Page 1
The biogeography of Western Australian
shallow-water barnacles
Diana S. Jones
Department of Aquatic Zoology, Western Australian Museum, Francis Street
Perth, Western Australia 6000, Australia
email: diana.jones@museum.wa.gov.au
Abstract – The barnacle fauna of tropical north-western Western Australia exhibits high species
diversity (101), a high incidence of tropical species (69) and a low species endemicity (6). In
comparison, the barnacles of warm-temperate south-western Western Australia show decreased
species diversity (44), a lower incidence of tropical species (13) and a higher species endemicity
(10). Western Australian barnacle distributions corroborate the general patterns demonstrated for the
shallow-water biota of northern tropical and southern warm-temperate provinces of Australia. In
south-western Western Australia, however, there is a stronger tropical component than could be
predicted by latitude, due to the influence of the Leeuwin Current, a poleward flowing, eastern
boundary current along the western coast of Australia. Members of the balanomorph families
Chthamalidae, Tetraclitidae and Balanidae dominate the tropical littoral north-west. Since
chthalamids are absent from warm-temperate south-western Western Australia, zonation shows a
simpler tetraclitid-balanid trend, with archaeobalanids also prominent on sheltered shores. In both
tropical and warm-temperate Western Australia the pedunculate component is larger in the sub-
littoral, the majority of species being epizoic on a variety of hosts. Species diversity in the warm-
temperate littoral and sub-littoral is not as great as that occurring in those zones in the tropics.
Pelagic species are all pedunculate.
INTRODUCTION
Information on the general distribution of Cirripedia in Australian waters was little known
prior to the latter part of the 20th century. In 1990 the barnacle fauna of Australian waters was
documented for the first time (Jones et al., 1990). Subsequently, a number of publications have,
more specifically, documented the barnacle fauna of the shores and shallow waters of Western
Australia (e.g. Jones, 1987; 1990a; 1990b; 1990c; 1991; 1992a; 1992b; 1993; Jones and Berry,
2000; Jones and Hewitt, 1995; 1996; 1997). These data, in association with those collected
during the Western Australian Museum/Woodside Marine Biological Survey of the Dampier
Archipelago, including the workshop at Dampier in July–August 2000 (Jones, 2003 in press),
enable a comparison to be made of shallow-water barnacle distribution and zonation patterns in
tropical and warm-temperate Western Australia.
In Australia it is generally recognised that northern tropical and southern warm-temperate
provinces exist (Wilson and Gillett, 1971; Wilson and Allen, 1987; Poore, 1994; O’Hara and
Poore, 2000). Tropical and warm-temperate provinces grade into each other in a broad overlap
zone along both the western and eastern Australian coasts. This region of overlap constitutes a
transitional zone, with the gradual replacement of a tropical fauna in the north by a
F.E. Wells, D.I. Walker and D.S. Jones (eds) 2003. The Marine Flora and Fauna of Dampier, Western Australia.
Western Australian Museum, Perth.
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480D.S. JONES
predominantly warm-temperate fauna in the south. A cold temperate province is absent in
Australia (Wilson and Allen, 1987).
Unlike South America and Africa, Australia has southward flowing warm currents on both
western and eastern coasts, i.e. the Leeuwin Current and the East Australian Current,
respectively. On the western coast the Leeuwin Current roughly parallels the East Australian
Current, the latter bringing warm waters southward to about 33°S before diverting as eddies into
the Tasman Sea (Cresswell, 1987).
The Leeuwin Current is different from other Southern Hemisphere eastern boundary
currents, such as the Humboldt Current of South America and the Benguela Current of South
Africa, which are northward flowing, cool and involve upwelling (Pearce and Cresswell,
1985; Morgan and Wells, 1991). The Leeuwin Current is an offshoot of the through-flow of
western Pacific water that has travelled through the Indonesian Archipelago before entering
the eastern Indian Ocean. During its transit through the Archipelago, this water has heated up
as well as gained a substantial low-salinity component as a result of heavy monsoonal rains in
the region. Leaving the Timor Passage it flows as a warm current across the Timor Sea, then
around North West Cape (21°47'S) as a poleward flowing, eastern boundary current along the
western coast of Australia, flowing south around Cape Leeuwin (34°22'S) and thence east into
the Great Australian Bight. This warm current of relatively low salinity water is a narrowish
body (~ 50 km wide and 200 m deep), flowing along the outer edge of the continental shelf
(Godfrey and Ridgeway, 1985). The Leeuwin Current is mostly quiescent in the austral
summer (November–February) but flow to the south intensifies in autumn (March), attaining
maximum flow in April–June, driven by a longshore sea level gradient (Godfrey and
Ridgeway, 1985). The current then abates and disappears in September–October. It is now
recognised that the flow of this current is strongly influenced by the El Niño Southern
Oscillation (Pearce and Phillips, 1988).
The existence of a warm Western Australian Current was postulated as early as the end of the
last century by Saville-Kent (1897) when attempting to explain the existence of exceptionally
diverse coral reefs at the Houtman Abrolhos Islands (28º–29ºS). This flow was later named the
Leeuwin Current by Cresswell and Golding (1980). The Leeuwin Current is now recognised as
a major influence contributing to the dispersal of tropical representatives of many taxa to the
south-western and southern coasts of Australia, farther south than could be predicted by latitude
(e.g. Wilson and Marsh, 1979; Maxwell and Cresswell, 1981; Veron and Marsh, 1988; Hutchins
and Pearce, 1994).
The present paper discusses the distributions of shallow-water (0–100m) barnacle species
occurring in tropical and warm-temperate Western Australia. These distributions are related to
the effects of the Leeuwin Current. The zonation patterns of barnacle species occurring in
tropical and warm-temperate Western Australia are also compared and discussed.
MATERIALS AND METHODS
A collection of 36 species of littoral and shallow-water barnacle species was made during the
expeditions, fieldwork and workshop associated with the Woodside Energy Ltd/Western
Australia Museum partnership to explore the marine biodiversity of the Dampier Archipelago
(Jones, 2003 in press). Data from these collections, as well as cirripede distribution data in
Western Australian waters contained in other Australian Museum collections and in literature
records, are included in the analysis presented in this paper.
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BIOGEOGRAPHY OF WESTERN AUSTRALIAN BARNACLES 481
RESULTS
Barnacle distribution
North-western Australia
In north-western Australia the barnacle fauna has been documented for the Kimberley area
(Jones, 1992b; Jones and Hewitt, 1997), the Dampier Archipelago (Jones, in press 2003), the
Muiron Islands and the eastern shore of Exmouth Gulf (Jones and Hewitt, 1996), and the
Montebello Islands (Jones and Berry, 2000). The Kimberley barnacle fauna (56 species) is
dominated by tropical Indo-west Pacific (46) and Indo-Australian (7) species. No species with
Australasian affinities were recorded. Only one barnacle species is endemic to Australia (Jones,
1992b; Jones and Hewitt, 1997) (Table 1).
A total of 49 barnacle species are presently recorded from the Dampier Archipelago (Jones,
in press 2003). Excluding cosmopolitan species, the majority are either Indo-west Pacific (27)
or Indo-Australian (8). Four species are Australian endemics and there are no species with
Australasian affinities (Table 1).
Thirty-nine species of shallow-water and pelagic cirripedes have been collected from the
Muiron Islands and the eastern shores of Exmouth Gulf (Jones and Hewitt, 1996). Additional
collecting at the Montebello Islands (Jones and Berry, 2000) added a further five species,
making a total of 44 from the North West Cape area. Of these, the majority are Indo-west
Pacific (38) or Indo-Australian (2). No endemic or Australasian species were collected (Table
1).
The barnacle fauna of the Dampier region shows many similarities with that documented for
the Kimberley. The latter, however, is more diverse in terms of species numbers and is
dominated, in terms of species diversity and numbers of individuals, by pyrgomatids and
archaeobalanids (Jones, 1992b). The Dampier fauna is similar to, although richer than, that
documented for the Muiron Islands and the eastern shores of Exmouth Gulf, and for the
Montebello Islands.
South western Australia
Thirty-one species of shallow-water and pelagic cirripede species were documented by Jones
(1990c) for the warm-temperate waters of south western Australia. Excluding a large
cosmopolitan element (16 species), a strong southern element (nine species) and a distinct
tropical component (six species) were present. The southern element consists of six species
endemic to southern Australia and three with Australasian affinities. The tropical component
Table 1
Affinities of intertidal and shallow-water barnacles of tropical north-western Australia (data from Jones,
1992b; Jones, in press 2003; Jones and Berry, 2000; Jones and Hewitt, 1996; Jones and Hewitt, 1997).
Ratios Pedunculata:Sessilia. C, cosmopolitan species; IWP, Indo-west Pacific species (extend from east Af-
rica to Hawaii); IA, Indian/Australian species (extend from the Indo-Malayan Archipelago, Australian and
New Guinea, to Japan); AA, Australasian species (occurring in Australian/New Zealand); AE, Australian
endemic species (only occurring in Australia).
Area in NW AustraliaSub order FamilyGenus SpeciesC IWP IA AA AE
Kimberley
Dampier Archipelago
North West Cape area
4 (3:1)
4 (3:1)
4 (3:1)
8 (3:5)
11 (5:6)
11 (5:6)
22 (3:19)
24 (7:17)
20 (5:15)
56 (3:53)
49 (9:40)
44 (5:39)
2 46
27
38
7
8
2
0
0
0
1
4
0
10
4
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482 D.S. JONES
(three Indo-west Pacific and three Indo-Australian species) was attributed to the influence of
the southerly flowing, warm water Leeuwin Current, which disperses tropical representatives of
many taxa to the southwest and southern coasts of Australia. The presence of Southern Ocean
species of the genus Lepas as well as fouling species (Balanus trigonus and Megabalanus
tintinnabulum) boosted the cosmopolitan element of the south-western Australian barnacle
fauna (Table 2).
Comparison of littoral and shallow-water barnacle faunas of tropical north-western and
temperate south-western Australia.
Distribution records from other Australian museum collections and the literature, additional
to as reported above, were also gathered (Jones, in press 2003). One hundred and one and 44
shallow-water barnacle species are now recognised from tropical north-western (Table 3) and
warm-temperate south-western Australian waters (Table 4), respectively.
Table 2
Affinities of intertidal and shallow-water barnacles of temperate south-western Australia (data from Jones,
1990c).
Sub orderFamilyGenus SpeciesC IWPIAAA AE
SW Australia
5 (4:1) 9 (5:4)16 (7:9) 31 (16:15)163336
Table 3
Taxonomic affinities of intertidal and shallow-water barnacles of tropical north-western Australia.
Order Pedunculata Order SessiliaTotal
Suborder
Families
Genus
Species
4
7
1
8
5
15
40
101
13
27
27
74
Table 4
Taxonomic affinities of intertidal and shallow-water barnacles of temperate south western Australia.
Order PedunculataOrder SessiliaTotal
Suborder
Families
Genus
Species
4
6
11
24
1
6
5
12
24
44
13
20
Table 5
Taxonomic ands biogeographic affinities of intertidal and shallow-water barnacles of tropical north-west-
ern and temperate south-western Australia. C, cosmopolitan species; IWP, Indo-west Pacific species; IA,
Indian/Australian species; AA, Australasian (ie Australian/New Zealand) species; AE, Australian endemic
species.
ProvinceSub orderFamily Genus SpeciesC IWPIA AA AE
Tropical
Temperate
5
5
15
12
40
24
101
44
26
21
45
10
24
3
0
3
6
7
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BIOGEOGRAPHY OF WESTERN AUSTRALIAN BARNACLES 483
Table 6
Biogeographic affinities of intertidal and shallow-water barnacles of tropical north-western Australia.
Sub order
Family
Genus
Species
C
IWP
IA
AA
AE
Heteralepadomorpha
Heteralepadidae
Heteralepas
japonica (Aurivillius, 1892: 125)
+
Paralepas
quadrata (Aurivillius, 1894: 30)
+
Iblomorpha
Iblidae
Ibla
cumingi Darwin, 1851: 183
+
Lepadomorpha
Oxynaspididae
Oxynaspis
celata Darwin, 1851: 34
+
Poecilasmatidae
Octolasmis
angulata Aurivillius, 1894: 22
+
aperta Aurivillius, 1894: 22
+
cf bullata Aurivillius, 1892: 125
+
cor Aurivillius, 1892: 124
+
lowei (Darwin, 1851: 128)
+
neptuni (MacDonald, 1869: 440)
+
warwickii (Gray, 1825: 100)
+
Dichelaspis
orthogonia Darwin, 1851: 130
+
Trilasmis
eburnea Hinds, 1844: pl. 21 fig. 5
+
Poecilasma
kaempferi (Darwin, 1851: 102)
+
Temnaspis
amygdalum (Aurivillius, 1894: 10)
+
excavatum (Hoek, 1907: 10)
+
fissum Darwin, 1851: 109
+
Lepadidae
Lepas
anatifera Linnaeus, 1758: 668
+
anatifera striata (de Graaf, 1952)
+
anserifera Linnaeus 1767: 1109
+
pectinata Spengler, 1793: 106
+
Conchoderma
auritum (Linnaeus, 1767: 1110)
+
chelonophilum (Leach, 1818: 412)
+
hunteri (Owen, 1830: 71)
+
virgatum (Spengler, 1790: 207)
+
Scalpellomorpha
Calanticidae
Calantica
sp. nov.
+
Lithotryidae
Lithotrya
valentiana (Gray, 1825: 102)
+
Balanomorpha
Chthamalidae
Caudoeuraphia
caudata (Pilsbry, 1916: 314)
+
Microeuraphia
withersi (Pilsbry, 1916: 312)
+
sp. 1
+
sp. 2
+
Chthamalus
malayensis Pilsbry, 1916: 310
+
Chelonibiidae
Chelonibia
caretta (Spengler, 1790: 394)
+
patula (Ranzani, 1818: 396)
+
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484 D.S. JONES
Table 6
(cont.)
Sub order
Family
Genus
Species
C
IWP
IA
AA
AE
testudinaria (Linnaeus, 1758: 668)
+
Platylepadidae
Platylepas
decorata Darwin, 1854: 429
+
hexastylos (Fabricius, 1798: 35)
+
ophiophilius Lanchester, 1902: 371
+
Stomatolepas
elegans (Costa, 1838: 117)
+
praegustator Pilsbry, 1910: 304
+
Coronulidae
Coronula
diadema (Linnaeus, 1767: 1108)
+
Cetopirus
complanatus (Mörch, 1852: 67)
+
Tubicinella
cheloniae Monroe & Limpus, 1979: 199
+
Tetraclitidae
Tetraclitella
costata (Darwin, 1854: 339)
+
divisa (Nilsson-Cantell, 1921: 362)
+
multicostata (Nilsson-Cantell, 1930: 2)
+
sp. nov.
+
Newmanella
vitiata (Darwin, 1854: 340)
+
Tetraclita
squamosa (Bruguière, 1789: 170)
+
Archaeobalanidae
Armatobalanus
allium (Darwin, 1854: 281)
+
arcuatus Hoek, 1913: 210
+
cf cepa (Darwin, 1854: 283)
+
filigranus (Broch, 1916: 8)
+
quadrivittatus (Darwin, 1854: 284)
+
quinquivittatus (Hoek, 1913: 216)
+
terebratus (Darwin, 1854: 285)
+
Solidobalanus
auricoma (Hoek, 1913: 198)
+
ciliatus (Hoek, 1913: 199)
+
Striatobalanus
amaryllis (Darwin, 1854: 279)
+
bimae (Hoek, 1913: 182)
+
tenuis (Hoek, 1883: 154)
+
Conopea
calceolus (Ellis, 1758: 853)
+
cymbiformis (Darwin, 1854: 221)
+
mjobergi (Broch, 1916: 7)
+
sp. 1 cf navicula (Darwin, 1854: 221)
+
sp. 2 cf dentifer (Broch, 1922: 326)
+
Archiacasta
spinitergum (Broch, 1931: 112)
+
Neocasta
glans (Lamarck, 1818, 398)
+
laevigata Gray, 1825: 103
+
Euacasta
antipathidus (Broch, 1916: 13)
+
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BIOGEOGRAPHY OF WESTERN AUSTRALIAN BARNACLES 485
dofleini (Krüger, 1911: 56)
+
porata Nilsson-Cantell, 1921: 348
+
zuiho (Hiro, 1936: 632)
+
Acasta
conica Hoek, 1913: 235
+
cyathus Darwin, 1854: 312
+
echinata Hiro, 1937: 70
+
fenestrata Darwin, 1854: 316
+
hirsuta Broch, 1916: 10
+
idiopoma Pilsbry, 1912: 294
+
japonica Pilsbry, 1911: 80
+
purpurata Darwin, 1854: 318
+
spongites (Poli, 1795: 25)
+
sulcata Lamarck, 1818: 398
+
Pectinoacasta
pectinipes (Pilsbry, 1912: 294)
+
Pyrgomatidae
Cantellius
pallidus (Broch, 1931: 118)
+
Savignium
crenatum (Sowerby, 1823: no pagination)
+
dentatum (Darwin, 1854: 369)
+
elongatum (Hiro, 1931: 154)
+
milliporae (Darwin, 1854: 367)
+
Nobia
conjugatum (Darwin, 1854: 364)
+
grandis Sowerby, 1839: 71
+
Pyrgoma
cancellata Leach, 1818: 171
+
Balanidae
Balanus
amphitrite Darwin, 1854: 240 (part)
+
cirratus Darwin, 1854: 241
+
reticulatus Utinomi, 1967: 216
+
trigonus Darwin, 1854: 223
+
sp.nov.
+
Megabalanus
ajax (Darwin, 1854: 214 (part))
+
cf ajax (Darwin, 1854: 214 (part))
+
rosa (Pilsbry, 1916: 61)
+
tintinnabulum (Linnaeus, 1758: 668)
+
Total
5
15
40
101
26
45
23
0
7
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486 D.S. JONES
Table 7
Biogeographic affinities of intertidal and shallow-water barnacles of temperate south-western Australia.
Sub order
Family
Genus
Species
C
IWP
IA
AA
AE
Heteralepadomorpha
Heteralepadidae
Paralepas
georgei Daniel, 1970: 33
+
quadrata (Aurivillius, 1894: 30)
+
Iblomorpha
Iblidae
Ibla
quadrivalvis (Cuvier, 1817: pl. 1 figs 15–16)
+
Lepadomorpha
Oxynaspidae
Oxynaspis
celata Darwin, 1851: 34
+
Poecilasmatidae
Poecilasma
dubium Hoek, 1907: 6
+
kaempferi (Darwin, 1851: 102)
+
Temnaspis
amygdalum (Aurivillius, 1894: 10)
+
fissum (Darwin, 1851: 109)
+
Octolasmis
angulata (Aurivillius, 1894: 22)
+
cor (Aurivillius, 1892: 124)
+
lowei (Darwin, 1851: 128)
+
neptuni (MacDonald, 1869: 440)
+
warwickii (Gray, 1825: 100)
+
Dichelaspis
orthogonia Darwin, 1851: 130
+
Lepadidae
Lepas
anatifera Linnaeus, 1758: 668
+
anserifera Linnaeus, 1767: 1109
+
australis Darwin, 1851: 89
+
hillii (Leach, 1818: 413)
+
pectinata Spengler, 1793: 106
+
testudinata Aurivillius, 1892: 123
+
Dosima
fascicularis Ellis & Solander, 1786: 197
+
Conchoderma
auritum (Linnaeus, 1767: 1110)
+
virgatum (Spengler, 1790: 207)
+
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BIOGEOGRAPHY OF WESTERN AUSTRALIAN BARNACLES 487
Scalpellomorpha
Calanticidae
Smilium
peronii Gray, 1825: 100
+
Balanomorpha
Pachylasmatidae
Tetrapachylasma
ferrugomaculosa (Jones, 1993)
+
Chelonibiidae
Chelonibia
testudinaria (Linnaeus, 1758: 668)
+
Coronulidae
Coronula
diadema (Linnaeus, 1767: 1108)
+
Cetopirus
complanatus (Mörch, 1852: 67)
+
Tetraclitidae
Epopella
simplex (Darwin, 1854: 353)
+
Tetraclitella
purpurascens (Wood, 1815: 55)
+
Archaeobalanidae
Solidobalanus
auricoma (Hoek, 1913: 198)
+
cilatus (Hoek, 1913: 199)
+
compressus (Hoek, 1913: 202)
+
Euacasta
dofleini (Krüger, 1911: 56)
+
Acasta
spongites (Poli, 1795: 25)
+
sulcata Lamarck, 1818: 398
+
Elminius
covertus Foster, 1982: 24,
+
modestus Darwin, 1854: 350
+
Balanidae
Balanus
amphitrite Darwin, 1854: 240 (part)
+
reticulatus Utinomi, 1967: 216
+
variegatus Darwin, 1854: 241
+
trigonus Darwin, 1854: 223
+
Austromegabalanus
nigrescens (Lamarck,1818: 391)
+
Megabalanus
tinnntinnabulum (Linnaeus, 1758: 668)
+
Total
5
12
24
44
21
10
3
3
7
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488 D.S. JONES
In tropical north-western Australia, 27 species are in the Pedunculata and the remaining 74 in
the Sessilia. The 27 pedunculate species are in 13 genera held in seven families within four
suborders. The 74 sessile species belong to 27 genera within eight families of one suborder
(Table 3). Of the 44 barnacle species recorded from temperate south-western Australian waters,
24 species are in the Pedunculata and the remaining 20 are in the Sessilia. The 24 pedunculate
species are in 11 genera held in six families within four suborders. The 20 sessile species belong
to 13 genera within six families of one suborder (Table 4).
The taxonomic and biogeographic affinities of the shallow-water barnacles of tropical north-
western and temperate southern waters of Western Australia are shown in Tables 5, 6, and 7.
The north-western barnacle fauna exhibits high species diversity (101), a high incidence of
tropical species (69) and a low species endemicity (6) (Tables 5 and 6). In comparison, the
barnacles of the southern temperate province exhibit decreased species diversity (44), a lower
incidence of tropical species (13) and higher species endemicity (10) (Tables 5 and 7)
Bathymetric zonation
Bathymetrically four barnacle species assemblages are recognised i.e. littoral, lower littoral
and sub-littoral, sub-littoral only, and pelagic (Foster, 1978; Jones, 1990c). The bathymetric
distribution of littoral and shallow-water barnacles of tropical north-western and temperate
south-western Australia are shown in Table 8.
Tropical north-western Australian shores
Members of the sessile balanomorph Chthamalidae, Tetraclitidae and Balanidae dominate the
littoral. The iblomorph Ibla cumingi and the scalpellomorph Lithotrya valentiana are the only
pedunculate shore representatives (Tables 8 and 9). In the tropical sub-littoral, whilst sessile
forms continue to dominate (58 species), the pedunculate component is much higher (17
species) than in the littoral (2 species). The majority of the tropical sub-littoral pedunculates are
epizoic species attaching to a variety of hosts, e.g. Oxynaspis celata attached to Antipatharia
spp, Octolasmis spp attached to decapod crustaceans and Trilasmis eburneum attached to
echinoids. Species diversity is greatest in the tropical littoral and sub-littoral (93 species)
compared to the temperate littoral and sub-littoral (35 species) (Table 8). Of the tropical pelagic
species, none is sessile. Numbers of tropical pedunculates almost equal those in temperate
waters (Tables 8 and 9).
Temperate south-western Australian shores
Members of the sessilian balanomorph Tetraclitidae and Balanidae dominate the littoral and,
on sheltered shores, the Archaeobalanidae may also be prominent. The iblomorph Ibla
quadrivalvis is the sole pedunculate shore representative (Tables 8 and 9). In the temperate sub-
littoral, the pedunculate component (14 species) is larger than in the temperate littoral (1
species). Similar to tropical sub-littoral pedunculates, most temperate sub-littoral pedunculates
are epizoic species attaching to a variety of hosts (e.g. Paralepas spp attached to Panulirus spp;
Poecilasma spp, Temnaspis spp and Octolasmis attached to a variety of decapod crustaceans).
Numbers of pedunculate and sessile species are almost equal in the temperate sub-littoral (14
and 12 species, respectively). Species diversity in the temperate littoral and sub-littoral (35
species) is not as great as that occurring in the tropical littoral and sub-littoral (93 species).
Pelagic temperate species have no sessile representatives. The pelagic pedunculates (9 species)
are almost equal in number to those found in tropical waters (8 species).
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BIOGEOGRAPHY OF WESTERN AUSTRALIAN BARNACLES 489
Table 8
Bathymetric assemblages of tropical north-western and temperate south-western Australian barnacles.
Littoral only
Pedunculata
Lithotrya valentiana
Sessilia
Caudoeuraphia caudata
Microeuraphia withersi
Microeuraphia sp. 1
Microeuraphia sp. 2
Chthamalus malayensis
Tropical
Temperate
Tetraclitella purpurascens
Elminius covertus
Littoral and sub-littoral
Pedunculata
Ibla cumingi
Sessilia
Newmanella vitiata
Tetraclitella costata
Tetraclitella divisa
Tetraclitella multicostata
Tetraclitella sp. nov.
Balanus amphitrite
Balanus amphitrite cf poecilosculpta
Balanus cirratus
Balanus cf poecilotheca
Megabalanus tintinnabulum
Tropical
Temperate
Ibla quadrivalvisElminius modestus
Balanus amphitrite
Balanus reticulatus
Balanus variegatus
Austromegabalanus nigrescens
Megabalanus tintinnabulum
Always sub-littoral
Pedunculata Sessilia
Tropical
Heteralepas japonica
Paralepas quadrata
Oxynaspis celata
Octolasmis angulata
Octolasmis aperta
Octolasmis cf bullata
Octolasmis cor
Octolasmis lowei
Octolasmis neptuni
Octolasmis warwickii
Dichelaspis orthogonia
Trilasmis eburnea
Poecilasma kaempferi
Temnaspis amygdalum
Temnaspis excavatum
Temnaspis fissum
Calantica sp. nov.
Chelonibia caretta
Chelonibia patula
Chelonibia testudinaria
Platylepas decorata
Platylepas hexastylos
Platylepas ophiophilius
Stomatolepas elegans
Stomatolepas praegustator
Coronula diadema
Cetopirus complanatus
Tubicinella cheloniae
Armatobalanus allium
Armatobalanus arcuatus
Armatobalanus cepa
Armatobalanus filigranus
Armatobalanus quadrivittatus
Armatobalanus quinquivittatus
Armatobalanus terebratus
Solidobalanus auricoma
Solidobalanus ciliatus
Striatobalanus amaryllis
Striatobalanus bimae
Striatobalanus tenuis
Page 12
490 D.S. JONES
Conopea calceolus
Conopea cymbiformis
Conopea mjoebergi
Conopea sp.1 cf navicula
Conopea sp. 2 cf dentifer
Archiacasta spinitergum
Neocasta glans
Neocasta laevigata
Euacasta antipathidus
Euacasta dofleini
Euacasta porata
Euacasta zuiho
Acasta conica
Acasta cyathus
Acasta echinata
Acasta fenestrata
Acasta hirsuta
Acasta idiopoma
Acasta japonica
Acasta purpurata
Acasta spongites
Acasta sulcata
Pectinoacasta pectinipes
Cantellius pallidus
Savignium crenatum
Savignium dentatum
Savignium elongatum
Savignium milliporae
Nobia conjugatum
Nobia grandis
Pyrgoma cancellata
Balanus trigonus
Megabalanus ajax
Megabalanus cf ajax
Megabalanus rosa
Temperate
Paralepas georgei
Paralepas quadrata
Oxynaspis celata
Poecilasma dubium
Poecilasma kaempferi
Temnaspis amygdalum
Temnaspis fissum
Octolasmis angulata
Octolasmis cor
Octolasmis lowei
Octolasmis neptuni
Octolasmis warwicki
Dichelaspis orthogonia
Smilium peronii
Tetrapachylasma ferrugomaculosa
Chelonibia testudinaria
Coronula diadema
Cetopirus complanatus
Epopella simplex
Solidobalanus auricoma
Solidobalanus ciliatus
Solidobalanus compressus
Euacasta dofleini
Acasta spongites
Acasta sulcata
Balanus trigonus
Pelagic
Pedunculata
Lepas anatifera
Lepas anatifera striata
Lepas anserifera
Sessilia
Tropical
Table 8
(cont.)
Pedunculata Sessilia
Page 13
BIOGEOGRAPHY OF WESTERN AUSTRALIAN BARNACLES 491
DISCUSSION
Barnacle distribution patterns
The tropical shallow-water biota of the northern Australian tropical province is continuous
with other parts of the Indo-west Pacific, and the Indo-west Pacific element constitutes the bulk
of this biota, with an Indo-Australian element also present on the western, and northern and
eastern coasts (Knox, 1963). As well as a high incidence of tropical species, the northern
Lepas pectinata
Conchoderma auritum
Conchoderma chelonophilum
Conchoderma hunteri
Conchoderma virgatum
Temperate
Lepas anatifera
Lepas anserifera
Lepas australis
Lepas hillii
Lepas pectinata
Lepas testudinata
Dosima fascicularis
Conchoderma auritum
Conchoderma virgatum
Table 9
Analysis of barnacle bathymetric assemblages of tropical north-western and temperate south-western
Australia.
Shore only
Pedunculata spp
1
0
1
Sessilia spp
6
2
8
Total nos spp
7
2
9
Tropical
Temperate
Total nos spp
Lower shore and sublittoral
Pedunculata spp
1
1
2
Sessilia spp
10
6
16
Total nos spp
11
7
18
Tropical
Temperate
Total nos spp
Always sublittoral
Pedunculata spp
17
14
31
Sessilia spp
58
12
70
Total nos spp
75
26
101
Tropical
Temperate
Total nos spp
Pelagic
Pedunculata spp
8
9
17
Sessilia spp
0
0
0
Total nos spp
8
9
17
Tropical
Temperate
Total nos spp
Table 8
(cont.)
PedunculataSessilia
Page 14
492 D.S. JONES
Australian biota exhibits low species endemicity and high species diversity – e.g. this has been
demonstrated for crustacean groups such as the stomatopods (Stephenson and McNeil, 1955),
the Penaeidae (Dall, 1957; Racek, 1959) and the Portunidae (Stephenson, 1972). Species
richness grades with latitude, with more species recorded in tropical than temperate waters (e.g.
demonstrated by Rowe and Gates, 1995 for echinoderms).
In Western Australia, North West Cape (21°47'S) and the offshore Houtman Abrolhos Islands
(28°19'–29°57'S) are generally considered to be the southern-most limit of the tropical marine
biota (Wilson and Allen, 1987). The southern limit of the tropical fauna in eastern Australia is
usually regarded as being somewhere between 26° and 27°S (Endean, 1957; Wilson and Gillett,
1971).
The north-western Australian barnacle fauna has been shown to be typical of the tropical
shallow-water biota of the northern Australian tropical province. Diversity decreases with
increasing latitude but there are no major distributional boundaries, with most Indo-west Pacific
species reaching as far south as North West Cape. This general distribution pattern has also been
documented for north-western Australian decapod crustaceans (Griffin and Yaldwyn, 1967;
Morgan, 1990), as well as echinoderms (Marsh and Marshall, 1983), corals (Veron and Marsh,
1985), molluscs (Wells, 1980), fishes (Blaber et al., 1985) and marine algae (Womersley, 1960).
Tropical and temperate faunas overlap on the western coast of Western Australia. There is
relatively high but decreasing species diversity, with the numbers of tropical species diminishing
with increasing latitude (Wilson and Allen, 1987). This has been documented for barnacles
(Jones, 1990b; Jones, 1993; Jones and Hewitt, 1995), as well as for other crustaceans (Jones,
1990d; Morgan and Wells, 1991; Jones and Morgan, 1993). The hydrological effects of the
Leeuwin Current on the south-western coast of Australia, including Rottnest Island, are well
documented (e.g. Hodgkin and Phillips, 1969; Cresswell and Golding, 1980). Due to the
influence of the southerly flowing, warm water Leeuwin Current, a significant tropical faunal
element is evident as far south as Rottnest Island (32°00’S) (Hodgkin et al., 1959; Black and
Johnson, 1983; Marsh, 1976; Hutchins, 1979; Wells, 1980; Jones, 1993a; Hutchins and Pearce,
1994). The Leeuwin Current can distribute the larvae of certain species around Cape Leeuwin
(34°22'S) and even eastward into the Great Australian Bight (Maxwell and Cresswell, 1981;
Wilson and Allen, 1987; Veron and Marsh, 1988; Walker, 1991). For example, Maxwell and
Cresswell (1981) recorded the occurrence of several demersal and pelagic tropical species in
the Great Australian Bight and suggested that the Leeuwin Current is responsible for the
presence of these species. Mass spawning during peak flow of the Leeuwin Current has been
used to explain the presence of corals at southern latitudes in south-western Australia (Wilson
and Allen, 1987), and may account for increased penetration of tropical species on the west
coast.
Compared to the northern Australian tropical province, the southern Australian warm-
temperate province exhibits decreased species diversity, high species endemicity, and a low
incidence of tropical species that decreases from west to east. In south-western Australia,
however, marine faunas show a stronger tropical component than could be predicted by latitude,
due to the influence of the Leeuwin Current. O’Hara and Poore (2000) also suggest that
localized regions of high winter temperatures in the Great Australian Bight between Esperance
and Eucla correspond with a region of low faunal exchange, perhaps allowing subtropical
species to persist further east than would otherwise be possible. Most of the temperate species
that occur along the south coast of western Australia reach as far west as Cape Leeuwin
(34°22'S) (Morgan and Jones, 1991).
Page 15
BIOGEOGRAPHY OF WESTERN AUSTRALIAN BARNACLES 493
The barnacle fauna of temperate south-western Australia clearly demonstrates this south-
western Australian pattern, as do other crustacean groups (Morgan and Jones, 1991; Stephenson
and McNeill, 1955), molluscs (Wells, 1980; Wilson and Allen, 1987), echinoderms (Clark,
1946; Rowe and Vail, 1982) and fishes (Wilson and Allen, 1987). In the temperate southern
Australian region, O’Hara and Poore (2000) have shown that species richness of Australian
marine echinoderms and decapods grades with latitude, from high in warm-temperate regions
around Sydney and Perth to low in cold-temperate southern Tasmania. Similarly, in South
African waters there is a progressive decline in invertebrate species richness from the warmer
south-east to the cooler north-west (Emanuel et al., 1992).
Barnacle zonation patterns
In tropical north-western Australia barnacles are a conspicuous component of littoral and
shallow-water environments but, conversely, are not a conspicuous component of such
environments in warm-temperate south-western Australia. On the shores of both tropical and
warm-temperate Western Australia members of the sessilian balanomorph families dominate.
There is only sparse pedunculate representation, with the three representatives (Lithotrya
valentiana and Ibla cumingi in the tropics and Ibla quadrivalvis in the warm-temperate) all
occurring in essentially refugial shore habitats.
In tropical north-western Australia, although sessile forms dominate the sub-littoral,
pedunculate forms become more numerous than in the littoral. In the warm-temperate sub-
littoral, however, whilst the pedunculate component is much larger than in the littoral,
pedunculates and sessile forms are almost equal in number, with pedunculates slightly
dominating. Species diversity is greatest in tropical north-western Australia in both the littoral
and sub-littoral, compared to warm-temperate south-western Australia.
No sessile species are pelagic in either tropical or temperate Western Australia. The temperate
pedunculates are slightly numerically dominant over the temperate pedunculates.
In north-western Australia zonation follows the chthamalid-tetraclitid-balanid trend
characteristic of the continental shores of tropical northern Australia (Jones, 1992b; Jones,
2003). Because chthamalids are absent from temperate south-western Australian shores (Jones,
1990c), zonation follows a simpler tetraclitid-balanid trend.
ACKNOWLEDGEMENTS
Sincere thanks are due to Woodside Energy Ltd for providing the generous funding for
the four-year Western Australian Museum/Woodside Energy Ltd Marine Biological Survey
of the Dampier Archipelago, of which this Workshop was a component. I also thank the
organisers and participants for a successful Workshop. I am especially indebted to Melissa
Hewitt, Western Australian Museum, and Professor Brian Morton, The University of Hong
Kong, for assistance with fieldwork. I also sincerely thank Mrs Angela and Dr Dick
Anderson for processing and databasing the barnacle specimens collected during the course
of the above. The co-operation of the following colleagues who provided data on various
cirripede collections in a number of Australian museums is also gratefully acknowledged –
Dr Penny Berents (Australian Museum, Sydney, New South Wales), Dr Gary Poore
(Melbourne Museum, Melbourne, Victoria), Dr Peter Davie (Queensland Museum,
Brisbane, Queensland) and Dr Wolfgang Zeidler (South Australian Museum, Adelaide,
South Australia).
Page 16
494D.S. JONES
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