MemoirsofMuseumVictoria62(1):91-99(2005) ISSN 1447-2546(Print) 1447-2554(On-line) http://www.museum.vic.gov.au/memoirs/index.asp Two new Middle Miocene spatangoids (Echinoidea) from the Murray Basin, SouthAustralia FrancisC. Holmes1 ChristopherAhYeeandJaniceKrause2 , ‘15KenbryRoad,Heathmont,Victoria3135,Australia,andInvertebratePalaeontology,MuseumVictoria,POBox666, Melbourne,Victoria3001,Australia([email protected]) 2P.O.Box581,Hamilton,Victoria3300,Australia Abstract Holmes,F.C.,AhYee,C.,andKrause,J. 2005.TwonewMiddleMiocenespatangoids(Echinoidea)fromtheMurray Basin,SouthAustralia.MemoirsofMuseumVictoria62(1):91-99. TwonewspatangoidtaxaaredescribedfromtheGlenforslanFormationcroppingoutintheMurrayRivercliffsnear Blanchetown, SouthAustralia. Onetaxon,Murraypneustesbiannulatusgen.etsp.nov., alargespeciesofspatangoid withtwo ‘peripetalous’fascioles (onecirclingthemarginandtheotherclosetothedistalendoftherelativelyshort petals),twodistinctsizesofaboralprimarytubercles,andadepressedapicalsystem.Theotherspatangoiddescribed, Spatagobrissusdermodyorumsp.nov.differsfromtheonlyotherfossilspeciesofthisgenusrecordedfromAustralia, S.laubei(Duncan, 1877),inhavingamuchshorterlabrum,markedlylargerperistomeandperiproctandlargerprimary tubercleswithintheperipetalousfasciole. Keywords Echinoidea,Spatangoida,Murrraypneustes,Spatagobrissus,newtaxa,MiddleMiocene,SouthAustralia Introduction Palaeontologycollection,MuseumVictoria(NMV).Inaddition to type material, the new species ofSpatagobrissus is repre- OfalltheAustralianTertiary sedimentarysequencesthatcon- sented by several specimens in private collections. Measure- tain extensive echinoid faunas, the 150 km ofMurray River mentsweremadewithadialcallipertoanaccuracyof0.1mm. cliffs, from Overland Comer, east of Waikerie, to Murray Parametersareexpressedasapercentageoftestlength(%TL) Bridge,SouthAustralia,haveoverthelast120yearsbeenmore ortestwidth(%TW). comprehensively studied by palaeontologists, students and amateurcollectorsthananyothersimilararea.IntheMiocene Ageandstratigraphy stratigraphic sequences alongtheMurrayRiverandelsewhere The Glenforslan Formation, synonymous with the Lower inAustralia, species belonging to the Spatangoida constitute Morganlimestone,conformablyoverliestheFinnissFormation approximately 50 percent of the recorded taxa of irregular echinoids. Inview ofthenumberofspatangoids occurring in and is ofearly MiddleMiocene (Batesfordian, Langian) agem. thissectionoftheMurrayRivercliffs,thediscoveryin2003of The thickness ofthe unit is relatively consistent at 13-15 threespecimensofanewgenusbelongingtothisorder, seem- although this is reduced in southern exposure due to post- inglyunrelatedtoanyothergenusknownfromthecontinent’s MiddleMioceneupliftandsubsequenterosion.Theformation fossilrecord,wasunexpected. is sublithifiedtolithified, weatheringtowhitish-creamcolour Thespecimenswerefoundwithin300mofeachotherina inoutcrop(LukasikandJames, 1998).Itiscomposedofcycles singlebedofthe GlenforslanFormation, cropping outonthe ofmollusc-bryozoan floatstone, with a microbioclastic pack- left bank of the Murray River, 7 km NNE of Blanchetown, stonematrix, gradingupwardinto Celleporariarudstone tops South Australia [Museum Victoria locality PL3203]. Further (infaunal bivalve and gastropod-rich microbioclastic matrix with large branching and sheeted Celleporaria) which also investigation ofthe surface exposure ofthe formation in the general vicinity of this discovery failed to produce any containpectens and oysters. Echinoids tendto befound ator additionalspecimensofthenewgenus. abovetherudstone-floatstonecontactatthebaseofthecycles, whiMcahtetrhiealsstuadnidesmeartehobdass.edS,peacreimheonusneudmbienrsthpereIfnivxeerdtePb,raotne bthreyzloaattnesrocfon1t-a3incimngldeenlgithc.atSeebdriamnecnhtisngareanpderuvnais-ilvaemliynmaorttslheede,t obscuring all physical sedimentary textures. The middle i 92 FrancisC.Holmes,ChristopherAhYeeandJaniceKrause Glenforslan Formation is interpreted as being deposited in peripetalous fasciole clear ofthe distal ends ofpairedpetals, relativelyshallowwaters,possiblylessthan10m,basedonthe clearlydistinguishthenewgenusMurraypneustesfromvirtu- presence of calcareous algae and mixotrophic foraminifers allyallothertaxaassignedtotheMicrasterina,inparticularthe (Lakasik,pers. com. 2005). Itformspartoftherichestwarm- 22generaincludedintheAsterostomatidaeFisher, 1966.Only waterbioticrecordfromsouthernAustraliaatatimeofmaxi- one of these genera, Asterostoma Agassiz, 1847, has tenta- mum transgression of the sea across the continental shelf tively been retained in the Asterostomatidae (together with (McGowranandLi, 1994,andpaperscitedtherein). Stomaporus Cotteau, 1888, agenus originally assignedtothe Thethreespecimensofthenewgenuswerefoundabout7.4 BrissidaeGray, 1855)bySmithetal.(2003).Oftheremaining mabovethebase oftheformation, approximately 2mabove 21 genera, Smith et al. have conditionally reassigned 13 to theLepidocyclinaZone. other families, only six of which are included in the Micrasterina.Theremainingeightarelistedasincertaesedisor Associatedfauna asbelongingtoanunnamedtaxon.Untiladetailedrevisionof these lattergenerais published, itis considered imprudentto NGlienneftoeresnlasnpeFcoiremsatoifonecwhiitnhoiinds50h0amveupbseternearmecaonrddeddowfnrsotmretahme assignthenewgenustoaspecificfamily. of PL3203 (Table 1), compared with 28 confirmed species knowntooccurwithintheMorganGroup(Glenforslan,Cadell, Murraypneustesgen.nov. andBryantCreekFormations). Typeandonlyknownspecies. Murraypneustesbiannulatussp. nov. Table 1. Echinoids recordedwithin the vicinity oflocalityPL3203. Letters in brackets indicate the frequency of occurrence of each Diagnosis. Large ovoid spatangoid with centrally depressed species.[A],abundant;[C],common;[F],fairlycommon;[U],uncom- adapical surface, apexwellposteriorofcentre.Apical system mon; [R],rare. Referencestoauthorsandsupportingliteraturecited ethmolyticwith4gonopores.Aboralprimarytuberclesof2dis- below, but not listed in the main textreferences, can be found in tinct sizes, smallandrandomly spaced, thelargerproximalto Holmes(1993). the ambitus. Labrum long, narrow, partially tuberculate, Cidaroida extending to third ambulacral plate. Two ‘peripetalous’ GoniocidarismurrayensisChapmanandCudmore, 1934[C] fascioles present; one marginal, passing above the periproct Arbacoida (pseudolateral); the second, rudimentary, non re-entrant and Murrayechinuspaucituberculatus(Gregory,1890)[F] clearofthedistalendsofpairedpetals.Intermittenthorizontal Temnopleuroida fasciolebandsalsooccurbetweenthe2‘peripetalous’fascioles. Cryptechinushumilior(Bittner, 1892)[C] Subanalfascioleincontactwithmarginalperiproct. OrtholophusmorganensisPhilip,1969[C] O.pulchellus(Bittner, 1892)[C] Etymology.FortheMurrayRivercliffs,theoriginofthefossils, Clypeasteroida and “pneustes”, a common suffix used for spatangoid MonostychiaaustralisLaube, 1869[A] echinoids.Gendermasculine. M.sp. ‘C’inHolmes, 1999[F] Remarks.Thefollowinganalysisofmorphologicalfeaturesof Scutellinoidespatella(Tate, 1891)[C] Spatangoida generasimilartoMurraypneustes gen. nov. isbasedprimarily BrissopsistateiHall, 1907[U] on Mortensen (1950a) and Smith et al. (2003). Although Brissussp.nov?[R] approximately 30 genera within the Micrasterina have been Cyclasterarcher (TenisonWoods, 1867)[C] investigated, only eight warranted further scrutiny; four cur- EupatagusrotundusDuncan, 1877[U] rently unassigned to a family by Smith et al. (2003), Eupatagussp.indet.[U] Elipneustes Koehler, 1914, Eurypatagus Mortensen, 1948, Hemiaster(Bolbaster)planedeclivisGregory, 1890[C] LinopneustesA.Agassiz, 1881,andPlatybrissusGrube, 1865; Loveniacf.forbesi(TenisonWoods, 1862)[C] threebelongingtotheMaretiidae(Lambert, 1905),Eupatagus Murraypneustesbiannulatusgen.etsp.nov.[R] L. Agassiz, 1847, Mazzettia Lambert and Thiery, 1915 and Pericosmuscompressus(Duncan, 1877)[R] Protenasterantiaustralis(Tate, 1885)[U] SpatagobrissusH.L.Clark, 1923;andMacropneustidaegenus Spatagobrissusdermodyorumsp.nov[F] LajanasterLambertandSanchezRoig, 1924. Using a broad comparison of 38 features, Linopneustes standsoutfromtheothersevengeneraashavingclosestaffin- SystematicPalaeontology ity with Murraypneustes; four of its recorded six species, L. longispinus (A. Agassiz, 1878), L. fragilis (de Meijere, OrderSpatangoidaClaus, 1876 1903),L. spectabilis (deMeijere, 1903), andL. brachipetalus SuborderMicrasterinaFisher, 1966 Mortensen, 1950b having distinct marginal peripetalous fascioles passing above the periproct. In the other two, FamilyIncertaesedis L. murrayi (A.Agassiz, 1879) andL. excentricus deMeijere, Remarks.Thecombinationofgenericfeatures,particularlythe 1903, the peripetalous fasciole is not marginal but somewhat presenceofamarginal ‘peripetalous’fasciole,intermittenthor- higher up on the test (Mortensen, 1950a: 221). Although izontal fasciole bands and a rudimentary non re-entrant L. fragilis has multiple fasciole bands around the ambitus, TwonewmiddleMiocenespatangoids 93 L. murrayi a rounded margin (Smith, pers. com. 2005) and MurrayRiverLock 1, Blanchetown, SouthAustralia [NMVlocality L. longispinus relatively shortclosingpetals; Murmypneustes PL3203]. isdistinguishedbyitscentrallydepressedaboralsurfaceform- Paratypes,NMYP312371andP312372fromthesamelocation. ingfourapices(domedonLinopneustes)withthehighestpoint Diagnosis.Asforgenus. ofthetestposteriortotheapicaldisk(anteriorinLinopneustes), the presence of two ‘peripetalous’ fascioles and intermittent Description.Testmoderatelylarge,ovoidinoutlinewithshal- hsopraiczeodntaalbofraaslciporleimbaarnydst,ubtewrocldeiss,titnhcetsliazregserofressmtarlilctreadntdoomtlhye lSopwecianmteenrsiorrasnuglecufsroamnd72potiont8e1d,mmsliignhtlleyngttrhu,ncwaittehdmpaosxtiermiourm. areaoutsidetheupper ‘peripetalous’fasciole,andaprominent width 81-86%TLoccurring at45%TLfromanteriorambitus. angularsubanalfasciole. Test44.1%TLhigh(uncompressedspecimen)withapexofall Basedonthesamecriteria,threeothergenerashowamod- specimens well posterior ofcentre, 63-67%TLfrom anterior erate degree of morphological similarity to Murraypneustes, ambitus. namelyElipneustes,Mazzettia,andEupatagus. Centreoftest on adapical surfacedepressedaround apical Elipneustes,however,canbedistinguishedbyitsverylong, diskandproximalendofpairedpetalstoformminorapicesin parallel-sided, open-ended, flush petals with conjugate pore interambulacra 1,4and5, andconjointly, araisedareaacross pairs;single,narrow,marginallysituatedperipetalousfasciole; ambulacrum III and interambulcra 2 and 3. Adoral surface peristome situated immediately below the apical disk; small mildly concave in vicinity ofperistome with ambulacrum III thickset plastron with minimal posterior swelling; and much slightly recessed anteriorly and the plastron forming a fairly larger primary tubercles scattered over all the aboral inter- pronouncedkeelposteriorly,terminatingattheanterioredgeof ambulacra. thesubanalfasciole. Mazzettia, a poorly known fossil genus, has an elongated Primary tubercles on adapical surface of 2 distinct sizes, heart-shapedtestwithalow sharpmargin, verylongweakly- bothwidelyandrandomlyspaced.Largerofthetworestricted bowedpetalsclosingdistally,norecordedperipetalousfasciole tothedistal35%oftheradiusoninterambulacrum5,and25% and only occasionally a weak shield-shaped subanal fasciole, elsewhere. Adorally, larger primary tubercles closely and anelongatedlabrumjustreachingtherelativelysmallplastron, evenly spaced throughout, with exception of naked areas in andrandombutcloselyspacedcoarsetuberclesaborally. ambulacraIandVandphyllodeplatesofambulacraII,IIIand Eupatagus, although possessing well developed peri- IV. Overlapping scrobicules on adoral surface form distinct petalous andsubanalfasciolesiseasilycategorisedbyitslack diagonal ridges. Small tubercles closely spaced immediately of an anterior sulcus, predominately short lanceolate closed below ambitus increase in size to that of larger primary petals, primary tubercles of varying density restricted to the tuberclesbelowcurvatureofmargin. Largeprimarytubercles, area within the peripetalous fasciole in aboral interlambracra perforatewithundercutmamelonandcrenulatedplatform,and 1-4, andvery small evenly spaced tubercles distally overthe maximum scrobicular diameter of approximately 2.5 mm, remainderofthe aboralsurface. Comparisonofotherfeatures twicesizeofsmallercounterparts.Ringofscrobiculartubercles isdifficultbecauseofvariabilityamongtheverylargenumber notalwayspresent. ofdescribed species. In particular, the eightAustralian fossil ‘Peripetalous’ and subanal fascioles present, the former, speciesvaryconsiderablyinprofile,aregenerallymoreround- though not continuously visible on any specimen clearly ed, and have an extremely wide range of primary tubercle passesaboveperiproct,whilelatterisdistinctlyangular,form- densities when compared with the type species on which the ing hexagonal outline where in contact with lower edge of genericdescriptionisbased. periproct. Intermittent horizontal fasciole bands also present Twoothergenera,PlatybrissusandEurypatagus,aredistin- laterallyaboveandparalleltomarginal ‘peripetalous’fasciole. guishedbytheircompletelackoffascioles,althoughtheformer Uppermost very narrow and more continuous fasciole, mayhaveasubanalfasciolepresentinjuvenilespecimens. although clear ofdistal ends ofpetals, appears to be a rudi- Lajanaster, although depressed aborally, has no anterior mentaryperipetalusfasciolenotindentedinterradially(Fig.3). sulcus,arelativelysharpambitus,amarkedlynarrowplaston, Apicalsystemanteriorofcentre,39.5-42.2%TLfromanter- and primary tubercles confined to the posterior column of iorambitustocentre ofdisk,levelorslightlybelowproximal anterior and lateral interambulacra within the peripetalous end ofpairedpetals. Ethmolyticm,mwith4 small closely spaced fasciole. gonopores approximately 0.3 in diameter, anterior pair Spatagobrissus was included in the comparative analysis, closer together than posterior pair. Detail of ocular plates primarily as a new species of the genus occurs at the same indeterminate. Hydropores numerous, approximately 70 locality(PL3203)asMurraypneustes.Inmostrespectsthefor- visible in one specimen, centrally located but extending merisverysimilartoEupatagusbuthasshorterpetalsandonly betweenposteriorpairofgonoporesandpossiblyocularplates smalltuberclesoverthewholeoftheaboralsurface. IandV(Fig.4A,B). Petalslanceolate,moderatelywideatmidlength,closeddis- Murraypneustesbiannulatus. sp.nov. tally, situated in gentle concave depressions incorporating adradialedgesofadjacentinterambulacraandcontinuingprox- Figures2A-F,3A-C,4A,B,5 imally across apical disk. Anteriorpaired petals shorter than Typematerial.Holotype,NMYP312370fromearlyMiddleMiocene posteriorpair, extending on average 60% ofthe radius meas- GlenforslanFormation(Batesfordian),MorganGroup,7kmNNEof ured along the surface ofthe perradial suture from centre of 94 FrancisC.Holmes,ChristopherAhYeeandJaniceKrause apicaldiskto ambitus;posteriorpairabout56%radius. Inner transverselyabovewell-roundedmarginwithambitus situated poresofpetals oval, outerpores slot-like, slightlycurvedand atabout30%TH.Adoral surfaceverymildlyconvexbutwith 50%wider.Porepairsnotconjugatebutlinkedbyafineridge prominentposteriorkeel causedby sharp rise ofambulacraI whichextendsalongbothsidesofeachpore(Fig.5).Maximum andVposteriortocentre. Inlateralview, posteriortruncation widthofinterporiferouszoneslightlymorethantwicewidthof coversabout35%TH. poriferous zone. Anterior paired petals diverge at approxim- Apical system ethmolytic with 4 genital pores, centre ately 135° and contain on average 23 pore pairs, posterior 31.3-38.6%TLfrom anteriorambitus (Fig. 7A). Pairedpetals petals297°and26pairs.Secondarytuberclesextendrandomly short (26.5-32.5%TL measured along surface of perradial acrossinterporiferous andporiferous zones andforadistance suturefromcentreofapicaldisk),narrow(7.0-9.0%TL),lance- outsidepetals withoutprimary tubercles. AmbulacrumIIInot olate, closing to closed,posteriorpairmarginally longerthan petaloid, basically flush with adjoining interambulacra for anteriorpair.Anteriorpairdivergeatabout 135°,posteriorpair about 50% radius, then gradually becoming concave towards 310°.Porepairsconjugate,innerporesoval,outer-tearshaped. anterior sulcus. Other details unknown, no sign of pores or Anteriorrow ofpore pairs in anteriorpairedpetals distinctly regularlyspacedtuberclesbeingvisibleonspecimens. narrower than posterior row and atrophied adapically. Peristome reniform, centre situated 27-30%TL from Interporiferouszoneupto1.5timeswidthofporiferouszoneat anterior ambitus, longitudinal dimension approximately widestpoint.AmbulacrumIII flush adapially, with 2 rows of 6.5%TL,transversedimension12%TL.Phyllodesshortandnot indistinctlongitudinallyorientatedporepairsandinterporifer- particularlywelldeveloped. ous zone containing few secondary tubercles and numerous Labrum long and narrow, averaging 20.5%TL, slightly miliaries. curvedatjunctionwithperistome andabuttingtheplastron at Peripetalous fasciole subcircular, not indented. Numerous centre ofthird pair of adjacent ambulacral plates. Numerous small,randomlyspacedperforate,crenulate,primarytubercles small tubercles adjacent to peristome with a few larger ones occur in interambulacra within fasciole. Outside fasciole, towardstheposteriorend(Fig.2C). tubercles in posterior halfof test very small, but anterior to Plastron closely tuberculate, width approximately 75% centre, gradually increasing in size towards interambulacra lengthmeasuredfromposterioredgeoflabrumtoanterioredge 2and3. ofsubanal fasciole. Strong posteriortaper suggests sixth and Peristome reniform, mildly sunken, width 16.0-19.3%TL, subsequent plates ofambulacraI and V indent behindpaired length8.9-10.3%TLwithanteriorborder23.4-26.8%TLfrom episternalplates. anterior ambitus. Phyllodes moderately developed with slot- Periproct opening marginal, not visible from above, tear shapedporesincirculardepressions.Labrumshortandwedge- shaped, slightly wider than high, set in a slight truncation shaped extending only to centre of second pair of adjacent approximately65°tothehorizontal. ambulacral plates. Anterior edge raised above surrounding Etymology, biannulatus(L)-two-ringed,referringtothepres- ambulacra and slightly projecting over peristome. Miliary tuberclespresentwithfewsecondarytuberclesinanteriorhalf enceoftwo ‘peripetalous’fasciolerings. (Fig.7B). Plastronlong,widthabout55-60%length,withambulacral FamilyMaretiidae platesindentingposteriorly. Subanalfasciolecirculartotrans- SpatagobrissusH.L.Clark, 1923 versely oval, enclosing 3 pore pairs each side of interradial suture, and with slight anteriorprojection atposterior end of Type species. Spatagobrissus mirabilis H. L. Clark, 1923, by prominent plastronal keel. Posterior edge offasciole margin- originaldesignation. ally clear ofperiproct opening. Adorally, ambulacra I and V relatively wide, covered with very fine, randomly spaced Diagnosis. SeeH.L.Clark(1923:402) miliarytuberclesuptosixthplate,thenbytuberclesofsimilar Spatagobrissusdermodyorumsp.nov sizetorestofadoralsurface. Periproct, tear-drop shaped, generally positionedvertically Figures6A-E,7A-C, 8A-E on truncated posterior margin but slightly visible from TypeMaterial.Holotype.NMVP312570fromearlyMiddleMiocene above on some specimens; height 17.1-19.1%TL, width GlenforslanFormation(Batesfordian),MorganGroup,inthevicinity 13.2-16.0%TL. BoflanNcMheVtowlnoc,alSiotuythPLAu3s2t0r3a,lia7. km NNE of Murray River Lock 1, Etymology.NamedforMichaelandMarieDermody,ownersof Paratypes, NMVP312571-P312373 from the same general area. GlenforslanStation. OthermaterialusedforstatisticalpurposesisheldinMuseumVictoria Remarks. Spatagobrissus dermodyorum sp. nov. differs pri- andprivatecollections. marily from the Middle Miocene Port Campbell Limestone Description. Test small, subcircular in outline with minimal species, Spatagobrissus laubei (Duncan, 1877), in having posterior truncation and no anterior sulcus. Specimens range narrowertestwithmore posteriormaximum width(Fig. 8A), 30.0-45.5 mm in length with maximum width 82-90%TL markedlylargerperistomeandperiproct(Fig.8D,E),andvery occurring 51-58 %TL from anterior ambitus. Maximum much shorter labrum (Fig. 8B). In addition, anterior paired height 49.5-60%TL at 52.6-65.5%TL from anterior petals are shorter and posteriorpetals longer (Fig. 8C), with ambitus. Adapical surface moderately inflated, evenly curved divergentangleoflattergreaterthanS. laubei.Aboralprimary TwonewmiddleMiocenespatangoids 95 tubercleslargerwithinperipetalousfascioleandinterambulacra Brissus, Cyclaster,Lovenia andPericosmus, occurintropical 2and3, whilefinetuberclesoutsidefascioleonposteriorhalf waters, andthree, Eupatagus, ProtenasterandSpatagobrissus oftestare muchsmallerindiameter.Adorally, plastronwider intemperatewaters (RoweandGates, 1995). Speciesofthese andlonger,andambulacraIandVnarrower. eightextant genera, withthepossible exceptionofCyclaster, TheextanttypespeciesSpatagobrissusmirabilisischarac- areknowntooccuratdepthsoflessthan20minAustralianor terisedby alarger(upto 110mmlong) andless inflatedtest, New Zealand waters. This depthrange is consistent with the moreposteriorlylocatedapical systeminlinewithmaximum sedimentary deposition ofthe GlenforslanFormation. On the width, greater area enclosed by peripetalous fasciole, shorter other hand, Indo-Pacific species of Linopneustes, including peristome, and periproct situated on an obliquely truncated L. brachypetalus, arefoundatdepthsexceeding270m(range surface below the ambitus. Primary tubercles within the 272-1788m),withonlytheWestIndiesspecies,L.longispinus, peripetalous fasciole ofS. mirabilis are also larger and more extending up into sublitoral waters (70-570 m) (Mortensen, closelyspacedthaninS. dermodyorum. 1950a). Spatagobrissus incus Baker and Rowe, 1990, an extant Basedontheavailableevidence,itisreasonabletoassume species endemic to southeast Australian waters, particularly Murraypneustesdermodyiwasabenthicfilterfeederinhabiting betweenFlindersIsland(Tasmania)andwesternSpencerGulf, relativelyshallow,warm(?sub-tropical)waters. SouthAustralia, has alargerandmoreroundedtest(upto 80 mmlong)and, similartoS. mirabilis,moreposteriorlylocated Acknowledgements apical system and greater area enclosed by peripetalous fasciole. WeareindebtedtoDavidHolloway(InvertebratePalaeontol- Compared with S. dermodyorum, it also has a much ogy,MuseumVictoria)forvaluableadviceandsupportduring wider and longer plastron and narrower adoral ambulacra thepreparationofthismanuscript, andtoKennethMcNamara I and V. Miskelly (2002: 156) noted two pairs of pore (WesternAustraliaMuseum),RichMooi(CaliforniaAcademy pairs occurineachsideofthe subanalfasciole, afeature also of Sciences) and Andrew Smith (Natural History Museum, recorded for S. laubei (McNamara et al., 1986: 80). This London),forsuggestingimprovements tothemanuscript.Jeff contrasts with the three pairs found on S. dermodyorum Lgruakpahsiicki(nPfeotrrmoa-tCiaonnadaan)disAsthhlanekyeMdifsokreplrloyvi(dBilnagckshpeeactihf,icNstSraWti)- (Fig. 1C). fordetailsofextantechinoidsfromAustralianwaters.Michael and Marie Dermody and Donald and Miriam Griffen Discussion (Blanchetown District, South Australia) are also thanked for ThemajordiagnosticfeaturesofMurraypneustes,particularly permissiontocollectontheirproperties.Asalways,ValHogan thepositionofthetwo ‘peripetalous’fascioles,theintermittent and Sandra Winchester (Library, Museum Victoria) were horizontal fasciole bands, and the random pattern and separ- helpfulwithaccesstoreferences. ation of two distinct sizes of primary tubercles across the aboral surface, give little indication ofits lineage. Certainly, References withintheAustralianCenozoicsequencesthatpredatetheearly Agassiz,A. 1878.ReportontheEchiniinNo.9(2)—Reportsonthe Middle Miocene Glenforslan Formation, all the 13 recorded resultsofdredging,underthesupervisionofAlexanderAgassiz,in generaofMicrasterinahavetheirperipetalousfasciolecloseor the GulfofMexico, by the United States Coast Survey Steamer in contact with the paired petals. Only the brissid Cyclaster “Blake”, Lieutenant-Commander C. D. Sigsbee, U. S. N., com- is recorded as having developed multiple fascioles in some manding.BulletinoftheMuseumofComparativeZoology,Harvard specimens(McNamaraetal., 1986: 68). University5: 185-196,pis 1-5. Similarly, there are no extant genera in Australian Agassiz,A. 1881.ReportontheEchinoidea.Reportsonthescientific waters that have any specific combination of characters that results of the voyage of H.M.S. Challenger during the years link them to Murraypneustes. Even the extant species 1873-76,Zoology3(9):1-321,45pis. Linopneustes brachipetalus, found off the east coast of Agassiz,L.in:Agassiz,L.,andDesor,E.1847.Catalogueraisonnedes families, desgenresetdesespecesdelaclassedesechinoderms. Australia, has long petals in contact with its marginal AnnalesdesSciencesNaturelles7: 129-168. peripetalousfasciole. Baker,A.L.,andRowe,F.W.E. 1990.Atelostomatidseaurchinsfrom The lack of anyjuvenile specimens ofthe new genus, or Australia and New Zealand waters (Echinoidea: Cassiduloida, indeed any marked variation in the size of the three known Holasteroida, Spatangoida, Neolampadoida). Invertebrate specimens, precludes any useful comment on the disposition Taxonomy4:281-316. andspecificfunctionofits somewhatunusualarrangementof Clark,H.L.1923.TheechinodermfaunaofSouthAfrica.Annalsofthe fascioles,assuchdevelopmenttakesplaceataveryearlystage SouthAfricanMuseum113:221^435. ofontogeny. Claus,C.F.W. 1876.GrundzugederZoologie.Thirdedition.Marburg Excluding Murraypneustes, only Hemiaster, of the nine andLeipzig,xii+1254pp. spatangoid genera known to occur in the Glenforslan CottBeaarut,heGl.emy18e7t5.AnDgeusilclrai.ptKiuonngldiegsaeScvheinnisdkeasVteetretinasrkieaspsdaeksadleimeisenSst Formation sequence which embraces locality PL3203 (Table Handlingar13(6): 1-48. 1),hasnoextantrecordinAustralianwaters.Theeightremain- Cotteau, G. 1888. Echinides eocenes de la province d’ Alicante ing generaaretoday almostexclusivelybenthic filterfeeders, (Espagne). Compte Rendu de VAcademie des Sciences Paris living inshore or on the continental shelf. Five, Brissopsis, 107(25):976-978. 96 FrancisC. Holmes,ChristopherAhYeeandJaniceKrause Duncan, P.M., 1877. On the Echinodermata of the Australian McGowran, B., and Li, Q. 1994. Miocene oscillation in southern Cainozoic(Tertiary)deposits.QuarterlyJournaloftheGeological Australia.RecordsoftheSouthAustralianMuseum27: 197-212. SocietyofLondon33(1):42-73,pis3,4. McNamara,K.J.,Philip,G.M.,andKruse,P.D. 1986.Tertiarybrissid Fell, H.B. 1963. New genera of Tertiary echinoids from Victoria, echinoidsofsouthernAustralia.Alcheringa10:55-84. Australia. Memoirs of the National Museum of Victoria 26: Meijere,J.C.H.de1903.VorlaufigeBeschreibungderneuen,durchdie 211-217. Siboga-Expedition gesamelten Echiniden. Tijdschrift der Fisher,A.G.1996.Spatangoids.Pp.U543-U628in:Moore,R.C.(ed.), NederlandscheDierkundigeVereeniging(ser.2)8: 1-16. Treatise on Invertebrate Paleontology Part U, Echinodermata 3. Miskelly, A. 2002. Sea urchins ofAustralia and the Indo-Pacific. GeologicalSocietyofAmericaandUniversityofKansasPress. CapricorniaPublications:Sydney. 179pp. Gray,J.E. 1855.CatalogueoftheRecentEchinida,orseaeggs, inthe Mortensen,T. 1948.ReportontheEchinoideacollectedbytheUnited collectionoftheBritishMuseum1:EchinidaIrregularia.Woodfall States Fisheries Steamer “Albatross” during the Philippine andKinder:Eondon.69pp.,6pis. Expedition, 1907-1910. Part 3. The Echinoneidae, Grube,A.E. 1865. Einige neue Seesterne des hiesigenzoologischen Echinolampadidae, Clypeasteridae, Arachnoididae, Laganidae, Museums. Jahresbericht der Schlesischen Gesellschaft fiir Fibulariidae, Urchinidae, Echinocorythidae, Palaeostomatidae, vaterlandischeCultur1865:35-37. Micrasteridae, Palaeopneustidae, Hemiasteridae and Spatanidae. Holmes,F.C.1993.Australianfossilechinoids:annotatedbibliography BulletinoftheUnitedStatesNationalMuseum100(14):93-140. andlistofspecies.OccasionalPapersoftheMuseumofVictoria6: Mortensen, T. 1950a. A monograph of the Echinoidea 5(1). 27-54. Spatangoida1. Protosternata, Meriodosternata,Amphisternata1. Koehler,R. 1914.Rectification.ZoologischerAnzeiger44: 191. Palaeopneustidae, Palaeostomaridae, Aeropsidae, Toxasteridae, Lambert,J. 1905.Echinideseoceniquesdel’Audeetdel’Herault.Pp. Micrasteridae, Hemiasteridae. C.A. Reitzel: Copenhagen. 432 129-164 in: Doncieux, L. (ed.), Catalogue descriptifdes fossils pp.,25pis. nummuliquesdeTAudeetdel’Herault.Premierepartie:Montagne Mortensen,T. 1950b.NewEchinoidea(Spatangoida). Videnskabelige NoireetMinervous.AnnalesdeTUniversitedeLyon17:1-184,5pis. MeddelelserfraDanskNaturhistoriskForeningIKj0benhavn 112: Lambert,J., andSanchezRoig,M. 1924. P.449in: LambertJ., and 157-163. Thiery,P.Essaidenomenclatureraisonneedesechinides6and7. Rowe,F.W.E.,andGates,J. 1995.Echinodermata.InWells,A.(ed.). LibraireL.Ferriere:Chaumont. Zoological Catalogue ofAustralia, Vol. 33. CSIRO Australia: Lambert, J., and Thiery, P. 1915. Descriptions des echinides des Melbourne,xiii+510pp. Terrains Neogenes du Basin du Rhone. Memoires de la Societe Smith, A.B., Stockley, B., and Godfrey, D. 2003. Spatangoida. In: Paleontologique Suisse (Abhandlungen der Schweizerischen Smith, A.B. (ed.), The echinoiddirectory. http:Avww.nhm.ac.uk/ PalaontologischenGesellschaft)41: 155-240,pis 13-17. palaeontology/echinoids[accessed19Jun2005]. Lukasik, J.J., andJames,N. P. 1998. Lithostratigraphicrevisionand correlation of the Oligo-Miocene Murray Supergroup, western Murray Basin, South Australia. Australian Journal of Earth Sciences45:889-902. Figure 1.A,B,generallocationmaps;C,mapoftheMurrayRiverbetweenWaikerieandSwanReach, SouthAustralia,showinglocalityofNMVlocalityPL3203,northofBlanchetown. TwonewmiddleMiocenespatangoids 97 Figure 2. Murraypneustes biannulatus gen et sp. nov. A-D, adapical, posterior, adoral and left lateral views ofholotype NMV P312370; E,adapicalviewofparatypeNMVP312371;F,leftlateralviewofparatype,NMVP312372.AllfromtheearlyMiddleMioceneGlenforslan FormationinthevicinityofNMVlocalityPL3203,northofBlanchetown,SouthAustralia. Figure3.FascioledetailsofMurraypneustesbiannulatusgen.etsp.nov.A,B,obliquelateralviewandambulacrumIVdetailabovemarginof paratype,NMVP312371;C,obliqueposteriorviewofparatype,NMVP312372.Scalebar10mm. FrancisC. Holmes,ChristopherAhYeeandJaniceKrause Figure5.Murraypneustesbiannulatusgen.etsp.nov.Detailofpore pairs and tubercles in and adjacent to ambulacrum V ofparatype, NMVP312372. Figure4.ApicaldiskdetailsofMurraypneustesbiannulatusgen.etsp. nov. A, holotype NMV P312370; B, drawing of paratype, NMV P312372, showing extent of hydropores (black circles), tubercles (white circles), and gonopores and oculars (black or stippled). Locationofposterioroculars(marked*)isassumed.Scalebars1mm. Figure6.Spatagobrissusdermodyorumsp.nov.A,B,D,adapical,adoral,andleftlateralviewsofholotype,NMVP312570;C,adoralviewof paratype,NMVP312571;E,posteriorviewofparatype,NMVP312572AllspecimensfromtheearlyMiddleMioceneGlenforslanFormationin thevicinityoflocalityNMVPL3203,northofBlanchetown,SouthAustralia.Scalebars10mm. TwonewmiddleMiocenespatangoids 99 Figure 7. Spatagobrissus dermodyorum sp. nov. A, detail ofpetals and apical disk ofholotype, NMV P312570; B, peristome and labrum tuberculationofparatype,NMVP312571;C,subanalfascioleandporepairsofparatype,NMVP312573.Scalebars1mm. {mm) length test Figure8.ComparativebiometricdataonspecimensofSpatagobrissuslaubei(Duncan,1877)fromtheMiddleMiocenePortCampbellLimestone, PortCampbell,Victoria(•),andS. dermodyorumsp.nov.fromtheearlyMiddleMioceneGlenforslanFormation,Blanchetowndistrict,South Australia(o).