An ecologic and biogeographic study of a new Tertiary land snail from mideastern Queensland (Pulmonata: Caryodidae)

7 Pages·1994·2.1 MB·English

AN EC0L0G1C ANDBIOGEOGRAPH1CSTUDY OFA NBW
TERTIARY LAND SNAIL FROM M1DEASTERN QUEENSLAND
(PULMONATA: CARYODIDAE)
JOHN STANISIC
Stamsic.J. 199406 01: An ecolngic and biogeographic study ofa tic* tertiary land snail
From midcastcrn Queensland. (Pulmonaia: Carvodidae) Memoirs of the Queensland
Muxtrum3S(\): 241-247 Brisbane. ISSN0079-8835.
Anewfossillandsnail,Praecaryodesantiquatagen.clsp.nov. isdescribedfromtheMiddle
lo Late Eocene deposits of the Rundle Formation, near Rockhamplon, mideastern
Queensland. The species is assigned to the Caryodidac. The phyletic, biogeographic and
eeolugical implications of this discovery are discussed. The new species suggests early
diversificationoftheCaryodidac.M^//^vc</.C«r>W/c/fle.Praecaryodesantiquaiasp.nov,,
l
ecology, btogeography, Eocene, landsnail.
John Sianisic, Queensland Museum, PO Box 3300, South Brisbane Queensland 4101
,
Australia; 14 February 1994.
Few Australian fossil land snails are known quence containing oil shale seams. These richly
and most arc from Pliocene-Pleistocene sedi- fossiliferous beds contain ostracods (Beaslcy,
mentsofsouthernAustralia(e.g.Strzclccki, 1845; 1945), fish (Hills, 1943), reptile fragments
Johnston, 1880; Ludbrook, 1978, 1984). Me- (crocodile and turtle) and flora (Foster, 1979:
Miehae! (1968) recorded Meracomelon lloydi Poster&Han-is, l981,Ruwett, 1988). Freshwater
from the Carl Creek limestones of north-west gastropods ( 'Planorbid-ae) are common in some
QueenslandandtheNorthernTerritory; Kershaw unils (Coshcll, 1983). The land snail remans
(1988ft) discussed several fossil land snails of werecontainedintheCiaystoneUnitoftheRam-
problematic affinityfrom Pleistocenedepositsin sayCrossingSeam withintheRundle Formation.
theKentGroupofislands, BassStrait;andHillet Little data are available to independently cor-
at. (1970) recorded Pedinogyra sp. (family relate theTertiary Basins ofcentral and northern
Caryodidae) fromalimestonequarryatMarmor, Queensland and most age determinations are
south of Rockhampton, mideastern Queensland based on long distance correlation with Tertiary
(hereafterMEQ).ThediscoveryofPraecaryodes sequences in Southern Australia. It has yet tobe
antiquatagen.clsp.nov. inMiddletoLateEocene demonstrated that the northern Australian Ter-
MEQ
deposits of is a significant addition to the tiary floras can be conveniently reconciled with
Australian fossil land snail fauna. known *porc-pollcn ranges from southeastern
Theageofthefossil and its relationshipstothe Australia. Given these shortcomings, the most
extant Gondwanan family Caryodidae provide a recent age estimate of the Rundle Formation is
basis for biogeographic speculation about the Middle to Late Eocene (Day et al.. 1983).The
evolutionofland snails in eastern Australia. The depositional environment of the Ramsay Cross-
climatic and habitat shifts which have occurred ing Scam was interpreted as shallow lacustrine
in eastern Queensland since the early Tertiary with lacustrine and pedogenic phases (Coshell,
have favoured the development of dry-adapted 1983)
Species, This provides an ecological framework Occurrence of land snails among the
fordiscussingcaryodid history in the region. predominantlyfreshwaterfossilremainssuggests
All material studied is in the collections ofthe that theyaredriftmaterialwhichhasbeenwashed
Queensland Museum (hereafterprefixed QMF). into the lagoon from surrounding areas
Henstridgc &. Misscn (1982) postulated a similar
STRATIGRAPHY, AGEAND mechanism to explain the high levels of car-
PALAEOENVIRONMENT bonaceous mailer among the oil shale.
Information on the contemporaneous vegeta-
NW
The Narrows Reds (Fig. 1), ofGladstone, tion present in the area surrounding the Rundle
MEQ,which includetheCurlewandRundleFor- Formationdepositionalenvironmentisnotavail-
mations and Worthington Beds (Henstridgc and able. However, the presence of large amounts of
Missen, 1982) are a Tertiary non-manne se- carbonaceous material in Ihe oil shales and the
.
242 MEMOIRSOFTHEQUEENSLAND MUSEUM
the more advanced sigmurethran pulmonatcs.
1SCCC
Tillier(I989)rejectedthisarrangementin favour
of a single family. Smith (1984, 1992) also
preferred to consider the Australian species as a
distinct family and this system is followed here.
The Caryodidae includes an unusual mix of
shelltypesrangingfromthedepressedlyconoidal
Anoglypta Martens, 1860 and giant bulimoid
Hedleyella Iredale. 1914 to the lenticular, thin-
shelled Pandofella Iredale, 1933 and the flat-
coiled, heavy-shelled Pedinogyra Albers, 1860.
This led Iredale (1937) to place these genera in
several different family units. Only thirteen ex-
tant speciesare known (Smith. 1992). All live in
\\_ds Gladstone
the moistforests ofeastern Australia.
Praecaryodesgcn.nov
CUHI£W FORMATION
: .... :!... Diagnosis
Shell large and subglobose with a flattened
MUfrrHTNCTON pawwnoM spire. Shell sculpture predominantly radial with
weak spiral grooves on the body whorl. Um-
bilicus absent. Aperture ovate; lip thickened and
FIG. 1.Locality mapshowing the Rundlc Depositand reflected.
theTertiarystratigraphicunitsintheNarrowsGroup,
Map adapted from Henstridge & Missen (1982). TypeSpecies
Scale line= 1km.
Praecaryodes antiquata sp.nov.
inference of high rainfall and humidity Etymology
&
(Henstridge Missen. 1982) suggest a tropical Acombinationofthe Latinpraemeaningearly
to subtropical closed forest. Fossil flora of the andthenominategenusofthefamilyCaryodidae.
younger Curlew Formation indicates deposition
in a low energy, freshwater swamp in an area of Praecaryodesantiquata sp.nov
marginal tropical/subtropical closed forest.
(Figs 2-5)
These interpretations correlate with a broader
perspective of vegetational change in eastern
MaterialExamined
Australiaduring theEocene (Kemp, 1981)point- Holotype: QMFM905, Rundlc, MEQ. Collected by
ing to a widespread closed-canopy rainforest
early in theepoch,becomingmorerestricted (i.e. SPoAuRtAhTeYrnPEPSa:cifQiMcFPIe3t4ro5l1e,um,QMAFugIu7s6t4,2[,9852.specimens.
replacedby sclerophyllouselements) in the Mid- Unit T claystone, Ramsay Crossing Seam, Rundlc
dle to Late Eocene when climate began to fluc- Formation,MEQ,opencut, L. CoshcJI.
tuate.
Etymology
SYSTEMATICPALAEONTOLOGY
Referring to the antiquity ofthe specimen.
Family CARYODIDAE Description
Thiele, 1926 Shell large (damaged and distorted), sub-
globose, diameter 41.94mm with 4% normally
Australian acavoid land snails havebeen tradi- coiled whorls, the last greatly expanded and not
tionally associated with the mesurethran family descending in Front. Spire and apex flat. Height
Aeavidae which also included a number of 28,30mm, Height of body whorl 21.62mm.
species from Africa. India and South America Protoconch 2 whorls, sculptured with fine,
(Zilch, 1959-60). Solem (1978a) separated the crowded, weakly protractively sinuatcd radial
Australian genera as a separate family, the ribs. Post nuclear sculpture of numerous, fine,
Caryodidae, and placed it and the Aeavidae near protractively sinuated radial ribs, becoming
A NEW TERTIARY LANDSNAIL 243
from the drier vine thickets of eastern
Queensland,hasalarge, globoseshell withoutan
umbilicusandwithconspicuouscancellatesculp-
tureon the body whorl. However theprotoconeh
is smooth and most importantly, the lip is noi
reflected.
Biogeogmpbically, these eastern Australian
camacnids are considered to be post-Miocene
northern immigrants (Bishop, 1981), hence ap-
pearing in the region much later than P. antt-
quota
DISCUSSION
PIG. l.Praecuryodes ontiffiwia sp now, QMF13905, The Middleto Late Eoceneage off antiquata
holotype. Drawing to show depressed shape and represents a marked chronological extension of
rhiekened lip(arrowed).[Drawingby Alison Hill]. the fossil record of land snails in Australia and
significantly predates previous reeouU
slightly less conspicuous on the body whorl, and (Pliocene/Pleistocene). The presence of a
irregularly spaced spiral grooves which arc par- reflected lip, a probable derived characterwithin
ticularlynoticeableonthebodywhorl. Umbilicus the Caryodidae, has important implication* fur
absent. Sutures weakly impressed. Whorls flat- thetimingofthe firstappearanceofthe familyon
tened above and below a weakly keeled the Australian land mass.
periphery.Apertureovate.Lipstronglythickened Abriefreappraisaloftherelationshipsofextant
and reflected Columella inserted obliquely. caryodid genera is a necessary preambi.
Parietal callusobscured. biogcographic considerations.
Remarks Caryodjd RELATIONSHIPS
In spite of the small amount of the mulctial Caryodids have an unusually diverse array of
&
availableforstudy,sufficientfeaturesarepresent shell four, yet Dartnall Dartnall (1972) con-
to make familial placement possible- The flat- cludedthat thegenerawerecloselyrelatedonthe
tened spire, sculpture, coiling pattern, whorl ex- basis of karyotype and reproductive anatomy.
pansion rate and reflected lip relate/*, antiquata Theirresultssegregated theTasmanian/W,c«'v;?
to the Caryodidae, in particular the fiat-coiled in and Caryodcs Albeis, 1850from the mailll
Pedinogyra, However, the absence of an um- Pygmipanda lredale, 1933^ Hcdleyclla and
bilicus is a major departure from this genus and Pedinogyra. Two remaining mainland genera -
indicatesapossibleconnection withthebulimoid Braziercsfa lredale, 1933 and Pandofella -were
caryodidgenera. In Tasmania a few examplesof not included in their study. Kershaw (1988a»b),
the bulimoid form do have an umbilicus and utilising fundamental features ofshell morphol-
occasionally,juveniles have an umbilicus which ogy, substantiallyagreed withthis interpret;i
is losi with growth. This may indicate that an The separation of Anoglypta and Carytxles
umbilicus was more common in ancestral forms from mainland caryodids is supported by
of these species and that the non-umbilical protoconeh features. Thenuclearsculptureofthe
lineage ofPraecaryodesis distinct from these. Tasmaiiian genera is characterised by distinct
Sphaerospira Morch, 867 (+Figuladru spiral cords which are irregularly and weaklycut
J
lredale, 1933, Varahadra Iredale, 1933 andBen- into long segmenls by radial grooves (Kershaw,
tositeslre&dle, 1933) (family Camaenidac) have 1988a,b). Mainlandcaryodidshavemoreregulai
numerous species living in the rainforests of decussate apical sculpture. In Braz.itresta and
southeastern Queensland (hereafter SEQ) and Pandofella this decussate nuclear sculpture is
MEQ (Smith, 1992). These genera also have almostgranular.Thesegenerahavethin,reduced
globose to subglobose shells which often lack an shells and anatomical featuresconsistent withan
umbilicus. However they have almost obsolete evolutionarymovetoslugdom.Thismayindicate
adult sculpture, smooth protoconeh, less c\- a close relationship between them even though
landedbodywhorlandweaklyreflectedlip.Xan Pando/i'lio has a genital diverticulum positioned
I
thomeion Martens, 1861 (family Camaenidac), similarly to that otAnoglypta (Stanisic, unpubl.).
244 MEMOIRSOFTHEQUEENSLAND MUSEUM
FIG. 3.Praecaryodesantiquata sp.nov. A-B, QMF13905, Rundle, MEQ, holotype; CD, QMF17642, Rundle
Formation, MEQ, paratypc(internal cast). Scaleunits inmm.
Solem(1978a)showedthatarrangementofinter- Shell form indicates specialisation to different
nal structures in slug-like taxa wereoften related adaptivezones.Thereducedsculpture, lowwhorl
to spatial readjustments resulting from a reduc- numbers, inflated body whorls and large aper-
tion inthe visceral humpratherthan representing tures ofHedleyella and Pygmipanda are adapta-
primary phylogenetic changes. Brazieresta and tions consistent with life in humid, moist forest.
Pandofella survive in cool rnoist, upland refugia IncontrasttheheavyandflatshellofPedinogyra,
in northern NSW(Brazieresta)and MEQrespec- with small, downwardly directed aperture and
tively and possibly represent highly specialised, higher whorl numbers is adapted for life in a
temperate relicts of a single (monophyletic) dry-forest habitat.
lineage within the family. Conchoiogically the position ofPedinogyra is
Pygmipanda, Hedleyella and Pedinogyra, dis- the most difficult to reconcile within the
play shell features which indicate a long period caryodids. The discoid shape, wide umbilicus,
ofdevelopment by each underdifferent ecologi- heavy shell with thickened lip and strong radial
calregimes.Pygmipandalivesinthemoist,warm apical sculpture are features which stand atodds
tocool, temperateforestsofcentral and southern with thoseoftheothergenera. Nonetheless, Ker-
New South Wales (NSW) and Victoria; Hed- shaw (1988b) emphasisedthatPedinogyra was a
leyella in the warm temperate and warm humid derived bulimoid taxon, citing the transition
subtropical forests of northeastern NSW and series within Hedleyella i.e. imperforate
SEQ; and Pedinogyra in the drier araucarian bulimiform [H. rnaconelti(Reeve, 1853)] to um-
microphyll vineforests and semi-evergreen vine bilicate, globose, sub-auriform [H. falconeri
thickets of northeastern NSW, SEQ and MEQ. (Gray, 1834)] asameansofderivingaflat-coiled
A NEWTERTIARY LANDSNAIL 245
hampton, MEQ. Subfossil Pedinogyra, possibly
Pleistocenein age arealsoknownfromtheCam-
moo and Olsen Caves, north of Rockhampton,
MEQ
(Stanisic, unpubl). These are all con-
sidered to be conspeciflc with an extant species,
Pedinogyra minor (Mousson, 1869) (-nanna
Iredale. 1937), which lives in the region.
Kershaw (1988a) discussed a fossil land snail
from the calcareous beds (?PIeistocene) of the
KentGroupofislands,BassStrait,whichmay be
related to Pedinogyra. Inspection of the shell
(Queen Victoria Museum, Launceston -
QVM:87.G.163) reveals an overall resemblance
to Pedinogyra. However, compared with thai
genus, ithas fewerwhorls,smallerumbilicus atld
vague traces of regularly spaced incised Spiral
lines cutting across fine radial ribs on the last ', i
whorl. The resulting canccllate pattern is not
equivalent to that seen in extantcaryodids but is
more commonly associated with the shells of
carnivorous Rhytididae. Although the peripheral
keel ofthe Tasmunian fossil is notpresent in any
living Australian rhytidid, it is possessed by
several New Zealand species ofthat family.
BlOGEOGRAPHICIMPLICATIONS
The presence ofa number ofderived caryodid
features (pronounced radial sculptureand thick
ened, reflected lip) in P. antiquata indicates that
the adaptive radiation of the Caryodidae in
centraleasternAustraliawasprobablycompleted
FIG. A,Praecaryodes antiquata sp.nov. QMF1345L much earlier i.e. beginning of the Tertiary or
RundleFormation.MEQ,paratype(internalcast).A, possibly Late Cretaceous. This would place
lop view; B, bottom view. Scale unitsinmm. familial origins contemporaneously with the
emergence of the advanced, sigmurcthran
shell from the more generalised bulimoid form. camacnids in northern land masses (Solcrn,
However, one featureofPedinogyrawhichisnot 1978b). Hence rather than being ancestral to the
easily explained in the context of general helicoid sigmurethrans. the caryodids (and
probablyacavoids ofAfrica and Indiaand South
caryodid patterns is the thickened reflected lip,
P. antiquatapossesses features whicharc tran-
sitionalbetweenthebulimoidcaryodidformsand
Pedinogyra - a depressedly glohose bulimoid
shell withsmall apertureand thickened, reflected
lip.Thelattercharacter,rarewithinextantforms,
indicates a possible close relationship with
Pedinogyra. P. antiquata may be the moistforest
ancestor of these modern-day, flat-coiled <:\
caryodids.
CaryodidFossil History llllllllllllllllll
ComparativelyIittleinformationisavailableon
othercaryodid fossils. Hill et aL (1970) recorded FIG. 5.Praecaryodes aniiquaia sp.nov., QMFI34SI,
a species of Pedinogyra from Pleistocene lime- paratype. Sectionedto show internal structure. Scale
stone deposits at Marmor, south of Rock- unitsin mm.
246 MEMOIRSOFTHEQUEENSLANDMUSEUM
America) may representanequivalentandparal- P. minor in the araucarian vine forests of
lel levelofsnail evolution in southern land mas- southeastern Queensland (with P, minor teptac-
ses. ingP antiquatainthedrierareasoftheGladstone
- Rockhampton region).
The colonising wave of northern camacnids
EcologicalImplications
The impetus for the initial radiation of the a{Sdpvhaanetraogsepiorfaosc.cl.a)swioonualldlhaatevrecbleiemnataibcleshtiofttsakteo
caryodids on ihe Australian land mass would wetter conditions and now occupy the
have been enhanced by the significant change in
microhabitats most probably favoured by P. an-
the nature of the flora from one dominated by
tiquata in the past.
conifers to one with a significant angiosperm
componentat theCretaceous/Tertiary boundary CONCLUSIONS
(Kershaw et aL 1991). The switch to a broad-
a
leaved,closed-forestwithapropensityforgreaiet
leaf fall would have favoured 4moisturc-loving* ThediscoveryofPraecaryodesantiquum inthe
snails by providing a new adaptive zone Of leaf sedimentsoftheRundleFormationhasimportant
Idler With moist to humid micro-environments implications for understanding the evolution of
and shelter.Subsequentdistributionofthe famrly the Caryodidae in eastern Australia. Shell fea-
in eastern Australia wouldhave been greatly in tures of this taxon indicate early diversification
ofthe family on the Australian land mass which
fluencedby heclimaticandvegeTationalchanges
thattookplatceinthelatterpartoftheTertiaryand rivals ihe appearance of the camaenids in
Quaternary {Kemp, 1981; Kershaw, 1981), The Laurasia. Climaticchangesince the Miocene not
only led to theextinctionofP. untiquuta butalso
diversification of eastern Australian rainforest
intotemperate,tropicalandsubtropicalfoinishad profoundly affected the evolution of the family
its beginning in the Mid-Boccnc (KershawctaL whichdisplaysapreferenceformoistforests.The
colonisation ofthe eastern Australian rainforests
1991). With the onset of the Miocene aridity by camaenids from the Papua New Guincan
phases 'drifil'1 subtropical rainforest dcvel-
where cdaphic profiles were suitable and in the region most likely usurped the adaptive /one
late Tertiary and Quaternary, mcsorherincle- (micro-habitat) previously held byPraecaryodes,
menis in the north were confined to montane ACKNOWLEDGEMENTS
rcfugia.
wePi,iearn;ttinqduactoaolelrivtedhaninMcEonQdittioodnasy.wVheigcehtawteiroen Thank* are due to ESSO (Australia) and \hc\r
was dominated by NothofagUS forest types jointventurepartnersSouthernPacificPetroleum
(Kemp. 1981), a conclusion supported by and Central Pacific Minerals for making the
palynological evidence from theRundledeposits material available forstudy.LeeCoshell(ESSO)
provided stratigraphic data for the paratype
p(lFoasytsers,om1e979sh)e.llSicghnairfaicctaenrtsly(cPo.mpaanrtaitqiuvaetlay tdhiisn- material, The assistanceofRon Kershaw, Queen
shell, more conspicuous sculpture) which today Victoria Museum, Tasmania in providing com-
arc found in Pedinogyrarotables (Reeve, 1852) parative fossil material is gratefully acknow-
and P, efftma. These species live in marginally ledged. The author is especially indebted to
wetterareas(BorderRanges,ConwayandClarke Andrew Ro/efelds forbringingthematerialtohis
Ranges respectivelyj attheperiphery of the mam attentionand foi providingdataand figuresonthe
stratigraphic elements of the paper. The
massofPedmogyru.
Queensland Museum Photography Section
CreationoftheFitm>ydryconidorwouldhav^
processed photographic illustrations. Finally
considerably changed conditions in the Rnck- thanksarcduetoJenniferMahoncyfortypingthe
hampton - St Lawience region and although
MEQ manuscript.
icfugia in (Sarina - Proserpine) enabled
caryodidstosurvive[Pedinogyraeffossa Iredale, LITERATURECITED
1937; Pandofella whitei (Hedley, 1912)] this
dtroatshteicexltoicnacltciohnanogfeP.inanetnivqiuratoan.meSnutbsperqoubeanbtlycolne-d BEASLEY. AW 1945. Ostracods of some Tcrtiaiv
ditions would have favoured the development BParsoicnese,diinhegistobfeatrhienRgoovnalIhSeoccioertryelOatfiQouneoefnsStlraunt*d
and dispersal ofthe comparatively dry-adapted, 56:95 (24
tfhitihck&er,Prsdmgocootnh,er1s9h3e3l)le-dP.Pead/i/nWo'glyrreadahlaev.ii1(9G3ri7f- BISHOofP.AuMstJr.ali1a9n81l.aTndhesnbaiilosg.coPpg.ra9p2h4y-9a5n4d,eIvnolKcuftliSotn,
. .
A NEWTERTIARYLAND SNAII 247
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