Table Of Content

A SIGNIFICANCEOFPOLARDINOSAURS IN GONDWAN THOMAS H. RICH Rich, T.H. 1996 12 20: Significance of polar dinosaurs in Gondwana. Memoirs ofthe QueenslandMuseum39(3)711-717.Brisbane. ISSN0079-8835. Polardinosaurs havebeen found at fourlocalities in the southern hemisphere and eight in the northern. Three groups of dinosaurs, neoceratopsians, ornithomimosaurs and oviraptorosaurs,previouslyknownonlyfromtheLateCretaceousofthenorthernhemispherewere present in the Early Cretaceous polar dinosaur fauna of southeastern Australia. Labyrin- thodontamphibianswerealsorepresent,butdonotoccurinthesamedepositsascrocodilians. Enlarged optic lobes ofthehypsilophodontid Leaellynasaura, suggestingenhancedability toseeunderlowlightconditions,istheonlyadaptationtoapolarenvironmentyetrecognised in adinosaur.Polardinosauriantaxaarenotuniqueatthefamiliallevel,butbelongtomore widespreadfamilies. Polardinosaurs probablylived undercoldclimatesquiteunlikethose tolerated by modern reptiles. QJ Early Cretaceous, southeastern Australia, dinosaurs, labyrinthodonts,palaeoclimate. Thomas H. Rich, Museum ofVictoria, P.O. Box 666E, Melbourne, Victoria, Australia; 5 April1996. Only one area within the pa—laeo-Antarctic Gondwanamay have been theplaceoforigin for Circle has produced dinosaurs southeastern theneoceratopsians, fortheyareknown nowhere Australia (Aptian-Albian, early Cretaceous). Four else priorto the Late Cretaceous. more are known to—have been close to the palaeo- Ornithomimosaurs are represented in the Al- Antarctic Circle southeastern Queensland, bian ofsoutheastern Australiaby femoradistinct Australia (early or middle Jurassic), Beardmore enoughtobaseanewgenusandspecies, Timimus Glacier, Antarctica (early Jurassic), James Ross hermani Rich & Vickers-Rich, 1994. As yet un- and Vega Islands, Antarctica (Campanian and described ornithosaur vertebrae are known from Maastrichtian,lateCretaceous)andNorthIsland, both the same Albian site that produced femora New Zealand (probably Maastrichtian, late from southeastern Australia as well as Aptian Cretaceous). In the Northern Hemisphere, eight sites a few hundred kilometres to the east. This areas (late Jurassic? to late Cretaceous) are material suggests apresenceforthisgrouponthe known to have produced dinosaurs within the Gondwana continents prior to when or- palaeo-Arctic Circle (Fig. 1). Most of the polar nithomimosaurs are best known in the northern dinosaur material from both hemispheres was hemisphere, the late Cretaceous. However, the acquiredduring thepasttwentyyears andmuch recent publication of Shuvosaurus inexpectatus ofitremains to be described (Table 1). Chatterjee, 1993 from the late Triassic ofTexas, Polar Gondwana may have served as both a implies that ornithomimosaurs have had a much refuge andbirthplace for somedinosaurgroups. longer history in the northern hemisphere than From the early or middle Jurassic of previously suspected. The late Jurassic Elaphro- southeastern Queensland has come a partial saurus bambergi from Tendaguru, Tanzania, skeleton of one of the earliest sauropods, formerly regarded as an ornithomimosaur, has Rhoetosaurusbrownii Longman, 1927. recently been allocated to the abelisaurs (Holtz, Asingle astragalus suggests thatthe well-known 1994). formAliosaurus may have persisted into the Ap- Oviraptorosaurs, previously represented ex- tian of southeastern Australia after having be- clusively in the late Cretaceous of the northern comeextinctelsewhereattheendoftheJurassic. hemisphere, appear to have been present in the Onthebasisofoneorafewbones,thepresence Albian ofsoutheastern Australia based on a par- of three groups previously known in the late tial surangular and a vertebra (Currie, Vickers- & CretaceousoftheNorthernHemispherehasbeen Rich Rich, 1996). suggestedintheearly Cretaceousofsoutheastern Associated with these dinosaurs is the most Australia. AnulnafromtheAptiantherebearsan unexpected andbestdocumentedtemporalrange uncannyresemblancetothatoftheMaastrichtian extensionofamajortetrapodgroup.Twodecades protoceratopsianLeptoceratopsgracilisfromAl- ago, labyrinthodont amphibians were thought to berta, Canada. This suggests that Polar have become extinct at the end of the Triassic. 712 MEMOIRS OFTHEQUEENSLAND MUSEUM * ir Ankylosaurla Aves Carnosaur Carnosaurfootprint SmallCarnosaurorTheropod Ceratopsidae > >*-^5r^ Hypsilophodontidae,5differentgenera Iguanodontidaefootprint Labyrinthodontamphibian Marsupialia Jf -r Piesiosaur Prosauropod Protoceratopslan Sauropod FIG. 1.Distributionofpolardinosaurs,SouthernHemisphere(thispage)andNorthernHemisphere(facingpage). Dinosaursites in polarlatitudes are notcommon, mosthavingbeen discovered in the last20years. None are fully studied. The higher diversity in southeastern Australia is probably due to greater collecting effort and easieraccessibility. When some ofthe othersites are more fully studied, they may provide similardiversity. BasemapfromSmith, Hurley &Briden(1981). SIGNIFICANCEOFPOLAR DINOSAURS IN GONDWANA 713 K ^ <ss§P-$r NorthwestTerritories, Canada TalkeetnaMountains Alaska GeologicalAgeofSiteorSites Dromaeosaurldae Hadrosauridae Hypsilophontidae J - Jurrassic, 145-208millionyears eJ - EarlyJurassic, 178-208millionyears IJ - LateJurassic,145-157millionyears eK - EarlyCretaceous,97-145millionyears IK - LateCretaceous,65-97millionyears r Multituberculata Ornithomimosauridae Oviraptosauridae Placentalla QuantityofFossilMaterial Greaterthan1,000bones 100-1,000bones Stegosauridae Tritylodontid Troodontidae Lessthan 100bones FIG. 1 (cont). Subsequent to that, they have been described (Dong, 1985) and late Jurassic of Mongolia from the early Jurassic of Australia (Warren & (Shishkin, 1991). Nearly complete lower jaws Hutchinson, 1983), middle Jurassic of China with teeth, pectoral girdle elements, and ver- 714 MEMOIRS OFTHEQUEENSLAND MUSEUM tebrae form the basis for recognition of their However, in the much more fossiliferous Lis- presence in the Aptian of southeastern Australia comb Bonebed locality on the Colville River, (Warren, Kool, Cleeland, Rich & Rich, 1991; Alaska, hypsilophodontids are all but unknown & Warren, Rich Vickers-Rich, inpress). despite the extensive sieving programme carried & Crocodilian-like in their overall morphology, out there (Clemens Nelms, 1993; pers. obs.). the latest survivors of this once dominant group The age of the site is Maastrichtan and by that ofamphibians may have found a final refuge in time there may not have been the variety of Aptian polarsoutheastern Australiabecause ofa hypsilophodontids there was earlier in the greater tolerance of cold, as is true of modern Cretaceous.Butthatdoesnotexplainwhytheone frogs and salamanders, than the crocodilians known to occurthere is not more abundant. which appeared there subsequently in theAlbian Today, there are no avian or terrestrial mam- when conditions became warmer. There is no malianfamiliesrestrictedtothepolarregions. As trace of an Albian labyrinthodont in the south- faras the available recordhas been analysed, the eastern Australian deposits where crocodilians same can be said of the polar dinosaurs as pre- & arerepresented by a few dermal scutes. viouslyindependentlynotedbyMolnar Wiffen Among polar dinosaurs from either hemi- (1994). Although new genera and species have sphere,onlyonefeatureofoneindividual has yet been recognised among them, they all belong to been interpretedas an adaptation tolife in ahigh families alsoknown at lowerlatitudes. latitude environment. This may reflect more the However, given the fragmentary nature of fact that very little is known about theseanimals muchoftheevidencefortheexistanceofvarious andlesspublished,thanthattheyrarelydisplayed groups, this could be an artefact. If one had a markeddifferencesbetweenthemselvesandtheir single tooth of a litoptern and that group was lower latitude contemporaries. The feature in otherwise unknown, depending on the species, question is the enlarged optic lobes ofthe brain the most parsimonious familial identification that occurred on the holotype of the hyp- might be Equidae. In that case one would be silophodontid Leaellynasaura amicagraphica parsimonious but one would be wrong. The & Rich Rich, 1989 from the Aptian of south- recent identification of protoceratopsians in the & eastern Australia. Hypertrophy of this structure Aptian ofAustralia (Rich Vickers-Rich, 1994) formed the basis for the suggestion that this could be aparallel case. On the basis ofa single animal hadenhancedabilitytoseeunderthelow bone, rather than propose an entirely new group lightconditionsthatwouldhaveprevailedduring of vertebrates which would share an uncanny the prolonged periods of continuous darkness resemblance in the form of the ulna to eachWinter. protoceratopsians but would in some other way Hypsilophodontid dinosaurs are generally a bedistinctfromthem,thespecimenwasallocated rareelementin mostdinosaurassemblages.Even tothisknowntaxonwhichextendeditsrecordnot where specimens are relatively common, as on only to another continent but also backwards in theIsleofWight, theirtaxonomic diversity is not time by at least 15 million years. At our present great. Southeastern Australia is a marked excep- stateofknowledge thatis an identificationwhich tion to thatgeneralisation. At leastsix species in isquiteplausiblebutcould well eventuallyprove five or six genera occur there;just over half the to be fundamentally in error. total dinosaurs recognised to date. Currie (pers. The Gondwanan polar dinosaur localities that comm.) has suggested that hypsilophodontids form groups ofthe same age are quite meagre in may have been primarily an upland group at theirtotal faunal lists, making comparisons quite lower latitudes, hence their general rarity there, preliminary. Itis fortuituous, however, thatinthe and are better represented in southeastern broadestterms,theyshareacommonfacies. Both Australia because ofits coolerconditions. early Jurassic sites are flood plain deposits and In this regard it may be noteworthy that the thethreelateCretaceousones, nearshoremarine. dinosaurknownfromthelateCretaceousofVega Itmaybe ofsome notethattheNewZealand late Island is a hypsilophodontid or dryosaurid (Mil- Cretaceous locality shares ankylosaurs and ner&Hooker, 1992) andthatofthefourfoundat dryosaurids/hypsilophodontidswiththetwosites Mangahouanga Stream, New Zealand, one is a intheAntarcticPeninsula,theonlytwodinosaurs probable dryosaurid, a family closely related to known to occur in the latterregion. & hypsilophodontids (Wiffen Molnar, 1989). There is no evidence such as tillites to suggest This explanation forthe apparent preponderance that continental ice sheets existed during the ofthese groups athighlatitude may wellbe true. Mesozoic at high latitudes (Frakes, Francis & SIGNIFICANCEOFPOLARDINOSAURS IN GONDWANA 715 TABLE 1. GondwanaPolarDinosaurLocalities. 1, Mt Kirkpatrick, Beardmore Glacier area, Plesiosaur Transantarctic Mountains, Antarctica, 165°E, Pterosaur 84°S Aves Rockunit. Falla Formation (fluviatile) Labyrinthodont: New genus and species (War- Age. Jurassic on stage-of-evolutionofdinosaurs ren etal., in press) & Palaeolatitude. 61°S (Smith, Hurley Briden Ceratodontidae: Ceratodus avus, Ceratodus 1981) nargun, Ceratodus sp., Coccolepis woodwardi, Palaeoenvironment. Forelandbasin flood plain Wadeicthys oxyops, Koonwarria manifrons, RecentReference. Hammer& Hickerson (1994) Leptolepis koonwarri Fauna. Cryolophosaurus ellioti, plesiomorphic allosaurid (Hammer, pers. comm. to Molnar 4, Antarctica, Antarctic Peninsula, James Ross 1995);Smalltheropod;Prosauropod;Tritylodon- Island, 57.9°W, 63.9°S tid; Pterosaur Rock unit. Santa Marta Formation (Marambio Group) 2, Australia, Southeastern Queensland, Taloona Age. Campanian Station nearRoma, 149°03±3'E, 26°05±3'S. Palaeolatitude. 65°S (Scoteseet al. 1988) Rockunit. WalloonCoalMeasures(InjuneCreek Palaeoenvironment. Nearshore marine Group) RecentReference. Gasparini,Pereda-S. &Molnar Age. Early Jurass\c [?Bajocian] (1996, this volume) Palaeolatitude. 65°S (Douglas & Williams Fauna. Nodosauridae & 1982), 56°S ( Smith, Hurley Briden 1981) Palaeoenvironment. Intracratonic flood plain 5, Antarctica, Antarctic Peninsula, James Ross Reference. Longman (1927) Island, 57.9°W, 63.9°S Fauna. Sauropoda: Rhoetosaurus brownei Rockunit. Hidden Lake Formation Age. Coni acian-Santonian 3, Victoria, Australia, numerous sites centred on Palaeolatitude. 65°S (Scoteseet al. 1988) 38°45'S, 143°30'E.(OtwayGroup)and38°40'S, Palaeoenvironment. Nearshore marine 145°40'E (Strzelecki Group) Recent Reference. Molnar, Angriman & Rock units. Strzelecki and Otway Groups Gasparini (1996, this volume) (fluviatile) Fauna. Theropod Age. Strzelecki Group Aptianexcept Koonwarra locality whichisAlbian; Otway GroupAlbian 6, Antarctica, Antarctic Peninsula, Vega Island, Palaeolatitude. StrzeleckiGroup, 77.8°S;Otway 57.5°W, 63/7°S Group, 66.8°S. Minimum estimates because of Rock unit. Cape Lamb Memberofthe Lopez de thermal overprinting between 75 & lOOmy BP BertodanoFormation (Whitelaw 1993). Age. Late Campanian - early Maastrichtian Palaeoenvironment. Rift valley flood plain Palaeolatitude. 65°S (Scoteseetal. 1988) & RecentReference. Rich Vickers-Rich (1994) Palaeoenvironment. Nearshore marine Ffltma.Hypsilophodontidae: Fulgurotherium RecentReference. Milner& Hooker (1992). australe Leaellynasaura amicagraphica, Atlas- Fauna. Hypsilophodontidae y copcosaurus loadsi, Victorian hypsilophodontid Type 1, Victorian hypsilophodontid Type 2, Vic- 7, New Zealand, North Island, Mangahouanga torian hypsilophodontidtype 3 Stream, 176°45'E, 39°S Ornithomimidae: Timimushermani Rockunit. Maungataniwha Sandstone Ankylosauria, cf. Minmi Age. LateCretaceous, Campanian-Maastrichtian Protoceratopsidae: Aff. Leptoceratops Palaeolatitude. 55°S (Scoteseetal. 1988) Oviraptorosaur? Palaeoenvironment Nearshore marine . & Carnosauria: Allosaurussp. RecentReference. Molnar Wiffen (1994) Testudines: Chelycarapookus arcuatus (tes- Fauna. Dryosauridae, Theropod, Ankylosauria, tudines) Sauropoda, Testudines, Plesiosauroidea, Testudines: Cryptodira Mosasauridae, Pterosauria 716 MEMOIRS OFTHEQUEENSLAND MUSEUM Syktus, 1992). However, it has been inferred on another four years, the major sites there may be thebasisoftheoccurrenceofdropstones aslarge worked out. Outcrops of the Wallon Coal as 3m across in finegrained marine sediments Measuresarerarein southeastern Queensland.In deposited during the late Jurassic and early the Mangahouanga Stream locality of New Cretaceous that winter seasonal ice did form at Zealandthefossil vertebratesare recoveredfrom highpalaeolatitudesinbothhemispheres(Frakes, loose boulders in the stream itself and dinosaur Francis & Syktus, 1992). remainsarequiterare.The RossIslandoccurren- In southeastern Australia, studies assessing the ces, like the New Zealand ones, are marine but mean annual palaeotemperature have been car- unlikethere,partialskeletonsratherthanisolated ried out on the sediments producing the bones have been found. Although Ross Island is dinosaurs. Palaeobotanical evidence based on unlikely to produce major concentrations of leafmargin and stomatal structure together with dinosaur remains, with persistent prospecting the overall composition of the flora have been overthe large available area, it may be expected taken to suggest a mean annual temperature of to continue to contribute significantly to our +10°C (Parrish et al., 1991) while an oxygen knowledge ofGondwana polardinosaurs. Clear- isotopeestimateis-6°C(Gregoryetal. 1989),the ly, to find significantly more information about difference between Chicago and Point Barrow, Gondwanapolardinosaursoutsidetheareaofthe Alaska, today. While the biological implications Falla Formation, new areas need to be inves- ofthesetwoestimatesarequitedifferent,theyare tigated such as the Cretaceous coalfieldsofNew concordantinthatthepalaeoclimatewasfarfrom Zealand (Rich, 1975). tropical. No matter what the palaeotemperature was, LITERATURE CITED polar dinosaurs would had to have adapted to CHATTERJEE,S. 1993.Shuvosourus,anewtheropod. prolonged periods of annual darkness each Research& Exploration9: 274-285. Winter. There have been suggestions that in the CLEMENS, W.A., & NELMS, L.G. 1993. Paleo- geological past the earth's rotational axis might ecological implicationsofAlaskan terrestrialver- have been significantly closer to being oriented tebrate fauna in latest Cretaceous time at high perpendicular to the plane of the ecliptic (e.g., paleolatitudes. Geology 21: 503-506. Douglas &Williams, 1982). Ifthishad occurred, CURRIE, P.J., VICKERS-RICR P., & RICH, T.H. the lengthofcontinuousdarkness each Winterat 1996. Possible oviraptorosaur (Theropoda, high latitudes would have been reduced. How- Dinosauria)specimensfromtheEarlyCretaceous ever, the criticism ofLaplace (1829) against the Otway Group of Dinosaur Cove, Australia. Al- deaergtrhe'essofbrlioqmuiittsyprheasveinntgorsihiefntteadtimonoraseitthdaonesaofveewr DONGcheZr.ing1a98250.: 7A3-7M9i.ddle Jurassic labyrinthodont a period ofabout 41,000 years (one of the com- (fSrionmobDraacshhyaouppsup,lacZeingtoincge,phaSliucshgueann.ePtrsopv.innocve..) ponents of the Milankovitch cycle), has never [Chinese with English abstract.) Vertebrata been refuted. Palasiatica23: 301-307. Ofall the polardinosaursites, that on the Col- DOUGLAS, J.G. &WILLIAMS,G.E. 1982. Southern villeRiver,Alaska(Fig. 1),hasthegreatestpoten- polar forests: the Early Cretaceous floras ofVic- tialtoyieldadetailedpictureabouttheseanimals. toria and their palaeoclimatic significance. Formorethan200kmontheleftbankofthatriver Palaeogeography, Palaeoclimatology, Palaeoecology39: 171-185. farreobmeaAultbiifaunlltyoeMxapaossteridcohutticarno.pFsotshsaitlrbaonngeesinhaavgee FRAKE'CSl,imLa.tAe.,MoFdReAsNCoIfSth,eJP.hE.an&erSoYzoKiTcU\S(,CaJ.mIb.r1i9d9g2e. been found at many places along that river. By University Press: Cambridge), xi, 274pp. contrast,thepotentialofsiteswhichareshownas GASPARINI, Z., PEREDA-SUBERBIOLA, X. & having less than 100 bones is much more MOLNAR, R.E. 19%, thisvolume. New dataon problematical. the ankylosaurian dinosaur from the Late The work on the polardinosaurs ofGondwana Cretaceous ofthe Antarctic Peninsula. Memoirs isjust beginning. Just how distinctive they were oftheQueenslandMuseum 39(3): 583-594. ufrnokmnotwheni.rlTohweerFlaatliltaudFeocromnatteimopnoraorfieAsntisaralcltbiucta GREGROIRCYH,, PR..VT..,&DROIUCTH,HITT.HT.,19C8.9B..,OxDyUgDeDnYi,sotIo.pRi.c, composition of carbonate concretions from the seems the most promising area at the moment in lowerCretaceous ofVictoria, Australia: implica- termsoffuturepotential.SoutheasternAustralian tions forthe evolution ofmeteoric waters on the outcrops are limited tojust 4 square kilometres Australiancontinentinapaleopolarenvironment. which have been thoroughly prospected. In Earthand Planetary Science Letters92: 27-42. SIGNIFICANCEOFPOLAR DINOSAURS IN GONDWANA 717 HAMMER, W.R. & HICKERSON, W.J. 1994. A Gondwana Origin? Research & Exploration 10: crested theropod dinosaur from Antarctica. 129-131. Science 264: 828-830. SCOTESE,C.R.,GAHAGAN,L.M.&LARSON,R.L. HOLTZ,T.R.Jr 1994.Thephylogeneticpositionofthe 1988. Plate tectonic reconstructions of the Tyrannosauridae: implications for theropod sys- Cretaceous and Cenozoic ocean basins. Tec- tematics.Journal ofPaleontology 68: 1100-1117. tonophysics 155: 27-48 LAPLACE, PS. 1829. 'MecaniqueCeleste', vol. II. SHISHKIN,M.A. 1991.Labirintodontizpozdneyyury LONGMAN, H.A. 1927. The giant dinosaur Mongolii. Palaeontological Zhurnal, no. 1, pp. Rhoetosaurus brownei. Memoirs of the 81-95.[AlabyrinthodontfromtheLateJurassicof MILNdQEiuRne,oesnAas.ulCra.nf&droMHmuOstOheeKuEumpRp9,:erJ1.-JC1.r8e1t9a9c2e.oAusnoorfntihtehoApno-d SMITHM'o,PnhAga.onBle.ir,ao.HzoUPiaRclLeEoPnaYtl,oelooAcg.oiMnct.ail&nJeonButRranIlaDlWE1oN:r,7l8Jd-.9C1.M.a1]p9s8'1.. tarcticPeninsula.JournalofVertebratePaleontol- WARR(ECNam,brAi.dAg.eU&niHveUrTsiCtyHIPrNesSs:ONCa,mbMr.iNdg.e)1.98130.2Tphp.e ogy 12:44A. MOLNAR, R.E., LOPEZ ANGRIMAN, A. & last labyrinthodont? A new brachyopoid (Am- GASPARINI,Z. 1996,thisvolume.AnAntarctic phibia, Temnospondyli) from the Early Jurassic Evergreen Formation of Queensland, Australia. Cretaceoustheropod.MemoirsoftheQueensland Museum39(3):669-674. Philosophical Transactions ofthe Royal Society, MOLNpAolRa,rRd.Ei.no&saWuIrFFfEaNu,naJ. 1f9r9o4m.ANleatweCrZeetaalcaenodu.s WARRTSEe.rHNi.,es&AB.,AV.3I,03CK:KOE1O-RL6S2,.-RL.I,CCHL,EEP.LA1N99D1,. MA.,nREIaCrHl,y Cretaceous Research 15: 689-706. Cretaceous labyrinthodont. Alcheringa 15: 327- PARRISH, J.T., SPICER, R.A., DOUGLAS, J.G., 332. RICH, T.H. & V1CKERS-RICH, P. 1991. Con- WARREN,A.A.,RICH,T.H.&VICKERS-RICH,P.in tinental climate near the Albian South Pole and press. The last, last labyrinthodont? Palaon- comparison with climate near the North Pole. tographica,Abt.A. Geological Society of America Abstracts with WHITELAW, M.J. 1993. Paleomagnetic Paleolatitude Programs23: A302. determinations for the Cretaceous vertebrate RICH, T.H. 1975. Potential pre-Pleistocene fossil localities of southeastern Australia-high latitude tetrapodsitesinNewZealand.MauriOra3:45-54. dinosaurfaunas. Journal ofVertebratePaleontol- RICH, T.H. & RICH, PV. 1989. Polar dinosaurs and ogy 13, supplementtoNumber3: 62A. biotas of the early Cretaceous of southeastern WIFFEN, J. & MOLNAR, R.E. 1989. An Upper Australia.NationalGeographicResearch5: 15-53. Cretaceous ornithopod from New Zealand. RICH,T.H. &VICKERS-RICH,P. 1994. Neoceratop- Geobios - Paleontologie, Stratigraphie, sians & Ornithomimosaurs: Dinosaurs of Paleoecologie 22: 531-536.