Integrative and Comparative Biology
Integrative and Comparative Biology, volume 52, number 3, pp. 366–387
doi:10.1093/icb/ics100
Society for Integrative and Comparative Biology
SYMPOSIUM
Australian Barnacles (Cirripedia: Thoracica), Distributions and
Biogeographical Affinities
Diana S. Jones1
Western Australian Museum, 49 Kew Street, Welshpool, Western Australia 6106, Australia
From the symposium ‘‘Barnacle Biology: Essential Aspects and Contemporary Approaches’’ presented at the annual
meeting of the Society for Integrative and Comparative Biology, January 3–7, 2012 at Charleston, South Carolina.
1
E-mail: diana.jones@museum.wa.gov.au
Synopsis Currently, 279 barnacle species are recognized in Australia waters. The barnacle fauna of tropical Australia
exhibits high species diversity (221), with a high incidence of tropical species (87 Indo-west Pacific [IWP], 16 West
Pacific and 65 Indo-Malayan), a low species endemicity (8), and 44 cosmopolitan and 1 Australasian species. Conversely,
that of temperate Australia shows lower species diversity (129), with a lower incidence of tropical species (26 IWP, 10
West Pacific and 25 Indo-Malayan), higher species endemicity (23), 37 cosmopolitan, 6 Australasian species, and 3
Australasian/Antarctic species. Distributions corroborate the general patterns demonstrated by the shallow-water biota
of northern tropical and southern temperate Australian biogeographic provinces. Tropical and temperate provinces grade
into each other in a broad overlap zone along both the western and eastern Australian coasts. This overlap zone is
essentially a transitional region, with the gradual replacement of a tropical barnacle fauna in the north by a predominantly temperate barnacle fauna in the south. Both western and eastern Australian coasts are bounded by major
poleward-flowing warm currents that have considerable influence on the marine flora and fauna, distributing tropical
species of many taxa much farther south than could be predicted by latitude. Currently, 16 barnacle species introduced
into Australian waters are identified, although this number may increase in the future due to new port developments and
increased shipping arrivals.
Introduction
The island continent of Australia, lying between 9–
448S and 112–1548E, has a vast coastline of 34,218
km. Bounded by the Indian, Pacific and Southern
oceans on its western, eastern, and southern margins,
and the Arafura and Timor seas on its northern
margin, its complex biogeographical positioning has
resulted in diverse terrestrial and marine faunas.
First collections of barnacles in Australia were
made during early French expeditions of discovery
(Bonnemains and Jones 1990) but Darwin’s monographs (1852, 1854) initiated the documentation of
the barnacles of temperate Australia. During the late
19th and early 20th centuries, various expeditions
added to the knowledge of this fauna (e.g., Hoek
1883, 1907, 1913; Krüger 1914; Broch 1916, 1922,
1931). Pioneering studies by Pope (e.g., 1943, 1945,
1958, 1965) increased knowledge of cirripede distributions in eastern Australia, and since the mid 1980s,
comprehensive field collecting throughout Australian
waters by Jones greatly augmented documentation of
the barnacle fauna (e.g., Jones 1987, 1990a, 1990b,
1991, 1992a, 1992b, 1992c, 1993, 1994, 1998, 2003,
2004, 2010; Jones et al. 1990; Jones and Hewitt 1995,
1996, 1997; Jones and Berry 2000).
Two hundred and seventy-nine barnacle species
are known in Australian waters. This article discusses
the distributions and biogeographic affinities of the
barnacles of the tropical and temperate waters of
Australia. Abbreviations are as follows: WA
(Western Australia), SA (South Australia), Tas.
(Tasmania), Vic. (Victoria), NSW (New South
Wales), Qld (Queensland), and NT (Northern
Territory).
ß The Author 2012. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved.
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Biogeography of Australian barnacles
Marine biogeographical zonation of
Australia
Currents
Four oceanic and coastal currents in the Australasian
region are significant in shaping the climate and
marine environmental conditions of Australia,
namely, the Indonesian Throughflow, the Leeuwin
Current, the East Australian Current (EAC), and
the Antarctic Circumpolar Current (Fig. 1).
The Indonesian Throughflow is a series of currents
that carry water westward from the Pacific Ocean to
the Indian Ocean through the straits and deep passages of the Indonesian Archipelago. This warm
tropical water influences the character of the
Leeuwin Current, a poleward flowing, eastern
boundary current off the western and southern
coasts of Australia, which is the world’s longest
coastal current (45000 km) (Cresswell and Golding
1980). It originates near the North West Shelf on
Australia’s northwestern coast and is a broad body,
50 km wide and 200 m deep, of warm, relatively
low salinity water 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 flows to the
south intensify in autumn (March), are strongest in
late autumn/early winter (April–June), and disappear
in September–October (Feng et al. 2003). In the
autumn/early winter, the Leeuwin Current accelerates, rounds Cape Leeuwin (348270 S 1168220 E) in
southwestern WA, and continues as an eastward
Fig. 1 Oceanic and coastal currents of Australia
367
shelf current, the South Australian Current, along
the southern coast of Australia (Ridgeway and
Condie 2004; Middleton and Bye 2007). As the
Leeuwin Current travels poleward, it breaks into a
series of southward and eastward flowing eddies
(Feng et al. 2005), eventually dissipating in the
Tasman Sea and Southern Ocean.
The Leeuwin Current disperses tropical representatives of many taxa (e.g., asteroids, holothurians,
tuna, and tropical reef fishes) to the southwestern
and southern coasts of Australia, farther south than
could be predicted by latitude (Maxwell and
Cresswell 1981; Hutchins and Pearce 1994). It is
very different from the other Southern-Hemisphere
eastern-boundary currents, the Humboldt Current of
South America, and the Benguela Current of South
Africa, which are northward flowing, cool, and associated with upwelling. The Leeuwin Current roughly
parallels the EAC, which brings warm waters southward to 338S (Newcastle, NSW) before diverting as
eddies into the Tasman Sea.
The EAC is a complex western boundary system in
the southwestern Pacific off eastern Australia
(Ridgeway and Dunn 2003; Ridgeway and Hill
2009). It flows southward from 258S (near Fraser
Island, Qld) and begins to dissipate beyond 338S,
with remnants continuing to drift south. It provides
both the western boundary of the South Pacific Gyre
and the linking element between the Pacific and
Indian Ocean gyres (Speich et al. 2002). The EAC
is strongest in the austral summer (November–
February). It is weaker than other western boundary
368
currents and a series of mesoscale eddies dominate,
producing highly variable patterns of strength and
direction of currents (Bowen et al. 2005; Mata
et al. 2007). Long-term data indicate that the EAC
has strengthened and extended further southward
over the past 60 years, so that tropical waters from
the Coral Sea region are forced further south, warming the Tasman Sea (Ridgeway 2007). Evidence from
biological sources indicates southward range extensions of biota (Edgar et al. 1997; Pittock 2003;
Thresher et al. 2003; Ling et al. 2008), which have
been attributed to enhanced flow (Edyvane 2003).
Thus, Australia is unique among continents in that
both the western and eastern coasts are bounded by
major poleward-flowing warm currents that have
considerable influence on marine flora and fauna
(Richardson and Poloczanska 2009).
The Antarctic Circumpolar Current is the dominant feature of the Southern Ocean. It connects the
Atlantic, Pacific, and Indian Oceans in an eastward
flow, allowing water, heat, salt, and other properties
to flow from one to the other and is considered the
powerhouse of the global climate (www.csiro.au/
Outcomes/Climate/Australasian Ocean Currents). It
is confined by land between Tasmania and
Antarctic and this region features high oceanic nutrient production.
Australian biogeographic provinces
In Australia, the Tropic of Capricorn lies at
238260 2200 S, with latitudes to the south in the southern zone and those to the north in the tropics
D. S. Jones
(Fig. 2). A northern tropical and a southern temperate biogeographic province are recognized, which
overlap extensively on both western and eastern
coasts (Wilson and Allen 1987; O’Hara and Poore
2000; Poore 2001, 2004; Poore and O’Hara 2007;
Poore et al. 2008; Waters 2010).
The northern tropical province has a tropical
marine biota that is continuous with other parts of
the IWP. It exhibits high species diversity, a high
incidence of tropical species and low endemicity at
the species level (Wells 1980; Wilson and Allen
1987). Conversely, the southern temperate province
has lower species diversity and harbors much higher
numbers of endemic species, due to their long history of geographic isolation from other temperate
regions over geological time. For example, approximately 95% of molluscan species, 90% of echinoderm species, and 85% of fish species are unique
to these southern waters Australia (Poore 2001).
This high endemism is also apparent in Australia’s
temperate macroalgae (Phillips 2001).
In general, species diversity decreases with increasing latitude but there are no major distributional
boundaries. On the western coast, most IWP tropical
species extend to North West Cape, WA (218470 S),
and some as far south as the Houtman Abrolhos
Islands (288190 -298570 S), and on the eastern coast
approximately to Point Vernon, Qld (258140 53 S,
1528 490 E) (Jones 2003, 2010). However, the importance of the major currents in structuring marine
communities can be seen in the biogeographic distributions of many species, functional groups, and
Fig. 2 Northern tropical and southern temperate biogeographic provinces of Australia
369
Biogeography of Australian barnacles
communities. For example, tropical species can occur
much farther south at latitudes inhabited by wholly
temperate species due to the effects of the Leeuwin
and the East Australian currents (Maxwell and
Cresswell 1981; Morgan and Wells 1991; Dunlop
and Wooller 1986; O’Hara and Poore 2000;
Edyvane 2003; Griffiths 2003).
Table 1 Barnacles (Cirripedia: Thoracica) of Australian waters
Subclass CIRRIPEDIA Burmeister, 1834
Superorder THORACICA Darwin, 1854
Order Ibliformes Buckeridge and Newman, 2006
Suborder Iblomorpha Newman, 1987
Family Iblidae Leach, 1825
Genus Ibla Leach, 1825
Distributions and biogeographic affinities of barnacles in Australian waters
Ibla cumingi Darwin, 1852
Currently, 279 barnacle species are known in
Australian waters (Table 1). Fifty-four are cosmopolitan (C) species, 92 have IWP affinities and 21 West
Pacific (WP), and 76 Indo-Malayan (IM) affinities.
Six species are Australasian (AA), occurring in
Australia and New Zealand waters, and two are
Southern Ocean species (SAA), occurring in
Australia, New Zealand, and Antarctica. Twentyeight species are endemic (AE) to Australian waters
(Tables 2 and 3).
Family Idioiblidae Buckeridge and Newman, 2006
Ibla quadrivalvis Cuvier, 1817
Subfamily Idioiblinae Buckeridge and Newman, 2006
Genus Idioibla Buckeridge and Newman, 2006
Idioibla pygmaea Broch, 1922
Subfamily Chaetolepadinae Buckeridge and Newman, 2006
Genus Chaetolepas Stüder, 1889
Chaetolepas calcitergum Buckeridge and Newman, 2006
Order Lepadiformes Buckeridge and Newman, 2006
Suborder Heteralepdomorpha Newman, 1987
Family Heteralepadidae Nilsson-Cantell, 1921
Northern Australian tropical province
Genus Heteralepas Pilsbry, 1907
The tropical barnacle fauna is continuous with other
parts of the IWP region and exhibits the high species
diversity, high incidence of tropical species, and low
species endemicity pattern. It consists of 221 species,
44 of which are C species. Eighty-seven have IWP
affinities and 16 have WP and 65 IM affinities. One
species has AA affinities and eight are AE species
(Tables 2 and 3).
A dominant IWP faunistic element in the northern Australian tropical province also has been documented in other groups, e.g., Thalassinidea,
Anomura, and Brachyura (78%) (Morgan 1990);
Penaeidae (Dall 1957; Racek 1959); Portunidae
(Stephenson 1972); Stomatopoda (Stephenson and
McNeil 1955); Echinodermata (77%) (Marsh and
Marshall 1983); Mollusca (71%) (Wells 1980);
fishes (Blaber et al. 1985); and marine algae
(Womersley 1960). Similarly, a low AE element has
been documented in brachyuran and anomuran
decapods (17–22%) (Griffin and Yaldwyn 1967;
Morgan 1990; Morgan and Wells 1991); echinoderms
(13%) (Marsh 1976; Marsh and Marshall 1983); molluscs (10%) (Wilson and Allen 1987); corals (0%)
(Wilson and Allen 1987; Veron and Marsh 1988),
and fishes (13%) (Wilson and Allen 1987).
Currently, there are no specific field data regarding the barnacle faunas of the remote tropical northern and northeastern coasts of Australia, where
collecting is logistically extremely difficult. Field
data from north-western Australia indicate 101 species in 40 genera within 15 families, including 26 C,
Heteralepas adiposa Zevina, 1982
Heteralepas cornuta (Darwin, 1852)
Heteralepas dubia Broch, 1922
Heteralepas japonica (Aurivillius, 1892)
Heteralepas utinomii Newman, 1960
Genus Paralepas Pilsbry, 1907
Paralepas dannevigi (Broch, 1922)
Paralepas georgei Daniel, 1970
Paralepas intermedia (Hoek, 1907)
Paralepas minuta (Phillipi, 1836)
Paralepas morula (Hoek, 1907)
Paralepas palinuri urea Newman, 1960
Paralepas pedunculata (Hoek, 1883)
Paralepas quadrata (Aurivillius, 1894)
Paralepas scyllarusi Utinomi, 1967a
Paralepas tuberosa (Nilsson-Cantell, 1932)
Family Malacolepadidae Hiro, 1933
Genus Arcalepas Jones and Morton, 2009
Arcalepas brucei Jones and Morton, 2009
Suborder Lepadomorpha Pilsbry, 1916
Family Oxynaspididae Gruvel, 1905
Genus Oxynaspis Darwin, 1852
Oxynaspis celata Darwin, 1852 [includes Oxynaspis indica
Annandale, 1910]
Family Poecilasmatidae Annandale, 1910
Genus Poecilasma Darwin, 1852
Poecilasma dubium Hoek, 1907
(continued)
370
D. S. Jones
Table 1 Continued
Table 1 Continued
Poecilasma kaempferi Darwin, 1852
Subgenus Anatifa Bruguière, 1789
Genus Glyptelasma Pilsbry, 1907
Lepas Anatifa anatifera anatifera Linnaeus, 1758
Glyptelasma carinatum (Hoek, 1883)
Lepas Anatifa anatifera dentata Linnaeus, 1758
Glyptelasma gigas (Annandale, 1916)
Lepas Anatifa anatifera striata de Graef, 1952
Glyptelasma gracile (Hoek, 1883)
Lepas Anatifa anserifera Linnaeus, 1767
Glyptelasma hamatum Calman, 1919
Lepas Anatifa australis Darwin, 1852
Glyptelasma orientale Calman, 1919
Lepas Anatifa hillii (Leach, 1818)
Glyptelasma pilsbryi Calman, 1919
Lepas Anatifa indica Annandale, 1910
Glyptelasma rectum (Pilsbry, 1907)
Lepas Anatifa pectinata Spengler, 1793
Genus Megalasma Hoek, 1883
Lepas Anatifa testudinata Aurivillius, 1892
Megalasma minus (Annandale, 1906)
Genus Dosima Gray, 1825
Megalasma striatum Hoek, 1883
Dosima fascicularis Ellis and Solander, 1786
Genus Temnaspis Fischer, 1884
Genus Conchoderma Olfers, 1814
Temnaspis amygdalum (Aurivillius, 1894)
Conchoderma auritum (Linnaeus, 1767)
Temnaspis bathynomi (Annandale, 1906)
Conchoderma chelonophilum (Leach, 1818)
Temnaspis excavatum (Hoek, 1907)
Conchoderma hunteri (Owen, 1830)
Temnaspis fissum (Darwin, 1852)
Conchoderma virgatum (Spengler, 1790)
Temnaspis kilepoae Zevina, 1968
Genus Alepas Sander-Rang, 1829
Temnaspis tridens (Aurivillius, 1894)
Alepas pacifica Pilsbry, 1907
Temnaspis tridens asymmetrica Broch, 1947
Order Scalpelliformes Buckeridge and Newman, 2006
Genus Octolasmis Gray, 1825
Suborder Scalpellomorpha Newman, 1987
Octolasmis angulata (Aurivillius, 1894: 22) [includes O. aperta
Aurivillius, 1894: 24]
Family Calanticidae Zevina, 1978
Octolasmis aymonini (Lessona and Tapparone-Canefri, 1874)
Octolasmis cf bullata (Aurivillius, 1892)
Genus Calantica Gray, 1825
Calantica affinis Broch, 1922
Calantica darwini Jones and Hosie, 2009
Octolasmis clubii Daniel, 1953
Calantica studeri (Weltner, 1922)
Octolasmis cor (Aurivillius, 1892)
Calantica trispinosa (Hoek, 1883)
Octolasmis geryonophila geryonophila Pilsbry, 1907
Genus Crosnieriella Jones, 1998
Octolasmis hawaiense Pilsbry, 1907
Crosnieriella acanthosubcarinae Jones, 1998
Octolasmis hoeki (Stebbing, 1894)
Genus Scillaelepas Seguenza, 1876
Octolasmis indubia Newman, 1961
Scillaelepas cf fosteri Newman, 1980 (see Buckeridge, 1999: 528)
Octolasmis lowei (Darwin, 1852)
Genus Smilium Gray, 1825
Octolasmis neptuni neptuni (MacDonald, 1869)
Smilium nudipes (Annandale, 1916)
Octolasmis nierstraszi (Hoek, 1907)
Smilium peronii Gray, 1825
Octolasmis scuticosta Hiro, 1939
Smilium sinense (Annandale, 1910)
Octolasmis warwickii Gray, 1825 (includes Dichelaspis equina
Lanchester, 1902)
Smilium zancleanum (Withers, 1953)
Family Lithotryidae Gruvel, 1905
Octolasmis weberi (Hoek, 1907)
Genus Lithotrya Sowerby, 1822
Genus Dichelaspis Darwin, 1852
Lithotrya dorsalis (Ellis, 1786)
Dichelaspis orthogonia Darwin, 1852
Lithotrya nicobarica Reinhardt, 1850
Genus Trilasmis Hinds, 1844
Lithotrya valentiana (Gray, 1825)
Trilasmis eburnea Hinds, 1844
Family Scalpellidae Pilsbry, 1907
Family Lepadidae Darwin, 1852
Subfamily Scalpellinae Pilsbry, 1907
Genus Lepas Linnaeus, 1758
Genus Scalpellum Leach, 1817
Subgenus Nonfurcata Memmi, 1980
Scalpellum inerme Annandale, 1905
Lepas Nonfurcata nonfurcata (Nilsson-Cantell, 1927)
(continued)
(continued)
371
Biogeography of Australian barnacles
Table 1 Continued
Table 1 Continued
Scalpellum stearnsii Pilsbry, 1890
Trianguloscalpellum hirsutum (Hoek, 1883)
Subfamily Meroscalpellinae Zevina, 1978
Trianguloscalpellum michelottianum (Seguenza, 1876)
Genus Litoscalpellum Newman and Ross, 1971
Trianguloscalpellum moluccanum (Hoek, 1883)
Litoscalpellum giganteum (Gruvel, 1902)
Trianguloscalpellum regium regium (Hoek, 1883)
Litoscalpellum intermedium (Hoek, 1883)
Trianguloscalpellum rubrum (Hoek, 1883)
Litoscalpellum juddi (Calman, 1918)
Genus Teloscalpellum Zevina, 1978
Litoscalpellum nipponense (Pilsbry, 1907)
Teloscalpellum ecaudatum (Calman, 1918)
Genus Alcockianum Zevina, 1978
Teloscalpellum gracile (Hoek, 1907)
Alcockianum alcockianum (Annandale, 1906)
Teloscalpellum latisculum (Newman and Ross, 1971)
Alcockianum persona (Annandale, 1916)
Order Sessilia Lamarck, 1818
Genus Gymnoscalpellum Newman and Ross, 1971
Suborder Verrucomorpha Pilsbry, 1916
Gymnoscalpellum tarasovi Newman and Ross, 1971
Family Verrucidae Darwin, 1854
Genus Annandaleum Newman and Ross, 1971
Genus Altiverruca Pilsbry, 1916
Annandaleum laccadivicum (Annandale, 1906)
Altiverruca gibbosa (Hoek, 1883)
Annandaleum lambda (Annandale, 1910)
Altiverruca navicula (Hoek, 1913)
Subfamily Arcoscalpellinae Zevina, 1978
Costatoverruca Young, 1998
Genus Arcoscalpellum Hoek, 1907
Costatoverruca pacifica (Buckeridge, 1994)
Arcoscalpellum dubium (Hoek, 1883)
Cristallinaverruca Young, 1998
Arcoscalpellum gryllum Zevina, 1981
Cristallinaverruca cristallina (Gruvel, 1907)
Arcoscalpellum inum Zevina, 1981
Genus Metaverruca Pilsbry, 1916
Arcoscalpellum mendelevii Zevina, 1981
Metaverruca halotheca (Pilsbry, 1907) [includes M. recta (Aurivillius,
1898)]
Arcoscalpellum pertosum Foster, 1978
Metaverruca sculpta (Aurivillius, 1898)
Arcoscalpellum sociabile (Annandale, 1905)
Newmaniverruca Young, 1998
Arcoscalpellum truncatum (Hoek, 1883)
Newmaniverruca albatrossiana (Pilsbry, 1912)
Genus Planoscalpellum Zevina, 1978
Genus Rostratoverruca Broch, 1922
Planoscalpellum planum (Hoek, 1883)
Rostratoverruca intexta (Pilsbry, 1912) [includes Altiverruca conchula
(Hoek, 1913)]
Genus Weltnerium Zevina, 1978
Weltnerium poculum (Hoek, 1907)
Suborder Balanomorpha Pilsbry, 1916
Genus Verum Zevina, 1978
Superfamily Pachylasmatoidea Utinomi, 1968
Verum australicum (Hoek, 1883)
Family Pachylasmatidae Utinomi, 1968
Verum candidum (Hoek, 1907)
Subfamily Bathylasmatinae Newman and Ross, 1976
Verum novaezelandiae (Hoek, 1883)
Genus Bathylasma Newman and Ross, 1971
Verum proclive (Hoek, 1907)
Bathylasma alearum (Foster, 1978)
Verum virgatum (Hoek, 1907)
Genus Tetrachaelasma Newman and Ross, 1971
Genus Anguloscalpellum Zevina, 1978
Tetrachaelasma tasmanicum Buckeridge, 1999
Anguloscalpellum pedunculatum (Hoek, 1883)
Subfamily Hexelasmatinae Newman and Ross, 1976
Genus Amigdoscalpellum Zevina, 1978
Hexelasma Hoek, 1913
Amigdoscalpellum costellatum (Withers, 1935)
Hexelasma arafurae Hoek, 1913
Amigdoscalpellum daschae Zevina, 1981
Hexelasma nolearia Foster, 1978
Amigdoscalpellum elegans (Hoek, 1907)
Subfamily Pachylasmatinae Utinomi, 1968
Amigdoscalpellum torbenbenwolffi Zevina, 1981
Genus Eutomolasma Jones, 2000
Amigdoscalpellum vitreum (Hoek, 1883)
Eutomolasma chinense (Pilsbry, 1912)
Genus Trianguloscalpellum Zevina, 1978
Eutomolasma japonicum (Hiro, 1933)
Trianguloscalpellum annandalei (Calman, 1918)
Eutomolasma maclaughlinae Jones, 2000
Trianguloscalpellum hamulus (Hoek, 1907)
Genus Pachylasma Darwin, 1854
(continued)
(continued)
372
D. S. Jones
Table 1 Continued
Table 1 Continued
Pachylasma scutistriata Broch, 1922
Stomatolepas transversa Nilsson-Cantell, 1930
Genus Tetrapachylasma Foster, 1988
Genus Cylindrolepas Pilsbry, 1916
Tetrapachylasma aurantiacum (Darwin, 1854)
Cylindrolepas darwiniana Pilsbry, 1916
Tetrapachylasma ferrugomaculosa Jones, 1993
Genus Stephanolepas Fischer, 1886
Superfamily Chthamaloidea Darwin, 1854
Stephanolepas muricata Fischer, 1886
Family Catophragmidae Utinomi, 1968
Family Coronulidae Leach, 1817
Genus Catomerus Pilsbry, 1916
Genus Coronula Lamarck, 1802
Catomerus polymerus (Darwin, 1854)
Coronula diadema (Linnaeus, 1767)
Family Chthamalidae Darwin, 1854
Coronula reginae (Darwin, 1854)
Subfamily Notochthamalinae Foster and Newman, 1987
Genus Cetopirus Ranzani, 1817
Genus Chamaesipho Darwin, 1854
Cetopirus complanatus (Mörch, 1852)
Chamaesipho tasmanica Foster and Anderson, 1986
Chelolepas Ross and Frick, 2007
Nesochthamalus Foster and Newman, 1987
Chelolepas cheloniae (Monroe and Limpus, 1979)
Nesochthamalus intertextus (Darwin, 1854)
Genus Tubicinella Lamarck, 1802
Genus Octomeris Sowerby, 1825
Tubicinella major Lamarck, 1802
Octomeris brunnea Darwin, 1854
Genus Xenobalanus Steenstrup, 1852
Octomeris intermedia Nilsson-Cantell, 1921
Xenobalanus globicipitus Steenstrup, 1852
Subfamily Euraphiinae Newman and Ross, 1976
Superfamily Tetraclitoidea Gruvel, 1903
Genus Caudoeuraphia Poltarukha, 1997
Family Tetraclitidae Gruvel, 1903
Caudoeuraphia caudata (Pilsbry, 1916)
Subfamily Austrobalaninae Newman and Ross, 1976
Genus Euraphia Conrad, 1837
Genus Austrobalanus Pilsbry, 1916
Euraphia hembeli Conrad, 1837
Austrobalanus imperator (Darwin, 1854)
Microeuraphia Poltarukha, 1997
Genus Epopella Ross, 1970
Microeuraphia withersi (Pilsbry, 1916)
Epopella simplex (Darwin, 1854)
Subfamily Chthamalinae Darwin, 1854
Subfamily Tetraclitellinae Newman and Ross, 1976
Genus Chthamalus Ranzani, 1817
Genus Tetraclitella Hiro, 1939
Chthamalus antennatus Darwin, 1854
Tetraclitella costata (Darwin, 1854)
Chthamalus malayensis Pilsbry, 1916
Tetraclitella divisa (Nilsson-Cantell, 1921)
Superfamily Coronuloidea Leach, 1817
Tetraclitella multicostata (Nilsson-Cantell, 1930)
Family Chelonibiidae Pilsbry, 1916
Tetraclitella pilsbryi Utinomi, 1962
Subfamily Chelonibiinae Pilsbry, 1916
Tetraclitella purpurascens (Wood, 1815)
Genus Chelonibia Leach, 1817
Subfamily Newmanellinae Ross and Perreault, 1999
Chelonibia caretta (Spengler, 1790)
Genus Yamaguchiella Ross and Perreault, 1999
Chelonibia patula (Ranzani, 1818)
Subgenus Yamaguchiella Ross and Perreault, 1999
Chelonibia testudinaria (Linnaeus, 1758)
Yamaguchiella Yamaguchiella coerulescens (Spengler, 1790)
Family Platylepadidae Newman and Ross, 1976
Subgenus Neonrosella Jones, 2010
Genus Platylepas Gray, 1825
Yamaguchiella Neonrosella vitiata (Darwin, 1854)
Platylepas coriacea Monroe and Limpus, 1979
Subfamily Tetraclitinae Gruvel, 1903
Platylepas decorata Darwin, 1854
Genus Tesseropora Pilsbry, 1916
Platylepas hexastylos (Fabricius, 1798)
Tesseropora rapax Jones, 1993
Platylepas ophiophilius Lanchester, 1902
Tesseropora rosea (Krauss, 1848)
Genus Stomatolepas Pilsbry, 1910
Tesseropora wireni (Nilsson-Cantell, 1921)
Stomatolepas dermochelys Monroe and Limpus, 1979
Genus Tetraclita Schumacher, 1817
Stomatolepas elegans (Costa, 1838)
Tetraclita squamosa (Bruguière, 1789)
Stomatolepas praegustator Pilsbry, 1910
Superfamily Balanoidea Leach, 1817
(continued)
(continued)
373
Biogeography of Australian barnacles
Table 1 Continued
Table 1 Continued
Family Archaeobalanidae Newman and Ross, 1976
Acasta idiopoma Pilsbry, 1912
Subfamily Archaeobalaninae Newman and Ross, 1976
Acasta japonica Pilsbry, 1911
Genus Armatobalanus Hoek, 1913
Acasta purpurata Darwin, 1854
Armatobalanus allium (Darwin, 1854)
Acasta spongites (Poli, 1795)
Armatobalanus arcuatum Hoek, 1913
Acasta sulcata Lamarck, 1818 [includes Acasta serrata Hiro, 1937b]
Armatobalanus cepa (Darwin, 1854)
Genus Pectinoacasta Kolbasov, 1993
Armatobalanus filigranus (Broch, 1916)
Pectinoacasta pectinipes (Pilsbry, 1912)
Armatobalanus quadrivittatus (Darwin, 1854)
Subfamily Elminiinae Foster, 1982
Armatobalanus quinquivittatus (Hoek, 1913)
Genus Hexaminius Foster, 1982
Armatobalanus terebratus (Darwin, 1854)
Hexaminius foliorum Anderson et al., 1988
Genus Striatobalanus Hoek, 1913
Hexaminius popeiana Foster, 1982
Striatobalanus amaryllis (Darwin, 1854)
Austrominius Buckeridge, 1983
Striatobalanus bimae (Hoek, 1913)
Austrominius adelaidae (Bayliss, 1988)
Striatobalanu krugeri (Pilsbry, 1916)
Austrominius covertus (Foster, 1982)
Striatobalanus tenuis (Hoek, 1883)
Austrominius erubescens (Bayliss, 1994)
Genus Solidobalanus Hoek, 1913
Austrominius flindersi (Bayliss, 1994)
Solidobalanus auricoma (Hoek, 1913)
Austrominius modestus (Darwin, 1854)
Solidobalanus ciliatus (Hoek, 1913)
Austrominius placidus (Bayliss, 1994)
Solidobalanus compressus (Hoek, 1913)
Family Pyrgomatidae Gray, 1825
Solidobalanus socialis (Hoek, 1883)
Subfamily Pyrgomatinae Gray, 1825
Solidobalanus solidus (Broch, 1931)
Tribe Hoekiini Ross and Newman, 1995
Genus Membranobalanus Hoek, 1913
Genus Australhoekia Ross and Newman, 1995
Membranobalanus cuneiformis (Hiro, 1936)
Australhoekia cardenae Ross and Newman, 2000
Genus Conopea Say, 1822
Tribe Pyrgomatini Ross and Newman, 1995
Conopea calceolus (Ellis, 1758)
Genus Cantellius Ross and Newman, 1973
Conopea cymbiformis (Darwin, 1854)
Cantellius acutum (Hiro, 1938)
Conopea dentifer (Broch, 1922)
Cantellius euspinulosum (Broch, 1931)
Conopea mjobergi (Broch, 1916)
Cantellius gregarious (Sowerby, 1823)
Conopea navicula (Darwin, 1854)
Cantellius iwayama (Hiro, 1938)
Subfamily Acastinae Kolbasov, 1993
Cantellius pallidus (Broch, 1931)
Genus Archiacasta Kolbasov, 1993
Cantellius secundus (Broch, 1931)
Archiacasta spinitergum (Broch, 1931)
Cantellius septimus (Hiro, 1938)
Genus Neoacasta Kolbasov, 1993
Cantellius sumbawae Hoek, 1913
Neoacasta glans (Lamarck, 1818)
Cantellius tredecimus (Kolosváry, 1947)
Neocasta laevigata (Gray, 1825)
Genus Creusia Leach, 1817
Genus Euacasta Kolbasov, 1993
Creusia spinulosa Leach, 1818
Euacasta antipathidus (Broch, 1916)
Genus Galkinia Ross and Newman, 1995
Euacasta dofleini (Krüger, 1911)
Galkinia indica (Annandale, 1924)
Euacasta porata (Nilsson-Cantell, 1921)
Genus Hiroa Ross and Newman, 1973
Euacasta zuiho (Hiro, 1936)
Hiroa stubbingsi Ross and Newman, 1973
Genus Acasta Leach, 1817
Genus Darwiniella Anderson, 1992
Acasta conica Hoek, 1913
Darwiniella conjugatum (Darwin, 1854)
Acasta cyathus Darwin, 1854
Genus Nobia Sowerby, 1823
Acasta echinata Hiro, 1937a
Nobia grandis Sowerby, 1839
Acasta fenestrata Darwin, 1854
Genus Arossella Anderson, 1993
Acasta hirsuta Broch, 1916
Arossella projectum (Nilsson-Cantell, 1938)
(continued)
(continued)
374
Table 1 Continued
Genus Pyrgoma Leach, 1817
Pyrgoma cancellata Leach, 1818
Genus Savignium Leach, 1825
Savignium crenatum (Sowerby, 1823)
Genus Trevathana Anderson, 1992
Trevathana dentatum (Darwin, 1854)
Genus Wanella Anderson, 1993
Wanella andersonorum Ross, 1999
Wanella milleporae (Darwin, 1854)
Subfamily Megatrematinae Holthuis, 1982
Tribe Pyrgominini Ross and Pitombo, 2002
Genus Pyrgomina Ross and Pitombo, 2002
Pyrgomina djanae Ross and Pitombo, 2002
Family Balanidae Leach, 1817
Subfamily Amphibalaninae Pitombo, 2004
Genus Amphibalanus Pitombo, 2004
Amphibalanus amphitrite (Darwin, 1854)
Amphibalanus cirratus (Darwin, 1854)
Amphibalanus improvisus (Darwin, 1854)
Amphibalanus littoralis (Ren and Liu, 1978)
Amphibalanus poecilotheca (Krüger, 1911)
Amphibalanus reticulatus Utinomi, 1967b
Amphibalanus variegatus Darwin, 1854
Amphibalanus zhujiangensis (Ren, 1989)
Subfamily Balaninae Leach, 1817
Genus Balanus Da Costa, 1778
Balanus trigonus Darwin, 1854
Genus Fistulobalanus Zullo, 1984
Fistulobalanus albicostatus (Pilsbry, 1916)
Fistulobalanus pallidus (Darwin, 1854)
Subfamily Megabalaninae Newman, 1979
Genus Austromegabalanus Newman, 1979
Austromegabalanus nigrescens (Lamarck, 1818)
D. S. Jones
45 IWP and 25 IM species, and 6 AE species (Jones
2003).
More specifically, the fauna of the vast and poorly
studied Kimberley region of WA (138440 S-188000 S,
1268470 E-1228150 E) contains 56 shallow-water species in 22 genera within eight families presently
documented (Jones 1992a; Jones and Hewitt 1997),
including 2 C, 46 IWP, 7 IM, and 1 AE species
(Jones 2003). At the Dampier Archipelago
(208200 S–208450 S, 1168240 E-1178050 E), 49 species in
24 genera within 11 families, including 10 C, 27
IWP, and 8 IM species, and 4 AE species are recorded (Jones 2003, 2004). In the North West
Cape area, 44 species in 20 genera within 11 families
have been documented from the Montebello Islands
(208270 S 1158310 E), the Muiron Islands (21.668S
114.328E) and the eastern shores of Exmouth Gulf
(218 550 S 1148230 E) (Jones and Hewitt 1996; Jones
and Berry 2000), including 4 C, 38 IWP, and 2 IM
species, no AE species being recorded (Jones 2003,
2004).
Tetraclita squamosa (Bruguière 1789) and
Caudoeuraphia caudata (Pilsbry 1916) are examples
of tropical barnacles commonly occurring in the littoral across the northern Australian tropical province. Tetraclita squamosa extends from Red Bluff,
Kalbarri, WA (278540 S 1148260 E), across the NT to
Point Vernon, Qld (258140 53 S, 1528 490 E) (Jones
1992a, 2004, 2010). Similarly, C. caudata extends
from the Dampier Archipelago, WA (208200 S
1168240 E), to Point Vernon (Jones 2004, 2010).
Examples of tropical endemic species are Calantica
darwini Jones and Hosie (2009) collected north of
Port Hedland, WA (188300 S 118836E to 188310 S
1188370 E, depth 196 km) and Crosnieriella acanthosubcarinae (Jones 1998) from northeastern Qld
(228270 S 1528150 E, depth 175m) (Jones 1998; Jones
and Hosie 2009).
Genus Megabalanus Hoek, 1913
Megabalanus ajax (Darwin, 1854)
Southern Australian temperate province
Megabalanus coccopoma (Darwin, 1854)
The southern Australian temperate barnacle fauna
exhibits lower species diversity, a low incidence of
tropical species, and high endemicity of species. It
comprises 129 species, of which 37 are C and 26
have IWP, 10 have WP, and 25 have IM affinities.
Six species have AA and 2 have SAA affinities, and
23 are AE (Tables 2 and 3).
In south-western Australia, 44 barnacle species in
24 genera and 12 families have been recorded (Jones
1990b, 2003), with 21 being C, and 10 I with WP
and 3 with IM affinities. Seven are AE species and
three have AA affinities. At Albany (358020 S
1178540 E), of 31 species documented, three are
Megabalanus concinnus (Darwin, 1854)
Megabalanus occator (Darwin, 1854)
Megabalanus rosa (Pilsbry, 1916)
Megabalanus tintinnabulum (Linnaeus, 1758)
Megabalanus validus (Darwin, 1854)
Megabalanus volcano (Pilsbry, 1916)
Megabalanus zebra (Darwin, 1854)
Genus Notomegabalanus Newman, 1979
Notomegabalanus algicola (Pilsbry, 1916)
Notomegabalanus krakatauensis (Nilsson-Cantell, 1934)
375
Biogeography of Australian barnacles
Table 2 Biogeographic affinities of Australian barnacles
Table 2 Continued
Biogeograpic affinities
Biogeograpic affinities
Genus
Species
Ibla
cumingi
C IWP WP IM AE AA SAA
Genus
IWP
Species
C IWP WP IM AE AA SAA
IWP
indubia
quadrivalvis
AE
lowei
Idioibla
pygmaea
AE
neptuni neptuni
IWP
Chaetolepas
calcitergum
AE
nierstraszi
IWP
Heteralepas
adiposa
cornuta
WP
AA
dubia
IWP
AE
utinomi
Paralepas
IM
dannevigi
minuta
weberi
IWP
orthogonia
IWP
Trilasmis
eburnea
IWP
Lepas (Anatifa)
IM
IM
IWP
anatifera anatifera
C
anatifera dentata
C
anatifera striata
C
anserifera
C
palinuri urae
WP
australis
C
pedunculata
WP
hillii
C
IM
quadrata
scyllarusi
WP
tuberosa
WP
Arcalepas
brucei
Oxynaspis
celata
Poecilasma
dubium
Glyptelasma
IWP
Dichelaspis
C
morula
warwickii
Lepas (Nonfurcata) nonfurcata
AE
georgei
intermedia
IM
scuticosta
C
japonica
C
AE
C
pectinata
C
testudinata
C
Dosima
fascicularis
C
Conchoderma
auritum
C
chelonophilum
C
IWP
kaempferi
C
hunteri
carinatum
C
virgatum
gigas
IM
Alepas
pacifica
gracile
IM
Calantica
affinis
hamatum
IWP
Indica
C
IWP
C
IWP
IM
AE
darwini
IM
orientale
studeri
IM
trispinosa
IM
pilsbryi
C
rectum
C
Crosnieriella
acanthosubcarinae
Megalasma
minus
C
Scillaelepas
cf fosteri
striatum
IWP
Smilium
nudipes
IM
Temnaspis
amygdalum
IWP
peronii
IM
sinense
IM
IM
bathynomi
excavatum
IWP
fissum
IWP
tridens
Lithotrya
C
IM
tridens asymmetrica
Octolasmis
Scalpellum
IWP
angulata
dorsalis
IWP
valentiana
IWP
IWP
stearnsii
IM
IM
intermedium
clubii
IM
juddi
Litoscalpellum
IWP
giganteum
C
WP
IM
WP
nipponense
geryonophila geryonophila C
hoeki
IM
inerme
aymonini
Alcockianum
alcockianum
Gymnoscalpellum
tarasovi
WP
hawaiense
C
nicobarica
cf bullata
cor
IM
zancleanum
IWP
kilepoae
AE
WP
persona
C
(continued)
IWP
IM
SAA
(continued)
376
D. S. Jones
Table 2 Continued
Table 2 Continued
Biogeograpic affinities
Genus
Species
Annandaleum
laccadivicum
IWP
lambda
IWP
Arcoscalpellum
Biogeograpic affinities
C IWP WP IM AE AA SAA
WP
dubium
Genus
Species
Tetrapachylasma
aurantiacum
Catomerus
C IWP WP IM AE AA SAA
WP
ferrugomaculosa
AE
polymerus
AE
gryllum
AE
Chamaesipho
tasmanica
inum
AE
Nesochthamalus
intertextus
IWP
AE
Octomeris
brunnea
IWP
mendeleevi
IM
pertosum
IWP
sociabile
Caudoeuraphia
AE
intermedia
IM
caudata
IM
truncatum
IM
Euraphia
hembeli
IWP
Planoscalpellum
planum
IM
Microeuraphia
withersi
IWP
Weltnerium
poculum
IM
Chthamalus
antennatus
Verum
australicum
IM
IM
candidum
proclive
IM
virgatum
IM
Anguloscalpellum
pedunculatum
Amigdoscalpellum
costellatum
WP
C
IWP
coriacea
IWP
C
IWP
ophiophilius
Stomatolepas
WP
dermochelys
C
elegans
C
praegustator
C
IWP
transversa
hamulus
hirsutum
IM
IM
Cylindrolepas
darwiniana
IM
Stephanolepas
muricata
Coronula
diadema
C
reginae
C
Cetopirus
complanatus
C
Chelolepas
cheloniae
C
IM
moluccanum
Teloscalpellum
C
testudinaria
IWP
torbenwolffi
Trianguloscalpellum annandalei
C
patula
hexastylos
WP
vitreum
caretta
decorata
IWP
elegans
regium regium
Platylepas
C
daschae
michelottianum
Chelonibia
IWP
novaezelandiae
AE
IWP
malayensis
C
rubrum
IM
ecaudatum
IM
gracile
IM
SAA
latisculum
C
C
IM
IM
Tubicinella
major
C
Xenobalanus
globicipitus
C
Austrobalanus
imperator
AE
Epopella
simplex
AE
Tetraclitella
Altiverruca
gibbosa
navicula
IWP
costata
IM
Costatoverruca
pacifica
IWP
divisa
IM
Cristallinaverruca
cristallina
IWP
multicostata
IM
Metaverruca
halotheca
IWP
pilsbryia
IM
sculpta
IWP
Newmaniverruca
albatrossiana
IWP
Rostratoverruca
intexta
IWP
Bathylasma
alearum
WP
Tetrachaelasma
tasmanicum
WP
Hexelasma
arafurae
Eutomolasma
chinense
IWP
vitiate
IWP
Tesseropora
rapax
AE
AA
rosea
IM
wireni
AA
nolearia
Pachylasma
coerulescens
IM
IM
WP
japonicum
AA
purpurascens
Yamaguchiella
Tetraclita
squamosa
IWP
Armatobalanus
allium
IWP
IM
arcuatum
maclaughlinae
IWP
cepa
IWP
scutistriata
IWP
filigranus
IWP
(continued)
(continued)
377
Biogeography of Australian barnacles
Table 2 Continued
Table 2 Continued
Biogeograpic affinities
Genus
Species
terebratus
IWP
amaryllis
IWP
IWP
secundus
IWP
septimus
IWP
sumbawae
krugeria
IM
tredecimus
IWP
Creusia
spinulosa
IWP
Galkinia
indica
IWP
auricoma
IWP
C
IWP
IM
compressus
IWP
socialis
solidus
Membranobalanus cuneiformis
calceolus
WP
Hiroa
stubbingsi
Darwinella
conjugatum
IWP
IWP
Nobia
grandis
IM
Arossella
projectum
IM
Pyrgoma
cancellata
IWP
Savignium
crenatum
IWP
C
IM
IWP
Trevathana
dentatum
IWP
dentifer
IWP
Wanella
andersonorum
IWP
mjobergi
IWP
milleporae
IWP
navicula
IWP
Pyrgominini
djanae
Amphibalanus
amphitrite
spinitergum
Neocasta
glans
IM
IWP
AE
IWP
a
improvisus
dofleini
IM
poecilothecaa
porata
IM
reticulatusa
zuiho
IM
variegatus
conica
IM
zhujiangensisa
C
IM
echinata
Balanus
trigonus
Fistulobalanus
albicostatusa
IWP
fenestrata
pallidus
hirsuta
IM
Austromegabalanus nigrescens
idiopoma
IM
Megabalanus
japonica
IM
coccopomaa
purpurata
IM
concinnusa
spongites
C
littoralisa
AE
antipathidus
cyathus
C
cirratus
IWP
laevigata
C
IM
AA
IM
C
IWP
C
AE
IWP
occator
C
IWP
a
IWP
rosaa
Pectinoacasta
pectinipes
IWP
tintinnabulum
Hexaminius
foliorum
AE
IM
IWP
ajax
sulcata
Austrominius
IM
cymbiformis
Archiacasta
Acasta
WP
pallidus
IM
ciliatus
Euacasta
C IWP WP IM AE AA SAA
bimae
tenuis
Conopea
Species
iwayama
IM
quinquivittatus
Solidobalanus
Genus
IWP
quadrivittatus
Striatobalanus
Biogeograpic affinities
C IWP WP IM AE AA SAA
IWP
IM
C
IM
validus
a
popeiana
AE
volcano
adelaidae
AE
zebraa
IWP
covertus
AE
algicolaa
IIWP
erubescens
AE
flindersi
AE
Cantellius
acutum
euspinulosum
gregarius
TOTAL
AE
placidus
cardenae
krakatauensisa
AA
modestus
Australhoekia
Notomegabalanus
IM
WP
WP
IWP
IM
(continued)
279
IA
54 92
21
76 28 6
2
Notes: C, cosmopolitan species; IWP, Indo-west Pacific species (extending from eastern Africa to Hawaii); WP, West Pacific species
(extending from eastern Australia to Hawaii); IM, Indo/Malayan species (extending from the eastern Indian Ocean, Indo-Malayan
Archipelago, Australia and New Guinea, to Japan); AE, Australian
endemic species (only occurring in Australia); AA, Australasian species (occurring in Australian and New Zealand); SAA, Australasian/
Antarctic species (occurring in Australia, New Zealand and
Antarctica).
a
Introduced species.
378
D. S. Jones
Table 3 Biogeographic affinities of barnacles of northern, southern, western, and eastern Australia
Species numbers C
IWP WP IM AE AA SAA
279
54 92
21
76 28 6
N. Australia 221
44 87
16
65 8
1
0
129
37 26
10
25 23 6
2
W. Australia 189
41 73
4
56 12 3
0
205
47 62
20
47 21 6
2
Australia
S. Australia
E. Australia
22
Notes: C, cosmopolitan species; IWP, Indo-west Pacific species (extending from eastern Africa to Hawaii); WP, West Pacific species
(extending from eastern Australia to Hawaii); IM, Indo/Malayan species (extending from the eastern Indian Ocean, Indo-Malayan
Archipelago, Australia and New Guinea, to Japan); AE, Australian
endemic species (only occurring in Australia); AA, Australasian species (occurring in Australia and New Zealand); SAA, Australasian/
Antarctic species (occurring in Australia, New Zealand and
Antarctica).
tropical IWP and six AE (Jones 1990b, 2003). A
higher endemic element has been documented in
other groups, e.g., decapods (77%) (Morgan and
Jones 1991); stomatopods (Stephenson and McNeill
1955); molluscs (95%) (Wells 1980; Wilson and
Allen 1987); echinoderms (90%) (Clark 1946; Rowe
and Vail 1982), and fishes (85%) (Wilson and Allen
1987).
IWP species representation in the southern
Australian temperate province decreases from west
to east while most temperate species occurring
along the southern coast of WA reach as far west
as 348270 S (Morgan and Jones 1991). Currently,
there are no comparable field data regarding the
shallow-water barnacle faunas occurring along
remote temperate southern and southeastern coasts
of Australia as, again, collecting is logistically extremely difficult.
Austrobalanus
nigrescens
(Lamarck
1818),
Catomerus
polymerus
(Darwin
1854),
and
Chthamalus antennatus (Darwin 1854) are examples
of barnacle species endemic to southern Australian
waters. Austrobalanus nigrescens occurs from
Kalbarri, WA (278420 S 1148100 E), across southern
Australia and northward to Double Island Point,
Qld (258560 S 1538110 E); Chthamalus antennatus
from Eucla, WA (318410 S 1288530 E), to Cooee Bay,
Qld (238080 S 1508450 E); and Catomerus polymerus
from the Eyre Peninsula, SA (34.058S 135.048E), to
Ballina Headland, NSW (288520 S 1538360 E)
(Jones1990b, 2010).
The overlap zones
The western and eastern coasts of Australia harbor
diverse, distinct barnacle faunas. The western
barnacle fauna comprises 189 species, of which 41
have C, 73 have IWP, 4 have WP and 56 have IM
affinities. Twelve species are AE and three have AA
affinities (Tables 2 and 3); that of the eastern coast of
Australia is composed of 205 species, of which 47 are
C, 62 have IWP, 20 have WP, and 47 have IM affinities. Twenty-one species are AE, six have AA, and
two SAA affinities (Tables 2 and 3).
In the western and eastern overlap zones, numbers
of tropical species diminish with increasing latitude
(Wilson and Allen 1987). In the western overlap
zone, the percentage of IWP barnacle species at
Shark Bay (258560 S 1138320 E) reduces from 55%
to 15% at Rottnest Island (328000 S 1158300 E)
(Jones 1990a, 1993; Jones and Hewitt 1995).
Similarly, a reduction in the IWP element is documented in other groups, e.g., 74% of the decapod
fauna at Shark Bay (Jones 1990c) and 39% of the
marine crustacean fauna of Rottnest Island (Jones
and Morgan 1993) are tropical IWP species.
A recognizable endemic component occurs in the
western overlap zone, but the proportion of endemics varies between marine groups, e.g., 18% of
the Shark Bay decapod fauna (Jones 1990c) and 48%
of the marine crustacean fauna of Rottnest Island
(Jones and Morgan 1993) are endemic, but less
than 10% of prosobranch molluscs (Wells 1980)
and 20% of shallow-water asteroids (Marsh 1976)
are endemic. At Shark Bay (Jones 1990a; Jones and
Hewitt 1995) and Rottnest Island (Jones 1993), 22%
and 23% of the barnacle species, respectively, are
endemic. Most of these endemic species have at
least part of their range in the western overlap
zone and often achieve their greatest numbers there
(Wells 1980; Wilson and Allen 1987).
Paralepas georgei (Daniel 1970), Tesseropora rapax
(Jones 1993), and Tetrapachylasma ferrugomaculosa
(Jones 1993) are examples of barnacle species endemic to the western overlap zone. Paralepas georgei
attaches to the gills of the Western Rock Lobster,
Panulirus cygnus, which itself is endemic to the western coast of WA. Tesseropora rapax and T. ferrugomaculosa have limited distributions at Rottnest
Island and along the mid-western coast.
The Leeuwin Current extends the southern latitudinal distributional limits of various marine taxa
down the western coast. The Houtman Abrolhos
Islands (288190 –298570 S) are generally considered to
be the southern-most limit of the tropical marine
biota (Wells 1980; Wilson and Allen 1987). Coral
reefs are richly developed and marine faunas occurring there are essentially tropical (Montgomery 1931;
Wilson and Marsh 1979; Wells 1980; Marsh and
Marshall 1983; Veron and Marsh 1988).
379
Biogeography of Australian barnacles
At Rottnest Island, Pocillopora damicornis (Linnaeus
1758) forms one of the most southerly developments
of reefs in the world and its associated symbiotic
decapod crustacean fauna is similar to that found
in many other tropical localities across the IWP
(Black and Prince 1983). A substantial proportion
of other marine faunas at Rottnest Island, including
zoanthids, echinoids, gastropods, and fishes, are of
tropical origin (Hodgkin et al. 1959; Black and
Johnson 1983; Hutchins 1994; Wells 1980).
When shallow-water and deep-water barnacles are
considered, of the 73 species of IWP barnacles
known to occur on the northern and western
coasts of WA, 10 reach Cape Leeuwin (348270 S
1168220 E) and 6 extend onto the southern
Australian coast (358S) (Jones 1990b, 1990c, 1992a,
1993, 2003, 2004; Jones and Hewitt 1995, 1996, 1997;
Jones et al. 1990; Jones and Berry 2000). Similar
trends can be demonstrated in other groups; of 308
tropical prosobranch gastropod species, 9 reach Cape
Leeuwin and 5 extend onto the southern coast (Wells
1980); of 318 hermatypic corals, 25 reach as far
south as Rottnest Island and 9 occur on the southern
coast (Veron and Marsh 1988); certain tropical echinoderm species extend into the Great Australian
Bight (Maxwell and Cresswell 1981).
On the eastern Australian coast, south-eastern
Queensland represents a transitional area between
the tropical and southern temperate provinces and
this is reflected in the composition of the barnacle
fauna. Seventy-three barnacle species are recorded,
with 22 C, 25 IWP, 9 IM, and 2 WP species (Jones
1992c, 2010). Three species have AA affinities and 12
are AE species. These figures demonstrate the influence of the tropical northern fauna and the 12 endemics reflecting the southern influence in this
transitional zone.
The tropical chthamalids, C. caudata (Pilsbry
1916) and Microeuraphia withersi (Pilsbry 1916),
extend from Point Vernon (258140 S 1528 490 E)
northward across the NT and to the Dampier
Archipelago, WA (208200 S 1168240 E), with
Chthamalus malayensis (Pilsbry 1916) further extending to Shark Bay (258560 S 1138320 E) (Jones 1990a,
2003, 2004, 2010). Conversely, their southern counterpart, the endemic Chthamalus antennatus (Darwin
1854) extends from Cooee Bay (238080 S 1508450 E)
southward then westward to Eucla, WA (318410 S
1288530 E) (Jones 2010). A similar pattern can be
demonstrated for the tropical tetraclitid, Tetraclita
squamosa (Bruguière 1789), from Point Vernon to
Red Bluff, Kalbarri, WA (278540 S 1148260 E), while
the southern Tetraclitella purpurescens (Wood 1815)
and Tesseropora rosea (Krauss 1848) extend from
Double Island Point (258560 S 1538110 E) and
Bustard Heads (248010 S 1518460 E) southward then
westward to Red Bluff, Kalbarri, WA, and Cottesloe,
WA (318590 S 1158 450 E), respectively (Jones 1990b,
2004, 2010). The northern tropical iblomorph, Ibla
cumingi (Darwin 1852), occurs from Point Vernon
northward, across the NT to Burnside Island,
Exmouth Gulf, WA (228060 S 114830.800 E), while its
southern temperate counterpart, Ibla quadrivalvis
(Cuvier 1817), extends from Currumbin (288080 S
1538290 E) to Bunbury, WA (338190 S 1158390 E)
(Jones 1990b, 2010). The endemic malacolepadid,
Arcalepas brucei (Jones and Morton 2009), is only
known from Moreton Bay, Qld (278280 0000 S,
1538280 0000 E) (Jones and Morton 2009).
Introduced species
Records of introduced barnacles in Australian waters
are not numerous. Pertinent literature documenting
fouling and introduced Australian barnacle species
was reviewed by Jones (1992b). Subsequent publications have documented introductions (Hass and
Jones 1999; Jones 2003, 2004; Huisman et al. 2008;
Wells et al. 2009; Yamaguchi et al. 2009), mainly
focusing on Western Australian introductions.
Information relating to introduced barnacle species
is also contained in unpublished reports to industry
and other stakeholders (D. S. Jones, unpublished
data). Currently, 16 species are recognized as introductions into Australian waters: Tetraclitella pilsbryi,
Striatobalanus krugeri, Amphibalanus improvisus, A.
littoralis, A. poecilotheca, A. reticulatus, A. zhujiangensis, Fistulobalanus albicostatus, Megabalanus coccopoma, M. concinnus, M. occator, M. rosa, M.
volcano, M. zebra, Notomegabalanus algicola, and N.
krakatauensis (Table 2). This number may well increase with a number of new ports being developed
in Australia and therefore a concomitant increase in
future shipping arrivals.
Conclusions
This brief overview of the distributions and biogeographic affinities of Australian barnacles presents all
data available to date. There are major gaps in information due to the vastness of the Australian
coastline (34,218 km), the logistics, and costs associated with accessing remote areas and the scarcity of
cirripede workers. However, comprehensive field collecting in WA and southeastern Queensland, plus
data from the literature and material in Australian
museum collections, allows some general statements
to be made.
380
Distributions corroborate the general patterns
demonstrated for the shallow-water biota of the
northern
tropical
and
southern
temperate
Australian biogeographic provinces. The barnacle
fauna of the northern Australian tropical province
is continuous with other parts of the IWP and exhibits high species diversity, a high incidence of tropical species, and low endemicity at the species level.
Conversely, the southern Australian temperate barnacle fauna exhibits lower species diversity, a low
incidence of tropical species, and high endemicity
of species. The IWP element constitutes the bulk of
the tropical Australian shallow-water barnacle fauna,
but representation of IWP species in the southern
Australian temperate province is low and decreases
from west to east.
Tropical and temperate provinces grade into each
other in a broad overlap zone along both the western
and eastern Australian coasts. This overlap zone is
essentially a transitional region, with the gradual replacement of a tropical barnacle fauna in the north
by a predominantly temperate barnacle fauna in the
south. Most tropical IWP species reach as far south
as North West Cape (218470 S) on the western coast.
The northern Australian tropical province thus extends to about 228S inshore and to about 298S at the
Houtman Abrolhos (288190 –298570 S). On the eastern
coast, the northern Australian tropical province extends to approximately Point Vernon, Qld (258150 S,
1528490 E). In eastern Australia, the northern limit of
temperate species is Cooee Bay, Qld (238080 S
1508450 E), while in the west it is Red Bluff,
Kalbarri, WA (278540 S 1148260 E).
The barnacle faunas of western and eastern
Australian coasts are diverse and distinct. On western
coasts, IWP, IM, and C species dominate and AE,
WP, and AA species have low representation, with
SAA species not represented. On eastern Australian
coasts, IWP, IM, C, AE, and WP species dominate,
with AA and SAA species having low representation.
Both the western and eastern Australian coasts are
bounded by major poleward-flowing warm currents,
which have considerable influence on the marine
flora and fauna. Tropical IWP barnacle species, and
many other taxa, are distributed to the southwestern
and southern coasts of Australia, much farther south
than could be predicted by latitude, or by the warm,
southward-flowing Leeuwin Current. A significant
tropical IWP element is evident as far south as
Rottnest Island (328000 S) and a number of tropical
species range farther south into the Great Australian
Bight (398000 S). While evidence is beginning to
emerge that southward range extensions of biota in
eastern Australia are attributable to an enhanced
D. S. Jones
EAC, no range extensions of barnacles have been
reported to date.
Sixteen barnacle species are currently recognized
as introductions into Australian waters, but this
number may increase with the development of a
number of new port facilities.
Acknowledgments
I sincerely thank Professor John Zardus for his tremendous efforts in organizing the symposium
Barnacle Biology: Essential Aspects and Contemporary
Approaches. I also thank Professor John Buckeridge
and an anonymous reviewer for pertinent comments
that significantly improved an earlier draft of this
article. I acknowledge the CSIRO, Australia, for
their kind permission to reproduce Figure 1. I also
thank all the conference organizers and the symposium participants for such a successful and enjoyable
meeting.
Funding
I wish to acknowledge the Society for Integrative and
Comparative Biology for providing generous funding
that allowed my attendance at the symposium.
Funding for the work associated with data collection
and completion of this paper has been generously provided through the following sources: the Western
Australian Museum (1980 to present); Australian
Museum Trust Postgraduate Scholarship (1984);
Associate Professorship, Musém national d’Histoire
naturelle, Paris (1994, 1997, 2000); Department of
Australian Heritage and CSIRO (1996–2005),
National Ports Survey Project (1996–2005);
Woodside Energy Ltd (1998–ongoing); Centre for
Research on Introduced Marine Pests (CRIMP),
CSIRO (1999); Gascoyne Development Commission
(1999); Senckenberg Museum DAAD Research
Fellowship (2000); Australian Heritage Commission
(2003–2006); Western Australian Fisheries (2006);
Australian Biological Resources Survey (2008–2012);
Chevron Australia (2009–ongoing).
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