Conservation of cave Fauna in Australia
Stefan Eberhard and Elery Hamilton-Smith
ACKMA Journal No. 23, June 1996. Pages 4 – 14.
INTRODUCTION
Despite the high level of interest in caves during the 19th century, and some promising early beginnings in the study of cave biology in Australia, research on cave fauna has been extremely limited until the last 30 years. Historically, Australian cave communities have been perceived as somewhat depauperate compared with the well-studied communities of Europe and North America, especially with regard to the representation of taxa which display a high degree of adaption to the cave environment. During the last decade in particular however, the traditional perception has been overturned by some dramatic discoveries of highly diverse communities containing ancient cave adapted species. It must be appreciated that many Australian karsts are poorly known biologically, and that much of the fauna remains undescribed. At the same time, some cave communities have been lost entirely and a great many more have been seriously degraded or are threatened by human activities. This article describes the major conservation issues and nature of impacts threatening Australian cave communities, and discusses the conservation strategies which can be applied to them.
BACKGROUND
In the prehistoric past Australia was part of the southern supercontinent Gondwana and was attached to Antarctica. The two lands shared a Gondwanan flora and fauna, and when the final separation between them occurred (45 million years ago), both lands were well-watered and supported cool temperate and subtropical forests. As the Australian continent separated from Antarctica and drifted northwards it began to dry out, its forests contracted and were replaced with arid-adapted vegetation. The formation of the Antarctic ice cap 15 million years ago saw the beginning of a series of marked climatic fluctuations which has greatly stressed the flora and fauna. Warm and wet interglacials alternated with very dry, cool and windy glacial stages, although the continent was not subject to extensive ice cover with the exception of a small area of the Eastern Highlands and the island of Tasmania. The cyclic fluctuation of the climate in the late Cainozoic, superimposed upon a generally increasing and spreading aridity over the last 2.4 million years, provided conditions under which subterranean refugia played an important role.
Many of Australia’s karst landscapes are quite ancient, and may have been available for colonisation by animals for a very long period of time. Palaeokarst features of Devonian age (350 to 400 million years ago) are known in New South Wales for instance, although subsequent episodes of burying, marine transgressions, flooding, sedimentation and major climate changes may well have eliminated faunas from that time. The occurrence of highly modified stygobiontic taxa belonging to morphologically conservative groups such as the Anaspidacea, imply that subterranean freshwater environments have been continuously available for a very long period, possibly since the end of the Palaeozoic era. The distribution of remipedes and their associated fauna can be explained by cave colonization along the shores of the Tethys Seaway during the Triassic and Jurassic (225-160 million years ago), whilst during the early Cretaceous the plate that is now Western Australia formed the eastern shore of Greater Tethys.
Karst, pseudokarst and groundwater localities are distributed across the Australian continent which encompasses climate types ranging from tropical through to temperate. Although areas of carbonate rock occur less frequently in Australia compared with the world average there are about 500 discrete localities documented. In addition there are cave and groundwater communities established within non-carbonate rocks such as basalt, sandstone, quartzite, acid igneous rocks, unconsolidated sediments and others. These include lava tunnels, sea caves, boulder and talus caves, acolian caves, and Interstitial environments.
THREATS TO AUSTRALIAN CAVE COMMUNITIES
Ever since European settlement of Australia a little over 200 years ago the natural environment has endured a legacy of neglect and abuse. This has been the result of a colonial consciousness which has fostered wholesale exploitation of the country's natural resources. Many karst and cave communities have suffered the brunt of Australian cultural tradition, however the last decade or so has witnessed a dramatic improvement in awareness as environmental issues have figured prominently within the social and political arena. Indeed, there is cause for optimism in the future outlook as a number of major conservation battles have been recently fought, and won, with karst issues being a key ingredient. There is little room for complacency however, because some highly significant sites have already been lost whilst others remain imminently threatened.
QUARRYING AND FLOODING
A number of caves, along with their resident fauna, have been obliterated through quarrying activities, whilst others have been inundated following the construction of water impoundments. Some of these sites were the focus of high profile conservation campaigns which ended up being settled in the law courts, however the hammer has fallen both ways in recent times. A conspicuous conservation campaign was waged at Mount Etna in Queensland where quarrying of limestone for cement had been steadily removing a number of significant caves (Bonyhardy 1993). Eventually, during the course of lengthy legal proceedings, speleologists resorted to occupying caves to prevent further blasting. This phase of the battle ended abruptly in 1988 when the quarry company illegally blasted-in the entrance of Speaking Tube Cave which was an important roost site of the endangered Ghost Bat, Macroderma gigas. The cave was later destroyed, although a change of government placed the remaining caves in the Fitzroy National Park.
The Mount Etna experience does not stand alone and a number of sites were lost before the importance of karst and cave communities was recognized outside of the speleological fraternity. Texas Caves in Queensland for example, were the type locality of a rare troglobitic silverfish and other undescribed invertebrates, but they were inundated by waters of the Glenlyon Dam (Archer 1978). In New South Wales, construction of the Burrinjuck Dam flooded the type locality of the rare Horseshoe Bat, Rhinolophus megaphyllus (Ryan 1965). More recently, during 1994 the government of South Australia permitted the blasting-in of a highly significant cave system at Sellicks Hill. Despite this setback there have been a number of successful conservation campaigns launched in other states. During the 1970s, the Colong karst in New South Wales and the Precipitous Bluff karst in Tasmania were both spared from quarrying proposals due to concerted efforts by speleologists and other minority groups. In the late 1980s cave archaeological sites on Tasmania Franklin River were a key piece of evidence which helped to save the river from being dammed. In 1992 a limestone quarry at Yessabah in New South Wales ceased operations following legal challenge by a speleologist who successfully demonstrated that he held a "vested interest" in the caves. His case was enhanced by the fact that one of the caves threatened was a major bat roosting site. Cave faunal values were a significant criterion in the nomination of Exit Cave in Tasmania for World Heritage status, whilst evidence of deleterious impacts of quarrying on this fauna contributed to closure of the operation in 1992.
LAND CLEARANCE AND AGRICULTURAL ACTIVITIES
Land clearance (deforestation) and agricultural activities (crop-raising, stock-raising, and forestry) are the most common and widespread threat facing karst biota in Australia. The impacts resulting from these activities are often subtle and difficult to quantify because much of the degradation occurred long before the biota had been recorded. The Impacts which potentially affect the biota include alteration to cave microclimates (including moisture, carbon dioxide and other parameters) and nutrient inputs, as well as changes to hydrological regimes and water quality, particularly sedimentation, nutrient enrichment and pollutants.
The extent of impacts can be gauged from comparative studies undertaken in South Australia on the effect of exotic pine (Pinus radiata) plantations on groundwater levels and atmospheric moisture regimes. In Sheathers Cave the water level dropped by approximately one metre over a five-year period following the establishment of a pine plantation above the cave, whilst conversely, the level in Mount Burr Cave rose significantly when the pine plantation above it was destroyed by wildfire (Grimes, Hamilton-Smith & Spate 1995: 16). A significant increase in humidity, which was associated with the re-generation of previously dormant speleothems, occurred at Naracoorte Caves following the removal of pines. Similarly, caves beneath pine plantations at Yarrangobilly in New South are less humid and contain less biota than those beneath indigenous forests (New South Wales 1983).
Observations have indicated a general decline in the numbers of cave dwelling bats in central and south-eastern New South Wales and, anecdotally, elsewhere. Throughout New South Wales there are many caves which previously had extensive use by large numbers of bats, but which now are used only infrequently by smaller numbers, or not at all. The evidence includes the absence of bats from large numbers of caves which contain piles of bat guano and roof staining from bat urine. The decline may be attributed, at least in part, to widespread deforestation which has severely restricted the area of suitable foraging habitat available to the insectivorous bats. Use of inappropriate insecticides may lead to the accumulation of residues in bats and consequent mortality (Dunsmore, Hall & Kottek, 1974), although direct evidence of this having occurred to cave dwelling species is lacking.
Local extinction of guano-dependent invertebrates has occurred at the sites which are no longer used by bats. One of the few documented examples involves the disappearance, between 1866 and 1895, of a large maternity colony of the Common Bent-Wing Bat (Miniopterus schreibersii) from Mount Widderin Cave in Victoria (Simpson & Smith 1964). The bats probably abandoned the cave due to the clearing of native scrubland, which also caused more rapid percolation of water into the cave. In turn, the loss of the bats impacted upon the invertebrate fauna of the cave, which now supports less than half the number of taxa recorded from other Miniopterus maternity sites in the region (Hamilton-Smith 1968).
Water table lowering in caves is acutely evident in Western Australia in the Augusta-Margaret River region, and at Yanchep where a uniquely rich community associated with underwater mats formed from tree roots is at risk (Jasinska & Knott 1991). The Cape Range stygofauna is also threatened by water abstraction.
Changes in flow regime and water quality become evident following alteration or removal of the native vegetation cover. The changes include an increase in water yield, thus exacerbating flooding in caves (Kiernan 1988). More rapid run-off after a precipitation event increases the flashiness and peak discharge of streams flowing into caves. Previously perennial stream courses may become ephemeral or intermittent. A limestone quarrying operation has caused significant adverse impacts to aquatic cave-dwelling fauna at Ida Bay in Tasmania. Changes in flow regime and water quality (including sedimentation, eutrophication, and toxins) are responsible for causing the extinction of fauna in Bradley Chesterman Cave (Eberhard 1995).
A number of recent Australian studies of surface waters implicate sedimentation as a primary cause of stream degradation and faunal decline, and this appears to be the case in underground streams also. In Exit Cave at Ida Bay in Tasmania for example, sedimentation originating from a limestone quarry has altered stream habitats and restricted the distribution of hydrobiid snails. During February 1992, when the quarry was operational, a study showed that snail abundances were significantly lower in sediment-affected streams compared with control streams. After closure of the quarry operation in October 1992 a rehabilitation program has sought to minimize further environmental degradation of the cave system by restoring natural inflow regimes and limiting the further influx of sediment. Continued monitoring of the snail populations now indicates similar abundances between the sediment-affected and control sites (Eberhard 1995). Since European settlement, many other stream cave systems in eastern Australia have been subject to increased rates of sediment deposition as a result of land-use activities within their catchments (Gillieson 1989).
The protection of water resources in general, and karst groundwater resources in particular, is an acute problem within rural communities throughout Australia. Ignorance and disregard of the nature of karst hydrological systems has severely compromised water resources and subterranean fauna in some localities. The impacts have been caused by improper rubbish disposal in sinkholes, and poor management of pollutants. Municipal disposal sites at Mole Creek in Tasmania and Buchan in Victoria both drain directly into karst aquifers. On one occasion, some 300,000 litres of waste milk was dumped in a cave at Allensford in Victoria (White 1976).
An extreme example is the dumping of thousands of sheep carcasses in Earls Cave, South Australia, and other dumping at this site over a long period (Aslin 1972a). One study in the region identified 26 other caves, 21 industrial drainage bores and a variety of other natural depressions as potential points of entry to the aquifer for pollutants (Aslin 1972b). The extent of the problem is indicated by the nitrate pollution of this aquifer being ten times in excess of safe levels (Waterhouse 1973). During the late 1980s there was a sudden and unexplained die-off of plant communities in one of the springs, Piccaninnie Ponds, which drain this aquifer. However, there is evidence of impacts to the biota occurring much earlier than this. The first European settlement in the region was at a site in Mount Gambier now known as the Town Cave, selected because of its ready access to good water. Woods (1862: 359-360) described this cave, saying that the water in the bottom of the cave was "….so deep as to give it, clear as it is, a deep sea-blue tint". He then added The water is full of a cypris and cyclops, the shells of which seem to strew the bottom. there is also much conferva, a shrimp-like brachiopod and a minute paludina. The site later became a drain for the waste of the growing town, and today the water level is lower, with accumulations of silt and rubbish ; the water is dirty and foul-smelling, and the aquatic community described by Woods has disappeared.
Nutrient enrichment may displace stygobiont communities and facilitate colonization of underground waters by epigean taxa. This has occurred at Bradley Chesterman Cave in Tasmania, although petroleum hydrocarbons have also been implicated in elimination of the original community at this site (Eberhard 1995). Pollution from sewage has been identified at both Mole Creek and Buchan, although action has been taken to eliminate the problem at the latter site. Subsidence in a limestone area also ruptured a sewage main in Tasmania (Kiernan 1988 : 9-10). Eutrophication is the major threat to stromatolite colonies in karst lakes and cenotes in Western and South Australia (McNamara 1992; Hallam & Thurgate 1992, Thurgate 1995). Run-ofr laden with agricultural fertilisers has increased the concentration of phosphates in Lake Clifton ten-fold. The resultant algal blooms have stifled continuing growth of the stromatolite colonies by reducing available light levels, as well as directly smothering them.
HUMAN VISITORS
Direct impacts to cave communities have been caused by human visitors who include guano ,miners, cave managers and tourists, recreational cavers, and scientists.
Traditionally, cave managers concerned with developing caves for public show have seen their conservation responsibility as being confined to the preservation and display of the most beautiful speleothems, and until recently, scant regard was given to the habitat requirements of the fauna. Cave microclimates have been significantly altered by the excavation of new entrances and connecting tunnels, or the enlargement of existing entrances. Additionally, entrances have been gated or sealed-off entirely, thus inhibiting access for bats and other trogloxenes, and altering the input of nutrients such as leaf litter and accidentals. At a number of sites, cave entrance gates have subsequently been modified to allow the free passage of bats, or to restore the original microclimatic conditions. The Alexandra Cave at Naracoorte, for example, was discovered by digging open the entrance, thus allowing entry to haphidophorid crickets which established a flourishing colony and created considerable interest amongst visitors. Later it was realized that the free air flow permitted through the wooden gate placed on the new entrance had desiccated the cave’s speleothems so a new air-tight door was installed to encourage their regeneration. This succeeded, but did of course result in loss of the cricket colony.
The metabolism of large numbers of human visitors affects cave microclimates by increasing carbon dioxide levels, moisture and temperature. Visitors also introduce lint, skin flakes, soil particles, food scraps, microflora, and fungi. These provide a rich food source which benefit some cavernicolous species but may also permit colonization by adventitious species. This phenomenon is evident at Jenolan Caves in New South Wales where large populations of springtails dwell alongside the tourist paths. Lampenflora growth also encourages colonization of the dark zone by invertebrates which are normally restricted to the twilight zone.
Other foreign materials which tend to accumulate in tourist caves include soot from carbide and other early combustible lighting sources, pitch from old lighting seals, verdigris from scraps of electrical wiring or from coins placed in 'wishing wells', broken light globes, scrap metal and timber (Bonwick & Ellis 1985). Old and disused timber left dumped in caves attracts cavernicolous species, and clean-up programs, although well intentional, can cause mortality to quite a few individuals when the timber is removed.
Recreational caving activities have caused widespread and profound disruption to bat populations, and colonies which are repeatedly disturbed may abandon their cave sites and not return to them, such as occurred at Cotter Cave in the Australian Capital Territory and elsewhere. Even minor disturbances of bats at their over-wintering sites and maternity roosts may be fatal. Caving activities have increased in popularity over the last 25 years especially, and may be part of the cause for the decline in bat populations throughout New South Wales and Victoria during this period, although hard data is lacking. Most cave visitors are simply unaware of the disturbance they may cause, but occasionally bats are deliberately killed.
During recent years however, bat conservation has been fostered by some land management agencies and caving groups. For example, a number of caves at Bungonia are closed annually by the New South Wales National Parks and Wildlife Service during critical periods of the bat breeding season and "hibernation". Furthermore, Horseshoe Bats are once more breeding in Anticline Cave in Victoria, as a result of the restraint that cavers (and the private landholder) have shown in limiting visits to the cave (Hamilton-Smith 1991).
Floor-dwelling invertebrates in many caves are widely under threat from recreational cavers who inadvertently trample them underfoot. (Hamilton-Smith 1968, Spate & Hamilton-Smith 1991). A side effect of intensive trampling is the compaction of soft floor sediment which may render it less suitable as invertebrate habitat. In extensively trampled caves the area of substrate inhabited by invertebrates has been greatly reduced, and the animals are confined to untrampled substrates alongside the walls, or in accessible crevices. This phenomenon is evident in many popular caving areas. Other microhabitats which are vulnerable to damage from trampling are pools and small watercourses, which often contain a distinctive and specialized fauna.
The final direct human threat to cave fauna concerns the activities of scientific researchers. At least part of the decline in the numbers of cave dwelling bats during the 1960s and 70s may be attributed to the pressure of specimen collecting and banding programs. The former activity is known to have decimated local populations on at least two occasions (Hamilton-Smith 1970). Participants in the latter scheme took particular care to minimize the disturbance they caused, and this research has yielded much useful information on the ecology of species (Simpson & Hamilton-Smith 1965). Some concern has been expressed about the potential for over-collecting of invertebrate specimens.
STRATEGIES IN CONSERVATION
Present understandings of the nature of karst landscapes emphasize the importance of the inter-relationships between environmental conditions prevailing on the surface and those underground. We argue that this is fundamental to the development of effective strategies for the conservation of subterranean faunal communities. This approach may involve, for instance, the management of water catchment areas which extend well beyond the geological boundaries of the karst outcrop. Aside from the highly publicized conservation campaigns which have been waged in attempts to conserve imminently threatened sites, the conservation strategies which have been applied in Australia to date include: legislative protection of threatened species and development of recovery plans for them; the protection of areas of sensitive habitat at individual cave sites; the location of karstlands within National Parks and other land reserves; and public awareness campaigns. Each of these strategies is discussed in turn below.
SPECIES PROTECTION
Virtually all bat species are protected under legislation throughout the Australian region. The legislation has constrained the shooting of bats, but wasn't implemented until long after the widespread loss of bat foraging habitat through land clearance activities had already occurred. The highest level of protective legislation is afforded to species which are considered endangered. In New South Wales for example, the bent-wing bats (Miniopteris spp.) are listed as vulnerable under Schedule 2 of the Threatened Species Conservation Act 1995. This means that forestry or other development proposals must, in addition to undertaking an environmental impact assessment (EIA), also carry out a fauna impact statement (FIS) if a Schedule 2 species is involved. This legislation potentially offers considerable security to the listed species, but for some reason, aquatic species are excluded from protection under the act.
Legislative protection of cave dwelling invertebrates has generally lagged far behind that for vertebrates although in 1971 a number of Tasmanian cave species were wholly protected under the Parks & Wildlife Act of 1970 (Tasmania). Although a remarkable step forward at the time, a number of problems have become evident. When the legislation was enacted the dozen or so species listed comprised, virtually, the known extent of Tasmania's cave fauna, so despite the discovery of many additional taxa in the meantime, it is still widely and erroneously believed by park managers and academic biologists alike that all cave fauna species are protected. A permit is required to collect any of the listed species, including those located outside of national parks or state reserves. The Threatened Species Protection Act of 1995 incorporates a number of the recently described taxa, but the majority remain undescribed and therefore cannot be listed. It could be further argued that some of the species are not threatened at all. The original proclamation was hurriedly instituted in reaction to fears of scientific over-collecting of specimens of one species which, ironically, is one of the most abundant and widely distributed. Although species protection of this kind may inhibit deliberate killing of individuals, it cannot be invoked to prevent trampling, pollution or other destruction of habitat.
A more sophisticated approach is the development and implementation of recovery plans for endangered species. This approach is multi-faceted and often involves habitat restoration and protection, public education programs, and enhanced breeding. Currently a recovery plan for Australian bats is being developed, and will include attention to cave-dwelling species. A few plans have been prepared for non-cavernicolous invertebrates, but there are formidable problems involved, including the lack of sufficient baseline information to identify species demanding such action, lack of research upon which to base plans and the complexity of implementation. Nevertheless, this approach must be pursued where it appears to offer the opportunity for more adequate conservation.
A substantial portion of Australia's invertebrate cave taxa remain undescribed, and they are likely to remain so in the foreseeable future. Thus, the taxonomic impediment will continue to be a major stumbling block in gaining protection for individual species. One solution may be to seek protection of ecological communities, which is provided for under the Australian Endangered Species Protection Act 1992, and at the time of writing a proposal has been submitted which seeks listing of cave communities as an Endangered Ecological Community under the Act (Eberhard and Spate 1995). At the very least, such recognition can be used as a basis for further conservation action.
HABITAT PROTECTION
The protection of areas of sensitive habitat within individual caves, coupled with an education program, has proved to be a pragmatic and successful conservation strategy which works much along the lines of a community recovery plan. Thus, in Mullamullang and Nurina Caves on the Nullarbor Plain, certain key habitat areas have been delineated with protective strings and signs which explain the reason for trying to exclude visitors from these areas (Poulter 1991, 1994). Similarly in Kubla Khan and Little Trimmer Caves in Tasmania, so-called substrate protection zones have been maintained by marking out pathways and "no-go" areas. This approach relies upon voluntary compliance by the cave's visitors of course, and its relative efficacy will depend on a number of factors such as the accessibility and popularity of the site, the presence of a dedicated management authority if any at all, and the type of people who visit the cave (e.g. speleologists cf the general public). There are many caves where the impacts of trampling are ongoing, and these sites would greatly benefit from some form of habitat protection.
LAND RESERVES
Reserves for the specific protection of caves, as opposed to cave communities per se, were first gazetted at Wombeyan in 1865, and then elsewhere later. Although some of these reservations were adequate, many were not, often only encompassing the cave entrance rather than the whole cave or karst system, and very rarely protecting the total catchment. Hence, substantial portions of some reserved cave systems extend beneath private land, state forest, or other land tenure, thus compounding the task of management and conservation. In some instances this problem has been solved by further land acquisition, but it may involve protracted negotiations to settle conflicts of interest.
Quite a number of important karst areas are located within National Parks or other conservation reserves, although more often than not their inclusion has been by fortuitous circumstance rather than deliberate intention. Consequently, some parts of the karst system, or its catchment, may not be included within the reserve boundaries. Such land protection is no panacea, and a range of other problems may arise, particularly in relation to fauna conservation. Lack of training for park managers in karst issues, combined with limited funding availability for karst-related conservation projects, is one of the more important. Many caves in National Parks are under great pressure from large numbers of recreational visitors and in some parks this problem has been addressed by limiting access through institution of a permit system. Some caves have been closed entirely and enforcement of the closure often requires the installation and maintenance of entrance gates which may interfere with access for bats.
A great many significant karst and cave communities are located on privately-owned land, or they are located in areas of state forest subject to logging operations, especially in Tasmania. Only in recent years, the forestry department of the Tasmanian government has played a leading role in conserving karst resources within production forests by developing a forest practices code which embraces karst. In addition, logging operations have been excluded from within some sensitive karst catchments, and operations have been deferred in other areas pending further investigations of their significance and vulnerability. For karstlands and cave communities under private ownership there is no legislative mechanism in place which specifically ensures their conservation, although recent developments pertaining to land clearance and catchment protection within New South Wales may prove to be relevant. Unfortunately, many karst communities on private land are already seriously degraded. The future of these communities depends to a large extent upon the attitude of the landholder, which varies between individuals.
Some caves and sinkholes continue to be utilized as rubbish dumps, whereas other landowners provide a high level of conservation management by excluding or limiting access to visitors. More often than not however, this action is based on sheer convenience and unsubstantiated fears of legal liability in case of a caving accident, rather than a genuine concern for conservation. The key to any conservation program is essentially one of enhanced understanding through education and interpretation at all levels, and insufficient attention has yet been given to the involvement of landowners as partners in conservation, particularly in respect to karst areas.
PUBLIC EDUCATION
It would appear that the future management and conservation of cave communities will rest both on a legislative footing and on better public recognition and understanding of the complexities of karst processes and karst environments (Eberhard & Spate 1995). The privatization of guiding services may result, although not invariably, to less emphasis on conservation and public education. However, collaboration between cavers, academic esearchers, and cave managers has proved to be a fertile starting point which has a positive flow-on effect to the general public. The development of a Minimum Impact Code for caving for example, has been adopted by some operators within the adventure tourism industry [Webb 1993). Another example is a pamphlet published by speleologists which has facilitated in raising awareness of the plight of cave dwelling bats in Victoria, whilst cooperation forged between the speleologists, a private landholder, and the state conservation department has enabled recovery of the Buchan population (Hamilton-Smith 1991; Friends of Buchan Caves 1993). Experience and expertise continues to evolve, which is transmitted through the auspices of organizations such as the Australian Speleological Federation and the Australasian Cave and Karst Management Association.
SUMMARY
Australia contains a great variety of karst landscapes and environments which are distributed across a broad range of climate types. Whilst much of the karst biota remains poorly known and described, there are subterranean communities of exceptional diversity and significance to zoogeographical and evolutionary studies. Unfortunately, some communities have become extinct, and a great many more have declined or become seriously degraded as a result of human activities. Karst biota continues to be threatened by deforestation, agriculture, quarrying, and cave visitors. Conservation strategies which have been applied include legislative protection of endangered species, land reserves, protection of sensitive habitats within individual caves, and public awareness campaigns. The current trend towards improved conservation and management of karst ecosystems depends upon a systemic approach to karst environments and processes, aided by further research and monitoring in combination with legislative protection of subterranean communities or taxa, and the fostering of public awareness.
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FOOTNOTE:
This paper is based upon preliminary material prepared for a chapter, of the same title, due to appear in Wilkens, Culver and Humphreys (eds.), Ecosystems of the World - Subterrenean Biota, to be published by Elsevier.