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Invertebrate SystematicsCSIRO Publishing
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Collingwood, Vic. 3066, Australia
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Fax: +61 3 9662 7611
Email: [email protected]
Published by CSIRO Publishing
for CSIRO and the Australian Academy of Science
w w w . p u b l i s h . c s i r o . a u / j o u r n a l s / i s
InvertebrateSystematicsCon t i nu i ng I n v e r t e b r a t e Ta x o n o m y
Volume 16, 2002
CSIRO 2002
All enquiries and manuscripts should be directed to:
P u b l i s h i n g
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CSIRO 2002 10.1071/IS02009 1445-5226/02/040555
Invertebrate Systematics, 2002, 16, 555570
IS02009Shor t- rangeendemi smi nAus tr a li aM .S. Har vey
Short-range endemism among the Australian fauna: some examples
from non-marine environments
MarkS.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 oforganisms 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
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 cornersof 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 etal. (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 assesssuch taxa. This paper is designed to give a basic overview of
8/3/2019 Harvey 2002
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556 M. S. Harvey
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.
Geographiccoverage
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 actualarea 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).
Theextentofshort-rangeendemismintheAustralianfauna
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 etal. (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.
Ecosystemsthatmayinduce short range endemism
Terrestrial ecosystems in Australia represent a wide array of
environments that were categorised into a nationalframework 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 etal.
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. Hopperetal. (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 freshwaterecosystems 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 indistant 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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Short-range endemism in Australia 557
Freshwater molluscan radiations in Australia have been
well documented, particularly those of the hydrobiids in
south-eastern and south-western Australia (e.g. Ponderetal.
1993; Milleretal. 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 ofBeddomeia, 12 species ofPhrantela, three species
ofNanocochlea and four species ofVictodrobia (Ponderet
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 theAustralian 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
OrderHAPLOTAXIDA
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
OrderONYCHOPHORA
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 genusNodocapitus Reid, which occur in
northern New South Wales, south-eastern Queensland
(N. barryi Reid andN. 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 commonlyfound in or under rotting logs. High molecular and
chromosomal divergence occurs within and between
populations usually with concordant morphological
divergence (Reidetal. 1995). Three species ofCephalofovea
that were found within the same log have been shown to have
8186% fixed gene differences (Reidetal. 1995).
Reidetal. (1995) and Gleeson etal. (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 extremeforms 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
OrderARANEAE
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;
Forsteretal. 1987; Platnick and Forster 1989; Forsteretal.
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. Raven1982, 1984a, 1994) has shown that some mygalomorph
8/3/2019 Harvey 2002
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558 M. S. Harvey
genera consist solely of short-range endemic species, with
many restricted to habitat isolates such as rainforest patches.
OrderSCHIZOMIDA
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 beenfound 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.
Brignolizomuswoodwardi (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
genusBrignolizomus 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 ina 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 ofBamazomus occur in
coastal limestone on the periphery of the range ofD. 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 etal.
1995; Stuart-Fox etal. 2001). Only twoNotozomus species
possess ranges greater than 50 km2.Notozomusrentzi 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 andthe females from the Ingham region were conspecific. Some
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 ofNotozomuswill 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,Julatenniuslawrencei 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 speciesare 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 andlittle-studied groups in the terrestrial fauna in Australia. This
Table 1. Species ofNotozomus 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 onezone.Notozomus curiosus Harvey from Mission Beach is not found in
a mountain rainforest zone and may be misplaced in the genus.
Mountain rainforest zones Species of Notozomus
1 Mt Finnigan N. daviesae, N. wudjl
2 Thornton Peak N. aterpes
3 Windsor Tableland N. majesticus
4 Carbine Tableland N. monteithi
5 Hann Tableland
6 Black Mtn N. maurophila
7 Lamb Range
8/9 Walsh/Hugh Nelson Range
10 Atherton Tableland N. rentzi*
11 Mt Bellenden Ker N. ker, N. rentzi*12/13 Malbon Thompson Range N. elongatus
14 Walter Hill Range N. raveni
15 Kirrima/Cardwell Range N. ingham*
16 Seaview Range N. ingham*
17 Hinchinbrook Island N. ingham*
18 Paluma/Bluewater Range N. spec
19 Mt Elliott
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Short-range endemism in Australia 559
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, Australeumajeekeli 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 caveC-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 ofAntichiropus 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.
Antichiropusvariabilis 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 of28,000 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 ofAntichiropus
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
ofAntichiropus millipedes are restricted to particular soil
types but these data have not yet been collected or tested.
Similar biogeographic studies on other Australianmillipede 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 etal. (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
OrderISOPODA
SuborderPHREATOICIDEA
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 ofEophreatoicus 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 thattheir dispersal capabilities are poor.
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560 M. S. Harvey
OrderDECAPODA
Family PARASTACIDAE
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 andGramastacus found in
south-eastern Australia andTenuibranchiurus restricted to a
small area of south-eastern Queensland and north-easternNew South Wales.Euastacus is found throughout mainland
eastern Australia and Cherax is widespread throughout
eastern and south-western Australia (Crandall etal. 1999).
Crandall etal. (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 theParastacidae 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 orEuastacus 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
andTenuibranchiurus, consist entirely of SREs, whereas two
others, Engaeus andEuastacus, have a preponderance of
SREs with 89% and 83% respectively. Horwitz and Adams
(2000) found that all species ofEngaewa from coastal creek
systems of south-western Australia were highly restricted indistribution, with the greatest area of occupancy of any
species being only a few hundred square kilometres. Three of
these species were highly localised withE. reducta Riek and
E. walpolea Horwitz & Adams occurring in areas of less
than 100 km2 andE. 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. Gramastacusinsolitus 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 genusParastacoides has been the
subject of several studies regarding its internal composition.
In the most recent study, Hansen etal. (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 andA.gouldi Clarkacross the north.
Table 2. Genera of Australian Parastacidae with numbers of
short-range endemic species (SRE)
Genus No. of species
No. ofwidespread
species
No. of SRE(
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Short-range endemism in Australia 561
Few of the 42 species ofEuastacus 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 andE. 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 etal. (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 thepresence 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 genusParanephrops within a clade that consists of
three Australian genera (Crandall etal. 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 etal. 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 spidersand 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 etal.
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 suitablehabitat 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 etal.
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.
Conservationofshort-rangeendemics
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 foundA. australis at nine
sites, particularly in the headwaters of streams with relativelylow disturbance. The implications of these studies are
8/3/2019 Harvey 2002
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562 M. S. Harvey
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. Ferrieretal. 1999; McKenzie etal. 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 Ferrieret
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 attentionmust 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 toBarry 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
Adams, M., and Humphreys, W. F. (1993). Patterns of genetic diversity
within selected subterranean fauna of the Cape Range peninsula,
Western Australia: systematic and biogeographic implications.
Records of the Western Australian Museum, Supplement 45,
145164.
Allen, G. R., Midgley, S. H., and Allen, M. (2002). Field Guide to the
Freshwater Fishes of Australia. (CSIRO Publishing: Melbourne.)
Attems, C. G. (1911). Myriopoda exkl. Scolopendridae. In Die Fauna
Sdwest-Australiens. Volume 3. (Eds W. Michaelsen and R.Hartmeyer.) pp.147204. (Gustav Fischer: Jena.)
Austin, C. M. (1996). Systematics of the freshwater crayfish genus
Cherax Erichson (Decapoda: Parastacidae) in northern and eastern
Australia: electrophoretic and morphological variation.Australian
JournalofZoology44, 259296.
Austin, C. M., and Knott, B. (1996). Systematics of the freshwater
crayfish genus Cherax Erichson (Decapoda: Parastacidae) in
south-western Australia: electrophoretic, morphological and habitat
variation.AustralianJournalofZoology44, 223258.
Austin, C. M., and Ryan, S. G. (2002). Allozyme evidence for a new
species of freshwater crayfish of the genus Cherax Erichson
(Decapoda : Parastacidae) from the south-west of Western
Australia.InvertebrateSystematics16, 357367.
Avery, L., and Austin, C. M. (1997). A biochemical taxonomic study of
spiny crayfish of the genera Astacopsis andEuastacus (Decapoda:Parastacidae) in south-eastern Australia.MemoirsoftheMuseumof
Victoria56, 543555.
Baehr, B., and Baehr, M. (1987). The Australian Hersiliidae
(Arachnida: Araneae): taxonomy, phylogeny, zoogeography.
InvertebrateTaxonomy1, 351437.
Baehr, B., and Baehr, M. (1998). New species and new records of
Hersiliidae from Australia, with an updated key to all Australian
species (Arachnida: Araneae: Hersiliidae). Sixth supplement to the
revision of the Australian Hersiliidae. Records of the Western
AustralianMuseum19, 1338.
Baehr, M. (1995). Revision of Philipis (Colepotera: Carabidae:
Bembidiinae), a genus of arboreal tachyine beetles from the
rainforests of eastern Australia: taxonomy, phylogeny and
biogeography.MemoirsoftheQueenslandMuseum38, 315381.Beesley, P. L., Ross, G. J. B., and Wells, A. (1998). Mollusca: The
Southern Synthesis. (CSIRO Publishing: Melbourne.)
Black, D. (1997). Diversity and biogeography of Australian millipedes
(Diplopoda).MemoirsoftheMuseumofVictoria56, 557561.
Blakemore, R. (1998). Retrovescus, a new genus of opisthogastric
earthworm from Tasmania.InvertebrateTaxonomy12, 655665.
Burbidge, A. H., Harvey, M. S., and McKenzie, N. L. (2000).
Biodiversity of the southern Carnarvon Basin. Records of the
Western Australian Museum,Supplement61, 1595.
Calder, A. A. (1996). Siphonaptera. In Zoological Catalogue of
Australia. Volume 28. Neuroptera, Strepsiptera, Mecoptera,
Siphonaptera. (Eds T. R. New, K. J. Lambkin and A. A. Calder.) pp.
136181. (CSIRO Publishing: Melbourne.)
Clark, E. (1936). The freshwater crayfishes of Australia. Memoirsof
theNationalMuseumofVictoria12, 3140.Clark, S. A., and Richardson, B. J. (2002). Spatial analysis of genetic
variation as a rapid assessment tool in the conservation management
of narrow-range endemics.InvertebrateSystematics16, 583587.
Cogger, H. G. (1994). Reptiles and Amphibians of Australia. (Reed:
Chatswood.)
Crandall, K. A., Fetzner, J. W., Jr, Lawler, S. H., Kinnersley, M., and
Austin, C. M. (1999). Phylogenetic relationships among the
Australian and New Zealand genera of freshwater crayfishes
(Decapoda: Parastacidae). Australian Journal of Zoology 47,
199214.
Darwin, C. (1859). The Origin of Species by Means of Natural
Selection. (Murray: London.)
Davis, A. R., Roberts, D., and Ayre, D. J. (1999). Conservation of
sessile marine invertebrates: you do not know what you have got
until it is gone. In The Other 99%. The Conservation andBiodiversity of Invertebrates. (Eds W. F. Ponder and D. Lunney.)
8/3/2019 Harvey 2002
10/17
Short-range endemism in Australia 563
pp. 325329. (The Royal Zoological Society of New South Wales:
Sydney.)
Doeg, T. (1997). Gone today, here tomorrow extinct aquatic
macroinvertebrates in Victoria.MemoirsoftheMuseumofVictoria56, 531535.
Doeg, T., and Reed, J. (1995). Distribution of the endangered Otway
StoneflyEusthenianothofagi Zwick (Plecoptera: Eustheniidae) in
the Otway Ranges.ProceedingsoftheRoyalSocietyofVictoria107,
4550.
Ferrier, S. (2002). Mapping spatial pattern in biodiversity for regional
conservation planning: where to from here? SystematicBiology51,
331362.
Ferrier, S., Gray, M. R., Cassis, G. A., and Wilkie, L. (1999). Spatial
turnover in species composition of ground-dwelling arthropods,
vertebrates and vascular plants in north-east New South Wales:
implications for selection of forest reserves. In The Other 99%. The
Conservation and Biodiversity of Invertebrates. (Eds W. F. Ponder
and D. Lunney.) pp. 6876. (The Royal Zoological Society of New
South Wales: Sydney.)Forster, R. R., Platnick, N. I., and Gray, M. R. (1987). A review of the
spider superfamilies Hypochiloidea and Austrochiloidea (Araneae,
Araneomorphae). Bulletin of theAmericanMuseum ofNatural
History185, 1116.
Forster, R. R., Platnick, N. I., and Coddington, J. (1990). A proposal and
review of the spider family Synotaxidae (Araneae, Araneoidea),
with notes on theridiid interrelationships.BulletinoftheAmerican
MuseumofNaturalHistory193, 1116.
Friend, J. A. (1982). New terrestrial amphipods (Amphipoda:
Talitridae) from Australian forests. AustralianJournalofZoology
30, 461491.
Friend, J. A. (1987). The terrestrial amphipods (Amphipoda: Talitridae)
of Tasmania: systematics and zoogeography. Records of the
AustralianMuseum, Supplement7, 185.
Gleeson, D. M., Rowell, D. M., Tait, N. N., Briscoe, D. A., and Higgins,A. V. (1998). Phylogenetic relationships among Onychophora from
Australasia inferred from the mitochondrial cytochrome oxidase
subunit I gene. Molecular Phylogenetics and Evolution 10,
237248.
Gray, M. R. (1994). A review of the filistatid spiders (Araneae:
Filistatidae) of Australia. Records of theAustralianMuseum 46,
3961.
Greenslade, P. J. M. (1994). Collembola. In Zoological Catalogue of
Australia. Volume 22. Protura, Collembola, Diplura. (Eds W. W. K.
Houston and P. J. M. Greenslade.) pp. 19138. (CSIRO Publishing:
Melbourne.)
Hamr, P. (1992). A revision of the Tasmanian freshwater crayfish genus
Astacopsis Huxley (Decapoda: Parastacidae). Papers and
ProceedingsoftheRoyalSocietyofTasmania126, 9194.
Hansen, B., and Richardson, A. M. M. (2002). Geographic ranges,
sympatry and the influence of environmental factors on distribution
of species of an endemic Tasmanian freshwater crayfish.
InvertebrateSystematics16, 621629.
Hansen, B., Adams, M., Krasnicki, T., and Richardson, A. M. M.
(2001). Substantial allozyme diversity in the freshwater crayfish
Parastacoides tasmanicus supports extensive cryptic speciation.
InvertebrateTaxonomy15, 667679.
Harvey, M. S. (1988). A new troglobitic schizomid from Cape Range,
Western Australia (Chelicerata: Schizomida). Records of the
WesternAustralianMuseum14, 1520.
Harvey, M. S. (1992). The Schizomida (Chelicerata) of Australia.
InvertebrateTaxonomy6, 77129.
Harvey, M. S. (1995). The systematics of the spider family
Nicodamidae (Araneae : Amaurobioidea).InvertebrateTaxonomy
9, 279386.
Harvey, M. S. (1998). Pseudoscorpion groups with bipolar
distributions: a new genus from Tasmania related to the Holarctic
Syarinus (Arachnida, Pseudoscorpiones, Syarinidae). Journal of
Arachnology26, 429441.Harvey, M. S. (2000a). Brignolizomus and Attenuizomus, new
schizomid genera from Australia (Arachnida: Schizomida:
Hubbardiidae). Memorie della Societ Entomologica Italiana,
Supplemento78, 329338.
Harvey, M. S. (2000b). A review of the Australian schizomid genus
Notozomus (Hubbardiidae). Memoirs of the QueenslandMuseum
46, 161174.
Harvey, M. S. (2001). New cave-dwelling schizomids (Schizomida:
Hubbardiidae) from Australia. Recordsof theWesternAustralian
Museum, Supplement64, 171185.
Harvey, M. S. (2002). The neglected cousins: what do we know about
the smaller arachnid orders?JournalofArachnology30. (In press.)
Harvey, M. S., and Humphreys, W. F. (1995). Notes on the genus
Draculoides Harvey (Schizomida: Hubbardiidae), with the
description of a new troglobitic species. Records of the WesternAustralianMuseum, Supplement52, 183189.
Harvey, M. S., and West, P. L. J. (1998). New species of Charon
(Amblypygi, Charontidae) from northern Australia and Christmas
Island.JournalofArachnology26, 273284.
Harvey, M. S., and Yen, A. L. (1989). Worms to Wasps: an Illustrated
Guide to Australias Terrestrial Invertebrates. (Oxford University
Press: Melbourne.)
Hawking, J. H. (1999). An evaluation of the current conservation status
of Australian dragonflies (Odonata). In The Other 99%. The
Conservation and Biodiversity of Invertebrates. (Eds W. F. Ponder
and D. Lunney.) pp. 354360. (The Royal Zoological Society of
New South Wales: Sydney.)
Hill, L. (1984). New genera of Hypselosomatidae (Heteroptera:
Schizopteridae) from Australia. AustralianJournal of Zoology,
SupplementarySeries103, 155.Hill, R. S. (Ed.) (1994). History of the Australian Vegetation:
Cretaceous to Recent. (Cambridge University Press: Cambridge.)
Hooper, J., and Kennedy, J. (2002). Small-scale patterns of sponge
biodiversity (Porifera) on Sunshine Coast reefs, eastern Australia.
InvertebrateSystematics16, 637653.
Hopper, S. D., Harvey, M. S., Chappill, J. A., Main, A. R., and Main, B.
Y. (1996). The Western Australian biota as Gondwanan heritage a
review. In Gondwanan Heritage: Past, Present and Future of the
Western Australian Biota. (Eds S. D. Hopper, J. A. Chappill, M. S.
Harvey and A. S. George.) pp. 146. (Surrey Beatty & Sons:
Sydney.)
Horwitz, P. (1990). A taxonomic revision of species in the freshwater
crayfish genus Engaus Erichson (Decapoda : Parastacidae).
InvertebrateTaxonomy4, 427614.
Horwitz, P., and Adams, M. (2000). The systematics, biogeography and
conservation status of species in the freshwater crayfish genus
Engaewa Riek (Decapoda : Parastacidae) from south-western
Australia.InvertebrateTaxonomy14, 655680.
Horwitz, P., Adams, M., and Baverstock, P. (1990). Electrophoretic
contributions to the systematics of the freshwater crayfish genus
Engaus Erichson (Decapoda : Parastacidae).InvertebrateTaxonomy
4, 615641.
Houston, W. W. K. (1994a). Diplura. In Zoological Catalogue of
Australia. Volume 22. Protura, Collembola, Diplura. (Eds W. W. K.
Houston and P. J. M. Greenslade.) pp. 139156. (CSIRO Publishing:
Melbourne.)
Houston, W. W. K. (1994b). Protura. In Zoological Catalogue of
Australia. Volume 22. Protura, Collembola, Diplura. (Eds W. W. K.
Houston and P. J. M. Greenslade.) pp. 117. (CSIRO Publishing:
Melbourne.)
8/3/2019 Harvey 2002
11/17
564 M. S. Harvey
Huber, B. A. (2001). The pholcids of Australia (Araneae: Pholcidae):
taxonomy, biogeography, and relationships. Bulletin of the
AmericanMuseumofNaturalHistory260, 1144.
Humphreys, W. F. (1993). The significance of the subterranean fauna in biogeographical reconstruction: examples from Cape Range
peninsula, Western Australia.Records of the WesternAustralian
Museum, Supplement45, 165192.
Humphreys, W. F. (1999). Relict stygofaunas living in sea salt, karst and
calcrete habitats in arid northwestern Australia contain many
ancient lineages. In The Other 99%. The Conservation and
Biodiversity of Invertebrates. (Eds W. F. Ponder and D. Lunney.)
pp. 219227. (The Royal Zoological Society of New South Wales:
Sydney.)
Humphreys, W. F. (2001). Groundwater calcrete aquaifers in the
Australian arid zone: the context to an unfolding plethora of stygal
biodiversity. Records of the Western Australian Museum,
Supplement64, 6383.
Humphreys, W. F., and Shear, W. A. (1993). Troglobitic millipedes
(Diplopoda : Paradoxosomatidae) from semi-arid Cape Range,Western Australia: systematics and biology.InvertebrateTaxonomy
7, 173195.
Hunt, G. S. (1985). Taxonomy and distribution ofEquitius in eastern
Australia (Opiliones: Laniatores: Triaenonychidae).Recordsofthe
AustralianMuseum36, 107125.
Hunt, G. S. (1992). Revision of the genus Holonuncia Forster
(Arachnida: Opiliones: Triaenonychidae) with description of
cavernicolous and epigean species from eastern Australia.Records
oftheAustralianMuseum44, 135163.
Hunt, G. S. (1993). A revision of the genusLomanella Pocock and its
implication for family level classification in the Travunioidea
(Arachnida: Opiliones: Triaenonychidae).RecordsoftheAustralian
Museum45, 81119.
Hunt, G. S. (1995). Revision of the harvestman genus Miobunus from
Tasmania (Arachnida: Opiliones: Triaenonychidae).RecordsoftheWesternAustralianMuseum, Supplement52, 243252.
Hunt, G. S., and Cokendolpher, J. C. (1991). Ballarrinae, a new
subfamily of harvestmen from the southern hemisphere (Arachnida,
Opiliones, Neopilionidae). Recordsof theAustralianMuseum43,
131169.
Jamieson, B. G. M. (1971). Earthworms (Megascolecidae:
Oligochaeta) from Western Australia and their zoogeography.
JournalofZoology,London165, 471504.
Jamieson, B. G. M. (1974). The indigenous earthworms
(Megascolecidae: Oligochaeta) of Tasmania.BulletinoftheBritish
MuseumofNaturalHistory (Zoology) 26, 203328.
Jamieson, B. G. M. (1994). Some ear thworms from the wet tropics and
from Bunya Moutains, Queensland (Megascolecidae: Oligochaeta)
of Tasmania.MemoirsoftheQueenslandMuseum37, 157180.
Jeekel, C. A. W. (1982). Millipedes from Australia, 1: Antichiropodini
from South Australia (Diplopoda, Polydesmida,
Paradoxosomatidae).BulletinZoologischMuseum, Universiteitvan
Amsterdam8, 121132.
Joseph, L., Moritz, C., and Hugall, A. (1995). Molecular support for
vicariance as a source of diversity in rainforest.Proceedingsofthe
RoyalSocietyofLondonB260, 177182.
Koch, L. E. (1977). The taxonomy, geographic distribution and
evolutionary radiation of Australo-Papuan scorpions.Recordsofthe
WesternAustralianMuseum5, 83367.
Lambkin, K. J. (1996). Mecoptera. In Zoological Catalogue of
Australia. Volume 28. Neuroptera, Strepsiptera, Mecoptera,
Siphonaptera. (Eds T. R. New, K. J. Lambkin and A. A. Calder.) pp.
123135. (CSIRO Publishing: Melbourne.)
Larson, H. K. (2001). A revision of the gobiid fish genusMugilogobius
(Teleostei: Gobioidei), and its systematic placement.Recordsofthe
WesternAustralianMuseum, Supplement62, 1233.
Lee, K. E. (1994). Earthworm classification and biogeography:
Michaelsens contribution, with special reference to southern lands.
MitteilungenausdemHamburgischenZoologischenMuseumund
Institute89(2), 1121.Main, B. Y., Harvey, M. S., and Waldock, J. M. (2002). The distribution
of the Western Australian pill millipede, Cynotelopus notabilis
Jeekel (Sphaerotheriidae). Records of the Western Australian
Museum20, 383385.
Mawson, P. R., and Majer, J. D. (1999). The Western Australian
threatened species scientific committee: lessons from invertebrates.
In The Other 99%. The Conservation and Biodiversity of
Invertebrates. (Eds W. F. Ponder and D. Lunney.) pp. 369373. (The
Royal Zoological Society of New South Wales: Sydney.)
McKenzie, N. L., Johnston, R. B., and Kendrick, P. G. (Eds) (1991).
Kimberley Rainforests of Australia. (Surrey Beatty & Sons:
Sydney.)
McKenzie, N. L., Halse, S. A., and Gibson, N. (2000). Some gaps in the
reserve system of the southern Carnarvon Basin, Western Australia.
Records of the Western Australian Museum, Supplement 61,547567.
Mesibov, R. (1994). Faunal breaks in Tasmania and their significance
for invertebrate conservation.MemoirsoftheQueenslandMuseum
36, 133136.
Mesibov, R. (1995). Distribution and ecology of the centipede
Craterostigmus tasmanianus Pocock, 1902 (Chilopoda:
Craterostigmomorpha: Craterostigmidae) in Tasmania. Tasmanian
Naturalist117, 27.
Mesibov, R. (1997). A zoogeographical singularity at Weavers Creek,
Tasmania.MemoirsoftheMuseumofVictoria56, 563573.
Mesibov, R. (1999). The Mersey Break: an unexplained faunal
boundary on the north coast of Tasmania. In The Other 99%. The
Conservation and Biodiversity of Invertebrates. (Eds W. F. Ponder
and D. Lunney.) pp. 246252. (The Royal Zoological Society of
New South Wales: Sydney.)Miller, A. C., Ponder, W. F., and Clark, S. A. (1999). Freshwater snails
of the genera Fluvidona and Austropyrgus (Gastropoda :
Hydrobiidae) from northern New South Wales and southern
Queensland.InvertebrateTaxonomy13, 461489.
Monteith, G. M. (1997). Revision of the Australian flat bugs of the
subfamily Mezirinae (Insecta: Hemiptera: Aradidae). Memoirsof
theQueenslandMuseum41, 1169.
Morgan, G. J. (1986). Freshwater crayfish of the genusEuastacus Clark
(Decapoda, Parastacidae) from Victoria.MemoirsoftheMuseumof
Victoria47, 157.
Morgan, G. J. (1988). Freshwater crayfish of the genusEuastacus Clark
(Decapoda, Parastacidae) from Queensland. Memoirs of the
MuseumofVictoria49, 149.
Morgan, G. J. (1989). Two new species of the freshwater crayfish
Euastacus Clark (Decapoda, Parastacidae) from isolated high
country of Queensland. Memoirsof theQueenslandMuseum27,
555562.
Morgan, G. J. (1997). Freshwater crayfish of the genusEuastacus Clark
(Decapoda, Parastacidae) from New South Wales, with a key to all
species in the genus.RecordsoftheAustralianMuseum23, 1110.
Moritz, C. (2002). Strategies to protect biological diversity and the
evolutionary processes that sustain it. Systematic Biology 51,
238254.
Mound, L. A. (1996). Thysanoptera. 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. 249332. (CSIRO Publishing: Melbourne.)
New, T. R. (1996a). Neuroptera. In Zoological Catalogue of Australia.
Volume 28. Neuroptera, Strepsiptera, Mecoptera, Siphonaptera.(Eds T. R. New, K. J. Lambkin and A. A. Calder.) pp. 1104.
(CSIRO Publishing: Melbourne.)
8/3/2019 Harvey 2002
12/17
Short-range endemism in Australia 565
New, T. R. (1996b). Strepsiptera. In Zoological Catalogue of Australia.
Volume 28. Neuroptera, Strepsiptera, Mecoptera, Siphonaptera.
(Eds T. R. New, K. J. Lambkin and A. A. Calder.) pp. 105122.
(CSIRO Publishing: Melbourne.) New, T. R., and Sands, D. (2002). Narrow-range endemicity and
conservation status: interpretations for Australian butterflies.
InvertebrateSystematics16, 665670.
OHara, T. D. (2002). Endemism, rarity and vulnerability of marine
species along a temperate coastline. Invertebrate Systematics 16,
671684.
Palma, R. L., and Barker, S. C. (1996). Phthiraptera. 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. 81247. (CSIRO Publishing: Melbourne.)
Pantin, C. F. A. (1960). Alfred Russell Wallace: his pre-Darwinian
essay of 1855.ProceedingsoftheLinneanSocietyofLondon171,
139153.
Pinder, A. M., and Brinkhurst, R. O. (1997). Review of the
Phreodrilidae (Annelida:Oligochaeta:Tubificida) of Australia.InvertebrateTaxonomy11, 443523.
Platnick, N. I. (1981). Spider biogeography: past, present, and future.
RevueArachnologique3, 8595.
Platnick, N. I. (2000). A relimitation and revision of the Australasian
ground spider family Lamponidae (Araneae: Gnaphosoidea).
BulletinoftheAmericanMuseumofNaturalHistory254, 1330.
Platnick, N. I., and Forster, R. R. (1989). A revision of the temperate
South American and Australasian spiders of the family Anapidae
(Araneae, Araneoidea). Bulletin of the American Museum of
NaturalHistory190, 1139.
Ponder, W. (1999). Using museum collection data to assist in
biodiversity assessment. In The Other 99%. The Conservation and
Biodiversity of Invertebrates. (Eds W. F. Ponder and D. Lunney.)
pp. 253256. (The Royal Zoological Society of New South Wales:
Sydney.)Ponder, W. F., and Colgan, D. J. (2002). What makes a narrow-range
taxon? Insights from Australian freshwater snails. Invertebrate
Systematics16, 571582.
Ponder, W. F., and Lunney, D. (Eds) (1999). The Other 99%. The
Conservation and Biodiversity of Invertebrates. (The Royal
Zoological Society of New South Wales: Sydney.)
Ponder, W. F., Clark, G. A., Miller, A. C., and Toluzzi, A. (1993). On a
major radiation of freshwater snails in Tasmania and eastern
Victoria: a preliminary overview of the Beddomeia group
(Mollusca:Gastropoda:Hydrobiidae). Invertebrate Taxonomy 7,
501750.
Poore, G. C. B., and Humphreys, W. F. (1992). First record of
Thermosbaenacea (Crustacea) from the southern hemisphere: a new
species from a cave in tropical Western Australia. Invertebrate
Taxonomy6, 719725.
Poore, G. C. B., and Humphreys, W. F. (1998). The first record of
Spelaeogriphacea (Crustacea) from Australasia: a new genus and
species from an aquifer in the arid Pilbara of Western Australia.
Crustaceana71, 721742.
Raven, R. J. (1982). Systematics of the Australian mygalomorph spider
genus Ixamatus Simon (Diplurinae: Dipluridae: Chelicerata).
AustralianJournalofZoology30, 10351067.
Raven, R. J. (1984a). Systematics of the Australian curtain-web spiders
(Ischnothelinae: Dipluridae: Chelicerata). AustralianJournal of
Zoology, SupplementarySeries93, 1102.
Raven, R. J. (1984b). A new diplurid genus from eastern Australia and
a related Aname species (Diplurinae: Dipluridae: Araneae).
AustralianJournalofZoology, SupplementarySeries96, 151.
Raven, R. J. (1994). Mygalomorph spiders of the Barychelidae in
Australia and the western Pacific. Memoirs of the Queensland
Museum35, 291706.
Reid, A. L. (1996). Review of the Peripatopsidae (Onychophora) in
Australia, with comments on peripatopsid relationships.
InvertebrateTaxonomy10, 663936.
Reid, A. L. (2000a). Eight new Planipapillus (Onychophora:Peripatopsidae) from southeastern Australia. Proceedings of the
LinneanSocietyofNewSouthWales122, 132.
Reid, A. L. (2000b). Description ofLathropatusnemorum, gen. et sp.
nov., and six new Ooperipatus Dendy (Onychophora:
Peripatopsidae) from south-eastern Australia. Proceedings of the
RoyalSocietyofVictoria112, 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. ZoologicalJournaloftheLinneanSociety114, 115138.
Riek, E. F. (1969). The Australian freshwater crayfish (Crustacea:
Decapoda: Parastacidae), with descriptions of new species.
AustralianJournalofZoology17, 855918.
Riek, E. F. (1972). The phylogeny of the Parastacidae (Crustacea:
Astacoidea), and description of a new genus of Australian
freshwater crayfishes.AustralianJournalofZoology20, 369389.
Ruhberg, H. (1985). Die Peripatopsidae (Onychophora). Systematik,
Okologie, Chorologie und phylogenetische Aspekte. Zoologica,
Stuttgart137, 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 WesternAustralianMuseum
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
WesternAustralianMuseum17, 447449.
Shear, W. A., and Mesibov, R. (1997). Australian chordeumatidan
millipedes. III. A review of the milliped family MetopiotrichidaeAttems in Australia (Diplopoda: Chordeumatida). Invertebrate
Taxonomy11, 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 QueenslandMuseum 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). InvertebrateTaxonomy12, 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
WesternAustralianMuseum, Supplement50, 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. AustralianJournal of Zoology 49,
119127.
Sturm, H., and Smith, G. B. (1993). New bristle tails (Archaeognatah:
Meinertiellidae) from Australia. Journal of the Australian
EntomologicalSociety32, 233240.
Tait, N. N., and Norman, J. M. (2001). Novel mating behaviour in
Florelliceps stutchburyae gen. nov., sp. nov. (Onychophora:Peripatopsidae) from Australia. Journal ofZoology, London253,
301308.
8/3/2019 Harvey 2002
13/17
566 M. S. Harvey
http://www.publish.csiro.au/journals/is
Tait, N. N., Stutchbury, R. J., and Briscoe, D. A. (1990). Review of the
discovery and identification of Onychophora in Australia.
Proceedings of theLinnean Society ofNew South Wales 112,
153171.Thackway, R., and Cresswell, I. D. (Eds) (1995). An Interim
Biogeographic Regionalisation for Australia: a Framework for
Establishing the National System of Reserves. (Australian Nature
Conservation Agency: Canberra.)
Theisinger, G., and Houston, W. W. K. (1988). Megaloptera. In
Zoological Catalogue of Australia. Volume 6. Ephemeroptera,
Megaloptera, Odonata, Plecoptera, Trichoptera. (Eds I. Campbell,
G. Theischinger, W. W. K. Houston, J. A. L. Watson, F. B. Michaelis,
C. Yule and A. Neboiss.) pp. 2332. (Australian Government
Publishing Service: Canberra.)
Walter, D. E., and Cond, B. (1997). Eukoeneniaflorenciae Rucker,
1903 (Arachnida: Palpigradi: Eukoeneniidae), Australias second
record of a cosmopolitan all-female species of palpigrade.
AustralianEntomologist24, 164.
Wilson, G. D. F., and Johnson, R. T. (1999). Ancient endemism amongstfreshwater isopods (Crustacea, Phreatoicidea). In The Other 99%.
The Conservation and Biodiversity of Invertebrates.
(Eds W. F. Ponder and D. Lunney.) pp. 264268. (The Royal
Zoological Society of New South Wales: Sydney.)
Wilson, G. D. F., and Keable, S. J. (2002). New genera of Phreatoicidea
(Crustacea: Isopoda) from Western Australia. Records of the
AustralianMuseum54, 4170.
Yager, J., and Humphreys, W. F. (1996).Lasionectesexleyi, sp. nov., thefirst remipede crustacean recorded from Australia and the Indian
Ocean, with a key to the world species. InvertebrateTaxonomy10,
171187.
Yeates, D. K., Bouchard, P., and Monteith, G. B. (2002). Patterns and
levels of endemism in the Australian Wet Tropics rainforest:
evidence from flightless insects. Invertebrate Systematics 16,
605619.
Yen, A. (2002). Short-range endemism and the Australian Psylloidea
(Insecta : Hemiptera) in the genera Glycapsis and Acizzia
(Psyllidae).InvertebrateSystematics16, 631636.
Zeidler, W., and Adams, M. (1990). Revision of the Australian
crustacean genus of freshwater crayfish Gramastacus Riek
(Decapoda : Parastacidae).InvertebrateTaxonomy3, 913924.
Manuscript received 8 March 2002; revised and accepted 25 June 2002.
8/3/2019 Harvey 2002
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Appendix1
.
(continued)
Phylum
Class
Orde
r
SREstatus
Aves
Anseriformes
None.
Apodiformes
None.
Caprimulgiformes
None.
Charadriiformes
None.
Cico
niiformes
None.
Colu
mbiformes
None.
Cora
ciiformes
None.
Cuculiformes
None.
Falconiformes
None.
Galliformes
None.
Grui
formes
None.
Passeriformes
None.
Pelecaniformes
None.
Phoe
nicopteriformes
None.
Podicipediformes
None.
Procellariiformes
None.
Psittaciformes
None.
Sphe
nisciformes
None.
Strig
iformes
None.
Struthioniformes
None.
Turn
iciformes
None.
Mammalia
Mon
otremata
None(Strahan1991).
Dasy
uromorphia
None(Strahan1991).
Peramelomorphia
None(Strahan1991).
Noto
ryctemorpha
None(Strahan1991).
Diprotodontia
None(Strahan1991).
Chiroptera
None(Strahan1991).
Rodentia
None(Strahan1991).