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Descriptions of a new genus and two new species of viviparid snails (Mollusca: Gastropoda: Viviparidae) from the Early Cretaceous (middle-late Albian) Griman Creek Formation of Lightning Ridge, northern New South Wales PDF

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Preview Descriptions of a new genus and two new species of viviparid snails (Mollusca: Gastropoda: Viviparidae) from the Early Cretaceous (middle-late Albian) Griman Creek Formation of Lightning Ridge, northern New South Wales

DESCRIPTIONS OF A NEW GEMS AND TWO NEWSPECIES OF VIVIPARIDSNAILS (MOLLUSCA: GASTROPODA: VIVIPARIDAE) FROMTHE EARLY CRETACEOUS (MIDDLE-LATEAI.BIAN)GRIMANCREEKFORMATION OFLIGHTNING RIDGE, NORTHERN NEW SOUTH WALES ROBERTJHAMILTON-BRUCE. BRIANJ SMITH & KAREN LGOWLETT-HOLMES HAMILTON BRUCE. RJ.SMITH. BJ &OOWLETT-HOLMES. KL. 2002.Descriptionsofa How genus and two new species of viviparid snails (Mollusea: Gastropoda: Vivip;ai((ae) from the Early Cretaceous (middle-lute Albianl Griman Creek Formation of Lightning Ridge, northern NewSouth Wales. Records ofihe SouthAustralianMuseum 35(2): 193-203. Fossil gastropods belonging to the family Viviparidae (Caenogastropodn tProsobranehuO) are described from Hie Early Cretaceous (middle- late Albian) Griman Creek Formation of Lightning Ridge, northern New South Wales From our research, this is the earliest definitive record of non-marine gastropods from Australia and amongst the oldest viviparid material in 'he world recorded todale. A newgenusand twonew speciesare described, andothermaterial jiiiihniaiile 10 |de extant genus Nniopala is recorded. Implications for the current distribution ofthe Viviparidae aieaddressed, "•Robert .1 Hamilton-Bruce, South Australian Museum, North Terrace, Adelaide,. South Australia 5000; Brian J Smith. Queen Victoria Museum. Wellington Street, Launeeston. Tasmania 7250, Australia; Karen L Gowlctt-Holmes, CSIRO Division of—Marine Research, GPO Bo\ 1538, Hobart, Tasmania7001. Australia; ^corresponding author email: Hamilton bruce.rohertts'saugov.sa.gov.au Manuscriptreceived 29July 2002. Opalised fossils have long been known from distributionofextant viviparid taxa arediscussed. the Lower Cretaceous (middle-laie Albian) The Viviparidae is a cosmopolitan group of freshwater deposits of the Griman Creek freshwater caenogasiropods, characterised by Formation, Lightning Ridge, New South Wales medium- to large-si«:d lurbiniform shells which (Smith 1999). To date, much of the published possess a rounded hody whorl, moderately high, work has focused on Fish (Kemp 1991; Kemp & pointed spire; wide, round aperture; and sub- Molnar 1981) and terrestrial tehapod remains spiral, horny operculum (Smith 1992).Thecurrent (Archer ct al 1985; Flannery et a| 1995; Molnar Australian distribution ofthe group is limited to a I'WJa, 1980b, 1991, Molnar & Gallon 1986; Rich few species occurring in lite large drainage basins et al 1989) with relatively few reports (Detimauet that span much of ihe arid centre and northern al 1992; McMichael 1957; Smith 1999) tropical regions ofthe continent summarising the vast quantities of invertebrate The fossilrecord forViviparidae is known from material recovered from the area. The recent the Jurassic toRecent {Vniparus Monitor!, 1810), opportunity to examine Lightning Ridee with a tentative report based on an internal shell gastropod specimens (brought to our attention by (Wemicia Cox, 1927) mould (possibly of marine Ben Kear. South Australian Museum, as part of origin) from the Lower Carboniferous ofEngland an ongoing assessment of vertebrate and (Brookes-Knight et al 1960). Within Australia invertebrate fossil material from the locality) from there are few records of fossil viviparids. the Collections ol the Australian Museum. Sydney, Elheridge (1902) described the earliest potential and several private individuals has prompted a Australian taxon, VMporus (?) atba-scopularis systematic appraisal of the gasiropods within ihe Elhendge (alsonotedby Newton(1915)) fromthe material recovered. This study describes Aprian marine deposits of Ihe Doncastcr Member specimens attributable to the family Viviparidae (Wallumbilla Formation \ensu Burton & Mason (Caenogastropoda (= Prosobranchta)), including a I998), White Cliffs, New South Wales This new genus and two new species. Indeterminate specimen is currently under examination by the material belonging to the exiant genus Noiopala authors and at present is regarded as being of is also recorded, and the implications for unclear viviparid affinity. Cotton (1935a) erected 194 RJ HAMILTON-BRUCE,BJSMITH, KLGOWLETT-HOLMES a species of Notopala (N. wanjacalda) from late present whorls across each side of the shell; the Pleistocene sediments along the Murray River point at which they met was assessed as being near Sunnyside, South Australia, and also noted a approximately the original apex ofthe shell (Cox second taxon (Notopala sp.) from the same 1960). All specimens are deposited in the deposits, which showed strong similarity to the Australian Museum (AM), Sydney. All extant N. hanleyi (Frauenfeld, 1862). Viviparid specimens were prepared using a diamond drill- snail shells from Early Cretaceous deposits in the bit attached via a flexible lead to a high-speed Lightning Ridge area were recorded but not rotary motor. Shell measurements were made to described by Dettman et al (1992) and Smith the nearest 0.05 mm using dial calipers. (1999); the latter also recorded possible representatives of the Naticidae, Thiaridae and Ellobiidae. Few other Australian non-marine Systematics gastropod fossils (all of Tertiary age) have been recorded (Archer et al 1994; Arena, 1997; ClassGASTROPODA Chapman 1937; McMichael 1968). ORTHOGASTROPODA Subclass SuperorderCAENOGASTROPODA MaterialsandMethods OrderARCfflTAENIOGLOSSA All specimens described herein are derived from the Lightning Ridge opal fields (exact mine SuperfamilyAMPULLARIOIDEA localities from which these specimens originated are unknown), Surat Basin, northwestern New Family VIVIPARIDAEGray, 1847 South Wales. The opal-bearing sediments in this area form part of the Griman Creek Formation, a unit dated as middle-late Albian in age (Dettman Diagnosis et al 1992). The deposit predominantly reflects a Medium to large dextral, turbiniform shells, coastal plain facies (Burger 1988; Dettman et al body whorl rounded, spire moderately high, 1992) situated in anEarly Cretaceous high latitude pointed; aperture wide, round, parietal fold zone (-70° S, Embleton 1984). Invertebrate present or absent; operculum horny, subspiral fossils, plant root impressions and vertebrate (modified afterSmith 1992). taphonomy suggest a freshwater estuarine to lacustrine setting (Dettman et al 1992; Molnar Remarks 1980a). Palaeoclimatic indicators imply strongly The above diagnosis follows Smith (1992), seasonal conditions with sea-level isotopic modified to accommodate the presence of a palaeotemperatures in the EromangaBasin/ Surat parietal fold in the new genus described below. Basin regions ranging from 11.9CC (northeast) to Viviparid snails are, as their name suggests, 16.3°C (southwest) (Dettman et al 1992; Stevens viviparous (live bearing) and are found in both & Clayton 1971). lotic and lentic systems throughout the world Designation of parietal fold position on the (Browne 1978). Within Australia the family is specimens was achieved by positioning the fold currently represented by the extant native genera within the aperture on a 360° compass setting Notopala, Larina and Centropala (Smith 1992). when a vertical line through the axis, juxtaposed A species accidentally introduced from Asia, to the column, is intersected by a horizontal line Bellamya heudei guangdungensis (Kobelt, 1906), corresponding to the midpoint of the aperture. recorded by Shea (1994) as established in New The adapical axis above the point of intersection South Wales, is not considered part of the is taken as zero degrees. The diameters of all Australian fauna in this study. The Australian shells were measured following the method of members ofthe family have undergone substantial Boycott (1928) and are defined as 'the greatest revision over many years (Cotton 1935a, 1935b; dimension that can be found starting with the Sheldon & Walker 1993; Stoddart 1982); the use edge of the lip to a point on the opposite side of of intraspecific shell variation and morphometric the shell on the last whorl'. To enable data by recent authors has resulted in a substantial extrapolation of the numbers of missing whorls, reduction in the numbers of accepted species. the incremental angle of the shells was judged Shell colouring and pattern and the form of the by drawing a line so as to touch each of the operculum, which are important characters in EARLYCRETACEOUS VIVIPARIDSNAIt.S 195 determining generic placement in this family, arc are in any way intended to suggest relationships characters lost in most fossils. between members of either the Notopala sp. The present specimens from Lightning Ridge presented here or the new genus and any extant are over 100 million years old (Albian [108- species. 97.5 MaJ) and totally opalised. While sculpturing on some specimens has been preserved, colour- bands and opcreula have not; therefore, only GenusAlbianopalin gen. nov. Structural shell characters could be used to assign them to taxa. Two major factors have been taken Diagnosis into account—before deciding to place them in Shell dextral, thick, solid, globose to Vivipandae the shell morphology, which fits subglobose, ventricose, three-five whorls, within the currently accepted diagnosis of the subumbilicale; aperture subovate, large, family widi only slight modification (the presence approximately cqua] to height of spire; parietal of the parietal fold in Ihe two new species); and fold present, simple. Operculum unknown. the freshwater depositional environment from which they originated. Whilst all the specimens Type species can be accommodated within the family Albianopalin benkeari sp. nov. Viviparidae, those possessing a parietal fold cannot be assigned to any currently recognised Etymology genus and are considered to belong to a new From the combination ofAlbian, n. referring to genus which is described below. Direct age ofthe LowerCretaceousopal-bearingdeposits comparisons between species, living or fossil, are of the Griman Creek Formation, and opalin. few (Brown 1980); however, due to the lack of Middle English, from apalus, Latin, alteration of colour patterns and opcreula as well as soft parts Greekopallios, for 'opal'. in these fossils, the only remaining methods are based on morphology. Therefore, we have applied Remarks parts of the morphometric data gathered by While this new genus has similarities to Sheldon and Walker (1993) to justify the Notopala (Cotton, 1935b), Albianopalin gen. nov. placement of these specimens within Viviparidae, is easily separated from Notopala and all other and in one case in the extant genus Notopala, and genera within the family by the presence of a to offer a method of comparison between extant parietal fold in the basal lip, currently a feature andextinct species. None ofthe data comparisons unique to this genus within the family. B RCil.JRE 1. AMFI22185Albaiw/uilin benkearisp. nov. in A,apertural and H.dorsal views. 196 RJ HAMILTON-BRUCE,BJSMITH, KLGOWLETT-HOLMES Etymology Named for Ben Kear, for his assistance in this study. Remarks Holotype unique. The position of the parietal fold separates A. benkeari sp. nov. from the other new species described below. The holotype specimen is undistorted, but the spire is incomplete. Extrapolation from the spire angle indicates that there may have been at least four and as many as five complete whorls originally. Being opalised, the state of preservation of the holotype is very good, with much of its detailed surface ornamentation preserved. Albianopalin Uzsmithae sp. nov. (Figs 3, 4, Appendix) FIGURE 2. AMF122185Albanopalin benkeari sp. nov. inapertural view showingangleofparietalfold. Diagnosis With the features of the genus; parietal fold simple, 112° from the vertical. Albianopalin benkeari sp. nov. (Figs 1, 2, Type specimen Appendix) Holotype: AMF122186 (Figs 3, 4, Appendix) Diagnosis Localityandhorizon With the features of the genus; parietal fold Lightning Ridge opal fields (precise site within simple, 70° from vertical axis. the opal fields unknown), northern New South Wales, Surat Basin, Griman Creek Formation, Typespecimen middle-late Albian. 'Holotype: AMF122185 (Figs 1, 2, Appendix) Description Localityandhorizon Shell, 15.1 mm high, 12.9 mm maximum Lightning Ridge opal fields (precise site within diameter, dextral, turbiniform, subglobose. the opal fields unknown), northern New South Teleconch with two complete whorls and broken Wales, Surat Basin, Griman Creek Formation, parts, pointing to the possible presence of further middle-late Albian. whorls. Whorls impressed. This particular specimen has undergone some dorsoventral Description compression which has resulted in distortion, Shell 17.7 mm high, 16.4 mm maximum especially at base of final whorl. Aperture large diameter, dextral, turbiniform, subglobose. (9.95 mm high), round; basal outer lip varicose Teleconch with three complete whorls and broken with no evidence of eversion. A single, basal mm mm parts, pointing to the possible presence of further parietal fold (3.75 long, 2.3 wide, whorls. Whorls impressed. Relatively evenly 2.2 mm deep) is present on surface of basal lip, spaced spiral prosocyrt ornamentation present on deepest side facing adaperturally (Fig. 4). Spiral many areas of teleconch. Aperture large prosocyrt ornamentation present on parts of mm (10.85 high), round; basal outer lip varicose teleconch, especially proximal to aperture. with no evidence of eversion. A single, triangulated parietal fold (3.0 mm long, 1.0mm Etymology wide, 2.0 mm deep) present on upper surface of We name this species for our colleague basal lip, deepest side facing adaperturally (Fig. Elizabeth (Liz) Smith of the Australian Museum 2). (AM), forherassistance in this study. EARLYCRETACEOUS VIV1PARIDSNAILS 197 HH A B FIGURE3. AMF122186Albunnpulinlizsmiilwesp, nov. in A,apertural andB,dorsal views. Remarks Holotype unique. The position of the parietal fold separates A. Uzsmithae sp. nov. from A. benkearl sp. nov. as described above. The holotype has undergone moderate dorsoventral compression, resulting in the specimen appearing stretched. Like A. benkeari sp. nov., the spire is incomplete, andextrapolation from [he spire angle indicates that there originally may have been at least four and as many as five complete whorls. Being opalised and having damage to parts ofthe surface of the teleconch, the state of preservation is good enough to preserve some of its detailed surfaceornamentation. Genus Notopala (Cotton, 1935b) Type specimen Type species: Paludina hanleyi (Frauenfeld. 1864)by original designation. Diagnosis Shell dextral, globose-conic, subumbilicaic, five whorls, ventricosc to angulate below the periphery; aperture subovate, large, about equal to FIGURE 4. AMFI22186 Albanopalin Uzsmithae sp. height of spire; parietal fold absent; operculum nov. inapcrluralview showingangleofparietal fold. corneous. 198 RJ HAMILTON-BRUCE, BJ SMITH, KLGOWLETT-HOLMES B FIGURE5. AMF13001 Notopalasp. inA, aperturaland B,dorsal views. Remarks lip varicose with no evidence ofeversion. These specimens are extremely significant. Until theirdiscovery, none ofthe currently living Remarks native Australian viviparid genera were known There is considerable variation in size, state of from deposits older than Plio-Pleistocene. This preservation, degree ofcorrosion and/ordamage to temporal range is nowextendedback tothe Lower the shells, opalisation, amount ofmatrix deposition Cretaceous (middle-late Albian). It is also on the shells, completeness of the aperture and interesting to note that another living species, the presence/absence of ornamentation. We have not lungfish Neoceratodusforsteri, has been recorded described these specimens as a new species at this from the Griman Creek Formation of Lightning stage. We feel that the exact taxonomic position of Ridge (Kemp & Molnar 1981). this material compared with other species in the genus requires further study. The preceding table Notopala sp. (Fig. 5, Appendix) (&TabWlael1k)e,rdia1g9r9a3m)(Faign.d6,gmroadpihfie(dFifgr.om7)Sheslhdoown Referredmaterial measurements of shell characters for each of the AMF13001 (Fig. 5), AMF122166-AMF122184 living Australian speciesofNotopala, andillustrate morphometric similarity of the specimens of (notfigured herein). Albianopalin nov. sp. as well as Notopala sp. Localityandhorizon described here to existing members of the genus. Lightning Ridge opal fields (precise site within However, the specimens of Notopala sp. vary the opal fields unknown), northern New South greatly in their physical condition, and it has not Wales, Surat Basin, Griman Creek Formation, been possible to take the full range of middle-late Albian. measurements fromeachofthe 20specimensbeing studied. The reason forthe inclusion ofthis data is Description not to demonstrate any relationship between extant Shell dextral, turbiniform, subglobose. species and the specimens from Lightning Ridge, Teleconch with two to three complete but to show that the physical characteristics ofthe impressed whorls; the incremental angle of the new material {Albianopalin sp. nov. and Notopala shell indicates that there were originally four or sp.) fit within the parameters for inclusion in the five whorls. Aperture large, round; basal outer familyViviparidae. EARLYCRETACEOUSVIVIPARIDSNAILS 199 ApertureLip tcngth (ALL) Umbilical "^ widih CVMB) Shell length (SHL) FIGURE 6. Diagram illustrating measurement parameters used in the morphomctric analysis (modified from Sheldon&Walker 1993). APL SHW UMB APW ALL CH Notopalasp.. gen. N. waterhousii >'.:••$:.•>:' "Banded" Shells N. essingtonensis Notopalasublineata (Cooper) N. alisoni Notopalasublineata (Darling) A. benkeari nov, sp, * I * N. hanleyi A. lizsmithae nov. sp. I = Standard deviation * single sample FIGURE7. Histogramshowingresultsofmorphometricanalysis.TaxaincludelivingspeciesofNotopala(modified from Sheldon & Walker 1993), Notopala sp. (AMF1300I, AMF122185-122184), Alhianopalin benkeari (AMF122185)andA. lizsmithae(AMF122186). 200 RJ HAMILTON-BRUCE, BJ SMITH. KLGOWLETT-HOLMES TABLE 1. Mean shell measurements (mm) for various living species of Notopala (*modified from Sheldon & Walker 1993) compared with those for Notopala sp. (AMF13001, AMF122185-122184), Albumopalin benkeari (AMF122185) and A. lizsmithae (AMF122186). Because not all specimens of Notopala sp. were complete, the number (n) ofspecimens used foreach individual parameter is indicated separately. Standard deviation (SD) rows were not includedforA. benkeariandA. lizsmitliaeasonly asinglespecimen hascurrently been recovered foreach ofthese taxa. Abbreviations: SHL, shell length; APL, apertural length; SHW, shell width; UMB, umbilical width; APW, aperturewidth; SPL, spirelength; ALL, apertureliplength. Species Number SHL APL SHW UMB APW SPL ALL *N. alisoni 20 21.59 11.91 16.3 02.59 09.77 14.78 05.35 (Brazier, 1979) SD 0.086 0.039 0.054 0.014 0.034 0.064 0.015 *N. essingionensis 20 21.70 12.00 16.58 1.96 10.10 14.47 6.11 (Frauenfeld, 1862) SD 0.082 0.038 0.067 0.016 0.038 0.067 0.025 *N. waterhousii 10 38.59 22.84 33.14 4.63 18.88 25.01 10.02 (Adams&Angus, 1864) SD 0.209 0.105 0.164 0.024 0.087 0.131 0.043 *N. hanleyi 42 16.67 10.50 13.83 1.29 8.52 11.00 5.57 (Frauenfeld, 1862) SD 0.520 0.026 0.036 0.00 0.025 0.037 0.013 *N. sublineata 28 18.32 10.84 14.53 2.05 9.21 12.04 5.24 (D)(Conrad 1850) SD 0.063 0.024 0.040 0.012 0.026 0.039 0.016 *N. sublineata 35 19.46 11.06 15.52 2.21 8.89 12.59 4.93 (C)(Conrad 1850) SD 0.058 0.027 0.039 0.010 0.018 0.040 0.009 'Banded' shells 15 22.15 13.18 18.10 2.06 10.69 14.6 6.42 SD 0.110 0.053 0.008 0.011 0.045 0.079 0.029 Notopala sp. 20 - 10.35 (19) 14.43 (20) 3.24(10) 9.20 (15) - 4.71 (9) SD - 2.563 3.053 0.616 1.80 - 1.071 — A. benkeari 1 17.7 10.85 16.4 3.35 9.85 6.2 A. lizsmithae 1 15.1 9.95 12.9 3.4 8.15 - 4.4 Discussion climatic conditions which characterised many Australianhigh latitudedepositional environments Albianopalin benkeari sp. nov., A. lizsmithae duringtheEarly Cretaceous. sp. nov. and Notopala sp. share the distinction of The implications of this study, combined with being the three oldest known definitively assigned subsequent investigations, could eventually members ofthe Viviparidae in Australia and thus demonstrate a Gondwanan and possibly serve to extend the range of the family in this Australian radiation formany ofthe species living region back to at least the uppermost Early around the globe today. Cretaceous. Since the family isconfidently known from theJurassic toRecentinEurope, Viviparidae Acknowledgments therefore appears to be an ancient pre-Jurassic group of probable Pangean origin. By the This paper would never have been initiated without Cretaceous, the family had diversified within the Ben Kear, who, since its inception, has more than Gondwananregioninto arangeofendemic genera earned oursincere thanks. We wish also tothank Philip and species. Ryan for examining the statistical methods employed. While there are numerous Cretaceous records Greg Brown and George Koulis ofthe Royal Adelaide offreshwater bivalves from Australia (Dettman et Hospital (RAH) helped considerably by contributing al 1992; Hocknull 1997; Jell & Duncan 1986; high-grade imaging. Elizabeth (Liz) Smith. Bob Jones, Ludbrook 1985; McMichael 1957), there are very Henk Godthelp, Tom Rich, Rolf Schmidt and Neville few for non-marine gastropods from the same Pledge supported this work and generously permitted period. The reasons for this apparent absence are access to their collections. Philip Clarke of the South Australian Museum, Adelaide (SAMA), gave financial unknown, but could be related to preservational support and encouragement. Anne Hamilton-Bruce of biases (with shells rapidly breaking up or The Queen Elizabeth Hospital (TQEH) contributed dissolving after death). Another possibly reason lateral thinking and assistance with editing. Chris lzzo could be a lack of gastropod species due to poor (zoology student at Adelaide University) assisted with tolerance of the strongly seasonal near-freezing measuring and statistical data analysis. Origin Energy, EARLYCRETACEOUS VIVIPARIDSNAILS 201 The Advertiser newspaper. Coober Pcdy Tomism Rozefields, A. 1992. Australian Cretaceousterrestrial Association, llie Souih Australian Museum and the faunas and floras: biostratigraphic and biogcograptiK WalcrhouscClubgaveothersupporttothisproject. implications. Cretaceous Research 13: 207-262. Ethendge. R 1902. A monograph of the Cretaceous invertebrate fauna ofNew South Wales. Memoirs of References theGeologicalSuneyofNewSouth Wales11: 1-98. Bmbleton, BJJ. 1984. Australia's global setting: past Archer, M, fl.Hirtery, TF, Ritchie, A & Molnar. RE. global settings. Irt JJ Veevers (ed). 'Phanerozoic 1985. First Mesozoie mammal from Australia. Earth History ol Australia', pp. 11-17. Clarendon Nature318 3f>3-3MS. Press: Oxford. Archer, M. Hand, S &Godthelp, H. 1994. 'Riversleigh: Flannery.TF.Archer, M.Rich.TH& Jones.R. 1995. A theStoryofAnimalsinAncientRainforest*ofInland new family of rnonotrcmes from the Cretaceous of Australia". Reed Hooks: Sydney Australia.Nature377: 418-420. Arena, DA. 1997. The palaeontology and geology of Hoeknull, S. A, 1997. 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Quarterly Wales 81 222-242. Notes. Geological Survey ofNew South Wales 107: McMichael, DF. 1968 Non-marine Mollusea from HI, Tertiary rocks in Northern Australia. Bureau of Chapman, F. 1937. Chert linieslone with Planorbis, Mineral Resources. Geolog( and Ge.uph\ncs from the Mount Elder Range. Western Australia. Bulletin 80: 13.1-160. ProceedingsoftheRo\'alSociety ofVictoria 50' 59- Molnar, RE. 1980a Procoelous crocodile from the NSW 68. Lower Cretaceous of Lightning Ridge Conon, BC. 1935a. Recent Australian Vtvipandae and MemoirsoftheQueenslandMuseum 20: 65-75. ;i fossil species. Records of the South Australian Molnar, RE 1980b. Australianlate Mesozoic Urrcsliial Museum5: 339-344. tetrapods: someimplications. Memoirsde lesSociele Ceuton. BC 1935b. The Australian viviparous river GeologiquedeFrance 139: 131-143. snatfs. Victorian Naturalist52: 96-99. Molnar, RE. 1991. Fossil reptiles in Australia. In P. t"o\, LR. I9W). 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A second Australian Notopala (Gastropoda: Prosobranchia: Cretaceous mammalian specimen from Lightning Viviparidae). Journal of the Malacological Society Ridge, New South Wales, Australia. Alcheringa 13: ofAustralia 14: 59-71. 85-88. Stevens, GR & Clayton, RN. 1971. Oxygen isotope Shea, M. 1994. The Chinese viviparid snail Bellamya studies on Jurassic and Cretaceous belemnites from heudei guangdungensis (Kobelt, 1906) in Australia New Zealand and their biogeographic significance. (Prosobranchia: Viviparidae). Molluscan Research New Zealand Journal of Geology and Geophysics 15:3-11. 14: 829-897. WWK Smith, BJ. 1992. Non-marine Mollusca. In Stoddart, JA. 1982. Western Australian viviparids Houston (ed). 'Zoological Catalogue of Australia' (Prosobranchia: Mollusca). Journal of the Vol. 8,pp. 1-398. AustralianGovernmentPublishing MalacologicalSociety ofAustralia5: 167-173. Service: Canberra.

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