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Invertebrate Systematics
C o n t i n u i n g I n v e r t e b r a t e Ta x o n o m y

Volume 16, 2002 CSIRO 2002

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Invertebrate Systematics, 2002, 16, 555570

IS02 09 SMho.Srt-HangrveyndemisminAustralia

Short-range endemism among the Australian fauna: some examples from non-marine environments
Mark S. Harvey
Department of Terrestrial Invertebrates, Western Australian Museum, Francis Street, Perth, WA 6000, Australia. Email: [email protected]

Abstract. The Australian fauna is assessed for short-range endemism at the species level, i.e. the prevalence of species with naturally small ranges of less than 10,000 km2. The phenomenon is found to be widespread and several groups are found to consist principally of short-range endemics: Gastropoda (snails and slugs, both freshwater and terrestrial), Oligochaeta (earthworms), Onychophora (velvet worms), Araneae (mygalomorph spiders), Schizomida (schizomids), Diplopoda (millipedes), Phreatoicidea (phreatoicidean crustaceans), and Decapoda (freshwater crayfish). The majority of taxa with high numbers of short-range endemics possess similar ecological and life-history characteristics, such as poor powers of dispersal and confinement to discontinuous habitats. The conservation of such groups is often hampered by poor taxonomic knowledge, but modern, comprehensive biotic surveys will be helpful in identifying short-range endemics.

Introduction Biogeographic analyses in which the distributions of organisms are examined and compared in detail have attracted considerable interest over the past 150 years. Indeed, detailed biogeographic observations first enabled Charles Darwin (Darwin 1859) and Alfred Wallace (see Pantin 1960) to deliberate upon the mechanisms that formed the morphological and behavioural differences between closely related but allopatric species. Distributional ranges of organisms are used in many facets of biology and have enabled researchers to explore more fully biological variation, life-history strategies in relation to local ecological conditions and geographical variation within a putative species. Distributional ranges of organisms are key elements of many guides to the worlds biota and maps in such publications can provide valuable clues to the correct identification of local forms, especially in well-studied groups such as flowering plants, butterflies and vertebrates. Evidence for small ranges among many of the more poorly studied taxa such as non-flowering plants and most invertebrates is sometimes regarded with suspicion and attributed to taxonomic ignorance, resulting in the dismissal of concerns for the well being of the taxon in question. Australia can be proud of its achievements in more fully assessing the biogeographic limits of its lesser-known fauna with landmark surveys of strategic species, often to gain further insights into other aspects of each species biology (see references in Ponder and Lunney 1999). Biological
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surveys in which a wide variety of taxa are collected for museum and herbaria collections represent the modern version of 19th century expeditions dispatched to all corners of the globe by wealthy industrialised nations in search of unusual specimens to fill their biological institutions. Modern surveys, however, have different goals in mind and use different tools with which to analyse the resulting data. Such contemporary examples include McKenzie et al. (1991) and Burbidge et al. (2000), in which a wide variety of terrestrial and freshwater taxa were systematically sampled to determine basic biodiversity patterns to assist in the assessment of local values for reserve establishment. Many modern surveys and published taxonomic works feature taxa that appear to be rare. The biology of these organisms is often little known and their precise distribution and life history are poorly documented. Much detailed research is often needed to uncover even the most rudimentary aspects of the missing knowledge. However, it has become clear from many recent coordinated surveys and taxonomic revisions in Australia that some taxonomic groups consist entirely of species whose distributions are naturally small. These taxa are often referred to as narrow-range endemics (e.g. Ponder 1999) or short-range endemics, and it has become clear in the last decade that there are many more short-range endemics (SREs) in the Australian fauna than previously suspected. The advent of electronic databases in which locality data, and other data, can be interrogated has revolutionised our ability to assess such taxa. This paper is designed to give a basic overview of
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some of the major non-marine faunal groups in Australia and assess each of them for the widespread presence of SREs. I have deliberately avoided a total survey of SREs and confined my review to those groups that predominantly consist of SREs. This unfortunately omits taxonomic groups such as beetles and mites, which comprise numerous SREs, but focuses attention on the lesser-known orders, which tend to receive less attention from biologists than their more abundant brethren. Geographic coverage It is difficult to define a short-range endemic, especially as most species have discontinuous ranges. I have here adopted a conservative approach and used a benchmark of 10,000 km2 as the maximum range for a short-range endemic, equating to a grid of 100 km 100 km. The actual area of occupancy of an SRE may be far less, but, as a result of inadequate surveying, it is virtually impossible to estimate such areas for most animal taxa in Australia today (but see Hansen and Richardson 2002). The extent of short-range endemism in the Australian fauna A review of the relevant literature on the Australian fauna indicates that short-range endemism within the bulk of a single major taxonomic unit is rare among the Australian fauna. Despite the occasional presence of short-range endemic species, there appear to be few SREs within the non-marine fishes, amphibians, reptiles, birds and mammals (Appendix 1). Exceptions include sporadic examples of species isolated in refugial habitats such as mountaintops, rainforest isolates or mound springs (Larson 2001). Similarly, there appear to be few insect or mite groups that exhibit extensive short-range endemism, despite the occurrence of numerous species that are restricted in their distribution either through geological and climatic variables (e.g. Yeates et al. 2002) or through ecological constraints (e.g. Yen 2002). Marine habitats seem to be devoid of higher taxa that are SREs, but the presence of SREs, especially in estuarine and coral-reef habitats, is widespread (e.g. Hooper and Kennedy 2002; OHara 2002). The largely continuous habitat and the widespread presence of planktonic larval forms among the majority of marine animals may prevent the evolution of SREs. However, in some sessile marine invertebrates, such as the colonial ascidians studied by Davis et al. (1999), there may be reduced gene flow between populations as a result of limited dispersal by the larval stage. For the purposes of this paper, I have excluded any further consideration of the marine environment. Ecosystems that may induce short range endemism Terrestrial ecosystems in Australia represent a wide array of environments that were categorised into a national framework of 80 bioregions by Thackway and Cresswell

(1995). Among the terrestrial fauna, there are numerous regions that possess short-range endemics. Mountainous terrains often harbour a variety of SREs largely owing to their topographic relief, which provides refugial habitats that are absent from the surrounding landscape (see Yeates et al. 2002). The widespread aridification and forest contraction prevalent during the Miocene through to the Pleistocene (Hill 1994) resulted in the fragmentation of populations. These refugia are important on a regional scale because numerous isolated and restricted species are found in such areas. Hopper et al. (1996) discussed some of the processes that may lead to the extreme diversification of the semi-arid Australian biota. Some freshwater habitats in Australia have a high proportion of SREs and many are restricted to individual river systems or drainage basins. Permanent freshwater ecosystems provide stable environments for a wide variety of taxa including many relictual lineages of Gondwanan affinities. Despite the wide array of new taxa being found in Australian freshwater ecosystems, the Australian limnological fauna is much better known than most terrestrial ecosystems, and it is possible to assess the distributions of many different faunal groups. Caves and other subterranean cavities provide suitable habitat for a wide variety of animals, both terrestrial and aquatic. These species are invariably evolved from an epigean ancestor and many cavernous environments contain relictual species whose nearest relatives nowadays exist in distant locales (Humphreys 1993, 1999). The aridification processes endured by Australia during the Tertiary led to a widespread contraction of the rainforests that once covered the country. In some areas with suitable geological conditions, the voids represented the sole place for some taxa to survive. The recent discovery of a high diversity of obligate stygal invertebrates existing in groundwater calcrete aquifers within the Australian arid zone (Humphreys 2001) highlights the largely untapped biodiversity that still awaits discovery. Most of the species thus far recorded in these habitats qualify as SREs because each aquifer comprises a discrete fauna. However, the higher taxonomic groups to which they belong also contain many widespread species and they have thus been omitted from the discussion below. Taxonomic survey Phylum MOLLUSCA Class GASTROPODA Remarks Molluscs are one of the most diverse group of living organisms, second only to arthropods in diversity. The highly endemic gastropod fauna of Australia has radiated widely both in freshwater and terrestrial ecosystems (Beesley et al. 1998).

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Freshwater molluscan radiations in Australia have been well documented, particularly those of the hydrobiids in south-eastern and south-western Australia (e.g. Ponder et al. 1993; Miller et al. 1999; Clark and Richardson 2002; Ponder and Colgan 2002). Numerous species in several genera have been detected and most species are SREs; some are even restricted to individual streams. A single radiation in Tasmania and eastern Victoria has generated at least 43 species of Beddomeia, 12 species of Phrantela, three species of Nanocochlea and four species of Victodrobia (Ponder et al. 1993). Further specimens representing unnamed species were also available to the authors, but were not described or formally named because of the enormous workload such a complete study would entail. Similarly, many species of terrestrial molluscs possess extremely restricted ranges. The number of SREs in the Australian fauna is large and many families consist entirely of SREs. Among the best-known studies are those of the late Alan Solem (see citations in Solem 1997), who recognised a vast array of species with highly restricted ranges. These restricted elements were particularly obvious in the drier regions of northern and western Australia, but similar patterns are found in land snails from the wetter regions of eastern Australia (see citations in Smith 1992). Phylum ANNELIDA Order HAPLOTAXIDA Remarks Although many earthworms are virtually cosmopolitan through anthropogenic dispersal, the terrestrial earthworm fauna of Australia contains numerous endemic elements showing strong Gondwanan links (Lee 1994). Recent revisions of many genera, in particular of the family Megascolecidae (e.g. Jamieson 1971, 1974, 1994, and citations within; Blakemore 1998), suggest that most native earthworm genera consist entirely of SREs, but that much detailed fieldwork and mapping is necessary to elucidate the fauna fully. Phylum ONYCHOPHORA Order ONYCHOPHORA Remarks Velvet worms, or onychophorans, are primitive soft-bodied relatives of arthropods. The Australian fauna consists solely of the Peripatopsidae, which also occurs in New Guinea, New Zealand, South America and southern Africa (Reid 1996). Prior to 1985 only six species were known from Australia (Tait et al. 1990). Subsequent revisionary treatments (Ruhberg 1985; Reid 1996, 2000a, 2000b; Tait and Norman 2001) have added nearly 70 new species, and additional, undescribed species are known (A.L. Reid, personal

communication). Each of the 32 named Australian peripatopsid genera occur in clearly proscribed ranges with very few disjunctions. The most notable break is exhibited by members of the genus Nodocapitus Reid, which occur in northern New South Wales, south-eastern Queensland (N. barryi Reid and N. inornatus Reid) and in mid-eastern Queensland (N. formosus Reid). Named Australian onychophorans are usually restricted to a discrete area with very few being known from more than 200 km2. The most widely distributed species appears to be Occiperipatoides gilesii (Spencer), which occurs throughout the Darling Range on the eastern outskirts of Perth, Western Australia, with occasional outlying populations on the low-lying Swan Coastal Plain. Onychophoran species tend to inhabit permanently moist habitats, usually in native forests, and are most commonly found in or under rotting logs. High molecular and chromosomal divergence occurs within and between populations usually with concordant morphological divergence (Reid et al. 1995). Three species of Cephalofovea that were found within the same log have been shown to have 8186% fixed gene differences (Reid et al. 1995). Reid et al. (1995) and Gleeson et al. (1998) suggest that the high levels of genetic divergence within the Australasian Peripatopsidae is the result of ancient radiations within the Australasian fauna, with some lineages pre-dating the separation of New Zealand and Australia. Onychophorans thus exhibit one of the most extreme forms of short-range endemism, with some species restricted to single localities and with high genetic differentiation that indicates poor mobility and a strong reliance upon permanently moist habitat for survival. Phylum ARTHROPODA Class ARACHNIDA Order ARANEAE Remarks The ability of many spiders to balloon has perpetuated notions that spiders are poor candidates for any serious biogeographic treatment (Platnick 1981). Indeed, many spider families and genera contain species that are widely distributed across the Australian landscape, indicating an ability to disperse that precludes them from short-range endemic patterns. This can be confirmed from examination of distributional data presented in recently published taxonomic revisions (e.g. Baehr and Baehr 1987, 1998; Forster et al. 1987; Platnick and Forster 1989; Forster et al. 1990; Gray 1994; Harvey 1995; Platnick 2000; Huber 2001). Nevertheless, members of some mygalomorph taxa exhibit patterns of short-range endemism, although not quite on the scale found in other invertebrate orders. Raven (e.g. Raven 1982, 1984a, 1994) has shown that some mygalomorph

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genera consist solely of short-range endemic species, with many restricted to habitat isolates such as rainforest patches. Order SCHIZOMIDA Remarks The arachnid order Schizomida consists of some 200 species in 37 genera (Harvey 2002). These small, fast-moving animals are primarily restricted to tropical and sub-tropical forests where they occur in leaf litter, under stones and logs or within caves. Since the late 1980s, 46 schizomid species have been described from northern Australia in seven genera (Harvey 1988, 1992, 2000a, 2000b, 2001; Harvey and Humphreys 1995). Several are known from cave ecosystems or their entrances, whereas the remainder are found in closed forests or nearby habitats. Although most species have been found at single localities, this does not appear to represent a collecting artefact because highly localised species are replaced in nearby suitable habitats by different species. Several species occur over more than 100 km2. Brignolizomus woodwardi (Harvey) has been recorded from several localities in south-eastern Queensland over an area of c. 1000 km2 but the acquisition of some recently collected specimens, courtesy of Mr Michael Rix, indicates that the genus Brignolizomus represents a complex array of putative species with much smaller ranges. On the other side of the country, the highly troglobitic Draculoides vinei (Harvey) from Cape Range peninsula in Western Australia, occurs in a network of caves and other subterranean voids situated in limestone formations. The schizomid occurs over a total surface area of some 100 km2 but allozyme data suggests that the populations are not panmictic and that the levels of genetic divergence may represent total or incipient speciation (Adams and Humphreys 1993). Three other congeneric species plus two species of Bamazomus occur in coastal limestone on the periphery of the range of D. vinei each from highly restricted isolated areas (Harvey 2001). A seventh species has been found recently in coastal limestone, but females are required to determine its systematic position. The rainforest-dwelling genus Notozomus from north-eastern Queensland was recently shown to be highly diverse at the species level (Harvey 2000b), with no sympatry between congeneric species. Such patterns are similar to those found in other invertebrate taxa occurring in the wet tropics (e.g. Hill 1984; Raven 1984b; Baehr 1995; Monteith 1997) and presumably result from similar vicariance events in the formation of the forests (Joseph et al. 1995; Stuart-Fox et al. 2001). Only two Notozomus species possess ranges greater than 50 km2. Notozomus rentzi occurs on the Atherton tableland south-west of Cairns and N. ingham occurs on Hinchinbrook Island and at three sites on the mainland, although Harvey (2000b) was unable to confirm whether the males from Hinchinbrook Island and the females from the Ingham region were conspecific. Some

Table 1. Species of Notozomus within the Wet Tropics of northern Queensland assigned to mountain rainforest zones (Monteith 1997) Only two species (denoted with asterisks) are found in more than one zone. Notozomus curiosus Harvey from Mission Beach is not found in a mountain rainforest zone and may be misplaced in the genus. Mountain rainforest zones 1 2 3 4 5 6 7 8/9 10 11 12/13 14 15 16 17 18 19 Mt Finnigan Thornton Peak Windsor Tableland Carbine Tableland Hann Tableland Black Mtn Lamb Range Walsh/Hugh Nelson Range Atherton Tableland Mt Bellenden Ker Malbon Thompson Range Walter Hill Range Kirrima/Cardwell Range Seaview Range Hinchinbrook Island Paluma/Bluewater Range Mt Elliott Species of Notozomus N. daviesae, N. wudjl N. aterpes N. majesticus N. monteithi N. maurophila

N. rentzi* N. ker, N. rentzi* N. elongatus N. raveni N. ingham* N. ingham* N. ingham* N. spec

rainforest zones within the Wet Tropics (Monteith 1997) currently lack records of schizomids (Hann Tableland, Lamb Range, Walsh/Hugh Nelson Range and Mt Elliott; see Table 1) and it can be predicted that species of Notozomus will eventually be found there as well. The peculiar N. curiosus Harvey, which was not confidently assigned to the genus Notozomus by Harvey (2000b), occurs at Mission Beach, a coastal site in the Wet Tropics region that lies outside of the mountain rainforest zones identified by Monteith (1997). Only one other schizomid occurs in the area, Julatennius lawrencei Harvey from Julatten in Carbine Tableland. Few other animal groups are known to exhibit such extreme isolation and speciation in the tropical rainforests of Queensland, despite the considerable recent attention given to the invertebrate and vertebrate fauna of the region. The vast majority of known Australian schizomid species are indisputably SREs, with most known from single localities. Those species with ranges greater than 50 km2 have somewhat doubtful identities, with insufficient material available of Brignolizomus woodwardi and Notozomus ingham to resolve the species-level taxonomy and Draculoides vinei populations showing marked genetic divergence. Class DIPLOPODA Remarks Millipedes are one of the most poorly understood and little-studied groups in the terrestrial fauna in Australia. This

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is despite their high levels of diversity at the ordinal level (Harvey and Yen 1989; Black 1997) and their abundance in soil and leaf litter habitats. The bulk of the published taxonomic studies have been based on limited opportunistic collections. Few monographic treatments of proscribed taxa based on a wide range of specimens have been published, making biogeographic conclusions somewhat difficult. Mesibov (1994, 1997, 1999) reported upon the distributions of selected Tasmanian species, noting their highly restricted and often disjunct distributions. Shear and Mesibov (1997) presented a review of the Australian members of the chordeumatidan family, Metopidiotrichidae, which was found to consist of 18 species in five genera distributed in eastern and south-western Australia. Most species were found to possess short ranges with the most widespread, Australeuma jeekeli Golovatch, ranging over most of eastern Tasmania. Humphreys and Shear (1993) and Shear and Humphreys (1996) studied the genus Stygiochiropus Humphreys & Shear, which consists of four species from the caves and other subterranean voids of Cape Range peninsula, Western Australia. Three species are known only from single caves, S. isolatus Humphreys & Shear from cave C-222, S. sympatricus Humphreys & Shear from cave C-111, and S. peculiaris Shear & Humphreys from Camerons Cave (C-452) near Exmouth, whereas S. communis Humphreys & Shear is widespread throughout the peninsula, occupying some 400 km2, but is sympatric with S. sympatricus in cave C-111. The genetic structuring determined by allozyme electrophoresis divided S. communis into three subregions that largely correlate with the deep gorges that intersect the Tulki limestone of the region (Humphreys and Shear 1993). Recent research (M. S. Harvey and P. L. J. West, unpublished data) into the paradoxosomatid genus, Antichiropus Attems, has shown a bewildering array of taxa, most of which possess extremely short ranges. Attems (1911), Jeekel (1982) and Shear (1992) described nine species from south-western Australia and South Australia, but these studies do little justice to the huge array of species found during surveys of terrestrial invertebrates in the region. To date, 90 species of Antichiropus have been found of which 81 are unnamed; however, this figure will surely climb higher because every field season (winter), additional species are found that have not been previously collected. Only two species possess ranges greater than 10,000 km2. Antichiropus variabilis Attems is a very common species on the Darling Range east of Perth with several isolated occurrences on the Swan Coastal Plain and specimens from as far south as Manjimup and Forest Grove. The main core of the distribution occupies c. 15,000 km2, but this is doubled when the outlying populations are considered. Antichiropus sp. PM1 occurs throughout the length of the northern portion of the wheat belt with a total area of occupancy of 28,000 km2. The remaining species are known from single

sites or have ranges up to c. 5000 km2. The factors that govern such wide-scale speciation within such a small area are poorly understood, but the lack of mobility of both juvenile and adult millipedes must be an overriding factor. In addition, the extremely seasonal life cycle of Antichiropus millipedes suggests that dispersal is limited, as they are only ever active on the surface of soil and litter during winter after suitable rain has moistened the surrounding environment this allows males and females to emerge to the surface and begin mating. Males die off soon after mating and females burrow down into the soil to lay their eggs. On hatching, the juvenile millipedes remain within soil and leaf litter habitats until the next winter. It is also possible that different species of Antichiropus millipedes are restricted to particular soil types but these data have not yet been collected or tested. Similar biogeographic studies on other Australian millipede genera will undoubtedly detect comparable results showing widespread allopatric speciation and extremely short ranges, especially in areas where soil and vegetation types vary considerably across the landscape. For example, Main et al. (2002) have documented the distribution of the sole Western Australian member of the order Sphaerotheriida, Cynotelopus notabilis Jeekel, which is restricted to tall forests in the high-rainfall zone along the south coast of Western Australia. Undoubtedly, other sphaerotheriids in eastern Australia will possess similar small ranges in suitable habitats.

Class CRUSTACEA Order ISOPODA Suborder PHREATOICIDEA Remarks Members of the freshwater isopod group Phreatoicidea occur only in permanent fresh waters such as lakes and springs, although some are semi-terrestrial and inhabit permanently moist habitats under stones (see Wilson and Keable 2002). Phreatoicideans are an ancient group and a freshwater species is known from the Triassic (Wilson and Johnson 1999). Three families are found in Australia, Phreatoicidae, Amphisopodidae and Hypsimetodidae, and most species have highly restricted distributions and are constrained by their narrow habitat requirements (Wilson and Johnson 1999). Wilson and Johnson (1999) summarised the known distribution of the Australian phreatoicideans based on a taxonomic inventory and some literature records. The vast majority of genera were found to be highly restricted and often allopatrically distributed. Furthermore, apart from the records of Eophreatoicus from Arnhem Land in the Northern Territory, all species are restricted to land units that have been above sea level since the middle Cretaceous, indicating that their dispersal capabilities are poor.

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Order DECAPODA Family PARASTACIDAE

Table 2. Genus

Genera of Australian Parastacidae with numbers of short-range endemic species (SRE) No. of No. of No. of SRE species widespread (<10,000 km2) species 005 002 014 001 035 042 022 003 001 125 00 00 00 00 04 07 09 03 01 24 005 002 014 001 031 035 013 000 000 101 % SRE

Remarks The freshwater crayfish of Australia are arguably the best-known non-insect group occurring in Australian freshwater ecosystems, with 115 named species in nine genera (Crandall et al. 1999). Eight of these genera are endemic to Australia and only Cherax extends further afield into southern New Guinea and associated islands. The other Australian genera are Engaewa, endemic to south-western Australia, Parastacoides and Astacopsis endemic to Tasmania, Engaeus, Geocherax and Gramastacus found in south-eastern Australia and Tenuibranchiurus restricted to a small area of south-eastern Queensland and north-eastern New South Wales. Euastacus is found throughout mainland eastern Australia and Cherax is widespread throughout eastern and south-western Australia (Crandall et al. 1999). Crandall et al. (1999) utilised DNA sequence data of the 16S region to propose a phylogeny for the Australasian Parastacidae and presented unrooted trees showing three major clades, one consisting of Engaewa, the second containing Cherax, Gramastacus, Engaeus, Geocherax and Tenuibranchiurus and the third containing Astacopsis, Euastacus, Paranephrops (from New Zealand) and Parastacoides. The species-level taxonomy of various genera within the Parastacidae has been the subject of numerous recent studies (see references in Crandall et al. 1999) that have greatly enhanced the initial efforts of earlier authors (e.g. Clark 1936; Riek 1969, 1972). Some studies have been conducted utilising only morphological characters, but Austin (1996), Avery and Austin (1997), Hansen et al. 2001 and Austin and Ryan (2002) have warned of the danger in relying exclusively on morphological data in the recognition of parastacid species, especially because morphological plasticity has resulted in differing phenotypes in different ecological settings. Nevertheless, these studies indicate that the Australian freshwater crayfish fauna largely consists of a range of short-range endemic species. Of the 124 recognised species, only 24 (19%) have ranges greater than 10,000 km2 (Table 2). Of these species, 16 belong to the genera Cherax or Euastacus and are generally larger, more mobile species with relatively broad ecological preferences and life history traits. The remaining 101 species (81%) possess narrow ranges of less than 10,000 km2 (Table 2). Four genera, Engaewa, Geocherax, Parastacoides and Tenuibranchiurus, consist entirely of SREs, whereas two others, Engaeus and Euastacus, have a preponderance of SREs with 89% and 83% respectively. Horwitz and Adams (2000) found that all species of Engaewa from coastal creek systems of south-western Australia were highly restricted in distribution, with the greatest area of occupancy of any
Engaewa Geocherax ParastacoidesA Tenuibranchiurus Engaeus EuastacusB Cherax Astacopsis Gramastacus Total
A

100 100 100 100 089 083 058 000 000 082

Including the eight unnamed species recognised by Hansen et al. (2001) and Hansen and Richardson (2002). B Avery and Austin (1997) failed to find any electrophoretic data to support the separation of E. bispinosus and E. yarraensis from E. armatus.

species being only a few hundred square kilometres. Three of these species were highly localised with E. reducta Riek and E. walpolea Horwitz & Adams occurring in areas of less than 100 km2 and E. pseudoreducta Horwitz & Adams known from only a single swampy headwater that is severely modified from habitat loss and degradation. Gramastacus, with the sole species, G. insolitus Riek, occurs in permanent swamps and creeks in western Victoria and south-eastern South Australia (Zeidler and Adams 1990). Although the eastern and western populations were electrophoretically distinct, there were insufficient differences in the allozyme data to reject a hypothesis of more than one species. Gramastacus insolitus occurs over a total area of some 25,000 km2, but the area of occupancy is somewhat discontinuous, which is greatly exacerbated by recent land clearing and the draining of swamps and other permanent wetlands for agricultural purposes. The western Tasmanian genus Parastacoides has been the subject of several studies regarding its internal composition. In the most recent study, Hansen et al. (2001) used allozyme electrophoretic data and detected between 11 and 19 species, most of which are morphologically cryptic and have highly restricted distributions. Indeed, of the 14 species subsequently recognised in the paper, only six have distributions greater than 500 km2, with the remainder occupying much smaller areas. In contrast, the other endemic Tasmanian genus, Astacopsis, has not been shown to possess similar cryptic speciation patterns and small distributions (Hamr 1992). Indeed Hamr (1992) found only three species that were mostly parapatrically distributed, with A. franklinii (Gray) in the eastern half of Tasmania, A. tricornis Clark in western Tasmania and A. gouldi Clark across the north.

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Few of the 42 species of Euastacus currently recognised (Morgan 1986, 1988, 1989, 1997; Short and Davie 1993) have large ranges. The most widely distributed species, E. armatus, is found throughout the MurrayDarling system, E. spinifer occurs along a large portion of eastern New South Wales and E. yarraensis occurs throughout south-central Victoria. The remaining species are extremely localised SREs and often restricted to single mountain ranges in the wet tropics. Horwitz (1990) and Horwitz et al. (1990) recognised 34 species of Engaeus from south-eastern Australia and Tasmania. Several species were found to be relatively widespread but 25 species were found to be highly restricted. The SREs mostly occur in eastern Victoria and northern Tasmania. Members of the genus Cherax are widespread throughout eastern, northern and south-western Australia (Austin 1996; Austin and Knott 1996), and a further 13 species have been recognised from New Guinea (Crandall et al. 1999). Of the 22 named Australian species recognised by Austin (1996), Austin and Knott (1996) and Austin and Ryan (2002), nine appear to be relatively widespread with 13 potentially representing SREs, with one from south-western Australia and the other three from eastern or northern Australia. The factors that govern this high level of short-range endemism among parastacids include poor powers of dispersal, long life cycles, slow maturation rates and their persistence from the Cretaceous as evidenced by the presence of confamilial genera in other parts of Gondwana (New Zealand, Madagascar and South America; Crandall et al. 1999). Indeed the phylogenetic placement of the New Zealand genus Paranephrops within a clade that consists of three Australian genera (Crandall et al. 1999) suggests that the major radiation of the parastacid fauna occurred prior to the break up of Gondwana during the Cretaceous, because long-distance oceanic dispersal in these freshwater creatures is impossible. Discussion Short-range endemism occurs within many different animal groups in Australia, but there appear to be few groups in which short-range endemism is the norm. Generally speaking, most vertebrates appear to be too vagile to initiate SREs (as here defined) or to maintain divergence between fragmented populations that may eventually lead to short-range endemism. Some fish, frogs and reptiles possess naturally small ranges (see Cogger 1994; Allen et al. 2002), but the vast majority of vertebrates do not conform to the definition of an SRE applied here. Of the groups surveyed to date, only certain invertebrate groups appear to possess widespread and uniform short-range endemism. These include freshwater and terrestrial molluscs, onychophorans, millipedes, certain arachnids such as mygalomorph spiders and schizomids and some crustaceans such as freshwater

isopods and decapods. Short-range endemism occurs in other groups, but is not uniform throughout the taxon. Evidence for short-range endemism in many groups may be lacking owing to the paucity of reliable taxonomic treatments and insufficient sampling to determine correct identities and range sizes of individual species. The majority of taxa with high proportions of SREs possess similar ecological and life-history characteristics, notably poor powers of dispersal, confinement to discontinuous habitats, slow growth and low fecundity. Many appear to be Gondwanan relicts (see Hopper et al. 1996) that have persisted from a time when the environment was more uniformly mesic (Hill 1994) and when moist habitats were more evenly distributed throughout the landscape. The aridification of Australia, which commenced during the Miocene, led to a major contraction of suitable habitat for many taxa. This habitat only persists in large areas along the eastern seaboard and in the south-west of this country, and elsewhere has been replaced by xeric vegetation types. The available water in such areas has also diminished with less reliable rainfall, higher run-off and evaporation. It is no coincidence that the majority of Australian freshwater fish species with highly restricted distributions (Allen et al. 2002) occur in the central drainage systems that are disjunct from other river systems, allowing the formation of distinct fish populations that have diverged sufficiently over time to form different species. Conservation of short-range endemics Comprehensive systematic reviews of Australian faunal groups often reveal the presence of short-range endemic species. This systematic research sometimes leads to the declaration of the species as threatened or endangered under state or federal legislation. Although the listing of some invertebrate species represent authentic examples of taxa in peril, others are based on insufficient sampling effort and, occasionally, doubtful taxonomic judgement. The Otway stonefly, Eusthenia nothofagi Zwick (Plecoptera: Eustheniidae), was listed as presumed extinct in 1993 by the Victorian Department of Conservation and Natural Resources (CNR) owing to the lack of recently collected specimens available to specialists. A subsequent survey of numerous potential habitats in the Otway Ranges by Doeg and Reed (1995) demonstrated that E. nothofagi was widespread throughout the forested regions of the Otway Ranges in a wide variety of stream types. Similarly, the Dandenong amphipod, Austrogammarus australis (Sayce) was listed as presumed extinct by the CNR in 1993 because the most recent collection was in 1911 and a subsequent search for the species at the heavily modified type locality was unsuccessful. Detailed searches by Doeg (1997) throughout the Dandenong Ranges found A. australis at nine sites, particularly in the headwaters of streams with relatively low disturbance. The implications of these studies are

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profound. Although the listing of species under state or federal endangered species programs is laudable, several important criteria need to be met prior to listing, including recent and comprehensive surveys in suitable habitats, as advocated by Mawson and Majer (1999). Generally speaking, invertebrates do not feature highly in nature conservation, especially during the process of reserve selection (e.g. Ferrier et al. 1999; McKenzie et al. 2000). The selection of terrestrial habitats for inclusion in nature reserve systems is largely governed by factors such as vegetation type and percentage of intact vegetation (Ferrier 2002), which remain important and easily measured parameters. Nevertheless, such measures are poor indicators of invertebrate assemblages, which generally have high geographical turnover in species composition (see Ferrier et al. 1999). The inclusion of SREs in conservation planning, both within formal networks of parks and reserves and in conservation-worthy habitats on private or leasehold land, may give slightly different prominence to some regions over others. There is little evidence that SREs, particularly invertebrate examples, are playing any significant role in conservation planning, but notable exceptions have become apparent, especially using well-known iconic species such as butterflies (New and Sands 2002). Human-induced changes in the abundance and geographic range of species are actively creating SREs. It is from this pool of species that most threatened species are selected for listing by state, national and international authorities. Particular attention must be given to the taxa listed above in conservation planning, because habitat loss and degradation will further worsen their prospects for long-term survival. The conservation of hot-spots displaying high concordance of SREs will ensure that the maximum number of such taxa is preserved, along with the underlying ecological processes that initially assisted in the formation or retention of the species (see review by Moritz 2002). Acknowledgments I am extremely grateful to Bill Humphreys, David Yeates and Bob Mesibov for their comments on the manuscript, and to Barry Hutchins for assistance with the list of freshwater fish orders. I also wish to thank Robin Wilson, Elizabeth James and the participants of the Short-Range Endemism in the Australian Biota symposium held as part of the joint meeting in Melbourne of the Society of Australian Systematic Biologists and the Australasian Evolution Society during July 2001, of which this paper formed a part. References
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Reid, A. L. (1996). Review of the Peripatopsidae (Onychophora) in Australia, with comments on peripatopsid relationships. Invertebrate Taxonomy 10, 663936. Reid, A. L. (2000a). Eight new Planipapillus (Onychophora: Peripatopsidae) from southeastern Australia. Proceedings of the Linnean Society of New South Wales 122, 132. Reid, A. L. (2000b). Description of Lathropatus nemorum, gen. et sp. nov., and six new Ooperipatus Dendy (Onychophora: Peripatopsidae) from south-eastern Australia. Proceedings of the Royal Society of Victoria 112, 153184. Reid, A. L., Tait, N. N., and Briscoe, D. A. (1995). Morphological, cytogenetic and allozymic variation within Cephalofovea (Onychophora: Peripatopsidae) with descriptions of three new species. Zoological Journal of the Linnean Society 114, 115138. Riek, E. F. (1969). The Australian freshwater crayfish (Crustacea: Decapoda: Parastacidae), with descriptions of new species. Australian Journal of Zoology 17, 855918. Riek, E. F. (1972). The phylogeny of the Parastacidae (Crustacea: Astacoidea), and description of a new genus of Australian freshwater crayfishes. Australian Journal of Zoology 20, 369389. Ruhberg, H. (1985). Die Peripatopsidae (Onychophora). Systematik, Okologie, Chorologie und phylogenetische Aspekte. Zoologica, Stuttgart 137, 1184. Shear, W. A. (1992). A new genus and two new species of millipedes from the Cape Range, Western Australia (Diplopoda, Polydesmida, Paradoxosomatidae). Records of the Western Australian Museum 15, 777784. Shear, W. A., and Humphreys, W. F. (1996). A new Stygiochiropus from a North West Cape (Western Australia) coastal plain cave (Diplopoda: Polydesmida: Paradoxosomatidae). Records of the Western Australian Museum 17, 447449. Shear, W. A., and Mesibov, R. (1997). Australian chordeumatidan millipedes. III. A review of the milliped family Metopiotrichidae Attems in Australia (Diplopoda: Chordeumatida). Invertebrate Taxonomy 11, 141178. Short, J. W., and Davie, P. J. F. (1993). Two species of freshwater crayfish (Crustacea: Decapoda: Parastacidae) from northeast Queensland rainforest. Memoirs of the Queensland Museum 34, 6980. Smith, B. J. (1992). Non-marine Mollusca. In Zoological Catalogue of Australia. Volume 8. (Ed. W. W. K. Houston.) pp. 1405. (Australian Government Publishing Service: Canberra.) Smith, G. B. (1998). Review of the Australian Nicoletiinae (Zygentoma:Nicoletiidae). Invertebrate Taxonomy 12, 135189. Smithers, C. N. (1996). Psocoptera. In Zoological Catalogue of Australia. Volume 26. Psocoptera, Phthiraptera, Thysanoptera. (Eds C. N. Smithers, R. L. Palma, S. C. Barker and L. A. Mound.) pp. 179. (CSIRO Publishing: Melbourne.) Solem, A. (1997). Camaenid land snails from Western and central Australia (Mollusca: Pulmonata: Camaenidae). Records of the Western Australian Museum, Supplement 50, 14611906. Strahan, R. (Ed.) (1991). The Australian Museum Complete Book of Australian Mammals. (Angus and Robertson: Sydney.) Stuart-Fox, D. M., Schneider, C. J., Moritz, C., and Couper, P. J. (2001). Comparative phylogeography of three rainforest-restricted lizards from mid-east Queensland. Australian Journal of Zoology 49, 119127. Sturm, H., and Smith, G. B. (1993). New bristle tails (Archaeognatah: Meinertiellidae) from Australia. Journal of the Australian Entomological Society 32, 233240. Tait, N. N., and Norman, J. M. (2001). Novel mating behaviour in Florelliceps stutchburyae gen. nov., sp. nov. (Onychophora: Peripatopsidae) from Australia. Journal of Zoology, London 253, 301308.

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Manuscript received 8 March 2002; revised and accepted 25 June 2002.

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Appendix 1. SRE status

Summary of short-range endemism in the major groups of Australian terrestrial and freshwater animals, excluding parasites

Phylum

Class

Order

Platyhelminthes Nemertea Nematomorpha

Turbellaria Enopla Gordioida

Mollusca

Bivalvia

Short-range endemism in Australia

Gastropoda

Tricladida Hoplonemertea Chordodea Gordea Unionoida Veneroida Architaenioglossa Sorbeoconcha Systellommatophora

Annelida

Oligochaeta

Basommatophora Eupulmonata Haplotaxida

Tubificida

Tardigrada

Hirudinea Tardigrada

Onychophora Arthropoda

Onychophora Arachnida

Arhynchobdellida Eutardigrada Heterotardigrada Onychophora Scorpiones

Araneae Amblypygi

Schizomida Pseudoscorpiones Opiliones

Palpigradi

Branchiopoda

Remipedia

Acari Notostraca Anostraca Cladocera Conchostraca Nectiopoda

Ostracoda

Podocopida

Unknown, but extremely likely. Unknown, but extremely likely. Unknown. Unknown. Some SRE (Smith 1992). None (Smith 1992). Many SRE (Smith 1992). Numerous SRE (Smith 1992; Ponder et al. 1993; Miller et al. 1999). The sole indigenous Australian member of this order, Prisma prismatica (Tapparone-Canefri), is widespread in eastern Australia and New Guinea (Smith 1992). The few indigenous Australian basommatophorans are mostly widespread species (Smith 1992). Numerous SRE (see text). Many indigenous terrestrial earthworms are extremely localised and most represent SRE (e.g. Jamieson 1971, 1974, 1994; Blakemore 1998) (see text). Pinder and Brinkhurst (1997) have shown that the Australian phreodrilids consist of some localised species and some widespread species. Unknown. Unknown, but unlikely. Unknown, but unlikely. Extensive, almost ubiquitous, presence of SRE (see text). The current taxonomic arrangement (Koch 1977) includes many species-complexes that once studied will eventually increase the size of the fauna extensively. Some of these species may then qualify as SREs (E. S. Volschenk, personal communication). Some SRE, mostly amongst the Mygalomorphae (see text). One species widespread in Queensland rainforests, two species known only from single localities (Harvey and West 1998). Extensive, almost ubiquitous, presence of SRE (see text). Very few (e.g. Harvey 1998), except for some troglobites. Many harvestmen possess small ranges (e.g. Hunt 1985, 1992, 1993, 1995; Hunt and Cokendolpher 1991), but often larger than SRE threshold of 10,000 km2. The known Australian fauna only consists of two introduced species (Walter and Cond 1997), plus a single specimen of a presumably endemic species from north-western Australia (W. F. Humphreys, personal communication). Some SRE. None. None. Some SRE. Unknown. The sole Australian remipede, Lasionectes exleyi Yager and Humphreys, is restricted to a single anchialine locality in Western Australia (Yager and Humphreys 1996). Few SRE. (continued next page)

567

Appendix 1.

(continued)

568

Phylum

Class

Order

SRE status

Maxillipoda Malacostraca

Copepoda Spelaeogriphacea

Thermosbaenacea

Few SRE. The sole Australian spelaeogriphacean, Mangkurtu mityula Poore and Humphreys, is restricted to aquatic stygal habitats in Western Australia (Poore and Humphreys 1998). The sole Australian thermosbaenacean, Halosbaena tulki Poore and Humphreys, is restricted to aquatic stygal habitats in the Pilbara region of Western Australia (Poore and Humphreys 1992).

Mysidacea Anaspidacea Isopoda

Amphipoda

Chilopoda

Decapoda Scolopendrida Geophilida Lithobiida Craterostigmatida

Symphyla Diplopoda

Scutigerigida Symphyla Polyxenida Sphaerotheriida

All Phreatoicidea are SRE (Wilson and Johnson 1999). Many indigenous slaters (Oniscoidea) will most likely be found to represent SRE. The Australian terrestrial amphipod fauna contains some localised species, but very few that possess distributions of less that 10,000 km2 (Friend 1982, 1987). Extensive presence of SRE, especially amongst Parastacidae (see text). None. Some SRE but Australian fauna poorly known. Few SRE. The sole Australian species, Craterostigmus tasmanianus Pocock, is restricted to Tasmania, where it is widespread (Mesibov 1995). Unknown, but unlikely. Unknown. Unknown, but unlikely. Unknown, but many of these large rolling millipedes will most likely be found to represent SRE, like Cynotelopus notabilis Jeekel (Main et al. 2002). Unknown, but many of polydesmids will most likely be found to represent SRE Unknown None (all julids in Australia are introduced species). Many SRE (see Shear and Mesibov 1997).

Pauropoda Collembola Protura Diplura Insecta

Polydesmida Spirobolida Julida Chordeumatida (= Craspedosomatida) Spirostreptida Siphonophorida Polyzoniida Pauropoda Collembola Protura Diplura Archaeognatha Thysanura Odonata

M. S. Harvey

Ephemeroptera Blattodea Isoptera Mantodea Dermaptera Plecoptera

Unknown. Unknown. Unknown. Unknown. Many species appear to be widespread, but others (e.g. many Neanuridae) appear to be SRE (e.g. Greenslade 1994). Few SRE (Houston 1994b). Many species appear to be widespread, but others appear to be SRE (e.g. Houston 1994a). Many species with small ranges (Sturm and Smith 1993). Smith (1998) found that Australian nicoletiids possessed small ranges. Most dragonflies and damselflies are strong flyers and many species are widely distributed. Even the sole species assessed by Hawking (1999) as Critically Endangered, Archaeophya adamsi Fraser (Gomphomacromiidae) was previously known from mid-east Queensland to mid-coastal New South Wales (Hawking 1999). Few mayfly species are widespread, and many appear to be locally distributed; thus some may eventually represent SRE. Nil, except for some troglobites. None. None. None. Many stoneflies have restricted ranges, but others are relatively widespread. (continued next page)

Appendix 1. SRE status

(continued)

Short-range endemism in Australia

Phylum

Class

Order

Orthoptera Phasmatodea Embioptera Hemiptera Thysanoptera Psocoptera Phthiraptera Megaloptera

Neuroptera Coleoptera Strepsiptera

Chordata

Cephalaspidomorphi Chondrichthyes Actinopterygi

Mecoptera Siphonaptera Diptera Trichoptera Lepidoptera Hymenoptera Petromyzontiformes Carchariniformes Anguilliformes Atheriniformes

Beloniformes Clupeiformes Cyprinodontiformes Osteoglossiformes Perciformes

Pleuronectiformes Salmoniformes Scorpaeniformes Siluriformes

Synbranchiformes

Sarcopterygii Amphibia Reptilia

Ceratodontiformes Anura Testudines Squamata Crocodylia

Few. Few. Few. Few. Few species appear to possess restricted distributions (Mound 1996). Few species appear to possess restricted distributions (Smithers 1996). Few species appear to possess restricted distributions (Palma and Barker 1996). Some dobsonfly species are widespread, but many appear to be locally distributed (Theisinger and Houston 1988); thus some may eventually represent SRE. Few Australian lacewings appear to possess restricted distributions (New 1996a). Some. Many strepsipterans are known only from single localities (e.g. New 1996b), but there is little evidence that this represents the real situation. Most of the Australian species are relatively widespread (Lambkin 1996), with few, if any, SRE. Few Australian fleas appear to possess restricted distributions (Calder 1996). Few. Few. Few. Few. Of the three Australian species, one, Mordacia praecox Potter, may be an SRE (see Allen et al. 2002). None (see Allen et al. 2002). None (see Allen et al. 2002). Some species are highly restricted, including some hardyheads (Craterocephalus), and some rainbowfish (Melanotaeniidae) (see Allen et al. 2002). None (see Allen et al. 2002). None (see Allen et al. 2002). The only species present in Australian waters are introduced (see Allen et al. 2002). None (see Allen et al. 2002). A few Perciformes possess restricted distributions (see Allen et al. 2002), especially amongst the Gobiidae (Larson 2001). None (see Allen et al. 2002). The only Salmoniformes in Australia are introduced (see Allen et al. 2002). None (see Allen et al. 2002). The only geographically restricted Australian catfish is Neosilurus gloveri Allen and Feinberg from the Dalhousie Springs, South Australia (see Allen et al. 2002). Of the four species of swamp-eels present in Australia, only one, the Blind Cave Eel, Ophisternon candidum (Mees), is highly restricted (see Allen et al. 2002). The sole species, Ceratodontus forsteri Krefft, occurs in a restricted area of Queensland (see Allen et al. 2002). Few (Cogger 1994). None (Cogger 1994). Some (Cogger 1994). None (Cogger 1994). (continued next page)

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Appendix 1. SRE status

(continued)

Phylum

Class

Order

Aves

Mammalia

Anseriformes Apodiformes Caprimulgiformes Charadriiformes Ciconiiformes Columbiformes Coraciiformes Cuculiformes Falconiformes Galliformes Gruiformes Passeriformes Pelecaniformes Phoenicopteriformes Podicipediformes Procellariiformes Psittaciformes Sphenisciformes Strigiformes Struthioniformes Turniciformes Monotremata Dasyuromorphia Peramelomorphia Notoryctemorpha Diprotodontia Chiroptera Rodentia

None. None. None. None. None. None. None. None. None. None. None. None. None. None. None. None. None. None. None. None. None. None (Strahan 1991). None (Strahan 1991). None (Strahan 1991). None (Strahan 1991). None (Strahan 1991). None (Strahan 1991). None (Strahan 1991).

M. S. Harvey