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CONSERVATION AND ENVIRONMENTAL MANAGEMENT OF SUBTERRANEAN BIOTA

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    ABSTRACT

    Research in caves is important, not only for instrinsicacademic reasons, but also for establishing manage-ment baselines, and for providing input into interpre-tative programs. Cave biota depend on outside foodsupply. Management of cave ecosystemsmust there-

    fore include protection of surface habitats. Cave eco-systems require an uninterrupted and unmodifiedtransport of organic matter through air, water andvisiting fauna. Movements of facultative cave ani-mals which are important food suppliers, especiallyat natural entrances, should be free of disturbance.An interdisciplinary approach is essential for study-ing caves and their biota, and for devising any suit-able conservation methods for management of natu-ral resources, which are unrenewable and unretriev-able. Subterranean and surface phenomena are inter-connected, the cave dwelling bats and swiftlets act-ing as intermediate fauna, linking the cave microeco-

    systems with the outside world.Management of caves for conservation should

    be based on maintaining ecological equilibria, propertechnical design, permit systems, and where neces-sary, on access restrictions

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    INTRODUCTION

    SPELEOLOGY, derived from SPELAEON, meaningCAVE and LOGOS, SCIENCE, is the science ofcaves and their environment. Their physical, biologi-cal and ecological aspects have been studied in depthonly in the past few decades. Speleology comprisesmany discrete fields of sciences, but avoids pigeon-holing, that may hamper an interdisciplinary ap-

    proach.According to the definition of the International

    Union of Speleology, a cave is a subterranean voidwhich can be entered by man. This definition impli-cates, that Speleology is a science, limiting itselfonly to underground spaces accessable to investiga-tors. This is very misleading, since caves enterable

    by man are only part of a much more complicatedsystem which exists in a matrix - in this case lime-stone consisting of innumerable cracks, fissuresand crevices, forming a labyrinth of interconnectedspaces, capable of accommodating some forms oflife.

    THE CAVE ENVIRONMENT

    INTERACTION WITH SURFACE CONDITIONSResearch methods of interdisciplinary sciences inSpeleology have awaken the importance of ecologi-cal interactions in the past 15 years. Hence concepts

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    developed in Speleology have been successful in de-vicing guidelines for environmental protection. For

    example, the concept of monitoring small changes inan uncomplicated microenvironment, such as a cave,may be useful in sounding a warning against poten-tially harmful broader changes at the surface, thatmight be masked by human intervention.

    Caves inhabitated by bats, swiftlets and otherfauna, which forage above ground, form a dynamic

    link with the surface. Fecal droppings and urine, dif-fering in quality and quantity, cause specific commu-nities to develop in the organic residue. These are ina state of dynamic equilibrium. A decline in numbersof these surface active fauna, due to disturbance oftheir above-ground environment, will have a directimpact on the cave fauna and flora, requiring adapta-tion to new dynamic equilibria or causing completedisruption, depending on the severity of disturbance.Water, air, silt, vegetation and animal remains, guanoare the means of interplay between the dark cave in-terior with the outside world. Without energy transferfrom the outside to the inside of caves, only a limited

    number of highly specialized cave fauna and floracan exist, maintaining a nearly closed ecological sys-tem in a darkest zones of caves. The energy requiredfor metabolism is then derived from minerals in thewall rock and sediment of cave floors. The basicfood cycle in such a cave condition, would dependon chemoautotrophic bacteria, serving as nutrient

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    material for cave dwelling animals with no inputfrom outside the cave.

    Scientists frequently encounter the close con-nection between outside phenomena and the interiorof caves, so that it is now a coined term, to explainthis as the surface-subsurface or subterranean inti-mate and dynamic interplay.

    The study of underground phenomena mayeven contribute to space biology, physiology and

    psychology. It might be possible, for instance, to re-cycle the human waste products that would accumu-late on a long space voyage, as a source of new nutri-ents. Study of the food cycles in caves may giveclues to possible ways of recycling waste products.

    The study of caves may provide a deeper in-sight into nature. Monitoring the delicate microenvi-ronment in caves may develop conservation methodsof the living world.

    CAVE ECOLOGY

    Till 1979, a space, deep below ground, in solid bed-rock, is considered the most sheltered environment,free from human disturbance. But a 1980, short sci-entific paper, published in France, challenges thisview of cave life. Juberthie, Delay and Bullion work-ing in the underground laboratory of the C.N.R.S.,reported cave limited fauna in cracks of rocks and

    screes situated just below the soil. They call this en-

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    vironment the Superficial Underground Compart-ment (S.U.C.). This is not only found in karst, but

    also in shales and sandstone. So the few, thinly scat-tered animals in the depths of caves are actually theadvance guard or frontier pushers of a much lessdeep-ranging underground community. Its discoveryhas profound implication for conservation of cavelife.

    To have an insight in the energy transfer, a

    study of the cave ecosystem and foodsupply is man-datory. Cave communities of specialized organismsliving in total darkness, towards which they haveadapted themselves, passes energy and usefulchemicals in a fairly organized way. The length offoodchains is limited by the energy input of the sys-tem.

    Cave communities depend mainly on the detri-tus from surface ecosystems. Organic material is alsoobtained from fecal matter of animals that periodi-cally look for food outside the cave, such as bats,cave rats, birds and crickets. In each case, bacterialdecomposers rely on a flow of organic material from

    sunlit areas, from plant remains in cave streams, or-ganic substances in cave silt, even from suspendedorganic matter in drip water. Thus the primary energysources exploited by cave organisms are surface de-rived organic material utilized by heterotrophic bac-teria, or minerals, broken down by autotrophic bacte-ria, the former being quantitatively the more impor-

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    tant factor. Decomposer bacteria in turn are eaten byprotozoans, which are utilized by aquatic cave dwell-

    ing flatworms, isopods, amphipods, which in turn arepreyed upon by larger aquatic fauna.Life in caves actually works on the same prin-

    ciples as in the highly complex outside world, butwith fewer variables in the ecologic equations. Cavestherefore provide ecologists with relatively simpli-fied, easily studied model ecosystems.

    The importance of cave streams carrying bitsof vegetation, has been emphasized by most Ameri-can biospeleologists. Organically rich mud aredumped onto the stram banks in caves during floods.Streams are particularly important in cave systems,developed in massive horizontally bedded limestone.

    Many such caves are found in Indonesia. Butwhen vertical joints and steeply inclining bedding

    planes are present as found in many continentalEurope and English karst regions, and also in themassive highland karst of Irian Jaya, the food supplyis usually carried into the caves by water tricklingdown fissures from the soil above. This is without

    doubt the source of food for fauna living in theS.U.C. It will vary in energy richness and chemicalcomposition with the type of the soil-cover fromwhich it comes. This in turn, partly owes its charac-ter to the surface vegetation and to human activitiesabove the caves. In some areas of Hawaii, wholecave communities have been wiped out in recent

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    times by forest clearance and crop cultivation.The routes along which energy travels in most

    ecosystems have been refined over many thousandsof years of evolutionary selection. Separate foodchains often meet and cross to form a network ofroutes along which energy flows through the commu-nity. Complex foodwebs allow continuous uninter-rupted flow of energy through the ecosystem, so thatif the population of one particular organism declines

    dangerously, its predators can switch to another foodsource, rather than starve.

    Every now and then, an established ecosystemmay be disupted by major pertubations such as cli-matic change, forest fires, human disturbance thatcause it to be thrown out of balance. If this happens,the main casualties will generally be the highly spe-cialized species, being slower to respond to a forcedchange in role than less specialized species, whichhave more variation in their gene pools and cantherefore adapt faster to changed circumstances.

    DISCUSSION

    Tropical cave biology is still in its infancy. The Brit-ish New Guinea Speleological Expedition of 1975was among the first European expeditions to studytropical biospeleology. Until recently, European andAmerican cave biologists believed, that only a very

    few creatures were adapted to tropical cave life, and

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    that cave evolved troglomorphic animals are onlypresent in significant numbers in caves of temperate

    latitudes. But deep in Selinum Tem, the largest cavefound in Papua New Guinea, cave biologists found acommunity of animals which consisted almost en-tirely of troglomorphic species, living in small flood-

    prone passages just above the phreatic level, whereslowly-draining flood waters deposit a harvest offood and where constant temperature and humidity

    prevail.Subsequent visits to the caves of Mulu

    (Sarawak, 1982) have revealed different, but equallyspecialized cave evolved communities in deep cavehabitats. Meanwhile Howarth (1973) begun to un-cover yet another specialized cave-evolved commu-nity in Hawaiis lava tubes. In most cave passages oflarge diameter, which contain big populations of batsor swiftlets and lots of guano, there are few, if any,troglomorphic species.

    It is often only in the remote flood-prone pas-sages, where most of the biospeleological interestlies! Chapman (1985) predicts with confidence, that

    many tropical karst areas, like the Gunung Sewukarst in Central and East Java, contains rich andcomplex troglomorphic faunas, rivaling those of

    New Guinea, Hawaii and Sarawak.The earliest description of cave animals from

    Indonesia was given by Jacobson E. (Febr., March,1911) who visited several caves in the Gunung Sewu

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    area, describing his experiences in collectiong animallife from subterranean water, obtaining among others

    some species of fish, crabs and prawns (1912; 513-516). The cave crabs were later examined by Ihle(1912; 177-182) and consist of the previously knownParathelpus convexa (de Man, 1879) a commonepigean species and a new species, described by Ihle:Sesarmoides jacobsoni (Ihle, 1912), collected at

    Ngingrong and Jomblang caves.The small white

    crabs reported by Waltham et al (1983; 90) frm un-derground water of Gunung Sewu could very well

    belong to this species. The prawns collected by Ja-cobson (1912) at Ngingrong cave consists of Macro-

    brachium lar (Fabricius, 1798) and Macrobrachiumpilimanus (De Man, 1879).

    The Gunung Sewu Cave Survey in 1982, ajoint British-Indonesian undertaking, sponsored bythe British Royal Geographical Society, collected afew subterranean prawns, repeated in 1983 by Williset al. These material were placed at the disposal ofHolthuis and proved to be a new species: Macro-

    brachium poeti, which are kept in collection of the

    Rijksmuseum van Natuurlijk Historie Leiden(RMNH Crust. D no. 35796 and D no. 35794). Theyare found in Luweng Jurangjero, about 100 m insidethe entrance from the main stream, from a static ca-nal, and from a percolation-fed pool with mud floor.Visits to caves at Tuban (Ngerong cave) by Buadi(1981), Segaranten (Cidolok caves) by Buadi and Su-

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    priatna (1981), Cigudeg (Sipahang cave) by Suyanto(1982), several caves at Nusakambangan isle and

    Gombong (Ratu and Lawa caves, Petruk cave) bySuyanto (1983) and some caves at Gombong andGunung Sewu by Notowinarno (1985) obtainedmany biospeleological specimen, some of which arestill waiting for determination, but none of them arethought to be troglobionts.

    The trend is for more and more species to be

    collected from tropical caves for identification, byfewer and fewer specialists capable of doing so. The

    prawns found by the English team in Gunung Sewu,for instance, had to be sent to Holland for identifica-tion. How many specimens lie idle on Indonesianshelves, waiting for specialists to identify them isanybodys guess.

    What is needed are detailed, systematic studiesof whole cave faunas and their relationship to thecave environment, not merely collection of a fewcave-evolved animals. A dead specimen is useless,unless it is accompanied by detailed description of itssurroundings, where and preferably how it lived; its

    behaviour, food, population and habitat characteris-tics. It is indeed impossible to record too much infor-mation.

    Bat guano maintains specialized cave ecosys-tems, known as guano ecosystems, which differsfrom one cave to the other, owing to the specializedcave organisms being specifically adapted to differ-

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    ent composition and granule size of bat dung . It istherefore essential to study the biochemistry of

    guano. Dung of insectivorous bats consists predomi-nantly of indigestible insect chitine, which breaksdown relatively slowly, and is therefore not an idealsubstrate for huge amounts of decomposers.

    Where frugivorous bats contribute to form softfruity dung, with plenty of carbohydrates, the guano

    becomes a very rich source of food, maintaining ex-

    tensive fauna of decomposers, scavengers and theirassociated predators. Guano of frugivorous bats ismainly decomposed by fungi; that of insectivorous

    bat guano is primarily attacked by bacteria.The relative proportion of the two types deter-

    mines the relative abundance of fungi and bacterialdecomposers and their respective micropredators. Inthe great Niah Cave in Northern Borneo, where 1million bats live with 4 million swiftlets (collocalia),the dung and corpses support an extraordinary rangeof life in which counts of 1 million arthropodes per 1m2 of dung are encountered, consisting of collembo-las, mites, millipedes, pseudoscorpions, beetles,

    cockroaches, isopods. In fermenting bat guano, at theNgerong river cave, we have counted about 100 verylarge cockroaches per square meter (Tuban cave ex-

    pedition, 1981).The data of cave fauna on given dates form

    bench marks for comparative studies to determinechanges in variety and quantity of these animals,

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    caused by detrimental factors to the ecosystem, likedeforestation, forest fires, quarrying. But as men-

    tioned before, the highly specialized fauna, found inthe deepest parts of the cave, in complete darkness,without temperature fluctuations, are the most sensi-tive to such changes.

    Individual species survive in an ecosystemonly as long as they have a place in their community.Damage to the ecosystem is repaired in an adjust-

    ment in the numbers of individual species, leading toa new level of dynamic equilibrium, where an effi-cient flow of energy through system is again reached.Human activities on the surface environment mayalready have deeply affected the S.U.C. community,even causing the local extinction of some cave ani-mals and collembola species.

    It is useless and uneconomic to protect indi-vidualrare species, if they are deprived of theire-cological niche. This is what popular conservationefforts have often done keeping spectacular ani-mals in captivity, while their habitat is destroyed. In-creasing public awareness and action to protect the

    whole ecosystem is fundamental to prevent speciesof flora and fauna becoming endangered or wipedout. Coneservation of bats, swiftlets and other imsecteating birds and rare cave dwelling animals meanconservation of their habitat. This permits completecave guano ecosystem conservation.

    The diversity of our natural environment

    should be preserved for future generations to enjoy.

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    It is a pity, that cave communities are low on the pri-ority list of most conservation programmes! Their

    man importance is as easily understood modal eco-systems, whose study can lead to a greater knowl-edge to manage our natural environment.

    Everyone appreciate that cave are habitats forbats and other insectivorous birds. Plant dispersal bybats, specializing e.g. in pollinating Mangrove treesor the act of chiropterochory, is of great importance

    in tropical forests. Nectivorous bats also play impor-tant roles in cross-pollination of these fruit plants.Many general of trees and shrubs, depend wholly ormainly upon these bats, which form an importantlinks in the complex interrelationships of a tropicalforest.

    Conflict of interest in karst areas, as shownthrough agricultural activities, quarrying of lime-stone, chalk production and cement factories in Indo-nesia, should be solved by a multidisciplinary, inter-governmental and environmental approach.

    Preservation of these unrenewable and unre-trievable habitats and ecosystems should beaccorded

    its full importance. Underground water resources inlimestone are also very important. Their replenish-ment depends on the management of rainwater catch-ments areas, including pattern of silvicultute.

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    RECOMMENDATION

    The Indonesian Federation of Speleological Activi-ties, as member of the International Union of Spe-leology, recommends to the Indonesian Governmenttrough a Panel Discussion (Karanganyar-1984),Seminar (Semarang-1984), First National Sympo-sium of Karst and Caves (Jakarta-1985), Second Na-tional Symposium of Karst and Caves (Jakarta-

    1996), Informal Meeting on karst silviculture(Yogyakarta-1985), and a Workshop on the Utiliza-tion of Caves (Bandung-1986), that :

    1. Data on karst and caves in Indonesia be collected,filed uniformly, computerized and reported to Scien-tifical Institutes, Departments (Ministries) and otherGovernmental Agencies, for further evaluation oftheir use and where possible, to promote their con-servation.

    2. The Indonesian public at large should be educatedto arouse their awareness of the Cultural, Physical,

    Biological and Ecological values of caves and theirenvironments.

    3. Plans to exploit karst areas for commercial rea-sons, must be assessed on a multidisciplinary andecological bases.

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    4. Management of biological resource in cavesshould be based on ecological principles. Biota of

    caves, fissure and crack systems, should be evaluatedand monitored. Availability of food resources(biotops) should be regularly monitored using cavecollembolae, pseudoscorpions and cave crickets at-fixed locations as indices of abundance. Bat popula-tion densities can be assessed at intervals, countingthese animals during their flight out of thecaves by

    means of serial photographs. Diversity of cave ani-mals, guano quality and quantity should also bemonitored. Should a cave contain outstanding

    biospeleological resources (such as great numbers ofbats and swiftlets, presence of blind troglobionticcave fishes and other cave troglomorphs), it should

    be reported to the Directorate General of NatureConservation, The Indonesian Scientific Institute,The Ministry of Population and Environment, TheMinistry of Interior and The Local Government.Such a cave should be closed to unauthorized indi-viduals and preferably gated. (A number of papersconcerning cave gating can be read from the pro-

    ceedings of the yearly symposia on cave manage-ment in the United States. The spaces between the

    bars should accommodate bats and swiftlets to flythrough and the structure should not impede with airflow. Access should be limited by requiring special

    permits or recommendations from above mentionedauthorities).

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    5. Should a cave, already frequented by tourists, con-

    tain these outstanding features in certain cave pas-sages, these locations should also be closed to public.It is therefore essential that a full assessment of cave

    biota, including their food chain(s) and their (micro)ecosystems should precede any attempt to opencaves for tourism and appropriate restrictions de-vised.

    6. Some caves should be recognized as undergroundlaboratories for the study of various aspects of biol-ogy, ecology, hydrology, sedimentology, paleontol-ogy and archaeology.

    Only by strictly adhering to these principles can weexpect that caves, their environment and their biota,and associated surface ecosystem can be protected,so that we and future generations in Indonesia get allthe benefit from these unique and outstandingly im-

    portant natural resources.

    REFERENCES1. Buadi and Supritana (1981) Catatan tentang kelelawar yangditemukan di dalam gua-gua di Cidolok, B.G.I. 2 (1) : 910.2. Buadi (1981) Biologi Gua Ngerong Laporan Ekspedisi keGua Ngerong : 21-243. Chapman P. (Nov. 1983) Cave Life part 9, Caves and Cav-ing, 2125.

    4. - (Sept. 1985) Cave Biology on Tropical Expeditions,

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    Caves and Caving: 2425.5. Christiansen K. and Bullion M., An Evolutionary and Eco-logical Analysis of the Terrestrial Arthropods of Caves in the

    Central Pyrenes, NSS Bulletin,40 : 103117.6. Ford T. D. and Cullingford C.H.D. (1976) The Science ofSpeleology, Academic Press : 362, 365, 397407.7. Holthuis L. B. (1984) Freshwater Prawns (Crustacea, Deca-poda, Natantia) from Subterranean Waters of the GunungSewu Area, Central Java, Indonesia, ZOOL. MED. Leiden 58(19) : 141148 ISSN 0024-0672.8. Howarth F. G. (1981), Non-Relictual Terrestrial Troglo-bites in the Tropical Hawaiian Caves. Proceedings 8thInt.Congress of Speleology (1) : 593.9. Juberthie C. and Delay B. (1981), Ecological and Biologi-cal Implication of the Existense of a Superficial Under-ground Compartment Proceedings 8thInt. Congress of Spe-leology : 203205.10. Ko R. K. T. (1981) Ekspedisi ke Gua Ngerong (Laporan)

    11. - (1984) Kehidupan Binatang Dalam guaLectureNotes.Finspac Basic and Advanced Courses.12. Moore W. G. (1978) Speleology, the Stufy of Caves,Zephyrus Press: 68, 7378, 8485, 111.13. Pack S. B. (1981) The Geological and Environmental Set-ting of Cave Faunal Evolution. Proceedings, 8th Int. Congressof Speleology: 501.14. Petty P. E. (1976) Subsurface Management as a Compo-

    nent of General Land Management. National Csve Mange-ment Symposium: 30-3315. Poulson T. L. and White W. B. (1969) The Cave Environ-ment. Science, 165 : 971981.16. Poulson T. L. (1975), Management of Biological Re-

    sources in Caves. National Cave Management Symposium,:4650.17. Stitt R. R. (1976) Human Impact on Caves. National Cave

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    Management: Symposium, 3650.18. Supriatna J. (1981) Biospeleologi Gua Selatan Cidolok.BGI (3) 1:14.

    19. Suyanto A. (1982), Catatan Binatang yang ditemukan didalam Gua Si Menteng BGI (4) : 57.20. - (1983), Biospeleologi Gua-Gua di Kabupaten Cilacapdan Kebumen.Laporan Survai ke Nusakambangan dan GuaPetruk:: 2740.21. Tuttle M. D. (1976) Gating as a Means of Protecting CaveDwelling Bats. National Cave Management Symposium: 7782.22. Waltham et al (1983) The Caves of Gunung Sewu, JavaCave Science 10 (2) : 5596.

    23. Yalden D. W. (1975), The Lives of Bat. A Demeter Press

    Book : 6668, 71.

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