Memoirs of the Museum of Vicloria 53(2): 227-266 (1992) 31 December 1992 https://doi.org/10.24199/j.mmv.1992.53.12 NEW CUCUMARI1D HOLOTHURIANS (ECHINODERMATA) FROM SOUTHERN AUSTRALIA, INCLUDING TWO BROODING AND ONE FISSIPAROUS SPECIES

By P. Mark OLoughlin and Timothy D. OHara

Department of Invertebrate Zoology. Museum of Victoria, Melbourne. Victoria 3000, Australia

Abstract

O'Loughlin, P.M. and O'Hara, T.D., 1992. New cucumariid holothurians (Echinoder- mata) from southern Australia, including two brooding and one fissiparous species. Memoirs of the Museum of Vicloria 53: 227-266.

Twelve new species of cucumariid holothurians are described from oft' the coast of southern Australia. Pemoams gen. nov. is erected for one species which has no intcrradial plates in the calcareous ring. Cucuvitrum gen. nov. (1 species). Squamocnus gen. nov. (1 species) and Apsolidiuin gen. nov. (3 species) are erected for species with combinations of body and ossicle form not represented in known genera. The other species are assigned to Neocnus, Trachythyone (2 species). Ocnus. Ncocucumis and Ncocucumella. The new species of Pcntocnus and Neocnus brood their young. The new species of Squamocnus is fissiparous. The new species of Cucuviirum is host to a copepod parasite (Cucumaricolidae).

Introduction Joshua (1914) recorded the Indo-Pacific cucu- Southern Australian holothurians are known mariid Plesiocolochints spinosus (Quoy and mainlv from the works of Ludwig (1874), Bell Gaimard, 1833) from "Victorian waters". The (1887), Joshua (1912. 1914), Joshua and Creed ill-defined locality data with the specimens in (1915), Erwe (1913). H.L. Clark (1914, 1938, the Museum of Victoria create doubt about its 1946). Hickman (1962, 1978), A.M. Clark occurrence in the region. (1966), Rowe (1976, 1982) and Rowe and Vail Joshua (1914) and Joshua and Creed (1915) (1982). Including those described herein. 23 recorded from southern Australia the New Zeal- cucumariid species are now known from the and species Pseudocucumis (=Neocucumella) Australian states of Victoria, Tasmania, South bicolumnatus (Dendy and Hindle, 1907). the Australia and from the south-west of Western South American species Cucumaria (=Neopsol-

Australia (Table 1 ). Of these, 5 are abundant and idium) convergens (Herouard, 1901), and the well known for the region: Psolicliella adhaerens subantarctic species Cucumaria (=Trachy-

Hickman, 1 962, Staurothyone inconspicua (Bell. thyone) squamata (Ludwig, 1 898). Rowe ( 1 982) 1887), Pentacta ignava (Ludwig, 1874), Cucu- recorded Ocnus calcareus (Dendy, 1896) from mella mutatis (Joshua, 1914) and Neoamphicyc- off south-western Australia. This material has lus tividus Hickman. 1962. These have been been re-examined by us and in all four cases has described in detail by Hickman (1962) (P. been assigned to new species. H.L. Clark (1946: ignava as P. austraiis), and current distribution 389) proposed the name Cucumaria squama- and habitat notes given by one of the us (M.O'L.) toides for the specimen of C squamata, but, as in Marine Research Group of Victoria (1984). A.M. Clark (1966: 345) pointed out, this name is Amphkyclus mortenseni Heding and Panning, a nomen nudum, failing to meet the criteria is 1 954. also described by Hickman ( 1 962), occurs under ICZN Article 13 and replaced herein. in deeper water in eastern Bass Strait. Extensive collecting of from the Some essentially tropical species extend southern Australian coast by us and with associ- around the coast of Western Australia into the ates since 1978 has revealed many new holothu-

Great Australian Bight (Rowe, 1 982). These are: rian species. Most of the specimens were col- Mensamaria inlercedens (Lampert, 1885); Pen- lected off algae or rocks in the rocky shallows, in tacta crassa (Ekman, 1918); Pentacta quadran- depths of 0-2 m, and representatives of these gularis (Troschel, 1846); and Pentacta anceps species were observed live before preservation. (Selenka, 1867). Cucumaria bicolor Bell, 1887 Other specimens, including materia) from the (see Heding and Panning, 1954: 92) and Cucu- continental shelf, were examined from the col- maria striata Joshua and Creed, 1915 (see A.M. lections of the Museum of Victoria (NMV), the Clark, 1966: 345) have been referred to the syn- Australian Museum (AM), the South Australian onymy of Mensamaria inlercedens. Museum (SAM), the Western Australian

227 228 1'. MARK O'LOUGHLIN AND TIMOTHY D. O'HARA

Museum (WAM). the Tasmanian Museum 1815) with ten tentacles and conspicuous pos- (TM), the New Zealand Oceanographic Institute terior processes on the calcareous ring is (NZOI) and the Museum of Comparative considered to be in the Phyllophoridae. Zoology, Harvard (MCZ). The within the is Two of the new species brood their young: the also currently confused and ill-defined, particu- Neocnus species in two marsupia in the anterior larly between the subfamilies Colochirinac and dorsal body wall, and the Penlocmts species in Cucumariinac. Panning (1949), in his revision pouches on the internal anterior body wall. of the Cucumariidae (which included the final Seasonal coelomic brooding has been reported breakup of old heterogeneous taxa such as Cucu- for two south-eastern Australian cucumariids: maria de Blainville, 1834 and Thyone Oken. Staurothyone inconspicua (Bell, 1887) and 1815), distinguished the Colochirinae from the Neoamphicyclus lividus Hickman, 1962 (sec Cucumariinae by the presence of cup-shaped Materia et al„ 1991). ossicles in the body wall of the former. However, many of the species in the Cucumariinae. as then Terminology defined, were subsequently found to have cups or cup-like derivatives, or at least plates similar Terminology predominantly follows Clark in form to genera in the Colochirinae. Panning and Rowe (1971) or Pawson (1970). Ossicle (1971) finally restricted the Cucumariinae to types include cups (pi. 4c), rosettes (pi. 70. tables two genera. Cucumaria and Cladodactyia (fig. 8c), buttons (pi. 9a), plates (pi. 7d), multi- Brandt, 1835 without, however, publishing layered ossicles (pi. 3b), pedicel endplates (pi. emended diagnoses for these genera or either 6g), bar-like ossicles (pi. 9e), mesh-like ossicles subfamily. (small plates in pi. 6h) and rods (pi. 2h). All Panning (1962, 1964. 1966, 1971) further podia or tube feet are referred to as pedicels. A restricted the diagnoses of many of the genera sole is a flattened, delimited modification of the within the Colochirinae. Consequently recent ventral body wall with peripheral pedicels which regional studies (Rowe. 1970; Rowe and Paw- do not extend to the introvert or anus (pi. 2b). son, 1985; Thandar, 1985, 1986, 1987; Gutt, The term radii is used in preference to ambu- 1990; this paper) have added a plethora of new lacra (following Pawson, 1970). Left and right genera as new or existing species could no longer are relative to facing anteriorly along the dorsal be satisfactorily placed in existing genera. This surface. has led to a more natural classification, recog- Order Grubc, 1840 nizing structural differences between ossicle (restricted by Pawson and Fell, 1965) types, but more research is necessary, par- ticularly on the derivation of the various cup Cucumariidae Ludwig, 1 894 ossicles.

relationships 1 Remarks. The between "cucu- Panning ( 97 1 ) attempted to divide the Colo- mariid" and "phyllophorid" holothurians have chirinae into three distinct groups, based on the been confused. Panning (1949) and Hedingand number of ossicle types present and their func- Panning (1954) define the Cucumariidae and tion. An "Ocnus" group with two types of body the Phyllophoridae predominantly on the basis wall ossicles, a "Pentacta" group with three, and of the number of tentacles, ten in the Cucumari- the genus Pseudocnus with special denticulate idae and more than ten in the Phyllophoridae. plates which function as cups, giving the skin

Pawson and Fell ( 1 965) consider that the nature adhesive traction. But this division is unnatural of the calcareous ring is fundamental, placing as many of the species in the "Ocnus" group genera with posterior processes in the Phyllo- have three types of body wall ossicles, not the phoridae and those without in the Cucu- two required, and Pseudocnus is not the only mariidae. However, Clark and Rowe (1971), genus to use denticulate body wall plates to

Hickman ( 1 978) and Cannon and Silver ( 1 987) achieve traction. have continued to use the older basis for classi- The phylogeny of the Cucumariidae poses fication. Pawson and Fell's classification is used many problems, particularly due to the apparent in this paper. Consequently the subfamily incidence of convergent evolution. Panning Thyonidiinae (e.g., Cucumella Ludwig and (1971), mainly after Pawson (1966), listed the Heding, 1935 and Neoamphicyclus Hickman, following characters as primitive: posterior 1962) with 15-30 tentacles and simple calcar- extensions on the calcareous ring; pedicels eous ring is included in the Cucumariidae, and restricted to the radii; and an imbricating the subfamily Thyoninae (e.g. Thyone Oken, armour of multi-layered scale-like ossicles i ' 1 '

NEW HOLOTHURIANS FROM SOUTHERN AUSTRALIA 229

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from all other gen- covering the whole body. No extant cucumariid clearly distinguish Pentocnus the Cucumariidae. The shallow, genus has all of these characters, but some gen- era within form of the plates suggests a era, e.g., Leptopentacta H.L. Clark, 1938, do saucer-like, spined relationship with the possess small extensions to the radial plates in possible form or functional cucumariid genera. the calcareous ring, and non-imbricating multi- cup ossicles present in other plates possibly layered ossicles. The denticulate body wall indicate some relationship with the genus Pseudocnus Panning, 1949. Colochirinae Panning, 1949 Some cucumariid genera have reduced inter- Pentocnus gen. nov. radial calcareous ring plates, but no other den- drochirotid genus lacks them completely. In the Diagnosis. Worm-like form, lacking sole or subfamily Thyonidiinae. Athyonidium Deich- modified ventrum; 10 irregularly developed mann, 1941 has very reduced interradials, often dendritic tentacles; calcareous ring discontinu- obscured by muscle; the resulting ring is also dis- ous, 5 spaced radial plates without posterior continuous. In Amphicyclus Bell, 1 884 the inter- prolongations, no interradial plates; pedicels radials are also reduced, but they adjoin the present on all radii, absent interradially. Body larger radial plates in a continuous ring. wall ossicles small, elongate, perforated. Hat or saucer-like, often denticulate plates, with blunt Pentocnus bursatus sp. nov. vertical spines; lacking true cups; pedicels with Plates la, 2e-g, Figure 1 similar plates to body wall, and endplates; ten- tacles with irregular perforated bar-like ossicles, Pseudocnus sp. - Roweand Vail, 1982: 224. small perforated plates and rods. 'cucumariid" sp - O'Loughlin, 1991: 225-226, fig. 4. Type species. Pentocnus bwsatus sp. nov. Material examined. Holotype. Victoria, Cape (five), in ref- Etymology. From the Greek pente Paterson, rocky shallows, algal and sponge epifauna, erence to the plates in the calcareous ring, with 14 Feb 1981, M. O'Loughlin, M. Nyhuis and C. Ocnus (masculine). Walker, NMV F57549. Paratvpes. Victoria, Cape Paterson, type locality Remarks. The lack of interradial plates in the and date. AM J22754( 1 ); NMV F58629( 1 juv); 1 8 Jan calcareous ring, the irregular development of the 1 980. NMV F54236( 1 ); rocky shallows, algal epifauna. tentacles, the delicate form of the ossicles, and 6 Mar 1983, NMV F53762(l): 29 Jan 1988. NMV the other possible paedomorphic characters, F54181(l).

B

= Figure 1 . Pentocnus bursatus gen. et sp. nov.: a, transverse section of body, scale bar 1.0 mm; b, 2 adjacent radial plates from the calcareous ring; c, side and top view of a dorsal body wall plate, scale bar = 0.01 mm. NEW HOLOTHURIANS FROM SOUTHERN AUSTRALIA 231

South Australia, Beachport, 1 39°59 37°29.3'S. 6'E tiated (0.8 mm long). The brood juvenile is well- 4 m, 14 May 1990. NMV F59207(l). differentiated (1.4 mm long). Both are together Other material. Western Australia, Rottnest I.. Cape in the coelom in a membranous sac which is Vlamingh. outer reef Hat, 9 Jan 1991, WAM 239- attached to the anterior, left ventral body wall. 91(1): Radar Reef, Sarxa.ssum/Cmoplwra mat, 1 3 Jan There is 1991. WAM 255-91(1). no change in the body wall at the point of attachment, nor any apparent aperture to the Description. Up to 15 mm long (live, extended). exterior. 1 mm wide; body worm-like, round in transverse NMV F54236 (10 mm long, tentacles with- section, lacking sole or modified ventrum, intro- drawn) has a dorsal gonad, 3 brood embryos and vert not distinct, mouth anterior, anus posterior; 2 juveniles. The gonad has 6 sac-like caeca, wall body thin, very extensible; 1 dendritic ten- graded in size, the 3 smallest with about 3 small tacles, vers' irregularly developed, about 6 fully white eggs, and the 3 largest each with one large developed, about 4 small or bud-like. One row of developing brown egg or embryo (up to 0.8 mm pedicels on each radius, with up to 18 pedicels in long). One brood embryo is in the coelom near a row, slightly more numerous ventrally; inter- the gonad. Two other brood embryos showing radial areas lacking pedicels. Calcareous ring radial pedicels, and 2 well-developed brood juv- comprises 5 spaced radial plates, no interradial eniles (up to 2.3 mm long) are associated with a plates; no posterior prolongations; anterior pro- thin-walled sac in the anterior left lateral jections constricted basally, widened and coelom. The sac is attached to the interradial rounded distally with apical notch; wide, deep, body wall where there is a body wall fold and rounded notch posteriorly, creating pairs of opening. NMVF54181 (8 mm long) has 2 undif- pointed projections (the 5 plates of the calcar- ferentiated embryos with firm brown ovoid eous ring are clearest in specimens NMV cases (1 mm long). Each embryo is in a thin- F57549. NMV F53762. NMV F59207, WAM walled sac which has a small growth attachment 239-91.) Dorsal madreporite; single, left to to the coelomic body wall. One is in the left ventral, polian vesicle. dorsolateral mid-body, the other in the right Body wall ossicles of one type: small, elongate, ventral anterior. At the point of attachment of perforated plates, with blunt vertical spines, the anterior embryo the body wall has broken often denticulate, flat or shallow saucer-like, up down to create a small opening. A gonad with a to 0.09 mm long: typical plate with 2 large cen- few small white eggs is present. AM J22754 (8 tral holes, one smaller hole near each end, 1-6 mm long, tentacles not withdrawn) has 2 undif- very small holes at ends; plates frequently den- ferentiated embryos (one in a gonad caecum, 0.7 ticulate at one end or one side; ossicles may- mm long), an embryo with radial pedicels, and a imbricate or be spaced in extended body wall. well-developed juvenile (2.2 mm long) in a Pedicels with body wall plates; endplates with membranous sheath in the left anterior coelom. irregular angular perforations, about 0.1 mm NMV F53762 (5 mm long) had 3 brood embryos wide. Tentacles with small irregular plates with (up to 1.5 mm long) in separate thin sacs large angular perforations, up to 0.07 mm wide; attached to a scar or fold in the body wall, one in abundant irregular rods with divided, twisted, the right dorsolateral coelom in the mid-body, 2 perforated ends, sometimes branched, typically close together in the left ventrolateral coelom 0.07 mm long; a few large, irregular, branched, anteriorly. There is only part of the gonad

perforated bar-like ossicles, up to 0. 1 3 mm long; remaining. NMV F59207 (5 mm long; tentacles a few elongate rosettes up to 0.05 mm long; some withdrawn) has one well-developed bursal juv- body wall plates, lacking spines. enile attached to the right anterior intracoelomic body wall, and a gonad with one large and a few- Live colour. Body and pedicels reddish brown; small white eggs. end rim of pedicels dark reddish brown; tentacle trunks brown, branches faintly reddish brown; Etymology. From of the Greek bursa (purse), colour persists in alcohol. with reference to the intracoelomic sacs in which brood juveniles develop. Reproduction. The holotype (6 mm long, ten- tacles withdrawn) has a dorsal gonad, 1 brood Distribution. Rottnest I., Western Australia, to gonad has 5 embryo and 1 brood juvenile. The Cape Paterson. Victoria. 0-4 m. sac-like caeca, graded in size, the smallest caeca with 1-2 small white eggs, the largest caecum Remarks. This very small species reproduces by long). brooding in intracoelomic with I brown egg or embryo (0.3 mm The sacs which are ran- brood embryo is dark brown, ovoid, undifferen- domly attached to the anterior body wall. 232 P. MARK O'l.OUGHUN AND TIMOTHY D. O'HARA

Embryos or juveniles may be present singly or tral tentacles reduced - does not exist in any together in the sacs. The very thin-walled sacs previously described cucumariid genus and war- lack ossicles and appear to be derived from the rants the creation of a new taxon. Of the other egg rather than from the body wall. At the point genera, Pseudoenus Panning, 1 949 and Pseudoc- of attachment of the sacs, with their developing nella Thandar, 1 987, are most similar. However, embryos or juveniles, the body wall may Pseudoenus lacks multi-layered ossicles and undergo change, is sometimes gathered into a Pseudoenella has ten equal tentacles, interradial small fold, and may break down to create a small papillae, and reduced cups. opening. Individuals produce few eggs but may Cueuvitrum rowei sp. nov. have all reproductive stages present, from small egg to fully-developed brood juvenile. Only one Plates lb, 3a-g, Figure 2 egg appears to mature at any one lime. All six Stereoderma sp. — A.M. Clark, 1966: 346-347, fig. adults exhibit bursal coelomic brooding. All 9. observed gonads contain eggs and are similar in Pseudoenus sp. — Rowe, 1982: 466, fig. 10.32b, pi. form, suggesting hermaphroditic or patheno- 32.2. — Rowe and Vail. 1982: 223. genic reproduction. The collection of brooding Material examined. Holotype. Victoria, Cape adults in January, February, March and May Patcrson, just below low tide level, 18 Jan 1980, M. suggests non-seasonal reproduction. O'Loughlin, T. O'Hara and J. Stephenson, NMV Ocnus sacculus Pawson, 1 983, from New Zeal- F57356. and, has a similar brooding habit. Intracoelomic Paratypes. Victoria, type locality and date, NMV sacs are attached to the dorsal intcrradial F5424l(5); Apollo Bay, Marengo, Hayley Point, just anterior body wall and contain up to 9 embryos. below low tide level, I 1 Jan 1980. NMV F54240(2). The internal sac walls are thin and transparent. Other material (partial list; all found as algal epi- The body wall forming the external wall of the l'auna, 0-2 m, unless otherwise stated; # indicates material with intracoelomic sac has fewer ossicles than elsewhere, and Paw- copepod parasites). son (1983: 228) suggests that "birth" occurs by rupture of the body wall. This species is easily distinguished from P. bursatus by the form of the calcareous ring, the form of the ossicles, body form, and distribution of pedicels. P. bursatus, known from only three widely separated localities, is found amongst algal tufts and sponge in the rocky shallows.

Cueuvitrum gen. nov.

Diagnosis. 10 dendritic tentacles, ventral 2 smaller; pedicels mostly confined to radii, a few scattered on dorsal and lateral interradii. Body wall ossicles numerous knobbed perforated plates, irregularly oval, some with the end or one side denticulate, some large multi-layered ossicles; tentacles and pedicels with bar-like perforated ossicles; pedicels with endplates. Calcareous ring simple, without posterior pro- longations; 10 plates tapered anteriorly, notched posteriorly.

Type species. Cueuvitrum rowei sp. nov.

Etymology. From the Latin vilrum (glass), in ref- erence to the glassy multi-layered ossicles in the body wall, with part of the family name, Cucu- mariidae (neuter).

Remarks. The combination of the following Figure 2. Cueuvitrum rowei gen. et sp. nov.: a, trans- characters - knobbed plates, multi-layered oss- verse section of body, scale bar = 1 .0 mm; b, radial and icles, pedicels mostly confined to radii, and ven- interradial plates from the calcareous ring. ,

NEW HOLOTHURIANS FROM SOUTHERN AUSTRALIA 233

Western Australia. Rottncst I., Strickland Bay, 20 elongate, slightly keeled midventrally, distinct Jan 1991, 400-91(1 juv); WAM Natural Jetty, 15 Jan dorsolateral and ventrolateral edges, body pen- 1991, WAM 393-91(1 juv); Cape Naturaliste. Eagle tagonal in transverse section; weakly developed Bay, 25 Feb 1975. WAM 556-89(13). oral valves formed by anterior radial body wall South Australia. Ceduna. Cape Vivonne, 16 Jan projections; mouth orientated anteriorly; exten- 1991, NMV F59209(4); Strcakv Bav, Point Wcstall, sible anal cone, often upturned; body wall firm, 15 Jan 1991, NMV F5921 l(i>; Greenlv 1., 28 Nov crystalline, with microscopic vitreous spots; 10 1976. SAM K 1 800( 1 ): Arno Bav. 22 Feb 1 988. 2-3 m, SAM K1813(l): Kangaroo I.. Emu Bav, 17 Jan 1990. dendritic tentacles. 2 ventral ones distinctly NMV F57550O): Gulf St Vincent, Normanville. 11 smaller; distinct modified ventrum, not a sole;

Nov 1 988. NMV F54386( 1 ): Robe. 9 Jan 1 990. NMV distinct introvert, lacking pedicels; 5 anal teeth. F575510). Pedicels in 2 rows on all radii, extending to Victoria. Cape Otway, Crayfish Bay, 31 Dec 1980, introvert and anus; ventrolateral and midven- NMVF54221(1 and 1 juv); 10 km NE of Apollo Bay, tral radii with up to 60 pedicels in each row; "Mullet Holes", 6 Jan 1981. NMV F57949(4): NMV dorsolateral series irregular, pedicels small; a F57948(l#); 10 Jan 1987. NMV F54218(5): NMV few small pedicels interradially on dorsal and F58640(l#): Port Phillip Bay, Altona, 14 Sep 1980, NMV F54228(8); NMV F58641(l#); Williamstown, lateral surfaces, used for grit attachment; single II Apr 1990. NMV F58643(l#): NMV F59208(5): very extensible small pedicels on each radius

38° 1 1 1 anteriorly and around anus. ring 7'S. 44°38'E, 3-6 m. 30 Mar 986. SPPS stn 3B 1 Calcareous NMV F57547(6#); Portsea Jetty, 4 m. 13 Mar 1975, lacking posterior prolongations; 5 radials with AM J 10857(1); Cape Schanck. Bushrangers Bay. 28 blunt anterior projections, often notched, large Mar 1981. NMV F54224(3 juv); Flinders, Mushroom notch posteriorly; intcrradials with pointed Reef. 16 Nov 1990, NMV F59206(l): Merncks, 27 anterior projections, wide shallow posterior May 1989, NMV F54395(2); Phillip I., Kitty Miller scallops. Madreporite dorsal; single polian ves- Bay, 13 Feb 1988, NMV F542 15(5); 1 kmEofHarm- icle left lateral; intestine runs dorsally from ers Haven. 300 m offshore, 4.5-6 m. 6 Mar 1 982. CPA mouth to mid-coelom, loops left back to mouth, stn 1 5. NMV F53772(2); Cape Paterson. 28 Apr 1 989. NMV F54394(5); NMV F58642(l#); Shack Bav. 4.5- then runs vcntrally to anus. Dorsal body wall ossicles of 5 types: 1) 12.2 m, 4 Mar 1 982, CPA stn 4, NMV F5377 1( I ); Wil- Abun- sons Promontorv, Hobbs Head, 12.5 m. 9 Feb 1982. dant perforated plates, heavily knobbed, irregu-

WPNPA stn 42,' NMV F53773(l juv); Rabbit I.. 23 larly oval, frequently thick and rounded at an Apr 1983, NMV F53783(2); eastern Bass Strait. end or edge, the opposite side or end denticulate; 39°0'S, 147°30.5'E. 28 m. 28 Jan 1971. NMV some plates fir-cone shaped, wide end smooth, F59203(l); Cape Everard. 8 Apr 1984. NMV narrow end denticulate; many plates with pro- F57943(2); Mallacoota. Bastion Point, 5 m, 6 Apr jections, bars, further developing layers, com- 1989. NMV F57363(6); Gabo 1.. 4 m, 4 Apr 1989. monly up to 0. 12 long; closely situated in NMV F573640). mm 2 or more layers within the body wall. 2) Scattered Tasmania. King 1., Rockv Cape, 2 m, 1 5 Mar 1 988. large multi-layered ossicles, irregularly oval, up NMV F5736 1 (5): Lulworth. Black Rock Point, 22 Nov 1982, NMV F53788(2); Waterhousc Passage. 23 Nov to 0.48 mm long, with 3-5 perforated knobbed 1982. NMV F53789(2): Bicheno, 7 m, 22 Mar 1988. layers of decreasing size, one end sometimes

NMV F57544(4): Swansea, 8 km S. 4 m, 2 1 Mar 1 988, denticulate. 3) A few irregular smooth perfor- F57360(5); F57548(2#); Eaglehawk NMV NMV ated plates, up to 0. 1 2 mm long. 4) A few irregu- Tinderbox, 29 Neck, 15 Feb 1991. NMV F592I60); lar smooth or lumpy or knobbed button-like May 1974, AM J 10855(1); D'Entrecasteaux Channel, ossicles, typically 0.07 mm long. 5) Rosettes in Esperance Point, 13 m, AM J22665(7#): Bruny I., some New South Wales specimens (AM J 10883, Adventure Bay. 15 m. 23 Apr 1991. AM J22667(3#); AM J 1 2735), up to 0.04 mm long. Plates of ven- Sadgrove Point, 7 Mar 1974. AM J 10856(1); South- trum similar to dorsum; multi-layered ossicles port, Lady Bay, 3 m, 1 1 Oct 1977, AM Jl 1164(1). New South Wales. Disaster Bav. Green Cape Light- less developed. Plates of introvert fewer, similar house. 16 m, 13 Feb 1973, AM J 12548(1); Twofold to body wall, more finely knobbed, none multi-

Bav, 3 m. 29 Mar 1985. AM J 1 9909(6); Eden, Cocora layered. Pedicels with irregular, bent or curved, Jervis Bay. 3 m, Point, 25 Nov 1 984. NMV F53786(3); perforated bar-like ossicles, often widened cen- 3-5 m, Oct 1977, 26Novl97l,AMJI2547(l);Bondi, trally, often denticulate and finely knobbed Port Jackson. 3 m, 29 Aug 1977, AM AM Jll 167(5): along outer edge, typically 0.14 mm long: end- J 1 0883(2); Manlv. 9 Oct 1979, AM J 1 2735(2); Nam- plates up to 0.25 mm wide, smaller perforations bucca Heads, 13 m, 1 1 Jan 1972. NMV F57555(l#); centrally; irregularly triangular, perforated, Coffs Harbour, 13 m, 22 Jan 1982, AM J 15470(2). curved, denticulate plates, typically 0.1 mm Description. Up to 21 mm long (tentacles partly wide; sometimes rosettes, 0.04 mm long. Ten- withdrawn), 4 mm wide and high; body tacles with flat to bar-like thick ossicles, 234 P. MARK O'LOUGHLIN AND TIMOTHY D. O'HARA

Tentacles with elongate, irregular, often widened centrally, per- rods or plates present posteriorly. perforated rods; pedicels without endplates. forated centrally and distally, sometimes thin Calcareous ring simple, without posterior pro- curved, often bent at centre, up to 0.26 mm long; longations, 10 plates, tapered anteriorly, undu- irregularly round to triangular, thin, perforated, lating posterior edge. Dorsal marsupia present. slightly convex, denticulate plates, up to 0.08 mm wide; rosettes, typically 0.04 mm Type species. Neocnus incubans Cherbonnier, long. 1972. Live colour. Body predominantly white, golden Remarks. This genus was previously known only in largest specimen, sometimes dark decking from the type species found on the Mediter- dorsally and laterally; dark vitreous spots dor- ranean coast off Tunisia. sally and laterally; tentacle branches pale yellow Two superficially similar genera, Pseudopso- to golden, with brown to black marking on ten- /w.sLudwig, 1898 and MicroclwerusGutt, 1990. tacle trunks extending to varying degrees onto also have simple ossicles, a sole, and pedicels introvert. Gold, yellow colours lost rapidly in predominantly restricted to the ventral radii. alcohol. However, Pseitdopsolus also has ten equal ten- vesicles, numerous, Reproduction. Some sac-like gonad caeca and tacles, four polian rare or plates in the body gonoducts have been observed in the right knobbed or smooth perforated dorsolateral coelom. Most of the many speci- wall, and is hermaphroditic. Microchoerus has mens are juveniles, and have been collected knobbed perforated plates, which reduce in during summer and autumn. density as body size increases, and lacks dorsal marsupia. Etymology: Named in recognition of the contri- Orbithyone H.L. Clark. 1938, known only bution by Dr F.W.E. Rowe to this paper and to from the type species O. megapodia H.L. Clark, research in Australia. 1938. found off Western Australia, is also characterized by an almost total lack of ossicles. Distribution. Rottnest I., Western Australia, to it differs in having distinctive cal- Coffs Harbour, New South Wales, including However, a Tasmania. 0-28 m. careous ring, numerous rosettes as well as rods in the tentacles, no sole, and numerous large pedi- parasites are Remarks. One, rarely two, copepod cels with well-formed endplates covering the sometimes present in the coelom of C. rowei whole body. specimens from Victoria, New South Wales and for Tasmania (see # in Material examined exact Neocnus bimarsupiis sp. nov. locations). Dr G.C.B. Poore (Museum of Vic-

toria, pers. coram.) has provisionally identified Plates lc, 2h, Figure 3 them as Cueumaricola sp. in the family Cucu- Neocnus sp. -- Rowe and Vail. 1982: 224. — maricolidae. They are long and narrow, some- tn.oughlin. 1991: 223-225, figs 2, 3. times extending more than half the length ofthe coelom, and carrying numbers of yellow eggs. Material examined. Holotype. Victoria, Apollo Bay, Rosette ossicles are present in the body wall of Marengo, Hayley Point, just below low tide level, 29 Dee 1974. some material from New South Wales. These M. O'Loughlin. NMV F54238. Paralvpes. Type locality, NMV F54239(120): AM specimens are otherwise similar to those from .122752(20). other regions, and to specimens from New South Other material (partial list; all collected as algal Wales without rosettes, and are thus included in epifauna, 0-2 m, unless otherwise stated). the current species. South Australia, Robe. 10 Jan 1988, NMV Most of the specimens have been collected as F54 183(40). algal epifauna and arc juveniles. A few adults Victoria, ("ape Bridgevvater, 20 .Ian 1979, NMV have been found attached to the undersurface of F53806(l): Cape Otway. Crayfish Bay, 31 Dec 1980. rocks. NMV F538 14(47); Apollo Bay. Marengo, Havlev Point, 11 Jan 1980. NMV F53808(75); Flinders, cast Neocnus Cherbonnier, 1972 of Mushroom Reef, 13 Jul 1990, NMV F58630(7);

Shorcham. 1 8 Oct 1980, NMV F538 13(4): Phillip]., E Diagnosis (emended from Cherbonnier, 1972). of Pyramid Rock. 22 Dec 1980, NMV F53810(l): Kil- Body short and stout with distinct sole; 10 den- cunda, 26 Jan 1987, NMV F53805(13); Manners dritic tentacles, ventral 2 smaller; pedicels con- Haven. 300 m oll'shore, 4.5-6 m, 6 Mar 1 982. CPA stn fined to periphery and mid-ventral radius of 15, NMV F53781(l); Cape Paterson. 29 Jan 1988. sole. Body wall ossicles rare, a few perforated NMV F54I82(8). NEW HOLOTHURIANS FROM SOUTHERN AUSTRALIA 235

Figure 3. Neocnus bimarsupiis sp. nov.: a, dorsal body view, showing paired anterior marsupia; b, ventral body view, showing extensible ventral tentacles; c, diagram of a marsupium, showing interior dorsal pedicels, with 4 extending through opening, scale bar = 0.5 mm; d, transverse section of body, scale bar = 1.0 mm; e, adjacent radial and interradial plates from the calcareous ring; f, tentacle rod ossicles, scale bar = 0.01 mm. 236 P. MARK O'LOUGHLIN AND TIMOTHY D. O'HARA

Tasmania, Lulworth, Black Rock Point, 22 Nov pouches sit radially within body wall, separated 1982, NMVF5378705). from coelom, exterior to dorsolateral radial muscle band; each pouch with central aperture; Description. Up to 6 mm long, 3 mm high, 2.5 series of up to 1 6 very small pedicels within each mm wide; body roughly cgg-shapcd; tentacles pouch, may extend out of the pouch aperture. orientated anteriorly or slightly upturned, anus Each brood pouch with up to 14 embryos or juv- posterodorsal; 2 dorsal brood pouches in eniles, uniform in size, differentiation evident at anterior body wall except in juveniles, each with 0.7 mm long; brood juveniles up to 1.5 mm external round aperture, and small interior pedi- long. cels which may extend through the aperture; body wall thin, soft; 10 dendritic tentacles, 2 Etymology. From the Latin bini (two) and mar- ventral ones small, slightly lobed, extensible to supium (pouch, ablative case), with reference to 3.0 mm long, with a pedicel-like attaching capa- the two brood pouches. bility; distinct sole; introvert not distinct from Distribution. Robe, South Australia, to Cape body wall. Sole with peripheral row of up to 20 Paterson, Victoria, and north-eastern Tas- large pedicels, up to 4 anterior ones aligned mania. 0-6 m. transversely; up to 4 irregularly on midventral radius; not extending to introvert anus; r and Remarks. This species is similar to A . ineubans. mostly no pedicels on lateral or dorsal surfaces, Both are small and epiphytic, brood their young except some within the dorsal marsupia and fre- in marsupia in the anterior dorsal body wall, and quently single small radial pedicels anteriorly, have tentacle rods as the main ossicle form. N. most prominent dorsolatcrally; up to 5 small ineubans differs in having small irregular perfor- pedicels around anus. Calcareous ring lacking ated plates as well as rods in the tentacles, rods of posterior prolongations; 10 plates with similar a different shape, rods and small plates in some anterior tapered projections, posterior scallops; posterior pedicels, rare large smooth perforated radials with small anterior notch. Madreporite plates in the body wall, and only one large mar- dorsal; single polian vesicle left lateral; intestine supium which contains up to 30 eggs or embryos ventral, with full loop extending coclom length, and juveniles. single additional short dorsal loop near anus. In A', bimarsupiis the method of transfer of Body wall mostly lacking ossicles, a few rods eggs or embryos from the gonad to the marsupia in dorsal, anal body wall; sole lacking ossicles. is not known. With the extensible attaching ven- Pedicels lacking endplates or ossicles. Tentacles tral tentacles, and the small suckered pedicels with abundant thin straight and curved rods, extending through the marsupium opening, typically 0.08 mm long; ends of rods swollen, there is a capacity for transfer of eggs or embryos with 1-6 small holes or notches, divided end of from gonopore to marsupium. Numbers of eggs rod often twisted together; rods rarely with side in a mature gonad sac, and of brood juveniles in or forks. branches Ossicles similar in juveniles. a marsupium, are similar. If there is a single action of transfer from Live colour. Body dark to very dark grey or one mature gonad sac to one marsupium, the action brown; sole, pedicels, tentacles to varying would be remarkably efficient. Since no brood juveniles degrees lighter in colour; colour persists in with fully developed tentacles alcohol. and mouth have been found in the marsupia, it appears that brood juveniles Reproduction. All observed gonads contain eggs complete their development to a stage of inde- and are similar in form, suggesting hermaphro- pendent nutrition outside the marsupium. ditic or pathenogenic reproduction. All adults N. bimarsupiis is found on algae, particularly (July to April) with similar mature gonads and brown algae, e.g. Zonaria angustata (Kuetz), in brood juveniles; gonad dorsal, with up to 8 sac- the rocky shallows, where it appears as a very like caeca, graded in size from bud to fully- small, dark, soft but tough, oval protuberance on developed; largest lie caeca against wall of brood the algal fronds. Its relatively large ventral pedi-

pouch, each caecum with to 1 3 large up yellow cels enable it to cling strongly to its substrate. eggs or embryos and a few small white eggs inter- spersed; large eggs or embryos frequently 0.6 Squamocnus gen. nov.

long, to 1 .2 long; small mm up mm caeca with up Diagnosis. 10 dendritic tentacles, ventral 2 white eggs. to 15 small Short anterior dorsal smaller; 5 oral valves; pedicels on ventral radii gonopore between bases gonoduct; of dorsal ten- and scattered on dorsal radii and interradii. pair. separate anterior tacle Two dorsal brood Body wall ossicles cups, knobbed perforated but- NEW HOLOTHURIANS FROM SOUTHERN AUSTRALIA 237 tons and large multi-layered ossicles; pedicels O. syracusanus (Grube, 1840) from Europe with perforated bar-like ossicles and endplates; has pedicels distributed on both radii and inter- tentacles with perforated bar-like ossicles, small radii, reduced ventral tentacles, reduced cups, mesh-like plates and rosettes. Calcareous ring knobbed buttons and multi-layered ossicles with simple, with undulating posterior margin and a pronounced denticulate end. It has been anterior tapers. recently referred to Pseudocnella by Thandar, type species. Squamociuts aureoruber sp. nov. 1987, but the other species in that genus have 10 equal tentacles and pedicels restricted to the Etymology. From the Latin squama (scale) in radii. reference to the multi-layered ossicles in the O. kerguelensis (Thcel, 1 886) with scattered body wall, with Ocnus (masculine). dorsal pedicels, reduced ventral tentacles, multi- Remarks. Several existing cucumariid genera layered ossicles, knobbed buttons and x-shaped have similar ossicles. Leptopentacta H.L. Clark, ossicles, is possibly related. Trachythyone amo- 1938 differs in having short posterior "tails" on kurae (Mortensen, 1925), from New Zealand, the radial plates in the calcareous ring and pedi- has cups, smooth and knobbed buttons and large cels restricted to the radii. Loiseitea Rowe and multi-layered ossicles in the dorsal body wall. Pawson, 1985 has deep complex cups, elongate Some of these multi-layered ossicles are calcareous ring plates and (at least on the type extended at one end into a "spine". Pedicels are species) large papillae on the radii. Trachy- biserially arranged on the radii with a few scat- thyone Studer, 1 876 has a similar body form, but tered on the mid-dorsal intcrradius. has smooth single-layered plates as the main oss- O.farquhan (Mortensen, 1925) and O. saccu- icle form. Ocnus Forbes, 1841 has similar lus Pawson, 1983, also from New Zealand, have knobbed buttons and cups, but lacks multi- similar ossicles to Squamocnus, but lack any layered plates and the pedicels are restricted to pedicels on the dorsal surface and probably the radii. deserve their own genus as Pawson (1983) Several little-known forms, previously suggests. referred to Ocnus or Trachythyone. are possibly We have not had the opportunity to examine better placed in Squamocnus. O. brevidentis these species so final placement cannot be deter- (Hutton, 1872), from New Zealand, is possibly mined with certainty. congeneric. Unfortunately, there has been con-

siderable variation in the descriptions of this Squamocnus aureoruber sp. nov. species. Dendy (1896) and Panning (1949) found multi-layered ossicles, knobbed buttons, Plates Id, 4a-h, Figure 4 cups and scattered dorsal pedicels in their speci- Trachythyone sp. — Rowe and Vail, mens. Mortensen (1925) described "large, 1982: 222 (part). — 6'Loughlin, 1991: 223, fig. I. smooth plates"; Pawson (1970) made no men- tion of large plates at all. Possibly more than one Material examined. Holotype. Victoria, Cape Paterson, just species is involved. below low tide level, 18 Jan 1980. M.

Figure 4. Squamocnus aureoruber gen. et sp. nov.: a, transverse section of body, scale bar = 1.0 mm; b, adjacent radial and interradial plates from the calcareous ring; c, top view of a dorsal body wall cup, scale bar = 0.01 mm. 238 P. MARK O'LOUGHLIN AND TIMOTHY D. O'HARA

interradials; 5 interradials O'Loughlin, T. O'Hara and J. Stephenson. NMV slightly higher than F54244. with posterior scallop, pointed anteriorly. Paratypes. Type locality and date, NMV Madreporite dorsal; single polian vesicle left lat- F54243(69); NMV F58622( 1*); Apollo Bay. Marengo. eral; intestine runs dorsally from mouth to mid-

1 .Ian 1980. Haylcy Point, just below low tide level. 1 coelom, loops back to ring, then runs ventrally to NMVF54242(11). anus. Other material (partial list: all found as algal Dorsal body wall ossicles of 5 types. 1) A sur- cpifauna, 0-2 m. unless otherwise stated: * indicates face layer of abundant small spinous cups, 2-3 tissiparous specimens). times as wide as deep, typically 0.04 mm long; South Australia. Streaky Bav, Point Westall. 1 5 Jan 1991. NMV F592I0(2): Robe. 10 Jan 1988. NMV typically rectangular, with 4 large and 4 small F54703(4): NMV F58623(2*); 9 Jan 1990. NMV corner holes; rim and cross thin, with short F57552(4*); Port MacDonnell. 8 Jan 1988. NMV spines pointing mainly vertically out from cup. F54705(13). 2) Some shallow cups, slightly irregular, fre- 31 Dec 1980. Victoria, Cape Otway. Crayfish Bay. quently 8-holed, only slightly spinous or lacking NMV F53801O5): Marengo. 2 Jan 1990, NMV spines. 0.04-0.09 mm long. 3) Large, imbricat- F57553(6*); 10 km NE of Apollo Bav. "'Mullet Holes". ing, knobbed, regularly perforated, often multi- 2 Jan 1988. NMV F54702(8): NMV F586260*): ossicles, predominantly 2-layered, very Lome, Grey Point. 14 Jun 1982. NMV F53798(l); layered Bushrangers Bay. 28 Mar 1981. NMV F53804(l): irregularly oval, typically 0.3 mm long, up to Flinders, E of Mushroom Reef 6 Mar 1982. NMV 0.5 mm long. Knobs on developing plates join to F53799(7); 13 Jul 1990, NMV F5863l(l); 11 Aug create bars and secondary layers. 4) Knobbed 1990, NMV F58638O0*); NMV F58635(l); Shore- and smooth buttons, frequently 4-holed. many F53796(3); ham, 18 Oct 1980, NMV NMV fairly regular, typically 0.09 mm long. 5) F58625(l*); Phillip I., Cowrie Beach. 27 Nov 1985. Smooth, slightly convex, irregularly oval, perfor-

NMV F53802(l 1 ): NMV F58628(l*): Hamicrs ated plates: holes large centrally, small peripher- Haven, 6 Mar 1982, CPA stn 23. NMV F53777(l); ally: typically 0.1 wide. Ventral body wall Cape Paterson. 14Feb 1981. NMVF53803(18): NMV mm F58624(l*); 28 Apr 1989. NMV F54393(24); NMV with cups, buttons, convex plates, large plates F586270*); Inverloch, Eagles Nest. 4.5-7.6 m. 5 Mar with secondary developments but not multi- 1982. CPA stn 3. NMV F53779(2). layered. Pedicels with endplates. 0.18 mm wide: Tasmania, Rocky Cape, 2 m, 15 Mar 1988. NMV Hat and elongate to bar-like ossicles, frequently F57365(l): Greens Beach, mouth of Tamar River. 7 bent and curved and widened centrally and dis- Mar 1981, F53790(2): Lulworth, Black Rock NMV tally. perforated, irregular, up to 0.18 mm long. Point, 22 Nov 1982. NMV F5379K2 juv); Bicheno, 7 Tentacles with irregular, large to small, bar-like m. 22 Mar 1988, NMV F59229(l). ossicles, some curved and bent and branched, Description. Up to 10 mm long (tentacles with- perforated, up to 0.2 mm long; some irregular, drawn), 1.7 mm high. 3 mm wide; body dome- convex, mesh-like, denticulate plates, 0.08 mm like with slight dorsolateral angles in transverse wide: rosettes, up to 0.05 mm long. section, flattened ventrally. rounded pos- teriorly, with extensible upturned anal cone: 10 Live colour. Body, including ventral surface and dendritic tentacles orientated anteriorly, 2 ven- pedicel walls, reddish orange often with slight tral ones small; body wall thin, microscopically dark flecking; ends of pedicels pale; tentacles scaly; distinct flat, very thin-walled ventrum, yellow, with dark brown on trunks and introvert; not sole; very thin-walled introvert, lacking all colour except dark markings lost rapidly in pedicels; 5 small radial oral valves which project alcohol. partly over introvert; anus with 5 fine teeth. Ventral radii each with 2 irregular rows of pedi- Reproduction. All observed gonads contain eggs cels, up to 40 in a row; pedicels large on mid- and are similar in form, suggesting hermaphro- ventral and ventral rows of lateroventral radii; ditic or pathenogenic reproduction. The mature distinctly smaller on dorsal row of lateroventral gonads have eggs or embrvos in January (NMV radii, near anus and introvert; not extending F58626), February (NMV F53803).' March onto introvert. Pedicels scattered dorsally, lat- (NMV F53799), July (NMV F58631) and erally, in irregular double rows on dorsal radii; August (NMV F58635). Gonad dorsal, up to 8 single very extensible small pedicels on each caeca graded in size, largest caeca with up to 15 anterior radial projection of body wall; 5 small yellow eggs or embryos interspersed with fewer extensible pedicels around anus. Calcareous ring small white eggs: eggs or embryos up to 0.7 mm lacking posterior prolongations: 5 radials with long; short anterior dorsal gonoduct: gonopore posterior notch, anterior taper and notch. between bases of dorsal tentacles. NEW HOLOTHURIANS FROM SOUTHERN AUSTRALIA 239

Some specimens show evidence of fissiparity. Trachythyone Studer, 1876 Fission commences with a transverse growth Diagnosis (based on species currently assigned change and constriction around the mid-body to the genus). 10 dendritic tentacles, ventral 2 (NMV F54243). The constriction deepens and usually smaller; pedicels on radii, sometimes begins to separate the anterior and posterior also scattered on dorsal and lateral interradii. halves (NMV F58623, F58624). The two ends Body wall ossicles cups overlying numerous finally separate by moving apart, as evidenced large perforated plates; plates usually smooth, by the long connecting thread of body wall rarely with some minor lumps, up to 0.95 mm remaining in one specimen (NMV F58622). long; often some knobbed buttons; cups small, Matching and recently separated anterior and shallow, sometimes spinous, sometimes posterior ends have been found together (NMV replaced by rudimentary x-shaped ossicles. Ten- F57553). Most large collections have one or a tacles and' pedicels with perforated rods or few with atypical round or short body form, and plates; endplates present. Calcareous ring lack a calcareous ring and tentacles in some cases simple, without posterior prolongations, 10 (NMV F57553) or the characteristic form of the plates, tapered anteriorly, posterior notches. posterior end in other cases (NMV F58623). One specimen is an posterior end with ring and Type species. Trachythyone muricata Studer, tentacles at a very earlv development stage 1876. (NMV F58625). Remarks. The genus Trachythyone was revived 1949 who assigned 21 species. He Etymology. From the Latin aureus (golden) and by Panning, subsequently (1964) severely restricted the ruber (red), in reference to the colour. genus, referring (1966) many of the original species to Leptopentacta H.L. Clark, 1938. Pan- Distribution. Streaky Bay. South Australia, to ning ( 1 964) suggested that one of the remaining Inverloch, Victoria, including northern Tas- crucifera (Semper, 1869), from the mania. 0-8 m. species, T. Indian Ocean, with its spiny cruciform plates, probably requires its own genus. T. amokurae Remarks. S. aureoruber can be distinguished 1925) from New Zealand, with its from the similar species listed above in the gen- (Mortensen, multi-layered ossicles, is not a Trachythyone (see eric remarks by the body wall ossicles, a combi- remarks). nation of well-developed cups, rare knobbed Squamocnus predominantly a Southern buttons and oval multi-layered ossicles. What is left is genus with about ten species. Several Evidence of rissipary in S. aureoruber has Ocean need revision. For example, T. parva (Ludwig, been found throughout its geographical range, in 1874) was originally described as having x- small and large specimens, and in summer and shaped ossicles in the external skin layer. How- winter. Direct observation of fission has not yet subsequent specimens with cups or flat been undertaken. Preserved material provides ever, "bowls" have also been referred to this species strong evidence that fissipary is a significant 1 by Panning ( 1 949, 1 964), Hernandez ( 982) and recruitment process. Mature gonads with eggs others. Various authors have also attempted to have been found in several specimens, indi- synonymize Oenus kerguelensis (Theel, 1886), cating that recruitment is both sexual and which has multi-layered ossicles, with T. parva. asexual. Evisceration has not been observed, an Clearly more than one species is involved. The and this phenomenon would not provide ring species from Kerguelen, T. muricata, T. squam- explanation for the absence of a calcareous speci- ata (Ludwig, 1898), T parva, O. kerguelensis and tentacles in some atypically short and O. ekmani (Ludwig and Heding, 1935), need mens. separation. Emson and Wilkie (1980: 161) noted that six clear The generic diagnosis above has been broad- holothurian species have been found to repro- species ened from that of Panning (1964) to include duce by fission, including two Oenus with lumpy plates or with knobbed but- In Oenus planet species from the Cucumariidae. on twist- tons. Several species have some minor lumps (Brandt. 1835) division usually occurs by pedi- the plates (e.g. T. parva sensu Panning, T. mac- stretching, muscles contract, and the ing or Pawson, 1962a, T bouvetensis (Lud- until the tissues phersonae cels pull in different directions wig and Heding, 1935)). However, the plates are are severed. never consistently knobbed to the extent of Specimens have been found predominantly Oenus or Pseudocnus. Several species (e.g. T. bob on algae in the rocky shallows. 240 P. MARK O'LOUGHLIN AND TIMOTHY D. O'HARA tonst (Mortensen, 1925)) have knobbed buttons 1982, NMV F54706(l); Harmers Haven, 18 May as a secondary ossicle type. As the primary oss- 1983, NMV F537840): Cape Paterson. 29 Mar 1982, NMVF54707(l);25Jan 1992, NMV F59228(4); Mal- icle type in these cases is smooth plates it seems lacoota, buried in sill, rocky shallows, 21 Jan 1981, unnecessary to exclude such species from NMV F5470K1). Trachythyone. Tasmania, Bicheno, 7 m, 22 Mar 1988, NMV The form of the ossicles is the main character F592270): Eaglchawk Neck, 15 Feb 1991. NMV distinguishing the species. '/'. iechleri (Lampert, F592I 5(3); Tinderbox, below low tide level, 29 May 1885) stands a little apart from the others, 1974, AM J 10863(1 and 5 slides); Southport, Sisters having large plates with only very few tiny per- Bay, 8 May 1982. NMV F53792(l). forations and some incipiently multi-layered Description. Up to 8 mm long (tentacles partly ossicles near the anus. Cups vary from predomi- extended), 3 mm wide; body predominantly nantly x-shaped ossicles (e.g. T. parva, 7. bol- round in transverse section to slightly pentag- '/'. lonsi, peruana (Semper, 1 868)), to well- onal and keeled; body about twice as long as developed cups with delicate cross and spinous wide; mouth orientated anteriorly; extensible rim (e.g. 7. squamata, T. nmricata). to shallow anal cone upturned; distinct modified ventrum, "bowl"-like cups with smaller perforations (e.g. not a sole, rounded not flat; introvert distinct, 7. parva sensu Panning, 7. macphersonae, T. thin-walled; 10 dendritic tentacles, 2 ventral mira (Ludwig and Heding, 1935), T. nina ones small; body wall fairly thick, 0.4 mm, not (Deichmann, 1930)), to small irregular cups (e.g. rigid, microscopically scaly. Lacking oral valves, 7. crassipeda CherbonnieTj 1961, T. bouvetensis, about 6 small anal teeth. Large pedicels on 7. Iechleri). ventral radii, extend up to introvert and anus, The genus Psolicliella Mortensen, 1925 has distinctly smaller near introvert and anus; mid- plates, rudimentary cups and scattered dorsal ventral scries an irregular zig-zag; ventrolateral pedicels. It can be distinguished from Trachy- radii with 2 rows, a ventral complete row and an thyonc by the presence of a sole and the low outer lateral incomplete row; up to 35 pedicels incidence of ossicles in the body wall. The mono- on each radius. Small scattered pedicels dorsally typic genus Neopsolidium Pawson, 1964, with and laterally, slightly concentrated on dorso- type N. convergens (Herouard, 1901), is possibly lateral radii into irregular zig-zag series; intro- a synonym of Trachythyone or Psolicliella. The vert lacking pedicels; 5 anterior thin extensible main ossicle type is smooth, or lumpy, perfor- pedicles; about 6 extensible anal pedicels. Cal- ated plates, 0.2-0.4 mm long, overlain by small careous ring lacking posterior prolongations; 5 complete cups. The genus is currently placed in radials with posterior notch, anterior taper and the Psolidac, but the form of the ossicles and the notch; 5 interradials with posterior scallop, lack of a true sole indicates that the species is pointed anteriorly. Single polian vesicle, left better placed in the Cucumariidae (Pawson, lateral. Dorsal madrcporite. pers. comm.).

Dorsal body wall ossicles of 3 types: 1 ) Numer- Trachythyone Candida sp. nov. ous large, thick, smooth plates, irregularly oval to elongate, up to 0.5 mm long, up to about 40 Plates lc, 5a-f, Figure 5a-c uniformly distributed perforations, some den- Psoliclium convergens, — Joshua, 1914: 6 [non ticulate at one end with pointed spines, smaller Neopsolidium convergent (Herouard, 1901)]. perforations at opposite rounded end. 2) Some Psoliclium sp. — H.L. Clark, 1946: 415. knobbed buttons, fairly regular, typically 4 Trachythyone sp. — Rowe, 1982: 466, fig, 10.31c. holes, up to 8, typically 0. 1 mm long. 3) Shallow — Rowe and Vail, 1982: 222 (part). cups, 4 times as wide as deep; typically 0.045 Material examined. Holotype. Victoria, Flinders, Jan mm long; more commonly oval than rectangu- 1912, coll. E.C. Joshua, NMV F59200. lar; typically 4, up to 8, perforations; rim and

Paratypcs. Type locality and date. F5920l( I NMV ); cross thin with numerous peg-like spines point- MCZ 1151 (slide of whole body mount); Phillip I., ing in all directions; cups form a surface layer in Kitty Miller Bay. rocky shallows, just below low tide body wall. Ventral body wall with cups and level, 8 Jan 1986, NMV F54229(i). elongate, narrow, slightly lumpy Other material (all found in the rocky shallows, 0-2 and knobbed, m. unless otherwise stated). perforated plates, typically 0.25 mm long. Pedi- cels with endplates, Victoria, CapcOtway, Moonlight Beach, 1 1-12 Mar up to 0.23 mm wide, and 1975, NMV F57947(2); Deep Sea Cove, 20 Dec 1983. abundant, oval to elongate, curved, denticulate, NMV F57945(I); Pebble Point, 18 Feb 1984, NMV often slightly lumpy, perforated plates, typically Flinders, E of F5795QO); Mushroom Reef, 6 Mar 0.2 mm long. Tentacles with irregular, curved NEW HOLOTHURIANS FROM SOUTHERN AUSTRALIA 241 a

Figure 5. a-c, Trachythyoite Candida sp. nov.: a, transverse section of body, scale bar= 1.0 mm; b, adjacent radial and intcrradial plates from the calcareous ring; c, top view of a dorsal body wall cup, scale bar = 0.01 mm, d-f, Trachythyone glebosa sp. nov.: d, transverse section of body, scale bar - 1.0 mm; c, adjacent radial and

intcrradial plates from the calcareous ring; f, dorsal body wall cups, scale bar = 0.01 mm.

and often bent, often widened centrally, perfor- Distribution. Western side of Cape Otway to ated, bar-like ossicles, up to 0.25 mm long; and Mallacoota, Victoria, and eastern Tasmania. 0- small, irregular, slightly convex, denticulate, 7 m. closely perforated plates, typically 0.06 mm wide; lacking rosettes. Remarks. Joshua's (1914) specimens, which he Live colour. Body uniform white or pale cream; identified as Rsolidium convergens, have been crystalline; no dark markings; tentacles trans- re-examined by us and form the basis of this new '/'. lucent, may be a few brown markings on species, Candida. The specimens are still in trunks. excellent condition and, as the largest available, have been made the types. They do not belong to Reproduction. Sexes separate. Male or female Psolidium Ludwig, 1 886 as they do not have a (from December to gonad in most specimens true sole and do possess the medium-sized, non- May). imbricating, smooth plates in the body wall typi- This species is similar Etymology. From the Latin Candida (glittering cal of Trachylhyone. to convergens which, as indicated white), in reference to the colour. Neopsolidium 1

242 P. MARK O'LOUGHLIN AND TIMOTHY D. O'HARA

white to yellow; above in the generic remarks, should probably Colour (preserved). Body colour white; tentacle be referred to Trachythyone or PsolidieUa. lumps on body wall and pedicels flecking on all inter- T. Candida can be distinguished from N. con- trunks dark. Dark brown vergent and all other Trachythyone species by radii of the W.A. specimen. form of the cups, which typically have a com- the Reproduction. Immature gonad tubules present plete rim and spines pointing in all directions on in the holotype. both cross and rim. Specimens have usually been found in silt Etymology. From the Latin glebosa (lumpy), in and under rocks in the shallows. reference to the white lumps over the body pedicels. Trachythyone glebosa sp. nov. Distribution. Michaelmas I., Western Australia Plates lg, 5g-j, Figure 5d-f to Kangaroo 1., South Australia. 3-15 m. Holotype. South Australia, Material examined. Remarks. Within Trachythyone, T glebosa is Kangaroo Vi vonne Bay, N side of Point Ellen, rock L, distinguished by the massing of body wall oss- and sand, 3-8 m, W. Zeidler and K.L. Gowlett- icles into visible white lumps, the elongate shape Holmes, 26 Jan 1989, SAM K1812. of the plates, and the very small irregular Paratype. Western Australia, W end of Michaelmas smooth

I., 35°02.9'S, 118°01.4'E, 15 m. 17 Apr 1986, NMV cups. F59226(l). Apsolidium gen. nov. Description. Up to 12 mm long (tentacles Diagnosis. Body stout with distinct sole and extended), 3 mm wide and high. Body wall thin; upturned anal and oral ends; 10 dendritic ten- body and pedicels with white lumps of massed tacles, ventral 2 smaller; pedicels on periphery ossicles, fewer ventrally; body rounded to and mid-ventral radius of sole, small ones scat- slightly pentagonal in transverse section; dis- tered dorsally and laterally. Body wall ossicles tinct modified ventrum; mouth orientated knobbed cups, overlying numerous perforated, anteriorly, anus posteriodorsally; 10 dendritic non-imbricating multi-layered ossicles, up to 1 . tentacles, 2 ventral ones distinctly smaller; dis- mm long, and lumpy or smooth button-like oss- tinct introvert, lacking pedicels; 5 small scale- icles. Tentacles with perforated plates, bar-like like anal teeth; lacking oral valves. Pedicels perforated ossicles; pedicels with perforated large; irregular paired rows to zig-zag series on 5 plates, endplates. Calcareous ring simple, with- radii; small scattered interradial pedicels dor- out posterior prolongations, 10 plates, with sally, laterally, lacking on ventral interradii. tapered anterior projections and posterior Pedicels small close to introvert and anus; 5 scallops. small ones around anus. Calcareous ring lacking posterior prolongations; 10 plates with tapered Type species. Apsolidium handrecki sp. nov. anterior projections, radially notched, posterior Etymology. From the Latin a (from), in refer- scallops. Single left lateroventral polian vesicle. ence to the differences from Psolidium (neuter). Dorsal madreporite. Dorsal body wall ossicles of 2 types. 1) Abun- Remarks. Apsolidium is similar to the genus dant large narrow plates, thick, smooth, perfor- Psolidium Ludwig. 1886 from the family Psoli- ated, rounded tapered ends, up to 0.6 mm long. dae. The psolids are characterized by the pres- 2) Cups small, irregular in shape, many with cen- ence of a sharply defined sole on the ventral tral cross and 4 holes, rim lumpy but lacking surface and an imbricating armour of large distinct knobs or spines, depth variable, most multi-layered ossicles, or scales, covering the about 0.024 mm long, some larger rectangular dorsal surface. In Psolidium the dorsal scales are ones about 0.04 mm long. Ventral body wall perforated by the scattered dorsal pedicels. The with cups, large narrow perforated plates up to three species of Apsolidium on the other hand 0.4 mm long. Pedicels with endplates, cups, have smaller non-imbricating multi-layered oss- curved narrow irregular perforated plates up to icles, similar to other cucumariid species such as 0.25 mm long. Tentacles with many small to the Cucuvitrum and Squamocnus species large, bent, perforated bar-like ossicles, up to described in this paper. 0.22 mm long; thin plates, irregularly oval, con- Several other cucumariid genera also have vex, denticulate, perforated, typically 0.056 mm well-defined soles. Of these, Pseudopsolus Lud- wide; abundant rosettes, typically 0.04 mm wig, 1898, Neocnus Cherbonnier, 1972 and long. Microchoerus Gutt, 1990 have reduced ossicles NEW HOLOTHURIANS FROM SOUTHERN AUSTRALIA 243 and virtually no dorsal pedicels. Psolidiella Mor- irregular, thick, smooth to knobbed, some bent, tenscn. 1 925 has scattered small plates and rudi- up to 15 holes, up to 0.2 mm long; button-like mentary x-shapcd cups. Neopsolidium Pawson, ossicles, fairly regular, typically 4-holed, typi-

1964, here also regarded as a cucumariid (see cally 0. 1 mm long; lacking multi-layered ossicles Trachythyone remarks), differs from Apsolidium and plates with secondary developments. Pedi- in having single-layered, often smooth, dorsal cels with endplates, 0.24 mm wide; cups; irregu- plates. lar narrow perforated plates, some finely knobbed, bent, curved, convex, up to 0.14 mm Apsolidium handrecki sp. nov. long. Tentacles with abundant rosettes, up to Plates 2a. 6a-i, Figure 6a-d 0.08 mm long; bar-like ossicles, small to large, straight or bent, perforated, some widened cen- Material examined. Holotype. Victoria, Merricks. trally, up to 0.27 long; fine mesh-like plates, rocky shallows. 27 May 1989. M. O'Loughlin. T mm rectangular, convex, denticu- O'Hara and C. Materia, NMV F54391. irregularly oval to Paratvpes. Tvpe locality and date, NMV F54392(l late, up to 0.08 mm long.

and 1 juv): 28 Jan 1983. NMV F53765(l); 29 Oct Live colour. Body grey, created by dense brown- 1980. NMV F53763(2). black flecking dorsally; sole pale grey with sparse Other material. South Australia. Evre Peninsula. flecking interradially; rim of sole cream; anal Arno Bav. jettv piles. 2-3 m, 22 Feb 1988, SAM K1815(2). cone dark at apex; tentacle trunks, introvert, oral yellow. Western Australia. Trigg 1.. off Caulerpa, 22 Nov disc dark brown; tentacle branches pale 1969. NMV F59225(2). Reproduction. Sexes separate. Holotype and Description. Up to 20 mm long (tentacles larger paratype (NMV F54392) with mature extended), 7 mm wide. 6 mm high; body stout female gonads; paratype (NMV F53765) with with oral extension and upturned anal cone; mature male gonad: juvenile (NMV F54392, 5 body wall thin, densely crystalline; 10 dendritic- mm long) with gonad development beginning. tentacles, 2 ventral ones distinctly smaller; dis- Gonoduct attached to dorsal body wall; gono- tinct oval sole, wider than body, margin pore between bases of dorsal tentacles. upturned; distinct narrow introvert, lacking Etymology. Named in recognition of the contri- pedicels; anus with 5 teeth. Up to 5 rows of pedi- bution by Mr Clarrie Handreck to the fieldwork cels on periphery of sole, none extending beyond of the Marine Research Group of Victoria and to sole to introvert or anus, 2 rows on midventral the collection of this species. radius, ventral interradii lacking pedicels; close cover of small pedicels dorsally, laterally, on oral Distribution. Merricks, Victoria, to Trigg I., SW extension and anal cone, used for attaching grit; Western Australia. 0-3 m. small radial pedicels around very extensible Remarks. At Merricks, Western Port, this mouth and anus. Calcareous ring lacking pos- species is found adhering to the top or side of terior prolongations; 5 radials with anterior pro- rocks in shallow water. jections, sometimes deeply divided, notched posteriorly; 5 interradials with pointed anterior Apsolidium densum sp. nov. projections, posterior scallops. Madreporite Plates 2b, 7a-h, Figure 6e-g dorsal; single polian vesicle left ventral. Dorsal body surface ossicles of 4 types. Material examined. Holotype. Victoria, 10 km NE of non- "Mullet Holes", under rock just below low 1) Numerous multi-layered ossicles, Apollo Bav. 11 Jan 1984, M. O'Loughlin, NMV imbricating, irregularly oval to elongate, up to tide level", F54237. 0.4 mm long, comprising regularly perforated Paratypcs. Tvpe locality and date, AM J227530); thick knobbed plates linked together. 2) Some type locality. 7*Jan 1985, NMV F53764(l); Victoria, perforated plates developing second- knobbed Flinders, Mushroom Reef, rocky shallows. 22 Jan ossicles, lumpy to ary layers. 3) Some button-like 1982. NMV F53766(l). smooth, thick, irregular, up to 8 holes, up to Description. Largest 40 mm long (tentacles with- 0.2 mm long. 4) A surface layer of fine-rimmed drawn), 18 mm wide, 12 mm high (mid-body); deep cups, typically oval, 4 holes, about as deep terminally smaller specimens more elongate; body stout as wide; arms of cross thin, branched, extension and anal cone upturned; knobbed, ends joined to varying degrees to form with oral body wall thick, up to 1.5 mm, densely crystal- thin, finely knobbed, sometimes incomplete cups; line: 10 dendritic tentacles, 2 ventral ones dis- rim; commonly 0.06 mm long. Sole with smaller; distinct oval sole, narrower than abundant elongate plates, frequently narrow. tinctly 244 I'. MARK O'LOlKillUN AND TIMOTHY D. O'HARA

Figure 6. a-d, Apsolidium handrecki gen. et sp. nov.: a, transverse section of body, scale bar- 5.0 mm; b, adjacent radial and interradial plates from the calcareous ring; c, top view of a dorsal body wall cup; d, side view of an incomplete cup, scale bar = 0.01 mm, e-g, Apsolidium dcnsitm sp, nov.: e, transverse section of body, seale bar = 5.0 mm; f, adjacent radial and interradial plates from the calcareous ring; g, top view of a dorsal body wall enp, seale bar - 0.01 mm. sp. nov.: h, transverse h-j, Apsolidium alvei section of body, seale bar = 5.0 mm; i, adjacent radial and inter- radial plates the calcareous from ring; j, side and top views of dorsal body wall cups, scale bar - 0.01 mm. NEW HOLOTHUR1ANS FROM SOUTHERN AUSTRALIA 245

e

b adjacent.radial and body, scale bar = 1 mm; occiduus sp. nov.: a, transverse section of Figure 7 a-d Ocnus cup, scale body wail button; d. top v.ew ot a dorsal body wall mferradial plates from the calcareous nng; c, dorsal

side and top v.ews of dorsal body wall cups, e, dorsal body wall button; f, ^l-tOmSfalcarettS (Dendy, 1 896): scale bar = 0.01 mm.

Knobbed per- periphery ot knobbed plates linked together. 2) body. Up to 6 rows of pedicels on thick plates with extending knobs, con- beyond sole to introvert or forated sole, none extending layers. ventral necting bars, developing secondary 3) anus, up to 4 rows on midventral radius, button-like ossicles, lumpy or smooth, close cover of small Some interradii lacking pedicels; 0.14 fairly regular, often 4-holed, commonly laterally, on oral body exten- pedicels dorsally, smooth, regularly perforated, grit. Cal- mm long. 4) Some sion and anal cone, used for attaching somewhat denticulate plates, 0.18 mm long. 5) lacking posterior prolongations; 5 careous ring oval with holes, rim projections, Knobbed cups, typically 4 radials with notched anterior continuous at corners, about twice as wide interradials, with slightly rarely notched posteriorly; 5 heavily as deep, cross and rim thick, only rim anterior projections, posterior lower pointed commonly 0.08 mm long. One large left ventral. knobbed, scallops. Single polian vesicle knobbed cup, 0.2 mm long. Sole with cups, Dorsal madreporite. with Massed smaller multi-layered ossicles. Pedicels ossicles of 5 types. 1 ) Dorsal body wall bar-like ossicles, endplatcs, 0. 1 8 mm wide; cups; ossicles, irregular, up to 1.1 mm multi-layered bent, sometimes curved, per- perforated thick thick, straight to long, comprising regularly 246 P. MARK O'LOUGHLIN AND TIMOTHY D. OHARA

forated, up to 0.34 mm long; perforated plates, Description. Largest 52 mm long, 33 mm high, irregularly triangular to elongate, marginally 27 mm wide (tentacles slightly extended); body denticulate or slightly knobbed, straight or bent, fat, rounded, slightly convex dorsally, deeply sometimes curved, 0.2 mm long. Tentacles with convex ventrally; short anteriodorsal and pos- abundant bar-like ossicles, large to small, irregu- teriodorsal oral and anal cones; body wall tough, lar, thick, bent, curved, some widened centrally, flexible; 10 dendritic tentacles, 2 ventral ones perforated, up to 0.38 mm long; perforated distinctly smaller; distinct short convex sole-like plates, flat to curved, irregular, up to 0.24 mm modified ventrum; distinct introvert, lacking long; abundant mesh-like plates, irregularly oval pedicels. Lacking oral valves. Ventral radii with to triangular, convex, denticulate, typically 0.08 3 short parallel series of small pedicels, widely mm wide; rosettes up to 0. 12 mm long. separated from introvert and anus, about 5 irregular rows on ventrolateral radii, up to 4 Live colour. Body white; some grey flecking rows on midventral radius; small pedicels scat- dorsally and on oral extension. tered sparsely and evenly over body, not on Reproduction. Sexes separate. NMV F53766 introvert or interradii of ventrum. Calcareous with male gonad; holotype with female gonad ring lacking posterior prolongations; 5 radials and small eggs. with posterior scallop, anterior taper and notch; 5 interradials with posterior scallops, pointed Etymology. From the Latin densum (thick), in anteriorly. Single elongate polian vesicle, left reference to the body wall. lateral.

Distribution. Apollo Bay and Flinders, Victoria. Dorsal body wall ossicles of 3 types. 1) Abun- 0-1 m. dant large plates, thick, smooth, very irregularly oval, closely perforated, often with developing Remarks. A. densum differs from A. handrecki in secondary layers and thickenings, up to 1.1 having a very thick body wall; a sole that is nar- mm long. 2) Button-like ossicles thick, rower than the width of the body; larger ossicles smooth, irregularly quadrilobed to elongate, with in the body-wall, pedicels and tentacles; more up to 8 knobs and other secondary developments on perforations, typically 0.1 mm long. 3) Deep cups, round to oval, rim plates in the sole; larger, shallower, heavily knobbed, cruciform knobbed cups; and white colour. centrepiece broad and smooth, 4-holed, some as deep as long, typically 0.045 long. Despite intensive searching in the rocky shal- mm Ventral body wall with cups; narrow plates, thin to thick, lows, this species has been found at only two sometimes lumpy, open ocean localities: near Apollo Bay and Flin- irregular, up to 14 perfor- ders, Victoria. ations, typically 0.24 mm long: lacking large plates. Pedicels with cndplates, 0.4 mm wide; Apsolidium alvei sp. nov. typical cups; irregular thin, narrow, perforated plates, some curved or bent, some widened cen- Plates If, 8a-f, Figure 6h-j trally, typically 0.24 mm long. Tentacles with Cueumaria squamata. Joshua — and Creed, 1915: small to large, bar-like ossicles, somewhat flat, 17. [non TfUchyxhyone squamata (Ludwig, 1898)). perforated, often bent with outer widening cen- Ciwumaria squamatoides H.L. Clark. 1946: 389. — trally, up to 0.36 mm long; typical cups; rosettes A.M. Clark, 1966: 345. rare. 'Ciwumaria 'squamatoides. — Rowe, 1 982: 466, fig 1 0.3 lb. Colour (preserved). Body white or pink, very Material examined. Holotype. South Australia, Yorke dark orally and anally; tentacle trunks very Peninsula, Marion Bay, washed up on beach, 6 Sep dark. 1978, W Zeidler. SAM K1824. Reproduction. Sexes separate: mature gonads Paratypcs. Tvpe locality and date, SAM K 1825(2). present in September; gonopore at inner base of Other material. South Australia, Yorke Peninsula, dorsal tentacle pair. Marion Bay, in Posidonia bed exposed at low tide, 28 Etymology. From the Latin alveus (belly of a Jan 1979, SAM K 1 82 1(1); Adelaide, between North Haven ship), in reference to the body form. and Largs Jetty, 300 m offshore, seagrass, 1 Dec

1 1 I 980, SAM K 823( ); Glcnelg, 1 1 1 m, 5 Feb 959, in Distribution. Yorke Peninsula to Encounter Bay, Posidonia. AM J22638(l); Port Noarlunga, I960, South Australia. 0-10 m. SAM K 1 822( I ); Encounter Bay, coll. Dr Verco. ident- ified as Ciwumaria squamata by Joshua and Creed, Remarks. A. alvei is referred to Apsolidium, and 1915, SAM K 1369(1). not Traehythyone. on the basis of the form of the NEW HOLOTHURIANS FROM SOUTHERN AUSTRALIA 247

body wall ossicles and the distribution of pedi- often occur in the same . Wc believe that cels. The body wall plates consistently develop it is appropriate to separate species which have secondary layers and the cups are deep with a multi-layered ossicles. Some of these species knobbed rim, similar to A. handrccki. The pedi- have been discussed under Squamocnus gen cels are arranged in up to five rows on the ventral nov. radii, and reduced and scattered dorsally, with- Within Panning's (1971) Ocnus species, O. out any signs of concentration on the laterodor- cakareus (and the following new species) differ sal radii. On the other hand, the secondary somewhat from the European species. O. calcar- developments on the body wall plates are not eus has cups based on a cruciform piece, rather developed to the extent of A, handrccki or A. than the three-rayed base of the cups in O. planci densum. The body form, including the small and O. Iacleus: and also has some additional sole-like ventrum. is also very different from the large Hat plates in the body wall. A future two other Apsolidium species. This correlates revision could separate the two groups into dif- with the known habitat of the three species: A. ferent genera. alvei prefers seagrass beds while .4. handrccki and A. densum live on rocks. The authors are not Ocnus occiduus sp. nov. satisfied that enough evidence currently exists to Plates Ih, 9a-f, Figure 7a-d assign A. alvei its own genus. The specimen, identified by Joshua and Creed (1915) as Cucumaria squamata, has been re- Ocnus cakareus. — Rowe. 1982: 446. tig. IO:32a examined by the authors and referred to this [non Ocnus cakareus (Dendy. 1896)1. species. H.L. Clark (1946) proposed the replace- Material examined. Holotype. South Australia. ment name C squamatoides for this specimen. Ccduna, Cape Vivonne, under rocks, 0-1 m, 14 .Ian However, it was never adequately described by- 1991. M. O'Loughlm. NMVF592I2. Joshua and Creed, nor examined by H.L. Clark. Paratvpes. Type locality and date. NMV

F592 1 3(6). The name squamatoides is therefore a nomen Other material. Western Australia. Roltnesl 1., nudum, failing to satisfy ICZN Article 13 as 1 254- North Point, rocky reef. 30 m. 1 I Jan 99 1 , WAM pointed out by A.M. Clark (1966: 345). 91(1); Cape Naturaliste, Eagle Bav. Apr 1964, WAM has been taken predominantly from A. alvei I 766-7 1( 1 ); Bunker Bav, 29 Jan 1975, WAM 425-78( ); in Gulf St Vincent. seagrass (Posidonia) beds Cowaramup Bay. 21 Oct 1989. WAM 880-89( I ); Flin- ders Peninsula. Salmon Holes, 20 Jan 1988, WAM Ocnus Forbes. 1841 161-88(1). Diagnosis (based on Panning, 1971). Body South Australia, Nuyts Archipelago, St Francis 1.. 24 KI8I 1(9); Franklin Is, 24 Feb 1983, cucumber-shaped, slightly angular: 10 dendritic Jan 1982. SAM SAM K 18 10(2); Streaky Bay, Point Westall, 15 Jan tentacles, ventral 2 smaller; pedicels in 1-2 rows 1991. NMV F592I90); Yorke Peninsula. Gleesons on radii. Body wall ossicles cups and small knob- Landing. 8 Nov 1976. SAM K. 18 16(1). bed perforated plates, often with 4 primary holes; pedicels with perforated plates and end- Description. Largest 32 mm long, 10 mm wide, 6 short, plates. Calcareous ring with posterior notches mm high (tentacles withdrawn); body and long tapers anteriorly. wide, low, to elongate or thin, slightly pentag- onal in transverse section, frequently twisted; plana (Brandt. 1 835). Type species. Ocnus body wall thick, finely crystalline, soft and flex- Remarks. Ocnus has traditionally been a large ible; mouth orientated anteriorly, anus pos- and heterogenous taxon. Panning (1971) restric- teriorly; 10 dendritic tentacles, 2 ventral ones (with synonym distinctly smaller; distinct thin-walled modified ted it to only 5 species: O. planci iacleus distinct introvert, lacking pedicels. O. brunneus Forbes, 1841) and O. ventrum; rows on 5 radii, more regular Forbes. 1841 from Europe; and with some Pedicels in paired ventrally creating distinct modified doubt: O. spvridophora H.L. Clark, 1925 from and distinct few small dorsal and lateral inter- Hawaii. O. viearius Bell, 1883 and O. cakareus ventrum; a pits created where pedicels (Dendy, 1896) from New Zealand. Unfortu- radial pedicels; small previously withdraw into body wall. Calcareous ring lack- nately "he did not assign other species, Meanwhilc posterior prolongations; 5 radials, with referred to Ocnus. to other genera. ing with scallops, tapered notched anterior pro- Rowe (1970) established Aslia for species posterior with posterior scallops, knobbed buttons with four holes only. This dis- jections; 5 interradials, with four pointed anterior projections. Dorsal madrcp- tinction is a little arbitrary as buttons holes orite. Single polian vesicle, left laterovcntral. holes and buttons with more than four 248 I'. MARK O'LOUGHLiN AND TIMOTHY D. O'HARA

plane of the cup rim (fig. 70- The dorsal body Dorsal body wall ossicles ol"4 types. 1 ) Masses of knobbed buttons, fairly regular, typically wall contains similar bar-like ossicles to O. occi- unfortunately 4-holed, oval, flat, knobbed centrally and per- duus. The tentacle ossicles are ipherally; typically 14 knobs, 2 central ones not extensively eroded, but no rosettes were normally larger than peripheral ones, mid- observed. The body form is more distinctly pen- lateral and mid-terminal knobs often smaller tagonal anteriorly. The pedicels emerge from than central ones or absent; largest buttons typi- body wall tubercles, and in no instances are cally 0.09 mm long. 2) Perforated bar-like oss- withdrawn into pits. These differences are slight, icles, thick, frequently bent or curved, up to but in the opinion of the authors, significant 0.29 mm long. 3) Rare large plates, irregularly enough to recognize a new taxon. oval, smooth to lumpy, regular rounded perfor- O. occiduus is usually found under rocks in the ations, up to 0.4 mm long. 4) Abundant cups, shallows. Despite intensive searching, this fairly regular, shallow, about 4 times as long as species has not been found east of Yorke Penin- deep; frequently oval and 4-holed, some rec- sula, South Australia. tangular and up to 8-holcd; capitate to peg-like Thyonidiinae Heding and Panning, 1954 spinelels centrally, peripherally, mostly pointing in all directions; cups typically 0.04 mm long. INeocucumis Deichmann, 1944 Ventral body wall ossicles as dorsally, but lack- Diagnosis (based on species currently assigned ing large plates. Pedicels with endplates, 0.22 to the genus). 20 dendritic tentacles, usually in mm wide; bar-like ossicles, curved or bent, 2 rings, 5 pairs of large intcrradial tentacles, widened centrally, perforated, up to 0.26 mm 5 pairs of small radial tentacles; pedicels in long; convex, irregular, denticulate, perforated double rows on radii; ossicles tables with round plates, 0.1 I mm long. Tentacles with small to or irregular perforated discs (0.05-0.45 mm large bar-like ossicles, straight and bent, some diameter) and 2-pillared spires with terminal widened centrally, perforated, up to 0.4 mm thorns; pedicel ossicles tables with elongate nar- long; convex mesh-like plates, irregularly oval to row discs and low 2-pillared spires, endplates, triangular, denticulate, typically 0.06 mm wide: rarely rods, rosettes; calcareous ring simple, rosettes, 0.07 mm long; some irregular, smooth, without posterior prolongations, 10 plates, slightly convex, perforated plates, 0.11 mm radials irregularly rectangular with a large pos- long. terior notch and 1-3 smaller anterior notches, Live colour. Body uniform white, grey on thin interradials at least 0.7 times as high as radials ventral surface and sometimes laterally; tentacle with a median posterior notch and sharp trunks dark brown to black; line tentacle anterior point. branches pale yellow. Type species. Cueumaria marioni Von Maren- Reproduction. Sexes separate; mature gonads zeller, 1877. present January and October; gonoduct right Neocueumis cauda sp. nov. dorsolateral; gonad tubules not forked. Plates 2e. lOa-f, Figure 8a-d Etymology. From the Latin occidaus (western), Material examined. Holotvpc. Victoria, 19 Sot" in reference to the western occurrence of this km Lakes Entrance, .WW'S, I48WF, 28 m, fine species in southern Australia. sand/shell substrate, 12 Aug 1989, G. Parry on FRY "Sarda". NMV F57952. Distribution. Rottnest I., Western Australia to Paralvpe. Type locality and date. Yorke Peninsula, South Australia. 0-30 m. NMV F57953(l). Other material. Remarks. Three small specimens of Ocnus cal- New South Wales, oft" Cape Three Points. 76-93 m, "Thetis". Feb 1898, AM careus from New Zealand (AM CJ 1 762) were .116358(4). examined for comparison. They conform to the descriptions and figures given by Dendy ( 1 896) Description. Holotype 12 mm long (tentacles and Mortensen (1925). They differ consistently fully withdrawn), 4 mm wide, tail 4 mm long; from the Australian species. The largest buttons paratype 9 mm long (tentacles fully withdrawn), have two central knobs which are normally sig- 3.5 mm wide, tail 2 mm long; body wall thin; nificantly larger than the 12 uniform peripheral body round in transverse section, with long thin ones (tig. 7e). The cups are slightly smaller, posterior tail; 20 dendritic tentacles, 5 outer typically 0.03 mm long; normally four-holed; pairs large, 5 inner pairs very small; lacking ring typically with spinelels pointing vertically to the of papillae around tentacles. Pedicels in 2 rows NEW HOLOTHURIANS FROM SOUTHERN AUSTRALIA 249

«=

radial J . , .„ f.a«svorc^«f>rtton.ofbodv scalebar- 1 .0 mm; b, adjacent and

KI8I7. KI802; k, SAM KI796; I, SAM 150 P. MARK O'LOUGHLIN AND TIMOTHY D. O'HARA

perforated ends in the ten- on all 5 radii, up to 80 pedicels on each radius; rod ossicles with wall tables pedicels slightly more developed ventrally; tacles, and the spire on the body interradial areas and introvert lacking pedicels; which is four-columned at the base. differ in 5 pedicles around anus. Calcareous ring plates All other species of Neocucumis irregularly- slightly separate; 5 radial plates with 4 pointed having table ossicles with very anterior projections, narrow angular waist, 2 shaped discs, denticulate discs, or with more or pointed posterior projections; 5 interradial fewer perforations on the disc. plates with single pointed anterior projection, The type specimens of A', eauda were dredged widened angular shoulders, 2 pointed posterior from coarse sediments. projections. Dorsal madreporite. Single left Neocucumella Pawson, 1962 lateral polian vesicle. Body wall ossicles of one type. Abundant Diagnosis (modified from Pawson, 1962b). 20 closely situated tables; discs rectangular to oval, dendritic tentacles, 5 outer pairs of large inter- frequently very irregular, typically 0.09 mm radial tentacles, 5 inner pairs of radial tentacles; long, typically with 4 large and 4 smaller corner pedicels in double rows on radii; body wall oss- holes, rarely up to 16 holes; spires v-shaped. 2 icles tables with regular oval discs with 4 large pillars fused distally, up to 0.04 mm high, typi- and 4 small holes, 0.05-0.10 mm long, spires cally 4 pairs of pointed terminal spines. Intro- with 2 pillars and terminal spines; pedicel oss- vert with sparse, narrowly oval, irregular tables, icles endplates, perforated supporting plates and curved discs; tentacle up to 1 2 holes, up to 0.08 mm long. Pedicels with tables, sometimes with endplates, 0.13 mm wide; tables with curved ossicles rods with widened, perforated, denticu- discs, frequently very irregular, many very nar- late ends; calcareous ring simple, without pos- row, widened in centre, rounded at ends, up to terior prolongations. 10 plates, radials notched 0.09 mm long, typically with 4 central holes and posteriorly and anteriorly, posterior ends diver- small terminal holes, spires similar to body wall gent, anterior ends truncate, interradials small, tables, shorter, up to 0.03 mm high. Tentacles inverted V-shaped, less than half as high as with numerous thin-walled, elongate, curved radials. plates, up to 0.18 mm long, perforations Tvpe speeies. Pseudocucumis bicolumnatus elongate; numerous very irregular mesh-like bendy and Hindle, 1907. plates, partly denticulate, typically 0.06 mm

wide. Remarks. Pawson ( 1 962b) distinguished his new genus Neocucumella from Neocucumis by the Reproduction. Holotype and paratype with dor- presence of regular eight-holed tables, and sal female gonad; gonad tubules filled with eggs; rounded ends to the interradial plates in the cal- up to 10 large tubules, 5 small; tubules elongate, careous ring. Another significant difference is each with a single fork mid-length. Gonoduct on the additional supporting perforated plates right dorsal body wall. which surround the endplate in the pedicels. The Colour (preserved). Uniform cream; tentacles collar of papillae surrounding the tentacle crown yellowish-green; tentacle trunks with transverse in Neocucumella is also present in at least one reddish-brown markings. Neocucumis species, N. watasei (Ohshima. 1915). Etymology. From the Latin eauda (tail), with ref- Of the other cucumariid genera which also erence to the long, thin posterior end of the possess two-spired tables, Mensamaria Clark, body. 1946 has 30 tentacles and Amphicvclus Bell. Distribution. Bass Strait, south of Lakes 1884 has 25. Entrance, to Cape Three Points, New South Neocucumella fracta sp. nov. Wales. 28-93 m. Plates 2d, lOg-j, Figure 8e-l Remarks. N. eauda is most similar to N. watasei (Ohshima, 1915) from Japan. N. watasei also has Pseudocucumis bicolumnatus. — Joshua and Creed. body wall tables with 4 large and 4 small holes 1915: 19. alternating around the table disc, and pedicel Mensamaria bicolumnata. — H.L. Clark, 1946: tables discs that curve away from the spire (Hed- 406. ing and Panning, 1954). N. watasei can be dis- Neocucumella bicolumnata. — Rowe. 1982: 467 tinguished from N. eauda by the presence of a [non Neocucumella bicolumnata (Dendy and Hindle. collar of papillae around the tentacles, irregular 1907)]. NEW HOLOTHURIANS FROM SOUTHERN AUSTRALIA 251

Material examined. Holotype. South Australia, some with up to 4 perforations terminally, some Spencer Gulf. Point Lowlv, 33°01'S. 137°48'E, 22 m, with clumps of small knobs, typically 0.08 mm Sep 1987, RV "Ngerin", SAM K1797. long. Paratvpes. Upper Spencer Gulf. E of Monument Hill. 32°50'S. 137"49'E. 17 m. 7-20 Feb 1986, SAM Colour (preserved). Uniform very pale yellow to K1817(l): E of Point Lowlv. 33WS. 137°48'E, 24 m, white. Sep 1987, SAM K1792(i): Point Lowlv, 33°01'S. 137°48'E, 22 m. Aug 1986. SAM K1796('l); 33WS. Reproduction. Sexes separate; male, female 137°50'E. 12 m. Sep 1987, SAM K1802(l): Fairway gonad tubules distinguishable in specimens col- 33°02'S, 137 <, 45'E, 18m, Feb 1987. Bank. SAM lected in February. Gonad, gonoduct dorsal; K 1793(2). gonad tubules not forked. Other material. South Australia, no locality or date, coll. Verco. identified as Pseudocucumis bieolum- Dr Etymology. From the Latin fracta (fragmentary), nata by Joshua and Creed. 1915, SAM K 1373(1); 14 with reference to the number of regular posterior lots from Upper Spencer Gulf, various stations, body ends collected. 32°50'-33"05'S, 137°40'-1 37°50'E, Feb 1986-Sep 1987, 10-24 m. SAM(5 and 14 •tails'). Distribution. Spencer Gulf, South Australia, to Victoria, eastern Bass Strait, 38°53.7'S, 147°55.2'E. eastern Bass Strait. 10-71 m. 71 m, shell/sand. 17 Nov 1981, BSS stn 171. NMV F592I40). Remarks. N. fracta is similar to the only other Description. Body (contracted, tentacles with- Neocucumella species, N. bicolumnata (Dendy drawn) up to 32 mm long, diameter 7.5 mm; and Hindle). known from New Zealand in 7-239 body rounded to slightly pentagonal in trans- m. N. bicolumnata, as described by Dendy and verse section; long thin posterior end. frequently Hindle(1907: 106) and Pawson (1963: 23; 1970: detached; lacking sole or modified ventrum; 32), differs in having consistently smaller body mouth anterior, anus posterior; 20 tentacles. 5 wall and tentacle ossicles, tentacle rods with pairs large, 5 inner pairs small. Fur-like collar of many more terminal perforations, no pedicels numerous fine papillae around the tentacle ring. on the introvert, and no curved table ossicles in 0.5 mm long. Lacking oral valves and anal teeth. the pedicel walls. The tables typically measure Pedicels in 2 rows on each radius; pedicels thin, 0.06 mm across and 0.02 mm high. The tentacle up to 80 in each row; pedicels extend across long rods, which have up to 20 terminal perforations, introvert: up to 15 small anal pedicels. Cal- measure up to 0.06 mm long. N. bicolumnata careous ring lacking posterior prolongations; 5 specimens up to 90 mm long have been found, radial plates with anterior notch, narrowed and are usually coloured red or brown in waist, widened posterior end, wider than alcohol. anterior end. with large notch; 5 interradials A specimen of N. bicolumnata from New Zeal- identified small, inverted V-shaped, with one anterior, 2 and (NZOI station B41, 13 mm long), posterior blunt pointed corners. Dorsal madrep- by Pawson (see Pawson, 1970: 32), was exam- is consistent with pre- orite. Usually single left lateral elongate tubular ined for comparison. It descriptions. It differs from N. fracta in polian vesicle (one specimen. SAM K 1 796. with vious smaller size of the body wall tables, 0.06 mm 4). the 0.36 wide; Body wall ossicles of one type. Abundant long; larger endplates. up to mm with curved discs; and closely-situated tables: discs oval, regular. 8 per- absence of pedicel tables "tooth" at the centre of forations, typically 0.08 mm long; spires with 2 the presence of a pointed of the radial plates in the cal- pillars, slightly tapered and joined distally, 0.04 the anterior notch Introvert and tentacle features mm high, typically with 8 blunt spines. Introvert careous ring. because of the condition with tables, some smaller and narrowly oval, could not be confirmed some with more than 8 perforations, typically of the specimen. 0.2 single Bass Strait specimen (NMV 0.06 mm long. Pedicels with endplates, up to The from the South Australian N. mm wide; curved supporting plates, elongate, F59214) differs in having larger tentacle rods, up curved, perforated, irregular, sometimes den- fracta material long, and smaller body wall tables. ticulate, typically 0.1 mm long, situated around to 0.15 mm long. The tables are similar in size to A'. endplates; tables with discs curved away from 0.06 mm to 20 bicolumnata. The presence of pedicels on the spire, narrower than body wall discs, up the form of the tentacle rods, the pres- perforations, typically 0.08 mm long, spire introvert, rarely ence of pedicel tables with irregular curved 0.03 mm high. Tentacles with thin rods, and the absence of a radial plate "tooth" in with short branches centrally or terminally. discs, 252 P. MARK O'LOUGHLIN AND TIMOTHY D. O'HARA

G., 1961. Deux nouvelles especcs the anterior notch, are all consistent with the N. Cherbonnier. d'holothuries dendrochirotes des cotes Bresil- fracta types. iennes. Bulletin du Museum National D 'Historic Joshua and Creed (1915) recorded a specimen Naturelle. Paris. (Serie 2) 33(6): 611-615. figs (SAM 1373) of N. bicotumnata collected by Dr K 1-2. Australia. This specimen, Verco from South Cherbonnier, G.. 1972. Neocnus incubans, nouveau belongs to the present upon examination, clearly genre et nouvelle espece d'holothurie dendro- species. chirote incubatrice de Mediterranee. Comptes Rendus des Seances de I Academic des Sciences, Acknowledgments Paris. (Serie D) 275(1): 225-227, figs a-q. 1957-1963. We are grateful to Dr Frank Rowe (recently of Clark, A.M., 1966. Port Phillip Survey, Memoirs of the National the Australian Museum) and to Dr David Paw- Echinodermata. Museum of Victoria 27: 289-384, 10 figs, 4 pis, 1 son (Smithsonian Institution) for their encour- tbl. agement, assistance with the systematics, and Clark, A.M., and Rowe, F.W.E., 1971. Monograph oj English translations of Panning's papers; to Dr shallow-water lndo-West Pacific echinoderms. of Vic- C.C. Lu and Ms Suzanne Boyd (Museum British Museum (Natural History): London, pp toria) for their support and ready assistance with vii + 238, 101 text-figs, 31 pis. the resources of the Department of Invertebrate Clark, H.L.. 1914. The echinoderms of the Western Zoology in the Museum of Victoria; to Dr Maria Australian Museum. Records of the Western Aus- Byrne for her assistance with reproductive tralian Museum I: 132-173, pis 17-26. than sea stars of biology; to Chris Rowley for his assistance with Clark. H.L.. 1925. Echinoderms other the tropical central Pacific. Bernice P. Bishop photography; to Mark Norman and Tania Museum Bulletin 27: 89-1 12. pis 9-11. Bardsley for assistance with the figures; to Karen Clark. H.L.. 1938. Echinoderms of Australia. An Hazeldene for assistance with labelling the col- account of collections made in 1929 and 1932. for the drawings of lection; to Margie Atkinson Memoirs ofthe Museum ofComparative Zoology. Neocnus bimarsupiis; to Mr Wolfgang Zeidler Harvard 55: viii + 1-596, 63 figs. 28 pis.

(SAM) and Dr Penny Berents (AM) for provid- Clark, H.L., 1 946. The echinodcrm fauna of Australia, ing research facilities and the loan of compara- its composition and origin. Publications of the tive specimens; to Loisette Marsh (WAM), Carnegie Institution 566: iv + 1-567. Lynne Albertson (AM), Lisa Hobbs (SAM), Paul Deichmann, E., 1930. The holothurians of the western Bulletin the Museum Anderson (NZOI), Alison Green and Elizabeth part of the Atlantic Ocean. of of Comparative Zoology 71(3): 43-226, 24 pis. Turner (TM) also for assisting with the loan of Deichmann, E.. 1941. Holothuroidea collected by the comparative specimens; and to Clarrie Hand- Valero III. during the years 1932-1938. Part 1. Christine Materia, Nyhuis, Joseph- reck, Marita Dendrochirota. Allan Hancock Pacific Ex- for their ine Stephenson and Keryn Walshe peditions 8: 61-153, text figs 1-6, pis 10-30. fieldwork. significant contribution to the Deichmann, E., 1944. Urodemas bifurcation, a new holothurian from South Africa, with revision of References a the genus Urodemas Selenka. Annals and Maga-

Bell, F.J., 1883. Studies in the Holothuroidea II: zine oj Natural History (Scries 1 1), 1 1: 731-737, 3 Descriptions of new species. Proceedings of the figs. 58-62.' Zoological Society of London 1883: Dendy, A., 1 896. Observations on the holothurians of Bell, F.J., 1884. Studies in the Holothuroidea III: On New Zealand. Journal of the Linnean Society. Amphicyclus, a new genus of dendrochiritous London (Zoology) 26: 22-52. pis 3-7. Holothurians. and its bearing on the classification Dendy. A., and Hindle, E., 1907. Some additions to of the family. Proceedings of the Zoological our knowledge of the New Zealand holothurians.

Society of London 1884:253-258, 1 tbl. Journal ofthe Linnean Society, London (Zoology) Bell. F.J., 1887. Studies in the Holothuroidea VI: 30: 95-125. pis 11-14. Descriptions of new species. Proceedings of the Ekman, S., 1918. Results of Dr. E. Mjoberg s Swedish Zoological Society ofLondon. 1887: 531-534. pi. Scientific Expeditions to Australia 1910-1913. 45. XIX Holothuriodea. Kungliga Svenska Vetens- Blainville, H.M.A. de, 1834. Manuel d'Aclinologie et kapsakademiens Handlingar 58(6): 1-70, 5 pis. de Xoophytologie. Paris. 188-197. Emson, R.H., and Wilkie, L.C., 1980. Fission and Brandt, J.F., 1835. Prodromus descriptionis animal- autotomy in echinoderms. Oceanography and ium ab H. Mertensio in orbis terrarum circum- Marine Biology, Annual Review 18: 155-250, 19

navigatione observatorum Petropoli I: 1-75, figs, 1 7 tbls.

I pi. Erwe, W., 1913. Holothuroidea. In Michaelsen, W., Cannon. L.R.G., and Silver, H., 1987. Sea cucumbers und Hartmeyer, R. (eds) Die Fauna Siidwest-

ofnorthern Australia. Queensland Museum: Bris- Australiens 4: 351-402, fig. 1. pis 5-8. bane, pp viii + 60. Forbes, E.. 1841. A history of British starfishes and NEW HOLOTHUR1ANS FROM SOUTHERN AUSTRALIA 253

Other of the class Echinodermata. John Ludwig, H., 1874 [1875]. Beitrage zur Kenntniss der van Voorst, London: xx + 267 pp. Holothurien. Arbeiten aus dem Zoologisch-Zoo- Goldfuss, G.A., 1820. Handbuch der Zoologie. tomischen Institut in Wiirzburg 2: 77-120, pis Nuremberg 1: 1-696, 2 pis. 6-7. Fahrt Grube. A.E., 1840. Aclinen. Echinodermen und Ludwig, H., 1886. Die von G. Chierchia auf der W'iirmer des Adriatischen und Miltlemeeres nach der Kgl. Ital. Corvette "Vettor Pisani" gesam- eiqenen Sammlungen heschrieben. Konigsberg: mclten Holothurien. Zoologische Jahrbucher 2: 33-43. 1-36, pis 1-2. 1894. Reports on an exploration off the Gutt. J.. 1990. New Antarctic holothurians (Echino- Ludwig, H., genera west coasts of Mexico, Central and South Amer- dermata) 1 . Five new species with four new charge of of the order Dendrochirotida. Zoologica Scripta ica, and off the Galapagos Islands, in Agassiz, the U.S. Fish Commission 19(1): 101-107, 19 figs. 4 tbls. Alexander by 1891, Lieut. Com- Heding. S.G., and Panning, A.. 1954. Phyllophoridae. Steamer "Albatross" during commanding. 12, Eine Bearbeitung der Polytentaculaten Dendro- mander Z. L. Tanner, U.S.N, the Museum of chiroten Holothurien des Zoologischen Museums The Holothurioidea. Memoirs of Zoology. Harvard 17(3): 1-183, pis in Kopenhagen. Spolict Zoologica Musei Haun- Comparative 1-19. siensis 13: 7-209, 102 figs. Ergebnisse der Ham- Hernandez, D.A., 1982. Holothuroidea des Siidwes- Ludwig, H„ 1898. Holothurien. burger Magalhaenischen Sammelreise 1X92/93, tatlantiks 1. Die Trachythyone-Arlcn. Mittei- Friederichsen and Co.: Ham- lungen aus dem Hamburgischen Zoologischen Band 1. 1-98. L. Museum undlnslitul 79: 251-261, 4 figs, 2 tbls, 2 burg. Ludwig, H., and Heding, S.G., 1935. Die Holothurien pis. der Dcutschcn Tiefsee-Expedition. 1. Fusslose Herouard. E., 1901. Note preliminaire sur les holo- Dendrochirote Formen. Wissenschaftliche thuries rapportees par I'Expcdition Antarctique und Ergebrisse der Deutchen Expedition Tiefsee auf Beige. Archives de Zoologie experimentale et gen- dem Dampher "Valdivia" 1898-9. 24: 121-214. erate, Paris. Notes 3(9): 39-48. E. Von, 1877. Beitrage zur Holothurien- Hickman, V.V., 1962. Tasmanian sea-cucumbers Marcnzeller. fauna des Mittlemeeres. Verhandlungen (K.-K.) (Holothuroidea). Papers and Proceedings of the 1- Zoologisch-bolanische Gesellschaft, W'ein. 27: Royal Society of Tasmania 96: 49-72, text figs 117-122, pi. V. 186. pis 1-2. Marine Research Group of Victoria, 1984. Coastal Hickman. V.V., 1978. Notes on three species of Tas- Victoria. An atlas selected manian sea-cucumbers including one species that invertebrates of of (Holothurioidea: species. Marine Research Group in association broods its young in the coelome. with the Museum of Victoria: Melbourne. 1 68 pp, Phyllophoridae. Caudinidae). Papers and Pro- 13 figs, 1 tbl. ceedings of the Royal Society of Tasmania 1 1 2: Materia, C.J., Monagle, J.F., and O'Loughlin, P.M., 29-37. 44 text-figs, pis 1-2. 1991. Seasonal coelomic brooding in southern Hutton, F.W., 1872. Catalogue of the Echinodermata species. Australian cucumariids (Echinodermata, Holo- of New Zealand, with diagnoses of the Pp. 301-307, 5 figs, 5 tbls in: Yanag- Wellington. 20 pp. thurioidea). James Hughes: Motokawa Notes on some New Zeal- isawa, Yasumasu, Oguro, Suzuki and Hutton, F.W.. 1 879 [ 1 878]. new (eds). Biology of Echinodermata. Balkema: Rot- and Echinodermata, with descriptions of the New terdam. species. Transactions and Proceedings oj Mortensen, T., 1925. Echinoderms of New Zealand Zealand Institute II: 305-8. and the Auckland-Campbell Islands. 1II-V. Joshua E.C., 1912.0nanewholothunanofthegenus Proceed- Asteroidea, Holothurioidea and Crinoidca. Taeniogyrus found in Port Phillip Bay. 79-8 pis Vidcnskabelige Meddelelser fra Dansk naturhis- ings the Royal Society of Victoria 25: 1, of figs torisk Forming i Kobenhavn 79: 26 1 -420, text 3-4 Holothuroidea with 1-70, pis 12-15. Joshua E.C., 1914. Victorian oj the Ohshima, 1915. Report on the holothurians collected descriptions of new species. Proceedings by the United States Fisheries Steamer "Alba- Society Victoria 27: 1-11, pi. L Roval of during the South Australian tross" in the Northwestern Pacific Joshua, E.C.. and Creed, E., 1915. species. summer of 1906. Proceedings ofthe United Slates Holothuroidea with descriptions of new 48: 213-291, pis 8-11. the Royal Society National Museum 7 ransactions and Proceedings oj Oken, L., 1815. Lehrbuch der Naturgeschichle. Part 3. Australia 39: 16-24, pis 2-4. of South xxviii + Holothurioidea. Zoologie. Leipzig and Jena, pp + 850 Lampert, K„ 1885. Die Seewalzcn. xviii. Semper ( . Eine Svstematichc Monographie. In O'Loughlin, P.M., 1991. Brooding and fission in shal- im Archipel der Phillipinen. Vol. 4_ (ed ), Reisen of southern Australia. Pp. pi., 3 low water echinoderms C.W. Krcidel's Verlag: Wiesbaden. 3 1 2 pp, 1 223-228, 5 figs, 1 tbl. in; Yanagisawa, Yasumasu, Suzuki and Motokawa (eds). Biology of zoologique ou choix Oguro, Lessorf RP 1830. Cenlurie imperjaitement Echinodermata. Balkema: Rotterdam. d'animaux rares, nouveaux ou Pallas, P.S., 1766. Miscellanea Zoologica. Hagaee: connues. Paris: 1-244, 80 pis. . :.

254 P. MARK OLOUGHLIN AND TIMOTHY D. O'HARA

revised classi- 152-156, pl.xi, figs 10-12. Pawson, D.L., and Fell, H.B.. 1965. A Panning, A., 1949. Versuch einer Neuordnung der fication of the dendrochirote holothurians. Familie Cucumariidae (Holothurioidea: Dendro- Breviora2\4: 1-7. chirota). Zoologische Jahrbuchcr, Abteilung fur Quoy. J. R.C, and Gaimard, J. P., 1833. Voyage de ' Systematik, Okologie und Geographic der Tiere decou vertes de I'"Astrolabe '. Zoologie, Zoophytes 78(4): 404-470, 62 figs. Paris: i-390. 26 pis. Panning, A.. 1962. Bermerkungen iiber die Rowe. F.W.E., 1970. A note on the British species of Holothurien-Familie Cucumariidae (Ordnung cucumarians. involving the erection of two new

Dendrochirota). 3, Teil. Die Gattung Pseudocnus nominal genera. Journal of the Marine Biological Panning, 1949. Mitteilungen aus dem Hambur- Association of the United Kingdom 50: 683- gischen Zoologischen Museum und Institut 60: 687. 57-80, figs 1-20. Rowe. F.W.E.. 1976. Restriction of the chiridotid Panning, A., 1964. Bermerkungen iiber die genus Trochodota Ludwig (1891) (Holothu- Holothurien-Familie Cucumariidae (Ordnung rioidea: Apodida). with the description of a new Dendrochirota). 4. Teil. Die Gattungen Stereo- species from South Australia. Transactions of the

derma, Slaurolhyone, und Trachythyone. Mittei- koval Society ofSouth Australia 1 00(4): 203-206.

lungen aus dem Ilamburgischen Zoologischen figs I -4. tbl 1 Museum und Institut (Kosswig-Festschrift) 61: Rowe, F.W.E.. 1982. Sea-cucumbers (class Holothu- 159-74, figs 1-10. rioidea). Pp. 454-474, figs 10:26-10:37, pis 29- Panning, A., 1966. Bermerkungen fiber die 32 in: Shepherd, S.A. and Thomas. l.M. (eds). Holothurien-Familie Cucumariidae (Ordnung Marine invertebrates ofsouthern Australia. Part I. Dendrochirota). 5. Teil. Die Gattungen Iletero- Government Printer: Adelaide.

thyone Pannning, 1949 und Leptopentacta H.L. Rowe, F.W.E. and Vail, L.L., 1 982. The distribution of Clark, 1938. Mitteilungen etui dem Ilambur- Tasmanian echinoderms in relation to southern gischen Zoologischen Museum und Institut 63: Australian biogcographic provinces. Pp. 219-

51-69, 8 text-figs, 4 pis. 225, 1 fig. in: Lawrence J. M. (ed). Echinoderms Panning, A., 1971. Bermerkungen fiber die Proceedings of the International Conference, Holothurien-Familie Cucumariidae (Ordnung Tampa Ray. Balkema: Rotterdam. Dendrochirota). 6. Teil. Die Gattungen um Ocnus Rowe, F.W.E.. and Pawson. D.L.. 1985. Loisettea Forbes. 1841 und um Pentacta Goldfuss, 1820. ampliictena, new genus, new species, from the Mitteilungen aus dem Hamburgischen Zoolog- sublittoral of northwestern Australia (Echinoder- ischen Museum und Institut 67: 29-51. text— figs mata: Holothuria). Proceedings of the Biological 1-5, pi. 3. Society of Washington 98(3): 672-677, figs 1-3,

Pawson, D.L., 1962a. A new from Mae- tbl 1.

quarie Island. Transactions ofthe Royal Society of Selenka, E., 1 867. Beitrage zur Anatomie und System- New Zealand, Zoology 2(7):' 47-48, pi. 1. atik der Holothurien. Zeitschr fur wissenschaft- Pawson, D.L.. 1962b. A new phyllophorid genus in the liche Zoologie, Leipzig 17: 291-374, pis 17-20. Holothuroidea. Transactions of the Royal Society Semper, C, 1868. Holothurien. Reisen im Archipel der of New Zealand, Zoology 2(1 1): 65-67, figs 1-2. Philippinen, Teil 2 Wissenschaftliche Resultate.

Pawson, D.L., 1963. The holothurian fauna of Cook Leipzig. Bd 1: 1-288, pis 1-40. Strait, New Zealand. Zoology Publications from Semper, C, 1869. Die Holothurien Ostafricas. In: Victoria University of Wellington 36: 1-38, 7 Decken, C.C. von der(ed.), Reisen in Ost Africa in pis. den Jahren 1859-1865. Leipzig and Heidelburg

Pawson, D.L., 1964. The Holothuroidea collected by 3(1): 119-122, 1 pi. Society southern the Royal Expedition to Chile, Studer, T., 1 876. Uber Echinodermen aus dem antar- 1958-59. Pacific Science 18(4): 453-470, figs 1- kischen Meere und zwei neue Seeigel von den 3. Papua-lnseln, gesammelt auf der Reise SMS Pawson, D.L., 1966. Evolution and phylogeny of the "Gazelle" um die Erde. Monatsbaichte d.k. Holothuroidea. Pp. U641-U646 in: Moore, R.C. Preussiche Akademie der Wissenschaften. Berlin: (ed), Treatise on Invertebrate Paleontology, Part 452-65. U(2), Echinozoa and Asterozoa. Geological Thandar, A.S., 1 985. A new southern African genus in Society of America and The University of the holothurian family Cucumariidae (Echino- Kansas. dermata: Holothuroidea) with the recognition of Pawson, D.L., 1970. The marine fauna of New Zeal- two subspecies in Cucumaria frauenfeldi Ludwig. sea cucumbers (Echinodermata: and: Holothu- South African Journal ofZoology. 20(3): 109-1 14,

roidea). New Zealand Oceanographic Institute 4 figs, 1 tbl. Memoir 52: 1-69, 10 text-figs, 2 pis. Thandar. A.S., 1986. A new genus and species of a D.L., 1983. sacculus Pawson, Ocnus new species dendrochirotid holothurian from South Africa. (Echinodermata: Holothuroidea) a brood protect- Zoological Society ofLondon (A) 210: 483-488. 1 ing holothurian from south-eastern New Zeal- fig. and. New Zealand Journal of Marine and Thandar, A.S., 1987. The status of some southern Freshwater Research 17: 227-230, 2 figs. African nominal species of Cucumaria (s.e.) 1

NEW HOLOTHURIANS FROM SOUTHERN AUSTRALIA 255

referable to a new genus and their ecological iso- ary developments, NMV F54704; b, dorsal body lation. South African Journal of Zoology 22(4)- wall multi-layered ossicle, NMV F53793; c, dor- 287-296, 7 figs. sal body wall spinous cups, larger 0.04 mm long, Theel. H.. 1 886. Report on the Holothurioidea. Part 2. NMV F53793; d, dorsal body wall knobbed but- Reports on the Scientific Results of the Voyage of ton, cups, and part of HMS "Challenger" 1873-76. Zoologv 39:1-290, a multi-layered ossicle, NMV F59220; e, ventral body wall and pedicel 1 6 pis. Troschel. F.H., 1846. Neue Holothurien-Gattungen. non-spinous cups, and flat and bar-like ossicles, Archiv fur Natuigeschichte. Berlin 12(1): 60-66. NMV F54704; f, dorsal body wall knobbed but- ton, NMV F53793; g, ventral pedicel flat and bar-like ossicles, NMV F59220; h, tentacle bar- Explanation of Plates like ossicles, NMV F59220.

Plate 1 Figures a-h, ventrolateral views of holo- Plate 5 Figures a-f. ossicles of Trachythyone types, scale bar = 5 a. Pentocnus mm: bursatus Candida sp. now, scale-bar = 0. 1 mm: a-b, dorsal gen. et sp. now. NMV F57549, 6 mm long; b. body wall perforated plates, some denticulate, Cucuvitrum rowei gen. et sp. now. NMV NMV F59215; c, ventral body wall cup, 0.05 F57356. 9 mm long; c, Neocnus bimarsupiis sp. mm long, holotype; d, ventral body wall cups, now, NMV F54238. 5 mm long; d. Squamocnus NMV F54701; e, ventral pedicel perforated aureoruber gen. et sp. now, NMV F54244. 1 plate, NMV F54701: f, tentacle ossicles, NMV mm long; e, Trachythyone Candida sp. now. F59215. NMV F59200. 8 mm long; f, Apsolidium alvci sp. Figures g-j. ossicles of Trachythyone glebosa now, SAM K 1 824. 54 mm long; g. Trachythyone sp. now, holotype, scale bar = 0.1 mm: g, dorsal glebosa sp. now. SAM K1812. 12 mm long; h. lump perforated plate; h, dorsal lump irregular

Ocntts occiduus sp. now, NMV F592 12. 23 mm cups; i, ventral pedicel endplate and curved nar- long. row perforated plates; j, tentacle bent bar-like ossicle and rosettes. Plate 2 Figures a-d. ventrolateral views of holo- types, scale bar = 5 mm: a, Apsolidium handrecki Plate 6 Figures a-i, ossicles of Apsolidium hand- gen. et sp. now, NMV F54391. 14 mm long; b, recki gen. et sp. now, type series, scale bar = 0.1 Apsolidium densum sp. now, NMV F54237, 35 mm: a, sole cups and button-like ossicles; b, sole mm long; c, NeocucUmis cauda sp. now, NMV lumpy and smooth button-like ossicles, cups,

F57952. 1 2 mm long; d. Neocueumella fracta sp. and curved pedicel plate; c, part of dorsal body now, SAM K1797, 25 mm long. wall multi-layered ossicle; d, side view of dorsal

Figures e-h. ossicles, scale bar = 0. 1 mm: e-g body wall developing cup, 0.05 mm wide; e, end Pentocnus bursatus gen. et sp. now: e, body wall view of dorsal body wall cup, 0.05 mm wide; f, perforated plate, NMV F59207; f. body wall per- ventral pedicel narrow perforated plates; g, ven- forated plates, type series; g, tentacle ossicles, tral pedicel endplate; h, tentacle bar-like ossicles mainly irregular rods, type series: h. Neocnus and mesh-like plate; i, tentacle rosettes. bimarsupiis sp. now. tentacle rods, NMV F54387. Plate 7 Figures a-h, ossicles of Apsolidium den-

sum sp. now, scale bar = 0. 1 mm: a, dorsal body Plate 3 Figures a-g, ossicles of Cucuvitrum rowei wall multi-layered ossicle, NMV F53766; b, dor- gen. et sp. now, scale bar = 0.1 mm: a. ventral sal body wall large knobbed cup, NMV F53766; body wall perforated plate with projecting knobs c, sole cups, NMV F53764; d, dorsal body wall and connecting bars, holotype; b. dorsal body smooth perforated plate, NMV F53766; e, dor- wall multi-layered ossicle, AM J 10883; c, dorsal sal body wall lumpy and smooth button-like oss- body wall knobbed perforated plates, NMV icles, NMV F53764; f, tentacle rosette, NMV F59221; d, dorsal body wall multi-layered ossi- F53764; g, pedicel bar-like ossicles, NMV dorsal h, tentacle bar-like ossicles and mesh- cles and knobbed plates, NMV F5922 1 ; e. F53764; body wall knobbed button-like ossicles, NMV like plates, NMV F53764. ossicles, holo- F5922 1 ; f, ventral pedicel bar-like 8 Figures a-f, ossicles of Apsolidium alvei type; g. tentacle ossicles, NMV F59221. Plate sp. n, holotype, scale bar = 0. 1 mm: a, dorsal button-like ossicles, Plate 4 Figures a-h, ossicles of Squamocnus aur- body wall large plate, and ventral body wall narrow plates and cup; eoruber gen. et sp. now, scale bar = 0.1 mm: a, cups; b, d, ventral body wall ventral body wall perforated plates with second- c, ventral body wall cups; 256 P. MARK O'LOUGHLIN AND TIMOTHY D. O'HARA

cups in side, end and ventral view; e, ventral pedicel endplate, thin perforated pedicel plates, and thick narrow body wall plates; f, tentacle bar-like ossicles.

Plate 9 Figures a-f, ossicles of Ocnus occiduus sp. nov., scale bar = 0. 1 mm: a, dorsal body wall buttons, SAM K181 1; b, dorsal body wall cups, SAM K1811; c, dorsal body wall perforated plate, SAM K 1 8 1 1 ; d, dorsal body wall bent bar- like ossicles, SAM K 1 8 1 1 ; e, pedicel bar-like oss- icles, SAM K1816; f, tentacle bar-like ossicles, mesh-like plates, rosette and button, SAM K1816.

Plate 10 Figures a-f, ossicles of Neocucumis cauda sp. nov., type series, scale bar = 0. 1 mm: a-c body wall and pedicel tables; d, introvert tables; e-f, tentacle plates. Figures g-j, ossicles of Neocucumella fracta sp. nov., scale bar = 0.1 mm: g-h, body wall tables, holotype; i, pedicel tables, holotype; j, tentacle rods, SAM K1373. NEW HOLOTHURIANS FROM SOUTHERN AUSTRALIA 257

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