J OURNAL OF C RUSTACEAN B IOLOGY, 32(4), 615-623, 2012
AN APPRAISAL OF THE DIVERSITY AND DISTRIBUTION OF LARGE
BRANCHIOPODS (BRANCHIOPODA: ANOSTRACA, LAEVICAUDATA,
SPINICAUDATA, CYCLESTHERIDA, NOTOSTRACA) IN AUSTRALIA
Brian V. Timms 1,2,∗
1 Honorary
2 Australian
Research Associate, Australian Museum, 6-9 College St, Sydney, NSW, 2000
Wetland and Rivers Centre, BEES, University of New South Wales, Sydney, 2052, Australia
ABSTRACT
The diversity of large branchiopods in Australia has not be reviewed since 1983 for fairy and brine shrimps (Anostraca), 2005 for
clam shrimps (Laevicaudata, Spinicaudata, Cyclestherida), and 1955 for shield shrimps (Notostraca). Presently five genera and 59
species of anostracans are known to occur in Australia (up from three genera and 29 species in 1983), with 35 described species
plus five known undescribed species of Branchinella, 18 described plus one known undescribed species of Parartemia, two described
species of Streptocephalus, two species of Australobranchipus, and two Artemia. Of these Branchinella clandestina and Streptocephalus
queenslandicus could be junior synonyms and hence invalid. A few species are widespread (B. australiensis, B. occidentalis, B. affinis), but
the majority are restricted to a state or two and many are localised. All species of Parartemia and a few Branchinella are halobionts, while
the remainder live in strictly fresh waters. Debate rages on whether one of the two international species of Artemia are native or introduced.
The degree of congeneric occurrences varies between genera and across the continent. The clam shrimp fauna is less well known with at
least two species of Lynceus, two of Eocyzicus, six of Caenestheria, two of Caenestheriella, an enigmatic species supposedly belonging
to Cyzicus, seven of Limnadia, two of Eulimnadia, eight of Limnadopsis, one of Eoleptestheria, and one of Cyclestheria. Two further
species of Lynceus, four of Limandia and four of Limnadopsis are being described. This makes 42 species, up from 26, but descriptive and
molecular work in progress suggest many more at least in the four genera being studied (Lynceus, Eocyzicus, Caenestheriella, Limnadia).
Eocyzicus parooensis and an undescribed species of Limnadia are halophilic. For the Notostraca, the supposedly widespread Triops
australiensis and Lepidurus apus viridis may represent many species separable molecularly.
K EY W ORDS: Artemia, Australobranchipus, Branchinella, Caenestheria, Caenestheriella, Cyzicus, Eocyzicus, Eoleptestheria, Lepidurus, Limnadia, Limnadopsis, Lynceus, Parartemia, Streptocephalus, Triops
DOI: 10.1163/193724012X634198
I NTRODUCTION
Studies on large branchiopods in Australia have lagged behind those in other inhabited continents. By 1900 only two
of its many fairy and brine shrimps had been described
though 12 of its clam shrimps were known. This imbalance was probably caused by the abundance of swamps with
clam shrimps near the fledging Sydney and Melbourne settlements and the paucity of fairy shrimps in eastern coastal
lowlands where most early settlers lived. The presence of an
Artemia in an early Sydney salt works had also been noted
and erroneously described as a new species (Timms, 2006).
Two notostracans were also found and described: Lepidurus
apus viridus (originally as L. viridus Baird 1850, also as its
synonym L. angasi Baird, 1866), and Triops australiensis
Spencer and Hall, 1896.
By 1940 the situation had not improved much for fairy
shrimps, though other groups were more intensively collected. Some collecting inland had added six fairy shrimps
(see Table 2), an endemic brine shrimp and two more erroneous descriptions of species of Artemia (Sayce, 1903),
nine more clam shrimp (Richter and Timms, 2005), and two
more Triops (later synonymised by Longhurst, 1955). An
∗ E-mail:
overlooked work during this period is a thesis by Richardson (1929) on clam shrimp in Western Australia (hereafter
WA). He accepted species described to that date and added
seven more, but none were formalised by publication and
now none of his specimens can be found; moreover, many of
his localities have been changed by settlement and no longer
support shrimp, so that these species have probably been lost
before they were formally named.
For the period 1940-2005, work continued on the clam
shrimp fauna with seven individual additions spaced over
the years (see Richter and Timms, 2005). At last, attention
was focused on anostracans but in just two publications: the
monograph of Linder (1941) added 13 species to the fauna,
almost all from WA; and the review of Geddes (1981) added
six species. Early in this period Longhurst (1955) simplified
notostracan taxonomy with his review, but his view of a
simple fauna in Australia of just two species is again being
challenged (see later).
During the most recent decade my colleagues and I have
tried to make amends for the past neglect of Australian large
branchiopods. Aided by intensive and extensive field work
to remote areas, 27 species (plus 5 undescribed taxa) of fairy
[email protected]
© The Crustacean Society, 2012. Published by Koninklijke Brill NV, Leiden
DOI:10.1163/193724012X634198
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JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 32, NO. 4, 2012
Table 1. Summary of number of species of Anostraca in Australia. *Brendonck et al. (2008) list species by family, but each family is represented by one
genus in Australia.
Genera
Artemia
Parartemia
Branchinella
Streptocephalus
Australobranchipus
Total
Linder
(1941)
Williams
(1980, 1981)
Geddes
(1983)
Brendonck et al.
(2008)*
Currently
recognised species
Known undescribed
species
1?
7
14
1?
1
8
19
1?
2
8
19
1
13+
32
2
23
29
29
48+
2
18
35
2
2
59
0
1
5
0
0
6
and brine shrimps have been described, 7 clam shrimps (plus
10 known undescribed taxa) have been added to species
lists for Australia, and the notostracan fauna is now thought
to be more complex. This work is still in progress. Here
I report on the present known diversity and distribution of
large branchiopods in Australia.
S YSTEMATIC R EVIEW
Anostraca
The Australian anostracan fauna is poor generically but
two genera (Branchinella, Parartemia) have radiated at the
species level so that overall richness is comparable to that
for many other zoogeographic regions, but not the huge
Palaearctic (Brendonck et al., 2008). This perception was
only reached recently (Table 1) as earlier assessments (Linder, 1941; Williams, 1980, 1981; Geddes, 1983) based on
more limited sampling did not reveal this specific diversity.
Both Linder (1941) and Geddes (1973a, 1981,1983) added
significant numbers of species to the fauna (see above), and
28 have been added in the last decade (Herbert and Timms,
2000; Rogers et al., 2007; Timms, 2001, 2002, 2005, 2008,
2010; Timms and Geddes, 2003; Timms and Hudson, 2009)
and the task is not finished yet (Table 1).
Ignoring Artemia whose distribution in Australia has been
largely influenced by man (Timms, 2011), only four species
(Branchinella affinis, B. australiensis, B. lyrifera, and B.
proboscida) are of wide distribution across the continent
(>50% of its area). Even for these there are significant areas
not included in their distribution (often the far north, the
forested east coast, and the isolated south, i.e., Tasmania
(hereafter Tas) (Tables 2, 3). The majority of species (53%)
occur in suitable habitat spread over 100 000 to 1 million
km2 or more (only 1.3-13% of the total area of 7.6 ×
106 km2 ), while 26 species (40%) are known only from
their type locality or nearby. Further collecting will possibly
enlarge the distributions of this later group, for instance
B. budjiti was initially collected only from northwestern
New South Wales (hereafter NSW), but now is known from
adjacent southern and central inland Queensland (hereafter
Qld) and northeastern South Australia (hereafter SA).
Most species live in drier inland areas where temporary
waters abound after rains. Of particular interest is the
Paroo region of northwestern NSW and southwestern Qld
(Fig. 1) where diversity is particularly high, with 18 species
of anostracans (Timms and Sanders, 2002; Timms, 2009).
Most of these species can be found in limited subareas,
e.g. the 30 km2 Bloodwood Station in NSW or 155 km2
Currawinya National Park in Qld. High diversity in this area
is encouraged by its variety of habitat types, there being an
abundance of various types of turbid claypans, clear water
grassy pools, treed swamps, and saline lakes. Diversity is
also high in WA (Tables 2-4), but this is due to its large area
and limited dispersal ability of Parartemia (Timms, 2002,
2010).
Interestingly, a few species occur on either side of
continent and not in between. This list includes Branchinella
pinnata, B. wellardi; in the case of B. compacta the two
metapopulations on each side of Australia are different
enough to be considered separate species (Timms, 2008;
T. Pinceel, personal communication). This suggests little
or no present interchange of genetic material between
populations on each side of Australia, though there could
be suitable staging habitat in central Australia, but limited
collecting there has not yet found intermediate populations.
In Parartemia, there are two species (P. acidiphila and P.
cylindrifera) with populations in southwestern Australian
and the Eyre Peninsula in South Australia separated by
the huge Nullarbor Plain (ca 1500 km) (Fig. 1). For these
species west and east populations are not morphologically
different, and also no molecular differences in the case of
P. cylindrifera (I. Kappas, personal communication), and
there is no possibility of intermediate populations on the
waterless plain. Hence, either speciation is extremely slow in
these cases, a scenario dismissed by Hebert et al. (2002) for
halobiont Parartemia, or there is at least some long distance
genetic mixing perhaps facilitated by long distance birdmediated dispersal (Green et al., 2008).
Species richness in each state is vastly different (Table 4),
strongly correlated with their areas (r = 0.9538). Given
Queensland and the Northern Territory’s (hereafter NT)
figures are below the trend line, more species could be found
there. What is more obvious is the high level of endemism
in WA vis-á-vis other states (Table 4). This has already been
commented upon separately for Branchinella (Timms, 2002,
2008) and Parartemia (Timms, 2010) and is influenced by
the isolation of the west from the remainder of Australia and
past climate changes.
A further difference between west and east Australia
is the level of congeneric occurrences in the freshwater
genus Branchinella. In the west, 19% of all occurrences
have two or more species within each site (diversity there
is due to the vast size of the state and the numerous
species of Parartemia), whereas in the east congeneric
617
TIMMS: THE DIVERSITY AND DISTRIBUTION OF LARGE BRANCHIOPODS
Table 2. Distribution of Branchinella species in Australia. *Indicates species is known only from its type locality and maybe
also nearby.
Species
Branchinella affinis Linder, 1941
B. apophysata Linder, 1941
B. arborea Geddes, 1981
B. australiensis (Richters, 1876)
B. basispina Geddes, 1981
B. buchananensis Geddes, 1981
B. budjiti Timms, 2001
B. campbelli Timms, 2001
B. compacta Linder, 1941
B. complexidigitata Timms, 2002
B. clandestina Timms, 2005
B. denticulata Linder, 1941
B. dubia (Schwartz, 1917)
B. frondosa Henry ,1924
B. halsei Timms, 2002
B. hattahensis Geddes, 1981
B. insularis Timms and Geddes, 2003
B. kadjikadji Timms, 2002
B. lamellata Timms and Geddes, 2003
B. latzi Geddes, 1981
B. longirostris Wolf, 1911
B. lyrifera Linder, 1941
B. mcraeae Timms, 2005
B. multidigitata Timms, 2008
B. nana Timms, 2002
B. nichollsi Linder, 1941
B. occidentalis Dakin, 1914
B. papillata Timms, 2008
B. pinderi Timms, 2008
B. pinnata Geddes, 1981
B. proboscida Henry, 1924
B. simplex Linder, 1941
B. tyleri Timms and Geddes, 2003
B. vosperi Timms, 2008
B. wellardi Milner, 1929
Branchinella n. sp. B
Branchinella n. sp. C
Branchinella n. sp. M
Branchinella n. sp. S
Branchinella n. sp. Y
Qld
NSW
Vic
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Tas
NT
SA
x
x
x
x
WA
x
x*
x
x
x
x*
x
x
x*
x*
x
x
x
x
x
x
x
x
x
x
x
x*
x*
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x*
x
x
x
x*
x
x
x
x
x*
x*
x
x
x*
x
x
x
x
x*
x
x
x*
x*
x*
occurrences are common, with 55% of collections in the
Paroo with more than one species (Timms, 2002; Timms
and Sanders, 2002), 54% of 13 sites in the Bulloo (an
endorheic catchment in Qld adjacent to the Paroo), 38% of
11 sites in northwestern Qld and 69% of 13 sites in central
Australia and adjacent northern SA (Timms, unpublished
data). For the brine shrimp Parartemia there are virtually
no congeneric occurrences anywhere (Timms et al., 2009).
The difference between the two genera is easily explained:
Branchinella feeds on planktonic algae, largely diatoms,
which vary in size, so niche specialisation is possible
(Timms and Sanders, 2002); whereas Parartemia feeds
largely on amorphous organic matter stirred from the bottom
(Marchant and Williams, 1977), so there is little opportunity
for niche diversification based on food particle size. There
is no easy explanation for different levels of congeneric
occurrences within Branchinella. Certainly it cannot be due
to greater species richness in the east as there are similar
numbers of species east and west (26 vs. 25 respectively,
Tables 2, 3). It is possible that the sparseness and greater
temporality of waters in the west vis-á-vis those in the centre
and east in wet La Nina years limits dispersal in the west and
hence the filling of niches.
As noted by Williams (1980, 1981), Parartemia is found
only in saline waters and Branchinella mainly in fresh waters. However some species of Branchinella are halobiont:
B. simplex (to 62 gL−1 ), B. buchananensis (to 42.6 gL−1 ), B.
papillata (to 14 gL−1 ), B. compacta (to 15.9 gL−1 ), B. nana
(to 13 gL−1 ), B. australiensis (to 11.2 gL−1 ), B. affinis (to
6.7 gL−1 ), and B. frondosa (to 4.2 gL−1 ) (Timms, 2009c).
New species H and M of Branchinella also are mildly salt
tolerant. Parartemia osmoregulate (Geddes, 1975) while at
least B. compacta and B. australiensis are osmoconformers,
their tissues tolerating the hyposaline conditions (Geddes,
618
JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 32, NO. 4, 2012
Table 3. Distribution of species of Parartemia, Streptocephalus and Australobranchipus. *Indicates species is known only
from its type locality and maybe also nearby.
Species
Parartemia acidiphila Timms and Hudson, 2009
P. auriciforma Timms and Hudson, 2009
P. bicorna Timms, 2010
P. boomeranga Timms, 2010
P. contracta Linder, 1941
P. cylindrifera Linder, 1941
P. extracta Linder, 1941
P. informis Linder, 1941
P. laticaudata Timms, 2010
P. longicaudata Linder, 1941
P. minuta Geddes, 1973
P. mouritzi Timms, 2010
P. purpurea Timms, 2010
P. serventyi Linder, 1941
P. triquetra Timms and Hudson, 2009
P. veronicae Timms, 1941
P. yarleensis Timms and Hudson, 2009
P. zietziana Sayce, 1903
Streptocephalus archeri Sars, 1896
S. queenslandicus Herbert and Timms, 2000
Australobranchipus parooensis Rogers et al., 2007
A.gilgaiphila Rogers et al., 2007
Fig. 1.
Map of Australia showing most places mentioned in the text.
Qld
NSW
Vic
Tas
NT
SA
WA
x
x*
x
x*
x*
x
x
x
x
x
x
x
x
x
x
x
x*
x
x
x*
x
x
x*
x
x
x
x
x
x
x
x
x
x
x
619
TIMMS: THE DIVERSITY AND DISTRIBUTION OF LARGE BRANCHIOPODS
Table 4.
State distribution of Anostraca.
State
Western Australia
Queensland
Northern Territory
South Australia
New South Wales
Victoria
Tasmania
Area (km2 × 106 )
Number of species
Endemic species
2.5
1.7
1.3
1.0
0.8
0.2
0.07
40
24
13
18
17
6
2
25
3
1
3
1
0
0
1973b). Presumably B. nana, B. affinis, and B. frondosa are
also osmoconformers, but it is not known how B. simplex
and B. buchananensis survive high salinities.
Two other genera of anostracans are found in Australia
but always in fresh waters (Table 3). Streptocephalus occur
across the northern two-thirds of the continent almost exclusively in summer only (author, unpublished data). Nominally there are two species, but in reality all populations
could be of one widespread species, comparative taxonomic
study being difficult due to no males ever being recorded for
S. archeri by Sars and their being rare for S. queenslandensis (see Herbert and Timms, 2000). The Australian endemic
Australobranchiopsis is represented by two species, neither common, occurring respectively in the western (A. parooensis) and northeastern (A. gilgaiphila) Murray-Darling
Basin. Both species are remarkable for their short life cycles, generally maturing and dying before various species of
Branchinella become well established (Rogers et al., 2007).
Parartemia, endemic to Australia, has undergone remarkable radiation with 18 known species and maybe a few
more, mainly in saline lakes in WA (Table 3). All species
of Parartemia use a lock and key mechanism in amplexus
and hence have distinctive antennal-head features in males
and thoracic modifications, including reduction/loss of the
eleventh thoracopod, in females (Rogers, 2002; Timms, in
press). Parartemia zietziana is known to tolerate salinities
to ca 300 g/kg, most species of Parartemia live in waters to ca 200-250 g/kg with just two (P. cylindrifica and
P. mouritzi) withstanding lower maximum salinities of ca
100 g/kg (Timms, 2009b). Even more remarkable is the ability of some species to live in acid waters (to pH 3); P. contracta survives in the absence of dissolved CO2 by having
an additional proton pump to produce ATP from endogenous CO2 (Conte and Geddes, 1988). The energetics of P.
zietziana has been elucidated (Marchant, 1978), this species
in highly saline lakes using >80% of assimilalation to osmoregulate. At times energy balance is negative, which accounts for its high mortality, inconsistent growth rates and
unpredictable recruitment (Marchant and Williams, 1977).
Much has been written about Artemia in Australia (summarised by Timms, 2011), and will not be repeated here.
Suffice it to say there are two species, A. franciscana Kellogg, 1906 limited to salt works, mainly in eastern and southern Australia and parthenogenetic populations mainly in WA
salt works and in many degraded lakes there (Timms, in
press). It may have been native to a few coastal lakes near
Perth before settlement, but now is spreading alarmingly,
unlike P. franciscana which is spreading elsewhere in the
world, but not yet in Australia (McMaster et al., 2007; Ruebhart et al., 2008)
Laevicaudata, Spinicaudata, and Cyclestherda
Clam shrimp are poorly known in Australia (Williams, 1980,
1981) and indeed not well studied or understood worldwide
(Brendonck et al., 2008). Some authors consider they are
of ‘sporadic occurrence’ and ‘by no means common’ in
Australia (Williams, 1980) while others like Weeks et al.
(2006) take a diametrically opposite view, at least for WA,
claiming they are ‘quite abundant and widespread.’ The
truth lies somewhere in between, with the Weeks et al.
(2006) opinion being inflated by the local abundance of
clam shrimps in many rock pools. Certainly their occurrence
is sporadic, but after good rains they can be common and
diverse as Timms and Richter (2002) showed for the Paroo
catchment in northwestern NSW.
Like many situations overseas, early descriptions of Australian clam shrimps are totally inadequate and for most
species there is no type material. Moreover early authors
often were ignorant of previous discoveries, none more so
than Henry (1924) who listed species known for Australia at
the time, but curiously omitted four described species (Limnadia badia, L. cygnorum, Eulimnadia feriensis, ‘Cyzicus’
rufa) from her list (Table 5). The Williams lists (1968, 1980,
1981) are more complete, but suffer from the uncritical acceptance of the presence of Cyzicus in Australia, when no
animal here (except the misnomer of ‘Cyzicus’ rufa Dakin,
1914) complies with the basic characters of this genus (see
below). Richter and Timms (2005) provide a modern synopsis of the fauna, increasing the list of known species to
26 (Table 5). However this list is far from complete, for instance Timms (2009b) recently recognised the presence of
Eoleptestheria and for almost all genera more species are
thought to occur in Australia than presently described (see
below). Adding to the uncertainty is the difficulty in identification of many species, e.g., Weeks et al. (2006) may have
misidentified Limnadia feriensis and L. sordida, and some
species of Limnadia and Caenestheria may be invalid. The
best estimate of the number of species present is approximately 50, which would make the Australian fauna at least of
similar richness to other major zoogeographic regions when
similar inaccuracies are corrected on their lists (Brendonck
et al., 2008).
Details for Australia are as follows.
Laevicaudata: Lynceidae: Lynceus.—Lynceus macleayanus
(King, 1855) and to a lesser extent L. tatei (Brady, 1886), are
widespread and common, mainly in longer lasting temporary
620
JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 32, NO. 4, 2012
Table 5. Summary of the number of species of Laevicaudata, Spinicaudata and Cyclestherida in Australia. *Brendonck et al. (2008) list species by families.
ˆ Many more species can be delinated molecularly (Schwentner, pers. comm.).
Suborders and
families
Genera
Henry (1924)
Laevicaudata
Lynceidae
Spinicaudata
Limnadiidae
Lynceus
2
(as Limnetis)
1 (as Paralimnadia)
Cyzicidae
Leptestheriidae
Cyclestherida
Cyclestheriidae
Total
Limnadia
Eulimnadia
Limnadopsis
‘Cyzicus’
Williams
Richter and Brendonck et al. Currently
Known
(1980, 1981) Timms (2005)
(2008)*
recognised undescribed
species
species
1
4
5
(as Estheria)
Eocyzicus
Caenestheria
Caenestheriella
Eoleptestheria
Cyclestheria
13
many omitted
waters. At least two further species are known from WA
alone (Zofkova, 2006; author, unpublished data). The genus
is in need of revision.
Spinicaudata: Limnadiidae: Limnadia.—There are seven
described species, but I recognise four undescribed species
and published and unpublished molecular work (Weeks
et al., 2009; M. Schwentner, personal communication)
suggests the presence of additional species. Three wellknown species live in rock pools: L. badia (Wolf, 1911)
in granite pools in southwest Australia, L. stanleyana King,
1855 in sandstone pools of the Sydney basin, and L. urukhai
Webb and Bell, 1979 in the granite pools of the mountains of
southeast Qld and northeast NSW (Timms, 2006). Limnadia
cygnorum (Dakin, 1914) lives in coastal southern WA,
while the status of L. grobbeni Daday, 1925, L. sordida
King, 1855 plus its synonym L. rivolensis Brady, 1886)
and L. victoriensis (Sayce, 1903) is unknown. It is possible
that most forms of what is called Limnadia in Australia
actually belong to an endemic genus (D. C. Rogers, personal
communication).
Spinicaudata: Limnadiidae: Eulimnadia.—There are two
described species, with E. dahli (Sars, 1896) widespread
across Australia, and E. feriensis Dakin, 1914 restricted to
WA. Unpublished molecular work (M. Schwentner, personal
communication) suggests there could be many more species
and in Richardson’s (1929) manuscript 5 new species were
proposed just for WA.
Spinicaudata: Limnadiidae: Limnadopsis.—This endemic
genus has recently been revised (Timms, 2009a) and now
includes two widespread species (L. birchii (Baird, 1860)
and L. tatei Spencer and Hall, 1896); and L. parvispinus
Henry, 1924 in eastern Australia and five additional species
in the west and northwest; L. multilineata Timms, 2009;
L. minuta Timms, 2009; L. occidentalis Timms, 2009; L.
paradoxa Timms, 2009; and L. pilbarensis Timms, 2009.
2
2
5
7
3
4
8
2
4
1
(C. rufa)
1
6
2
2
)
)13
)
)
)
)10
)
1
1
not included
23
26
25
2
2
7
4
2ˆ
8
4
2ˆ
7
2
1
1
32
10
The genus name Limnadopsium for L. tatei was not accepted
and L. brunneus Spencer and Hall, 1896 was declared a
nomen dubium. Molecular data by Weeks et al., 2009 and
Schwenter et al., 2011 suggest additional species. Three
from eastern Australia and one from central Australia, are
presently being described (Schwentner et al., in press).
Spinicaudata: Cyzicidae: Eocyzicus.—Presently two species
are known, both of wide distribution: Eocyzicus parooensis
Richter and Timms, 2005 occurs in hyposaline waters and
is the most salt tolerant of all clam shrimps (Timms,
2009b); and E. argillaquus Timms and Richter, 2009 prefers
turbid waters. Molecular work in progress (M. Schwentner,
personal communication) suggests there could be roughly
10 species in Australia. Richardson’s (1929) manuscript
recognised a new species; indeed he was the first to realise
this genus occurs in Australia, not Richter and Timms
(2005).
Spinicaudata: Cyzicidae: Cyzicus.—If Cyzicus is defined as
those cyzicids in which there is a deep occipital notch and an
associated acute extension of the occipital part of the head
together with a broadly spatulate rostrum in males (Daday,
1914, 1923, 1925, 1926, 1927; Mattox, 1959), then this
genus has so far not been recognised in Australia. However,
one species Cyzicus (= Estheria) rufa Dakin, 1914 remains
officially placed there as the description is insufficient to
place it elsewhere (Richter and Timms, 2005). Like many
clam shrimps described before 1950, it has not been sighted
since discovery.
Spinicaudata: Cyzicidae: Caenestheria.—Of the six described species, three are readily recognised: C. lutraria
(Brady, 1886) is widespread; C. sarsii (Sayce, 1903) (or
maybe it is C. ellipitica Sars, 1897) occurs in northern WA;
and C. berneyi (Gurney, 1927) in inland Qld, all in turbid
waters. The validity of others (C. dictyon (Spencer and Hall,
1896) and C. rubra (Henry, 1924) is questionable and at
TIMMS: THE DIVERSITY AND DISTRIBUTION OF LARGE BRANCHIOPODS
least one undescribed species is known (author, unpublished
data).
Spinicaudata: Cyzicidae: Caenestheriella.—Two species are
known: the widespread and ubiquitous C. packardi (Brady,
1886), and C. mariae (Olesen and Timms, 2005) limited
to rock pools in WA. When described, Spencer and Hall
(1896) indicated three varieties of C. packardi, which
Richter and Timms (2005) thought insufficiently separated
to list as subspecies. However M. Schwentner (personal
communication) has molecular evidence suggesting there
are many species and subspecies presently encompassed
within the name C. packardi. Richardson (1929) proposed
a new species for WA, as well as recognising the presence of
C. packardi.
Spinicaudata: Leptheseriidae: Eoleptestheria.—This was
first recognised in Australia by Timms (2009c), the species
thought to be the Eurasian E. ticenensis (Balsamo-Crivelli,
1859) perhaps recently arrived in tropical Australia. This
may not be so as one museum collection has since been
found to be 100 years old instead of recent (J. Taylor,
personal communication). Certainly though it is spreading
southwards as it is now found in the Paroo where it was not
collected 20 years ago.
Cyclestherida: Cyclestheriidae: Cyclestheria.—The circumtropical Cyclestheria hislopi (Baird, 1859) occurs across
northern Australia and southwards along the Qld coast to
about Rockhampton (Timms, 1986). This species is different
from other clam shrimps in that it typically lives in permanent vegetated waters and reproduces by parthenogenesis. In
2011 it was found in northern inland NSW (M. Schwentner
and B. Timms, unpublished data) and so could be moving
southwards or perhaps reclaiming lost territory (Williams,
1980).
Notostraca
According to Longhurst (1955) there are just two species of
shield shrimps in Australia, Lepidurus apus viridus Baird,
1850 and Triops australiensis australiensis Spencer and
Hall, 1896. Although both are widespread in temporary
pools, T. australiensis occurs in drier inland areas and L.
apus in the far southwest and much of the southeast of
the continent (Williams, 1968; Williams and Bushby, 1991;
Tyler et al., 1996). Before Longhurst’s review there was
a confusing array of species names available for both, a
situation which may repeat itself if present molecular work
is accepted. Thus there is a hermaphroditic taxon of Triops in
the Paroo (Murugan et al., 2009), a halophilic taxon in WA
salt lakes (Timms et al., 2007; B. Vanschoenwinkel, personal
communication) and at least three taxa in rock pools on
inselbergs in WA and the NT (B. Vanschoenwinkel, personal
communication). Historical records (e.g. Main, 1967), and
hearsay (B. Knott, personal communication) suggest Triops
used to occur occasionally in rock pools on inselbergs in the
inner wheatbelt, but not now (Timms, 2006).
D ISCUSSION
This assessment leads to four salient conclusions: 1) an imbalance in the rate of discovery of fairy and brine shrimps
621
vis-á-vis the various groups of clam shrimps, 2) species richness in Australia is not as depauperate as generally believed
and in fact matches those of most biogeographic regions,
3) only a few species are widespread across the continent
and many are localised, and 4) molecular studies have so far
largely confirmed species and generic distinctions and in future offer further delineation of present hidden diversity.
Initially, far more clam shrimps were described from
Australia than fairy and brine shrimps, though in recent
years this has been reversed. These recently discovered
anostracans were found by exploring in remote regions,
and in the future more species of all large branchiopods
may be described due to new species being detected by
molecular analysis rather than by traditional morphological
means, both as predicted by Adamowicz and Purvis (2005)
in their assessment of world diversity of branchiopods. At
this moment there is evidence of such further new species
among the clam and shield shrimps by this route, but only
one fairy shrimp so far.
Until this assessment, Australia’s large branchiopod fauna
could have been considered depauperate (Williams, 1980,
1981; Brendonck et al., 2008) but not anymore. Adamowicz
and Purvis’s (2005) ‘uplift factor of 1.1 for differential
knowledge of diversity among biogeographical regions’ was
severely underestimated for large branchiopods in Australia,
for since their analysis date of 2003 the number of described
species has increased by a factor of 1.6 (36 to 58 valid
species) for fairy shrimps and by 1.3 (23 to 32 species)
for clam shrimps. This is due to my activities over the last
decade in which I have explored remote areas for diversity,
thus largely catching up on earlier exploration in other
continents.
Having some species widespread across the continent and
others localised is not unusual among the biogeographic
regions, but the proportion of species known only from
their type localities is inordinately high at an estimated 30%
for fairy shrimps and 15% for clam shrimps. Obviously,
further exploration is needed to fill in the gaps and from the
inequalities between states exposed in Table 4, Qld and NT
need further study. In addition there is a large area of remote
inland WA and also northwestern SA which could harbour
further species of brine shrimps (Timms and Hudson, 2009;
Timms, 2010). A major inhibiting problem for these remote
areas is impossible land access after episodic rains when
the shrimps appear, so that further discoveries will be much
slower than in the most recent decade.
Adamowicz and Purvis (2005) estimated diversity of
large branchiopods will increase by a factor of 1.7 due
to splitting of existing species by molecular analysis. As
shown above this is commencing among clam and shield
shrimps, but hardly yet among anostracans. Relationships
within Branchinella and Parartemia have been surveyed using molecular techniques and almost all species have been
proved valid (Remigio et al., 2001, 2003; Kappas et al.,
2011; T. Pinceel, personal communication) Exceptions include Branchinella clandestina which lies within the variability shown by B. affinis (T. Pinceel, personal communication) and P. contracta populations which differentiate
into two groups (Kappas et al., 2011). In a separate study,
fragmented populations of B. longirostris showed different
622
JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 32, NO. 4, 2012
patterns in their genetic constitution and morphological variation, but it was still concluded the various populations belonged to a single species, but with an exceptionally high intraspecific diversity (Zofkova and Timms, 2009). T. Pinceel
(personal communication) showed two species pairs (B. halsei and B simplex; B. mcraeae and B. pinderi) to be congruent molecularly, but I do not accept this as their morphology
is quite different (see Timms, 2004).
ACKNOWLEDGEMENTS
This paper was presented at the seventh International Large Branchiopod
Symposium in Taipai, Taiwan, September 10-12, 2011 organised by Professor Lien-Siang Chou. I thank Ilias Kappas, Brenton Knott, Tom Pinceel,
Martin Schwentner and Bram Vanschoenwinkel for sharing unpublished
data with me, Christopher Rogers for helpful comments on the manuscript
and two anonymous referees for advice.
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R ECEIVED: 23 November 2011.
ACCEPTED: 20 February 2012.