Memoirs of the Museum of Vicloria 56(2):435-439 ( 1997)
DIVERSITY IN CENTRAL AUSTRALIAN LAND SNAILS (GASTROPODA: PULMONA TA)
BRONWEN SCOTT
Department of Zoology, James Cook University of North Queensland. Townsville, Qld 4811, Auslralia
Abstract
Scott, B., I 997. Diversity in central Australian land snails (Gastropoda: Pulmonata).
Ml'moin of //1e Museum of Victoria 56(2): 435-439.
Much of the_ land snail diversity in central Australia is due 10 camaenids (70 of 83 spp.).
�any .ca1;1aen1d species have restricted distributional ranges so that. although land snail d1vers1_ty In Central Australia is relatively high, local diversity is generally low. The fauna of the Knchauff Range in the Central Ranges is an exception to this general rule as the Finke Gorge National Park contains at least 25 species of snails, 30% of the central Australian fauna. Similar areas in the Central Ranges. such as the George Gill Range and eastern MacDonnell Ranges have a significantly lower diversity of land snails. It is postulated that high levels of diversity and endemicity in central Australian camaenids are the product of
vicariance events related to Tertiary and Quaternary changes in climate.
Introduction
Over 80 species ofland snails from eight families have been recorded from central Australia (Solem, I 989, 1991, 1993; Scott, in prep.). The land snail fauna is dominated by the families Camaenidae and Pupillidae s./., both of which are widespread across mainland Australia.
Other families are represented in this area by no more than two species each. It has been suggested that the relatively small number of families in the central Australian fauna (com
pared with the greater complement in coastal areas) is the result of dispersal from coastal centres of origin to the interior, combined with the effect of filtering due to fluctuating climatic conditions (Solem, 1993). Problems arise, how
ever, with dispersal hypotheses as explanations of the origin and distribution of organisms as they are ad hoc narratives used to describe specific cases and, as such, are generally untest
able.
Hypotheses concerning vicariance biogeogra
phy provide alternative interpretations of distri
butions. In that model of historical biogeogra
phy, it is assumed that taxa and the areas they occupy have evolved together (Nelson and Plat
nick, 1981 ). Reconstruction of a phylogeny allows the reconstruction of the history of a region. Biogeographic hypotheses erected using data from one group of organisms can be tested with those from another. Land snails are excel
lent subjects for such biogeographic studies as they are 'vicariant conformers' (Springer, 1981:
230), responding rapidly to environmental dis
ruptions (Gould, 1969).
435
Origins and diversification of the Central Australian land snail fauna
It has been suggested that many families rep
resented in central Australia were Tertiarv invaders from Asia and that the apparen·t reduction in diversity from north to south and from east to west is a result of dispersal from tropical centres of origin (Solem, 1959, 1992, 1993). The most diverse of those families, the Camaenidae. is postulated to have entered Aus
tralia in several waves (Solem, 19 59, I 992, I 99 3;
Bishop, 1981 ), the products of the earliest wave having the most extensive distribution and those of the most recent restricted to the eastern rain
forests (Solem, 1959). This interpretation is based on the assumption ofmonophyly of Amer
ican and Australasian camaenids but this assumption is probably incorrect (Scott, 1996).
It is possible that the Australasian component of this family is closely related to the Asian Brady
baenidae (Scott, 1996) and is of Gondwanan origin (Scott, in press).
Central Australian camaenids show a high degree of regional endemisrn but two genera in the subfamily Sinumeloninae, Sinume/on and Pleuroxia, have ranges that extend beyond the Centre. Species of these genera occur in the Centre and in southern and parts of western Aus
tralia (Solem, 1992, 1993). This has been interpreted as the result of primary differen
tiation of ancestral taxa in the Centre, followed by southerly and westerly migrations to give rise to local radiations (Solem. I 992, 1993). Present distributions may, however, also be explained as the result of tectonic or climate-induced vicar-
https://doi.org/10.24199/j.mmv.1997.56.35 28 February 1997
ianceeventswhichcaused populations ofances- traltaxatobecomeisolatedandthus to giverise tomoderntaxa. Preliminarystudiesonthephy- togeny and cladistic biogeography ofthe Sinu- meloninae indicate that fauna of southern and westerncoastal areaswasseparated fromthatof central Australia early in the evolutionary his- tory ofthe group (Scott, in prep.). Within the central Australian region, some montaneareas, such as the Central Ranges, may have become
isolated from adjacent areas not long after the Centre-SW split.Vicariance eventscausingdis- ruptions of these magnitudes may have been associated with cyclical climate change in the Caenozoic, when periods of relative dryness associated with global cooling alternated with
warm andhumid pluvialstages. Alternation of arid grasslandsandextensivewater bodiesinthe plains and valleys may have presented signifi- cant barrierstoformerly widespreadsnail taxa.
Whichofthetwoextremesmayhave playedthe
moresignificant rolecannot be determined, but they may haveacted synergistically.
Congenersofthe central Australianbulimulid Bothriembryon spencen(Tate, 1894) are found insouthern and westernpartsofthe continent.
as well as the south-eastern coast ofTasmania (Smith, 1992). There is generally little spatial overlap betweentheCamaenidaeandBulimuli- daeas camaenids do notoccurinTasmaniaor the extreme south-west of the mainland and Bothriembryonisnotfound inthenorthoreast.
However, although the distributions of Bothriembryon and the two camaenid genera, Sinumelon and Pleuwxia, do not correspond entirely, broad geographical congruence between the three genera is such that future phylogenetic and biogeographic analysisofone genus will provide a hypothesis available for testingwith the othertwo genera.
Patterns ofdiversity in the Central Ranges landsnail fauna
AlmosthalfoftheCentralAustralian speciesof land snail havebeen recorded fromthe Central Ranges (MacDonnell, Krichauff and George
Gill Ranges)(Smith, 1992; Solem, 1993; Scott, in prep.), which cover 12% ofthe area ofthis region, while Finke Gorge, in the Krichauff Range, alonecontainsalmosta thirdofthe cen- tralAustralian land snail species (Table 1).The
snailfauna oftheCentralRangesiswell-known as collecting sites are readily accessible, but direct comparison between the Central Ranges andadjacent areasmaypresentproblemsassites
furtherawayfromAliceSpringsmayhavebeen
lessthoroughly explored.
Camaenid species comprise 66% ofthe land snail fauna ofthe Central Ranges and demon-
strate a high level ofendemism compared with species in other families (Table 2). The non- camaenidspeciesofthatfaunaarelargelywide- spread, shared with otherpartsofCentral Aus- tralia and, in several cases, also with coastal areas.
The pattern of diversity demonstrated by camaenidsinthe Central Rangescan beseen in
otherareassuchasthe Kimberlev andadjacent ranges (Solem. 1979, 1981a, l'981b, 1984b, 1985, 1988a, 1989). Many of these endemic species have narrow ranges which are usually allopatric to congeners (Solem, 1988b). This appears to be a common pattern among land snails(Solem, 1984c)anditisrarefortwoshort- rangeCentralRanges endemicstooccursympa-
trically,althoughthey are frequentlyfound with other more widely-distributed species. Geo- graphicrangesdo notappeartoberestricted by competitionasthey areneverparapatric and,as narrow-range endemics do not come into con- tactwith eachother,therecan benoevidence of repressive interactions.
Pleuroxia adcockiana (Bednall, 1894) is the sole representative ofthe genus in the Central Ranges andtheedges ofitsdistributioncoincide with thoseofthe Central Rangesbiogeographi- cal area. The four speciesofSinumelon in this area are moderate-range endemics and divide the Central Ranges into four units: farwestern MacDonnell Ranges. Krichauff and James Ranges(S.exposition Iredale. 1937).nearwest- ern MacDonnell Range and Alice Springs (S.
bednalli Ponsonby, 1904), eastern MacDonnell Range (S. dulciensis Solem, 1993) and George
Gill Range (S. gillemis Solem. 1993) (Solem, 1993).
The genera Gramdomelon and Semotrachia show very high degrees ofendemism. Threeof five species ofGramdomelon are found in the CentralRanges and twoofthose areknownonly fromsinglelocalities, at opposite ends of Finke Gorge (Solem, 1993; Scott, in prep.). Semotra- chiaisan extremelydiverse genus,with 18ofits
28speciesknownonlyfromthe CentralRanges (Solem, 1992; Scott, in prep.). Many (16) of these species are narrow-range endemics, con- fined to singlegorgesinthe easternand western MacDonnell Ranges, or from rock walls in the Krichauff, James and George Gill Ranges.
Combinationofpatternsofdistribution from these species suggeststhatalthough the Central
DIVERSITY IN CENTRAL AUSTRALIAN LAND SNAILS 437
Table 1. Land snail diversity in Central Ranges (data compiled from Solem, 1989, 1991, 1993;
Scott, in prep.) (atotal numberofspecies recorded from theCentral Ranges = 39; btotal
numberofspecies recorded from central Australia = 83).
No. ofspecies3 % ofspecies recorded from central Australia11
"Western MacDonnell Range Eastern MacDonnell Range Krichauffand James Ranges Finke Gorge
George Gill Range
23 14 27 25 9
27.7 16.8 32.5 30.1 10.8
Table 2. Land snail diversity at selected sites in the Central Ranges (sites areapproximately equivalent in size) (datacompiled from Solem, 1984a, 1989, 1991, 1993; Scott, in prep.)
endemic camaenids
non-endemic camaenids
endemic others
non-endemic others
KrichauffRange Finke Gorge
W MacDonnell Range Glen Helen
Serpentine Gorge
—
Ellery Big Hole Simpson Gap—
Fenn GapAlice Springs
Eastern MacDonnell Range George Gill Range
Rangesrepresent acoherent biogeographicunit, this unit can be divided into seven areas, each with its own evolutionary history. The Alice Springs region, eastern MacDonnell Ranges, Krichauff and James Ranges, and the George GillRangeeachrepresent singleareas,whilethe western MacDonnell Range is a composite of
three sub-areas.
Evolutionin the Central Ranges TheCentralRangesare subparallelstrike ridges,
running approximately east-west, and arecom- posed principally of sandstone and quartzite with some dolomite and shale(Mabbutt, 1967;
Thompson, 1991).The MacDonnellRangesare dividedinto easternandwesterncomponentsby theTodd River andthe western range is separ- ated from the Krichauff and James Ranges immediately to its south by Missionary Plain.
TheKrichauffandJamesRangesarecontinuous with the eastern MacDonnell Rangeasonearm ofageosyncline; the western MacDonnell Range forms the other arm (Thompson, 1981).
14
1 3 12
1 3 5
2 3 6
1 3 6
4 4 6
2 1 6
Extreme climate changes ofthe Tertiary are registeredin thegeomorphology ofthe Central Ranges. The ranges rise abruptly from plains and, in many areas, arecapped with duricrust, indicating the pre-erosional valley floorsofthe Tertiary. Exposureofthe mountain rangesand the subsequent cutting of gorges through the rangeshas probably occurred only overthelast 20 million years(Twidale, 1994). The presence ofnumerouswatercourses,which cut across the rangesratherthan runparalleltothem,indicate the importanceof waterin producingthisland- scape. Several significant waterways are found
in this area, including the Finke River and its
tributariesinthewestern MacDonnelland Kri- chauff Ranges,andtheRossRiverinthe eastern
MacDonnell Range.
Coincidenceofwaterwayswith the limits of biogeographic areas suggests that rivers may have been important in the dissection and iso- lationofancestral populations.Water-cut gorges anddepositional plainsprovide evidence ofsig- nificant pluvial phases, during which snails
might have been restricted to montane islands.
438 B. SCOTT
Increasing aridity during glacial phases may havecontributed to theisolationof populations and subsequent speciation (e.g. Haffer, 1982;
Winter, 1988; Bush and Colinvaux, 1990) but thenumerousshort-rangeendemiclandsnails in otherwise contiguous mountain ranges suggest that it is the presence ofwater, ratherthan its
absence, which is the most important factor in speciation here.
Implications for land snailconservation Short-range endemic invertebrates arc often neglected when areas are selected for conser- vation. Economic imperatives ensure that sur- veys focus on readily-observed and readily- identified taxa,suchasvascularplantsand ver- tebrates with little attention directed towards otherorganisms. As the rangeofa snail species
may be less than 0.5 km2 (Solem, 1988b), the inclusion of narrow-range invertebrate taxa in protected areas is as likely to be the result of goodluckasitisofgoodplanning.Awarenessof the possible existence of such taxa combined with an understanding ofthe history ofan area will greatly increase the chances ofshort-range endemics being located and subsequently pro- tected from extinction.
Acknowledgments
ThankstotheConservation Commissionofthe NorthernTerritoryforpermissiontoworkinthe
MacDonnellRanges, FinkeGorge and Watarrka National Parks,andforassistancewithvehicles.
Thanks especially to Dennis Matthews and Darren Schunke ofFinkeGorge National Park forextraordinarylocal knowledge andindefati- gableenthusiasmandhumour,toSuzanneBoyd and Alan Yen ofthe Museum of Victoria for allowingmetojoin themalacologicalunitofthe expedition, and to James True ofJames Cook Universityforinvaluable assistanceon the first trip. Funding to attend this conference was provided by the Zoology Department, James Cook University.
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