Helictite, (2012) 41: 75-85 Preliminary notes on the Cavernicolous Fauna of Judbarra / Gregory Karst Area, northern Timothy Moulds1,3* and Peter Bannink2 1 The University of Adelaide, Environmental Biology Department, Evolution and Ecology Laboratory, North Tce, Adelaide, South Australia, 5001. 2 Centre for Tropical Agriculture, DERM, PO Box 156, Mareeba, , 4008. 3 now at GHD, Terrestrial and Marine Ecology, 239 Adelaide Tce, Perth, Western Australia, 6004. *Corresponding author [email protected]

Abstract The Judbarra / Gregory Karst Region is situated in the Judbarra / Gregory National Park, west of Timber Creek, Northern Territory. Several large joint controlled maze caves occur within the area, developed within and below a prominent dolomitic layer (the Supplejack Member). The caves are predominantly shallow in depth (< 15 m below the surface) but are occasionally developed deeper as multi-level systems, reaching the aquifer. Two biological surveys from the largest caves have revealed 56 morphospecies from 43 families, 19 orders, and 7 classes. All collecting was undertaken in the northern dry season (April to September) and consisted predominantly of opportunistic collecting. The diversity of invertebrates collected from the Judbarra / Gregory karst comprised non-troglobionts (48 species, 86%), troglobionts (5 species, 9%), stygobionts (2 species, 3%), and trogloxenes (1 species, 2%). Five of the species are considered to be potential troglobionts, and two potential stygobionts as indicated by troglomorphisms such as elongate appendages and reduced or absent eyes. The five troglobiont species are an isopod (Platyarthridae: Trichorhina sp.), a scorpion (Buthidae: Lychas? sp. nov.), a pseudoscorpion (Geogarypidae: Geogarypus sp. nov.), a millipede (Polydesmida: sp.), and a planthopper (Meenoplidae: sp.). The two stygobiont species are a hydrobiid snail (Hydrobiidae: sp.), and an amphipod (Amphipoda: sp.). The troglobiont scorpion is only the second collected from a cave environment from continental Australia. Keywords: Biospeleology; troglobiont; stygobiont; scorpion; tropical.

Introduction Kimberley, including tainisopid and phreatoicid isopods, and atyid (Page et al., 2008; Humphreys, 2008). Australia’s numerous tropical karst areas (Webb et al., 2003; Figure 1) have been the subject of biological There are few studies of cave fauna in the Northern studies over the past few decades, with the number of Territory, with previous research incorporating arthropod studies increasing during the last decade, mainly within diversity within broader management plans for Cutta the calcrete and alluvial aquifers of the Pilbara and Cutta and Kintore Cave Reserves near Katherine Yilgarn regions of Western Australia (Guzik et al., 2008; (Hamilton-Smith et al., 1974; Hamilton-Smith et al., Humphreys, 2008). Among the most notable studies 1989; Parks and Wildlife Commission of the Northern on the invertebrate diversity and evolution in tropical Territory 2000); a taxonomic study (Dumont & Maas, Australia is the extensive work on the Cape Range of 1985) describes new species endemic to Cutta Cutta Western Australia (Harvey et al., 1993; Humphreys, Cave and the immediate karst area, but does not discuss 1991; Humphreys, 1993; Humphreys, 2000; Humphreys the Tindall karst in its entirety. A more recent study, & Shear, 1993) and the Undara lava tubes and Chillagoe investigating the stygofauna of the Ngalia Basin calcrete tower karst in northern Queensland (Roth, 1988; Roth, aquifers, west of Alice Springs in the Northern Territory, 1990; Hoch & Howarth, 1989a; Hoch & Howarth, 1989b; revealed the presence of stygobitic dytiscid diving Howarth & Stone, 1990), with both these areas found to , isopods and bathynellids (Balke et al., 2004; be rich in subterranean-adapted invertebrates including Taiti & Humphreys, 2001; Watts & Humphreys, 2006; numerous endemic species. Other karst areas receiving Cho et al., 2006). biological study include the Ningbing and Jeremiah Hills near Kununurra (Humphreys, 1995), Barrow Island Here we report on the invertebrate fauna of (Hoffman, 1994; Humphreys, 2000, 2002), and Pilbara Bullita Cave in the Northern Territory, situated in the aquifers (Karanovic, 2006). A new and several Judbarra / Gregory National Park, west of Timber Creek. species of bathynellids have been described from the far The cave is a shallow, joint controlled maze cave, situated eastern Kimberley near Lake Argyle and surrounding in a monsoonal climate, experiencing dry winters and areas (Cho et al., 2005). Stygofauna from the alluvial wet summers (Martini & Grimes, 2012). Bullita Cave aquifers associated with the Ord River irrigation scheme and the karst of the Judbarra / Gregory National Park have also been sampled (Humphreys, 1999). Some were first discovered by Europeans and explored in 1990 groups of stygofauna are notably absent from the (Storm & Smith, 1991) and exploration has continued

© The Authors, 2012. Journal compilation © Australian Speleological Federation Inc., 2012 Helictite, 41, 2012. 75 Grimes: Sandstone karsts

Figure 1: Major karst and basalt areas in northern Australia. Major carbonate aquifer basins are shown in light grey throughout much of the intervening 20 years (Kershaw, passages, sediment banks, near sumps, and bat guano 2012), although they were known to the Aboriginal deposits. Aquatic sampling sites included gour pools and people of the area previously. The discovery of permanent sumps. Judbarra / Gregory Karst Area presented an opportunity to investigate the subterranean fauna of a new and totally Specimens were assigned categories of cave unexplored karst area in tropical Australia. dependence based upon degree of morphological adaptation to the cave environment (troglomorphisms). Initially cave invertebrates were collected during expeditions held between 1991 and 1994 by members of All material has been lodged with the Northern the Top End Speleological Society (TESS) and Canberra Territory Museum of Arts and Sciences (NTMAS), and Speleological Society (CSS). Additional sampling was the Western Australian Museum (WAM). undertaken in 2006 by Timothy Moulds as a dedicated Temperature and humidity were recorded at the biospeleological survey of the karst region, including collection sites in BAA 4, BAA 35 and BAA 38, using a both the Central Karst Area (Bullita Cave) and the Dingo whirling hygrometer. Block (Northern Karst Area), situated to the north of Limestone Creek (Figure 2). Results Methods The combined surveys for the Judbarra / Gregory karst area have recorded 56 morphospecies from 43 families, Invertebrate Sampling 19 orders, and 7 classes (Table 1). The species collected are predominantly non-troglobionts (48 species, 86%), Opportunistic sampling was undertaken between 1991 troglobionts (5 species, 9%), stygobionts (2 species, 3%), and 1994 and a fauna survey in 2006, all undertaken in and trogloxenes (1 species, 2%) (Figure 3). Specimens the northern dry season (April to September). Collecting that showed obvious troglomorphisms were classified was undertaken in three different cave systems within as troglobionts or stygobionts, while other species the Judbarra / Gregory Karst area: the Central karst area were assigned non-troglobiont status as the distinction (BAA 22 and BAA 35); the Dingo System (BAA 38) between accidental and troglophile was unable to be located to the north of Limestone Gorge; and BAA 4, a determined without extensive surface collecting. limestone block located to the east of the Dingo Block (Figure 2). Specimens were collected using forceps, Five of the species are potentially troglobionts, and aspirators and wet paint brushes, and aquatic fauna two are potentially stygobionts due to troglomorphisms were collected using hand held dip nets. Sampling such as elongate appendages and reduced or absent eyes. was undertaken at multiple locations and microhabitats The five troglobiont species are an isopod (Platyarthridae: in each cave to provide a broad representation of the Trichorhina sp.), a scorpion (Buthidae: Lychas? sp. nov.), invertebrates present and to enable some comparison a pseudoscorpion (Geogarypidae: Geogarypus sp. nov.), of diversity, community structure and ecology between a millipede (Polydesmida: sp.), and planthopper nymphs caves and ultimately with other karst areas. Cave micro- (Meenoplidae: sp.). The Trichorhina isopod, polydesmid habitats investigated included cave entrance areas, cave millipede and planthopper nymphs are white to a very

76 Helictite, 41, 2012. Judbarra / Gregory Karst

Figure 2: Location of cave invertebrate sampling sites

Helictite, 41, 2012. 77 Moulds & Bannink: Cave Biology MAGNT * MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT Museum WAM WAM MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT MAGNT * MAGNT MAGNT MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT MAGNT MAGNT MAGNT MAGNT MAGNT MAGNT MAGNT MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT Acquisition / ID Number BAA35:12 BAA27:02 BAA27:34 BAA27:19 BAA27:03 BAA35:19 BAA27:64 T82731 T80206 BAA04:10 BAA27:14 BAA27:72 BAA27:35 A000332 BAA04:16 A000473 BAA35:01 BAA35:15 BAA27:14 BAA27:07 BAA27:44 BAA27:36 BAA27:01 BAA27:04 BAA27:66 BAA27:11 P037764 P007834 P007839 P007835 P007840 P007840 P007836 P007837 P007842 P007841 BAA04:03 BAA27:41 P007838 BAA04:12 BAA27:12 Zone Cave Twilight Transition Transition Twilight Deep Deep Twilight Transition Transition Deep Transition Transition Entrance Entrance Entrance Transition Transition Transition Transition Entrance Entrance Transition Transition Transition Deep Deep Transition Transition Transition Transition Transition Transition Transition Transition Transition Transition Transition Transition Transition Transition Transition Deep t rogloxene Classification T NT NT NT NT Troglobiont NT NT NT Troglobiont? Troglobiont? NT NT NT NT NT NT NT NT NT NT NT NT NT NT Stygobiont Troglobiont NT NT NT NT NT NT NT NT NT NT NT NT NT NT Stygobiont + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + (Dingo) BAA 38 BAA + + + + + + + + + + + + + + + + BAA 35 BAA (Bullita) + + + + + + + + + + + + + + + BAA 22 BAA (Bullita) + + + + + + + + + (E of BAA 4 BAA Dingo) sp. sp. nov. sp. sp. sp. sp. Species sp. sp. sp. sp. A sp. sp. Table 1: Invertebrates recorded from the Judbarra / Gregory Karst Table Endacusta sp. C sp. B Blatella sp. sp. sp. sp. sp. Geogarypus sp. nov. Lychas sp. sp. sp. sp. Argiope? sp. sp. Philoponella sp. sp. Mesida Psilochorus Panjange Holthuisiana transversa sp. T richorhina sp. Torresitrachia Mesodontrachia aff. fitzroyana Stenopylis coarctata Westracystis lissus Westracystis Charopa Discocharopa aperta Eremopeas interioris Gastrocopta macrodon Gastrocopta cf. pediculus sp. B Pupoides pacificus sp. A Gyliotrachela australis sp.

Taxa Gryllidae Blattellidae Entomobryidae Geogarypidae Buthidae Family: ? Amaurobiidae ? Hersilidae Theridiidae Araneidae Sparassidae Ctenidae Uloboridae Lycosidae Nesticidae Tetragnathidae Pholcidae Sundatelphusidae Platyarthridae Camaenidae Camaenidae Helicodiscidae Helicarionidae Endodontidae Endodontidae Subulinidae Pupillidae Pupillidae Pupillidae Reduviidae (Harpactorinae) Pupillidae Hydrobiidae Order Orthoptera Blattodea Collembola Polydesmida Scutigerida Geophilida Scolopendrida Pseudoscorpionida Scorpionida Araneae Amphipoda Hemiptera Class NT– Non Troglobiont. Troglobiont. NT– Non

Insecta Collembola Diplopoda

Chilopoda

Arachnida

Crustacea

The specimens collected are located at the Western Australian Museum (WAM) and the Museums and Art Galleries of the Northern Territory (MAGNT). The symbol ( * ) (MAGNT). Territory Art Galleries of the Northern and the Museums Australian Museum (WAM) The specimens collected are located at the Western indicates the specimen has not yet been provided with an accession number and collection instead. 1 Gastropoda

78 Helictite, 41, 2012. Judbarra / Gregory Karst

pale yellow in colour and eyeless which indicates troglobiont status. The status of the planthopper nymphs remains uncertain as no adult specimens were collected Museum MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT MAGNT * MAGNT to confirm the presence of troglomorphisms. The degree of troglomorphisms varies between taxa with the pseudoscorpion potentially a troglobiont although this remains uncertain (M. Harvey, pers. comm., June 2011). The two potential stygobiont species were collected Acquisition / ID Number BAA27:21 BAA27:48 BAA27:61 BAA35:20 BAA22:04 BAA27:20 BAA27:17 BAA27:73 BAA27:46 BAA35:17 BAA35:16 BAA27:22 BAA35:10 BAA35:13 from a sump in BAA 38 and are a hydrobiid snail (Hydrobiidae: sp.), and an amphipod (Amphipoda: sp.). Zone Cave The majority of species were recorded from the Twilight Twilight Transition Entrance Transition Entrance Entrance Entrance Deep Entrance Twilight Twilight Twilight Transition Transition Zone (31 species, 55%), followed by the Entrance Zone (10 species, 18%), Twilight Zone (8 species, 14%) and the Deep Cave Zone (7 species, 13%) (Figure 4). Classification NT NT NT NT NT NT NT NT Troglobiont NT NT NT NT NT The cave invertebrates collected were mostly (20 species, 36%), followed by (15 species, + + + + + + + + + + + 26%), gastropods (12 species, 21%), (4 (Dingo) BAA 38 BAA species, 7%), centipedes (3 species, 5%) and millipedes and collembolan each represented by a single species (Figure 5). + + + + + BAA 35 BAA (Bullita) Cave environmental conditions + + + Measurements of the cave environments showed BAA 22 BAA (Bullita) that air temperatures were fairly constant throughout the cave, with variations generally less than a degree in most parts of the cave (Table 2). However, due to + (E of

BAA 4 BAA the generally open nature of the caves in this area with Dingo) multiple entrances, and numerous grikes and daylight holes (Figure 6), much of the cave forms part of the twilight zone where daily fluctuations of temperature and humidity may be greater. Humidity is lower near entrances as would be expected, and higher deeper into caves near sumps where the permanent to semi-

sp. permanent water ensures humidity remains at or near

Species saturated levels. The higher humidity areas are generally sp. where the troglobiont species were recorded. Due to the open nature of much of the Judbarra / Gregory Karst, Table 1: Invertebrates recorded from the Judbarra / Gregory Karst Table sp. sp. sp. sp. Omorgus sp. Ochthephilum Dihammus mixtus sp. sp. sp. Euander Physopelta australis sp. B Pirites ? sp. humidities are often lower than would be expected when compared to other tropical karst, such as Cape Range (Humphreys, 1991) and in Katherine (Tindall Limestone), where humidity is saturated, or nearly so, within a few metres of cave entrances. Taxa

Discussion Muscoidea Formicidae Gelechioidea Family:? Trogidae Staphylinidae Cerambycidae Curculionidae Meenoplidae Anthocoridae Lygaeidae Largidae Reduviidae (Emesinae) The diversity of invertebrates collected from the Judbarra / Gregory karst is quite high considering the lack of systematic collecting undertaken within the system. The entrance and twilight zones are habitats for Order approximately one third of all species recorded. These zones are relatively large and common throughout the Diptera Hymenoptera Lepidoptera Coleoptera Central karst due to the predominance of open grikes in the Judbarra / Gregory karst (Martini and Grimes 2012). The highest diversity of species was from the Transition

Class Zone, (which comprises the majority of all dark zone) NT– Non Troglobiont. Troglobiont. NT– Non The specimens collected are located at the Western Australian Museum (WAM) and the Museums and Art Galleries of the Northern Territory (MAGNT). The symbol ( * ) (MAGNT). Territory Art Galleries of the Northern and the Museums Australian Museum (WAM) The specimens collected are located at the Western indicates the specimen has not yet been provided with an accession number and collection instead. 1 due to air movement through the predominantly open

Helictite, 41, 2012. 79 Moulds & Bannink: Cave Biology

the Judbarra / Gregory also contains what may be only the second species of troglobiont scorpion on the Australian mainland, collected from the deep zone of BAA 22. The Gastropoda were the third group that dominated the invertebrate diversity of Judbarra / Gregory karst, comprising 12 species that, apart from a single species, were collected from the Dingo Cave system (BAA 38). It is highly likely that many of these species occur more widely across the Judbarra / Gregory Karst area, as they have not been as extensively sought beyond this one karst block.

Figure 3: Ecological classification of invertebrates from the Terrestrial molluscs are commonly found in limestone Judbarra / Gregory Karst (number of species and percent regions in northern Australia, usually amongst the leaf composition). litter of remnant monsoon vine rainforest. Snail shells are commonly encountered within cave sediments, probably deposited during wet season floods. Eleven species of terrestrial snail shells were found, located within the entrance, twilight, and transition zones of the caves. Two of these, Gyliotrachela sp. and Mesodontrachia sp., appear to be new species endemic to the area (V. Kessner, pers. comm.). The other nine species have been reported from various regions of northern Australia (Solem, 1981, 1982, 1984, 1985, 1989). The shell of a twelfth species of gastropod was located in the sump regions of BAA 38. It has been identified as belonging to the Figure 4 Location of invertebrates from the Judbarra / aquatic family Hydrobiidae (W. Ponder, pers. comm.), Gregory Karst (number of species and percent composition). although no live specimens have yet been collected; thus it can not be confirmed as a definite stygobiont until live specimens become available for examination. Members of this family are found in streams in southern and eastern Australia, but isolated populations have been recorded from mound springs in the Lake Eyre basin and stygobiont species from Ngalia Basin, NT. This specimen would represent the northern extent of known distribution of this family.

One species of terrestrial isopod from the superfamily Oniscoidea was located within the transition zone, and was often found in association with fig leaf litter and is considered to be a troglophile. A second, minute (5 mm), Figure 5: Composition of Judbarra / Gregory Karst blind, white specimen, collected in the deep cave zone invertebrates by Class (number of species and percent of BAA 38, belongs to the family Platyarthridae and composition). appears to be a troglobiont. Trichorhina australiensis, karst area substantially reducing the amount of deep Wahrberg 1922, is the only other recorded species from cave zone. this family on the Australian continent (H. Dalens pers. comm.), although approximately thirteen troglomorphic The diversity of species recorded showed insects taxa are currently being described from calcretes of the to be the most diverse with 20 species, although Yilgarn region in Western Australia (S. Javidkar, pers. 12 of these are considered to be accidentals and comm. 2011). therefore of little interest when determining the true diversity of the Judbarra / Gregory Karst invertebrate The subterranean waters located in BAA 38 contained fauna. Only a single insect species is classified as a an undescribed freshwater amphipod. In recent years the troglobiont (Meenoplidae sp.). The next most diverse diversity of subterranean amphipods has been found to group was arachnids with 15 species, predominantly be much larger than originally thought, with field surveys comprised of (13 species) that included several in the Pilbara and Yilgarn regions of Western Australia families (Sparassidae, Ctenidae, Lycosidae) commonly discovering numerous undescribed species (Barnard encountered as troglophiles in tropical karst areas in & Williams ,1995; Finston et al., 2004; Finston et al., Australia (Moulds, unpub. data). The fauna of 2007; Cooper et al., 2007; Bradford et al., 2010). Little

80 Helictite, 41, 2012. Judbarra / Gregory Karst

Table 2: Temperature and humidity records from the caves Cave No System Date Cave Zone Distance (m) Air Temp oC Humidity (RH%) BAA38 Dingo 28/09/93 Transition 100 (H) 24.0 75 BAA38 Dingo 29/06/92 Deep (sump) 150 (H) 23.0 90 BAA38 Dingo 12/06/93 Twilight 20 (H) 24.0 45 BAA38 Dingo 06/07/93 Entrance 8 (V) 24.0 40 BAA38 Dingo 08/07/93 Deep (sump) 32 (V) 24.0 95 BAA38 Dingo 16/10/93 Deep (sump) 32 (V) 22.5 90 BAA04 E of Dingo 01/09/91 Transition 8 (V) 23.0 70 BAA35 Bullita 23/06/94 Transition 8 (V) 24.0 75 BAA35 Bullita 09/07/93 Transition 8 (V) 24.5 70 (H) = Horizontal distance into cave from an entrance (V) = Vertical distance into cave from an entrance

Figure 6: Roots and leaf litter beneath skylights in Frontyard section of Bullita Cave. [A. Pryke] comment can be made regarding this species until it is two subterranean-adapted scorpions have previously described and placed within a taxonomic framework. been described from Australia. A very troglomorphic species, Aops oncodactylus (Urodacidae) was recently The common freshwater crab Holthuisana transversa, described from Barrow Island, a limestone island off the occurs widely within rivers across northern Australia coast of Western Australia (Volschenk & Pendrini, 2008) (Bishop, 1963), and has been collected from Grants Cave that was connected to the mainland in recent geological (KAB17) near Katherine, Northern Territory (P. Bannink time. The other most proximal troglobiont scorpion to unpub. data), caves of the east Kimberley (Humphreys, Australia is Liocheles polisorum from Christmas Island 1995), Niggle Cave (C15) near Camooweal, and in the Indian Ocean, an isolated seamount. The single northern Queensland (Iken, 1980). A population of this specimen exhibits reduced eyes, elongate appendages species was observed in BAA 22 over three consecutive years in cave passage sediments annually flooded at and generally reduced pigmentation (E. Volschenk, Limestone Gorge. This species is expected to be present pers. comm. 2011). The specimen was collected from in numerous karst systems in northern Australia where a vertical piece of moist flowstone in the deep zone of suitable habitat is present; it is considered a troglophile. BAA22 (Figure 2). This species represents the 23rd cavernicolous and troglomorphic scorpion in the world The presence of a troglobiont scorpion within the and the second from mainland Australia (Volschenk & Judbarra / Gregory Karst is extremely significant, as only Pendrini, 2008; Lourenço & Pham, 2010; Lourenço &

Helictite, 41, 2012. 81 Moulds & Bannink: Cave Biology

Pham, 2012). Mexico presently has 57%, or 13 species in Western Australia (unpub. records), while other of the world’s troglomorphic cavernicolous scorpions, Endacusta species have been recorded from Chillagoe in now followed by Australia with 13% (3 species), and Queensland (Sullivan, 1984). Vietnam 9% (2 species). Three species of cockroach from the family Only a single species of pseudoscorpion has been Blattellidae were collected. These species were recorded from Judbarra / Gregory karst, despite the commonly associated with leaf litter and organic numerous areas examined over multiple years. This detritus in the caves and all appear to be accidentals is surprising as this order is very commonly found in or troglophiles, being encountered near entrances subterranean habitats in all climatic zones of Australia and showing no evidence of troglomorphism. The (Edward & Harvey, 2008; Harvey, 1991; Moulds, 2004; relatively commonly encountered family of troglobiont Moulds et al., 2007). Two specimens of a geogarypid cockroaches from northern Australia subterranean (Geogarypus sp. nov.) were collected from the surface of environments (Pilbara, Cape Range, Undara and vertical walls in the southern portion of the Central karst Chillagoe amongst others), the Nocticolidae, have so far area (Figure 2). This represents an undescribed species not been observed in the Judbarra / Gregory karst region. that may potentially be a troglobiont as it possesses elongate appendages, especially the chelal fingers, and reduced pigmentation, although it does not have reduced Conclusions eyes (M. Harvey, pers. comm. 2011). The majority of species do not exhibit any troglomorphisms and were found in the vicinity of Several species of reduviid hemipterans were collected cave entrances and twilight zones. Seven potential from multiple caves within the karst region. These are troglobiontic and stygobiontic species have been predatory species that prey upon invertebrates. Two collected including a hydrobid snail, a buthid scorpion, of the reduviids belong to the subfamily Harpactorinae a geogarypid pseudoscorpion, a polydesmid millipede, which is commonly encountered in caves throughout a platyarthrid isopod, an amphipod, and a meenoplid the Northern Territory (P. Bannink, unpub. data). A planthopper. This is a significant number of troglomorphic further two emesine reduviids were found, including a species to be found from limited collecting in an large (8 cm) troglophile (Pirites sp.?) which has also extensive tropical cave system, and further collecting, been reported from the Kimberley (T. Moulds, unpub. especially if undertaken in a co-ordinated and systematic data), Cape Range (W. Humphreys, pers. comm.) and manner, will undoubtedly reveal additional species. The Queensland (Howarth 1988). Physopelta australis majority of these species are known from single records, (family Largidae) is an accidental, and was found and further investigation may increase their known local congregating in large numbers at the entrance to BAA 38. distribution. Multiple specimens of a pale meenoplid homopteran (planthopper) have been collected in the deep sump The absence of formal descriptions for many of passages of BAA 38 and from moist drip holes within the species collected from the Judbarra / Gregory Karst large sediment banks where tree roots were exposed in hampers the ability to provide meaningful comparisons BAA 35. No adult specimens were recorded, thus the with other nearby karst areas. The only nearby karst area species can not be confirmed as a troglobiont. This is where any general subterranean biological knowledge the only potential troglobiont species recorded from the exists is Cutta Cutta Cave, in Tindall Limestone near Judbarra / Gregory karst that is known from multiple Katherine, and in the east Kimberley (Humphreys, 1995). caves and indeed karst blocks. Troglobiont planthoppers It could be expected that some of the troglomorphic feed on the sap from subterranean tree roots, and have species may share affinities with the Katherine karst area, been recorded from numerous karst areas in Australia due to the general similarities with epigean invertebrate including Cape Range (Hoch, 1993), Undara, and fauna and similar troglophilic species encountered in both Chillagoe (Hoch & Asche, 1988). Troglobiont species areas. The further identification and description of new from this family have been collected from caves in the taxa from the Judbarra / Gregory karst remains important Katherine area (Tindall Limestone) (P. Bannink, unpub. so that the significance and degree of endemism can be data). more accurately determined.

The only orthopteran collected is an undescribed Acknowledgments species of gryllid cricket, Endacusta sp. nov. (G. Brown, pers. comm.). Twenty one species of Endacusta have We would like to recognise the Traditional Owners been described in Australia, four of which have been of the Limestone Gorge area of Judbarra / Gregory recorded from the Northern Territory, including E. National Park. This survey would not have been possible major (Alice Springs), E. koopinya, E. pardalus and without the endless amount of voluntary assistance E. paraboora (Otte & Alexander, 1983). Members of given by members of the Top End Speleological Society the Top End Speleological Society have observed and (Karen Magraith, Guy Bannink and Bruce Swain), collected several undescribed species from this genus and the Canberra Speleological Society, who explored from caves in Katherine and the Ningbing Ranges and mapped many of the cave systems, and collected

82 Helictite, 41, 2012. Judbarra / Gregory Karst many of the specimens. TESS would also like to thank Dumont, H.J. & Maas, S., 1985: Mesocyclops the Parks and Wildlife Commission of the Northern cuttacuttae n. sp. from a cave in Northern Australia Territory for permitting access to the karst region, and (Crustacea: Copepoda, Cyclopoida). The Beagle, local resident rangers Keith Claymore (1991-1993) Occasional papers of the Northern Territory and Andrew Hartwick (1994-1996) for their logistic Museum of Arts and Sciences 1: 115-122. support. A special thanks to Arthur Clarke for providing Edward, K.L. & Harvey, M.S. 2008: Short-range a summary of his unpublished data, Dr A. Wells, Dr endemism in hypogean environments: the G. Brown, Dr R. Willan and Dr A.J. Bruce of the pseudoscorpion genera Tyrannochthonius and Northern Territory Museum of Arts and Sciences for Lagynochthius (Pseudoscorpiones: Chthoniidae) in their patience and assistance in dealing with the material the semiarid zone of Western Australia. Invertebrate and also to Mr V. Kessner for identifying the terrestrial Systematics 22: 259-293. gastropods. Dr Mark Harvey and Dr Erich Volschenk Finston, T.L., Bradbury, J.H., Johnson, M.S. & Knott, provided identification and notes on the pseudoscorpion B. 2004: When morphology and molecular markers and scorpion material respectively. The two referees conflict: A case history of subterranean amphipods provided excellent suggestions which greatly improved from the Pilbara, Western Australia. this paper. The authors would like to thank Dr Susan Biodiversity and Conservation 27: 83–94. White and Ken Grimes for the opportunity to publish this paper in the Judbarra / Gregory Special Issue of Helictite. Finston, T.L., Johnson, M.S., Humphreys, W.F., Eberhard, S. & Halse, S. 2007: Cryptic speciation in two widespread subterranean amphipod genera References reflects historical drainage patterns in an ancient landscape. Molecular Ecology 16: 355–365. Balke, M, Watts, C.H.S., Cooper, S.J.B., Humphreys, W.F. & Vogler, A.P. 2004: A highly modified Guzik, M.T., Cooper, S.J.B., Humphreys, W.F., stygobiont diving of the genus Copelatus Cho, J-L. & Austin, A. 2008: Phylogeography (Coleoptera, Dytiscidae): and cladistic of the ancient Parabathynellidae (Crustacea: analysis based on mitochondrial DNA sequences, Bathynellacea) from the Yilgarn region of Western Systematic Entomology, 29: 59–67. Australia. Invertebrate Systematics 22: 205-216. 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Sparassid , in Odyssey block of Bullita Cave System. [K. Grimes]

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