Australian Barnacles (Cirripedia: Thoracica), Distributions and Biogeographical Affinities

D. S. Jones

Seventy-four littoral and shallow-water (0–100 m) barnacle species from south-eastern Queensland are documented. Data from new collections of littoral and shallow-water barnacle species made during the February 2005 workshop, are combined with older SE Queensland barnacle collection data from Australian museums. South-eastern Queensland represents a transitional area between southern temperate and tropical zones and this transitional position is reflected in the composition of the barnacle fauna. The number of Australian endemic species (12) is relatively high, reflecting the influence of the southern fauna, but the fauna is dominated by species with Indo-west Pacific (25), cosmopolitan (22) and, to a lesser extent, Indo-Japanese (9) affinities, with Australasian (2), western Pacific (2) and Indo-Australasian (1) species minor components. One new species, Arcalepas brucei Jones & Morton, 2009, was collected during the Workshop. Neonrosella subgen. nov. is proposed to replace Rosella...

The subfamily Acastinae contains a diverse group of barnacles that are obligate symbionts of sponges and alcyonacean and antipatharian corals. Integrating morphological and genetic (COI) data to compare against known species, this paper reports on nine species of sponge-inhabiting barnacles of the subfamily Acastinae, including three undescribed species (Acasta caveata sp. nov., Euacasta acutaflava sp. nov., and E. excoriatrix sp. nov.) and three species previously not recorded in Australian waters (A. sandwichi, Pectinoacasta cancellorum, and P. sculpturata). The new species are distinguished from similar species by a suite of morphological characters as well as genetic distances. A lectotype for Pectinoacasta cancellorum is designated. Sponge hosts were identified for all specimens where possible and are represented by 19 species from eight families and five orders.

Aim To relate patterns of distribution of marine echinoderms and decapods around southern Australia to major ecological and historical factors.Location Shallow-water (0–100 m) marine waters off southern Australia, south of 30° S.Methods (1) Record the presence/absence of known echinoderm and decapod species in cells of c. 1° latitude and longitude, along the coast of southern mainland Australia and Tasmania. (2) Describe patterns in species composition, species richness and endemism through gradient analysis, ordination and cluster analysis. (3) Relate these patterns to distance and temperature gradients, the area of continental shelf, the average size of species range, and known historical factors.Results Species composition varied with both latitude and longitude. Species richness was relatively constant from east to west but graded with latitude from high in the warm-temperate regions around Perth and Sydney to low in cool-temperate southern Tasmania. Species richness was not related to the area of continental shelf or average species range size. Species turnover was not correlated with rates of temperature change. It was problematic to separate distance from temperature gradients, but there was evidence that the southern distribution limits of some species are related to minimum sea surface temperature. Within the taxonomic groups surveyed, evolutionary radiation has been largely limited to a few cosmopolitan species-rich genera.Main conclusions There are historical as well as ecological hypotheses explaining the latitudinal gradient of marine species richness in southern Australia: (1) the continual invasion and speciation of species of tropical origin as Australia has split from Gondwana and drifted northward; (2) progressive extinction of some Gondwanan cool-temperate species at the limits of their range; (3) low level of immigration of additional cool-temperate species; and (4) some in situ endemic speciation.

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. For permissions please email: journals.permissions@oup.com. 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. 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