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INTRODUCTION The Biodiversity Series This report is the fourth in the Biodiversity S~ries produced by the Department of the Environment, Sport and Territories. The series intends to provide information about aspects of Australia's considerable biodiversity, including its global and national significance, and its conservation and management status. Many of the papers iri the Series will consist of reports prepared for the Biodiversity Unit on a range of biodiversity related subjects.. This report has been written by scientists within the CSIRO Division of Wildlife arid Ecology under commission to the Biodiversity Unit. It investigates the significance of refugia to biodiversity in the arid and semi -arid zones of Australia, and examines the management issues confronting them. Some 70 per cent of Australia is included in the study area. The area is of vital economic importance and is also significant as a reservoir of biodiversity, much of which (particularly plant and invertebrate diversity) remains to be discovered. Within this vast dry area pockets - exist which, because they retain moisture, or because they have been isolated from climatic or human-induced changes, act as refugia or refuges. These refugia contain concentrations of plants and animals, sometimes in unique ecosystems. Many of these organisms are relicts of millennia old faunas and floras and as such are of very great scientific interest. Many refugia are places to which populations of plants and animals retract in the face of adverse environmental conditions such as Widespread drought, and still others have acted as refuges from land clearing and exotic animal invasions of the last two centuries. This report examines the 76 refugia which can be identified in arid and semi-arid Australia . from all available literature and describes their· known significance'for biodiversity . conservation, including endangered species, species of evolutionary importance, and species· which are restricted to these areas. Land uses and management issues are also identified. It is noteworthy that in s()me regions, fewer refugia than expected were found in references, perhaps reflecting a lack of data. An appendix explores a methodology for identifying ecological refugia. The authors have made a significant contribution to understanding the importance· of refugia in the drier areas of Australia. This report is a valuable addition to the Biodiversity Series. Department of the Environment, Sport and Territories 3 1. TERMS OF REFERENCE The report is to consolidate and summarise information on refugia in Australia's arid and semiarid areas, their value and role in conserving biological diversity and in maintaining the ecological sustainability of surrounding areas, and the management requirements for their long term .sustainability. The project was designed to cover the points outlined below. 1.1. Identity of known biological refuges Location, extent, type (e.g. ephemeral wetland, mound spring, cave, gorge, .etc.). , 1.2. Importance for the conservation of biological diversity Species known to be endemic to the area. Rare or endangered 'species found there. Significant populations. 1.3. Importance for ecological sustainability of surrounding areas Importance for maintenance of natural populations,· including those species which help regulate pest outbreaks. Role in reducing land degradation, including role as seed banks. 1.4. Management requirements Threats posed: type and extent. Potential to reverse degradation or eliminate threats. Current land uses which conflict or may conflict with biodiversity conservation. Type of land management. Gaps in knowledge are delineated. In describing the importance for the conservation of biodiversity, estimates of importance are included, and the known and potential values of each refuge are noted. A summary 6f management r~quirements is also provided. Status as wetland sites of importance (including Ramsaf), and national or other conservation parksor reserves. Importance as refuges in dry years, as areas from which recolonisation can OCCUf. Importance for relict species or populations. Degree of degradation, if any. 7 Biological-Refugia in Australia Refugia are identified by reference numb,ers, eg WA 1; SA 13; NSW 5 etc, as in text (pages 71-131). See also State maps at beginning of Sections 4-8. . L WA 13 (( WA 11 WA WA19 WA 14. WA21 WA2 WA2B. NT2 • WA1B~WA26 WA30. . WA23 2111. WA 27 /". WA 12 • NT13 • WA1 • WA3 WA4: WA15 • WA20 • WA22 • WA17 SA3 WA~ o, 600 , Kilometres 8 SYDNEY · . CANBERRA o 9 2. EXECUTIVE SUMMARY Several types of refugia are defined to take into accounuhe fact that different concepts may be involved in the idea of a refuge, In evolutionary . ..--tenns, a refuge is a region in which certain organisms persist during a period in which most of the original geographic range becomes uninhabitable because of climatic change. However, the term refuge may be used in other ways, particularlyto mean a region to . which sgedes retract for short periods (i.e. for a numbeiOf years at the most) when large parts of their preferred habitats become uninhabitable because of drought or other effects. Yet another type of refuge comprises regions to which threatened species have retreated because of enVironmental changes set in train by European settre;ent. Our investigations have led us to the following nine categories: • . Islands • Mound springs • Caves • Wetlands • Gorges • . Mountain ranges • Ecological refuges • Refuges from exotic animals • Refuges from land clearing Information on the distribution of plants and animals in arid and semi-arid Australia was surveyed in order to determine the identity and location of foci of biological diversity. Subsequently, this data-set was usedto identify locations thought to constitute refugia because they possess unusual aggregations of species considered endemic, relictual, threatened, or otherwise significant. The chief refuge valu,e of each is noted, and comments provided on threatening processes within the refugia, and on land tenure. Although the emphasis of the study, by force of -circumstance, is on the species component of biological diversity, the refugia that are identified are likely to incorporate at least some values inherent in genetic and ecosystem diversity. An attempt is made to provide a measure of. the importance of each refuge that might be used to compare among them. As is the case with most ranking procedures, this one cannot be considered as concrete, objeclioe orfinal. It is merely a starting-point for discussion. 10 The number of refugia identified in arid and semiarid Australia was 76; there were 32 in Western Australia, 13 in South Australia, 14 in the Northern Territory, 8 in Queensland, and 9 in New South Wales. Some refugia possessed values appropriate to more than one category, but when classed according to their primary characteristics the distribution of refugia among the categories was as 'follows, Islands, 9; Mound springs, 4; Caves,J; Wetlands, 36; Gorges, 5; Mountain ranges, 12; EcologicaI.refuges, 2; Refuges from exotic animals, 3; and Refuges from land clearing, 2. Our attempt to rank the refugia showed that Wetlands tended to be classed as of lower importance compared to the other categories because they possessed fewer notable species. A preliminary analysis on a test region of central Australia shows that it would be possible for Landsat imagery to be used to identify areas of persistent vegetational greenness; such areas might function as shortcterm drought refugia for certain organisms. Management issues affecting refugia are discussed. It is concluded that they are identical to those facing arid Australia asa whole, land degradation resulting from over-grazing by domestic stock and by feral animals, alterations in hydrology, removal of habitats by clearing at the fringes of the semi-arid zone, . depredations by exotic predators, and uncontrolled fire. Thus, better management of refugia is dependent upon mitigation of the same forces operating throughout the landscape, but improved management isespecially necessary in these refugia because the organisms within them frequently have no other home. 3. METHODS 3.1. Definitions - What is a Refuge? It is imperative that we be clear from the beginning about what constitutes a refuge in arid and semi-arid Australia. The Oxford dictionary defines "refuge" as "place of shelter from pursuit or danger or trouble". In ecological or conservation terms, though, this definition needs to be refined in order to encompass several important varieties of refuge springing from quite different sources. In evolutionary terms, a refuge is a region in which . certain types or suites of organisms are able to persist during a period in which most of the original geographic range becomes uninhabitable because of climatic change. The resulting refuge contains high frequencies of endemic species, because the species in it tend to respond to the contraction of ringe by evolving differences from their original, widespread stock (e.g. Brown and Gibson 1983). Well-known examples include patches of tropical forests in many parts of the world, refugia for plants in the Arctic, and - of importance to us - organisms confined to isolated, moist environments in arid regions. Such ospecies are frequently refe~ed to as relicts. However, this category of refuges in evolutionary time does not include all phenomena that we need to describe. Ecologists frequently use the term refuge in other ways, particularly to mean a region in which a species or suite of species persist for short periods when large parts of their preferred habitats become ' uninhabitable because of unsuitable climatic or ecological conditions (e.g. drought, flooding or biologically-driven collapses in food supply). Such 'c, ecological refugia may operate for less than one .!leneration, or just a few, but may nevertheless be especially significant in arid Australia because of the 'continent'spropensity to drought. Yet another ;~ategory of refuge comprises regions in which \ /Jhreatened species occur; in this case, a species has ·fT~.treated because of factors ultimately to do with ;:?~nviro~me~ changes set in train by European 's.ettlement. Por purposes of description, and ···.a.rticularly for management action, it is vital to i~"ntify the correct category of refuge. Our 'y"stigations hav.e led us to the following set of leo'ogb.ries, as defined below. . .. 0 0 '.-'-,------.~-- .\ ;,',:-:, . " 3.1.1. Islands Refuges in historical time from introduced predators and competitors and from changing land-use due to European use that has otcurred on the adjacent mainland. Often rich in endangered mammals and endell)ic species of mammals, reptiles and plants. Also offshore islands from Barrow Island in the north-west to Kangaroo Island in the south-east were refuges from predation by Aborigines and dingoes in pre-European times (Abbott 1980). 3.1.2. Mound springs Localised habitat with Regional endemics of aquatic oinvertebrates (isopods, ostracods, and hydrobiid molluscs) and fish (where springs form wetlands).. Springs are areas with concentrations of species with limited range and specialised habitat requirements; as more taxonomic work is undertaken, more of the biota is being recognised as regionally endemic. They may also provide a localised habitat for waterbirds that ·is not affected by seasonal variations in rainfall. The Dalhousie Springs are a well-known example. 3.1.3. Caves Provide a refuge in evolutionary time for species that formerly occupied the litter layer of the forest floor of rainforest at a time when the climate of Australia was much wetter and rainforest extended into areas / that are now arid or semi-arid. The Nullarbor caves constitute examples of such refuges. 3.1.4. Wetlands Provide permanent or semi-permanent wetland habitat in arid or semi-arid environments to the benefit of waterbirds and aquatic fauna and flora. Includes long-lived pools in river systems as well as lakes, and can be broadened to incorporate intermittent wetlands such as Lake Eyre. 3.1.5. Gorges Provide a specialised microclimate for plants and animals that allow the continued survival of forms that were far more widespread in the evolutionary ·past when climates were )Vetter. These;reas often "provide additional refuge from present-day' ecological forces such as fire. A well-known example is Palm Valley in central Australia. 11 3.1.6. Mountain ranges Provide a mixture of refuges in evolutionary 'time through the presence of sheltered environments and geographically isolated habitats in which speciation may take 'place, but also provide run-off water and plant nutrients which may produce relatively resource-rich areas functioning a~refugia in ecological time-frames. 3.1.7. Ecological refugia As noted in the previous category, the presence of relatively dependable supplies of moisture and nutrients may provide n;fugia for animals dependent upon regular plant production for persistenq; in the uncertain Australian climate. 3. 1. 8. Refuges from exotic animals These refuges in historical tiine include sites other than islands. The refuges have effectively been created by human activities, and include environmental zones atJhe edge of the range of exotic species (e.g. the northern fringes of the ranges of rabbits and foxes) and localised areas where species have been able to evade the impact of these species (e.g. rugged cliff habitat where animals can escape from the effects of introduced predators and competitors). 3.1.9. Refuges from land clearing Areas where species or suites of species have survived because clearing has been proved uneconomic due to-aridity or infertility. This phenomenon is most prominent at the semi-arid fringes o(the arid zone. 3.2. Definitions - What Constitutes the Arid and Semi-Arid Zones? 3.2.1. Boundaries The arid and semi-arid zones are widely recognised to constitute those areas which receive less than 250 mm annual average rainfall on the southern margin and, in the north where evaporation is greater, less than 500 mm (Williams and Calaby 1985; Figure 3.1). We adopt this definition and the boundaries shown by Williams and Calaby. 3.2.2. Subdivisions Some geographic structure is crucial as a basis for discussion of refugia within the vast area constituted by arid and semi-arid Australia. Rather than create our own sub-divisio'ns, or use older regions such as those of Beard (1969) even though they may be established in the literature, we chose to use the Interim Biogeographic Regionalisation for Australia (!BRA) ofThackway and Cresswell (1994). The' advantage of doing so, i.e. of presenting the information in a context in which future discussion of it can readily proceed, far outweighs the slight disadvantage that may spring from forthcoming minor adjustments to the boundaries of the Interim Regions. Nevertheless, it should be borne in mind .. ·that these Regions are still subject to modification and refinement, and are best considered as a con';enienl framework for discussion rather than as absolute environmental categories. The !BRA was derived by compiling the best available data and information about each State and Territory, including field knowledge, published resource and environmental reports, and biogeographic regionalisations, as well as continental data sets. The !BRA was developed specifically for the National Reserves System Cooperative Program, and the focus is on continental landscape assessments. It is acknowledged that such a regionalisation cannot be the sole criterion for allocating conservation priorities. The Biogeographic Regions occurring in , arid and semi-arid Australia are shown in Figure'3.1. 3.3. ALiterature Survey Designed to Identify Foci of Biological Diversity and to Determine the Location of Refugia We approached the task of describing refugia by scanning as much scientific and technical literature as possible within the time available to us. For each Biogeographic Region, we accumulated references on natural history and ecology and extracted from them information relevant to potential,foci of biological diversity. We made p~rticular efforts to obtain material from the "grey literature" represented by publications of very limited distribution but which frequently contain much detailed survey or ecological information. 'The information sought concerned: 12 " , • species listed as endangered (E) or vulnerable (V) by ANZECC (hom that shown in Australian Nature Conservation Agency 1994); • species that are thought to be endemic to the Region under consideration; • relict populations (whatever the cause of relictuality); • other populations considered by authors of the material to be si.gnificant; • and geographic areas identified by the authors as constituting refugia. This material is presented, Region-by-Region and State-by-State, in sections 4-8, in the order Western Australia, South Australia, Northern Territory, Queensland, and New South Wales. When this material was brought together, we inspected it to determine whether significant records of species or groups of species fell into a pattern that might reflect the presence of any of the categories of refuges specified in section 3.1. These data were then summarised into identification of the refugia occurting in each Region, again grouped according to State or Territory, in sectio'ns 10-14.. Our assessment has been inclusive rather than critical; instead of excluding doubtful cases, we have tended to list them. This approach has been adopted because we envisage this Report as a starting point for more detailed field assessment of the refuges identified herein 'purely from a literature survey.. Sections 15 and 16 discuss some general aspects of the refugia and of the management issues emerging . from our analysis. Finally, an Appendix (section 18) examines the possibility'-.of using satellite imagery to identify dependably productive areas that may function as short-term refugia for certain types of organisms. If imagery could be analysed in this way, it might expedite the identification of ecologically significant parts of the landscape requiring special management. Figure3.1. The Australian continent showing the boundary of the arid and semi-arid region (heavy line; from Williams and Calaby 1985) and the relevant Biogeographic Regions (from Thackway and Cresswell 1994). Interim Biogeographic Regionalisation for Australia Regions are shown which fall wholly or partly within the arid a.nd <om'_,r,rl zones (indicated by the heavy black line). 13