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Records of the Western Australian Muswm Supplement No. 64: 37-47 (2001). Subterranean biodiversity in New South Wales: from rags to riches Mia E. Thurgatel', Jane S. Gough 2 , Andy Spate 2 and Stefan Eberhard 3 lJenolan Caves Reserve Trust, Locked Bag, Jenolan Caves, New South Wales 2790, Australia e-mail: [email protected] lNew South Wales National Parks and Wildlife Service, PO Box 2115, Queanbeyan, New South Wales 2620, Australia 3CaveWorks, PO Witchcliffe, Western Australia 6286, Australia *For correspondence Abstract - The karst areas of New South Wales, Australia, support a diverse invertebrate subterranean fauna with up to 422 taxa recognised to date, of which 83 are obligate subierranean taxa. The terrestrial fauna is dominated by arachnids and insects and includes 45 terrestrial obligate taxa. A rich stygofauna has also been identified that contains 38 stygobites, dominated by crustaceans. Five karst areas are particularly important as focal points for biodiversity. Jenolan, Wombeyan, Wee Jasper and Stockyard Creek contribute significantly to the total subterranean fauna, while Wombeyan and Wellington are exceptional for their stygobitic communities. High levels of richness and endemism, as well as relictual distributions and ancient lineages highlight the significance of the fauna. INTRODUCTION Recent research has demonstrated that some Australian karst areas support rich subterranean invertebrate faunas, particularly in the States of Tasmania, Queensland and Western Australia (Howarth and Stone, 1990; Ooran et al., 1997; Humphreys, 2000a). This paper focuses on the issue of subterranean biodiversity in another region of Australia, namely the karsts of New South Wales and the Australian Capital Territory (hereafter included with New South Wales). Hamilton-Smith (1967) compiled the first systematic review of the total Australian cavemicolous fauna and found that the karsts of New South Wales had a comparatively richer fauna than elsewhere on the continent. Gray's (1973) subsequent review of the Australian cave spider fauna added further to the tally provided by Hamilton-Smith. In all, these two papers listed 84 cavemicolous species from New South Wales. In the two decades following the publication of these reviews, new records and descriptions of subterranean invertebrates from New South Wales accumulated slowly and survey efforts lagged behind other areas in Australia. However, some significant increases in knowledge resulted from baseline surveys of particular karsts. For example, the results of intensive surveys at Jenolan Caves almost doubled the known subterranean diversity of the state (Gibian et. al., 1988; Greenslade, 1989). Sufficient information had been published by the early 1990s to indicate that a rich fauna would be uncovered by further surveys. However, this information was widely scattered, difficult to access, and had not been collated. Furthermore, a long-held preoccupation with obligate forms (of which relatively few had been discovered) did not allow an appreciation of the diversity of the entire spectrum of cavernicolous invertebrates. Consequently, a perception arose that the fauna of New South Wales was depauperate and insignificant in comparison to that being discovered elsewhere in Australia (Eberhard and Spate, 1995). In 1995 a reconnaissance survey was completed covering many karst areas in New South Wales (Eberhard and Spate, 1995). The results of this and more recent surveys have markedly increased the known diversity of the subterranean fauna. We use the metaphor 'from rags to riches' to highlight this finding and to dispel former erroneous perceptions of a depauperate fauna. The information provided in this paper is largely drawn from an analysis of these surveys, and many of the results are reported for the first time. Specifically, we present information regarding the composition, distribution and diversity of the fauna and highlight its significance. CHARACTERISTICS OF THE SUBTERRANEAN FAUNA The present knowledge of the subterranean faunas of New South Wales is far from complete owing to the limited survey and taxonomic effort. DOI: 10.18195/issn.0313-122x.64.2001.037-047

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Records of the Western Australian Muswm Supplement No. 64: 37-47 (2001).

Subterranean biodiversity in New South Wales: from rags to riches

Mia E. Thurgatel', Jane S. Gough2, Andy Spate2 and Stefan Eberhard3

lJenolan Caves Reserve Trust, Locked Bag, Jenolan Caves, New South Wales 2790, Australiae-mail: [email protected]

lNew South Wales National Parks and Wildlife Service, PO Box 2115, Queanbeyan, New South Wales 2620, Australia3CaveWorks, PO Witchcliffe, Western Australia 6286, Australia

*For correspondence

Abstract - The karst areas of New South Wales, Australia, support a diverseinvertebrate subterranean fauna with up to 422 taxa recognised to date, ofwhich 83 are obligate subierranean taxa. The terrestrial fauna is dominatedby arachnids and insects and includes 45 terrestrial obligate taxa. A richstygofauna has also been identified that contains 38 stygobites, dominated bycrustaceans. Five karst areas are particularly important as focal points forbiodiversity. Jenolan, Wombeyan, Wee Jasper and Stockyard Creek contributesignificantly to the total subterranean fauna, while Wombeyan andWellington are exceptional for their stygobitic communities. High levels ofrichness and endemism, as well as relictual distributions and ancient lineageshighlight the significance of the fauna.

INTRODUCTIONRecent research has demonstrated that some

Australian karst areas support rich subterraneaninvertebrate faunas, particularly in the States ofTasmania, Queensland and Western Australia(Howarth and Stone, 1990; Ooran et al., 1997;Humphreys, 2000a). This paper focuses on the issueof subterranean biodiversity in another region ofAustralia, namely the karsts of New South Walesand the Australian Capital Territory (hereafterincluded with New South Wales). Hamilton-Smith(1967) compiled the first systematic review of thetotal Australian cavemicolous fauna and found thatthe karsts of New South Wales had a comparativelyricher fauna than elsewhere on the continent. Gray's(1973) subsequent review of the Australian cavespider fauna added further to the tally provided byHamilton-Smith. In all, these two papers listed 84cavemicolous species from New South Wales.In the two decades following the publication of

these reviews, new records and descriptions ofsubterranean invertebrates from New South Walesaccumulated slowly and survey efforts laggedbehind other areas in Australia. However, somesignificant increases in knowledge resulted frombaseline surveys of particular karsts. For example,the results of intensive surveys at Jenolan Cavesalmost doubled the known subterranean diversityof the state (Gibian et. al., 1988; Greenslade, 1989).Sufficient information had been published by the

early 1990s to indicate that a rich fauna would be

uncovered by further surveys. However, thisinformation was widely scattered, difficult toaccess, and had not been collated. Furthermore, along-held preoccupation with obligate forms (ofwhich relatively few had been discovered) did notallow an appreciation of the diversity of the entirespectrum of cavernicolous invertebrates.Consequently, a perception arose that the fauna ofNew South Wales was depauperate andinsignificant in comparison to that being discoveredelsewhere in Australia (Eberhard and Spate, 1995).In 1995 a reconnaissance survey was completed

covering many karst areas in New South Wales(Eberhard and Spate, 1995). The results of this andmore recent surveys have markedly increased theknown diversity of the subterranean fauna. We usethe metaphor 'from rags to riches' to highlight thisfinding and to dispel former erroneous perceptionsof a depauperate fauna. The information providedin this paper is largely drawn from an analysis ofthese surveys, and many of the results are reportedfor the first time. Specifically, we presentinformation regarding the composition, distributionand diversity of the fauna and highlight itssignificance.

CHARACTERISTICS OF THESUBTERRANEAN FAUNA

The present knowledge of the subterraneanfaunas of New South Wales is far from completeowing to the limited survey and taxonomic effort.

DOI: 10.18195/issn.0313-122x.64.2001.037-047

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38 M.E. Thurgate, }.S. Gough, A. Spate, S. Eberhard

Currently many genera and most species remainundetermined. Most caves have been sampled onlyonce so data on annual, seasonal and longer-termvariations of population and community dynamicsis lacking. There are many distributional gaps in the

data as only 10% of caves, and 70% of cavernouskarst areas have been surveyed (Eberhard andSpate, 1995). Owing to the taxonomic limitations wehave adopted a conservative approach and treatmultiple specimens, not clearly assigned to a

- ----------------------J-Keyo Karst areas

Le_BiOdiversilylc>cal pointsNorthern NSW

ooStockyard Creek

O-mWillioYessabah

o

illi

oo oGloucester

~--4-- South Eastern Highlands

~"~"~

''0

<" N--~ W*E100 0 100 200 km

E J

S

oWellington Caves

o

Figure 1 Distribution of cavernous karst areas in New South Wales including localities mentioned in the text. Notethat the karst areas fall into two broad groups (northern and southern), with most being concentrated in andaround the South Eastern Highlands. Areas supporting a high subterranean invertebrate diversity arehighlighted.

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Subterranean biodiversity in New South Wales

particular species, as a single taxon. Hence, it islikely that the diversity of the fauna is far greaterthan is presented here.

CompositionInsects and arachnids predominate the karsts of

New South Wales accounting for three-quarters ofthe total fauna. The most diverse insect orders areColeoptera, Collembola, Diptera and Lepidoptera.The arachnid fauna is dominated by Araneae, butincludes Acarina, Opiliones and Pseudoscorpiones.Crustacea, Myriapoda and Gastropoda account fora further 21% of the fauna.

Terrestrial faunaInsects, arachnids, gastropods and myriapods

dominate the terrestrial fauna. They form distinctcommunities associated with particular habitatssuch as twilight and entrance zones, plant detritus,sediment banks, rock surfaces, tree roots and batguano (Smith, 1982; Eberhard and Spate, 1995;Eberhard, 1998). Little is known about the ecologyof these communities. Some specialised taxa arerestricted to a particular type of habitat (eg. tineidmoths in bat guano and fulgoroid planthoppers ontree roots), while others are generalists and occuracross a range of habitats.The faunas associated with bat guano deposits

were amongst the earliest to be collected andstudied, and consequently are better understoodthan most. The guano-based communities in NewSouth Wales typically include spiders,pseudoscorpions, mites, tineid moths, springtails,pscopterans and beetles (Eberhard and Spate, 1995;Hamilton-Smith and Eberhard, 2000). One of thefew Australian studies on the structure anddynamics of an invertebrate cave community(Harris, 1973) was based on guano communitiesfrom Carrai Bat Cave at Stockyard Creek. Thislocation (and all others mentioned in the text) isshown in Figure 1.

Aquatic faunaCrustaceans, especially amphipods and syncarids,

dominate the subterranean aquatic fauna. Other(minor) taxa include oligochaetes, mites, hydrobiidgastropods, paludicolan flatworms, and variousinsect orders. Distinctive communities areassociated with flowing waters, pools, edgeenvironments, and different substrate types such ascoarse sediments or tree root mats (Eberhard, 1993,1998; Thurgate and Gough, unpublished). Thecomposition of the subterranean aquatic fauna ofNew South Wales is discussed in greater detail inThurgate et al. (2001).

Obligate faunasThe New South Wales faunas display varying

39

degrees of dependency on subterraneanenvironments. We have adopted the approach ofHumphreys (2000b) to define dependency. Hencefor terrestrial faunas, troglobites are forms onlyfound in subterranean environments whiletroglophiles occur in both hypogean and epigeanenvironments. Both troglobites and troglophilesspend their entire lifecycle underground.Trogloxenes enter caves actively or passively, andnormally feed on the surface. Subterranean aquaticforms are classified as stygobites, stygophiles andstygoxenes using similar criteria.Some authors have used the degree of

morphological adaptation as an additional criterionto identify cave-dependent faunas (see Camacho,1992). Many terrestrial obligates found in earlierNew South Wales surveys (particularly the spidersand insects) display little or no adaptation. Thisprompted Hamilton-Smith (1967, 1971) to introducethe term second-level troglophile to distinguish "thoseforms living out their total life-cycle within caves,known only from caves, but not exhibiting anymodification to the cave environment" (Hamilton-Smith, 1971: 64). Use of this term detracts from thefact that these forms are an important component ofthe obligate fauna, so we do not distinguishbetween troglobites and second-level troglophiles inthis paper. However, as the term persists in theAustralian literature we indicate taxa previouslyidentified as second-level troglophiles in theAppendix.In contrast to the terrestrial obligate fauna, most

stygobionts are highly troglomorphic (lacking eyesand pigment). Many also have Pangaean andGondwanan affinities (see Thurgate et al., 2001) orare distributional and/or phylogenetically relicttaxa (Hamilton-Smith and Eberhard, 2000; Serov,personal communication, 2000; Wilson and Johnson,1999). This suggests that some karst regions haveacted as long-term refuges (sensu Humphreys,2000b) for stygofauna (Eberhard and Spate, 1995).The distribution of karst areas that contain obligatefaunas is shown in Figure 2.Based on current estimates, up to 20% of New

South Wales subterranean invertebrates are obligatespecies, while similar proportions are troglophilesand stygophiles. For the terrestrial faunas, mostobligate species are found in the Araneae (11species), Pseudoscorpiones (8 species) andColeoptera (6 species). Amongst the aquatic faunastygobites occur mostly in the Amphipoda (16species) and Anaspidaceae (12 species).

Distribution patternsThe karsts of New South Wales occur as a broad

band of scattered outcrops in the east of the State.Most are located in and around the South EasternHighlands, while a smaller cluster occurs innorthern New South Wales (Figure 1). The South

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40 M.E. Thurgate, J.S. Gough, A. Spate, s. Eberhard

Key to Fauna Recorded from Kar~t Areas I

1::,. StygobitesD Troglobites+ Troglobites and Stygobites

DAshford

+Timor

Stockyard CreekD Sulcor oD~illi illiKunderang Brook • U Y ssabah

DCo boyne

1::,.Wellington1::,. Bakers Swamp

Bowan Park .t:.D BorenoreCanomodine + fII Cliefden & Walli

r+JenolanAbercrombie J+ t L.EJ Tuglow

aun er +Colongwombeyan~Little W mbeyan Creek

ia

100 200 km2

o100i

+Talmo

weeJasp~e~Cooleman PlainYarrangobilly

SFigure 2 Distributions of karsts in New South Wales that contain obligate fauna. Sites marked 'troglobite' may include

a number of taxa that are considered to be second-level troglophiles. A full list of obligate taxa and theirdistribution is provided in the Appendix.

Eastern Highlands are characterised by cool, moistclimates, and these karst systems are impounded(surrounded by non-carbonate rocks) and of lowrelief. In contrast, the karsts of northern New SouthWales are characterised by a warm, seasonallymoist climate, and most occur as small, isolated

-------

outcrops often rising above the surroundinglandscape.The broad-scale distribution patterns of the fauna

reflect the characteristics of the karsts themselves.The disjunct nature of the karsts is reflected bysimilarly highly disjunct distribution patterns

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Subterranean biodiversity in New South Wales

amongst the subterranean fauna. Most of the faunais concentrated in the karsts of South EasternHighlands, which also contain a higher proportionof aquatic communities and obligate taxa.The northern and southern karsts each contain

distinctive faunal elements, and it has beensuggested that the gap between these areas mayhave acted as a barrier to dispersal (Hunt, 1992).For example, the genus Speotarus occurs only inthe north of the State whilst Notospeophonus isfound only in the south. Interestingly theMycetophilidae, represented by the single speciesArachnocampa richardsae Harrison, 1966, arerelatively common in abandoned mines, moistcanyons and gullies in the South EasternHighlands, and generally in rainforest environ-ments. Only in the north of the State are theyfound in caves and then only at Gloucester andStockyard Creek. Both areas contain caves withrunning streams and, unlike most of the State'skarsts, are beneath rainforest, so it is unlikely thatthe gap in karst distribution has been significant inthis instance.Most arachnids are scattered in disjunct

populations across the State, but there is evidenceto suggest a dispersal barrier has influenced someof this fauna. Colcarteria and Heteropoda andPholcidae are found only in northern karsts. Incontrast, Kaiya, Badumna and Australomimetus arepredominantly southern in distribution. Similarly,within the harvestmen, Megalopsalidae is broadlyscattered, but Triaenonychidae is mostly confinedto karsts in the South Eastern Highlands.The genus Holonuncia (Triaenonychidae) forms

a complex of 12 species in New South Wales(Hunt, 1992), of which six species occur only insurface environments. Three species are found inboth epigean and hypogean environments,however cave populations display minoradaptations and may be genetically isolated fromsurface congeners. Holonuncia sussa, H. dewae andH. weejasperensis are obligate cavernicoles (Hunt,1992). A similar distribution pattern may beevident amongst other arachnids (e.g. Laetesiaspp., Archaearanae and Pholcidae). Theseepigean/hypogean species-complexes may proveuseful in testing biogeographic and evolutionaryhypotheses about the Australian cave fauna(Eberhard, 1998).

EndemismCave communities in New South Wales possess a

very high degree of local endemicity as is typical ofmany karst areas worldwide (Culver, 1982). Apreliminary analysis by Thurgate et al. (in press)found that up to half of the total subterranean faunaof the State is restricted to a single karst area. Thereare also a number of centres of endemism, notablyWombeyan and Jenolan in the south and the

41

Macleay Valley, between Yessabah and StockyardCreek, in the north (Figure 1).Amongst the terrestrial fauna, SOme species are

endemic to a karst area but occur in several caves(e.g. Australotettix carraiensis, Trinemura anenome andProgradungula carraiensis). Amongst the beetle fauna11 of 25 families contain taxa that are localendemics. Some subterranean invertebrates have aneven more restricted distribution, being limited tosingle caves (e.g. Speotarus lucifugus princeps,Teraphis cavicola and Laetesia weburdi). Culver (1986)identifies single cave endemism as a featurecommon to dissected karsts such as those found inNew South Wales.Endemism is also extremely high in the aquatic

faunas, particularly amongst the amphipods andsyncarids. Of the 63 stygobionts, 27 are restricted tosingle cave systems. The crustacean fauna found atWombeyan is almost exclusively endemic to thatarea and most species are restricted to single caves(Thurgate et al., 2001).

DIVERSITY OF NEW SOUTHWALESSUBTERRANEAN FAUNA

The subterranean invertebrate fauna of NewSouth Wales includes 422 species from 269 generaand 166 families. Most of this richness isconcentrated in the terrestrial fauna (359 taxa),particularly amongst the insects and arachnids. Thedominant insect orders are Coleoptera, Collembolaand Diptera, while Araneae is the most diversearachnid order (Table 1). The aquatic communities

Table 1 Richness within taxonomic groups recordedfrom New South Wales caves. Valuesrepresent the minimum number of taxapresent based on available taxonomicinformation. Based on the New South WalesCave Invertebrate Database (Eberhard andSpate, 1995) and unpublished records.

Taxa Number of Number of Number offamilies genera species

ArachnidaAcarina 12 12 17Araneae 34 66 111Pseudoscorpiones 7 9 15Opiliones 2 3 14

InsectaColeoptera 25 42 61Collembola 9 34 39Diptera 14 15 20Other insect orders 26 28 36

Gastropoda 9 14 23Crustacea 14 28 45Myriapoda 3 5 19Minor groups 11 13 22

Totals 166 269 422

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42

are less diverse (63 taxa), and Amphipoda (20 taxa)and Anaspidacea (12 taxa) are the richest orders.At least 83 obligate taxa have been recorded from

30 karsts in New South Wales (see Appendix andFigure 3). This includes up to 45 terrestrial obligatesand 38 stygobites. Spiders (11 taxa), pseudo-scorpions (8 taxa) and beetles (6 taxa) are the richestgroups in this instance. Once again, the Amphipodaand Anaspidacea (16 and 12 taxa, respectively)dominate the aquatic fauna as these two groupsinclude most of the stygobites.The abundance of individuals is generally low in

both aquatic and terrestrial environments.Exceptions occur where there is a stable andabundant food supply, such as occurs in guanodeposits and in aquatic plant root mats. Forexample, Harris (1973) recorded very high numbers(up to 34 X 106/m2) of the guano mite, UrooboveIlacoprophila in association with bacteria, protozoans,nematodes, spiders and insects at Stockyard Creek.In most karst areas the stygobitic fauna is cryptic

and in low abundances, but at Wellington,syncarids, amphipods and isopods are found inhigh densities and can easily be observed (Spate etal., 1999). Similarly, caves at Wombeyan supportrich and abundant crustacean communities whereroot mats are present (S. Reilly, personalcommunication, 2000).Comparisons with Tasmanian karsts provide a

temperate-zone context for examining the richnessof the· New South Wales' fauna, particularly asTasmania is already recognised for its highsubterranean biodiversity (Clarke, 1997; Doran etal., 1997). Within this context, it is interesting to findthat the diversity of the fauna in New South Walesis comparable with that of Tasmania, particularly athigher taxonomic levels and for stygobitic fauna(Table 2).Such comparisons need to be treated with caution.

The setting, nature and extent of each state's karstareas and their environmental history are verydifferent, as is the associated fauna. However, it isnow apparent that New South Wales contributes asignificant component of Australia's totalsubterranean biodiversity, and that the levels of

Table 2 Taxonomic and ecological richness of thesubterranean invertebrate fauna of New SouthWales compared to Tasmania.

Variable NSW! Tas2

Number of families 166 179Number of genera 269 271Number of species (taxa) 422 643Number of terrestrial obligates 45 108Number of stygobites 38 38

I Eberhard and Spate (1995) and unpublished records to April 2000.20arke (1997) and personal communication.

M.E. Thurgate, J.S. Gough, A. Spate, S. Eberhard

taxonomic and ecological richness are much higherthan has been appreciated previously.

FOCAL POINTS OF BIODIVERSITYIn a worldwide review, Culver and Sket (2000:

11) have used "an arbitrary cutoff of 20 or moreobligate subterranean species for a site" to beidentified as a biodiversity 'hotspot'. No single sitein New South Wales is known to meet this criterion.However, Wombeyan must be considered acontender for this title with 19 confirmed obligatetaxa, and a further seven that may fall into thiscategory (see Appendix).The use of a single numerical measure based on

obligate forms is problematic as this reveals littleabout overall diversity and community dynamics.There are also problems of scale and of cavedefinition that confound the use of such arbitrarymeasures, problems readily apparent in Culverand Sket's review. Given our present knowledge,and without putting any numeric thresholds inplace, several areas in New South Wales appear tostand out as focal points for subterraneanbiodiversity.In the broader geographical context, and when

the total subterranean fauna is considered, karstareas in the South Eastern Highlands areparticularly diverse. Rich communities of bothterrestrial (notably Holonuncia, Collembola,Carabidae and Chilopoda) and aquatic fauna(Amphipoda and Syncarida) are present. Thenorthern karsts are also rich in terrestrial faunaincluding Gastropoda (Charopidae), Araneae(Pholcidae and Heteropodidae) and Hemiptera. Theavailable evidence suggests that aquaticcommunities of the northern karsts are less speciesrich than those found in the South EasternHighlands.The overall richness (including obligate forms)

and numbers of local endemics of some individualkarst areas is quite high. The richest sites areJenolan (126 taxa, 11 obligate and 30 endemic),Wombeyan (83 taxa, 19 obligate and 25 endemic),Wee Jasper (62 taxa, 11 obligate and 13 endemic)and Stockyard Creek (55 taxa, 6 obligate and 9endemic). Collectively these sites support more thanhalf of the State's subterranean fauna. Most otherkarsts contain less than 30 taxa and have fewendemics or obligate forms.The higher richness at Jenolan, Wombeyan, Wee

Jasper and Stockyard Creek may reflect the greatersurvey efforts in these areas. However, the largesize, greater relative reliet and highly cavernousstructure at the first three locations may alsoinfluence richness by providing a wide range ofhabitats for colonisation. It has also been suggestedthat these karsts have been maintained as moistrefuges over geologically significant ~e periods,

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Subterranean biodiversity in New South Wales

thus facilitating the survival of relictual populations(Eberhard and Spate, 1995).Most other New South Wales karst areas have a

lower environmental buffering capacity owing totheir smaller size, lower relief, fewer and smallercaves, or fewer habitats. Pleistocene arid periodsmay have completely desiccated smaller karsts thuseliminating the pre-existing subterraneaninvertebrates (Eberhard and Spate, 1995). This mayhave been particularly true for terrestrial faunasthat have been more exposed to the effects ofclimate change than their aquatic counterparts.Caves at Wombeyan contain diverse subterranean

communities, and overall this location can beconsidered a focus of stygobitic diversity. At least25 groundwater taxa have been collected fromWombeyan of which up to 15 taxa are stygobites.Of particular interest is the high diversity ofamphipods, of which there are at least ten species(Eberhard and Spate, 1995) representing half of theknown stygobitic amphipod diversity recorded forthe State.A number of karst areas are notable for species

richness within a single order. Eberhard (1993)identified ten myriapods from one cave during asingle survey at Deua, and also found ninedifferent beetles at Willi Willi. Intensive surveyand taxonomic efforts at Jenolan have alsorevealed a particularly rich collembolan fauna of34 taxa from 14 caves. This includes taxa fromNeanuridae and Neelidae, which have not beenrecorded in other New South Wales caves(Greenslade, in press).

CONCLUSIONSThis paper clearly demonstrates that the

subterranean biodiversity of New South Wales hasprogressed from 'rags to riches' with the recognitionof the rich and diverse fauna. Clearly, the fauna ofNew South Wales makes a significant contributionto the subterranean biodiversity of Australia. Manynew families, genera and species have beendiscovered in the last five years alone. Overall,levels of endemism amongst the subterranean faunaare very high and many taxa show relictual anddisjunct distribution patterns. Some of the fauna hasPangaean and Gondwanan affinities and will beimportant in testing phylogenetic and evolutionarytheories. The high diversity of stygofauna,previously unsuspected in New South Wales, is alsoan important discovery (Eberhard and Spate, 1995).The information presented here is a preliminary

account of the significance of the State's fauna, andthere is scope for much important research in thefuture. At the present time, we have species lists fora few selected caves in many scattered karst areas,with many have yet to be investigated. Little isknown of the phylogeny or ecology of the fauna

43

that has been collected, and there are manytaxonomic groups yet to be described. This lag intaxonomic research is of particular concern and it isvitally important that funding be directed to thisarea if a full appreciation of the subterranean faunais to be gained.The future conservation and management of karst

ecosystems in New South Wales must rest on afoundation of better public understanding, aided byfurther research and monitoring, in combinationwith legislative recognition of rare and threatenedcommunities and species. To date, efforts in theseareas have mainly focussed on obligate faunas.While we acknowledge that these are an importantcomponent, the identification of these faunas isproblematic, particularly if morphological criteriaare used. We advocate the adoption of a wholeecosystem approach for recognising and protectingthe diversity of the State's subterraneaninvertebrates.

ACKNO~EDGEMENTS

We wish to thank Bill Humphreys for organisingthe symposium at which this paper was presented.Buz Wilson, Peter Serov, Arthur Clarke, JohnBradbury, Winston Ponder, Mike Gray and BarryMoore provided invaluable information that hasbeen incorporated into this paper. We thankAlexander Herr for comments on early drafts of themanuscript, and our reviewers whose commentsgreatly enhanced the final paper. We alsoacknowledge the assistance of numerous cavers,divers, land managers and taxonomists withoutwhom the New South Wales cave invertebratedatabase would not exist.

REFERENCESCamacho, AI. (1992). A classification of the aquatic and

terrestrial subterranean environment and theirassociated fauna. In AI. Camacho (ed.), The NaturalHistory of Biospeleology: 57-103. Monografias MuseoNacional de Ciencias Naturales, Madrid.

Clarke, A (1997). Management Prescriptions for Tasmania'sCave Fauna. Report to the Tasmanian RFAEnvironment and Heritage Technical Committee,Hobart.

Culver, D.e. (1982). Cave Life: Evolution and Ecology.Harvard University Press, Massachusetts.

Culver, D.e. (1986). Cave faunas. In M.E. Soule (ed),Conservation Biology: The Science of Scarcity andDiversity: 427-423. Sinauer Associates, Massachusetts.

Culver, D.e. and Sket, B. (2000). Hotspots ofsubterranean biodiversity in caves and wells. JournalofCave and Karst Studies 62: 11-17.

Doran, N.E., Eberhard, S.M., Richardson, AM.M. andSwain, R. (1997). Invertebrate biodiversity andconservation in Tasmanian caves. Memoirs of theMuseum ofVictoria 56: 649-653.

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Eberhard, S. (1993). Survey ofFauna and Human Impacts inthe Jenolan Caves Reserve. Report to Jenolan CavesReserve Trust, Jenolan Caves.

Eberhard, S. (1997). Cave fauna of New South Wales:nomination of key threatening processes, endangeredecological communities, endangered populations andthreatened species for inclusion on the Threatened SpeciesConservation Act 1995. Report to Scientific Committee,New South Wales National Parks and WildlifeService, Queanbeyan.

Eberhard, S. (1998). Cave invertebrates. In J. Bauer and P.Bauer (eds), Under Bungonia: 74-83. J.B. Books, OakFlats.

Eberhard, S. and Spate, A. (1995). Cave InvertebrateSurvey: Towards an Atlas ofNSW Cave Fauna. Report toDepartment of Urban Affairs and Planning and theAustralian Heritage Commission, Canberra.

Gibian, M., Smith, G. and Wheeler, L. (1988). InterimReport on the Survey of the Invertebrate Fauna of JenolanCaves. Report to Jenolan Caves Reserve Trust, JenolanCaves.

Gray, M.R. (1973). Survey of the spider fauna ofAustralian caves. Helictite 11: 47-75.

Greenslade, P. (1989). Collembola from Jenolan Caves.Report to New South Wales National Parks Service,Canberra.

Greenslade, P. (in press). Diversity and latitudinalgradients in composition of Australian cavecollembolan faunas including notes on some effectsof disturbance. Pedobiologia.

Hamilton-Smith, E. (1967). The arthropoda of Australiancaves. Journal of the Australian Entomological Society 6:103-118.

Hamilton-Smith, E. (1971). The classification ofcavernicoles. National Speleological Society Bulletin 33:63-66.

Hamilton-Smith, E. and Eberhard, S. (2000). Conserv-ation of cave communities in Australia. In H.Wilkens, D.e. Culver and W.F. Humphreys (eds),Ecosystems of the World, Volume 30: SubterraneanEcosystems: 647-664. Elsevier, Amsterdam.

Harris, J.A (1973). Structure and dynamics of a cavepopulation of the guano mite Uroobovella coprophila(Womersley). Australian Journal ofZoology 21: 239-275.

M.E. Thurgate, J.S. Gough, A. Spate, S. Eberhard

Howarth, F.G. and Stone, F.D. (1990). Elevated carbondioxide levels in Bayliss Cave, Australia: implicationsfor the evolution of obligate cave species. PacificScience 44: 207-218.

Humphreys, W.F. (2000a). The hypogean fauna of theCape Range peninsula and Barrow Island,northwestern Australia. In H. Wilkens, D.e. Culverand W.F. Humphreys (eds), Ecosystems of the World,Volume 30: Subterranean Ecosystems: 581-601. Elsevier,Amsterdam.

Humphreys, W.F. (2000b). Background and glossary. InH. Wilkens, D.e. Culver and W.F. Humphreys (eds),Ecosystems of the World, Volume 30. SubterraneanEcosystems: 3-14. Elsevier, Amsterdam.

Hunt, G.S. (1992). Revision of the genus HolonunciaForster (Arachnida: Opiliones: Triaenonychidae) withdescription of cavernicolous and epigean speciesfrom eastern Australia. Records of the AustralianMuseum 44: 135-163.

Smith, G. (1982). Broad similarities in the invertebratefauna in NSW caves. Highland Caving Group Journal 3:26-30.

Spate, A, Gough, J. and Thurgate, M. (1999). Karsticgroundwater ecosystems in the Murray Darling andOtway Groundwater Basins. Proceedings of the MurrayDarling Basin Groundwater Workshop 1999: 70-78.Murray Darling Basin Commission, Griffith.

Thurgate, M.E., Gough, J.S., Clarke, AK., Serov, P. andSpate, A. (2001). Stygofauna diversity anddistribution in Eastern Australian cave and karstareas. Records of the Western Australian Museum,Supplement No. 64: 49-62.

Thurgate, M.E., Spate, A. and Herr, A. (in press).Diversity at depth: the New South Wales caveinvertebrate story. Helictite.

Wilson, G.D.F. and Johnson R.T. (1999). Ancientendemism among freshwater isopods (Crustacea,Phreatoicidea). In W. Ponder and D. Lunney (eds),The Other 99%. The Conservation and Biodiversity ofInvertebrates: 264-268. Royal Zoological Society ofNew SouthWales, Mosman.

Manuscript received 17May 2000, accepted 17March 2001.

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Subterranean biodiversity in New South Wales

Appendix: Obligate Subterranea~Invertebrate Fauna of New South Wales.

45

The following list identifies those taxa that are obligatesubterranean species. These have been subdividedaccording to the classifications used by Eberhard andSpate (1995) as indicated by the following codes: Sb =stygobite; Tb = troglobite; Tp2 = second-level troglophile.Where codes show (?) this classification is probable butas yet unconfirmed. The karst areas are shown by lettercode at the end of each entry. Area codes are as follows:A =Abercrombie; AS =Ashford; B =Bungonia; BN =Borenore; BP =Bowan Park; BS =Bakers Swamp; CB =

Comboyne; CG = Colong; CL = Cliefden; CN =Canomodine; CP =Cooleman Plain; DE =Deua; 15L =Talmo; J =Jenolan; JA =Jaunter; KB =Kunderang Brook;LW =Little Wombeyan Creek; MA =Marble Arch; S =Sulcor; SC =Stockyard Creek; T =Tuglow; TR =Timor;W =Wombeyan; WA =Walli; WE =Wellington; WJ =Wee Jasper; WW =Willi Willi; WY =Wyanbene; Y =Yarrangobilly; YE = Yessabah. Based on the New SouthWales Cave Invertebrate database (Eberhard and Spate,1995) and unpublished records.

Phylum PlatyhelminthesClass TurbellariaOrder Tricladida

Family, genus and species undetermined

Phylum NemertinaFamily, genus and species undetermined

Sb? CP;J;W

Sb? W

F. Hydrobiidae

Phylum MolluscaClass Gastropoda

Austropyrgus sp. novoAustropyrgus sp. 1

Sb A;B;LW;WSb? W

r

F. CtenidaeF. GradungulidaeF. Linyphiidae

F. MicropholocommatidaeF. StiphidiidaeF. SynotaxidaeF. TextricellidaeF. Theridiidae

F. AtemnidaeF. Cheliferidae

F. Chthoniidae

Triaenonychidae

F. UrodinchychidaeF. Laelapidae

F. Rosensteiniidae

Phylum ChelicerataClass ArachnidaOrder Araneae

gen. novo (near Janusia) sp. novo TbProgradungula carraiensis Forster and Gray, 1979 Tp2gen. novo sp. novo TbLaetesia weburdi (Urquhart, 1889) Tp2gen. novo (near Laetesia) sp. novo TbChnsmocephnlon sp. 1 TbProcambridgea cavernicola Forster, 1973 Tp2gen. novo sp. novo TbTextricella sp. novo Tb?Icona sp. novo 1 Tp2Icona sp. novo 2 Tbkonasp.nov.3 Tb

Order PseudoscorpionesOratemnus cavernicola Beier, 1976 Tb?Protochelifer cavernarum Beier, 1968 Tp2Protochelifer sp. near cavernarum Tp2Pseudotyrannochthonius jonesi Chamberlin, 1962 TbPseudotyrannochthonius sp. TbSathrochthonius tuena Chamberlin, 1962 TbSathrochthonius sp. near tuena TbTyranochthonius cavicola Beier, 1962 TbTyranochthonius sp. near cavicola Tb

Order OpilionesHolonunda cavernicola Forster, 1955 Tp2H. dewae Hunt, 1992 TbH. sussa Hunt, 1992 TbH. weejasperensis Hunt, 1992 TbHolonuncia sp. novo Tb?

Order AcarinaUroobovella coprophilaWomersley, 1960 Tp2Cosmolaelaps sp. Tp2Hypoaspis sp. Tp2Nycteriglypus dewae Womersley, 1963 Tp2

KBSCJ;W;YJYCNW;WJ;WYMA;WYCPB; CG; MA; T; W; J; SCCLY

TibooburraAS;CL;TRA; BN; S; SC; TR; W; WA; WJ; YEunknown cave, Blue MountainsSCDE; J; T; W; WJTB15L

J; TR; TWCGWJTR, CL, A, WY

CB;SC;W;WJWJCBW

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Phylum CrustaceaClass Ostracoda

Family, genus and species undetermined

Class CopepodaFamily, genus and species undetermined (sp. 1)Family, genus and species undetermined (sp. 2)

M.E. Thurgate, J.S. Gough, A. Spate, S. Eberhard

Sb? J; WJ

Sb? CP; J; W; WJSb? J

Familynov.

F. Psammaspididae

F. Neoniphargidae

F. Janiridae

F. Styloniscidae

Family Phreatoicidae

Class MalacostracaDivision SyncaridaOrder Anaspidacea

Gen. novo 1 sp. novo 1Gen.nov. 1 sp.nov. 2Psammaspides sp. novo 1Psammaspides sp. novo 2Psammaspides sp. novo 3Psammaspides sp. novo 4Psammaspides sp. novo 5Psammaspides sp. novo 6Psammaspides sp. novo 7Psammaspides sp. novo 8Psammaspides sp. novo 9Psammaspides sp. novo 10

Division PericaridaOrder Amphipoda

Jasptorus solepti Bradbury and Williams, 1997Neocrypta georginae Bradbury and Williams, 1997N. moniae Bradbury and Williams, 1997N. primaris Bradbury and Williams, 1997N. robinae Bradbury and Williams, 1997N. simoni Bradbury and Williams, 1997Neoniphargus richardi Bradbury and Williams, 1997N. secus Bradbury and Williams, 1997Wombeyanus botulosus Bradbury and Williams, 1997Gen.nov.1 sp.nov.1Gen.nov.1sp.nov.2Gen.nov.1sp.nov.3Gen.nov. 1 sp.nov.4Gen.nov.2sp.nov.Gen. novo 3 sp. novoGen. novo 4 sp. novoGenus and species undetermined

Order IsopodaSuborder Asellota

?Heterias sp.

Suborder OniscideaGenus and species undetermined

Suborder PhreatoicideaCrenoicus sp. novo

Sb TRSb WWSb BSSb BPSb CLSb CPSb JSb I5LSb WJSb WESb WSb WY

Sb WJSb ASb BPSb WESb CNSb JSb WSb WSb WSb WSb WSb CGSb TRSb WSb WSb WSb? J;W

Sb CP;WE

Tb? B; CP; J; JA; T; W

Sb J; WJ

Phylum UniramiaClass SymphylaOrder Syrnphyla

Family, genus and species undetermined Tb? B; CP; Y; W

F. Entomobryidae

F.OncopoduridaeF. SminthuridaeF. Neanuridae

Class InsectaOrder Collembola

Coecobrya sp. near hoeftiSinella coeca groupOncopodura sp.Adelphoderia sp.Kenyura sp.

Tb? JTb JTb? JTb CG; CL; J; T; W; WJ; YTb J

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Subterranean biodiversity in New South Wales 47

Order ThysanuraF. Nicoletiidae Trinemura anemone Smith, 1988 Tb B

Order ColeapteraF. Carabidae Notospeophonus jasperensis ssp. Tp2 A

N. jasperensis jasperensis Moare, 1964 Tp2 WJN. jasperensis vicinus Maare, 1964 Tp2 BSpeotarns lucifugus lucifugus Moore, 1964 Tp2 ASS. lucifugus princeps Maare, 1964 Tp2 WWTeraphis cavicola Moare, 1977 Tp2 Y

0rder DipteraF. Cypselosomatidae Cypselosoma australis McAlpine, 1966 Tp2 SC