Higher Taxon PISCES

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Higher Taxon PISCES

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31 December 2005 - Douglass F. Hoese, Dianne J. Bray, John R. Paxton & Gerald R. Allen

Introduction

Fishes are the largest and most diverse group of vertebrate animals, with an estimated 32,500 extant species (Nelson 2006), about equal to the number of all other vertebrates combined. In Australia, they are found in almost all aquatic habitats. About half of the fish species known at the time from Australian waters were treated in the work Zoological Catalogue of Australia Volume 7 Pisces (Paxton et al. 1989). That work has now been revised and incorporated into Volume 35 of the Zoological Catalogue of Australia series. In Paxton et al. (1989), the fish fauna of Australia was estimated to number 3,600 species, and it was predicted that by 2005 the number of known species would grow to 4,500. Currently the list of known species stands at 4,482 described species and approximately 300 species for which identity is as yet to be determined.

The classification presented here follows largely the work of Nelson (1994). Although there have been several changes subsequent to this classification, it does provide a useful reference point. If more recent studies are available, we have attempted to update the relevant sections. This work has changed many of the families recognised in various presentations and in the analyses in the first fish Catalogue (Paxton et al. 1989). In the interests of stability, the works of Nelson (1994) and Eschmeyer (1998) are followed here for family name spellings. Nelson's (1994) classification has been revised (see Nelson 2006). However, this later work of Nelson's was published too late to be incorporated into the current catalogue except to cite the reference where appropriate. The classification in Nelson (2006) is similar to that followed here, although the order of some families is different.

The orginal work of Paxton et al. (1989) included a total of 1,814 named species in 198 families. Changes to that orginal work include the addition of 400 species, changes to about 140 species scientific names and the recognition of about 15 species as junior synonyms.

The information presented in this Catalogue generally follows the format and style of the previously published volume. That work (Paxton et al. 1989), however, includes a more detailed discussion of general issues.

Geographical coverage in this volume is similar to that used in Paxton et al. (1989), coinciding with Australia's Exclusive Economic Zone (EEZ) (see Figure 1). Species from Elizabeth and Middleton Reefs off the east coast of Australia are included, based on the surveys undertaken by Gill & Reader (1992) and Oxley et al. (2004). Lord Howe Island, which is part of the Economic Zone, is included here using the works of Francis (1993), Francis & Randall (1993) and Speare et al. (2004), although it was not considered in Paxton et al. (1989). The Territory of Ashmore and Cartier Islands is included, although Paxton et al. (1989) included species from that Territory in Western Australia. Allen et al. (1989) recorded 747 species from the area, but they included reefs outside of the Territory. Allen (1993) recorded 569 species from reef, shore and epipelagic environments from Ashmore and Cartier Islands, whereas in this work 591 species have been recorded. Kospartov et al. (2006) estimated 651 species from Ashmore Reef and Cartier Island. Estimates vary, depending on whether offshore and deeper benthic fishes are included. Although we have updated most of the records concerning the Territory of Ashmore and Cartier Islands, some records may have been missed, thus causing some species still to be listed as from Western Australia.

Fishes from other Australian external territories are not included. These areas are covered by various checklists: Cocos Keeling Island - Allen & Smith-Vaniz (1994); Christmas Island - Allen & Steene (1979); Macquarie Island - Williams (1988); and Norfolk Island - Francis (1993) and Francis & Randall (1993).

We have attempted to include all species that have been recorded in the literature as reported from Australia. In addition, this Catalogue completely updates the taxonomic groups included in Paxton et al. (1989). However, the literature on fishes is so extensive that undoubtedly some works may have been missed. Also there is a considerable literature for which it is not possible to verify species identifications. Therefore, we have not treated some literature, particularly where there is simply a list of species. We have endeavoured to include references up to the end of 2005 and new species descriptions covering Australian species before the middle of 2006. Most museums around Australia kindly assisted by providing access to collections and to distribution records and these recordes are included, even if unpublished. However, records have not been included if we considered the identifications to be unreliable. We have respected intellectual property, and have not included species where specialists requested we not include them, in order for those workers to document occurrences of the species in Australia in the scientific literature. In a few cases new records are included based on material found in various museums in Australia and the bases of those records are included.

A major listing of Australian fishes was compiled by CSIRO Marine Laboratories in Hobart (Yearsley et al. 1997); the listed species were not annotated. However, Gordon Yearsley kindly provided information regarding the bases of the records. We have used that list to cross check the species presented in this Catalogue, and have noted some discrepancies which we were unable to resolve. In most cases, the discrepancies relate to species for which there is some uncertainty about their taxonomic status. In some cases, the CAAB list has recorded species as unidentified, but we have included records based only on published literature records. The CAAB list includes new undescribed species and species of uncertain identity. Because we only include described species, there are numerous species in the CAAB list that we do not include. Ultimately, some of those species may be documented as occurring in Australian waters.

Hutchins (2001) compiled a checklist of just over 3,000 species of Western Australian fishes (including Ashmore and Cartier Islands). This Catalogue records 2,745 species from Western Australia. Hutchins' list included over 250 unnamed species, and probably 200-300 Ashmore Reef species not recorded for the mainland. For some groups there are also some duplicate listings of species under junior synonyms or species simply listed as 'species a', 'species b', etc.

Introduced fishes are included in this Catalogue. Although most of these are freshwater introductions (McKay 1984, 1989; Arthington & McKenzie 1997), eight marine species also appear to have been introduced into Australian waters (Paxton & Hoese 1985; Clements et al. 2000; Lockett & Gomon 2001). The list of freshwater introductions in this work is undoubtedly incomplete. Releases of aquarium fishes have become very common, particularly in Queensland, and we have not tried to document all these introductions, many of which have failed to establish breeding populations. Consequently, introduced freshwater species are included only where breeding populations appear to have become established.

Translocation of freshwater fishes within Australia is also common and is discussed by Arthington & McKenzie (1997) and Russell et al. (2003). We have not attempted to document all such translocations here.

Historical Review

The first major publication on Australian fishes was White's 1790 Journal of a Voyage to New South Wales. Eleven species were described and figured from, Botany Bay, New South Wales. Since that time several partial and comprehensive checklists of Australian fishes have been published, including the works of Macleay (1881a, 1881b, 1881c, 1881d, 1884), McCulloch (1930), Whitley (1964a) and Paxton et al. (1989).

Macleay's (1881a-d, 1884) Descriptive Catalogue of the Fishes of Australia included a brief description and a statement of distribution for each Australian species. McCulloch's (1930) Checklist of Australian Fishes (completed and published by G.P. Whitley after McCulloch's death in 1925) provided full synonymies for both genera and species, and distribution by State for each species. Many of the synonymies listed by McCulloch are still recognised today. Whitley's (1964a) name-list of Australian fishes included only the author and date for each species. Ian Munro of the CSIRO produced a 'Handbook on Australian Fishes' between 1956 and 1961 (see Munro 1956–1961) as supplements published in the Australian Fisheries Newsletter; Munro's work covered less than one third of the species known at the time.

Comprehensive works covering Australian State ichthyofaunas include Castelnau (1872) for Victoria; Ogilby (1886) and McCulloch (1922) for New South Wales (later editions of McCulloch's work published in 1927 and 1934 are identical to the first edition, but are preceded with additions by Whitley (1934)); Ogilby (1916), McCulloch & Whitley (1925), Marshall (1964) and Grant (1965, 1975, 1982 1991) for Queensland; Waite (1921, 1923), Scott (1962) and Scott et al. (1974) for South Australia; Lord (1923, 1927), Lord & Scott (1924) and Last et al. (1983) for Tasmania; and, Whitley (1948) and Allen (1985) for Western Australia.

Major regional works produced for a broader audience include Allen & Swainston (1988) and Allen (1997) for tropical north-western Australia; Randall et al. (1990, 1997) for the Great Barrier Reef; Hutchins & Swainston (1986) and Gomon et al. (1994) for southern Australia; Kuiter (1993) for south-eastern Australia; Kuiter (1996) and Grant (2002) for numerous marine species of Australia; and Gloerfelt-Tarp & Kailola (1984) and Sainbury et al. (1985) for trawled fishes from north-western Australia. If there was any uncertainty as to the identity of any of the species in these publications they are not included in this Catalogue. Four publications by Carpenter & Niem (1999a, 1999b, 2001a, 2001b) record some species from Australia on distribution maps, but it is often not clear what is the basis of the distribution record. Consequently, species from those works have been added only where there is a clear indication that the respective authors for the various chapters have mentioned Australia in the text. In many cases there are discrepancies between the text and the maps.

Freshwater species are treated by several workers, including Whitley (1964c) Lake (1971, 1978), McDowall (1980, 1996), Cadwallader & Backhouse (1983), Merrick & Schmida (1984), Leggett & Merrick (1987), Allen (1989), Larson & Martin (1990), Allen et al. (2002){fish1newdh60} and Pusey et al. (2004).

References to other earlier works too numerous to list here, covering partial faunas or important commercial fishes, can be found in Whitley's (1964a) Survey of Australian Ichthyology. Also, Paxton & McGrouther (1997) reviewed the growth and development of the fish collections of Australia and the general historical aspects of Australian ichthyology.

Summary of Current Knowledge of Systematics of Australian Fishes

In the present classification 327 fish families are recognised in the Australian fauna. The 26 most speciose families account for 50% of the species. Seventy-seven families contain only a single species; some of the families are monotyic, but most are represented by only a single species in Australian waters. Two hundred and thirty families (70% of families) each contain fewer than 11 species, accounting in total for only 16.5% of the species. Only nine families have more than 90 species (Table 1). Interestingly, the estimates of the total number of species in these same families were typically much lower in 1989. The 1989 figures were based on total number of estimated species including those undescribed, whereas the figures given in the second column are based on described species only. Consequently, the Gobiidae figure appears only slightly higher, but inclusion of undescribed and uncertain species would add about 50 additional species. The Myctophidae changed little, but little work has been conducted on the group in Australia since that time. The large increase in species numbers for Labridae resulted from the inclusion of Scaridae and Odacidae as subfamilies. For other families, the earlier estimates are under by 3-20%, which is an indication of the general uncertainty of the state of knowledge of fishes in Australia. The greatest relative increase in species numbers occurs in Macrouridae and is largely due to intensive work undertaken on this family in recent years by several workers in New Zealand, Australia, the USA and Russia.

It was estimated that for the families included in the 1989 Catalogue, more than 10% of species were unidentified to species or known to be new and undescribed (Paxton et al. 1989). Although a large number of these species have now been identified or described as new, additional new (described and undescribed) and as yet unidentified species are being discovered, resulting in an increase of roughly 50 named species per year, a rate that has remained virtually unchanged since 1964 (Figure 2). Macleay (1881a-d, 1884) recorded 1,291 species, based in part on the prodigious cataloguing efforts of Cuvier & Valenciennes (1822-1849) and Günther's Catalogue of Fishes (1859-1870) published by the British Museum (Natural History). The increase in species to obtain McCulloch's (1930) total of 2,023 species was in part due to the deep water collecting undertaken aboard the HMCS Thetis (Waite 1899) and, in particular, the FIS Endeavour (McCulloch 1911, 1914, 1915, 1916, 1926; Norman 1926), as well as the increasing number of working Australian ichthyologists during that period. In the 34 intervening years to Whitley's (1964a) total of 2,447 species, many of the new fish species records were due to sampling of the immense Great Barrier Reef fauna (Whitley 1932a, 1964b; Woodland & Slacksmith 1963).

In more recent years, extensive work by Frank Talbot and students from One Tree Island and Lizard Island has increased significantly the amount of material that has been available for numerous taxonomic studies. Similarly, fish taxonomists in various State and Commonwealth institutions have increased significantly the amount of material now available in Australian museum collections (Paxton & McGrouther 1997). Because of the rapid rate of increase in the known fauna, we have not attempted to estimate the number of unknown species for many of the families covered in this Catalogue.

The exceptional increase in Australian fish species numbers, an 83% increase since 1964, is primarily the result of a much larger number of ichthyologists currently working in Australia, more effective collecting methods and collecting from previously unsampled habitats, and the increased study of Australian fishes by overseas scientists.

In 1964, there was one fish taxonomist associated with each of three Australian museums (Australian Museum, Queen Victoria Museum and Western Australian Museum) and CSIRO Fisheries. In 1989, each of the eight Australian museums (those listed above plus Museum Victoria, Museum and Art Gallery of Northern Territory, Queensland Museum, South Australian Museum and Tasmanian Museum) and CSIRO Fisheries had at least one ichthyologist on staff or as an associate, two had two and one had three ichthyologists. By 2006, however, the number of research scientists employed as ichthyologists in those institutions had declined to five. The increase in scientists during the 1970s and 1980s resulted in a significant increase in collections. The fish collection at the Australian Museum numbered about 70,000 specimens in 1968. In 1997, the collection had grown to about 450,000 juvenile and adult fishes and perhaps an equal number of larval specimens (Paxton & McGrouther 1997), and at the end of 2004 that number was some 635,000 juvenile and adult specimens (McGrouther 2006). Increases have occurred also in most other collections, so that the total collection of Australian fishes now held in nine different Australian institutions is estimated at more than one million juvenile and adult fishes.

Extensive fish collecting with scuba and rotenone began in Australia in 1965. The previously unsampled habitats of coral and rocky reefs between five and 45 metres yielded numerous new species and records. The sustained sampling of coral reef fishes from the Great Barrier Reef, first at One Tree Island and later at Lizard Island, resulted in the present day distributional limits of many Great Barrier Reef species. These two localities are the only two comprehensively collected areas of the Great Barrier Reef and even today the larger species from each have not been sampled adequately. The northern limits for many of the Great Barrier Reef species have been extended as a result of two expeditions involving the collaborative efforts of the Australian Institute of Marine Science, Australian Museum, Museum and Art Gallery of the Northern Territory, Queensland Museum and the Field Museum of Natural History. The expeditions explored the waters between Lizard Island and the Coral Sea off the northern tip of Australia. Few collections have been made between Lizard Island and One Tree Island, and throughout much of the Coral Sea. Collections along coastal areas have been made by most State museums, but there are still extensive gaps in our knowledge of coastal fishes. During the early 1970s, exploratory fishing in deep water was begun by New South Wales Fisheries using the research vessel FRV Kapala, and later by the CSIRO, using the vessels FRV Courageous, FRV Franklin, FRV Soela and FRV Southern Surveyor. More recently, fishing in depths to 1,500 metres has yielded numerous new records of both benthic and midwater species; specimens are housed within the collections of State museums and CSIRO. Although collections have been made of freshwater fishes, few comprehensive collections exist for many of the rivers in Australia. In recent years, large larval fish collections have been developed, and considerable effort has been made to preserve material in alcohol for future genetic studies.

Finally, in the last 30 years many Australian specimens have been included in revisionary studies by overseas ichthyologists, primarily as a result of loans from Australian collections. The number of taxonomic problems associated with Australian fishes far exceeds the capacity of ichthyologists working presently within Australia and continuing progress will be made through the efforts of the worldwide ichthyological community.

ZOOGEOGRAPHY

The following analysis is based on data presented in this Catalogue. It is largely a description of characteristics of the fauna, rather than an attempt to apply modern biogeographic techniques, which are far beyond the scope of this work. Data are incomplete for some species because, largely, the distribution patterns have not been updated for about half the species in Paxton et al. (1989). Consequently the following analysis is a snapshot based on data that are approximately 5-15 years old. Discussions of the various categories analysed are included in the following section, Notes on the Database.

Distribution patterns of Australian fishes are complex. Little work has been done to identify breeding areas of marine species. Juveniles of tropical species often occur in temperate areas and in some cases isolated populations can occur in southern waters (Hutchins 1993, 1994).

Major zoogeographical reviews of the Pacific region include Ekman (1953), Briggs (1974), Springer (1982) and Randall (1998). Santini & Winterbottom (2002) proposed that the south-west Pacific, Australia and the western Indian Ocean may have been the centres of origins of tropical coral reef fishes of the Indo-Pacific, with the Indo-west Pacific being a highly derived area.

Zoogeographical reviews dealing largely with Australia include Whitley (1932b) and Wilson & Allen (1987). Zoogeographical reviews dealing largely with Australia include Whitley (1932) and Wilson & Allen (1987). Whitley (1932b) recognised six provinces, three tropical northern (Damperian west from the tip of Cape York to near Geraldton, WA; Solanderian east from the tip of Cape York for the Great Barrier Reef; and Banksian for the Queensland coast in the same area south to Fraser Island), one subtropical south-eastern (Peronian from Fraser Island to the middle of Bass Strait), and two southern temperate (Flindersian from Geraldton to the middle of Bass Strait and Maugean for Tasmania). Wilson & Allen (1987) reviewed the major components and distribution of the Australian marine fauna. Their study, based on fishes, molluscs, echinoderms and corals, included geological history and the effects of ocean currents. They recognised two broad regions, a northern tropical region and a southern temperate region, with eastern and western overlap zones. They also recognised a series of 13 overlapping areas that represent the common distribution patterns of shallow water benthic animals. Subsequently, collaboration between the Commonwealth and State governments led to the development of the Interim Bioregionalisation of Australia (Commonwealth of Australia 2005). That work recognised several major regions, with the States further subdividing each of those regions. Development of the regions generally did not include analysis of animal distribution patterns. The resultant patterns have been of use for management puposes in establishing marine protected areas, but the biological significance of the regions is unclear. The biogeographical regions recognised here are shown in Figure 1.

The biogeography of Australian freshwater fishes is reviewed by Whitley (1947), McDowall (1981) and Unmack (2001).

Extralimital Distributions

Broadly, 70% of the Australian species also occur in other areas of the west Pacific, 26% in the central Pacific and 40% in the Indian Ocean (Table 2).

Extralimital distributions arranged by mutually exclusive categories are shown in Table 3. Although most species not endemic to Australia are widely distributed, particularly in the Indian Ocean and western Pacific, a number of distinctive patterns of more restricted distributions are apparent. For example 273 species are found elsewhere only in the south-west Pacific (New Zealand, Lord Howe Island, Norfolk Island, Kermadec Islands; with some species extending to the Solomon Islands and New Guinea). Similarly 144 species are found elsewhere only in New Guinea. Of the 369 circumglobal species, 112 are known only from the Southern Hemisphere. Thirty-eight species are found elsewhere only in the North-west Pacific (Japan, China, Korea), and 104 species have disjunct distributions (i.e. not found in contiguous areas). The species with disjunct distributions are often oceanic or benthic-deepsea species and their distribution patterns probably reflect inadequate sampling. Ten species, known primarily from reefs on the outer edge of the Great Barrier Reef and the Coral Sea, are found also in the central Pacific; some were originally considered by Springer (1982) to be restricted to the central Pacific.

Australian Distributions

The biogeographical regions within Australia as used here are modified from those used in Paxton et al. (1989). For the analysis of distribution within Australia, distributions were modified as part of the updating of that work, and are similar to regions identified in the Interim Bioregionalisation of Australia (Commonwealth of Australia 2005).

The Australian fish fauna is predominantly a marine tropical fauna (Table 4). In total 68% (3,029 species) of the total native species occur in tropical marine environments, 27% (1,222 species) in temperate environments and 9% (390 species) occur in either subtropical or both temperate and tropical environments.

Most species are known from inshore marine environments (63%), with 25% from offshore environments (beyond continental shelf) (Table 5). In Table 5, continental shelf species are included in inshore, but they are also listed separately, because it was not possible to distinguish between 'inner shelf' and 'outer shelf' species. A small number of species occur in both offshore and inshore waters and most of these are known from the outer Continental Shelf or are pelagic migratory species.

Only 5.8% of native Australian fishes are found in fresh water. Of the total of 283 species found in fresh water, 25 are introduced, 166 are restricted to fresh water, 91 are also found in estuarine environments and 31 in marine environments, characteristic of the predominantly marine origin of the Australian freshwater fish fauna (Table 6). Almost 68% of the freshwater species occur in the tropics, 32% in temperate environments and 2.7% in subtropical environments. Of the species found in estuarine environments 61% are also found in marine environments, 24% also occur in freshwater and 20% are largely restricted to estuarine environments. A small number of species (28) occur in freshwater, estuarine and marine environments.

Coral and rocky reef species dominate the fauna with 46% of Australian species found in these environments, followed by soft bottom marine species at 34%, pelagic fishes found outside of estuaries equal 15% and estuarine species make-up 10%. Some species occur in two of these environments (see Tables 7).

Much of the tropical marine fauna occurs on coral reefs, which are well developed in northern Australian seas. A multi-year project funded by Conservation International involving the preparation of distribution maps for every shallow (< than 60 m depth) coral reef fish in the Indo-west and central Pacific region reveals that the Australian fauna comprises 1,722 coral reef fishes, a total that is exceeded only by Indonesia (2,097 species). The largest concentration of coral reef species, consisting of 1,468 species, is found on the Great Barrier Reef (G. Allen, unpublished data). Note that these figures differ from those outlined in the Catalogue in Tables 7 and 10. The higher total number of coral reef species (1,722 versus 1,610 in Table 7) obtained in the Conservation International study is probably due to slightly different methods of allocating species to the coral reef category. The lower number obtained for the Great Barrier Reef (1,468 versus 1,625 in Table 10) is largely due to the inclusion of trawl species in the Great Barrier Reef region in the Catalogue.

Endemism within Australia

Overall, 24% of the Australian fish fauna is endemic. Most environments have a high level of endemism (Table 7). Freshwater environments have the highest level of endemism (61%), followed closely by rocky reef environments (55%), then soft bottom marine environments (33%), followed by estuarine environments (29%). The lowest level of endemism occurs on coral reefs (8%) and in pelagic marine environments (7%). Similarly endemism is highest in temperate and subtropical environments (47 and 46% respectively) and lowest in tropical environments (14%) and in species found in both tropical and temperate environments (17%) (see Table 8).

Endemism of fish species by political jurisdiction is provided in Table 9. Western Australia has the highest proportion of Australian endemic species, with 57% of the Australian endemic species found in the State. The Northern Territory has the lowest percentage of Australian endemics (15%), whereas other states are similar in having 25-37% of the Australian endemics. The high figure for Western Australia is undoubtedly due in part to the large area. Western Australia also has the highest percentage of species endemic to a single state (7.1%), followed by Tasmania (6.0%), South Australia (3.7%), Queensland (2.6%), Northern Territory (2.1%), New South Wales (1.3%) and Victoria (0.2%). However, the percentage of species occurring within a State that are endemic to Australia shows a different pattern. South Australia has the highest percentage of species of any State that are Australian endemics (58.5%), followed by Victoria (48.9%), Tasmania (42.8%), Western Australia (22.3%), New South Wales (21.7%), Queensland (14.8%) and finally the Northern Territory (14.7%). The areas covered for each State in this analysis are larger than the jurisdictional areas for management and the State responsibility for the species would involve a smaller percentage. However, because most of the endemics are inshore and freshwater fishes, the figures are approximations of State responsibilities.

Endemism by geographical categories is given in Table 10 for inshore species and Table 11 for oceanic (offshore) species.

Endemism by geographical zone for inshore fishes is generally low for any particular region. The highest number of endemics confined to one zone is 27 for the N coast. The highest percentage of endemism within a region is that recorded for the Coral Sea (3.7%), but the total number of species known from the area is low due to limited sampling and thus the endemism figure is likely to be artifically high. The next highest percentage of endemism within a region is 2.1% for the Lower W coast. The figures for most zones are between 1 and 2%, with the lowest mainland area (0.3%) for the Great Australian Bight and the Gulf of Carpentaria, suggesting that these two regions are not well defined based on endemism. The Territory of Ashmore and Cartier Islands has no species restricted to the Territory. Combining regions increases the percentage of endemism, but generally three or more regions need to be combined to achieve a figure of 10% or more. Combining the Central E coast and Lower E coast actually decreases the percentage of endemism to 1.9%, primarily due to the large number of tropical species that are commonly found within the Central E coast. The percentage of endemism in shore fishes ranging from Victoria and Tasmania to and including the SW coast reaches 9.8%. Adding the Central E coast and Central W coast increases the figure to 17%.

Almost 15% of species found in offshore waters are currently regarded as endemic. Similarly, 15% of Australian endemic species occur in offshore waters. About half of these species also occur in inshore waters. The high number (71) of apparently endemic oceanic species is probably related to intensive collecting activity in the Australian region; it is likely that many of these species are more widespread. For example many of the endemics are known from single samples, such as in the family Liparidae, where 29 endemic species are known from only one location.

Endemism is high in freshwater fishes, with 56% of the total species occurring in freshwater endemic to Australia. Figures are given in Table 12 for the various drainage divisions. Endemism is highest for native freshwater species restricted to freshwater. Endemism varies from 45% for the Lake Eyre Basin to 6% for SE coastal. Three drainage divisions (Bulloo River basin, W plateau and S Gulfs) have no regional endemics (Figure 3). All drainage divisions have a high proportion of Australian endemics, with Lake Eyre and Bulloo River basin having the highest percentage of Australian endemics (83.9 and 85.7% respectively). The other drainage divisions have between 31.5 and 71.4% of their species endemic to Australia.

Of the 166 species restricted to freshwater 79% are endemic to Australia. All 35 species found outside of Australia occur in New Guinea, but only seven are widely distributed in the west Pacific (mostly gobiids). Of the total species found in freshwater, 68 also occur in estuarine or marine environments. Twenty-three of these are commonly found in both freshwater and estuarine environments. Eight species occur primarily in fresh water, but are migratory, breeding in the sea (eels), or with marine larval stages which migrate into fresh water (galaxiids). The remaining species are primarily marine, but sometimes occur in lower reaches of rivers, in fresh water.

The number of endemics occurring in any broad habitat varies considerably, generally being lowest in tropical regions, particularly marine environments (Figure 4). Breaking down endemism by broad areas (tropical and temperate) and broad habitats (freshwater, marine and estuarine) indicates that endemism varies considerably by habitat and area (Table 13). Similarly, a breakdown of inshore versus offshore environments indicates that endemism is low in offshore environments (Table 14, Figure 5). The higher endemism among temperate offshore fishes probably is an artifact, reflecting the greater collecting effort in temperate envionments of Australia. The more inshore fauna shows the highest percentage of endemism in temperate areas (73% for temperate reef species and 66% for temperate estuarine species). However, among tropical species, estuarine and Continental Shelf representatives have a similar percentage of endemism (16% and 17% respectively), whereas reef species have a low percentage of endemism (8%). The figures for tropical species are also likely to be artefacts of collecting effort. Reef environments have been far better sampled and studied in the Indo-Pacific region than estuarine and continental shelf areas.

Certain groups of fishes tend to have high levels of endemism in Australia. Table 15 lists the larger families (with more than 20 species), which have more than 20% of its taxa endemic to Ausralia. The families Syngnathidae and Labridae have the greatest number of endemics. Freshwater families, such as Eleotridae and Galaxiidae also have a high level of endemism. The high level for Liparidae is probably an artifact of collecting effort as mentioned above.

Introduced Fishes

Currently only eight marine species are thought to have been introduced into Australia (Table 16). Three of the species have been taken on only one or two occasions and are not thought to have become established. The established species include temperate species (three gobiids and two tripterygiids). Because most tropical species are widely distributed, it would be difficult to identify an introduced tropical marine species that is native to the western Pacific.

Table 17 lists the species of freshwater species that are known to have established populations, at some point in time. The list is based on some information that is several years old and is probably incomplete; some species may no longer have established populations. Some species were deliberate introductions for pest control or angling.

Introduced freshwater fishes (Table 12) in some areas account for a high percentage of species known from the drainage. The Murray-Darling drainage has the highest percentage of introduced species (33.3%), followed by SW coastal (32%), S Gulfs (27.8%), SE coastal (20%), NW coastal (16.7%), NE coastal (11.7%), Lake Eyre Basin (6.5%) and N coastal (1%). Two drainages (N Gulf and Bulloo River basin) have no introduced species.

Acknowledgements

A large number of people helped with the preparation of this Catalogue. We thank all the various workers in Australian museums and Commonwealth organisations: the late John Glover and Terry Sim (South Australian Museum); Martin Gomon and Tania Bardsley (Museum Victoria); Barry Hutchins and Sue Morrison (Western Australian Museum); Jeff Leis, Mark McGrouther, Kerryn Parkinson, Sally Reader and Tom Trnski (Australian Museum); Rolly McKay and Jeff Johnson (Queensland Museum); Peter Last and Alastair Graham (CSIRO); and Barry Russell and Helen Larson (Museum & Art Galleries of the Northern Territory) for providing information on their collections and allowing access to specimens and records under their care.

Gordon Yearsley kindly provided information from the CAAB files and provided valuable feedback on species information and literature. We are grateful to Libby Moodie and Judie Recher who helped to compile information for this Catalogue.

Unpublished information and reviews of various chapters were kindly provided by Emperor Akihito, Kunio Amaoka, Eric Anderson, Masahito Arai, Lynnath Beckley, Kent Carpenter, John Caruso, Peter Castle, Francois Chapleau, Dan Cohen, Bruce Collette, Leonard Compagno, Ross Feltes, Carl Ferraris, Nicola Fox, Malcolm Francis, Tom Fraser, Ron Fricke, Ron Fritzsche, Javad Ghasemzadeh, Bob Gibbs, Tony Gill, Martin Gomon, Ofer Gon, Graham Hardy, Barry Hutchins, Yuji Ikeda, Walter Ivantsoff, Tomio Iwamoto, Akihisa Iwata, Dave Johnson, Jeff Johnson, Patricia Kailola, Les Knapp, Rudie Kuiter, Helen Larson, Peter Last, Matthew Lockett, Doug Markle, Keiichi Matsuura, John McCosker, Bob McDowall, Mark McGrouther, Katsusuke Meguro, Peter Møller, Randy Mooi, Hiroyuki Motomura, Tom Munroe, Ed Murdy, Joe Nelson, Jørgen Nielsen, John Olney, Nikolai Parin, Chris Paulin, Frank Pezold, Ted Pietsch, John Pogonoski, Stuart Poss, Jack Randall, Bill Richards, Clive Roberts, Tyson Roberts, Barry Russell, Kunio Sasaki, Yuri Sazonov, Koichi Shibukawa, Bill Smith-Vaniz, Vic Springer, John Stevens, Andrew Stewart, Bruce Thompson, Jim Thomson, Tom Trnski, Peter Unmack, Peter Whitehead, Alan Williams, Jeff Williams, Rick Winterbottom and Thosaporn Wongratana. Jeff Leis provided advice and kept us informed about several recent references. Bill Eschmeyer provided valuable information about types and literature.

Particular recognition must go to Ken Graham of New South Wales State Fisheries. Over many years Ken has contributed to the Australian Museum significant collections of deepwater fishes, collected from the FRV Kapala off New South Wales, as well as providing valuable assistance in the identification of these remarkable fishes. Valuable specimens have also been provided by other Fisheries staff from New South Wales and Western Australia. The efforts of Peter Last, Gordon Yearsley, Alastair Graham and other staff of CSIRO Marine Laboratories in Hobart have increased significantly our knowledge of fishes in Australian waters. Ship time is a precious commodity in Australia, but we have been fortunate to have had boat time on a number of vessels including the R/V El Torito previously owned by Walter Starck, the R/V Lady Basten of the Australian Institute of Marine Sciences, the FRV Kapala formerly of New South Wales Fisheries and the R/V Sunbird formerly of Lizard Island Research Station. We and other Australian workers have participated on various legs or cruises of the FRV Franklin, FRV Southern Surveyor, FRV Courageous and FRV Soela of CSIRO; Kaiyo Maru of Japan; Dmitry Mendeleev of USSR; Hai Kung of Taiwan; the FRV Tangaroa of New Zealand; and, HMAS Kimbla, HMAS Cook and HMAS Sprightly of the Royal Australian Navy. Our fellow curators in other Australian institutions have been most supportive. Finally, we wish to acknowledge Rudie Kuiter of Melbourne, Victoria and Roger Steene of Cairns, Queensland, for their major contributions, through photography, collection and fieldwork, to the understanding of the fish fauna of Australia.

Keith Houston of the Australian Biological Resources Study provided valuable assistance in early stages of preparation of this Catalogue in the database Platypus. We also wish to thank the former directors of ABRS, Ian Cresswell and Mary Colreavy, for supporting the completion of the Cataogue; the Fisheries Research and Development Corporation for funding this project; and Alan Snow of Seafood Services Australia Ltd and the Fish Names Committee for aiding in the coordination of the common names.

We particularly thank Pam Beesley and Alice Wells for their patience and perseverance in bringing the Catalogue to completion.

Database Notes

Synonymies listed here are not comprehensive. Generic synonymies are not given, except for replacement names or where confusion exists over names of similar spellings. For species junior, synonyms are only included if the type locality is from Australia, if the name has been in widespread use in Australia or if the type species of a genus known from Australia is a junior synonym of a species known from Australia.

In the preparation of this work, original literature was consulted to determine species and genus available name data. The publication Catalog of Fishes by Eschmeyer (1998) was invaluable for cross checking data, and in many cases was the only published information regarding type registration numbers.

Since publication of the Zoological Catalogue of Australia Volume 7 Pisces in 1989 a number of type catalogues have been published: Hutchins & Smith (1991) listed Western Australian Museum types; Nijssen et al. (1993) revised the type catalogue for the University of Amsterdam; and Fricke (1991, 1992) provided information on types from the Staatliches Museum für Naturkunde, Stuttgart, Germany. The Australian Museum type list is available from the Australian Museum Website, and the Day types from the Australian Museum are treated by Ferraris et al. (2000).

Publication dates for J. Bennett are treated in Pethiyagoda et al. (1994). Publication dates for W.C.H. Peters are treated by Bauer et al. (1995). Dates for the Bleeker papers are covered in Pieters & Dickinson (2005), but it appeared too late to update all publication dates of Bleeker here.

CAAB Numbers

Codes for Australian Aquatic Biota (CAAB) is a continuously maintained and expanding 8-digit coding system for aquatic organisms in the Australian region. It is maintained by CSIRO Marine and Atmospheric Research (Yearsley et al. 1997). Initially developed to cover fishes and selected other organisms of research or commercial interest, it has more recently been expanded to provide more comprehensive coverage of a number of aquatic groups, as information is available. CAAB is used by a large and expanding number of government and industry groups for marine biology and fishery purposes and has replaced almost all other national and locally designed coding systems. The CAAB list includes some commercial species imported into Australia as well as Lord Howe Island species that do not occur on the mainland. The first two digits indicate the main group (37 for fishes), the next three digits are a code for the family and the last three digits are the species number. The numbering system for families was originally established by the Australian Museum in the late 1960s, covering 471 families. Subsequent taxonomic work has resulted in some families being synonymised with other families, resulting in the disappearance of some numbers. In other cases new families have been recognised. Because the family numbering system was originally consecutive in taxonomic order, new numbers are no longer added for families. As a result when a new family is recognised, the new family will retain the number of the family in which the species were previously placed.

Common Names

Common names have been entered from major books on Australian fishes. Over 10,000 common names have been used for Australian fishes. Names are included here for almost all species. Names from works published overseas are generally not included, except for standard FAO names or where no name could be found in the Australian literature. Myers (1999) work was used for names of small coral reef fishes, for which no name had been found in the Australian literature. Previously there were no standards for common names, and as a result, variations on a single name can lead to up to six different spellings, such as Longspine Fish, Long-spine Fish, Long-spined Fish, Longspined Fish, Long Spine Fish or Long Spined Fish. Authors of major books have often been trained overseas and there has been an increasing tendency for overseas names to appear in these works. For example, Randall et al. (1990) use the name rabbitfish for species commonly referred to as spinefoot in Australia, which is also the accepted FAO name for many of the species. Similarly the name snapper is commonly used for Lutjanidae in the USA and is increasingly being used for some species in Australia. It is also common for authors of books to make up new names for species that already have an Australian common name. Different names are often used for the same species in different parts of Australia. In some cases the same name has been given to two or more species. As a result the number of common names included here far exceeds the number of known species and available names.

Over the last two years a Standards Name List for Australian fish has been developed by scientists from museums throughout Australia and CSIRO Marine and Atmospheric Research in conjunction with the Fish Names Committee administered by Seafood Services Australia. The List is available on the internet at http://www.marine.csiro.au/caab and as a publication (see, Yearsley et al. 2006). The Standards Name List is currently under review and the names of some species are likely to change.

The common names included here are in alphabetical order with the agreed standard name placed first and in bold. For a few species, a standard name has not been determined and where there are common names for these they are listed in alphabetical order. For a few species there are no standard or common names, and for other there are no common names, except for the agreed standard name. The standard names provided here are consistent with the most recent Standards Name List (see Seafood Services Australia 2006).

Museum Acronyms
The museum acronyms used in this Catalogue are listed in Appendix II. The list has been standardised by the Australian Biologial Resources Study across all taxonomic groups. In a few cases, the abbreviations do not match the standards used in ichthyology and herpetology developed by Leviton et al. (1985).

Distribution Categories

Under this heading are listed the drainage divisions, coastal zones, oceanic zones and States shown on the map of Australia (Fig. 1). These zones differ in some cases from those used in the first fish volume, particularly in New South Wales and Western Australia. The arrangement is also not consistent with the previous version, where regions were listed in a clockwise manner. There are some inconsistencies, particularly in Queensland where the boundaries between the Great Barrier Reef and the NE coast zones are difficult to distinguish. Distributional limits including latitudes (for the east and west coasts) and/or longitudes (for the north and south coasts) are presented following the general categories, for most species. These limits are taken mostly from the literature and records in various museum collections in Australia. We have relied extensively on collections in the Australian Museum and Western Australian Museum, but material and records were also examined in the Museum and Art Gallery of the Northern Territory, Queensland Museum, Museum Victoria and the CSIRO Marine Laboratories. It should be noted that there was not sufficient time to update all the distribution patterns of species recorded by Paxton et al. (1989).

The distributional extremes of a species may give a biased picture of the true centre of distribution, particularly where young of coral reef species are found in southern New South Wales due to the action of the East Australia Current in the summer. More detailed distributional analyses based on home ranges or breeding ranges, while needed, are beyond the scope of this work. There is also a boundary effect; where a species only just crosses into the next zone, it is recorded in that zone in this work.

Each species was designated as tropical, temperate or subtropical based on its Australian distribution. Some species occur in both tropical and temperate climatic zones, but those primarily tropical species with seasonal vagrants in more temperate latitudes were only listed as tropical. Subtropical species are defined as those occurring on the east or west coasts, but not widespread in southern or northern waters. Because of the problem with currents influencing distributions of the juveniles, assigning a poorly known species to one of the categories was subjective.

Extralimital Distribution Categories

Each species was placed initially in one of 28 mutually exclusive zoogeographic categories, which have been reduced here to 22 (see Table 3). The extralimital zoogeographic categories are based only on the non-Australian distribution; thus a species occurring in Western Australia, Queensland, the Philippines and Japan is considered west Pacific and not east Indo-west Pacific. Definitions of the categories follow Paxton et al. (1989) and are illustrated in Figure 6. Most of the categories are logical and self-explanatory, being single areas or combinations of the major zoogeographic areas. Allocation of a species to the various categories often was subjective, taking into account the overall distribution of the species, particularly where there was some overlap in the distribution areas (such as south Pacific and south-west Pacific). For example, most species from the south-west Pacific ranged from Lord Howe Island eastward to New Caledonia or Fiji, but a few species in this category reached as far north as New Guinea. However, species included in the New Guinea category are those only found in New Guinea, outside of Australia. Similarly species in the category north-west Pacific were only found in that region outside of Australia, but wide ranging species from the west Pacific also might occur in that region.

Ecological and Biological Descriptors

Most of these are habitat descriptors and are presented for each species in order of increasing depth or distance from shore. We have attempted to improve the consistency and broaden the categories, but the information presented here is still not comprehensive.

Biological References

We have attempted to include a broad range of taxonomic and biological references, including life history information, for each species. Although the literature references are not complete, they do provide a useful source for locating additional references. For each species the relevant page number is given for the three most important previous Australian checklists or catalogues on Australian fishes — McCulloch 1930; Munro 1956–1961; Whitley 1964a. For the latter two works the species number is given. The name if different from present usage, is also given for all works. For example, in the reference Whitley, G.P. (1964). [53] (1818) — square brackets enclose the page number and round brackets enclose the species number and a comment if the species name was different from the senior synonym. If the species has been recorded from Australia since Whitley (1964), that reference is given. Those three previous checklists are abbreviated in the references sections as author and date only. Rarely, a reference to a distributional or taxonomic problem with the species is also presented. References for higher classification are included in order, suborder and family sections. For many species the literature records may be found in the references for the genus. It was not possible to list all literature under individual species because of the format used here.

Australian Biological Resources Study

Australian Faunal Directory