Harvey 2002

8/3/2019 Harvey 2002

1/17

Invertebrate SystematicsCSIRO Publishing

PO Box 1139 (150 Oxford St)

Collingwood, Vic. 3066, Australia

Telephone: +61 3 9662 7629

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

8/3/2019 Harvey 2002

2/17

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

3/17

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).

8/3/2019 Harvey 2002

4/17

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

5/17

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

8/3/2019 Harvey 2002

6/17


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.

8/3/2019 Harvey 2002

7/17

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(

8/3/2019 Harvey 2002

8/17


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

9/17

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


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.


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.


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


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.


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).


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



Melbourne.)

Houston, W. W. K. (1994b). Protura. In Zoological Catalogue of



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.)


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


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


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


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.


(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.


(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


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.


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.)


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.


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


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


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

14/17

8/3/2019 Harvey 2002

15/17

8/3/2019 Harvey 2002

16/17

8/3/2019 Harvey 2002

17/17

570 M. S. Harvey

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).

Harvey 2002

Documents