The distribution and patterns of species diversity of land snails in eastern Australia

8 Pages·1994·1.7 MB·

by  John Stanisic

THE DISTRIBUTION AND PATTKRNS OFSPECIES DIVERSITY OF
LAND SNAILS IN EASTERN AUSTRALIA
JOHNSTANISIC
Sianisic.J. 1994 06 30: Thedistribution and patterns ofspeciesdiversity of land snails in
eastern Australia. Memoirs of(he QueenslandMuseum 36(1): 207-214. Brisbane. ISSN
0079-8835.
For coastal and near-coastal eastern Australia (Torres Strait islands to the New South
Wales-Victorian border), itispredictedthat the total number ofspecies ofland snails will
exceed 700(presently less than400are known)when laxonomic revisionsarc completed
Most ofthissnaildiversity isconcentrated inacoastal stripless than200km wide. Species
diversity and endemicity is greatest for rainforest. The climate induced attrition ofmesic
communitiessincetheMiocene(morepronounced inthe PlctStOCCftC),theirsurvival within
refugiaand theirsubsequentradiation anddispersal undermore favourableconditions, are
fundamental ro explaining how these diversity patterns arose Endemicity and species
diversity of land snails in eastern Australia is also high on limestone outcrops. In an
extraordinary situation in the Macleay Valley, northeastern New South Wales, rainforest
andlimestoneSirejuxtaposed, resultingindiversity levels which areexceptionally highon
aworld scale.
Species composition within areas of high snail diversity indicates that these snail com-
munitiesevolvedthroughalongtermaccumulationofLaxaratherthanlocalisedradiations
In a few cases, e.g. isolated dry vine thickets, between-site diversity is increased by
geographicreplacementoftaxa.
Today the distribution of land snails in eastern Australia strongly reflects rainforest
biogeography; furthermorelandsnailcommunitystructurecorrelates with rainforeststruc-
tural types. This suggests that land snails can have strong predictive value in identifying
climatic refugia, ^jj/utmdSruiih, MoIIuum. eastern Australia, biodiversity, communities,
limestone. romfi>rc\i, <ti,\ftihvttotl
John Stumsu, Queensland Museum. HO Box3300, South Brisbane. Queensland. 4101,
Australia;21 November 1993.
The study of land snail diversity in eastern For the past thirteen years the Queensland
Australia is in its infancy. Faunal checklists and Museum Malacology Section has beenengaged
species' descriptions have not been comple- inasystematic collection programmeextending
mentedby 'follow-up* surveyworkandrevision- from the Torres Strait islands to theNew South
arystudies.TheworksofCox (1868)and Iredale Wales/Victorian border in an effort to redress
(1937a, 1937b, 1938) still form the basis ofler- these shortcomings. More than 1000sites(asite
restrial malacological knowledge for this region, is an area <lkmfc in size) (Fig. 1) have been
ConlemporarycontributionsofSmith&Kershaw sampled by hand-collecting live and dead
0979, 1981) focused on Victorian and Tas- specimens and sorting retrieved leal litter (land
manian species and more recently Smith ( 1992) snails usually possess a hard shell, which
produced a revised checklist of all Australian remains in the litter after death and allows for
land snails. Attempts at 'modern' biogeographic post-mortem sampling). Resultsindicatethatthe
syntheses arc limited to McMichael & Iredale diversity of the east-coast land snail fauna has
(1959). Solem (1959), Bishop (1981) and Smith been grossly underestimated. While a great deal
(1984). With the exception of some rainforest oftaxonomicworkneedstobecompleted before
&
surveys of the mid-1970's (e.g. Broadbent acomprehensivebiogeographicsynthesiscanbe
Clark, 1976) and a survey of the New England presented, preliminary findings make it possible
district, New South Wales (Simpson & Stanisic, to promote these often-neglected animals in the
1986), there has been a lack of comprehensive biodiversity debate. This paper examines some
Heldworkdirectedatrecordinganddocumenting aspects of land snail diversity in eastern
the east-coast land snail faunj. Australia which have come to light during the
208 MEMOIRS OFTHEQUEENSLAND MUSEUM
course ofthe study.
The major physiographic features ofthe study
areaare theGreat Dividing Rangeand a series of
coastal ranges which rise from a narrow coastal
plain to provide barriers loonshore, rain-bearing
winds. Consequently, in addition to drier
sclerophyll vegetation, these mountains support
rainforestthatvaries in structure andextentfrom
north to south (Webb & Tracey, 1981) and from
sea level to mountain summit. In more westerly
areas, rainforest in the form ofdry vine thickets
occurs sporadically on volcanic soils and rock
outcrops, in particular limestone. Field work has
beenfocusedoncensusingthe snailcommunities
within these habitats, which occur mainly in a
narrow coastal strip less than 200km wide.
The following abbreviations have been used in
the text: NSW, New South Wales; NEQ, north-
eastern Queensland; MEQ, mideastern
Queensland; SEQ, southeastern Queensland.
LAND SNAILS AND RAINFOREST
The.association betweenland snailsandmoist,
closedforestsisanancientone. Landsnailsmake
their first appearance in the fossil record in the
Upper Carboniferous coal beds of Europe and
North America and ithasbeensuggested (Solcm
& Yochelson, 1979) their early radiation was
closely tiedtothe appearanceandproliferationof
angiosperms. The basis for the association is
ecological. Those factors which favour rain-
forests (high nutrient soils, moisture) are also
those which favour land snails. Rainforests pro-
vide the additional benefit of shelter. In eastern
Australia this bond has been more strongly rein-
forced by climatic and geologic events which
shaped present-day, east-coastphysiography.
Rainforest was once more widespread in
Australia than today (Martin, 1981). With the
onsetofarid episodes in the Miocene (Galloway FIG 1.Distribution of land snail collecting sites in
& Kemp, 1981), rainforest either disappeared or eastern Australia, Queensland Museum 1980-1993
was restricted in distribution in many areas (Pig. (Ca, Cairns; Ma, Mackay; Ro. Rockhampton; Br,
Bn^hane: Sy, Sydney
2). In the east, uplift of the Great Divide and ):
volcanicactivitycombinedtoprovide favourable bodied land snails, would have been placed
moisture-soil conditions which ameliorated the underconsiderable stress. In some cases extinc-
effects ofcontinental drying and allowed mesic tions would have occurred.
communities to persist. Subsequently rainforest The fact that most land snails in eastern
has fluctuated greatly in extent (Kershaw, 1980, Australia (over 90%) now occur in rainforest
1981) and the extreme drying events of the indicates just how critical the persistence of
Quaternary would have seen rainforest retreat to moistrefugiawastotheirsurvival in this region.
rcfugia such as moist uplands and gaily heads, Within the rainforest vegetation mosaic, snail
emergingonlyinwetterperiods(Webb&Tracey, diversity and community composition differ
1981). The animal communities within them, in markedly with latitude and from site to site. The
particularthe strongly moisture-dependent, soft- regions which have the greatest number of
DIVERSITY PATTERNSOFLANDSNAILS 209
FAMILY Named Estimated the rock provides a protective niche from
new wildfires so that in many instances snail com-
species^
species* munities, quite distinct from those in the sur-
1Ivdmcenidac 4 5 rounding forest, have developed and been
maintained. Landsnailswhich inhabit thesespe-
Helicinidae 5 3
cialrefugiaalsobenefitfrom an ample supply of
Cvclophoridae 2 2
calcium. Endcmicity and specialisation in these
Pupinidae 19 10 limestone snail communities indicate long-term
Diplommatinidae 4 3 isolation (Stanisic, 1990). In anexceptionalcase
Aehatinellidae 3 1) in the Macleay Valley, NENSW, subtropi-
cal/warm temperate rainforestand limestoneare
Pupillidae 20 4
juxtaposed resulting in extraordinarilyhigh land
Enidae
1 ] snail diversity.
Mcpaspiridac
1 !
Succineidae 3 1 FAUNAL COMPOSITION
Subulinidac
1
Rhytididae 27 8
Caryodidae 9 There are 22 families represented in the east-
coast native land snail fauna composing more
Hclicodiscidac
1 than 670 species of which about half are un-
Punctidac J3 7 described (Table I). This compares with es-
Charopidae 74 200 timates ofatotal Australian faunaof504spa
A1horacophondac 1 3 in 25 families (Smith, 1992) and illustrates the
enormous diversity (mostly undescribed) ofthe
Cystopeltidac 4 2
east-coast fauna. It is probable that final species
Helicanonidac 46 40 numbers for this region will exceed 700. The
Camaenidae 105 35 main contributors to this biodiversity are the
Corillidac 1 Charopidae, Camaenidae and Helicarionidae.
The Helicarionidae and Charopidae have their
Ralhouisiidae 2
greatest expression in eastern Australia but the
TOTAL 347 325 Camaenidae are more diverse in other areas of
Australia (see Solem. 1992, for overview). The
TABLE I. Faunal composition ol' eastern Australian Australian opcrculatc land snails
land snail diversity ("^based on various sources; (Hydrocenidae. Heiieinidae, Cyclophondae,
*based on undescribed species identified in the col- Pupinidae and Diplommatinidae)arealso large-
lectionsofthe Queensland Museum). ly confined to the cast-coast rainforests. The
stable mnisnire regimeand volcanically derived
species are those with dissected topography
acidic sinIs have provided an ideal environment
(Sleep gorges and high mountains) that support
forslugevolutio—n andthree familiesofslug taxa
diverse rainforest vegetation communities (e.g.
are represented the Ralhouisiidae, Cystopel-
Wet Tropics, Border Ranges). Moisture stability tidae and Athoracophoridae. The first is an
mid the availability of diverse niches have
obligate rainforest group while the latter two
provided an ideal environment for the evolution have representatives m transitional wcl
and persistence ofcomplex snail communities.
sclerophyll forestaswellasclosedforests.Semi-
slugs, belonging to the family Helicarionidae,
LIMESTONE
display an even greaterdiversity here with over
30 species present, some restricted to mountain
Limestoneoutcrops (Fig. 2). in particularlarge tops. The great majority of these are rainforest
tower karsts such as those present in the Chil- dwellers with only a few species occurring in
lagoe-Palmerville region, NEQ, north of Rock- drier sclerophyll forests.
hampton, MEQ, and Jenolan, NSW, are Dry-adaptedgroups suchasthe Pupillidaeand
significant secondary habitats for land snails. Punctidac have several representatives in the
Theysupportremnantsofwet-adaptedvegetation warm to hot humid fOTCSK but are more diverse
in otherwise dry, sclerophy ll-dominatcd fn the drier vine thickets and forests, and
countryside. Moisture is trapped in crevices and c Icropfiyll forest.
210 MEMOIRS OPTHE QUEENSLAND MUSEUM
SITEDIVERSITY
Landsnaildiversity levelsatindividualcollect
ing sites have been found to be highly variable
and range from about 5 species per site in dry
sclcrophyll forests to over 40 species per site in
some rainforest patches (Fig. 3). Greatest diver- dryo.rrldor
i
sity has been recorded in the subtropical rain-
forests of southeastern Queensland. Binna Burra
(BorderRanges) and Buoloumbu Creek (Conon-
daleRanges)haveyieldedinexcessof40species.
From thissubtropical zenith diversitydiminishes \(^ *A drycorridor
both tothe north (tropical forest) and to the south
(temperate forests) In most cases increased dis-
tance from the coast correlates with a marked
drop in diversity so that subcoastal rainforesl
patches (= dry vine thickets) average about 10
srcpfeucgiieas.suEcxhcaesptthioosnesatoCcacrunrarivnonisGoolragteed, MmEoiQs,t CarGnoragrevoni*5\\<f*1*I _
and Ml Kaputar (Nandcwar Range), NSW, and C? limestone
onlimestoneoutcrops. HenceatChillagoe,NEQ,
more than 25 species have been recorded on ^* rainlbrc.si ,12.
limestone in otherwise snail depauperate .«• appro*, subfaunalboundary
NSW
countryside. In southern limestone out-
crops located in areas of sclcrophyll woodland
(e.g. Abcrcrombie, Yarrangobilly, Wombeyan. Ml Kaputar24| r, \| .Si
Jenolan) also record above average site diversity
levels indicating the probability of highly
localised endemism In NENSW average rain- d
20A-
forest site diversity levels are generally 10-20
species but in the Macleay River valley, where
limestone occurs within rainforest, species num-
bers at some sites (Yessabah, 36; Mt Sebastopol,
26) are much higher owing to the presence of
limestone endemics (Stanisic, 1990). There are I
few areas ofthe world where site diversity levels
exceed 30 species (Solem, I9S4). In this context
eastern Australia is an area of exceptional land
snail diversity.
MOSAIC DIVERSITY Fid 2.Distribution of'majorcoastal rainforest blocks
and areas oflimestone outcrop in eastern Australia.
Doited linessignify possibleboundariesfordivision
EsDcisasrepcmteednttopo(gOrialpiheyr.ass1oc9i8a2t)edwaintdhthseeGvreeraet oSlcrluabn;d2s,naIirlosniRnatongseub.f3a,unPaullmuenrit\si(llAe-,J4,).C1h,iLlolcakgeore;bi5e,
palaeoclimatic regimes have combined to WetTropica; 6, Ureenvale;7,Broken River; 8,Con-
produce a diverse pattern of vegetation com- way Range. 9, Eungclla; 10, The Caves; II, BuK
munities ineastern Australiathatsupportequally burin; 12 Gympie; 13, BorderRanges; l4,Ashford,
diverselandsnail assemblages. The resultis rela- 15, Dorrigo; 16. Manilla- Tarnworth; 17, Carrai-
tively high diversity within comparatively small Wernkimbe; 18, Maeleav Valley; 19. Barrington
areas even though diversity at individual sites is CTooWpsf:a-2Y(ia.SHSi;ll2B3o.d-MCoalpotnagi.nsA.RaCt;ap2C1,Ylolrtka;warBr,a;W2e2i,
not high. Tropics; C, hinasleigh Uplands; D, mideastcrn
Mt Bcllendcn Ker is the second highest moun- Queensland; E. Brigalow Lands; P, southeastern
tain (alt. = 1560m) in theWetTropics, NEQ. The Queensland; G, Border Ranizes-norlhcastcrn NSW,
eastern face is eovered in rainforest which is K.NcwEnglandTablelands;[.SouthernTablelands;
alhtndmally stratified into a numberof structural J,south eoastal New South Wales.
DIVERSITY PATTERNSOFLANDSNAILS 211
tionismostmarkedintheWetTropicsbutisalso
presentatothereast-coast localities.
The usual patterns of land snail distribution,
which strongly reflect the rainforest/woodland
vegetation mosaic are complicated by the
presence of limestone (Fig. 2). A large number
of outcrops occur along the Great Divide in
NSWbutthoseintheMacleayValley,NENSW,
are perhaps the most significant. They are
situatedinthemidstoftemperateandsubtropical
rainforest and even the most easterly outlier
(Yessabah) is vegetated by rainforest in spite of
being surrounded by much drier countryside
(Floyd, 1983). Fifty sites (Fig. 5) were sampled
inaareabounded by theHastings River(south),
Nambucca River (north) and the Great Divide
(west). Rainforest sites (those without lime-
stone) yielded comparatively low numbers of
species (mean 5.64 species/site) but limestone
sites werefarricher(31.00species/site). In con-
trast eucalypt forest sites were relatively snail
poor(mean3.00species/site).Thehighdiversity
foundon the limestonesresultsfrom acombina-
tionofwidespread species found insurrounding
rainforest and eucalypt woodland, and a sig-
nificant number of limestone endemics (see
Stanisic, 1990, for some examples). Some
species are confined to individual outcrops.
Total diversity was approximately 85 speciesin
thesampledareaandwascomposedofanumber
of quite distinctive assemblages which have
developed underdifferent micro-environments.
In the vine thickets west of Townsville to
Sarina, NEQ, there is evidence ofgeographical
replacementofspeciesbetween widely separate
rainforest patches. Individual patches have low
species numbers (5-10 species) but diversity
overthe totality ofpatches exceeds this because
ofallopatricspeciation. Solem (1984)reporteda
similar phenomenon in northwestern Australia.
REGIONALDIVERSITY
FIG. 3.Comparison of land snail diversity levels at
selected sites in eastern Australia (Ca. Cairns; Ma,
Mackay; Ro, Rockhampton; Br, Brisbane; Sy, Syd- Iredale (1937a) first applied the concept of
regional diversity to Australian land snails. Al-
ney).
though this scheme has received some support
types varying from complex mesophyll vine (McMichael& Iredale, 1959;Smith, 1984)there
forest in the lowlands to simple microphyll vine has been criticism of its predictive value
fern forest and thicket near the summit (Fig. 4). (Bishop, 1981). Horton (1973) suggested that
Sampling at selected altitudes produced 45 any zoogeographtc subdivision of Australia
speciesoverall sites.However,few species were should consider isolating barriers as well as the
presentatall altitudes (Fig. 3). Greatestdisparity climate and vegetation ofan area. The proposed
occurred between the base (100m) and the sum- methodology (based on bird distribution) has
mit(1560m)sites which hadonly 5 of37 species particular relevance to land snails in eastern
in common. This pattern ofaltitudinal stratifiea- Australia where theirevolution has been driven
:i: MEMOIRSOFTHEQUEENSLANDMUSEUM
Sllca lOOm Sootn 10S4ia IMOin
ilptlnhctT'
100m "JO 1
1 SOOai ffl 12/26 Spcclnf>Wr«l
!OS«io lt» 7/31 &/17
* ICm| ryil iBaOm 33 6/3J io/au
FIG.4.AltitudinalstraiificaiionoflandsnailcommunitiesonMtBellendenKer, NEQ(12/26meansthatalotal
of26species were collected at 100m/500msitesand 12 werecommon toboth).
by climate-based changes to mesic vegetation colonising northern element. The southern ele-
communitiessince theMiocene. Afulloutlineof ment has more relicts in the north (mostnotably
a scheme ofland snail subfaunal units in eastern among the Charopidae) than vice versa.
Australia will be presented elsewhere but the Charopids are numerically dominant in many
approximateboundariesoftheunitsarcshownin placesespecially inareasofhighdiversitywhere
Figure 2. Theproposed scheme divides the east- site numbers can range 8-12 species. Sites with
ern Australian land snails into a number of large numbersofcharopids(whetherinthenorth
smaller subregionaj units Csubfaunas*) defined orsouth)orwithany narrowendemic landsnails
by acoincidenceofspecies ranges and separated are significant (Stanisic, 1990) because they in-
byspeciesrangeendpoints(fauna!breaks).These dicate long-term moisture stability.
faunal breaks express past and present environ-
mental discontinuities and may coincide with DISTRIBUTION PATTERNS
present climatic barriers (dry corridors) or, less
obviously, reflect amore complex history ofen-
vironmentalsifting.Initialinvestigationsindicate Solem (1984) predicted thaithe medianlinear
that the proposed land snail subfaunas in range forall land snail species wouldbeconsid-
Queenslandshowconsiderableconcordancewith erably less than 100km (and probably less than
the natural biogeographic regions outlined by 50km). Stanisic(1990) snowed that a numberof
Stanton& Morgan(1977)which weredefinedby eastern Australian Charopidae had much more
climate, vegetation and landform. extensive ranges (150-200km). Most cast-coast
rainforest land snails show considerablebreadth
COMMUNITY STRUCTURE of distribution within major rainforest tracts
(Wet Tropics; mide3stern Queensland;
araucarian vine forests of southeastern
The species composition of land snail com- Queensland; Border Ranges etc.). Upland
munities present at any site usually consists of refugiaand limestoneoutcrops havethegreatest
narrow rangeendemicsandspecieswhich have a number of narrow range endemics. In driervine
more widespread distribution. The proportion of thicketsspeciescan rangeoverconsiderabledis-
narrow range endemics is greatest in refugia tancebutoccursporadicallyin isolatedandscat-
(moist uplands, limestone outcrops) whereas tered thickets. Species which live in eucalypt
widespread species tend to dominate drier vine woodland tend to have the widest ranges. These
thicketsandeucalypt woodland. These land snail features highlight the importance of refugia for
communitiesappeartobearesultofanaccumula- land snail survival, the significance of the rain-
tionofspeciesratherthan localisedradiations. In forest ecosystem in providing long-term mois-
only a few cases can minor local radiations be ture stability and the broad environmental
identified,however,the widerpicture isoneofan adaptability ofdry-adapted forms.
ancient southern element complemented by a While it is possible tocomment informatively
DIVERSITY PATTERNSOFLANDSNAILS 111
on local species richness, discussion of species to the limited activity periods of land snails in
abundance(numbersofindividuals)remains lar- drier environments. The effects of competitive
gelyqualitativeandspeculative. Noformalstudy interaction between and among species is un-
ofland snail abundance (in time and space) has known as are the effects ofdensities on popula-
yet been attempted in eastern Australia. tion. There is a desperate need for basic
Limestonesitesand the araucarian vineforests ecological studies of eastern Australian land
of southeastern Queensland have the greatest snails with particularemphasis on the aspect of
numbers of species living sympatrically. How- abundance.
ever, because land snails have fairly specific
mierohabitairequirementsthelocalabundanceof CONSERVATION
a particular species will vary greatly depending
o(mnictrhoehadbeigtraetediovferesintvyi)r.onHmeennctealinheMteEroQgenleairtgye apApesarintdoicbaetoraspootfenctliiamlaltyicsirgenfiufgiicaanltagndrosunpaiolsf
camaenids are dominant overall but reach organisms in the biodiversity/conservation
greatest numbers in rocky talus slopes. On lime- debate. In spite of the large numbers of lift-
stone outcrops the density of individuals is en- described species, land snails represent a
hancedby thepresenceofshelteredsoutheasterly manageable taxonomic unit for use in land
aspects along driplines. Many of the enviion- managementdecisions.Todaythcirdistrihutions
ments withhighsnail numbersareseasonallydry reflect rainforest biogcography and in many in-
and fluctuations in local abundance may occur stances land snail community composition cor-
hutthishasnotyetbeennotquantified. IntheWet relates strongly with rainforest structural type>.
Tropicslargecamaenidsdonotappeariohavethe This suggests that land snails can have strong
sMaEmQe local abundance as those which Occur in predictive value in identifying potential reserva-
so thai locating large numbers of 10- tion areas,
dividualsisdifficult.Similarsituationshavebeen
NENSW
encountered in the wetterforestsof and ACKNOWLEDGEMENTS
SEQ. Reasons for this are not obvious. Solem
(1984) suggested that there might he. advantages
Dataon theland snailsofthe Macleay Valley,
NENSW, were collected as part of a larger
project on the limestone Charopidae of NSW
funded by ABRS. This financial assistance is
gratefully acknowledged. These data were col-
latedandanalysedbyCraigEddie. Fieldworkto
Mt Bellenden Ker. NEQ, was funded by the
American-based Earthwateh organisation.
Thanks are also to Jennifer Mahoney for typing
the manuscript.
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