Table Of ContentMemoirs ofthe Museum ofVictoria 56(2):591-596 (1997)
CONSERVATION OF CAVE FAUNA: MORE THAN JUST BATS
David P. Slaney and Philip Weinstein
Department ofZoology, James Cook University, Townsvillc. Old 481 1, Australia
Abstract
Slaney, D.P. and Weinstein, P., 1997. Conservation ofcave fauna: more than just bats.
Memoirs ofthe Museum ofVictoria 56(2): 591-596.
Theexpansionofbiospeleologyasascienceandcavingasa recreationalactivityhaveled
toacorrespondingincrease in pressureon cavesand theirecosystems. Thethreats include
direct human disturbance from visitation, and indirect modifications tocave habitats and
the surrounding environment. Despite the high profile ofconservation issues such as the
threat topopulationsofghost bats, littlework hasbeencarriedout in Australiatoassessthe
impactofhumansoncaveecology.Theaim ofthisstudywastobrieflyreviewtheeffectsof
disturbance on populations of cave invertebrates, and to investigate the impact that
sampling may have upon them.
We recorded the weekly total ofindividualsofeach macroiitvertebrate speciescollected
over one month ofcontinuous trapping with a variety ofmethods in Rope LadderCave,
Fanning River Caves. North Queensland. The results indicated that numbers ofpseudos-
corpions, cockroaches, rhizophagid beetles, and pselaphid beetles declined significantly
overthesamplingperiod. Samplingofcave invertebratesforscientific purposes maythere-
fore have an impact on the population dynamics ofcave species, and may consequently
affecttheecologyofthecaveitself.Werecommendthatthegeneralecologyoforganismsina
cave be investigated beforeany intensive sampling iscarried out, and that where possible,
sampling with replacement should be used.
Introduction pods from Bayliss lava tubeat Undara, seven of
which were newspecies(two represent newgen-
Cave environments are unique habitats that are era) and 24 ofwhich were troglobites (Howarth
sufferingfrom increasingpressures placed upon and Stone, 1990). We have discovered further
them. Caves contain a wide range of invert- new species, including a pseudoscorpion, and
ebrate taxa, such as crickets, cockroaches, milli- several sibling species of cave cockroaches
pedes, amphipods, isopods, and arachnids. (Weinstein and Slaney, 1995). Caves are not
Many of these organisms show morphological onlyimportantwith respecttodocumentingand
modifications (troglomorphies) that are not preserving biodiversity, but are important for
found in corresponding surface dwelling studying adaptation, speciation, and species
(epigean) species. Troglomorphies include interactions. They provide us with natural lab-
reduction or loss of eyes, wings, and bodily oratories in which we can frame and test evol-
pigmentation, and attenuation of appendages
utionary hypotheses.
(Barr, 1968; Culver, 1982, Kane and
Richardson, 1985). Many of these species are Caveorganismsare particularly vulnerable to
relicts, having few or no closely related epigean disturbanceasthey livewithindiscretehabitats,
species. Althougha largeamountofresearch has with isolated island like distributions. Species
been carried out on Australia's temperate caves are particularly vulnerable when endemic
(Hamilton-Smith, 1967, 1987; Richards, 1971; speciesareconfined tooneortwocaveswithin a
Eberhard, 1993; Eberhard el al., 1991), few karst region. For example, near Chillagoe, Qld,
studies have looked at our tropical caves. Not foursmall limestonetowersoccurwithina2km-
until the early 1980s did entomological studies areawithin which several endemiccaveadapted
delvedeeperinto North Queensland caves. This arthropods are found, with some being restric-
work led to the discovery ofa rich and diverse tedtosingletowers(Hochand Howarth, 1989b).
rangeoftropical cave fauna including sandflies, Populationsizesareoftensmall,andduetotheir
plant hoppers, assassin bugs, and cockroaches isolated distribution may exhibit a limited gene
(Lewis and Dyce, 1983; Hoch and Howarth, pool and restricted gene flow, which may result
1989a. b, Malipatil and Howarth, 1990; Roth, in severe bottle necks, further increasing their
990). Oneexpeditionby Howarth and Stone in vulnerability (Barr and Holsinger, 1985;
11985 recorded over 40 species of cave arthro- Caccone, 1985; Culver, 1986). Thesmall sizeof
591
592 D.P. SLANEY AND P. WEINSTEIN
caves, compared to surface habitats, also lost as a result ofdam constructions (Eberhard
reduces the resilience of such species to and Spate, 1995). Alternatively, lowering the
disturbance. Disturbancesmaynotonlyresult in water table can have a similar effect on cave
the extinction of these organisms, but in the invertebrates. In Yanchep, Western Australia,
destruction of the unique cave habitat itself. groundwater pumping is threatening aquatic
Disturbances can be categorised as either cave species (Jasinska and Knott, 1991). In the
indirect(broughtaboutbymodificationstocave Naracoorte caves, South Australia, the transpi-
habitats and the surrounding environment) or ration from overlying pine plantations has
direct (brought about by human visitation to reduced the amount of water seeping into the
caves). caves, resulting in cave desiccation.
Indirect disturbance Quarrying
Deforestation Inadditiontochanginglocalflowregimesand
Clearingofvegetationfortimber,mining,and increasing sedimentation rates, quarrying of
road construction changes local hydrology, and limestone for cement and other building
causes severe erosion and increased frequency materials(eg marble) often entails thecomplete
and intensity of flooding (Lichon. 1993). In destructionofacaveandevenentirekarsttower
NSW and Victoria approximately 60% of the systems. Blastinginthevicinityofcavescanalso
karst regions have had their native vegetation causeseverestructural damagetothem, altering
removed or severely modified (Eberhard and the microclimate within. Limestone operations
Spate, 1995), while in Tasmania, most caving at Mount Etna, Queensland, haveledto numer-
areas are covered by intact temperate forest ous caves being destroyed, including ghost bat
which is underthreat by clearing for the wood- (Macrodermagigas) maternity caves (Eberhard
chipping industry (Lichon, 1993). Populations and Spate, 1995), and presumably their associ-
of glow worms in Flowery Gully caves, Tas- ated invertebrate cave communities.
mania, were wiped out as a result of clearing Pollution
vegetation,anactwhichmadeaoncepermanent Caves are often used for dumping ofagricul-
subterranean stream become intermittent, in tural, industrial and public waste. The occur-
turn lowing the high humidity in the caves rence of sudden large influxes of nutrients (eg
required by the glow worms (Lichon, 1993). dead farm animals)can leadtothe introduction
Clearing ofvegetation near caves also removes of surface invertebrates and allow some cave
the majorsourceofnutrient input, dramatically species to out-compete others, upsetting the
lowering the food available to cave delicate ecological balance (Chapman, 1993).
invertebrates. From 1900to 1976wasteandwashwaterfroma
The exposure of soils surrounding caves cheese factory and abattoir in Yahl, Mount
caused by clear felling, quarrying, and farming Gambier, South Australia, was discharged into
increases the rate of erosion, resulting in high an unconfined limestone aquifer. The effluent
levelsofsediment deposition in caves(Kiernan, contaminated groundwater, resulting in
1988). High levels of erosion at an Ida Bay abnormally high nitrate levels in the surround-
quarry, in Tasmania, has caused clay to be ing limestone karst (Slaney and Ragusa, 1990).
washed into surrounding caves (Lichon, 1992). At Mole Creek, Tasmania, stock access to karst
Thedepositionofclayincavepassagesnormally regions and dairy effluent runoff"has increased
containinggravelsubstrateshasledtothelossof nutrient levels, resulting in high bacterial popu-
cave invertebrates adapted to living in such lationsincavewaters(Lichon, 1993). Pesticides
habitats (Eberhard, 1995). Scouring of organ- and fertilisers may also be washed into caves
isms from their stream habitat can also occur from surrounding agricultural land (Chapman,
during periods of flooding (Chutter, 1969), 1993, Lichon, 1993). Cave entrances may not
while at Mole Creek Caves, Tasmania, land only be blocked by waste but infilled to make
clearing has resulted in rapid subsidence and way for farming, roads and housing develop-
collapse ofcave systems (Lichon, 1993). ments. Such blockages lead to a decrease in the
Changing water levels amount of nutrient input into the cave ecosys-
Flooding induced by land clearance or by the tem and result in the depletion or extinction of
construction ofdams can obliterate entire cave cave invertebrates (Culver, 1986).
communities. Cave communities at Texas in Tourism
Queensland, Burrinjuck in NSW, Dartmouth in Constructionofpathsand walkways, lighting,
Victoriaand Lorinna in Tasmaniahaveall been gates on cave entrances, and widening of cave
CONSERVATION OF CAVE FAUNA 593
entrances fortourism, create a numberofprob- also impactoncaveorganismsthrough thetoxic
lems for cave organisms (e.g., Webb, 1984). calcium carbide by-product ofthese torches.
These constructions lead to the alteration and Generally, caves as discrete confined habitats
loss of habitat, upon which highly adapted tend to have a carrying capacity of visitation,
organisms depend, by altering cave tempera- above which level of disturbance collapse of
tures, humidities and atmospheric composition cave communities is likely to occur (Howarth
(e.g., Pugsley, 1984). Artificial lighting used for and Stone, 1982, Spate and Hamilton-Smith,
tourism can enable the establishment ofplants 1991). Field data on cave fauna distributions in
that would not be able to survive in the normal Hawaiian lava tubes by Howarth and Stone
conditions of little to no light. In turn, intro- (1982) show that species diversity and popu-
duced plants may provide habitat for surface lation levels are inversely proportional to the
dwelling invertebrates, thus altering cave com- level ofvisitation and human disturbance. Dis-
munity composition (Howarth, 1982; Eberhard turbance of bat colonies brought about by
andSpate, 1995). Constructionofgateswith low human visitation, (particularly by bat
sills have a tendency to trap leaf litter which researchers !) has contributed to the decline of
would otherwise be washed into caves, as has many colonies (Tuttle, 1979). Bat decline from
occurred at the entrance to Kubla Khan Cave in humandisturbancehasbeenobserved incentral
Tasmania (Spate and Hamilton-Smith, 1991). and south-eastern NSW caves (Hall and Duns-
Changes to cave microclimates also affect bat more, 1974). In turn, the loss ofbats has led to
maternity sites and associated cave invert- the loss ofendemic guanophilic invertebrates.
ebrates(Tuttle, 1979).Theplacementofairtight
doors to control desiccation of speleothems in Impact ofscientific studies
AlexandraCaveat Naracoorte, South Australia, Despitethehigh profileofconservation issues
has wiped out populations of rhaphidophorid such as the threat to populations ofghost bats,
cave crickets (Hamilton-Smith, 1987). little work has been carried out in Australia to
assess the impact of humans on cave ecology,
with no studies on the effect that trapping may
have on cave invertebrate populations. Key
Direct disturbance (1978)and Heath(1987)claimedthatthereisno
Damagetocavesand the lossofcave fauna may evidence of collecting ever affecting a popu-
becauseddirectlybyhuman visitation.Visitsby lationofinsectsdetrimentally. However,declin-
thegeneral public, cavingclubs, and researchers ingpopulationsofthecollectedBathurstCopper
result in the trampling of some cave invert- Butterfly suggest otherwise (Dexter and Kitch-
ebrates (Spate and Hamilton-Smith, 1991), and ing, 1993), and we provide evidence in this
the compaction of cave floor sediments occu- paper that intensive sampling of cave invert-
pied by others (e.g., crickets and beetles. ebrates can deplete populations. Culver (1986)
Middleton, 1979). At Mount Widderin Cave. statedthatworldwide,oneofthedangerstocave
Skipton, Victoria, an invertebrate community fauna is the appalling numberofspecies threat-
living on unconsolidated substrate has ened by over-collecting for scientific purposes,
disappeared as a result of soil compaction by but he cites few data to support this statement.
high numbers of human visitors (Spate and With this in mind weanalysed past datawe had
pHaacmtiilotno,n-wSamliktihn,g19t9h1r)o.uIgnhaddciatvieonptoooslosil cmoamy- ofabutnaai,netdo aisnsesisnvtehsetigiamtpiaocnts tohfat toruorpicsaalmplcaivneg
adversely affect aquatic invertebrates. In Tas- may have had on populations of cave invert-
mania, rare psammaspid and syncarid crus- ebrates.
taceansare confined tosuch habitats(Eberhard, During August 1993 we surveyed the
1993), and alterations of turbidity in their macroinvertebrate fauna ofRope LadderCave,
waters may critically alter their environment. FanningRiverCaves, in tropical north Qld. and
Cavevisitorsalso posea potential threattocave recorded the weekly total ofindividuals ofeach
invertebratesbyintroducingorganismsfromthe macroinvertebrate species collected over one
surface and from other caves which may out- month ofcontinuous trapping with four differ-
compete and displace the extant cave fauna. ent methods concurrently (pitfall only, baited
Cave visitation may also result in the trampling pitfall, leaflitterdry, and leaflitterwet, with no
ofplant roots, leadingto the lossofroot feeding replacement;WeinsteinandSlaney, 1995). Four
invertebrates and their predators (Howarth, species were trapped in sufficient numbers to
1982). Cavevisitorsusingacetylenetorchesmay look for trends in their frequency of capture.
594 D.P. SLANEY AND P. WEINSTEIN
Pseudoscorpions, cockroaches, rhizophagid Queensland have an exceptional abundance of
beetles, and pselaphid beetles showed a signifi- cave species (Humphreys, 1993, Howarth,
cantdeclineinnumberoverthesamplingperiod 1988). LegislativelyonlyTasmaniaandWestern
(from 12, 44. 680 and 18 individuals to 5, 25, Australia recognises cave invertebrates as
250 and 4 individuals respectively). Given that important, but even then only on a species by
sampling occurred at the same time ofday and species basis (New. 1984; Eberhard and Spate,
that we found no seasonal effects (e.g., periodic 1995; Humphreys, pers. comm.). The formu-
rainfall) impacting on these populations during lationofconservationpolicies/strategiesforpro-
the study period (Weinstein, 1994; Weinstein tecting cave invertebrates is difficult due to the
and Slaney. 1995),theprobabilityofrecordinga lack oftaxonomic and ecological data. The lack
simultaneous decrease in populations of four ofsuch data is obviously not a problem unique
different species by chance alone is 0.0625 tocaves, but isamajorshortfallwhichthreatens
(0.54). Thus, the data suggest that our sampling the conservation of Australia's invertebrate
of cave invertebrates adversely affected their faunaonthewhole(New, ms.).Cavesmorethan
numbers. most other ecosystems can play an important
Past studieshavealso indicatedthatsampling roleinmakingboththepublicandpolicymakers
may have adverse effects on cave populations, aware of this problem, as they tend to have a
with pitfall trapping reducing cave beetle popu- relatively high local profile (e.g.,Jenolan Caves,
lations (Peck, 1975, 1976). All that may be Undara Lava Tubes, and Chillagoe Caves, all of
required to extinguish the local population ofa whicharerichincavefaunaandprovideimport-
cavedwellingspeciesisthecarelessoraccidental ant local landmarks).
desertion of a pitfall trap (Vandel, 1965; Oneoftheproblemsfacingcaveresearchersis
Howarth, 1982).Asingletrapinarestrictedarea determining whether insect species are rare, as
may not only trap a species to extinction, but they areoften difficult toobserve due tocamou-
may subsequently affect the ecology ofthe cave flagingadaptations,the crypticmicrohabitatsin
itselfthrough changes in the population dynam- which they live (cracks or rock piles), and the
icsofthesurvivingcavefauna.Apossiblewayof inaccessibility of the cave in the first place.
minimising such impacts is to sample with Further, individuals within each population
replacement, or iftrap sites are to be left in situ maybeable to find refuges from which theycan
for an extended period oftime, to use a 'trap' continually recolonise other cave regions, thus
which allows forthe free movement ofindivid- demonstrating patchy distributions both in
uals. We have designed a survey tool which space (location) and time (seasons). Howarth
meets both of these requirements (Slaney and (1983) stated that there is a requirement for
Weinstein, in press). Awet leaflittertrap(which experimental ecological studies to determine
simulates nutrient influx into drytropical caves what factors limit cave species distribution,
duringmonsoonal rains) maintainsthe viability what are the significant perturbations, and how
of organisms, thus allowing their release back thesedisturbancesaffect cavecommunities. For
into the cave environment following identifi- example, long term ecological research is
cation and counting. Because this trap is not required to establish methodologies for dis-
only conservation friendly but is also more tinguishingbetween short term population fluc-
effectiveand moreefficient than othersampling tuations and longer term irreversible changes
techniques (Weinstein and Slaney, 1995), we (Howarth and Ramsay, 1991).
recommend it as the survey tool of choice in Generally, biologists recognised that insect
these fragile ecosystems (Slaney and Weinstein, species are endangered to the extent that their
in press). habitats are endangered, and that their conser-
vation can be accomplished only by conserving
their habitats (Key, 1978). With cave ecosys-
Implications
temsthe surfaceand subsurface drainagebasins
Compared withthe USA, Australia isextremely are coupled, forming a highly integrated unit
depauperate in numbers of caves (Jennings, (White el al., 1995), and the conservation of
1983). Despite this, Australian caves are of cavefauna would thus be best achieved by pro-
international significance, containing a diverse tecting the cave ecosystem and the surrounding
range of cave organisms. Cave habitats in catchment area (not just around the cave
Tasmania are amongst the richest temperate entrance)from thesortsofdisturbanceswehave
cave communities (Eberhard and Spate. 1995), outlined in thispaper(includingscientists!). We
while tropical caves in Western Australia and recommendthatecosystemstabilityand vulner-
CONSERVATION OF CAVE FAUNA 595
ability to disturbance, as well as the general Eberhard,S. M., 1995.Impactofalimestonequarryon
ecology of organisms in a cave, be considered aquatic cave fauna at Ida Bay in Tasmania.
beforeanyintensive samplingiscarriedout. We NPWS. Hobart.
further recommend that any sampling be done Eberhard. S.M., Richardson, A.M.M. and Swain, R.,
1991. The invertebrate cavefauna ofTasmania.
with replacement using techniques similar to
Report to the National Estate Office, Canberra.
our own leaflitter traps (Slaney and Weinstein, Eberhard, S.M. and Spate, A., 1995. Cavecommuni-
in press). For cave communities where direct ties. NSW NPWS submission to the Endangered
visitation is having an effect on populations of Species Unit of the Australian Nature Conser-
cavespecies,accessmustberestricted,andtour- vation Agency.
ismshouldbeestablished only in caveswithfew Hall,L.S.andDunsmore,J.D., 1974.Asurveyofcave
species. Where possible, caves immediately dwellingbats.AustralianSpeleologicalFederation
adjacent to such tourist caves should have Newsletter 64: 9-10.
restricted access to allow organisms to retreat Hamilton-Smith, E., 1967. The Arthropoda of
Australian caves. Journal of the Australian
into a readily available refuge. EntomologicalSociety 6: 103-118.
The faunas of many caves which are under Hamilton-Smith, E., 1987. Karstcreatures: the fauna
threat eitherdirectlyorindirectlyhavenotbeen of Australian karst. Australian Ranger Bulletin
investigated at all, and may neverbe discovered 4(3): 9-10. —
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habitats, always bearing in mind that conser- Fulgoroidea). Systematic Entomology 14: 179-
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