral holes of the tables are uniform black; introvert beige with two rows of black-brown small er (fig. SE) than in tables from small specimens. One tube feet along each ambulacrum; colou r in alcohol same but specimen (49 mm long) has a mixture of table bases with faded; body wa ll beige. Twenty tentacles in 2 circles: exter­ large and narrow hol es; rim of di sc smooth or with few short nal circle of I 0 large tentacles, inner circl e of 10 small tenta­ blunt spin es; table discs 60- 120 llJll ac ross, table height 50- cles. 150 11m; the small er the specimen the taller the tables; spire Calcareous ring simil ar to spec imens from Sulawesi (see of the tables with 4 pillars united by 2-6 cross beams, and MASSIN , 1999: fig . 72a-b). ending in a small crown of short spines. End plates of tube Body wa ll with only rosettes, 15-35 f.UTI long (fig. 6A). Tube feet II 0-240 f.UTI across (fig. SF), larger ventrally. Introvert feet with simil ar rosettes to those from body wall , 15-40 f.UTI with, miliary granul es, round or elliptical, 40-80 llJll across long (fig. 6B), in add iti on, inegular rods, 80-130 f.UTI long '' 68 CLAUDE MASSJN

R

0 0 0 -- B 0

2mm A

50J..tm B-H -----

Fig. 5. Phyl/ophorus ( Phyllothuria) hypsipyrga (MARENZELLER, 188 1). A: calcareous ring (R: radial; lR: interradial); B: tables fro m dorsal body wall ; C: tab les fro m ventral body wall; 0 : tables from dorsal tube feet; E: tables from ventral tube feet; F: tube feet end plate; G: miliary granules from introvert; H: rods from tentacles. II New record s of Dend rochirotida (Echinodermata: Holothuroidea) fr om Papua New Guinea 69

(fig. 6C) located close to end pl ate (fi g. 6D) whi ch is 300-320 tubular, 9 mm long with long posteri or processes (fig. 7 A) )-UTI in diameter, and a few tables reduced to discs, perforated made up of a mosaic of pieces; radial pl ates with an anteri or by 4 holes. Introvert with tables only (fig. 6E), spires 40-50 slit for the attachment of the retractor muscle; interradial )-UTI hi gh, discs 80-100 ~1 across, perforated by 4 large cen­ pl ates with an anteri or notch; retractor muscles 5 mm long; tral holes and 10-45 small peripheral holes. In tube fee t of tentacles fully retrac ted. Poli an vesicle 3.8 mm long. introvert, irregul ar perforated rods, 30-155 ~1 long (fi g. 6F), Madrepori c pl ate and gonad not observed. numerous rosettes, 30-65 f.lm long (fi g. 6G) and tables (fi g. Ossicles sparse in body wall and tube feet. Body wall with 6H); disc of tabl es round or ovate with smooth margin , 4 tabl es onl y, di scs 70 ~n across, perforated by 4 large central large central holes and 12-30 small, peripheral holes; spire holes and 4 small peripheral holes (fig. 7B); rim of disc 20-25 )-UTI hi gh, made o f 2 pillars ending in a bundle of blunt smooth; no pillar or 2 pill ars united by a single cross beam spines; end plates 600-630 ~1 across. In the tentacles, ro­ and ending in 2 diverging spines, sometimes bifurcate settes onl y, large at base (fig. 6J), small in second ary distall y (fig. 7B). Tube feet with tables with curved, perfo­ branches and close to extremities (fi gs. 6K, L); in the di stal rated di scs and a spire made up of 2 pillars fu sed at apex in a extremities, no ossicles. single point (fig. 7C); end pl ates small , 45-50 ~1 across, some star- shaped (fig. 7D). Introvert with numerous rosettes, HABITS 20-50 ~11 long (fig. 7E).

Body always buried in the sand , onl y tentacles and introvert REMA RKS visible; tentacles coming out at sunset and at night. T he specimen collected in Papua New Guinea presents many REMARKS affin it ies with Th yone bico rnis O HS HI MA , 19 15 as reported from China (LIAO & CLAR K, 1995). It differs mainly in the The specimens of Massinium magnum from Papua New form of the tabl es from the tube feet. The ones from China Guinea are similar to those specimens from Sulawesi de­ have a spire made of 2 separate pillars whereas the one from scribed by M ASS IN ( 1999, fi g. 72). They di ffer slightly by the Papua New Guinea has a spire made of 2 fused pillars (tack­ height of the spire of the introvert tables (shorter for the like). Since tables of the tube feet in some Thyone species specimens from Papua New Guinea) and by the perforated may present both forms of spire ( T. coma fa CHERB ONN IER, rods of the tube feet of the introvert (rods perfo rated by a few 1988, T crebrapodia CHERBONNI ER, 1988) this difference large holes fo r specimens from Papua New Guinea versus cannot be retained at the specific level. numerous small holes for specimens from Sulawesi). The specimen observed shows also affinities with T. The species is new to the fauna of Papua New Guinea. crebrapodia (particularly the specimens described by LI AO & PAWSON , 2001 ) and T. longicornis CHERBONN IER, 1988. However, in the original description of T. crebrapodia, Thyoninae PANNING, 1949 CH ERBONN IER mentioned tables in the introvert whereas in Thyone JAEGER, 1833 the specimen from Papua New Guinea onl y rosettes are Thyone bicornis 0HSHI MA, 19 15 present. The specimen observed di ffe rs from T. longicornis Fig. 7A-E by the much smaller size of the two points of the body wall table spires and by the greater number o f small holes in the Thyone bicornis OHSHIM A, 19 15 : 270, pl. 10, fig. 24a-d; tube feet end pl ates. The species is new to the fauna of Papua CH ANG & LlAO, 1964: 26, text fi g. ; LIAO & A.M. CLARK , New Guinea. 1995: 503, fig. 305a-e; LI AO, 1997: 198, fi g. 117a-e; W ON & RHO, 1998: 9, fi g. 2A-H; LANE et al. , 2000: 491. Non Thyone bicom is; YANG, 1937: 6, fig. 3a-g. Sclerodactylidae PANNING, 1949 Ciadolabinae REDING & P ANNING, 1954 TYPE LOCALITY Cladolabes BRANDT, 1835 Cladolabes acicula (SEMPER, 1868) Suruga Bay, south mid-Japan Cue umaria acicula SEMPER, I 868: 54, pl.15, fig. 11. MATE RI AL EXA MI NED Cladolabes acicula; M ASS IN, 1996: 42, fig. 29A-D, 30A-C (synonymy and li st of record s); IRSNB IG 25848/9, Hansa Bay (M adang Province, Papua Cladolabes aciculus; LIAO & A.M . CLAR K, 1995: 489, fig. New Guinea), coli. B . TURS H & G. SEG HERS, 17.x. l 978, 295a- d; LI AO , 1997: 18 1, fig. 105a-d; LANEetal.,2000:490. dredgin g at 40-60 m on sandy-muddy bottom; one specimen. TYPE LOCA LITY DESCRIPTIO N Viti Island (Fiji) Specimen small , 23,5x9 mm; body curved, banana-shaped, with mouth and anus sli ghtly dorsal; tube feet narrow, cylin­ MATERI AL EX AMI NED drical, sparsely scattered all over the body wall. Skin thin, soft, translucent; colour in alcohol greyi sh. Calcareous ring IRSNB IG 26700/199, Durangit Reef (Hansa Bay, Madang '' 70 CLAUDE MASS IN \iG-@-?}-@ ®-~-eYB@ D

50~m A-L

Fig. 6. Massiniummagnum (LUDWIG, 1892). A: body wall rosettes; 8 : tube feet rosettes; C: rods from tube feet; D: tube feet end plate; E: tables from introvert; F: irregular rods from introvert tube feet; G: rosettes from introvert tube feet; H: tables from introvert tube feet; 1: rosettes from tentacle base; K: rosettes from tentacle secondary shaft ; L: rosettes close to the smallest rami fications of the tentacles. II New records of Dendrochirotida (Echinodermata: Holothuroidea) from Papua New Guinea 71

50 Jlm B-E

Smm A

Fig. 7. Thyone bicornis OHSHIMA, 1915. A: calcareous ring (R: rad ial; lR: interradial); 8: tables from body wall; C: curved tables from tube feet; D: tube feet end plates; E: rosettes from introvert.

Province, Papua New Guinea), co li C. MASS IN, 05.xi.l983, Cladolabes schmeltzii (LUDWIG, 1875) by diver under a block of Leptoria plu)'gia (ELLIS & SOLANDER, 1786) at 7 m depth, one specim en; IRSNB IG Thyonidium Schmeltzii LUDWIG , 1875: 94, fig. 20a-b. 27598/36, Tab Island (Madang's Reef, Madang Province, Cladolabes schmeltzii; Ll AO & A.M. CLARK, 1995: 490, fig. Papua New Guinea), coli. C. MASSIN , 18.ix.l989, by diver 296a-b; LIAO, 1997: 183 , fig. I 07a-b; MASS IN , 1999: 96, fig; under a bl ock of Leptoria phrygia at 3 m depth, one speci­ 80a-e, 81 , 113d (synonymy and list of records); LANE et al., men. 2000: 490.

DESCRIPTION TYPE LOCALITY

Living specimens bright orange with brown-black tentacles; Bowen (Great Barrier Reef, Australia) in alcohol completely white; body cy lindrical, 74x30 mm and 68x20 mm. Tube feet in rows along ambulacra only, MATERIAL EXAMINED densely crowded ventrally, sparse dorsally ; mouth and anus terminal. Oss icles identical to the ones illustrated by MASSIN IRSNB IG 27598/61, patch reef NW of Guzem Isl and (1996: fig. 29, 30) for a specimen from Ambon. (Madang's Reef, Madang Prov ince, Papua New Guinea), co li. C. MASSIN, 23.ix.l989, by diver under a block of REMARKS Hydnophora exesa (PALLAS, 1766), 3 m depth, one speci­ men. Throughout its wide distri bution area (from Mauritius to Fiji and Hawaii) Cladolabes acicula is very co nstant in its colour DESCRIPTION pattern and oss icl e assemblage. The species is new to the fa un a of Papua New Guinea. Specimen small , 21 x8 mm , cy lindrical with mouth and an us '' 72 CLAUDE MASSIN

terminal. Colour in alcohol whitish yellow; tube feet in rows Afrocucumis stracki MASSIN, 1996 along the ambulacra only, more numerous ventrally than Fig. 8A-E, pl. I dorsally. Tentacles partly withdrawn, only the outer circle of 10 tentacles visible; anus without anal teeth. Ossicles identi­ Afrocucurnis stracki MASSIN, 1996: 40, fig. 28A-G. cal to the ones figured by MASSIN, 1999 (fig. 80). Pseudocucumis africanus; ENGEL, 1933: 16, fig. 15a-b, 16a-d. REMARKS

Cladolabes schmeltzii is very constant in its ossicles compo­ TYPE LOCAL!TY sition throughout its distribution area (see illustrations of Ambon (Indonesia). HEDING & PANNING 1954, CLARK & ROWE 1971 , CANNON & SILVER 1987, LIAO & CLARK I995, MASSIN 1999). The species is new to the fauna of Papua New Guinea. MATERIAL EXAMINED IRSNB IG 27598/31 , Tab Islands (Madang's Reef, Madang Afrocucumis DEICHMANN, I944 Province, Papua New Guinea), coil. C. MASSIN, 18.ix.l989, Afrocucum.is africana (SEMPER, I868) by diver under a block of Syrnphyllia radians EDWARDS & HAIME, 1849 at 3 m depth, 2 specimens; IRSNB IG 27598/ Cucumaria africana SEMPER, I868: 53, pl. I5, fig. 16. 45, patch reef on the left side of Guzem Island (Madang's Afrocucwnis ajricana; MASS IN , 1999: 96, fig. 79 (synonymy Reef, Madang Province, Papua New Guinea), coli. C. and list of records); SAMYN, 2003: 8, fig. I A-C, fig. 51 A. MASSIN, 2l.ix.I989, by diver under a block of Hydnophora microconos (LAMARCK, 1816) at 2 m depth, 4 specimens ; IRSNB IG 27598/53, patch reef N.W. of Guzem Island MATERIAL EXAMINED (Madang's Reef, Madang Province, Papua New Guinea), IRSNB IG 26700/110, Laing Island K3-4-5 (Hansa Bay, coli. C. MASSIN, 22.ix.l989, by diver under a block of Madang Province, Papua New Guinea), coli. C. MASSIN , Favites flexuosa (DANA, 1846) at 3 m depth, 2 specimens; 18.x.1983, by hand on the reef flat at low tide, under stones, 4 IRSNB IG 27598/76, patch reef in front of Guzem Island spec1mens. (Madang's Reef, Madang Province, Papua New Guinea), coli. C. MASSIN, 28.ix.1989, by diver under a block of Cyphastrea microphthamla (LAMARCK, 1816), 5 m depth, TYPE LOCALITY one specimen; IRSNB IG 27754/ 101 , patch reef in front of Querimba Island (Mozambique) Riwo Island (Madang's Reef, Madang Province, Papua New Guinea), coli. C. MASSIN, 14.ix.l990, by diver under a block DESCRIPTION of Goniastrea sp. at 5 m depth, one specimen; IRSNB IG 27754/103, patch reef in front of Riwo Island (Madang 's Specimens small, from 7x9 mm to 124x 14 mm, barrel­ Reef, Madang Province, Papua New Guinea), coli. C. shaped; living specimens brown-black dorsally as well as MASS IN, 14.ix.1990, by diver under a block of Agariciidae at ventrally; in alcohol, completely white, translucent; large 3 m depth, one specimen; IRSNB IG 27754/104, patch reef body wall ossicles visible in the translucent body wall. Tube in front of Riwo Island (Madang's Reef, Madang Province, feet large, in 2 rows on each ambulacrum. Mouth and anus Papua New Guinea), coli. C. MASSIN, 14.ix.1990, by diver terminal. Ossicles identical to those illustrated by MASSIN under a block of Pavona clavus (DANA, 1846) at 4 m depth, ( I996; fi g. 27) from a specimen fTom Ambon. Large lenticu­ one specimen; IRSNB IG 27754/ 118, patch reef in fro nt of lar plates 200-290 1J111 across. Guzem Island (Madang's Reef, Madang Province, Papua New Guinea), coil. C. MASSIN, I9.ix.I990, by diver under a REMARKS block of Platygyra sp. at 3 m depth, 3 specimens.

The size of the large lenticul ar plates and the absence of mil­ DESCRIPTION iary granules in the tube feet are characteristic of Aji-ocucumis africana and readily separates A. africana from Specimens from 23 to 55 mm long and 7 to I4 mm wide at A. stracki MASS IN, I996. Up to now A. africana has mai nly mid-body; cylindrical, tapering mai nly distally; colour of liv­ been found intertidally (see among others OHSHIM A 1916; ing specimens brown; colour in alcohol brown-beige with ROWE & DOTY 1977; SLOAN et al. 1979; CHERBONNIER white close to the mouth and the an us which are both tenni­ 1988; LlAO & CLARK 1995; MASSIN 1996; LANE et al. 2000; nal ; tube feet large, long, numerous, scattered all over the SAMYN 2003 , present study) whereas A. stracki lives in shal­ body wall at mid-body; rows (2) visible close to the mouth low waters ( 1-5 m depth). The species is new to the fauna of and the anus for most specimens ; only few specimens with Papua New Guinea. rows visible over the whole length. In most specimens, anal papillae surround anus. Tentacles full y withdrawn; genital papillae not observed. Contrary to the type materi al, poste­ rior processes of calcareous ring fragmented (fig. SA). Re­ tractor muscles prominent, attached at I /3 of body length I I

New records of Dendrochirotida (Echinodermata: Holothuroidea) fTom Papua New Guinea 73

5mm 100 J..tm 50 J..tm A B,C D

Fig. 8. Afrocucumis srracki MASS IN , 1996. A: calcareous ring (R: radi al; lR: interradial); B: perforated plates from body wall and tube feet ; C: tube feet end plate; D: C-shape rods from tentacles. A: adult; 8-D: juvenile.

from anterior end ; Polian vesicle, si.ngle, large; stone canal 140 !lJn across. In tentacles a few C-shaped rods (fig. 8D, E, single. pl. I H), I 00-135 !lJn long, with or without perforated ex­ Typical ossicles as large lenticular plates, 300-375 !lJn tremities. In the brooding female retractor muscles of the cal­ across, in the body wall ; miliary granules in the tube feet; careous ring reduced. rods with numerous perforations at their extremities in the tentacles. The number of miliary granules in tube feet highl y REM ARKS vari able within the species and not related to size of speci­ men. By their size and form of the ossicles the I 5 specimens from Four females with gonad well developed containing very Papua New Guinea are identical to the holotype and paratype large oocytes (nearly one mm across) and one female with (MASSIN, 1996, fig. 28A-G) from Ambon. doli olaria larvae in the gonad. One female with brooding ju­ Brooding is a well-known phenomenon among veniles, 1.6-1.8 mm long, approximately 80 free in the coe­ Dendrochirotida. O'LOUGHLIN (1994) and ALCOCK & lomic cavity and about 30 still in the gonads. All the oocytes, O'LOUGHLI N (2001) menti oned 32 cucumariid species, all doliolaria larvae or juveniles are at the same stage of devel­ inhabiting cold or temperate waters, with this behaviour. opment in the gonad of a female. All the material was col­ Some phyllophorids, a sclerodactylid and many psolids are lected in September ( 1989 & 1990) . also known to brood (O'LOUGHLIN, 1994); nearly all brood­ Juveniles with 10-25 tube feet in 3 rows ventrall y (the 2 pos­ ing species inhabit cold or temperate waters. Afrocucwnis terior tube feet particularly large: pl. I A) and 2-4 tentacles africana has been mentioned by OHSHIMA ( 1916) as a brood­ (pl. I B); calcareous ring reduced to a narrow ribbon, visible ing species in the temperate waters of Ushibuka (Kyushu, through the transparent body wall , of I 0 pieces: 5 radial (pl. Japan). This brooding activity has never been mentioned for 1C , left piece, !D) and 5 interradial (pl. l C, ri ght piece) made specimens of A. africana fro m tropical waters where the spe­ up of irregularly branching rods forming a tridimensional cies is very common. (MASSIN, 1999; SAMYN 2003) The mesh (pl. I C, D); contrary to the adults, radi al plates of juve­ closely related species A. stracki is the first tropical brooding nil es have anterior projecti ons and no posterior processes; Dendrochirotida reported in the li terature. ossicles of the body wall and tube feet are plates (fig. 8B, pl. Release of the juveniles from the coelomic cavity has never 1E -F), 100- 160 I-UTI across with large perforati ons; develop­ been observed but is supposed to occur through the anus or in g plates with 4 centrall y located knobs (pl. IE), large plates through a tear of the body wall. The synchronism in the de­ with 4 knobs or 4 short pillars (Fig. 8B , pl. I F) sometimes velopment of the juveniles and the very reduced retractor united by a crown; close to the tube feet end plate, perforated muscles of the incubating female suggest that it could also plates are rod-li ke (Pl. IJ); end plates (fig. 8C, pi.I G), 11 0- happen by autotomy of the aqua-pharyngeal bulb. I I 74 CLAUDE MASSIN

P l. 1. Afrocucwnis stracki MASS IN, 1996. 1. 6 mm long juvenile. A: vent ral view; B: frontal view with two tentac les visible; C: calcareous ring, radial (R) and interradial (IR ) pi eces; D: calcareous ring, radi al piece; E and F: ossicles from body wall ; G: end pl ate from tube fee t; H: ossicles from tentacles; J: ossicles from tube feet. 11 New records of Dendrochirotida (Echinodermata: Holothuroidea) from Papua New Guinea 75

Ossicles of the juveniles are rather different from those of the mouth without valves; anus surrounded by 5 anal papillae. adults, a well known phenomenon among holothurians Tentacles 10, ventral 2 smaller. Calcareous ring 1/4 of body (MASS IN et a!., 2000). The calcareous ring is also very differ­ length with paired long posterior processes (fig. 9B); radial ent between juveniles and adults. The tridimensional mesh of plates with an anterior groove for the insertion of retractor the calcareous ring of the juveniles is very similar to the one muscle; posterior processes long, narrow, made of numerous illustrated by CHERBONNIER (1953: pl. I i) for Leptosynapta pieces (fi g. 9B); interradial plates, with an anterior notch, milll.tla (BECHER, 1906), by BILLET (1988: fig. 18A-E) for made up of a mosaic of pieces extending to I /3 of the length Ypsilothuria bitentaculata attenuata (R. PERRIER , 1902) and of the radial plates (fig. 9B). Gonad pr~:;se nt in a 2 1 mm long by M ASS IN et al. (2000: Fig. 2A for Holothuria (Metriatyla) specimen, made up of a few undivided, thick tubules. scabra JAEGER, I 833. This structure of the early calcareous Body wall packed with perforated, rounded calcareous ring, present in three of the main holothurian orders plates, 20-60 1-lJn across (fig. 9C); the majority smooth , only (Aspidochirotida, Dendrochirotida and Apodida), must be some are nodulous, with perforations partly obscured (fig. considered a plesiomorphic character. 9D). Tube feet with 2 kind of ossicles: elongated, perforated The specimens were all found in shallow waters (2-5 m plates (up to 165 1-lJnlong)(fig. 9E), smallest ones often nodu­ depth) under coral blocks or in crevices located in the dead lous (fig. 9F); arched rods with a central spire-like perforated part of the coral. At Ambon they have been collected at simi­ process 45-55 l-lJ11 high (fig. 9G); end plates more or Jess 60 lar depth: 1-3 m (MASS IN 1996). They are not linked to a 1-lJn across (fi g. 9H). Tentacles with fine, straight rods, 25-50 particular coral species. 1-lJn long with forked and/or perforated extremities (fig. 91). The species is new to the fauna of Papua New Guinea. Its In one specimen, tentacle deposits also include a few X­ distribution is restricted to the eastern part of Indonesia (Am­ shaped or Y-shaped ossicles (fig. 9K), irregular rods (fig. bon, Misool) and Papua New Guinea. 9L), nodulous rods (fig. 9M) and rosettes (fig. 9N).

REMARKS Placothuridae PAWSON & FELL, 1965 Placothuria PAWSON & FELL, 1965 The specimens present many affiniti es with Placothuria Placothuria cf. molpadioides (SEMPER, 1868) molpadioides (SEMPER , 1868) in their general aspect, the po­ Fig. 9A-N sition of the tube feet, the calcareous ring, the ossicles of the tube feet and tentacles. However, the very large, thick body Ocnus molpadioides SEM PER, I 868: 55, pl. I 5, fi g. I 3; wall plates (250-800 l-lJ11 across) described by LIAO & A.M. LAMPERT, 1885: 133; T HEEL, 1886: I 18. CLARK (I 995) and illustrated by LIAO ( 1997) were not ob­ Cue umaria molpadioides; LU DW IG, I 889- I 892: 344. served. T his could be linked to the size of the specimens as Stolus molpadioides; PANN ING, I 949:· 463; A.M. CLARK & the present specimens range from 14 to 30 mm in length ROWE, I 971: 182; LlAO & A.M . CLARK, 1995: 497, fig. while those from China were 50 to 136 mm long (see SEM­ 302a-c, pl. 23, fig. 2. PER, I 868; LIAO & A.M. CLARK, 1995). It is known that Placothuria molpadioides; LIAO , I 997: 224, fig. I 34a-e; ossicles do change with advancing age, becoming more LANE et al., 2000: 49 I; LI AO & PAWSON, 2001: 83. spiny or nodulous (MASSIN, 1994) so that the absence of large plates in the specimens from Papua New G uinea could be explained by their small size. Because of the uncertainty TYPE LOCALITY in the identi fication the specimens are reported as China. Placothuria cf. molpadioides until a complete growth series of this species becomes available. MATERIAL EXA MI NED This species was only known from China (LIAO & A.M. CLARK , 1995; LANE et al., 2000) and is new to the fauna of IRSNB IG 2584811 , Hansa Bay (Madang Province, Papua Papua New Guinea. · New G uinea), coli. B. TURSH & G. SEGHERS, 15.x. I 978, dredging at 50 m on sandy-muddy bottom, one specimen. IRSNB IG 25848/4, Hansa Bay (M adang Province, Papua Discussion New Guinea), coli. B. TURSH & G. SEGHERS, 17.x. l 978, dredging at 50 m on sandy-muddy bottom, 4 specimens. The Dendrochirotida o f Papua New G uinea now include 12 species (see table I): II of them in the present work and DESCRIPTION Pentacta quadrangularis (LESSON, 1830) cited by BROUNS & HEIJS ( 1985). If we assume that in Papua New Guinea the Specimens small , from 14x3 mm to 30x4.5 mm (width meas­ Aspidochirotida are represented by at least 30 species (esti­ ured at mid-body). Body U-shaped, tapering at both ends, mati on based on surrounding areas: 35 for the Ton·es Strait, pentagonal in cross section (fig. 9A). Skin tough, gritty to the CLARK, I 921; 32 species for Sulawesi, MASS IN, 1999) and touch. Colour in alcohol: body whitish, tube feet yellowish. the Apodida by 13 species ( 13 for the Torres Strait, CLARK, Tube feet restricted to the ambulacra, in a single row along 192 1; l 5 species for Sul awesi, MASS IN, 1999), the the bivium and in 2 rows along the trivium (up to 3 rows at Dendrochirotida, with 12 species, represent 22% of the mid-body); base of the tube feet conical, rigid as body wall, holothuroid diversity along the coast of Papua New Guinea. tip of the tube feet cylindrical, soft. Mouth and anus terminal, It is well known that Aspidochirotida are characteristic of \\ 76 CLAUDE MASS IN

' IR

0,

0

~~------~------=~ c;::::::::::=~::::==::;:,:::::::=::=:::::::

1 mm 5mm 8 A 50 J..lffi 20 J..lffi C-H J-N

Fig. 9. Placothuria cf mo/p adioides (SEMPER, 1868). A: dorsal view of a specimen; B: calcareous ring (R: radial; IR: interradi al); C: smooth perforated pl ates from body wall ; 0: nodul ous perforated plates from body wall ; E: smooth perforated plates from tube feet; F: nodulous perforated plates from tube feet; G: arched rods from tube feet; H: tube feet end plate; J: rods from th e tent acles; K: X- or Y- shaped rods from tentacles; L: irregul ar rods from tentacles; M: nodulous rods from tentacles; N: rosettes fro m tentacles. II

New records of Dendrochirotida (Echinodermata: Holothuroidea) from Papua New Guinea 77

Taxon in current assignment Taxon in original assignment Family

Leptopentacta imbricata (SEMPER, 1868) Ocnus imbricatus SEMPER , 1868 Cucumaridae

Co lochim s quadrangularis (LESSON, 1830) Holothuria quadrmzgularis LESSON, 1830 Cu cumari id ae

Plesiocholodhirus australis (LUDW IG, 1875) Co /ochints australis LUDW IG, 1875 Cu cumariid ae

Sto/us rapax (KOEHLE R& YA NE Y, 1908) Cucunw ria rapax KOEH LER & VA NEY, 1908 . ' Phyllophoridae Phyllophonts (Phyllothuria) hypsipyrga (MARENZELLER, 188 1) Orcula hypsipy1ga MARE NlELL ER, 188 1 Phyll ophori dae

Massiniw11magnum (L UDW IG, 1882) Thyonidiummagnum LUDWI G, 1882 Phyllophoridae

Thyone bicom is 0 HSH IMA , 19 15 Thyone bicom is 0 HSHI MA, 19 15 Phyllophori dae

Clado/abes aciwla (SEMPER, 1868) Cucumaria acicu/a SEM PER , 1868 Sclerodactylidae

C/adolabes sc/uneltzii (LUDW IG, 1875) Thyonidium Sc hmeltzi i LUDW IG, 1875 Sclerodactylidae

Afi·ocztcumis qfricana (SEMPER, 1868) Cuculllaria qfricana SEMPER, 1868 Sclerodactylidae

Afrocucumis stracki MASS IN, 1996 Afrocucumis stracki MASS IN, 1996 Sclerodactylidae

Placo tlwria c.f molpadioides (SEMPER , 1868) Ocnus molpadioides SEM PER, 1868 Pl acothuridae

Table I: Li st of th e Dend rochiroti da known from Papua New Guinea

clear tropi cal waters whereas Dendrochirotida prefer temper­ Island , Kosrae Island and Guam is most probabl y linked to ate and cold waters (BAKUS, 1973). In tropical Indo-West their weak dispersion capacity and also to the rarity or ab­ Pacific shall ow-waters (± 20° N to 20° S) Dendrochirotida sence o f appropriate bi otopes around these remote islands to represent from 0 to 35 % of the shall ow-waters holothuroid harbour Dendrochirotida . di versity (see table 2). The low proportion of Dendrochirotida along the tropical The very low proportion or absence of Dendrochi rotida east coast of Afri ca is related, according to SAMYN & around remote islands as Aldabra, Cocos Islands, Ashmore T ALLON (in press), not onl y to their weak dispersion capacity Reef, Cartier Island, Rowley Shoals, Scott Reef, Christmas but also to the prevalent currents and to the presence of

Area % Dendrochirotida A uthors

Chri stmas Island 0 MARS H 2000

Kenya and Pemba Island 2 SAMY N 2003

Cocos Islands 3 MARSH 1994

Ashmore Reef & Cartier lsi 4 MARSH eta/. 1993

Kosrae lsi (Eastern Caroline Islands) 4 KERR 1994

Guam (Mariana Island s) 5 ROWE & DOTY 1977, KERR et a/. 1992

Somali a 6 TORTO NESE 1980

Aldabra (Seychelles ) 6 SLOA N et a/. 1979

Row ley Shoals & Scott Reef 7 MARSH 1986

Sou th Vietnam 18 LEVIN 1999

Sul awes i (Indo nes ia) 18 MASS IN 1999

Madagascar 20 Cl-IERBO NN IER 1988

Arabian Sea 2 1.4 SAMY N 2003

Pap ua New Gu inea 22 Present stu dy

Gul f of Aden 23.7 SAMYN 2003

Torres Strait 24 H.L. CLARK 192 1

Northern Austra li a 35 CANNON & SILVE R 1987

Table 2: Percentage of Dendroc hi ro tid a amongst holothu rian species in trop ical Indo-West Pac ific shallow-waters (±20°.N to 20°.S). 78 CLAUDE MASS IN ''

downwellings. CHERBONNIER, G. 1946. Sur un e holothurie de Quoy & Gaimard , Sampling techniques may also be taken into account to ex­ type d'un nouveau ge nre: Plesioco/ochirus. Bulletin du Museum pl ain , at least partly, the low proportion of Dendrochirotida d 'Histoire Naturelle de Paris (2) 18: 280-286 observed in some areas. In Papua New Guinea 42% (5112) of CHERBONN IER, G. 1953. Recherches sur les sy naptes (holothuri es the dendrochirotids have been obtained by dredging sandy­ apodes) de Roscoff. Archives de Zoo/ogie experimenrale el muddy bottoms between 40-60 m depth. The data, among generate 90: 163- 186. others, of SLOAN et al. ( 1979), TORTONESE ( 1980), MARSH CHERBONN IER, G., 1988. Echinodermes: Holothurides. Faune de (I 986), MARSH (2000) and SAMYN (2003) are based on di ver Madagascar 70: 1-292. and/or shore collecting onl y and very often in shall ow-waters CLAEREBOUDT, M., Cl. MASSIN & J. BOUILLON. 1990. A general (0-20 m depth). Sampling through dredging of the above survey of Laing Island environment (Papua New Guinea). Indo­ mentioned areas will possibly increase their dendrochirotid Malayan Zoology 6: 1- 23. diversity. CLARK , A.M. , 1982. Echinoderm of Hong Kong. In : B.S. MORTON & C.K. TSENG, (eds), The Marine Flora and Fauna of Hong Kong and Sou thern China, Proceedin gs of th e I st Intern ational Ma rine Biological Works hop. Hong Kong University Press, Hong Kong, Acknowledgement vo l. I: 485-50 I. CLARK, A.M. & F.W.E. ROWE , 197 1. Monograph of shall ow-water It is a pl easure to thank Drs F.W.E. ROWE, Y. SAMYN and A. Indo-West Pacific echin oderm. Trustees of th e British Museum of TH ANDAR for critically reviewin g the manuscript and im­ natural Hi sto ry, London , i-vii + 238 pp, pi s 1-31. proving the English; Dr P.M. O 'LOUGH LI N fo r its comments; CLARK, H.L. 192 1. The echinoderm fauna of Torres Strait: its com­ Dr D. V ANDENSPEGEL fo r the SEM photos; Dr Ch. DE position and its ori gi n. Department of Ma rine Biology of th e RIDDER fo r allowing me to study the material coll ected by Ca rn egie Instituti on of Was hington X: i- viii + 223 pp, 38 pis . Prof. B. TURSH and Mr G. SEGHERS in 1978. It is also a CLARK , H.L. 1938. Echinoderm from Australi a. Memoirs of the pleasure to thank the numerous persons who helped me col­ Museum of Comparative Zoology Harvard 55 (VIII): i-viii + 596 lect material and particularly S. LUXFORD, M. pp, pis 1-28 . CLAEREBOUDT, J. M ALLEFET and H. PILATE. The expedi­ DANIEL, D. & B.P. HALDER. 1974. Holothuroidea of the Indi an tions to Papua New Guinea were fin anciall y supported by Ocean with remarks on their distribution. Journal of the Marine several projects of the Belgian Funds for Joi nt and Basic Re­ Biological Association of India 16(2): 412-436. search, by the King Leopold III Foundation for Exploration DEICHMANN , E. 1941. 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