Karst Wetlands Biodiversity Ani) Continuity Through Major Climatic Change: an Example from Arid Tropical Western Australia

KARST WETLANDS BIODIVERSITY ANI) CONTINUITY THROUGH MAJOR CLIMATIC CHANGE: AN EXAMPLE FROM ARID TROPICAL WESTERN AUSTRALIA

W.F.Humphreys krrestrial InvertebrateZoology, Museum o.f l,{atural Science, WesternAustralian Museum, Francis Street,Perth, WA6000, Australia; E-mail: [email protected]. gov. au

Abstract

Subterraneankarst wetlands,recently included as a wetland type under the Ramsarclassification, are introducedhere to the wetlandsbiodiversity literature.The generalcharacteristics of karst land- scapesexplain the connectionswith non-karsticwetlands, the often diffuse limits to karst wetlands and their vulnerability to exploitation and pollution. Less than l0oA of the Earth shows distinctive karst landforms but on a global basiskarst wetlandssupport the greaterpart of subterraneanbiodi- versity.This introduction is amplified by a detailed account of a single karst wetland Cape Range peninsula,Western Australia. Cape Range,the only orogenicTertiary limestone in Australia, occurs on a continent relatively devoid of karst. It lies in the arid tropics where surface water flows are mostly episodic and frequently saline. The Cape Range karst wetlands contain both freshwater and anchialine wetlands whose natural accessis through a few karst windows and caves.The anchialine system shows marked stratification of their physico-chemicaland biological profiles. The lack of surfacewaters and the particular geographical context means that much of the groundwater fauna represents ancient geographical or phyletic relicts. A general case is made supporting the persistenceof groundwaterfauna in karstic systemsthrough geologicaleras.

Introduction

The 6th Conferenceof ContractingParties to the RamsarConventionl (Brisbane 1996) voted to include subterraneankarst wetlandsas a wetland type under the Ramsarclassification system. This inclusionhas far reachingimplications, not only for karst regions,but also for a wide rangeof wetlands,and for groundwatercon- servationgenerally. Karst hydrologicalsystems are extensive,occur in many guis- es, are vulnerable,and frequentlysupport unique phyletic and distributionalrelict- ual faunas(see Humphreys. in pressa). In addition,they havedrainage characteristics (Ford and Williams 1989) that serveto influencethe hydrologicalchanges experiencedby wetlandsremote from the karst systemsthemselves.

I Conventionon Wetlandsof InternationalImportance especially as Waterfowl Habitat (RamsaqIran 197r).

BiodiversiQ in wetlands: assessment,function and conservation, volume l edited by B. Gopal, W.J.Junk and J.A. Davis, pp. 227-258 @ 2000 Backht:s Publishers. Leiden. The Netherlands 228 WE Httmphrevs

The biota of karstlandscomprise a wide rangeof epigeantaxa, including calci- phytesand calciphile lineages such as molluscs - which canbe both highly diverse andlocally endemic (Solem 1981a, 1981b, 1984,1985, 1997, Woodruff and Solem 1990)- and,in karstwetlands, phreatophytic vegetation that makesdirect use of the groundwaterand which canhave a major influenceon the local hydrology(Tromble 1977).But it also includesthe specialisedfauna restrictedto subterraneanaquatic systems- thesewill be referredto generallyas stygobitesor stygofauna(Gibert et al. 1994)in contradistinctionto troglobiteswhich are essentiallyterrestrial fauna restrictedto subterraneanair-filled voids (the term troglobitesis sometimesused sensulqto to encompassall hypogeanenvironments). Stygofauna can alsobe both highlydiverse and locally endemic (e.g. Holsinger 1978, Longley 1981, Holsinger and Culver. 1988,Wilson and Ponder1992, Stoch 1995,Bradbury and Williams 1996a, 1996b, 1997a, 1997b, Knott and Jasinska 1998, Humphreys 1999a, Humphreys,in pressb, Wilson andKeable 1999).

What is Karst

The unusualgeomorphic features of the Kras - known as Karst in the period of the Austro-Hungarianempire - region on the Italo-Slovenianborder have become known as 'karst phenomena'.The term karst denotesthe distinctivestyle of terrain resulting predominantlyfrom the dissolutionof rock by natural waters,hence the term 'solublerock landscape'is sometimesused. Owing to their solubilitykarst is most fully developedin carbonaterocks suchas limestoneand dolomite,and evap- oraterocks suchas gypsum.Such areas are characterrzedby sinking streams,caves, encloseddepressions, fluted rock outcropsand large springs(Fig. 1, Ford and Williams 1989).The integrity of such landscapesis dependentupon the mainte- nanceof the natural hydrologicalsystem and they are potentiallyhighly sensitive,, comparablein this respectto desertsor coastalmargins. Whether the coincidence of karst,desert and coastalsystems, as in the CapeRange karst discussedbelow, makesthe areaextraordinarily vulnerable is a moot point.

The Limits to Karst Wetlands

The dissolutionof carbonaterocks occursmost rapidly with running water and at the interfacesbetween waters of differing chemicalcomposition, and especially betweenfreshwater and saltwater(Ford and Williams 1989).In mostregions water tables,at somestage, have been lower thanat presentowing, inter alia, to previously drier climateor lower sealevel, hence, for this reasonalone, karst formationsare found commonly well below the current water table. Even in arid zonesthe lower partsof karstare usually below the watertable (phreatic zone) and may containboth lentic(groundwater) and lotic (pressuretubes) wetland systems. Even in thoseparts of the karst abovethe local water table (vadosezone) there may be lotic wetlands, as found for example,in undergroundstream passages. Wheresurface exposures of wateroccur in karstlands,the normalrange of wetland featuresis producedbut,, in addition,a numberof distinctivefeatures - some- times with specializedfaunas - may also occur associatedwith karst.Amongst theseare anchialine pools (Maciolek 1986, Brock et al. 1987,Thomas et al. 1991, Karst wetlands biodiversity and continuiQ through ntajor climatic chctnge 229

Fig. l. The comprehensivekarst system.(Fig 1.1 of Ford and Williams 1989;with permission).

1992,I1iffe1992, in press,Sket 1996), springs and spring brooks (Botosaneanu 1998), mound springs(Williams 1965,Harris 1992,Ponder et al. 1995,Knott and Jasinska 1998),tufa (travertine)dams (Julia 1983,Marias 1990,Pentecost 1992, Drysdale and Head 1994,Humphreys, Awramik andJebb 1995), calcretes (Humphreys 1999a) and blue holes(Stock 1986b,Ilitre 1992),details of which will be presentedonly where they are relevantto the examplepresented later in this chapter. Furthermore,open conduit flow permits large-scalemovements within karst which may facilitatedynamical processes - which may themselvesbe extrinsicto the karst system- supportingthe fauna associatedwith the karst. For example,in the EdwardsAquifer (artesian),Texas (Kuehn and Koehn 1988,Longley 1981, 1992),and in Movile Cave (karstic) in Romania(Sarbu and Popa 1992,Sarbu, in press,Sarbu et al. 1996),sulphides, respectively of petroleumand magmatic origin, supportchemoautotrophic ecosystems (Poulson and Lavoie,,in press).Analagous systemsare found associatedwith both hydrothermal(Childress and Fisher 1992) and cold deepsea vents (Arp and Fisher 1995,Scott and Fisher1995), with fauna in the latter also utilising methaneharvesting bacteria (MacDonald et al. 1989). Chemoautotropyhas also been demonstratedin FrasassiCave (Sarbu et al., in press),and strongly indicatedin anchialinesystems (Pohlman et a1.,in press, Humphreys1999b). Karst wetlandsmerge imperceptiblyinto groundwater- water which doesnot interact in the short term with surface waters - and with flow paths beneath (hyporheic)and alongside(epirheic zone))surface water courseswith which short- term interactionstake place through upwellings and downwellings(Stanford and Ward 1988,Gibert et al. 1994,Ward et al.,in press)which haveprofound effects on 230 WF Htmphreys stygalecology (e.g. Dole-Olivier et. al. 1994,Danielopol et al., in press).The fauna typical of karstwetlands may extendinto lavatubes (Wilkens et al.1986),riverine gravels(Boulton 1993),and into immediatelysub-littoral systems such as subma- rine springs (Danielopol and Bonaduce 1990). In addition, through Gyben- Hetzbergsystems - in which freshwaterlens overlie seawater intrusions(Ford and Williams 1989) they mergewith the marinesystems especially in coastsexhibit- ing anchialine(var. anchihaline) waters which may supportextremely diverse, often relictualfaunas (Thomas et al. l99l,Iliffe 1992,in press,Thomas et al. 1992,Sket 1996,Yager and Humphreys 1996).Anchialine systemsare near coastal subter- raneanmixohaline waters that are undertidal influencebut haveonly subterranean connectionswith the sea;they typically occur in volcanicor limestonebedrock and have limited surfaceexposure (Stock et al. 1986).Such waterstypically exhibit markedchanges in the depthprofiles of numerousphysico-chemical parameters (Iliffe, in press,Yager and Humphreys1996, Humphreys 1999b). Karst wetlandsinclude two distinctprocesses that integrateto producethe char- acteristickarst landscapes. For, in additionto the dissolutionprocesses defining karst, there are also constructionalkarst featuresinvolving the accumulationof new car- bonatemasses that rangein scalefrom minor speleothemdeposition to massivetufa depositson hillsides.Such depositions may involveboth biogenicand physicochem- ical processes(Humphreys, Awrwnik and Jebb 1995).Associated coral reef com- plexesmay be consideredan integralparl of the karst system(Hamilton-Smith et al. 1998),as, for example,the largeNingaloo reef fringing the CapeRange peninsula. Calcretesare a form of carbonaterock generallyomitted from discussionsof karst. Calcretesoccur as soil calcretesand groundwatercalcretes and are especially importantin the Australiancontext as they form underarid climates(annual rainfall <200 mm) with high potentialevaporation (>3000 mm per year;Mann andHorwitz 1979). Groundwater calcretes often develop solution features and exhibit the extremelyhigh hydraulictransmissivity showing that the voids and cavitiesfoim a conduit systemtypical of karst aquifers;for example,the Millstream aquiferon the Pilbaracraton of WesternAustralia (Bamett and Commander1985), and in calcretes associatedwith the palaeodrainagebasins of inlandAustralia (Sanders 1974, Mann and Deutscher1978, Jacobson and Arakel 1986,Humphreys 1999a). Groundwater caleretesin the Pilbaraand theYilgarn haverecently yielded a rich relict freshwater groundwaterfauna with Gondwanan(Harvey 1998,W.F. Humphreys, unpublished), and Pangaeanaffinities, including the crustaceanorder Spelaeogriphacea,previously known only from cavesin SouthAfrica andBrazil (Pooreand Humphreys1998) and diving beetles(Dytiscidae: Watts and Humphreys1999, in press).

Global Significanceof Karst Wetlands

About l2o/oof the earth'sdry ice-freeland comprisescarbonate rocks (Fig. 2), which are the principle karst rocks, and between7-10% showsdistinctive karst landforms and/or significant karst groundwatercirculation (Ford and Wilhams 1989).The globaldistribution of karstis uneven(Table l) - karstregions are mainly distributedon the continents(North America,Asia and Europe)and on seamount islands(Antilles, Oceania),and are lesswell representedin Africa and Australia (Juberthieand Decu 1994b)and the other Gondwananfragments. Karst wetlands biodiversity and continttitlt through major climatic change zJl

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TableI. Proportion of the land surfaceof various countriesthat are karstic. Data from Juberthieand Decu (1994b).

Country %oKarst

Malta r00.0 LesserAntilles (many of) r00.0 Cuba I5.0 Slovenia 40.0 Panama 30.0 France 25.0 Italy 25.0 Albania 25.0 PuertoRico 25.0 Bulgaria 22.s Spain 22.0 Yugoslavia(formerly) 20.0 China 20.0 Austria 17.0 USA 15.0 Canada t2.0 Belize 8.0 Czechoslovakia(formerly) 2.5 Hungary 1.5 Romania t.4 !a Poland I --)

Ninety-sevenpercent of the world's freshwateris containedin the groundwaterof alluvial aquifersand karstic systems,compared with a mere two percentin rivers and lakes(LVolich 1914).In manyparts of the world groundwateris the sourceof most potablewater and often of the major part of industrialand agriculturalwater. About 25% of the world's populationis suppliedlargely or entirely by karst waters (Fordand Wifliams 1989). This enormouslyimportant resource- evenignoring the other numerousand var- iedvalues of karsticsystems (e.g. see Ford and Williams 1989,Davey 1984) - is both heavily exploitedand r,.ulnerableto pollution. The surfaceof karst is often sparsely coveredby soil andthe undergounddrainage with its characteristicopen conduit flow, makeskarst susceptibleto the ingressof pollutantswhich, for the samereasons, are difficult to containonce they haveentered the system(Notenboom et aI. 1994). The managementof karst areas requires special knowledge - particularly becausesurface geomorphology and groundwater drainage are frequently discordant - and managementtypically needsto extend well outside the karst area itself (Kiernan 1988).The flow characteristicsin karst often precludethe simple applicationof Darcianhygrological models, especially as the hydraulicconductiv- ity of karstsystems is markedlyscale dependent (Ford and Williams 1989).For the samereasons monitoring wells in karst terrains generallydo not work (Quinlan, Daviesand Worthington 1992),nonetheless, both are frequentlymisapplied to karst systems. Tworecent volumes provide a valuableinsight into the significanceof karstwet- landsto stygofauna.The first is the 'EncyclopediaBiospeleologica'(Juberthie and Decu 1994a,1998) which providesglobal coverageof both terrestrialand aquatic Karst wetlands biodiversity and continuity through major climatic change ZJJ hypogeanfauna from both a systematicand biogeographicalperspective, and deals with the hypogeanenvironments nation by nation, including karst systems.These volumesprovide more detailedinformation on the distributionof karst within each 'Stygofauna country and the characteristicsof its hypogeanfauna. The secondis Mundi'(Botosaneanu1986) which is a compilationof theworld's stygofauna to that date.I haveused it to obtainan estimateof the significanceof karstwetlands to stygofauna globally, despitethe inevitable bias resulting from the concentrationof work on stygofaunain Europeand North America. Conservatively,I estimatethat about58% of the worlds stygofaunaare associatedwith karst wetlands(Table 2). The nearly 2000 recordsinvolved in thesechapters, which compriseonly about a fifth of the chaptersin the book, are indicativeof the biodiversity containedin the stygofauna.Although stygofaunahas been sparsely sampled and little researched,it is clearthat karst systemsharbour a major part of the hypogeanwetlands biodiversity.Within them,karst windows between epigean and hypogean environments offer a fascinatingglimpse at the subtleselective pressures leading to the evolutionof the morphological,behavioural and physiologicaladaptations to subterraneanlife (Culveret al. 1995).

Table2. The distribution of stygofaunaspecies in karst and non-karsthypogean habitats. Amongst the l9 major taxa a conservativeaverage of 58oh(+31.7) are found in karst wetlands.Data were extracted from chapters taken at random - excluding predominantly marine groups - from Botosaneanu(1986) and classified according to the habitatdescription given in the endpapers.Karst categoriesinclude habitatsA-G, I, T and U; non-karstinclude habitatsH, K-S,V and Z; habitats denoteda andb wereexcluded as they could not be thuscategorised. This is a conservativeestimate as someof the non-karstclassifications may actuallybe from karst(e.g. artesian springs). The com- bined numberdoes not equalthe numberof speciesin eachgroup becausein many casesa species is found in both karst and non-karsthabitats. A speciesis included even if the habitat is denotedas probable.

Chapter Karst Non-Karst %oKarst

5. Porifera 3 0 100 8. Microturbellaria sub.fw.l 26 50 34 9. Turbellaria:tricladida 79 81 49 a I 1. Rotatoria J 87 J 1A a- 16. Nematoda- continentalsub. aquatic IJ 22 JI 23. Mollusca- continentalsub. aquatic 249 85 75 33. Copepoda:Cyclopoda - continentalsub. fw. 93 5l 62 35. Copepoda:Harpacticoida - Continentalsub. fw. 170 255 40 t1t 39. Syncarida 27 lJl t7 40. Mysidacea 25 I 96 49. Isopoda:Oniscidea t6 0 100 50. Isopoda:Phreatoicidea I 6 t4 54. Amphipoda:Melitid grouping 37 59 39 55. Amphipoda:Niphargus group 117 183 39 56. Amphipoda:Crangonyctid - Holarctic 82 6l 57 6 I . Amphipoda:Liljeborgiidae 0 100 62. Amphipoda:Pardaliscidae 1 50 63. Amphipoda:Sebidae 0 r00 aa 86. Pisces )I 9 80

I sub. fw. - subterraneanfreshwater. 234 WE Huntphreys

Protection

The potentialand realised impacts of humanactivities on groundwaterfauna are sum- marisedprimarily from the latestauthoritative review by Notenboomet al. (1994). Groundwatermanagement includes the needto protectits ecologicalintegrity but this is difficult in karstareas because the dispersivityof pollutantscan be very high, and the "filter" phenomenacan be deficient or even totally absent(Notenboom et al. 1994).Once pollutants have reached aquifers, sorption and degradation processes are generallylow and residencetimes long. Togetherthese make groundwaterenviron- mentsrulnerable and difficult to rehabilitate.Organic pollution can changeground- water foodwebsbecause it createsfavourable conditions for colonisationby detritiv- orousepigean organisms. Under these conditions, the uniqueadaptations of many sty- gobitesto oligotrophicconditions is overriddenby pollution andthey arereplaced by epigeanspecies (Notenboom et al. 1994,Malard 1995). Anchialinesystems, such as much of the coastalarea of the CapeRange peninsula discussedin the following example,which often containendemic fauna, are extra- ordinarilyvulnerable to evenslight organicpollution (Iliffe et al. 1984),while being tolerantof a wide rangeof physico-chemicalconditions (Sket 1996,Humphreys 1999b).The terrestrialfauna (troglobites), which providessome food for aquaticbiota in the limestoneaquifeq also reflectschanges in the percolationinput of increasing pollution by wastes,fertlhzers, and toxic chemicalsQ.{otenboom et al. 1994).

Cape Rangepeninsula, Western Australia: an example

In this chapterno attemptis madeto review karstwetlands on a global basis.On the contrary,owing to the novelty of their inclusionin the Ramsarwetland classifi- cationsystem, one structurally quite diverse system will be examinedin somedetail to flesh out someof the essentialcharacteristics of suchsystems. Some comment will be madeon the relativediversity of this systemvis-d-vis other systems globally. The systemin questionis the CapeRange peninsula of northwesternAustralia, one of the world's more diversekarst faunaprovinces. As an exampleit servesboth to emphasisethe diversityof the associatedfauna and its diverseorigins, as well as the diversity of wetlandtypes containedwithin the karst itself. The exampleserves to indicatethat such systemsmay havecontinuity in both spaceand time throughout long periodsof geologicaltime.

Australia

Althoughareas of carbonaterock occurless frequently in Australiawhen compared with the world average,about 500 discreetlocalities are documented(Hamilton Smith and Eberhard in press).In Australia eight of the 698 wetlandsrated as nationallysignificant (ANCA 1996)comprise inland subterraneankarst but only threeinclude obligate subterranean fauna - Loch McNess(ANCA 1996 p. 891) in southwesternAustraha, the SpringTower Complex (ANCA 1996:p. 311) in northern Queensland"and the Cape Range SubterraneanWaterways (ANCA 1996:p. 774) in northwesternAustraha. The latter,despite being in an arid region,is the only wetlandlisted principally for its subterraneanaquatic fauna. Karst wetlands biodiversity and continttiQ throttgh ma.jor climatic change 235

The listing, ?Snationally significant, of only eight karst wetlandsin Australia beliesthe significancethere of karstwetlands. Although recentwork hasshown that groundwaterfauna is both diverseand widespreadin Europe and North America (Marinonieret aL.7993,Gibert et al. 1994),it is poorly known inAustralia. It is moderatelydiverse in Tasmania(Eberhard et al 1991),New SouthWales (Eberhard and Spate 1995),,and in the Kimberley, WesternAustralia (Humphreys 1995a). Elsewherein WesternAustralia the stygofaunais highly diverse in the Pilbara (Poore and Humphreys 1998, Humphreys, 1999a)and in parts of the Camarvon Basin (CapeRange peninsula and Barrow Island:Humphreys 1993b 1994, 1999a, Bradbury and Williams 1997b).Owing to the novelty of the inclusionof karst wet- landsunder the RamsarConvention, it may be expectedthat many significantareas of karstwetland will be recognisedworldwide as they are assessedand as knowl- edgeof their stygofaunaimproves. As will be shownbelow, a karst region of moderatesize may supportnot only a greatdeal of geneticdiversity within taxa,but speciesswarms and diversecommuni- ties within habitats,as well as diversehabitats. One karst arcamay supportan inter- mixed affayof hypogeansystems - terrestrial,freshwater, marine and anchialine.

Cape Rangepeninsula

Arid zonesare consideredto be sparselypopulated with troglobitessensu lato owing both to a lack of water (Howarth 1980) and the low food input from xeric plant communities(Peck 1978).Hence, a priori, the arid northwestof Australia might be thoughtperipheral to debateon specialisedsubterranean animals, as well as to discussionon both wetlandsbiodiversity and persistence.Yet the subterranean fauna of the CapeRange peninsula in northwesternAustralia is amongstthe more diversein the world andis recognizedto containrelict faunaof the highestconser- vation status(Morton et al. 1995). The diversity of the subterraneanfauna of the CapeRange peninsula is partly a resultof thejuxtaposition of terrestrial,freshwater and anchialinesystems. Within the compositionof this faunaare echoesof orogenicand eustaticevents, climatic change,and of past connectionswith other parts of Australia, easternGondwana and evenPangaea (Harvey 1993,Harvey et al. 1993,Humphreys 1993a-d Poore and Humphreys1992,Yager and Humphreys1996). While I dealhere only with the aquaticcomponent of the fauna, it should be borne in mind that this rich aquatic fauna occurs immediately beneath a diverse terrestrial fauna of obligate cave dwellers(Humphreys 1993a 1993d\ in pressb) which will, in partat least,form part of the food web (Humphreys1999b, Humphreys and Feinberg1995). Here,I discussthe diversityof both the hypogeanbiota and the habitatsof this arid zone subterraneanwetland the affinities of its fauna and its ancientorigins. The thesisis developedthat thesesubterranean wetlands, being well bufferedfrom the vicissitudesof most types of surfacechange, can survive little alteredthrough erasof geologicaltime. Suchcommunities may be expectedto be very resilientto the perturbationsresulting from Recentclimatic change,despite their apparentvul- nerability to other anthropogenicchanges (Iliffe et al. 1984)and the wide variety of physico-chemicalenvironments that they may inhabit(Humphreys l994,Yager and Humphreys1996, Sket 1996,Humphreys 1999b). 236 WF Humphrel,s

RegionalSetting

To appreciatethe areait is necessaryto placeit in regionaland geologicalcontext. The CapeRange peninsula projects northwards into the Indian Oceanon the western shoulderof Australiajust within the tropics(Fig. 3). The westcoast is fringed by the Ningaloo coral reef lying on Australia's narrowest continental shelf (Kendricket al. 1991).Barrow Islandlies 170km northeast,on the shallowNorth West Shelf, part of a continuousseries of shelvesextending around northern Australia to New Guinea.Cape Range and Barrow Island are anticlinesof Terttary marinelimestones of Oligoceneto Mioceneage (Wyrwoll et al. 1993,McNamara andKendrick 1994).The CapeRange anticline, rising to an altitudeof 310 m, start- ed to fold in the Late Miocene(Malcolm et al. l99l) and hasbeen fully emergent since at least the Pliocene(Wyrwoll et al. 1993),possibly earlier (Humphreys 1993c).Cape Range is the only Tertiaryorogenic limestone in Australia;limestones of this type are the host rocks for much of the most biologically interestingkarst in the world (Spateand Little 1997).The emergentmarine terracesand the coastal plain, cut into the Miocene limestones,are coveredby Plioceneand Quaternary depositsof alluvial and colluvial materialsincluding carbonate based conglomer- ates,calcarenites, siliceous dunes and sandplains (Wyrwoll et al. 1993). The Cape Range SubterraneanWaterways (hereafter 'the waterways')occupy the entire coastalplain and the lower foothills of the CapeRange peninsula north- ward of a line joining NorwegianBay on the west coastand the Bay of Rest on Exmouth Gulf (ANCA 1996).A closelyrelated system occurs on Barrow island (Fig. 3). They occurin the Quaternarydeposits of the coastalplain and extendinto the underlyingTertiary limestones and those of the foothillsof CapeRange. In the waterwaysfresh water overlayssea water that has penetratedthe aquifer at depthand the watertabledeclines towards sea level nearthe coast(Allen 1993). It is a Ghyben-Herzbergsystem in which a freshwaterlens floats on a wedgeof sea water(Fig. 4). In a perfectGhyben-Herzberg system under hydrostatic equilibrium, the layer of freshwateris about 40 times as thick as the height of the water table abovesea level (Fordand Williams 1989).In practice,as in CapeRange, there is a zone of mixing where the water is variously brackish and this zone may be very thick, up to 30 m in the waterways(Yager and Humphreys 1996, Humphreys 1999b).If the groundwaterflow is intercepted,for exampleby a borefield"then it will causethe saltwaterinterface to moveinland and upward(Davidson 1995). The coastalplain watersare, in places,anchialine, that is they haveno surface connectionwith the seabut fluctuatewith marinetides, contain sea-derived species, have limited exposureto the open air, and have noticeablemarine and terrestrial influences(Stock et al. 1986).Anchialine waters, to which accessis gainedthrough karst windows, are typically highly stratified"with fresh water - brackishwater in arid areas overlying seawater at depth (Fig. 5). This stratification is commonly associatedwith major changesin otherphysico-chemical parameters such as oxy- gen concentration(becoming less oxic with depth; Sket 1996, Yager and Humphreys1996, Humphreys 1999b), and redox potential. Consequentlythe nearcoastal sections of the waterwaysgrade from marinenear the coast and at depth, to freshwaterinland and near the surface,with a variable Karst wetlands biodiversity and continuiQ throttgh major climatic change LJ/

Fig. 3. The regional setting of the Cape Range Peninsulaand Barrow Island. The Precambrian PilbaraCraton has not undergonemajor regionaldeformation since 2400 Ma (Trendall 1990)and has been, emergentcontinuously for >600 Ma. This is fringed to the west by middle Mesozoic deposits(c. 100 Ma). A seriesof anticlinesof CretaceousandTertiary limestones occurs furtherto the west.Compiled from Hocldng et al. (1987)and Wyrwoll et al. (1993).Drawn by C. Larvrence.

zone of mixing in between. Such anchialine systems are noted both for their relict faunasand their speciesrichness (Sket 1981 1996),and they are the subjectof widespreadconservation assessment (Sket 1981, Maciolek 1986, Brock et al. 1987, Ridgleyand Chai 1990,Martin et al. 1991,Thomas et al. l99l,lliffe 1992,Martin et al. 1992,Thomas et al. 1992,Bailey-Brock and Brock 1993). The waterwaysoccur beneath a diversearid zonevegetation comprising mostly Eucalyptarsscrub and Ficus, and, on the plain, principally Triodia grasslandand 238 WE Htmphreys

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100 |50 2oo Concentratim (lrl4)

Fis. 5. Depth profile of the physico-chemical environmenrof cave C-28 in the Cape Range SubterraneanWaterways.

chenopods(Keighery and Gibson1993). Stable isotope composition shows that the energyutilized by the faunain one easternpart of the rangeir p..oo-inantly from d1ep9rrooting "planti c-3 plants but with a significant contributionfrom c-+ (Davies1996). While both the terrestriaranO aquatic components feed on the specialisedsubterranean fauna,they. also make opportunisticuse of epigeanfaina e-n-tering the stygal.realm through surface op.ningr, both natural (Humphreys and artificial and Feinbergrgg5, w.F. Humphrlys,unpublished). Accessto waterwithin cape Rangep.op.. is ilmliea to five cavesin which the rvatersurface is at gO an altitudeof aboui t m. Thesecaves contain a melitid amphi- showingmarked 99{ geneticdiscontinuity across gorges (Humphreys and Adams 1991,Adams and Humphreys lgg3). It was prou"aut!on.. drt oi the speciose lurr-pnreys ]?.?-3t) melitid community on the coastalplains (Bradbury and williams 1996a)but from which it is now disiunct. The subterranean realm above the water table contains a very rich, endemic troglobitecommunity which comprisesat least41 species(31 gd.ru; and consti- tLltesmostly a relict rainforest-floor fauna,with boih temperai-eand tropical ele- tnents(Harvey et al. 1993,Humphreys 1993a 1993c). This atteststo a much wetter 240 WF Humphreys past,probably in the late Miocene (Humphreys1993c). Together with the stygofauna of at least22 specres(12 genera)they make the Cape Range/BarrowIsland systemone of the worlds richer assemblagesof subterraneanfauna. This is especially the caseas the areahas been the subjectof very little researcheffort com- paredwith other areasrecognised for the diversity of their hypogeanfauna, such as the AppalachianMountains in the United States(e.g. Ilolsinger and Culver 1988), or Mexico (e.g.Reddell and Mitchell l97l).It containsll.2o/o of the describedspe- cialisedsubterranean fauna of the World'stropics, 15. 1o/oof the aquaticand 8.6% of the terrestrialfauna (based on datafrom Peckand Finston1993).

Methods

Samplingand descriptiveeffort has focusedon the subterraneanfauna and so char- acterisationof the biota of the few openpools is poor.Sampling has been conduct- ed from the surfaceand by cavediving, using netsof varioustypes (mesh size 125- 350 pm), baitedtraps and by hand from open anchialinepools, caves, traditional wells (humanoccupation of the areaextends to 33',000years BP; Morse 1993),pas- toral wells and boresconstructed for otherpurposes (water abstraction, piezome- ters,,exploration, monitoring and shot holes), rarely by filtering the output of pumps.The physico-chemicalenvironment was determinedas describedelsewhere (Humphreys1994, l999b,Yager and Humphreys1996). For the purposeof this paper,it is convenientto considerthe Cape Range SubterraneanWaterways together with Barrow Island"despite the lack thereof karst windows and where accessto the stygofaunais mainly through 40 to 50 m deep boreholes.The shallowNorth WestShelf would havebeen exposed through much of the Pleistoceneand into the early the Holocene.There are both terrestrialand aquaticsubterranean species in commonbetween Barrow Islandand the eastcoast of the Cape Range peninsula (Adams and Humphreys 1993, Harvey and I{umphreys1996) suggesting that the emergentshelf was suitablehabitat for the troglobitessensu lctto (Humphreys 1993c).

Biodiversityin an Arid Zone Wetland

The waterwayshould be discussedagainst the backgroundthat the areais both arid and hot, and made effectivelymore arid owing to the karst terrain.The mean daily temperatureis c. 27'C with the mean monthly maximum temperatureexceeding 35'C for four monthsand the meanrelative humidity is persistentlylow (Vine et al. 1988).Consequently the annualevaporati on (3219mm) exceedsthe sparseprecip- itation(median:280 mm) by morethan an orderof magnitude(Vine et al. 1988). Rainfall in the regionresults from at leastfour processes(Gentilli 1972)resulting in very low predictabilitybetween both seasonsand years (Humphreys et al. 1989). The fixed red sanddunes of Pleistoceneage (Wyrwoll et al. 1993)that overlieparts of the north and southparts of the range,and beneathwhich both troglobitesand stygobitesare found (W.F.Humphreys, unpublished), attest that previouslythe area hasbeen even more arid. Conversely,at othertimes the areahas been substantially wetter, supportingrainforest, as is evident from the lowland rainforest affinities (Harveyet al. 1993,Humphreys 1993a 1993c, in pressb) of the terrestrialtroglo- bite faunain both CapeRange itself and on the adjacentcoastal plain. Karst wetlands biodiversity and contintrity throttgh major c'limatic change 241

Stygofauna

A minimum of 26 stygofaunaltaxa occur in the wetlands(Table 3) includingthe only two speciesof blind fish in Australia.The remainingspecies are mainly crustaceans and the faunaappears predominantly to be endemicto the CapeRange and Barrow Island systems.The fauna includesa great diversity of amphipods(Bradbury and Williams 1996a, 1996b, 1997a, 1997b)- the only known representativesin the southernhemisphere of the classRemipedia (Yager and Humphreys 1996), the order Thermosbaenacea(Poore and Humphreys1992), and the generaHaptolana (Bruce andHumphreys 1993) and Danielopolina (Baltanas and Danielopol 1995). The crus- taceanfauna includesmarine (melitid and hadziid amphipods),euryhaline (bogidiellid amphipods)and freshwater(syncarids, which probably evolvedbefore the break-upof Pangaea)lineages, as well as major stygobiontlineages that are widely disjunct from congenericspecies found on either side of the North Atlantic (e.g. Remipediaand Thermosbaenacea)and/or which belong to lineagesclearly Tethyan in origin (thaumatocyprididostracods, Haptolana). Six genera are apparently endemic to the region (Humphreys 1993b, 1994, Knott 1993, Bradbury and Williams 1996al997b,Yager and Humphreys1996). The anchialinefauna consistsof the samegroups of crustaceansas recorded from anchialinecaves of the Caribbeanregion (Bahamas,,the YucatanPeninsula of Mexico, and Cuba;Yager 7987a,1987b, 1994)and the CanaryIslands (Lanzarote; Wilkens et al. 1986, 1993).It comprises,amongst others, remipedes, cirolanid isopods,atyid shrimp, ostracods,gammarid amphipods and thermosbaenaceans.

Table3. The composition of the stygofaunaof northwesternAustralia. About half of the fauna has beenfound since 199I . The fauna is found variously in cavesin CapeRange itself (R), on the coastal plain borderingCape Range (P), and on Barrow Island(I). * denoteshighly stygomorphicspecies.

Major taxon Genus and Species Locality

Turbellaria Microturbellaria PI Polychaeta:Spionidae Gen.nov. et sp. nov.R. Wilson, pers.comm. 1996 P Ostracoda:Halocyprida * Danielopolinasp.nov. Baltanas & Danielopol,1995 P Ostracoda:Cypridacea Aglaiella sp.nov.K. Wouters,pers. Comm. 1995 PI Ostracoda:Cytheracea Darwinula sp. K. Wouters,pers. Comm. 1995 P Remipedia:Nectiopoda x Lasionectesexleyi Yager & Humphreys P Syncarida:Bathynellacea Atopobathynellasp. nov. H.K. Sminke,pers. comm. 1994 IP Isopoda:Cirolanidae * Haptolana pholeta Bruce & Humphreys PI Amphipoda: Melitidae * Norcapensismandibulis Bradbury & Williams R * Nedsia douglasi Barnard & Williams P * N. strakraba,N. hurlberti, N. humphrevsi,N. uriJimbriata, I N. Jiagitis, lrl. macrosculptilis, N. culptilis all Bradbury & Williams Amphipoda: Hadzitdae * Liagoceradocttssubthalassicls Bradbury & Williams I * Liagoceraclocttsbranchialis Bradbury & Williams P Amphipoda:Bogidiellidae + Bogidommaatrstralis Bradbury & Williams I Thermosbaenacea * Halosbaenatulki Poore & Humphreys PI Decapoda:Atyidae * St1;giocarislancifera Holthuis P * Stygiocaris styli.fera Holthuis PI Pisces:Eleotridae * Milyeringa veritas Whitley P Pisces:Synbranchiformes * Ophisternoncandidurz (Mees) P 't A1 L+Z- WE Httmphrevs

This aquaticcommunity compositionhas become predictable in anchialinehabitats (Holsinger1989, 1992, Poore and Humphreys1992, Humphreys 1993b, Yager and Humphreys 1996) despitethe great disjunction from congenersthat are mostly found in subterraneanhabitats on either side of the North Atlantic (e.g.,Poore and Humphreys1992, Bruce and Humphreys1993, Yager and Humphreys1996). The affinities of the fauna are consideredpredominantly to lie with the Tethysocean (Table 4; Humphreys \993b, Knott 1993) that openedbetween the continentsof Gondwanaand Laurasiaand which persistedfrom the Triassicuntil the late Eocene (200-40Ma; Smithand Briden 1977).Itwill be arguedlater that this may represent a surviving Jurassiccommunity. The stygofaunaoccupies waters with a wide rangeof physico-chemicalcharacteristics.While somespecies appear to be restrictedto watersbeneath a strongpycnocline (Lasionectesexleyi, Danielopolina sp. nov., Liagoceradocusbranchialis; Yager and Humphreys 1996), others, such as Stygiocaris sQlifera and Milyeringa veritas, are variously found in habitatsranging from open anchialinepools in fulI sunlight,in canalsin freshwatercaves, and beneath a strongpycnocline at a depthof 32 m in seawaterbeneath several layers of hydrogen sulphide in almost anoxic waters.The rangeof physico-chemicalcharacteristics of the water from which various specieshave been collected is given elsewhere(Humphreys 1999b). The fauna is predominantlyeuryhaline. The meansalinity (total filterablesolids, mg L-1) of the water at siteswhere O. candidumwas sampled was 3520 (range710-7700; n : 8 samplesites), M. veritas 8168 (250-26500;n: 20), Segiocaris spp. 2859 (310- 20000;n:25), H. tulki2577 (610-7700; n : 19),H. pholeta1783 (660-3300; n : 6), I{orcapensismandibulis in CapeRange s.s. 437 (390-500;r : 3), melitid amphipod, coastalplain 3622 (660-l1000; n: 19).

SurfaceFauna

The waterwayshave very restrictedsurface expression, limited to one substantial drownedsinkhole (cenote: Yager and Humphreys1996) and perhaps15 karst windows,no more than 2 m across,into the anchialinesystem. The stygofaunaldiver- sity, high by world standards,is complementedby an epigeanfauna which inhabits thesekarst windows and alsocontains some disjunct species (see Appendix). Although only two algaehave been characterisedthese karst windows support substantialprimary productionfrom greenplants in the photic zone.The nitrifica- tion and sulphurbacteria, which occur nearthe pycnocline(Yager and Humphreys 1996,Humphreys 1999b), may make somechemoautotrophic contribution to the energysupply for this anchialinefauna (Humphreys1999b) as occursin a similar systemin Yucat6n(Pohlman et al., in press).Anchialine atyids on Hawaii,in addition to filter feedinglike their freshwaterrelatives (Fryer 1977),also scrapethe ben- thic algallcyanobacterialcrust ingesting diatoms, cyanobacteria and algae(Bailey- Brock andBrock 1993).Hence, it is likely thatsuch systems are more complex than currentlyappreciated. The faunaldiversity in the karstwindows includesstrictly marineforms, suchas the circumtropicalsyllid polychaeteSphaerosyllis centroamericana, and estuarine elements,such as the mollusclravadia sp.and possibly the amphtpodGrandidierella sp.,elsewhere known only from the saltingsat Dampier,WesternAustralia (J. Lowry. Karst wetlands biodiversity and continuity through major climatic change 243

Table4. The affinities of generafrom the stygofaunaof northwesternAustralia. T : Tethyan,G : Gondwanafl,P : Pangean

Taxon Genus Distribution Affinities

Ostracoda:Cypridacea Aglaiella Egypt,Mozambique, Sri Lanka (K. Wouters,pers. comm. 1995) Halocyprida Danielopolina T WestIndies, Canary Is., Galapagos, Atlantic abyssal Syncarida:Bathynellacea Atopobathynella G SE Australia,New Zealand Chile Thermosbaenacea Haktsbaena T WestIndies, Colombia, Canary Is. (Poore& Humphreys1992) Amphipoda:Melitidae* l,ledsia spp. T (Knott 1993) Hadziidae Liagoceradocus T W. Pacific, Madagascar,Canary Is. Bogidellidae Bogidomma P Endemicgenus Isopoda:Cirolanidae Haptolana T Cuba, Somalia (Bruce & Humphreys 1993) Decapoda:Atyidae Stygiocaris T Madagascar(Banarescu I 990) Remipedia:Nectiopoda Lasionectes T Turks & Caicos,West Indies (Yager& Humphreys1996) Pisces: Perciformes:Eleotridae Milyeringa Endemic genus,affinities unknown Synbranchiformes Ophisternon ?T Circum tropical (Mexicancaves)

*Melitids are possiblyancestral to the Hadziids(Barnard and Barnard 1983),almost all of which are blind stygobiontswith a Tethyandistribution.

pers. comm. 1994).Freshwater elements are also representedsuch as thiarid molluscs, water mites, dragonflies, gerrids and dytiscids and feral tropical aquarium fish. The meiofauna of the area is essentiallyunknown. One substantial surface waterway, Yardie Creek, crosses the coastal plain and extends as a short gorge, less than two kilometre long, extending into Cape Range. This overdeepenedgorge - the only extensivepermanent inland water within c. 100 km - cuts through the waterways and must be in hydrological continuity with them. However, the nature of this connection has not been characterized although tidal fluctuations, as is generally found in the anchialine system (W.F. Humphreys and R.D. Brooks, unpublished), do occur in the creek when the bar is closed (N. McGregor, pers. comm. 1996).The creek, which is generally closed by a sandbar that is breached by exceptional tides andlor rainfall - and when dugong,,Dugong dttgon (Mtiller), may feed there (N. McGregor!,pers. comm. 1996) - is a mixoha- line system that ranges from fully marine to freshwater. Despite the large extent of this waterway within an arid zone,no systematicsam- pling has been undertaken there. The biota is predominantly marine, including a remnant of a formerly more extensivemangal (Kendrick and Morse 1990), and fish, at least one species,Crctterocephalus capreoli Rendahl (Atherinidae), having pop- ulations in the creek that are genetically divergent from those in the adjacent sea(R. Watts, pers. comm. 1996). In the less saline upper reaches,and at a small spring-fed freshwaterpool raised above the general level of the creek, there is a fringing vegetation of emergent aquatics,chiefly Typhadomingensis Pers. (Typhaceae) and Schoenoplectuslitoralis Schrader (Cyperaceae).Both are disjunct by severalhundred kilometres from their 244 WE Humphrelts respectivemain range(Keighery and Gibson1993). There is alsoa disjunctpopu- lation of MelanoidesSp., ? freshwaterthiarid gastropod(Slack-Smith 1993).

Age and Persistenceof the StygofaunalCommunity

Here I discussthe ageand persistenceof the subterraneancommunity now inhabit- ing the waterwaysand showthat they could haveexisted in this habitatfor very long periodsdespite great climatic changes(Quilty 1994).The generalstability of the areahas attributesfavouring long-term persistence of a subterraneanfauna. It is in close proximity to the Pilbara Craton, emergentsince the Silurian at least. Additionally,it hasbeen in continualjuxtaposition, since the Cretaceous,to both an emergentshoreline and to the continentalshelf, and throughoutthis period hashad a carbonatedepositional environment (Hocking et al. 1987). I argue that the composition of the anchialine fauna from Australia to the Caribbean,and the presenceof obligatestygal species widely disjunctfrom their congeners,supports an early vicarianceas a resultof seafloorspreading and sug- geststhe survivalto the presentof a Triassicor Jurassiccommunity. This community wasprotected in its stygalrealm from changesat the surface.I go on to discuss the generalsignificance of groundwaterfauna regionally in permitting the persistenceof biota throughclimatic and otherchanges at the surface.

TethyanAlJinities and Age of the Fauna

A number of crustaceansoccur as congenericspecies on either side of the North Atlantic which belong to higher taxonomic ranks (e.g. Remipedia,Thermosbae- nacea)whose biological characteristics are suchthat they that wereconsidered to be especiallyconvincing examples of Tethyanrelictual distributions (Newman 1991). Thesetaxa arerestricted to a rangeof hypogeanhabitats, have few young and the free-livinglarval stageshave been suppressed. They areextreme K-strategists (Stock 1986a)- the specieshave very limitedpower of dispersal,a potentialfurther limited by their restrictionto a range of hypogeanhabitats and often specialisedenviron- mentalrequirements. For example,remipeds have only beencollected by cavedivers in anchialinecaves below a densityinterface (Yager and Humphreys1996). These specialisedrequirements enhance the alreadyhigh potentialof troglofaunasensu lqto to be discriminatingpalaeogeographical monitors (Danielopol 1980, Sbordoni 1982). Furthermore,the presenceon either side of the North Atlantic of congeneric speciessuch as Halosbaena and Monodella (Thermosbaenacea),Stygogidiella, Pseudoniphargus,PsammogammerLts, Gevgeliella and Spelaeonicippe(Amphi- poda), Curassqnthura(Isopoda) and Speleonectes(Remipedia), which have the samestygal lifestyles, provides support for the hypothesisthat the vicariancewas dueto the openingof theAtlantic. The hypothesisis more convincingbecause their distributionon a reconstructedJurassic landmass is relativelymodest (Fig. 6). A numberof speciescongeneric with membersof the classicallyTethyan relict faunafound on eitherside of the North Atlantic, or with otherwisewidely disjunct distributions, are also known from the Cape Range system of northwestern Australia(Fig. 3); for example,Halosbaena (Canaries and Caribbean),Hctptolana (Cuba, Somalia),Lasionecles (T[rks and Caicos),Liagoceradocus (Canaries and Karst wetlands biodiversitlt and continttiQ through major climatic change 245

Fig. 6. The relativelocation of the continentalplates and the extentof the epicontinentalseas in the Jurassic(151 Ma) when the TethysSea first connectedthe presentCaribbean, Mediterranean and IndianOcean regions. Even with this reconstructionthe generaHalosbaena (Thermosbaenacea; H), Haptolana (Isopoda: Cirolanidae;I) and Lasionectes(Remipedia; L), known from northwest Australiahave widely disjunctdistributions variously including Somalia, the Canaryand Caribbean islands.The plate map was producedusing Scoteseand Denham(1988) and simplified.

Pacific) and Danielopolina (North Atlantic and easternPacific; Baltanasand Danielopol 1995).Being obligatestygofauna, the generaare likely to havebeen presentas stygofaunabefore the isolationof the Australianplate. At leastone of theselineages is known to havebeen present in marine cavesat this time. Fossil ancestorsof Danielopolinaoccur in submarinefissure and cavefills showingthat they had alreadycolonised para-anchialine systems by the Late Jurassic(Aubrecht and Kozur 1995).Shouldthese five independentlineages prove to be monophylet- ic, thenthe ageof vicarianceof thesewidely disjunctcongeneric species may, perhaps,be soughtin the congruenceof areacladograms. Attempts are currentlyin progressto testthis hypothesisusing molecular clocks. The distributionof remipedsand their associatedfauna can be explainedby cave colonizationalong the shoresof Tethysduring the Triassicand Jurassic(225-160 246 WF Huntphrevs

Ma BP; Cals and Boutin 1985),and subsequentdispersal after the break-upof Pangaeain the Mesozoic(Wilkens et al. 1986)through sea floor spreadingand con- tinentaldrift (Stockand Longley 1981, Iliffe et al. 1984,Hart et al. 1985,Wdgele and Brandt 1985). During the early Cretaceousthe plate that is now Western Australiaformed the easternshore of GreaterTethys (Howarth 1981). For both Halosbaena (Wagner 1994, H.P. Wagner, pers. comm.) and Danielopolina(Baltanas and Danielopol1995) cladistic analysis suggests that the Australianpopulations were isolated from thosein theAtlantic regionprior to the vic- arianceresulting from the openingof the Atlantic itself. This sequenceis consistent with that of the break-upof Pangaeaproposed by Howarth(1981) and the dissolution of Gondwanaduring which the separationofAfrica fromAntarctica(i.e. of Westfrom EastGondwana, 142-133 Ma BP) predatedthe openingof theAtlantic Basinbetween Africa and SouthAmerica(127 Ma BP; Partridgeand Maud 1987).Bythe Upper Cretaceous(c. 80 Ma BP, probablyeven by the Middle Cretaceous,100 Ma BP; Candeand Mutter 1982)Australia was not evenjoined by epicontinentalseas to land- massesother than Antarctica (Howarth 198 1). This concordanceis consistentwith the hypothesisthat the generarafted on the tectonicplates and in which casethere has beenremarkable morphological conservatism (cf. Pooreand Humphreys 1992, Bruce andHumphreys 1993,Yager and Humphreys 1996) following an earlyvicariant event. The very predictabilityof the highertaxonomic composition of this community (seeabove), with widely vicariantcommon genera, coupled with the clearTethyan affinities of the fauna,suggest that the community has survivedsince the opening of the Atlantic andthat we aredealing with a survivingJurassic community. Underthis hypothesis,the faunais much too old to havecolonised the Tertiary limestonesof CapeRange,, and they would haveto havecolonised older substrates and subsequentlymigrated to their presentlocation. Being obligatestygofauna, they must perforcehave migrated within a matrix that was of necessitycontinuous in spaceand time. What factorsfavour this route? Inland from CapeRange, the PilbaraCraton - continuouslyemergent since the Precambrian(Hocking 1990)- is fringedwith Mesozoicrocks which initially could havebeen colonised by the fauna and from which subsequentmigration within the matrix could have populatedthe developingCainozoic rocks (Humphreys 1993b 1993c).That the areaseems to be suitablefor suchmigration - acrossthe emergent North WestShelf in thePleistocene - is supportedby the currentdistribution of both terrestrialand aquatictroglobites (see above) as well as the geneticaffinities of StygiocarisstyliJbra (Adams and Humphreys1993). However, the areaalso has other attributesmaking it especiallyfavourable for the survivalof an ancientrelict fauna.

TectonicContext and Palaeo-environment

The Exmouth Sub-basin,in which Cape Range lies, was initiated in the Early Jurassic(c. 200 Ma BP) and was in contactwith the waters of the Neotethys (Apthorpe1994) and hasremained predominantly a marginalmarine environment. It hasbeen continuously in closeproximity to the continentalmargin since Greater Indiabroke away in theEarly Cretaceous(c. 140Ma BP).By themiddle Cretaceous (c. 85 Ma BP) the separationof the tectonicplates had advancedsufficiently to permit full oceaniccirculation, which, combinedwith the warmerwater resulting Karst wetlands biodiversity and continuity throttgh major climatic change 1^1 from the northwardmovement of AustraliaPlate, led to the dominationof reef and platform carbonatesfor the remainderof the Cretaceous.This marine carbonate depositional environment continued throughout the Catnozotc(Malcolm et al. 1991).An eastwestcompressional phase of tectonismin the Late Miocene,that resultedfrom the subductionof theAustralian plate beneath the Asian plate,led to the formation of the Cape Range and Rough Range anticlinesby inversion and thrustingalong the basin margin (Malcolm et al. l99l). A fault trendingnorth-east across the baseof the peninsula(roughly from Point Cloatesto the Bay of Rest;Fig. 3) marksthe boundarybetween the Gascoynesub- basin and the Exmouth sub-basin(Hocking et al. 1987).The Cretaceousand Cainozoic depositsin the latter are underlainby Carboniferousand earlier rocks, whereasin the former they are underlain by Jurassicrocks, as are those in the Barrow sub-basinand the Dampier sub-basin.This boundaryclosely approximates the known southerndistribution of the CapeRange stygofauna.

LongeviQ

The parsimoniousexplanation, then, is that we are dealingwith a survivingJurassic communitythat hasgradually separated from the Atlantic provincethrough sea-floor spreadingand rafting on the tectonicplates. Even were the faunaMiocene in origin - proposedby sometheories on the origin of anchialinefaunas (Por 1986,Stock 1986a, 1990)that do not incorporatethe highly disjunctdistributions of obligatestygofauna (Humphreys,in pressa) - the longevityof theseaquatic systems is still considerable. To accountfor the presenceof a clearly old faunain TertiaryCape Range, I con- sideredthe community to have migrated acrossthe North West Shelf during the Miocenefrom adjacentto the PilbaraCraton (Humphreys 1993b,1993c). However it now seemspossible that this phasecould have occurred much earlier,in the middle Cretaceous,and that the community could subsequentlyhave remained more or less in situ stayingwithin the matrix near the surfaceas the carbonatesediments accumulated.Nonetheless, several elements of the stygofaunahave recently been found,as predicted (Humphreys 1993b, 1993c), in aquifersof the CarnarvonBasin where they abut the Pilbara Craton (W.F. Humphreys,unpublished). There they (Haptolana, Stygiocaris, Halosbaena) are found in alluvial aquifers overlying Miocenelimestone, in the cobblebeds deposited by riversentering the coastalplain from the PilbaraCraton, and evenpenetrating river gravelsonto the cratonitself as far as groundwaterdivides (W.F.Humphreys, unpublished). At presentelements of the stygofaunaare known from an altitudeof c.300m to 3l m below sealevel. The fauna is sampledparticularly from the crystallineTulki Limestoneand as the range is an anticline,it dips beneaththe sea aroundthe peripheryof the peninsula.Marine tides arc apparentin the caves1.5 km inland (Fig. 7; W.F.Humphreys and R.D. Brooks,unpublished) and in waterbores up to 3.5 km inland (Forth 1973)indicating open conduit flow of the groundwaterwith- in the coastalplain. This tidal movementwill serveto expandthe zoneof mixing at the saltwaterinterface and transport nutrients and energythough the coastalanchia- line aquifer.The latter is demonstratedby the transportof heat associatedwith the tidal movementin a cave(Fig. 8), suchheat transport may in itself haveecological consequences(Shepherd et al. 1986).This movementis thoughtto occurin karsti- 248 WE Humohrevs fied Tulki Limestoneas it dips beneathsea level. Owing to Pleistocenesea level changes,the limestoneis likely to be karsticto greaterthan 100m below sealevel, channelshaving formed at the changingwater table, especiallyby mixture corro- sion.Divers have penetrated caves to 31 m below sealevel, deeper than the present depthof the seaover much of the North West Shelf. Someof the more recentsurface changes impinging upon the areaare evident. During the Quaternary oceanswere lowered by 120-140 m during maximum glacialsand elevatedby 5 to 8 m in the warmestinterglacials. Thus for most of the Pleistocenethe North WestShelf would havebeen exposed above sea level and had a similar geomorphologyto the presentcoastal plain. The distributionand genetic compositionof the presentfauna indicatesthat this extensivecoastal plain provided suitablehabitat for both the terrestrial and aquatic subterraneanfauna. Such eustaticchanges would havealtered the geographicextent of this wetlandwhich is now pushedagainst the foothills of CapeRange and is aboutas constrainedas it has ever been.The fixed red dunes and the relict rainforesttroglofauna provide evi- denceof both wetterand drier climatesin the past(see above). It is hypothesisedthat the elementsof this community have survived,by virtue of its subterraneanexistence, owing to the ameliorationof the vagariesof the surface,geomorphological and geologicalchanges, and dueto their being ableto move through the matrix of the rock while remaining protectedin the hypogeanrealm, successivelyadjusting to vertical and lateral shifts in the location of its habitat.

GeneralCase

Thereis wide supportingevidence from othertaxa and areasthat troglobitessurvive in their protectedunderground ecosystems with remarkably little morphological

1200 1180

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Fig. 7. Semidiurnaltidal movementsin cave C-28, an anchialinecenote on the Cape Range peninsula, 1.5 km inland from the Indian Ocean.Data were recordedhourly using pressuretransducer and data logger for 10 days in 1996 fDataFlow, Noosaville, Queensland](W.F. Humphreys and R.D. Brooks, unpublisheddata). Karst wetlands hiodiversity and continuity through major climatic change 249

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Fig. 8. The relationshipbetween relative water level (X mm) and water temperature("C) in C-215, a strongly tidal cave,for the period 300-339 siderealdays 1996.The water is 23.8 m below ground level and would in the absenceof tidal action, be expectedto have a stablewater temperatureon a daily basis.The tidal cycle is about half that shown in Figure 7 but of similar form. Temperature: 0.023x,- 1.18x 10-5Xz + 16.58(r2 :0.46,Fs 2,934: 403.1,P<0.0001) (WF. Humphreysand R.D. Brooks, unpublisheddata). changethrough long periodsof geologicaltime. As no ordinal taxon of terrestrial troglobitesis restrictedto the hypogean,the most compellingevidence comes from aquatic taxa. A rangeof widely distributedfreshwater lineages is known whosebiogeography is explicableif they enteredfreshwater and were subsequentlydispersed by tectonic plate rafting, for example the syncarids(Schmink 1974, Boutin and Coineau 1987),spelaeogriphaceans (Poore and Humphreys1998), and bogidiellid (Karaman 1982,Stock 1977 1983) and crangonyctidamphipods (Holsinger 1986),plus at least four orders of isopods (phreatoicids,aselloids, calabozoids and ?proto- janiroids:Wdgele 1990). It appearsthat suchhypogean animals and even,as suggestedhere, the communities in which they occur, can survive the myriad perturbationsthat would have impingedon a givenlocale through geological time - tectonic,orogenic and eusta- tic events,climatic, atmosphericand oceanicchanges, and of pastconnections with other parts of Australia, easternGondwana and even Pangaea.As such, anthro- pogenicglobal warming and its consequencesis unlikely to causeany seriousprob- lems to these hypogean wetlands, in sharp contrast to the epigean wetlands. Nonetheless,these hypogean wetlands are highly vulnerableto more direct anthropogenicchanges (e.g. Notenboom et al. 1994),having a low capacityto copewith disturbance,and which are difficult, if not impossible,to restoreonce degraded (Yuan 1988).Owing to the characteristicsof karstregions, and wherethe surface and groundwaterdrainage are often discordant(Kieman 1988,Ford and Williams 1989,European Commission 1995), management has to be holistic and often to extendbeyond the confinesof the karst itself. 250 WF Humphrel,s

Acknowledgements

I thank the numerouspeople who have supportedme in the field and the laborato- ry and who have provided information, taxonomic expertiseand encouragement. The work has been variously funded by Federaland Stateagencies, private trusts and companies.All are acknowledgedfully in more appropriatepublications.

Appendix

High leveltaxonomic groupings of taxacollected in openwater bodies that do not comprisepart of the stygofauna.The referencesdenote the sourceof determinations other than the author.

Rhizoclonium ?tortuosum (Dillw.) Kuetz. (Chlorophyta: Cladophoraceae); Lamprothamniumpapulosum (Walk.) J. Gr. (Charophyta:Characeae); Euplotes sp. and Parameciumsp. et al. (Protista,,Knott 1993); various undetermined foraminifera(D. Haig, pers. comm. 1995),microturbellaria, rotifers, nematodes,, oligochaetes,ostracods and harpacticoidcopepods; molluscs lravadia (Iravadia) sp. ?L ornata (Iravadiidae:status ambiguous: Slack-Smith 1993) andMelanoides sp. (Thiaridae; Slack-Smith 1993); polychaetesSphaerosyllis centroamericana Hartmann-Schroder1959 and Typosyllus(Ehlersia) cf. broomensrsHartmann- Schroder1979 (Syllidae);ostracod Aglaiella Daday 1910 sp. nov. (K. Wouters, pers. comm. 1995); the cyclopoid copepodsMetacyclops mortoni Pesce,De Laurentiis and Humphreys 1996, Microcyclops varicans G.O. Sars,Apocyclops dengizicus (Lepechkine), Halicyclops longifurcatus Pesce, De Laurentiis and Humphreys 1996, and Halicyclops spinifer Kiefer 1935; aorid amphipod Grandidierellasp. nov. (elsewhereknown only from saltworksconcentration pools in Dampier;J. Lowry, pers.comm. 199\; water mrte Coaustraliobateslongipalpis (Lundblad) (Acarina: Hydracharina:Hygrobatidae); fragments of trichopteraand odonata(Humphreys and Feinberg 1995); girrid bug Limnogonussp. (nymph, from distributions it is probably Limnogonusfossarumgilguy Andersen and Weir, in press:T.A. Weir,pers. comm. 1996);chironomrd Chironomus (Kie/ferulus) intert- inctus Skuse;dytiscid beetle Copelatusiruegularis Macl.; feral guppiesPoecilia reticulataPeters 1859 (Pisces: Poeciliidae). The latterare in an overflowpool in the borefieldsupplying Kailis Fisheries(conductivity 335 mS m-1)and in the tank at UpperBulbarli Well (conductivity110 mS m-l).The formeris maintainedas a stock wateringpoint andthe guppieswere reportedly introduced to controlmosquitos at the recommendationof the W.A. WaterAuthority (G. Passmore;pers comm.). The gup- pies clearlythrive underthese conditions and the populationdensity is immense.The populationscould be spreadby peopleor by flooding of the naturaldrainage line. If they enternatural freshwater or brackishwaters they arelikely to eliminatethe native fauna(G. Allen, pers.comm. 1993).Several other species of feral tropical aquarium fish have recently been seen in the anchialine system, including cyclids (W.F. Humphreysand D.B. Noltie, unpublished)but they do not seemto have become established(pers. obs. 1999). Karst wetlands biodiversity and continttity throttgh major climatic change 251

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