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New cucumariid holothurians (Echinodermata) from southern Australia, including two brooding and one fissiparous species PDF

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Preview New cucumariid holothurians (Echinodermata) from southern Australia, including two brooding and one fissiparous species

Memoirs ofthe Museum ofVicloria 53(2): 227-266 (1992) NEW CUCUMARI1D HOLOTHURIANS (ECHINODERMATA) FROM SOUTHERN AUSTRALIA, INCLUDING TWO BROODING AND ONE FISSIPAROUS SPECIES By P. Mark OLoughlin and Timothy D. OHara Department ofInvertebrate Zoology. Museum ofVictoria, 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 ofthe Museum ofVicloria 53: 227-266. Twelve new species of cucumariid holothurians are described from oft' the coast of southern Australia. Pemoamsgen. nov. iserected forone specieswhich 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 otherspecies areassigned to Neocnus, Trachythyone(2species). Ocnus.Ncocucumisand Ncocucumella.Thenewspecies ofPcntocnusandNeocnusbroodtheiryoung.ThenewspeciesofSquamocnusisfissiparous. The new species ofCucuviirum is host to a copepod parasite (Cucumaricolidae). Introduction Joshua(1914)recorded the Indo-Pacificcucu- 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 ofVictoria 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 theNewZeal- 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). Ofthese, 5areabundantand idium) convergens (Herouard, 1901), and the well known forthe region: Psolicliella adhaerens subantarctic species Cucumaria (=Trachy- Hickman, 1962,Staurothyoneinconspicua(Bell. thyone)squamata (Ludwig, 1898). Rowe (1982) 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 fourcases 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- C and habitatnotesgiven byoneoftheus(M.O'L.) toides for the specimen of squamata, but, as in Marine Research Group of Victoria (1984). A.M. Clark(1966: 345)pointedout, thisname is Amphkyclus mortenseni Heding and Panning, a nomen nudum, failing to meet the criteria 1954. also described by Hickman (1962), occurs under ICZN Article 13 and is replaced herein. in deeper water in eastern Bass Strait. Extensive collectingofechinoderms from the Some essentially tropical species extend southern Australian coast by us and with associ- around the coast ofWestern Australia into the atessince 1978 has revealed many new holothu- GreatAustralian Bight(Rowe, 1982). Theseare: rian species. Most of the specimens were col- Mensamaria inlercedens (Lampert, 1885); Pen- lected offalgaeorrocks 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- mariastriataJoshua and Creed, 1915 (seeA.M. lections ofthe Museum ofVictoria (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),theNewZealandOceanographic Institute terior processes on the calcareous ring is (NZOI) and the Museum of Comparative considered to be in the Phyllophoridae. Zoology, Harvard (MCZ). The taxonomy within the Cucumariidae is Twoofthe new speciesbrood theiryoung: the also currently confused and ill-defined, particu- Neocnusspecies 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 breakupofold heterogeneoustaxasuch 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 thebodywall oftheformer. However, manyofthespeciesintheCucumariinae.asthen Terminology defined, were subsequently found to have cups orcup-like derivatives, orat least plates similar Terminology predominantly follows Clark and Rowe (1971) or Pawson (1970). Ossicle in form to genera in the Colochirinae. Panning (1971) finally restricted the Cucumariinae to types includecups(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. (psomdailaloprlattuebseifneetpi.are6hr)efaenrrdedrotdosas(ppie.di2che)l.s.AlAl resPtarnicntiendgth(e19d6i2a,gno1s9e6s4.of1m96a6n,y 1o9f71t)hefguerntehrear sole is a flattened, delimited modification ofthe within the Colochirinae. Consequently recent ventral bodywall with peripheral pedicelswhich 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 ofnew lacra (following Pawson, 1970). Left and right generaas neworexistingspeciescould no longer are relative to facing anteriorly alongthe dorsal be satisfactorily placed in existing genera. This surface. has led to a more natural classification, recog- Order Dendrochirotida 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, 894 1 ossicles. Remarks. The relationships between "cucu- Panning(1971)attempted todivide the Colo- mariid" and "phyllophorid" holothurians have chirinae into three distinct groups, based on the been confused. Panning (1949) and Hedingand number ofossicle types present and their func- Panning (1954) define the Cucumariidae and tion. An "Ocnus" group with two types ofbody the Phyllophoridae predominantly on the basis wall ossicles, a"Pentacta"groupwith three,and ofthenumberoftentacles, ten in theCucumari- 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 ( 1965)considerthat the nature adhesivetraction. 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 (1978) and Cannon and Silver (1987) 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 manyproblems, particularlyduetotheapparent 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 a 3 3 g &C © S£i?- £a .J©3j «3^ Sa <*U .5 3©O—N P'OO ~3©$'— <3 a ~§ SwSOO- Sar~. ,Js. .3a U3 <R3 3 3 3o 5u l^3--3go --u3S—wean -©5 don S 3 •I a ft.il r cS a,-S C cs 3 41 53 o 3 5 p "^3 O © a *nO—os s^\cj §' 2^ 3<a3 g3 S*5-'<aS3"C^Sro2-o2 *<pj MUCiO ©© •2-_*3' 5a!—oi «1 fat co •S32'C*c3"o—nC_uO- -g32 JDC $<53 "«©5 OI ft, ft, II ^3CSuC5S ^§> <3 o Ly- •-_ =2 PSS;Oo1^N S. « c ^ s =33 3J35 Si S"-1 ^cj 52 eft• 3 Bgo 'cS^. 55d "3j.; 2c; 1s.1s "5 £ jgac g § a O c 3STo—o ^aj _^^ _s •Si « .32 5g o *g3a5 t2j-3j\jJl ;~©^33 ^k^ao""J U o S3tte9c-« 2 .©sp g S«!U OS33 13'-o 3T3 55 ft. ft, s; ft, fe; o Z _0J ~ E cc tn oo O OCh3 fNNO| oo—oo eoo—noo oc;/>i cc R § BO doUg-V a_s £o s2o2-acoa TTc(0c33a30 £OM"atEo"nS• O?30N3 J—aEcdr « t! Ovai,s4OJO K& oc>• II00 6-S a> O jitjio; ^t33*a 3co/.l -^18^3 ©3 "3> p g I g I & c1^> 3 •5 C3 p 5 s 2 n6o S"3 AS —Si S 3 a 1 3 K g 3 "fS JH ^ 0a. & C.39o5 £§~3 c^ ^O sa. 33 ~«S, "LkS25,^~tS5.^"-gjo;.^"t2S*. gs sS «Sa «23 fS2t, f.St2. 1c= ~3 ~f«5335e3j;,3n«S2S3;jl 6 < 230 P. MARK O'LOUGHLIN AND TIMOTHY D. O'HARA coveringthe whole body. Noextant cucumariid clearly distinguishPentocnusfrom all othergen- genus has all ofthese characters, but some gen- era within the Cucumariidae. The shallow, era, e.g., Leptopentacta H.L. Clark, 1938, do saucer-like, spined form ofthe plates suggests a possess small extensions to the radial plates in possibleform orfunctional relationshipwiththe the calcareous ring, and non-imbricating multi- cupossicles present in othercucumariid genera. layered ossicles. The denticulate body wall plates possibly 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- Diagnosis. Worm-like form, lacking sole or drochirotid genus lacks them completely. In the modified ventrum; 10 irregularly developed subfamily Thyonidiinae. Athyonidium Deich- dendritic tentacles; calcareous ring discontinu- mann, 1941 has very reduced interradials, often ous, 5 spaced radial plates without posterior obscuredby muscle;theresultingringisalsodis- prolongations, no interradial plates; pedicels continuous. InAmphicyclusBell, 1884the inter- present on all radii, absent interradially. Body radials are also reduced, but they adjoin the wall ossicles small, elongate, perforated. Hat or larger radial plates in a continuous ring. saucer-like, often denticulate plates, with blunt Pentocnus bursatus sp. nov. vertical spines; lacking true cups; pedicels with similar plates to body wall, and endplates; ten- Plates la, 2e-g, Figure 1 tacleswith irregular perforated bar-likeossicles, Pseudocnus sp. - Roweand Vail, 1982: 224. small perforated plates and rods. 'cucumariid" sp - O'Loughlin, 1991: 225-226, Type species. Pentocnus bwsatus sp. nov. fig. 4. Material examined. Holotype. Victoria, Cape Etymology. From the Greekpente (five), in ref- 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 Rcteaenlmtcaaacrrlkeesos.u,stTrhhienegd,eltlahiceckaitorefrefigonurtleamrrrodafedivtaehlelopoplsamstiecenlstesio,nfattnhhdee a61n9M8d0ad.raNte1M.9AV83M,F5JN422M237V65(4(1F)51;3)r;7oN6c2kM(ylVs)h:aFl52l89o6w2sJ,9a(nal1g1ja9ul8ve)8p;.if1Na8uMJnaaVn. the other possible paedomorphic characters, F54181(l). B Figure 1.Pentocnusbursatusgen.etsp.nov.:a,transversesectionofbody,scalebar= 1.0mm;b,2adjacentradial plates from the calcareous ring; c, side and top view ofa dorsal body wall plate, scale bar = 0.01 mm. NEW HOLOTHURIANS FROM SOUTHERN AUSTRALIA 231 4 mS,out14h MAuasytra1l9i9a0,.BNeaMchVpoFr5t9,20377°(2l9)..3'S. 1 39°59 6'E dtiiaftfeedre(n0t.i8atmedm(1lo.n4g)m.mThleonbgr)o.oBdojtuhveanrieletoigsewtehlelr- VlOatmhienrghm.ateoruitaelr.WreeesfteHrant,Au9stJraalnia,19R9o1t,tnWesAtMI..C2a3p9e- in the coelom in a membranous sac which is 9119(911).:WRaAdaMrR2e5e5f-,91S(a1r)x.a.ssum/Cmoplwramat, 13Jan Tathtearceheisdntoocthheanagneteirniotrh,elbefotdyvewnatlrlalatbothdeypwoailnlt. D1emscmripwtiidoen;.bUopdytwoo1r5m-mlimkel,ornogu(nlidvei,netxrtaennsdveedr)s.e oexftNaertMitoaVrc.hmFe5n4t2,3n6or(1a0nymamppalronegn,ttaepnetratculreestwoitthh-e section, lackingsoleormodified ventrum, intro- drawn) hasadorsal gonad, 3 brood embryosand vertnotdistinct,mouth anterior, anusposterior; 2 juveniles. The gonad has 6 sac-like caeca, body wall thin, veryextensible; 1 dendriticten- 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 4smallorbud-like. Onerowof developing brown egg or embryo (up to 0.8 mm pedicelson each radius, with upto 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 broodjuv- 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 NMV mm eous ringNMarVe clearest NinMVspecimens WAM cases (1 long). Each embryo is in a thin- F57549. F53762. F59207, walled sac which hasa 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 ossiclesofonetype: small,elongate, ventral anterior. At the point ofattachment 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 mm AM to 0.09 long: typical plate with 2 large cen- few small white eggs is present. J22754 (8 mm tral holes, one smaller hole near each end, 1-6 long, tentacles not withdrawn) has 2 undif- very small holes at ends; plates frequently den- ferentiated embryos(onein agonad caecum,0.7 mm ticulate at one end or one side; ossicles may- long), an embryo with radial pedicels, and a mm imbricate or be spaced in extended body wall. well-developed juvenile (2.2 long) in a Pedicels with body wall plates; endplates with membranous sheath in the left anterior coelom. mm NMV mm irregular angular perforations, about 0.1 F53762(5 long) had 3brood embryos mm wide. Tentacles with small irregular plates with (up to 1.5 long) in separate thin sacs mm large angular perforations, upto 0.07 wide; attached toa scarorfold in thebodywall, 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 mm 0.07 long; a few large, irregular, branched, anteriorly. There is only part of the gonad mm NMV mm perforated bar-like ossicles, uptom0.m13 long; remaining. F59207 (5 long; tentacles afewelongaterosettes upto0.05 long;some withdrawn) has one well-developed bursal juv- body wall plates, lacking spines. enileattachedtotherightanteriorintracoelomic Live colour. Body and pedicels reddish brown; body wall, and a gonad with one large and a few- small white eggs. end rim ofpedicelsdark reddish brown; tentacle trunks brown, branches faintly reddish brown; Etymology. From of the Greek bursa (purse), colour persists in alcohol. with referencetothe intracoelomicsacsinwhich Reproduction. The holotype (6 mm long, ten- brood juveniles develop. tacles withdrawn) has a dorsal gonad, 1 brood Distribution. Rottnest I., Western Australia, to embryo and 1 brood juvenile. The gonad has 5 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 with brown eggorembryo (0.3 mm long). The brooding in intracoelomic sacs which are ran- I 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 orjuveniles may be present singly or tral tentacles reduced - does not exist in any together in the sacs. The very thin-walled sacs previouslydescribedcucumariid genusand war- lack ossicles and appear to be derived from the rants the creation ofa new taxon. Ofthe other egg rather than from the body wall. At the point genera, PseudoenusPanning, 1949 and Pseudoc- ofattachment ofthe sacs, with their developing nellaThandar, 1987,aremostsimilar. 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 maybreakdown tocreatea small papillae, and reduced cups. opening. Individuals produce few eggs but may Cueuvitrum rowei sp. nov. have all reproductive stages present, from small eggto fully-developed brood juvenile. Only one Plates lb, 3a-g, Figure 2 egg appears to mature at any one lime. All six — Stereodermasp. A.M. Clark, 1966: 346-347, fig. adults exhibit bursal coelomic brooding. All 9. — observed gonads contain eggs and are similar in Pseu—doenussp. Rowe, 1982: 466, fig. 10.32b, pi. form, suggesting hermaphroditic or patheno- 32.2. Rowe and Vail. 1982: 223. genic reproduction. The collection ofbrooding adults in January, February, March and May Material examined. Holotype. Victoria, Cape Patcrson,just below low tide level, 18 Jan 1980, M. suggests non-seasonal reproduction. NMV OcnussacculusPawson, 1983, from NewZeal- OF'5L7o3u5g6h.lin, T. O'Hara and J. Stephenson, and, hasasimilarbroodinghabit. Intracoelomic Paratypes. Victoria, type locality and date, NMV sacs are attached to the dorsal intcrradial F5424l(5); Apollo Bay, Marengo, Hayley Point, just NMV anteriorbody wall and contain upto 9 embryos. below low tide level, I 1 Jan 1980. 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 ofthe l'auna, 0-2 m, unless otherwise stated; # indicates sac has fewer ossicles than elsewhere, and Paw- material with intracoelomic copepod parasites). son (1983: 228) suggests that "birth" occurs by rupture ofthe body wall. This species is easily distinguished from P. bursatusbytheformofthe 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, irregularlyoval, somewiththeend orone 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; 10platestaperedanteriorly, 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 ofthe 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- versesectionofbody,scalebar= 1.0mm;b,radialand icles, pedicels mostly confined to radii, and ven- interradial plates from the calcareous ring. , NEW HOLOTHURIANS FROM SOUTHERN AUSTRALIA 233 WesternWAuAstMralia. Rottncst I., Strickland Bay, 20 elongate, slightly keeled midventrally, distinct Jan 199W1,AM 400-91(1 juv); Natural Jetty, 15 Jan dorsolateral and ventrolateral edges, body pen- 1991, 393-91W(1AMjuv); Cape Naturaliste. Eagle tagonal in transverse section; weakly developed Bay, 25 Feb 1975. 556-89(13). oral valves formed by anterior radial body wall South Australia. Ceduna. Cape Vivonne, 16 Jan 1991, NMV F59209(4); Strcakv Bav, Point Wcstall, projections; mouth orientated anteriorly; exten- 15 Jan 1991, NMV F5921 l(i>; Greenlv 1., 28 Nov sible anal cone, often upturned; body wall firm, S19A7M6.KS1A81M3(Kl)1:80K0a(n1g):aArronooI.B.aEv.mu22BFaevb, 11978J8a.n2-1399m0,. dcreynsdtrailtliicne,tewnittahclmesi.cro2scvoepnitcralvitorneeosusdsipsottisn;ct1l0y NMV F57550O): Gulf St Vincent, Normanville. 11 smaller; distinct modified ventrum, not a sole; Nov 1988. NMVF54386(1): Robe. 9 Jan 1990. NMV distinct introvert, lackingpedicels; 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- "NMMuVlFle5t42Ho2l1e(s"1,a6nJdan1 j1u9v8);1.1N0 MkmVNFE57o9f49A(p4o)l:loNBMayV, tral radii with up to 60 pedicels in each row; F57948(l#); 10 Jan 1987. NMV F54218(5): NMV dorsolateral series irregular, pedicels small; a 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 N38M°1V7'S. 144°38'E,3-6m.30Mar 1986.SPPSstn3B1 anteriorly and around anus. Calcareous ring F57547(6#); Portsea Jetty, 4 m. 13 Mar 1975, lacking posterior prolongations; 5 radials with AM J10857(1); Cape Schanck. Bushrangers Bay. 28 blunt anterior projections, often notched, large Mar 1981. NMV F54224(3juv); Flinders, Mushroom notch posteriorly; intcrradials with pointed seRBNMtreaansMeyyf,HV1.a51.1v3F19eN568Fn49M.e3N,b9Vo34N0v1(F0M9558)1mV38;97,9o7N0FNf2,f5(MMs42hNV3)Vo9;rM5eFCF(,V52a584)p46.;Fe24552-19PP(65h2al(i0tm5#le6).l)r(;i;sl6po1)SnM:Ikh..a,amM2rcKEe8kir1otA9nBtfp8acyH2rvk..aMs1i,Cr49l.8Pml259eA-7.-r aismtccnholateeulenltrolihrpeoufsrtt.nosMlpmaavritcdoednjrr-teaecrlcpa;otolielroliyinnosttmt,eeo,stdalwionoinruoedsspae.slr;ulsensfhsitanlbgdlalooecrwkspaotlploloiymsatoneufrvtriehoos,m-r 12.2m,4Mar 1982,CPAstn4,NMV F53771(I);Wil- Dorsal body wall ossicles of5 types: 1) Abun- sons Promontorv, Hobbs Head, 12.5 m. 9 Feb 1982. dant perforated plates, heavily knobbed, irregu- WPNPA NMV stnN42M,'V F53773(l juv); Rabbit I.. 23 larly oval, frequently thick and rounded at an Apr 1983, F53783(2); eastern Bass Strait. endoredge,theoppositesideorenddenticulate; NMV 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. narrow end denticulate; many plates with pro- F57943(N2M);VMallacoota. Bastion Point, 5 m, 6 Apr jections, bars, further developing layers, com- N19M89V. F57364F05)7.363(6); Gabo 1.. 4 m, 4 Apr 1989. monly up to0.12 mm long; closely situated in 2 N19MT8a2V,smNFa5nM7iV3a6.F1K5(5i3)n7:g8L8u1(.l,2w)Ro;orctWhka.vteBCrlaahpcoeku,sRc2ocmPk,asPs1oa5ignMet.a,r22321NN98oo8vv. otloarrm0g.eo4rm8eulmltaimy-elrlasoynwegir,tedhwiiontshstihc3el-eb5so,dpieyrrrwfeaoglrulal.ta2re)ldySkconavotaltb,ebreuedpd 1982. NMV F53789(2): Bicheno, 7 m, 22 Mar 1988. layers of decreasing size, one end sometimes NNMMVVF5F75574346(04()5:);SwaNNnsMMeVVa, 8Fk57m54S.8(42m#,);21EMaagrle1h9a8w8k, adtenetdipclualtaetse,.u3p)tAo 0.fe1w2 mirmregluolnagr. 4s)mAooftehwpierrrfeogru-- ANMEedsacpvykee,rn1at9n1u7c5r4ee,FeAPBboaMiyn.1t9J,911510.18m35.5m(21,3);AAFDpM'5rE9nJ21t92Ir926e160c.6)a5A;s(t7MTe#ia)nuJ:dx2e2BCr6rhb6uao7nnx(ny,3e#2l)I.9,;, AoslsaosrMmiceslmeNso,eotwtyhpSioocuartllhlyuW0ma.pl0ey7smosmrpmemkcnlioomnbegbn.es5d()ARboMustetJot1nt0-e8ls8i3ki,en Sadgrove Point, 7 Mar 1974. AM J 10856(1); South- J12735), upto0.04 long. Platesofven- port, Lady Bay, 3 m, 1 1 Oct 1977, AM Jl 1164(1). trum similar to dorsum; multi-layered ossicles NewSouth Wales. DisasterBav. Green Cape Light- lessdeveloped. Platesofintrovert fewer, similar house. 16 m, 13 Feb 1973, AM J12548(1); Twofold to body wall, more finely knobbed, none multi- PBoaivn,t,32m5.N2o9vMa19r841.98N5M. VAMF5J37189690(93()6;)J;erEvdiesnB,aCyo.c3omr,a plaeyrefroerda.tePdedbiacre-llsikweitoshsiicrlreesg,uloafrt,ebnewnitdoernceudrvceedn,- J2A61M0N8o8Jv3l(ll29)17;6l7(M,5aA)nM:lJvPIo.2rt594JO7acc(tkls)1o9;n7B.9o,3nAdmi,M,N2JM93-1VA257u3mg5,(12O9)c7;t7,N19aA7m7M-, atprllaaoltnleygs,ouuoptfetteron0e.dd2ge5en,mtitmcyupilwcaiatdleely,as0nm.da1l4lfemirnmepleyrlfoknongr:oatbeibnoednd-s bucca Heads, 13 m, 1 1 Jan 1972. AM F57555(l#); centrally; irregularly triangular, perforated, Coffs Harbour, 13 m, 22 Jan 1982, J15470(2). mm curved, denticulate plates, typically 0.1 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 elongate, irregular,oftenwidened centrally, per- rodsorplatespresent posteriorly.Tentacleswith forated centrally and distally, sometimes thin perforated rods;pedicelswithoutendplates. curved,often bent atcentre, upto0.26 mm long; Calcareous ring simple, without posterior pro- irregularly round to triangular, thin, perforated, longations, 10 plates, tapered anteriorly, undu- slightly convex, denticulate plates, up to lating posterior edge. Dorsal marsupia present. mm mm 0.08 wide; rosettes, typically 0.04 Type species. Neocnus incubans Cherbonnier, long. 1972. Live colour. Body predominantly white, golden Remarks. Thisgenuswaspreviouslyknownonly in largest specimen, sometimes dark decking from the type species found on the Mediter- dorsally and laterally; dark vitreous spots dor- ranean coast offTunisia. 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- Reproduction. Some sac-like gonad caeca and tacles, four polian vesicles, rare or numerous, gonoducts have been observed in the right knobbedorsmooth perforated platesin thebody 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 ofthe 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, echinoderm research in Australia. 1938. found off Western Australia, is also Distribution. Rottnest I., Western Australia, to characterized by an almost total lack ofossicles. Coffs Harbour, New South Wales, including However, it differs in having a distinctive cal- Tasmania. 0-28 m. careousring, numerousrosettesaswellasrodsin the tentacles, no sole, and numerous large pedi- Remarks. One, rarelytwo,copepod parasitesare cels with well-formed endplates covering the sometimes present in the coelom of C. rowei whole body. specimens from Victoria, New South Wales and Tasmania (see # in Material examined forexact 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- — maricolidae. They are long and narrow, some- Neocnus sp. -- Rowe and Vail. 1982: 224. timesextendingmore than halfthe length ofthe tn.oughlin. 1991: 223-225, figs 2, 3. coelom, and carrying numbers ofyellow eggs. Material examined. Holotype. Victoria, Apollo Bay, Rosetteossiclesare present in thebodywallof Marengo, Hayley Point,just below low tide level, 29 sspoemceimmeantserairael oftrhoemrwiNseewsiSmoiultahr tWoaltehso.seThferosme DePear1a9l7v4p.esM..TOy'pLeoulgohcalliint.y,NNMMVVFF5544223389.(120): AM otherregions,and tospecimensfrom NewSouth .122752(20). Waleswithout rosettes, and arethus included in epiOftahuenra,m0a-te2rima,lu(nplaerstsiaolthliesrt;wiaslel sctoaltleedc)t.ed as algal the current species. South Australia, Robe. 10 Jan 1988, NMV Most ofthe specimens have been collected as F54183(40). algal epifauna and arc juveniles. A few adults Victoria, ("ape Bridgevvater, 20 .Ian 1979, NMV havebeen found attached tothe undersurfaceof F53806(l): Cape Otway. Crayfish Bay, 31 Dec 1980. NMV rocks. F53814(47); Apollo Bay. Marengo, Havlev NMV Point, 11 Jan 1980. F53808(75); Flinders, cast Neocnus Cherbonnier, 1972 of Mushroom Reef, 13 Jul 1990, NMV F58630(7); NMV DBioadgynosshiosrt(aenmdensdteoudtfwriothm dCihsetrinbcotnnsioleer;,1019d7e2n).- oSfhoPrycrhaammi.d1R8ocOkc.t21298D0e,cNM19V80,FN53M8V13(F45)3:8P1h0i(lll)ip:].K,ilE- cunda, 26 Jan 1987, F53805(13); Manners dfriinteidc tteontpaecrleisp,hevreyntraanld2msimda-lvleenrt;rpaeldircaedlisucsono-f H15a,veNn.M3V00Fm537ol8l1'(shlo)r;e,C4a.p5-e6Pma,te6rMsoanr. 129982J.aCnP1A98s8t.n sole. Body wall ossicles rare, a few perforated NMV F54I82(8). NEW HOLOTHURIANS FROM SOUTHERN AUSTRALIA 235 Figure 3. Neocnusbimarsupiissp. nov.: a, dorsal body view, showingpaired anteriormarsupia; b, ventral body view, showingextensible ventral tentacles; c, diagram ofa marsupium, showing interiordorsal pedicels, with 4 extending through opening, scale bar= 0.5 mm; d, transverse section ofbody, 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 Description. Up to 6 mm long, 3 mm high, 2.5 muscle band; each pouch with central aperture; mm wide; body roughly cgg-shapcd; tentacles seriesofupto 16 verysmall pedicelswithin each pouch, may extend out of the pouch aperture. orientated anteriorly or slightly upturned, anus Eachbrood pouch with upto 14embryosorjuv- posterodorsal; 2 dorsal brood pouches in anteriorbodywallexcept injuveniles, each with enilems,muniform in size, differentiation evidenmtmat 0.7 long; brood juveniles up to 1.5 external round aperture, andsmall interiorpedi- cels which may extend through the aperture; long. 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 mm 3.0 long, with a pedicel-like attachingcapa- the two brood pouches. bility; distinct sole; introvert not distinct from Distribution. Robe, South Australia, to Cape body wall. Sole with peripheral row ofup 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 and anus; Remarks. This species is similar toAr. ineubans. mostly no pedicels on lateral or dorsal surfaces, Both are small and epiphytic, brood theiryoung except somewithin thedorsal marsupia and fre- in marsupia intheanteriordorsal bodywall,and quently single small radial pedicels anteriorly, have tentacle rods as the main ossicle form. N. most prominent dorsolatcrally; up to 5 small ineubansdiffers in havingsmall irregularperfor- pedicels around anus. Calcareous ring lacking ated platesaswellasrodsin thetentacles, rodsof 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 upto 30eggsorembryos 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 eggsorembryos from thegonad tothe marsupia in dorsal, anal body wall; sole lacking ossicles. isnotknown. With theextensibleattaching ven- Pedicels lacking endplates orossicles. Tentacles tral tentacles, and the small suckered pedicels with abundant thin straight and curved rods, extending through the marsupium opening, mm typically 0.08 long; ends of rods swollen, there isacapacityfortransferofeggsorembryos with 1-6 small holes or notches, divided end of from gonopore to marsupium. Numbers ofeggs rod often twisted together; rods rarely with side in a maturegonad sac, and ofbroodjuveniles in branches or forks. Ossicles similar in juveniles. a marsupium, are similar. If there is a single Live colour. Body dark to very dark grey or action oftransfer from one mature gonad sac to brown; sole, pedicels, tentacles to varying onemarsupium,theactionwould be remarkably degrees lighter in colour; colour persists in efficient. Since no brood juveniles with fully developed tentaclesand mouth havebeen found alcohol. in the marsupia, it appears that broodjuveniles 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 broodjuveniles; 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, softbuttough,ovalprotuberanceon developed; largest caeca lieagainstwall ofbrood thealgal fronds. Its relatively large ventral pedi- pouch, each caecum with up to 13 large yellow cels enable it to cling strongly to its substrate. eggsorembryosanda fewsmallwhiteeggsinter- spersed; large eggs or embryos frequently 0.6 Squamocnus gen. nov. mm mm long, upto 1.2 long;smallcaecawith up Diagnosis. 10 dendritic tentacles, ventral 2 to 15 small white eggs. Short anterior dorsal smaller; 5 oral valves; pedicels on ventral radii gonoduct;gonoporebetween basesofdorsalten- and scattered on dorsal radii and interradii. tacle pair. Two separate anterior dorsal brood Bodywallossiclescups, knobbedperforatedbut-

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