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A review of the Tertiary fossil Cetacea (Mammalia) localities in Australia PDF

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Preview A review of the Tertiary fossil Cetacea (Mammalia) localities in Australia

MemoirsofMuseumVictoria61(2): 183-208(2004) ISSN 1447-2546(Print) 1447-2554(On-line) http://www.museum.vic.gov.au/memoirs/index.asp A review ofthe Tertiary fossil Cetacea (Mammalia) localities inAustralia ErichM. G. Fitzgerald School of Geosciences, P.O. Box 28E, Monash University, Clayton, Victoria 3800, Australia ([email protected])andGeosciences,MuseumVictoria,GPOBox666E,Melbourne,Victoria3001, Australia(efitzger@museum,vie.gov.au) Abstract Fitzgerald,E.M.G.2004.AreviewoftheTertiaryfossilCetacea(Mammalia)localitiesinAustralia.MemoirsofMuseum Victoria61(2): 183-208. The stratigraphy, age, lithology, and vertebrate faunaofall 56 pre-Pleistocene fossil Cetacea-bearing localities in Australiaarereviewed.ThemajorityoftheselocalitiesoccurinthestateofVictoria,andareMioceneinage.Themost completecetaceanfossilshavebeenrecoveredfromcoastalexposuresoftheUpperOligoceneJanJucFormation,south- westofTorquayintheTorquayBasin(Victoria).TheinadequatelyknownfossilrecordofcetaceansinAustraliaisdue toalackofresearch,andnotalackofpotentiallyfossiliferousrockoutcrop. Keywords Cetacea,Archaeoceti,Mysticeti,Odontoceti,Australia,localities,fossilrecord,Tertiary,Oligocene,Miocene,Pliocene Introduction TheAustralianfossilrecordofcetaceansissopoorlyknown Australiahasanincompletelyknownfossilrecordofcetaceans bsiegcnaiufsiecanlitttfloesssiylstceemtaatciecanpsrohsapveectbienegnhadsisbceoevnerceadrrbiyedacocuitd.eAnlt,l (whales, dolphins, and porpoises; Order Cetacea). The oldest often by amateur palaeontologists or members ofthe public. ffoossssiill rceectoarcdeabnesinfgrobmesAtusktnraolwina afrreomEatrhleyLOaltiegoMcieoncee,nwei-tEhartlhye Icneittaicaleasntsepshatvoeadpvraenvcieourselsyeabrechenared:is1c,otvoerideedntiinfyAwushterraelifao;ss2i,l Pliocene (Fig.l for correlation of cetacean-bearing strati- determinethefaunalcompositionsoftheselocalitiesbasedon graphic units). The majority of Australian fossil cetaceans fossilsinmuseumandothercollections;3,conductacensusof described (but not necessarily published) have been derived the sedimentary geology ofthese localities; and 4, determine from only a few locations. These fossil sites occurwithin the thegeologicalageofthefossil-bearinglocalities. Paleogene-Neogene marine sedimentary basins along the This review aims to document localities inAustralia from southern margin ofVictoria. Other described fossil cetaceans which pre-Pleistocene fossil cetaceans have been recovered. have been collected from South Australia and Tasmania, Because this review is principally based on museum and although fossil cetaceans from these states are less well representedinmuseums. universitycollections,itisunlikelythatalllocalitieshavebeen recognised. Severallargeprivatecollections existthatinclude Mahoney and Ride (1975) provided a synopsis of fossil significant material. Ifaccurate locality and stratigraphic data cetacean genera and species described from Australia with informationonthehistoryofcollectionandstudy.Theydidnot oofccuforsswiilthcetthaecseeanspleoccialmietnise,s iitsimsulcikhelylartgheart.tThehirsearlenviuemwbeirs includeallcetaceansininstitutionalpalaeontologycollections. intended to serve as a companion to Rich and co-workers’ rFeovrideywcseof(1A9u8s2tara,lia1n98f2obss)ilpcuebtlaicsehaends biamspeodrtoanntmutsaexuomnocmoilc- (1991)reviewofTertiaryterrestrialmammallocalities. lections. Bearlin (1987, 1988) continued studiesonAustralian fossilCetacea, withanalyses ofNeogenemysticete specimens Overviewoflocalitiesandthefossilrecord andtheirprovenance.Inallofthesestudies,researchwascon- The fossiliferous localities in Australia that have yielded ductedprimarilyonspecimensinMuseumVictoria,Melbourne cetaceansoccurwithintheoffshoreandonshoremarine,andin (NMV),andtoalesserextentonfossilsintheSouthAustralian one case freshwater, sedimentary basins in south-eastern and Museum and the School of Earth Sciences, University of southerncentralAustralia. These areas ofPaleogene-Neogene Melbourne. The fossil cetaceans in the palaeontology collec- sedimentationcorrespondtothe: Otway,Torquay,PortPhillip, tions ofthe Tasmanian Museum, Hobart and Queen Victoria andGippslandbasinsinVictoria; StVincent,Murray,Gambier Museum,Launcestonremainlargelyunknown. andLakeEyrebasinsinSouthAustralia;andtheBassBasinin 184 E. M.G. Fitzgerald Figure 1. Correlation chart of Oligocene-Pliocene Australian fossil cetacean-bearing stratigraphic units. Abbreviations: Fmn=Formation; Lst=Limestone;Mbr=Member;Sst=Sandstone.Brokenlinesindicateuncertainmaximumandminimumagerangesofstratigraphicunits.Ages, chronostratigraphy,andplanktonicforamzonesmodifiedfromAlleyetal.(1995)andHoldgateandGallagher(2003). Tasmania (Figs 2, 3). The most abundant and diverse fossil stem-grouptoothedmysticetes(Bertaetal., 2003),thearchaic records of cetaceans occur in the Otway, Torquay, and Port baleen-bearing “cetotheres” in the Miocene, early records of Phillip basins of Victoria. However, the most complete and BalaenopteridaeintheMid-LateMiocene(Bearlin, 1988),and best-preserved fossil cetacean material is derived from the thetransitiontoarelativelymodernmysticetefaunacomposed TorquayBasinincentralcoastalVictoria. of balaenids and balaenopterids in the Late Miocene-early The most northern localities are those in the Lake Frome Pliocene(Bearlin, 1987, 1988;Fordyce, 1991). areainnorth-easternSouthAustralia(31°S), whereasthemost TheAustralianfossilrecordofOdontocetiisknownonlyin southern localities occur offshore southern Tasmania (45°S). broadoutlines.Thisreflectsthepaucityofcompleteskulls,pre- The stratigraphicrecordofcetaceans ismostcompleteforthe serving diagnostic features, in museum anduniversity collec- Miocene, with several localities in South Australia, Victoria, tions. There areonlytworecords offairly complete anddiag- and Tasmania representing the Early, Middle, and Late nostic fossil odontocete skulls in Australia: an undescribed Miocene (seebelow). However, mostMiocene sites areeither squalodontidfromVictoria(Bearlin, 1982), andProsqualodon MiddleorLateMiocene in age. TheLate Oligocenerecordis davidis (Flynn, 1923, 1948) from Tasmania. The skull ofthe becoming better known, with a higher diversity ofcetaceans latterspecimenisnowlost(MahoneyandRide, 1975) andthe than previously recognised (Fitzgerald, 2003, 2004). It is formerskulllackstherostrumandmostofthedorsalsurfaceof important to note that most of the Late Oligocene fossil thecranium.However,theremainderoftheholotypespecimen cetaceanscanprobablybereferredtoMammalodontidae,anda ofP.davidis,consistingofmuchofthepostcranialskeletonand new undescribedgroupofarchaictoothedmysticetes. Nofos- mandibles,is stillextantinthecollectionoftheDepartmentof silcetaceanshavebeenrecordedfromtheEoceneofAustralia Geology, University ofTasmania. Most odontocetefossils are (Fordyce, 1982a, 1991). Otherpoorlydocumentedstratigraph- representedbyisolatedmandibles,teeth,perioticsandtympan- ic intervals are theEarly Oligocene, earlyEarly Miocene and ic bullae. An exception is an incompletely prepared, partially LatePliocene. articulated, postcranial skeleton from the Late Oligocene of InAustralia, the fossilrecord ofmysticetes is muchbetter Victoria (NMV P48861). Although lacking cranial elements known than that for odontocetes (Bearlin, 1987; Fitzgerald, (apartfromseveralteeth),themorphologyofthescapulaindi- 2004).TheAustralianLateOligocene-LatePliocenerecordof catesthatthis specimenmayrepresentaprimitiveeurhinodel- Mysticeti is particularly informative because it documents phinidodontocete. TertiaryfossilCetacealocalitiesinAustralia 185 Figure2.TertiarysedimentarybasinswithfossilCetacealocalitiesin Victoria and Tasmania. Modified from Abele et al. (1988) and HoldgateandGallagher(2003). Taxonomy Figure3.TertiarysedimentarybasinswithfossilCetacealocalitiesin Systematicbiologistshaveyettoreachaconsensusoncetacean SouthAustralia.ModifiedfromAlleyetal.(1995). systematics andtaxonomy. This is largely due to the publica- tion ofnumerous divergentphylogenetic hypotheses, utilising bothmolecularandmorphologicaldata sets, overthe lasttwo decades (e.g., Amason and Gullberg, 1994; Geisler and Luo, studieswereinfluencedbythetaxonomicreferencesofthetime 1996; Heyning, 1989, 1997; Messenger and McGuire, 1998; and contemporary philosophical approaches to systematics. Msiclhienmkeosviwtecrheerteaasl.s,es1s9e9d3;byZGheoius,ler19a8n2d).SaHnidgehresr(c2l0a0s3s)ifbiuctattihoen Tgihveenfatuhnealpaluicstistpyroefsennetwedmhaetreeriianlre(falpeacrtttfhreosmebcieatsaecse.anHso)weavdedre,d more conventional classification of Fordyce and Muizon tomuseumcollectionsoverthelast30years,itisunlikelythat (2001) is followed herein. The classification ofnon-cetacean the taxonomy requires major reassessment. Ofmore singular mammals follows McKenna and Bell (1997). For non- importance is the incomplete nature ofthe faunallists, dueto mammalian tetrapods, osteichthyans, and most chondrich- the lack of identification of many specimens. It is highly thyans,theclassificationofCarroll(1988)isused. Somemod- probablethatthefaunallistsrepresentartificiallyimpoverished ifications to chondrichthyan taxonomy presented in Carroll faunas,particularlyofmarinetetrapodsandosteichthyanfish. (1988) and Kemp (1991) have been made following a recent Forcetaceans and othermarine tetrapods, faunal lists have beencompiledfrommypersonalobservationsandconsultation reviewbyPurdyandcolleagues(2001).Fossilspeciesofgreat- of published (Bearlin, 1988; Fordyce, 1982a, 1984, 1991; toothed sharks arereferredtothegenus Carcharodon, andnot Pledge, 1985) and unpublished (Bearlin, 1987) studies. Carcharocles,followingGottfriedandFordyce(2001). Terrestrialandfreshwatervertebratefaunaswereadoptedfrom Rich and others (1991), and unpublished data provided by Faunallists K. Piper. Chondrichthyanandosteichthyanfaunaswereadapt- FewAustralian workershave studiedCainozoic marineverte- edfrominformationinKemp(1978, 1991),Pledge(1985)and brate faunas in detail, with notable exceptions being Bearlin Stinton(1958, 1963)aswellasmyownobservations.Although (1987), Chapman and Cudmore (1924), Chapman and Pledge (1967) reviewed South Australian Tertiary chon- Pritchard (1904, 1907), Fordyce (1984, 1991), Kemp (1970, drichthyans, his published localities for chondrichthyans can- 1982, 1991),andPledge(1985).Kemp(1991)outlinedthevar- not be correlated with South Australian cetacean localities. iousbiasesaffectingthetaxonomiccompositionsofchondrich- Manyoftheidentificationsofmarinetetrapods,andtheirtaxo- thyanfaunas inAustralia, andhis conclusions may be applied nomicaffinities,arepreliminary. tocetaceans.MostofthesubstantialTertiarymarinevertebrate Information on the Australian Cainozoic fossil record of holdings of Museum Victoria were obtained prior to 1980. marine birds may be found in: Finlayson (1938), Glaessner Indeed, most ofthe collectionofCainozoic marinevertebrate (1955), Jenkins (1974, 1990), Simpson (1957, 1959, 1965, fossils dates from the first halfofthe twentieth century. It is, 1970), Rich (1975), Vickers-Rich (1991), Vickers-Rich and therefore, inevitable that identifications and alpha taxonomic Rich(1993)andWilkinson(1969).Formoredetailedinforma- 186 E. M.G. Fitzgerald tion on Cainozoic fossil marine fish see Chapman (1913, 1917a, 1930), Chapman and Cudmore (1924), Chapman and Pritchard(1904, 1907)andKemp(1991). Localities Forthepurposesofthisreview,alocalityisdefinedasalimit- ed area where fossils have been derived from one particular stratigraphicunit. However, thereis somevariationintheuse ofthe term “locality” when applied to sites thathave yielded fossil cetaceans. For example, cetacean fossils have been recovered from an area around Bird Rock, Torquay, which encompassesoutcropintheshoreplatform,andinthecliffsec- tions, at varying heights above the base of the cliff. Furthermore,fossilsmayhavebeenderivedfromanareainthe immediate vicinity of the Bird Rock stack, or up to 150 m south-westofBirdRock.Thiscontrastswithlocalities suchas CliftonBank,alongMuddyCreek,wherethereisonlyasmall areaofoutcrop withinalimited area. Inmostcases, fieldcol- lectiondataforspecimensisnotdetailedenoughtoenablepre- Figure4.Areaswithfossilcetacean-bearinglocalitiesinVictoriaand ciselimitstobeplacedonthe area ofalocality. Hence, these Tasmania. Solid circles representfossil Cetacea localities. Modified localitieshavearelativelybroaddefinition. fromAbeleetal.(1988)andHoldgateandGallagher(2003). Stratigraphy Unlessotherwiseindicated,stratigraphicnomenclaturefollows that of Abele and colleagues (1988) and Holdgate and Gallagher(2003)forVictoria,andAlleyandco-workers(1995) forSouthAustralia.Determinationofthestratigraphicposition ofmostlocalities wasbasedonworkinAbele andcolleagues (1988,andreferencestherein),datapresentedbyAlleyandoth- ers(1995),andLukasikandJames(1998). AustralianfossilCetacealocalities VICTORIA(Fig.4) 1.TheOtwayBasin(Fig.5) Figure 5. Fossil Cetacea localities in the Otway Basin, western 1.1DuttonWay,Portland Victoria.SolidcirclesrepresentfossilCetacealocalities. Geographic location. This site consists of several disjunct pointsalongsouth-south-eastfacingseacliffsandbeachonthe north-west side ofPortland Bay, north of Portland (38°19'S, Material.Allworn,isolatedelements: cranialfragments,peri- 141°38'E). Fragmentarymaterialhas alsobeencollectedfrom otics,tympanies,teeth,vertebrae,ribfragments. reefsandtheseafloorinPortlandBay,inwaterdepthsofupto Fauna. Carcharodon megalodon, Carcharodon carcharias, 100m. Isurus hastalis, Isurus sp., Palorchestidae, Vombatidae, Stratigraphic position. The fossiliferous horizon that has Macropodidae, Ektopodontidae, ?Phocidae, Balaenidae, yielded fossil cetaceans has been identified as a phosphatic Balaenoptera sp., Megaptera sp., Physeteridae, Ziphiidae, nodulebedatthebaseoftheWhaler’sBluffFormation. Delphinidae: 2species. Age. Early Pliocene. The onshore Whaler’s Bluff Formation 1.2ArchSite,GrangeBurn(Fig.6forlocalities 1.2-1.4) hasbeendatedtozonesN18-N19(Mallett, 1977).Datingofthe offshoredepositsoftheWhaler’sBluffFormationindicatethat Geographic location. Resting on aquartz porphyrybar atthe sedimentation continued into the mid-Pliocene (zones N19- baseofalowcliffonthesouthernbankofGrangeBurn,oppo- N21) (Holdgate andGallagher, 2003). Itislikelythatmost of siteanaturalarch,immediatelynorthof“TheCaves”property, the fossil cetacean material was derived from the nodule 8 km west of Hamilton, western Victoria (near 37°43'30"S, horizonandthereforehasamaximumageofearliestPliocene. 141°56'0"E) (Bearlin, 1987). Lithology. The main lithologies present include abasal phos- Stratigraphic position. Bochara Limestone Member, Port phatic nodule bed, fossiliferous clays, oysterbeds, and sandy Campbell Limestone, unconformably overlying Devonian limestones (HoldgateandGallagher,2003). RocklandsRhyolite. TertiaryfossilCetacealocalitiesinAustralia 187 been recorded from the base of the Muddy Creek Marl, at Clifton Bank, which indicate that this memberlies above the top ofthe Cyatheacidites annulatus Zone (Harris, 1971); this suggests thattheMuddy CreekMarlMemberisyoungerthan 15Ma. Gill(1957) suggestedaMiddleorLowerMioceneage (Balcombian)fortheMuddyCreekMarl. Lithology.Richlyfossiliferousgreysiltymarl. Material.Fossilcetaceanelementsaregenerallyverywellpre- served, although no articulated skulls and skeletons have yet been recovered from this unit. Elements preserved include vertebrae,ribs,andperiotics. Fauna. Carcharias sp., Carcharias taurus, Odontaspis acutis- sima, Isurus hastalis, Isurus retroflexus, Isurus desori, Isurus planus, Isuruscf.paucus,Isurussp., Carcharodonmegalodon, Carcharodon sp., Galeocerdo aduncus, Notorynchus primi- genius, Dasyatidae, Heterenchelys regularis, Muraenesox obrutus, Hypomesus glaber, Merluccius fimbriatus, Trachichthodes salebrosus, Sebastodes fissicostatus, Sillago pliocaenica, Diodonformosus,Balaenidae,Scaldicetuslodgei, cf.Physetersp.,Odontocetiindet. Figure 6. FossilCetacealocalities intheGrange Bum area, westof Hamilton in westernVictoria. Solid circles represent fossil Cetacea 1.4Forsyth’sBanktoFossilRockStack localities. Geographiclocation. Riverbankandriverbedexposuresalong andin Grange Burn, south-eastof“The Caves” homestead, 8 kmwestofHamilton(37°43'42±03"S, 14r56'40±04"E. Age. Early Middle Miocene. The foraminifera Lepidocyclina howchiniinthematrix suggests a zoneN8-N9 (Batesfordian) Stratigraphic position. Grange Burn Formation, uncon- ageforthecetaceanfossilsfromthislocality(Bearlin, 1987). formably overlying the Muddy Creek Marl. The base of the GrangeBurnFormationismarkedbyaphosphaticnodulebed Lithology.Yellow-brownbryozoalcalcarenite. horizon (Gill, 1957; Dickinson et al., 2002; Holdgate and Material. Onepartialskeleton. Gallagher,2003).InmostsectionswherethetopoftheGrange BurnFormationisexposed,abasaltlayerdisconformablyover- Fauna. Carcharodonmegalodon, Isurushastalis, Pelocetussp. liesthemarinesediments(Gill, 1957).Furthertothesouth-east alongtheGrangeBum, aterrestrialpalaeosolfaciesispresent 1.3CliftonBank betweentheGrangeBurnFormationandthebasalt. Geographic location. Low cliffs above the riverbank on the Age. The Grange Bum Formation has generally been con- westernandeastern sides ofMuddy Creek(southofthejunc- sidered as Kalimnan (Early Pliocene) in age, due to the com- tion ofGrange Burn and Muddy Creek), and due west ofthe position of the rich invertebrate macrofauna (Gill, 1957). “Clifton” property, located 8 km west of Hamilton (near Foraminifera dates indicate a zone N17 age (Mallett, 1977). 37°43'30"S, 141°56'0"E). ThebasaltabovetheGrangeBurnFormationhasbeendatedto Stratigraphic position. Muddy Creek Marl Member, Port 4S.r/3S5rMdaateussfirngomKt-hAerba(sTeuronfbutlhle Getraanl.g,e1B9u65r)n.FMoormraetiroencenitnldyi,- BCuarmnp,beel.gl.LaitmePsatt’osneG.ulInlymo(rseeecGoimlpl,le1t9e57s)e,ctitohnesMaulodndgyGrCarnegeek cetatael.d,a2040.20)-.5T.0hMesaemdaatxaiimnduimcaatgeeafnoreatrhleiefstorEmaartliyonPl(iDoicceknienasgoen MarlisconformablyunderlainbytheBocharaLimestone(Gill, fortheGrangeBurnFormation. 1957), and disconformably overlain by the Grange Burn Formation. Lithology. Shelly marl and sandy to pebbly limestone (HoldgateandGallagher,2003). Age. Middle Middle Miocene to early Late Miocene. Sr/Sr dating ofshells from 1 mbelow thebase ofthe Grange Burn Material.AsistypicalofmostMio-Pliocenenodulebedfossil Formation, within the Muddy Creek Marl, yielded an age of vertebratematerialinVictoria,fossilsareoftenrolled,polished, 10.8 Ma (Dickinson et al., 2002). The presence of the andbroken.Almostallspecimensrepresentisolatedelements, foraminiferan Orbulina suturalis indicates a zone N10 age with associated material being very rare. Typically preserved (MiddleMiocene,Serravallian, 12-14Ma)(Abeleetal., 1988). elements include partial rostra, cranial fragments, isolated However, the Muddy Creek Marl can range as high as zone periotics and tympanic bullae, teeth, incomplete mandibles, N16(upto~8Ma)(Singletonetal., 1976).Acaciapollenhave vertebrae,andribs. 188 E. M.G. Fitzgerald Fauna. Heterodontus cainozoicus, Carcharodon megalodon, 1.7LeighRiver Carcharodon carcharias, Isurus escheri Isurus hastalis, Myliobatis sp., Edaphodon sweeti, Isch,yodus cf. dolloi, RGievoegrr,apahbiocutlo5catkiomn.nOourtthcroofpoSnhetlhfeoreda,st4er6nkbamnknoorftthh-eeaLsetigohf Kurrabi sp., Phocidae, “Cetotheriidae”, Balaenoptera sp., cf. Geelong(near38°00'S, 143°58'E). Scaldicetus sp., cf. Physeter sp., cf. Mesoplodon sp., ?Delphinidae. Stratigraphicposition. Middle section ofthe Gellibrand Marl (Abeleetal., 1988;Dickinsonetal., 2002). 1.5SpringCreek Age.ProbablyEarlyMiocene.Inthenorth-eastpartofthePort Geographiclocation.BedofSpringCreek,nearMinhamite,40 CampbellEmbayment,theGellibrandMarlrangesinagefrom kmsouth-eastofHamilton(near37°59'S, 142°20'E). Late Oligocene to Middle Miocene (Abele et al., 1988). OutcropalongtheLeighRiverisEarlyMioceneinage, asthe Stratigraphicposition. Unnamedunit(Abeleetal., 1988).This Late Oligocene-aged base of the formation is not exposed, unit is probably laterally equivalent to the Grange Burn and the youngest Middle Miocene (Bairnsdalian) section Formation. Gill (1957) mentioned a Kalimnan-aged location has probably been eroded, ornot deposited, due to the Leigh “from Goodwood station nearMinhamite Railway Station 25 River locality being at the embayment margin (Abele et al., miles(40.2km) SEofHamilton”(p. 152). 1988). Age. Presumed Late Miocene-Early Pliocene. T.A. Darragh Lithology.Marl,calcareous silt,clay,andsandwithminorcal- (pers. comm, to G.G. Simpson, cited in Simpson (1970)) carenitelayers. has suggested a Cheltenhamian (Upper Miocene) or older age for the Spring Creek beds near Minhamite. Material. Onewell-preservedisolatedtooth. Simpson (1970) suggested that the Spring Creek locality Fauna.Isurussp.,Cetaceaindet. was equivalent in age to the Black Rock Sandstone at 1.8HopkinsRiver Beaumaris. Lithology. Fossiliferous green-grey marly fine sand Geographic location. Northendofoutmcrop in aquarry onthe m west bank ofthe Hopkins River, 150 south ofthe Princes approximately 1 thick(Abeleetal., 1988). HighwaybridgeovertheHopkinsRiver,nearAllansford,about Material. Worn and polished isolated elements: periotics and 10kmeastofWarrnambool(near38°23'S, 142°35'E). indeterminatebonefragments. Stratigraphicposition. Port Campbell Limestone sensu stricto Fauna. Pseudaptenodytesmacraei, Balaenopterasp. (Abeleetal., 1988;Tickelletal., 1992). Age. Middle-Late Miocene, zones N11-N17 (Holdgate and 1.6Kawarren Gallagher, 2003). Geographiclocation. Old“Alkemade’sQuarry”, slightlynorth Lithology.Yellowfine-grainedcalcarenite. ofKawarrenrailway station,onthe steepnorthbankofLoves Creek, Kawarren, about 19 km south ofColac (near 38°29'S, Material. A fairly well-preserved posterior part of mandible 143°35'E). withtwolargeconicalteethinplace. Stratigraphicposition. Clifton Formation (Abele et al., 1988; Fauna,cf.Scaldicetussp. McHaffieandInan, 1988). 1.9Gibson’sSteps Age. In the north-east margins of the Port Campbell Embayment,theCliftonFormationisLateOligocene(Abeleet Geographiclocation. CliffatGibson’sSteps,approximately 12 al., 1988). km east of Port Campbell, western Victoria (near 38°40'S, 143°07'E). Lithology. The Clifton Formation is generally a medium- coarse-grainedcalcarenite,withabout10%quartzandlimonite Stratigraphic position. Port Campbell Limestone (Bearlin, sand (Tickell et al., 1992). At the Kawarren Quarry, this unit 1987;Abeleetal., 1988;Tickelletal., 1992). consists of friable, pale yellow limestone interbedded with Age.Bairnsdalian,MiddleMiocene,zoneN10(Bearlin, 1987). hardercrystallinebands (McHaffieandInan, 1988).Allofthe vertebrate fossils have been recovered from the more friable Lithology.Yellow-greyfine-grainedcalcarenite. calcarenitelayers. Material. Onepartialskeleton Material. Generallywell-preservedisolatedelements, exhibit- Fauna. “Cetotheriidae”. ing alow degree ofrolling. Elementspreservedincludeteeth, ribsandbonefragments. 1.10Curdie Fauna. Carchariassp.,Isurussp., Isurusdesori, Carcharoides Geographic location. “Kurdeez” Quarry (Victorian sp., Carcharoides totuserratus, Carcharodon angustidens, AgriculturalLimeLtd), 5kmnorth-north-westofCurdie,near ?Mysticetinewfamily. Timboon(approximately38°27'S, 142°56'E). TertiaryfossilCetacealocalitiesinAustralia 189 Stratigraphicposition. PortCampbellLimestone. Lithology.Sandybryozoalcalcarenitewithathindiscontinuous basallayerofphosphaticnodulesandquartzpebbles (Abeleet Age. Balcombian, correlated with zone N10, Serravallian, MiddleMiocene(Bearlin, 1987). al., 1988). Material. Onetooth. Lithology. Calcarenite. Material. Onepartialskeleton. Fauna.Prosqualodonsp. (=Parasqualodonwilkinsoni). Fauna. ?Balaenopteridae. 2.TheTorquayBasin(Fig.7) 1.11PrincetownBeach 2.1 SplitPoint(Fig.8forlocalities2.1-2.7) eGaesotgorfapPhoritcClaomcpatbieolnl.,NweeasrteProninVtictRoorniaald(,38°P4r2i'nSc,et1o4w3n°,09'1E8).km AGieroegyrsapIhnilect,lo4ca9tikonm. Tsoowuatrh-dwsesltighotfhoGuseee,loantgSpl(inteaProi3nt8,°2n8e'Sa,r 144°06'E). Stratigraphic position. Nodule bed within the Clifton Stratigraphicposition. PointAddisLimestoneMember(Webb, Formation. 1995). Age. Late Oligocene. Below the unnamed nodule horizon, shells have given Sr/Sr dates of 27.4 Ma (Holdgate Age.LateOligocene. and Gallagher, 2003). Immediately above the nodule bed Lithology.Yellowsandybryozoalcalcarenite. Sr/Sr dates average -24.0 Ma. Foraminifera from the Clifton Formation indicate a zone P21b-P22 age (Late Material. One incomplete anterior caudal vertebra, and one worntooth. Oligocene). Fauna. Carcharias macrotus, Carcharoides totuserratus, Lithology. Thin horizon of phosphate and limonite nodules Cetaceaindet. withinlimestone-sandylimestonematrix. Material. Onewornandpolishedincompleterib. 2.2PointAddis Fauna. Odontaspissp.,Cetaceaindet. Geographic location. Cliffs on southern side ofPoint Addis, south-west of Torquay, central coastal Victoria (38°23'S, 1.12CastleCove 144°15'E). Geographic location. One oftwo possible locations: (1) type Stratigraphicposition. PointAddisLimestone.Mostvertebrate localityfortheCalderRiverLimestonealongthesouth-eastern fossilshavebeencollectedfromthebaseoftheuppermember bthaenkcooafstth1e.C7akldmersoRuitvhe-re,ansotrotfh-CwaessttleofCHoovred.eBrnotVhalsei,teosra(r2e)oonn Wofebtbhe,P1o9i9n5t).Addis Limestone (Nicolaides andWallace, 1997; thewesternsideofCapeOtway,andeastofPointReginald,in the Aire district (near 38°47'S, 143°25'E). Etheridge (1878) Age. Late Oligocene (Abele et al., 1988; Holdgate and notedthattheholotypetoothofParasqualodonwilkinsoniwas Gallagher,2003). foundwestoftheAireRiver. Lithology. Ferruginous intraclastic conglomeratewithabraded Stratigraphic position. Calder River Limestone. Fordyce shelly and vertebrate skeletal components (Nicolaides and (1982a) suggested that the holotype specimen of Wallace, 1997; Webb, 1995). This horizon directly overlies a Parasqualodon wilkinsoni was derived from the Calder regionally extensive hardground (Nicolaides and Wallace, River Limestone, despite the fact that there is no direct 1997;Webb, 1995). evidence that would indicate such a derivation. However, Material. Isolated elements: teeth, tympanic bullae, rarepost- there are two lines of evidence that indirectly suggest the cranialremains.Thematerialisusuallybrokenandworn. provenance ofthe holotype ofParasqualodon wilkinsoni'. (1) the holotype tooth preserves distinctive features suggesting Fauna.Mammalodontidae. affinities with Prosqualodon davidis (indeed, this isolated tooth may be congeneric with P. davidis, or even con- 2.3BellsHeadland specific); the Prosqualodontidae is only known from Late Oligocene-earliest Miocene deposits; (2) McCoy (1867a) Geographiclocation. 300msouth-westofBellsBeach, south- statedthattheholotypetoothwascollectedfromsandybedsat westofTorquay(near38°22'S, 144°16'E). Castle Cove on the Cape Otway coast; the only fossiliferous Stratigraphic position. Lower beds of the Point Addis marinesedimentsinthisarea,whichareofsuitableagetoyield a prosqualodontid tooth, are those of the Calder River Limestone(Abeleetal., 1988). Limestone. Age.LateOligocene. Age.LateOligocene. Lithology. Sandybryozoalcalcarenite(Abeleetal., 1988). . 190 E. M.G. Fitzgerald Figure 7. Fossil Cetacea localities in the Torquay and Port Phillip basins,centralcoastalVictoria. SolidcirclesrepresentfossilCetacea localities.ModifiedfromHoldgateandGallagher(2003). Material. Fossilpreservationis generallyverygood. Cetacean wFiegsutreof8.ToLraqtueayO,liignocceennteraflossciolasCteatlacViecatolroicaa.litSiheasdoendtahreeacsoarsetpsroeustehn-t material consists of: one skull and associated ear bones; coastal outcrop of the Oligocene-Miocene Torquay Group. Solid associatedvertebrae,andbonefragments. circlesrepresentfossilCetacealocalities. Fauna.Mysticetinewfamily; genusandspecies2. Stratigraphicposition. JanJuc Marl,basedonthepresence of 2.4BellsBeach themolluscsLiratominaintertextaandChionehalli(whichare Geographiclocation. Onshoreplatform,lowtidemark,baseof restrictedtotheJanJucMarl,T.A.Darragh,pers.comm.). low cliffs at north-east end of Bells Beach, south-west Age.LateOligocene. ofTorquay,centralcoastalVictoria(38°22'S, 144°17'E). Lithology. Grey-yellow glauconitic and pyritic carbonate- Stratigraphicposition. PointAddisLimestone cementedcalcarenite. Age.LateOligocene Material. One partially articulated incomplete skeleton: skull, Lithology.Yellowbryozoalcalcarenite(Abeleetal., 1988). mandibles,teeth,tympanies,thyrohyal,atlas,axis,cervicalver- tebra,ribs, scapulae,radius. Material. One partially articulated incomplete skeleton, isolatedperiotics,andvertebrae. Faunal.Mysticetinewfamily,genusandspecies 1. Fauna. Carcharodon angustidens, Mammalodontidae genus andspeciesindet 1 2.7BirdRock 2.5RockyPoint Geographic location. Bluff adjacent to Bird Rock stack and cliffs to the south-west, and shore platform exposures from Geographic location. Low cliffs at Rocky Point, small head- BirdRockinthenorth-east,extending250mtothesouth-west; landatnorthern-mostendofBellsBeach, about250 mNE of south-west of Torquay, central coastal Victoria (38°20'54"S, BellsBeach, south-westofTorquay(near38°22'S, 144°17'E). 144°18'35"E). Stratigraphicposition. ?LowerJanJucMarl. Stratigraphic position. Upper Jan Juc Marl (Holdgate and Gallagher,2003;Webb, 1995). Age. ?LateOligocene. Age. Late Oligocene. Siesser (1979) suggested that the Lithology.Yellow-orangesiltyandsandymarls. Oligocene-Mioceneboundary occurs 2.5 mbelowtheJanJuc Material.Indeterminatebonefragments. Marl-PueblaClaycontact.Liandcolleagues(1999)notedthat thelocalnannoplanktondatumusedby Siesser(1979) toesti- Fauna. Cetaceaindet. mate the position of the Oligocen-Miocene boundary, is no 2.6Deadman’sGully longervalidbecausethe ageofthisdatumis -24.5 Ma, while the recognised date of the Oligocene-Miocene boundary is Geographic location. Cliff exposures 600 m south-west of 23.8-23.9Ma(Berggrenetal., 1995).Liandothers(1999)pro- Fishermans Steps, near Deadman’s Gully, south-west of videdforaminiferalbiofaciesevidenceindicatingthatthetopof Torquay,centralcoastalVictoria(near38°20'S, 144°18'E). the Jan Juc Marl corresponded to the Late Oligocene-Early TertiaryfossilCetacealocalitiesinAustralia 191 Miocene boundary. Furthermore, the Jan Juc Marl-Puebla Material. The completeness and quality of preservation of Clay contact has yielded Sr/Sr dates of 23 ± 1 Ma (Kelly fossil cetacean material varies. However, the vast majority et al., 2001), consistent with the Oligo-Miocene boundary of cetacean fossils consist of disarticulated and isolated ele- occurring at the Jan Juc Marl-Puebla Clay contact, and not ments. Some fossilspreserve fine surface detail onbones and within the Jan Juc Marl. Foraminifera from the Jan Juc Marl teeth, whereas others are highly polished and worn with no correlate this formation with international foram zones surfacedetailpresent. P21-P22, and therefore indicate a Late Oligocene age Fauna. Carchariastaurus, Carchariasmacrotus, Carcharoides (Holdgate and Gallagher, 2003). Sr/Sr dates have yielded an a(Dgiecokfin2s3o.n9,-2270.042;MaHoflodrgtahteeJaanndJuGcalMlaarglheerx,p2o0s0e3d).atTBhiersdeRdoactka tspo.t,usIeLrraamtunsa,caCtatriccah,aLroadmonnaasnpg.u,sCtaidrecnhsa,rhIisnuursusspd.,esGoaril,eoIcseurrduos sp., Notorynchus cepedianus, Myliobatis sp., Diprotodontidae supportaLate Oligocene (Chattian) ageforthe JanJuc Marl, and indicate that the beginning ofthe Early Miocene occurs c(oDl.livPeircik,erMinagmmaanldoTd.onRicshp.,perMsa.mmcaolmom.d)o,ntcifd.aMeamspm.alionddeot.n, immediately above the top of the Jan Juc Marl, in the “Cetotheriidae”,Mysticetinew genus,Mysticetifamilyindet., PueblaClay. cf. Squalodontidaeindet.,Odontocetifamilyindet. Lithology. Siltyglauconitic marl andclayey sandy glauconitic calcarenite (Abele, 1979; Glover, 1955; Holdgate and 2.9OceanGrove Gallagher,2003;RaggattandCrespin, 1955;Webb, 1995). Geographic location. Near Ocean Grove, on the Bass Strait Material.Fossilpreservationisgenerallyfairlygood,however sideoftheBellarinePeninsula(near38°15'S, 144°3FE). fossilsfromtheupperbedsexposedinthebluffareoftencor- roded. Typical elements preserved include ribs, vertebrae, Stratigraphicposition. PueblaClay(Abeleetal., 1988). teeth, tympanies, and periotics. Less common are skulls and Age.EarlyLongfordian(EarlyMiocene). partiallyarticulatedskeletons. Lithology. Calcareoussand(Abeleetal., 1988). Fauna. Heterodontus cainozoicus, Carcharias elegans, Carcharias taurus, Carcharias sp., Odontaspis incurva, Material.Bonefragments. Carcharoides totuserratus, Carcharodon angustidens, Isurus Fauna.Cetaceaindet. desori, Isurusplanus, Isurus sp., Dasyatis sp., Myliobatis sp., Megalops lissa (Stinton, 1958), Pterothrissus pervetustus 3.ThePortPhillipBasin(Fig.7) (Stinton, 1958), Heterenchelys regularis (Stinton, 1963), Astroconger rostratus (Stinton, 1958), Urconger rectus 3.1BatesfordQuarry (Stinton, 1963), Merluccius fimbriatus, Gadus refertus Geographic location. Australian Cement Company quarry (Stinton, 1958), Ophidion granosum (Stinton, 1963), south of Batesford, on the western bank of the Moorabool Trachichthodes salebrosus (Stinton, 1958), Cleidopus carver- River,westofGeelong(38°06'S, 144°17'E). nosus (Stinton, 1963), Sillago pliocaenica (Stinton, 1958), Coelorhynchus elevatus, Xiphias sp., Aves indet., Mysticeti Stratigraphic position. Batesford Limestone; unconformably family indet., Mammalodon colliveri (Pritchard, 1939), overliesPalaeozoicgranite,andgradesconformablyupintothe Mammalodon sp.,Mammalodonnewspecies 1,Mysticetinew overlying Fyansford Formation (Abele et al., 1988; Holdgate family,Prosqualodonsp., ?Eurhinodelphinidae. andGallagher,2003;Webb, 1995). Age. The lower 21 m of the Batesford Limestone is 2.8WaurnPondsQuarry Longfordian(LowerMiocene)inage,whereastheupper 12m Geographic location. Waurn Ponds Quarry, operated by Blue tohfethueppfeorrmmoasttioLnoiwsetrhetoLelpoiwdeorcyMcildidnlae-bMeiaoncnegnteypBeatseescftoirodniaonf Circle Southern Cement Ltd, south of Waurn Ponds (near localmarinestage(HoldgateandGallagher,2003). 38°12'S, 144°16'E). Lithology.Fossilcetaceanshavebeenrecoveredfromthebasal Stratigraphicposition.WaurnPondsLimestoneMemberofthe beds, which are composed of calcareous sand and gravel. JanJucFormation; alateralequivalentofthelowerpartofthe The upper Batesfordian part of the formation consists of Point Addis Limestone (Holdgate and Gallagher, 2003; biocalcarenite(Abeleetal., 1988;Bowler, 1963;Webb, 1995). NicolaidesandWallace, 1997). Material. In both upper and lower parts of the Batesford Age.LateOligocene(-24-27Ma)(Abeleetal., 1988;Holdgate Limestone, cetaceanfossils are generally well preserved with andGallagher,2003;NicolaidesandWallace, 1997). only slight mineralisation of original bone. The majority of Lithology. Bryozoal calcarenite with some interbedded marls. cetaceanfossilshavebeenfoundintheBatesfordian-ageupper These sediments are capped by a subaerial exposure surface beds.Ribfragmentsandvertebraearethemostcommonlyrep- thatislaterally equivalentto a similarhorizon atPointAddis. resented elements, with tympanic bullae being poorly repre- At Waurn Ponds Quarry, this horizon is heavily cemented sented. Elements of the appendicular skeleton and cranial (Abeleetal., 1988; Holdgate and Gallagher, 2003; Nicolaides remainsareveryrare. Onlytwoskeletonsrepresentedbyassoc- andWallace, 1997;Webb, 1995). iatedelementshavebeenrecoveredfromBatesfordQuarry. 192 3.5 E. M.G. Fitzgerald Fauna. Orectolobus sp., Carcharias taurus, Carcharodon Beaumaris megalodon, Isurus desori, Isurus hastalis, lsurus oxyrinchus, Geographic location. Coastal exposures located east of Isuruscf. oxyrinchus, Isurusplanus, Isurus retroflexus, Isurus Rickett’sPointonthewestshoreofBeaumarisBay,north-east cf.paucus,Isurussp.,Lamnidaeincertasedis,Carcharhinuscf. shoreofPortPhillipBay(37°59'S, 145°03E). brachyurus, Carcharhinus sp., Galeocerdo aduncus, Sphyrna sp., Notorynchus primigenius, Pristiophorus lanceolatus, Stratigraphicposition. BasalBlackRockSandstone (Abeleet Labrodon sp., Spheniscidae, Diprotodontidae indet., “Ceto- al., 1988;HoldgateandGallagher,2003). theriidae”indet.,Physeteridae,?Squalodontidaenewgenusand new species. Age.CheltenhamiantoKalimnan(latestLateMiocene-earliest EarlyPliocene)(Abeleetal., 1988;Dickinsonetal.,2002;Gill, 3.2MooraboolRiver 1957;HoldgateandGallagher,2003;Mallett, 1977). Geographic location. North-west of Geelong (near 37°56'S, Lithology. Mostfossilvertebrateshavebeenrecoveredfroma layer of ferruginous and phosphatic nodules in a matrix of 144°09'E). quartz sand and gravel. This nodulebedoccurs atthebase of Stratigraphicposition. LowerMaudeFormation (Abeleetal., the Black Rock Sandstone. Less abundant fossil vertebrates 1988). occurinthe sedimentsimmediately abovethenodulehorizon, Age. Latest Late Oligocene-earliest Miocene (zones P22-N4) this layer consisting ofcalcareous sandstone and sandy marl (Abeleetal., 1988;HoldgateandGallagher,2003). (Abeleetal., 1988; Gill, 1957; Singleton, 1941). Material. Cetaceanfossilsderivedfromthenodulebedconsist Lithology. Shellybryozoalcalcarenite. offragmentary, isolatedelementsthatareusuallyhighlyworn Material.Cetaceanmaterialconsistsofonetympanicbullaand andpolished,havingundergonesignificantpost-mortemtrans- bonefragments. port. Thesefossilsincludeindeterminatebonefragments,ribs, Fauna. Heterodontus cainozoicus, Carcharias sp., Isurus sp., vertebrae, forelimb elements, teeth, rostral and cranial frag- cf.Mammalodontidae. ments, periotics, tympanic bullae, and incomplete mandibles. Vertebrate fossils from the sandy beds overlying the nodule 3.3NorthShore, CorioBay horizon are better preserved than the nodule material, with a lesserdegree ofreplacementofboneby secondary mineralis- Geographiclocation. OutcropalongnorthshoreofCorioBay, ation. The sandy bed material includes ribs, vertebrae, teeth, nearGeelong(near38°06'S, 144°24'E). mandibles,periotics,tympanicbullae,middle-earossicles, and Stratigraphic position. Fyansford Formation (Abele et al., rarepartiallyarticulatedskeletons. 1988). Fauna. Heterodontus cainozoicus, Carcharias taurus, Age. Youngest part ofthe Fyansford Formation, representing Carcharodon megalodon, Parotodus benedenii, Isurus desori, zoneN12(Bairnsdalian,MiddleMiocene)(Abeleetal., 1988). Isurus hastalis, Isurus retroflexus, ILamna, Megascyliorhinus sp., Carcharhinus cf. brachyurus, Carcharhinus sp., Lithology. Basal sandy calcarenite passing upwards into cal- Galeocerdo aduncus, Pristiophorus lanceolatus, Myliobatis careoussiltandclaywithsandycalcareniteinterbeds(Abeleet sp.,Edaphodonmirabilis,Edaphodoncf.mirabilis,Edaphodon al., 1988). sweeti, Ischyodus cf. dolloi, Sillago pliocaenica (Stinton, Material.Bonefragments. 1958), Lactarius tumulatus (Stinton, 1963), ?Trionychidae, Pseudaptenodytes macraei, IPseudaptenodytes minor Fauna.Lamnasp.,Cetaceaindet. Spheniscidae, Diomedea thyridata (Wilkinson, 1969),, Zygomaturusgilli (Woodburne, 1969; Stirton, 1967),Kolopsis 3.4Curlewis sp. cf. K. torus (Rich et al., 2003), Phocidae (Fordyce and Geographic location. Cliffsections along the northern side of Flannery, 1983), “Cetotheriidae” indet., cf. Balaena, the Bellarine Peninsula, near Curlewis (near 38°ll’S, Balaenidae indet., Balaenoptera sp., Megaptera sp., 144°30E). Physetodonbaileyi(probablyPhyseteridaeindet.), Scaldicetus macgeei,Physeteridaeindet.,Ziphiidae,Delphinidaeindet. Stratigraphic position. Fyansford Formation (Abele et al., 1988). 4.GippslandBasin(Fig.9) Age.EarlyMiddleMiocene(Abeleetal., 1988). 4.1MerrimansCreek Lithology. Calcareous clay and marl with thin bryozoal m Geographiclocation.Approximately25 fromthesurface,in calcareniteinterbeds (Abeleetal., 1988). the Gippsland Cement Quarry onMerrimans Creek, about 19 Material.Bonefragments. km south-east ofRosedale, Gippsland,Victoria(near38°15'S, 146°51E). Fauna. Heterodontus cainozoicus, Isurus hastalis, Trachichthodessalebrosus(Stinton, 1958),Diodonsp.,Cetacea Stratigraphic position. Gippsland Limestone Formation indet. (HoldgateandGallagher,2003).

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