Records of the Western Australian Museum Supplement No. 64: 85-95 (2001).

Maintenance of karst biodiversity, with an emphasisupon Australian populations

Elery Hamilton-Smith

Chair, IUCNIWCPA Working Group on Cave and Karst Protection

P.O. Box 36/ Carlton South, Victoria 3053/ Australiae-mail: elery@alexia.net.au

Abstract - Biospeleology - the stJldy of cave-dwelling organisms - has longbeen recognised as making a major contribution to the understanding of bothevolutionary and adaptive processes and community ecology. A focus on thebroader concept of the karst environment as a whole leads to consideration ofa much wider range of inter-dependent organisms, ranging from bacteria andother microbiota, through a wide range of terrestrial, freshwater andanchialine invertebrates, various vertebrates, and karst-dependent plantassociations adapted to life on alkaline soils, often with cyclic aridity. Thispaper reviews the Australian biota, identifies the major threats to itscontinuing biodiversity and discusses the importance of, and potentialstrategies for, protection. It concludes by identifying major priorities forresearch and protective action.

INTRODUCTIONAlthough there has already been considerable

world attention given to the conservation ofunderground fauna (e.g. Tercafs, 1992), this hasbeen focussed on bats (e.g. Stebbins, 1988) andtroglobitic invertebrates (e.g. Juberthie, 1995). Thereports of both a symposium organised by the KarstWaters Institute (Sasowsky et al., 1997) and a recentIUCN/World Bank meeting (Vermeulen andWhitten 1999) are probably the only two suchdocuments to deal with the total range of karstbiota. A more detailed world review which includesa section on conservation, but which still tends tomaintain a primary focus upon bats andinvertebrates (Wilkens et al. 2000), is expected toappear in the coming year.

Within Australia there has been an increasingliterature dealing with the conservation of karstbiota, but again generally concentrated ontroglobitic invertebrates or bats (e.g. Clarke, 1997;Duncan et al., 1999; Eberhard, Richardson andSwain, 1991; Eberhard and Spate, 1995; Eberhardand Hamilton-Smith, 2000; Humphreys, 2000;Poulter, 1991; Slaney and Weinstein, 1997).

The Karst ContextFor present (and most other) purposes, it is

important to think of karst as not just caves orlandscaper but rather as / . . . a karst system,incorporating component landforms as well as liferenergy, water, gases, soils and bedrock' (Eberhard,1994: 8). Its integrity depends upon the preservationof the dynamic interaction between these variouscomponents, as abnormal perturbation in anyone

of them will have implications for all others. Thisdynamic/holistic approach to definition has nowbeen widely adopted (e.g. Kiernan, 1995; Bozovic,1997; Sket, 1997; Yuan, 1988)

Many rocks, including much unmetamorphosedlimestone, have a degree of porosity to water whichhydrologists refer to as primary or intergranularporosity. Karstified rocks are more much moreporous, due to solutional processes, oftencommencing along joint or bedding planes. Thesefirst cavities or protocaves provide for secondaryporosity, and a greatly increased water movement.In turn, caverns develop and where this hasoccurred the increased porosity provides forturbulent flow and often very rapid movement ofimmense bodies of water. However, this usually co­exists with much slower flows through the still­existing protocaves or other small voids [for acomprehensive review, see Gillieson (1996)].

From the biological perspective, it is vital torecognise that any karst system will have aspectrum of void dimensions, ranging from whatHowarth (1983) has termed microcaverns. Howarththen uses the term mesocaverns for spaces rangingfrom 0.1 cm tOr say, 20 cm which cannot generallybe entered by humans and macrocaverns for spaceslarge enough for human entry or inspection. Thedifferentiation between the first two of these levelsof cavern is based in their hydrological propertiesand their relationship to flow patterns ofgroundwater (Ford and Williams, 1989). There is noequivalent conceptual boundary betweenmesocaverns and macrocaverns.

Much of the study of karst biota has focussed

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upon the macrocaverns, simply because these arethe only caverns directly accessible to biologists.Those investigating aquatic species have oftencollected from springs, but mayor may not haverecognised the extent to which those springs are fedby mesocaverns. However, Howarth and manyother workers have demonstrated the immenseimportance of the smaller voids as a habitat forinvertebrates; much of the terrestrial cave fauna isin fact a mesocavernous fauna, adapted to live inthese minute cavities with their relatively highhumidity and high level of carbon dioxide. Further,many caverns have no direct entrance to thesurface, and comprise vast networks of smalltunnels near or below the watertable, and theseprovide the habitat for a complex aquatic faunawhich has only recently even been recognised inAustralia.

This distinction may well be vital in assessing anykarst areas from a biological perspective. Anoutstanding example occurs at the Cape Rangekarst in north-western Australia, where proponentsof quarrying had argued that the area in which theyplanned to quarry had no caves, and thereforequarrying at that site did not threaten theremarkable cavernous fauna of the region. Even asuperficial examination showed that the rock wasriddled with mesocaverns, and so provided for apotentially rich fauna indeed (Hamilton-Smith et al.,1998).

As already emphasised, most biospeleologicalwork has focussed upon the subterranean fauna,particularly bats and troglobitic invertebrates.However, there is a wide range of other extremelyimportant and often neglected karst-dependentspecies:• microbiota: nanobia, fungi, bacteria, algae and

other protozoa• plants, including bryophytes and a wide range

of vascular plants, each of which are adapted toand require an alkaline environment, and manyof which are also adapted to survival incyclically arid conditions

• invertebrates which depend upon the karstvegetation, or in some other way are limited tokarst terrains

• vertebrates which depend upon specificvegetation associations, or utilise the karstterrain for shelter.

These are, of course included in regional studiesfrom those parts of the world, e.g. Europe, NorthAmerica, where immense regions are comprised ofcalcareous rocks, but the influence of the karstenvironment may well betaken for granted and notsubjected to systematic analysis.

Importance of karst biotaThe need for preservation of obscure, or often

even large and familiar, species of plants and

E. Hamilton-Smith

animals is sometimes questioned. However, theintrinsic right of all species to survival is now beingmuch more widely accepted as a basic principle inenvironmental management. One Australianspeleologist (Poulter, 1991) has attracted someattention to this issue by using the phrase 'Caverights for Troglobites!' The central discussion of thispaper is therefore not so much on why we shouldensure the survival of species, but how this mightbe best implemented.

However, it is useful to also note some of theanthropocentric arguments which might usefullysupport efforts in protection. Karst biota providesa natural laboratory of great richness which cansupport research into evolutionary processes,environmental adaptation, geoclimatic history andpopulation dynamics to name but a few areas.Certainly the remarkable discoveries of diversecave-adapted communities in northernQueensland (Howarth and Stone, 1990) and fromCape Range to the Kimberley in Western Australia(Eberhard and Humphreys, 2000) have made asignificant contribution to the understanding ofevolutionary processes and Australasian geo­climatic history. The contribution of maintainingbio-diversity within the overall gene reservoir hasbeen argued extensively in other contexts. Cavemicrobiota hold specific promise in the verypractical and economically valuablepharmaceutical industry.

Then human interest in many karst species ­watching the spectacle of bat flights, the occurrenceof karst-dependent and often very beautiful plantspecies, the intriguing and often beautiful characterof many cave invertebrates and doubtless manyother possibilities - all give a more humandimension to their importance. It has been argued(Danielopol, 1998) that that more attention shouldbe paid to the aesthetics of cave biota through itsrelationship with art - an idea which seems toresonate with the recent discoveries of cave bearskulls arranged in aesthetically pleasing patterns byour forefathers of 75,000 years ago (Lascu, 1996),and the further evidence from even very early rockart.

The pattern of Australian karst biotaAnything like a comprehensive review does not

yet exist. Probably the broadest review is containedin a paper by Eberhard and Humphreys (2000).

The micro-biota include, as in any community, animmense diversity of bacteria, fungi, algae,protozoans and other forms. There has been aninterest in cave microbiota since the pioneeringwork of Dudich in the 1930s and this is reviewed inthe Encyclopaedia Biospeologica (Juberthie and Decu,1994). Chapman (1993) provided a very accessiblediscussion of what was then known about the roleof microbiota in the ecology of cave cOJl).In.unities.

Maintenance of karst biodiversity

However, recent discoveries in other continentshave generated immense interest, and a newgeneration of research (e.g. Northup et al., 1997a). Astriking example which demonstrates the extent ofcurrent interest amongst karst scholars in theUnited States can be seen by reviewing the abstractsof papers presented at the 1998 convention of theNational Speleological Society (1998).

Regrettably, we still know very little about thiscomponent of the Australian karst biota. Managershave paid attention to lampenflora in tourist caves,and developed techniques for prevention or controlof its development (e.g. Johnson, 1979). Bacterialproduction of carbon dioxide has been recognised,and at Bungonia Caves is associated withprecipitation of iron minerals (James, 1994a, 1994b).The role of algae in the deposition of phototropicspeleothems is widely recognised (Spate, 1997), buthas only been studied in the stromatoliticspeleothems of Jenolan and some other New SouthWales areas (Cox et al., 1989; James et al., 1994).Sulphur-metabolising bacteria have beenrecognised at Bungonia Caves (James, 1994b) and inWeebubbie Cave on the Nullarbor Plain (James andRogers, 1996). It is now clear that microbiota play amuch greater role than previously thought in thefood web of subterranean communities, inspeleothem development and in other geo­mineralogical processes within karst, includingspeleogenesis.

Most biological investigation in karst has focussedupon the considerable spectrum of terrestrialinvertebrates, ranging from the interstitial speciesof soils and micro-caverns to the well-known andwidely studied troglobites and troglophiles. Thesepose a special problem in protection because of theremarkable degree of adaptive radiation andendemism - in the Australasian region it is notunusual for species to be confined to a single caveor single karst outcrop. The Australian fauna hasbeen described within the context of regionalbiogeography by Eberhard and Hamilton-Smith(2000) and more broadly by Eberhard andHumphreys (2000).

The aquatic environment occurs in a range offorms. Streams and rivers are important in manykarst provinces, especially in the impounded karsts(or Karst Barre) of the Eastern highlands. Jennings(1985) draws special attention to this phenomenon,where relatively small areas of karst are completelysurrounded by impervious rocks, and so exhibitvarious distinctive characteristics. Widespread andoften diffuse groundwater aquifers are well-knownin extensive karsts. Island karsts have a distinctivepattern of water movement, and may demandspecial consideration. In many island and coastalkarsts, the anchialine zone, where fresh and salt­water mix, provides for a distinctive faunalcommunity, as, for example, in Cape Range in

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Western Australia, and many Pacific and Caribbeancountries. It is only in recent years that theimportance of these various habitats has beenrecognised in Australia, and this is currently theleading edge of Australian biospeleologicalresearch.

Trogloxenes, particularly birds and bats, haveattracted an immense amount of study, and weknow that the survival of many species isdependent upon the availability of appropriate caveenvironments. It is also clear that the trogloxenesprovide the greater mass of food inputs to manycave ecosystems. Only one Australian bird, theswiftlet Aerodramus spodiopygius (Forster, 1845), canbe considered truly be cave-dependent (Pecotich,1980, 1982; Smyth et al., 1980) but many bats ar€normally cave-dwelling (Churchill, 1998; Duncan etal., 1999). In turn, the guano of cave-dwellingswiftlets and bats provide food for distinctiveinvertebrate populations.

The surface vegetation of karst is often distinctiveand supports a wide range of karst-endemicspecies. Only the eastern European countries have along-standing recognition of the special features ofkarst vegetation and a long record of research.Again, there is a high degree of localised endemism,due at least in part to the diversity of microclimatein karst areas.

At least in Australia, localised endemism is welldeveloped in the tropics but appears to be absent orgenerally of minor significance in temperate areas.However, this may well reflect the extremelylimited attention to this issue. In CentralQueensland, the semi-evergreen vine thicket forest(Webb and Tracey, 1970) is a relict of the Indo­Malaysian flora which was at one time far morewidespread in Australia and is now only found onthe Central Queensland limestones. It ischaracterised by tolerance of seasonal aridity andlack of resistance to wildfire, which has led tostands elsewhere in Northern Australia beingreplaced by Eucalyptus forest. At Chillagoe onefinds both semi-evergreen and deciduous vinethicket forests along with other associations on thenon-karst land units (Godwin, 1983). Again, theseforests survive because the karst surface providesprotection from wildfire. The composition of theflora is shaped by the alkalinity of the soils, semi­arid climate, and for the larger trees (particularlythe figs), access to ground-water. The deciduousforest is not only visually striking, but is the onlyextensive deciduous association in Australia.

The vegetation of Cape Range is remarkablydiverse, with both tropical and temperate species,many of which are at the northern or southernlimits of their range, a complex patterning andinter-relationship of four different plantcommunities, and a number of disjunct occurrences(Keighery and Gibson, 1993). Similarly in the East

88 E. Hamilton-Smith

Threats to the survival of karst biotaMany of the threats to karst biota are the large­

scale events which threaten the very integrity oreven survival of the karst itself. These have alreadybeen reviewed in Watson et al. (1997) and so theyare summarised here without extended discussion:

Total Destruction as a result of mining, quarry-of Karst ing, submersion beneath

water storages

Kimberley one finds an U open savanna" vine thicketflora (McKenzie, personal communication), thestudy of which is only in its early stages. Thisassociation provides the habitat for a strikingexample of local endemism amongst camaenid landsnail species and genera, with some species beingrestricted to a single hill. The 27 land snail speciesrecognised have a geographic range averaging onesquare km (Solem, 1988; Woodruff and Solem,1990). Although Solem described this as '...perhaps the greatest concentration of short rangerestricted endemic species found anywhere in theworld...', it must be pointed out that a number ofcomparable occurrences have now been identifiedin the rain forests of south-east Asia (Vermeulenand Whitten 1999, Vermeulen, personal commun­ication)

In more temperate areas, there are few karst­preferring species of vascular plants, but manybryophytes do appear to favour karst over otherenvironments. Studies by Downing (1992, 1997,1998), Downing and Selkirk (1993), Downing et al.,(1991, 1995, 1997) show that in limestone areas,bryophytes are both more diverse and morenumerous on limestone than on adjoining rocks.Species found on limestone appear to be respondingto the alkaline conditions, the relative surfacearidity of karst and the diversity of micro-habitats.Many species are also found in arid areas,particularly on alkaline soils. Not surprisingly, thisis not dissimilar in character to the patterns ofvascular plant distribution on many tropical karsts.

This distinctive vegetation together with the oftenhighly dissected terrain of karst leads in turn to anespecially attractive surface environment forspecific species of invertebrates, reptiles, birds andmammals. At least some species may be whollyconfined to karst regions, but again, there is a greatlack of knowledge.

Pollution

Human Entry tocaves or otherutilisation

Sewage and domestic drain­age, farm or industrial wastes,hydrocarbons from fuelspillage, microbial pollution.

military use, religious ob­servances and monuments,sanitoria, burial, manufactur­ing, dwelling sites, farming,wine-making, smuggling,research, tourism, concertauditoria, recreation andtourism

This listing does not make any judgement aboutthe desirability and acceptability of any specificpractice; it simply points to a range of phenomenawhich will have a threatening impact of greater orlesser degree. Many of the phenomena mentionedare of long-standing practice, are culturallyapproved, and often extremely desirable in theirown right. But all demand due assessment when anew activity is contemplated or initiated, andcontinuing re-assessment through their continuingexistence.

Threats of specific significance to the biota of karstinclude all of the above, many of which may have aboth direct and indirect impacts. For instance,clearing of vegetation obviously destroys the flora,some of which will recover given the opportunityfor re-vegetation, but other elements of the floramay never return. But the destruction of vegetationalso impacts upon soil quality and the biologicaldynamics of the soil. This in turn changes the waterregime within underlying caves and impactsdirectly upon microflora. The whole food chain ofthe fauna thus changes, and again species may wellbe destroyed. Generally, studies of karst biota haveonly taken place long after disturbance of thesurface environments, and so many of thecommunities upon which we have based ourunderstandings have already been verysignificantly modified and may well have sufferedimpoverishment. Thus, we must recognise thatthere may well be very special opportunities forresearch and protection in undisturbed areas.

Changes in water levels, either drying out ofcaves or flooding, quarrying, changes in land useand pollution are all likely to result in loss of fauna.These activities may well take place at aconsiderable distance from the site which isimpacted, and thus may destroy a population in anotherwise protected area.

Examples in Australia include:• extinction of the glow worm population at

Flowery Gully, Tasmania as a result of landclearing and reduced runoff (Lichon, 1993)

• loss of fauna, particularly Hydrobiid snails,from silting of passages in Exit Cave, within theTasmanian World Heritage Area, as a result of

monoculture forestry, quarry­ing, land clearance, construc­tion, waste disposal or otherland fill; war; de-watering orlowering of water table;extractive industries includingspeleothem harvesting, guanomining, removal of karren,birds nest harvesting, etc.

Major land orhydrologicaldisturbance

Maintenance of karst biodiversity

quarrying outside of the WHA boundary(Eberhard, 1997)

• loss of virtually all fauna from BradleyChesterman Cave as a result of the samequarrying operation along with eutrophicationand toxic pollution (Eberhard, 1997)

• flooding of Texas Caves in Queensland killedthe total karst community, including anendemic silverfish (Archer, 1978; Smith andShipp, 1978)

• disappearance of a large population of Bent­winged bats (Miniopterus) from Mt WidderinCave, Skipton, Victoria as a result of landclearing (Simpson and Smith, 1964), and aresulting decline in the associated invertebratecommunity (Hamilton-Smith, 1968), followedby virtual extinction of invertebrates as a resultof excessive trampling of the floor.

Even when an area is placed under protection,the development activities and constructions whichprovide for visitors and a range of othermanagement initiatives can have a drastic impact.The construction of roadways, car-parks andbuildings, if not undertaken with wisdom andsensitivity, can have remarkably destructiveimpacts on the karst environment. In brief,declaration of protected areas is not enough in itself- it must be accompanied by environmentallysensitive management.

Pollution, whether by soluble or liquid substancesor by increased sedimentation, is likely to havesevere impacts, often over very large areas andJuberthie (1995: 36-39) provides an excellent reviewof this problem in relation to troglobitic faunas.However, there is some evidence that recovery fromwater-borne pollution may occur relatively quicklyas new populations enter the system fromunpolluted watersheds (Lewis, 1996). One of themore insidious and potentially disastrous forms ofpollution is the eutrophication of major aquifers bythe use of agricultural fertilisers. In two regions ofAustralia where stromatolites are living in karstlakes or cenotes, this has resulted in the growth ofdense mats of invasive species of algae and otherwater plants and in turn this reduces thepenetration of sunlight to the point where the algaeresponsible for stromatolite development may wellbe killed (MacNamara, 1992; Thurgate, 1995).

Most recently, an extremely rich and diverseguanophile community in the Bat Cave, Naracoorte,South Australia (Hamilton-Smith, 1972) has beenalmost totally eliminated. This event is still underinvestigation, but it appears to the result of aninsecticide which has been proven to occur in theupper layer of guano deposits in the cave. It is thuspart of the changes in environmental conditions dueto changes in land use with increasing agricultureand new technology in insect eradication.

Another common source of pollution results from

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the development of pathways and other structuresfor tourist access to caves; potentially dangerouspollutants are often introduced to the cave. Theseinclude copper from discarded waste left byelectrician, zinc and cadmium from galvanisedmetals and hydrocarbon spillage. A recent reviewof this problem at least raises a warning to thoseresponsible, and develops a series of proposals forimproved practice (Spate et al., 1998).

Human entry to caves may have drastic impacts.Part of the problem is that many people seespeleothems as the karst resource of mostimportance and totally ignore the value of the cavefloor. As a generalisation, in spite of the very realaesthetic value of speleothems, they are bothincredibly abundant and of limited other value.Floors, on the other hand, are often an incrediblelibrary of natural records of the past - layeredsediments, pollens, sub-fossils from many phyla ofthe animal kingdom, and human or proto-humanbones or artefacts. From the perspective of thispaper, they are one of the more important biotopes,often the key habitat for both microbiota and adiversity of invertebrates.

In caves which have been developed for tourism,pathways have usually been laid on the cave floorwith no regard to what may be destroyed in theprocess. However, with proper path construction,the environment may well provide more effectivelyfor continuing survival of biota than mightotherwise be the case. As noted above, at MtWidderin a remarkable relict community ofguanophilic invertebrates which had survived for100 years after the departure of the bats on whichthey had depended for food (Hamilton-Smith, 1968)was wiped out in a couple of years byindiscriminate trampling of the cave floor. A simpleelevated pathway system would have preventedthis catastrophe.

Turning to threats which are specific to biota, twoproblems seem to be pre-eminent. The first,regrettably, is over-zealous or poorly plannedresearch and collecting (Slaney and Weinstein,1997). Many of us have encountered this problem,and it is alarming that often those responsible arescientists. There is probably a much smallerproblem with collecting for commercial purposes ­although in Europe, the market for troglobiticspecies, especially beetles, has been a significantproblem, while in Thailand, the survival of Kitti'sbat (Craseonycteris thonglongyai Hill) was severelythreatened until the government stepped in tocontrol collection and trafficking, apparently withreasonable success.

The second is the problem of invasive species.The ubiquitous cockroaches Periplaneta americanaL. (Yussof, 1997: 7-8, 29-30) and P. australasiae F.(Price and Steiner, 1999) have invaded the BatuCaves of Malaysia in enormous numbers and both

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have been recorded from other cave sites inMalaysia and Thailand. Regrettably, this invasionwas well established by the time it was properlyrecognised, but seems to have been associated withthe opening of the cave for visitors in 1974, andmay have been carried in with building materialsor even on the feet of visitors. It has beensuggested that this invasion may well be largelyresponsible for the major decline in populations ofboth the Malaysian cave cockroach, Pycnoscelusstriatus Kirby, and the remarkable Liphistiusbatuensis Abraham spiders for which this cave wasfamous.

At another level, Downing (1997, 1998, alsoDowning et al., 1997) has described the way inwhich invasive species have led to a decline in theendemic species of bryophytic flora on karst. Moreobviously, the vine thicket forests of NorthernAustralia have been severely damaged by theinvasion of both Lantana and Rubber Vine.

There are doubtless hundreds of other examplesand many may be unrecognised because theinvasion occurred prior to relevant biologicalresearch. As a possible Australian example, thedominant ubiquitous scavenger beetle Alphitobiusdiaperinus Panz. in Bat Cleft Cave at Mt. Etna,Queensland, may be part of the original fauna, butis more likely to have arrived in the cave sinceWestern settlement (Hamilton-Smith, 1970). Thereis certainly some invasion by microbiota, butagain, there is too little research to detail this inAustralia.

Ensuring ProtectionThere is a hierarchy of possible protection

strategies which may be utilised (Eberhard andHamilton-Smith, 2000).

Legislative protection ofspeciesWhile many countries have in fact enacted

species protection of a number of vertebrates andplants, and this has been utilised in the protectionof cave fauna in at least both Slovenia (since 1922)and Croatia, Tasmania and Western Australia arethe only state of Australia where cavernicolousinvertebrates have been proclaimed as protectedspecies, one of which is also coverd byCommonwealth legislation. However, theeffectiveness of this approach has been questionedon a number of grounds: species must beindividually identified in the legislation and thismay pose taxonomic difficulties while manyspecies of cavernicoles are undescribed and socannot be listed; enforcement is virtuallyimpossible; and species protection cannot beinvoked to prevent the destruction of habitat.There are now moves to develop legislativeprotection of specific ecological communities, andif able to be implemented successfully, this will be

E. Hamilton-Smith

an effective response to some of the difficultiesinherent in species protection, and may offerexcellent possibilities in relation to karstcommunities. Several subterranean communitieshave been declared protected in WesternAustralia, including troglobitic, stygal and rootmat ecosystems.

Recovery planningfor threatened speciesThe development of systematic recovery plans

for species or groups of species is being developedin some countries. Although the effectiveness ofthis is often constrained by lack of research andthe very complexity of ecological communities,there may well be occasions on which it might bevery usefully invoked. The Bat Action plan(Dunean et al., 1999) is an excellent and relevantAustralian example. The rehabilitation of the IdaBay quarry site in Tasmania also provides aspecific example of planned action to restore acave community (see below).

Protection of SpeCifiC Habitat AreasIn itself, this group of strategies provides a

considerable hierarchy of approaches. At thesmallest level, it includes actions such as:• track marking in caves to prevent trampling,• development and voluntary observance of

minimum impact codes for cavers orresearchers. This is an important initiativewhich targets populations of cavers andresearchers. An Australian example is providedin Watson et al. (1997) while many other fineexamples have been developed, certainlyincluding those of the United Kingdom andSwitzerland.

• closure of caves or karst areas (often at specificseasons for the special purpose of protectingcrucial bat sites). One such example, voluntarilyinstituted by cavers, resulted in the recovery ofa bat population after a 20-year absence(Hamilton-Smith, 1991).

• finding means to minimise, control or eliminateinvasive species

One of the dilemmas which we currently face isthat having recognised the importance ofmicrobiota, we lack the detailed knowledge todevelop sound protection programs. But bothinvasive species and pollution represent clearthreats, and the latter must be more broadly definedthan is often the case. Invasive species are also asignificant threat. Pollution includes anyimportation of organic matter, even including theskin flakes, hair and lint left behind by any humanentry. It also includes any materials, e.g. clotheswhich are not freshly laundered, or dirty boots,which may serve to carry invasive species ofmicrobiota. Northup et al. (1997b) have d~velopeda

Maintenance of karst biodiversity

series of practical guidelines for use by cavers andresearchers. These are unlikely to be widelyobserved in many situations - and some caves havealready been so impacted that any damage hasalready been done. However, they may be of greatimportance in entering and assessing new andpreviously uninvestigated cave systems. There is afurther major threat in alteration of air movementsin any previously closed or severely constrictedcave.

However, area protection may also be developedon a broad-scale level, and the proper location oflimestone quarries is an excellent example of this.Quarrying is one of the major threats to the integrityof karst in many countries, and often demonstratesa virtually irreconcilable conflict of interest. Majordisputes have occurred in Australia at Mt Etna(Queensland), Yessabah, Colong, Bungonia andWombeyan (New South Wales), Sellick's Hill (inSouth Australia) and Ida Bay (Tasmania). But fiveof these Australian examples are no longer beingactively quarried, and three are receiving extensiveand thorough rehabilitation. Steps are now underway to develop a greater recognition ofconservation values and to negotiate both co­operative planning on extraction and the use of newtechnology which will ameliorate impacts upon thebiota.

Rehabilitation and RestorationWhich brings us to the growing importance of

rehabilitation and restoration. Both tourismmanagers and speleologists have been involved inaesthetic restoration of caves, often withconspicuous success, and in turn this may wellrestore cave habitats.

More importantly, there are now outstandingexamples of restoration of total karst habitat,including:• Waitomo Cave, New Zealand, where the

famous glow-worm population was seriouslythreatened by degradation of the wetlandhabitats which produced the rich population ofDiptera upon they depended for food,

• The Horse Cave and Hidden River system inKentucky, U.S.A., where cessation of thepractice of using the cave for sewage and wastedisposal had led to an extensive recovery of thefaunal community (Lewis, 1966),

• Ida Bay Caves, in the Tasmanian WorldHeritage Area, where cessation of quarryingand rehabilitation of the former quarry led toa remarkably rapid recovery of bioticcommunities. The decision was made to notutilise the artificial fertilisers demanded byforest ecologists as their use would have hadnegative impacts on the troglobitic and otherunderground biota. This projectdemonstrates in various ways the importance

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of adapting rehabilitation technology to aresearch-based understanding of the siteconcerned, rather than seeking any set ofrigorous and universalised procedures(Eberhard, 1997; Houshold, 1997; Gillieson,1995).

Declaration ofNational Parks and other Protected AreasThis is clearly the option which has the best

potential to protect total environmental systems,such a karst province. However, several warningsmust be noted:• It is all too common for a government to

declare parks, encourage (even just by makingthe decision) the public to visit them, yet notprovide adequate resources for safeguardingof park values. The result may well be anincrease in degradation of the environment.In other words, declaration is not enough: itmust be accompanied by adequate resourcesfor management and protection of parkvalues. There are often major issues here inthe lack of exp~rtise on issues in karstmanagement, and consequently a very realneed for development of professionaleducation for karst managers.

• The boundaries of karst drainage systems oftendo not co-incide with surface drainage systems.Unless the total watershed is included withinpark boundaries, this may lead to significantchanges in the water regime within the park asa result of off-park actions (Neale, 1985).Waitomo (New Zealand) provides a well­documented example, where the karst systemwas seriously threatened by increasedsedimentation. However, this also demonstratesthat where it is not realistic to include the totalwatershed within park boundaries, a problemmay be solved by negotiation withneighbouring landholders and/ or usingavailable planning ordinances (Simmons andLohrey, 1985).

On the more positive side, not only has there beenan immense increase world-wide in the total areaunder some form of permanent resource protection,but there has been a steady increase in globalexpenditure on resource protection and a rapidlygrowing body of research and knowledge.

International agencies have provided a valuableresource in supporting this development, whileinternational treaties have provided for WorldHeritage Recognition, usually with substantialstrengthening of management. The recentrecognition, initially in Australia (Australian NatureConservation Agency, 1996), of some major karstaquifers as wetlands under the Ramsar Conventionprovides a further tool for protection and this iscurrently under examination in Eastern Europealso.

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Public EducationThe importance of public education can never be

neglected. The development of more effective karstprotection, in the long run, depends upon thelegitimisation of governmental action by the public asa whole. This must encompass a spectrum from theinclusion of karst understandings in science education,the education of park managers, cavers , tourismoperations and others actively involved in karstutilisation, land owners, to wider public educationthrough interpretation at parks, journalism, electronicmedia and popular books for all ages.

Some Key IssuesIt is clear from discussion throughout this paper that

many aspects of the Australian karst environmenthave not been adequately investigated. In particular,all too little is known of the surface flora and fauna.We have come to realise that if we are to develop asatisfactory understanding of the surface ecology ofkarst that we must recognise the extent to which thereis often a mosaic of micro-habitats, each with theirown microclimate, soils and biotic community. Thusat Jenolan Caves it has proved necessary to develop aspecial series of maps developed in order to providefor recording of this data (Thurgate and Gillieson,1999). Similarly, we have already referred to thefindings of Keighery and Gibson (1993) on the CapeRange flora. In brief, overall generalisations are notenough - we need to understand the complex natureof, and interrelationships between, the various micro­habitats.

There are also very few cave communities wherewe have developed an understanding of theecology. Again, although it is vital to accuratelydetermine species, we also need to understand thevarious community structures, and where a numberof discrete communities occur in one cave system,the relationships between them.

Turning to conservation, it now appears that manycave communities may not be as vulnerable as weonce thought. If, as discussed above, the primaryhabitat of many troglobitic invertebrates is in themesocaverns, then these are not likely to besignificantly impacted by human entry and impact inthe macrocaverns. Lewis (1996) has certainly invokedthis as one source of the recovery populations in theHidden River System in Kentucky. But we still needmuch better public understanding, more rigorouscontrols over both levels and pollution of karstaquifers and much more adequately karst-sensitiveenvironmental impact assessments.

ConclusionThis paper provides an all too brief overview of

some key issues in karst protection. Australia isfortunate in that some of us commenced workingactively to promote better protection andmanagement of karst over 40 years ago - it has been

E. Hamilton-Smith

a long road, so we have both a lot of experience anda lot of mistakes from which we have learned more.We still have a long way to continue travellingalong that road, even turning back at regularintervals to revisit parts already travelled.

Regrettably, one of the things we have learned isthat resource protection is a continuing strugglewith human greed and global industrialisation(Bonyhardy, 1993: 145-146). Although the last 25years have seen immense advances in protection,we have also seen reversals by governments whoare all too readily persuaded that mining, tourismand other industrial activities matter more thanpreservation of our natural heritage. So protectionis not just a matter of knowledge, it also demandscontinuing political will and political skills.

ACKNOWLEDGEMENTS

A broad-ranging review of this kind always owesa great debt to one's colleagues who are toonumerous to name individually but they are valuedand appreciated none the less. In this case, theauthor also expresses sincere thanks for the verythorough and critical review by the referees. Theirefforts have contributed a great deal to the qualityof the paper.

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