The Azooxanthellate Scleractinia (Coelenterata: Anthozoa) of Australia
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© Copyright Australian Museum, 2004 Records of the Australian Museum (2004) Vol. 56: 259–329. ISSN 0067-1975 The Azooxanthellate Scleractinia (Coelenterata: Anthozoa) of Australia STEPHEN D. CAIRNS Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, PO Box 37012, Washington, DC 20013-7012, United States of America [email protected] ABSTRACT. A total of 237 species of azooxanthellate Scleractinia are reported for the Australian region, including seamounts off the eastern coast. Two new genera (Lissotrochus and Stolarskicyathus) and 15 new species are described: Crispatotrochus gregarius, Paracyathus darwinensis, Stephanocyathus imperialis, Trochocyathus wellsi, Conocyathus formosus, Dunocyathus wallaceae, Foveolocyathus parkeri, Idiotrochus alatus, Lissotrochus curvatus, Sphenotrochus cuneolus, Placotrochides cylindrica, P. minuta, Stolarskicyathus pocilliformis, Balanophyllia spongiosa, and Notophyllia hecki. Also, one new combination is proposed: Petrophyllia rediviva. Each species account includes an annotated synonymy for all Australian records as well as reference to extralimital accounts of significance, the type locality, and deposition of the type. Tabular keys are provided for the Australian species of Culicia and all species of Conocyathus and Placotrochides. A discussion of previous studies of Australian azooxanthellate corals is given in narrative and tabular form. This study was based on approximately 5500 previously unreported specimens collected from 500 localities, as well as a re-examination of most of the types and previously reported specimens from the Australian region. Fifty-six species are recorded as new to Australia; 183 state range extensions are listed; and 96 worldwide bathymetric range extensions are noted. In order to characterize the Australian fauna, all 703 known azooxanthellate species were tabulated as to coloniality, method of attachment, and depth range: 187 species are colonial, 516 solitary; 373 are attached, 265 free, and 54 transversely dividing; and 200– 1000 m is the most common depth range. Compared to all azooxanthellate species, those from Australia have a slightly higher percentage of species that are solitary and unattached (or transversely dividing), due to a disproportionate number of species in the families Flabellidae and Turbinoliidae. Bathymetrically they are typical of the worldwide fauna. Sixty-seven species are endemic to the Australian region. Both UPGMA cluster analysis and MDS ordination reveal two main regions: a northern tropical region and a southern warm temperate region, consistent with zonation patterns of shallow-water marine invertebrates. CAIRNS, STEPHEN D., 2004. The Azooxanthellate Scleractinia (Coelenterata: Anthozoa) of Australia. Records of the Australian Museum 56(3): 259–329. www.amonline.net.au/pdf/publications/1434_complete.pdf 260 Records of the Australian Museum (2004) Vol. 56 Azooxanthellate corals comprise half of the species and A particularly successful cruise was made by the R.V. genera of the order Scleractinia (Cairns, 1999b), and, with Franklin (06/88/1–22) in August, 1988, which collected the publication of this paper, consist of 703 species. Two many and varied deep-water Scleractinia that are now hundred thirty-seven species, or one-third, of the deposited at the Australian Museum. Unfortunately, specific azooxanthellate species occur off Australia, making it one station numbers were not recorded for all specimens of this of the richest regions in the world for this type of coral. The cruise. These undocumented stations are referenced as purpose of this paper was not to re-describe the azooxan- Franklin 06/88/x so as not to lose the unique records from thellate coral fauna of Australia, as most of the species have this cruise. All stations were made between 10–12°S and been adequately described and figured within the last 15 144–145°E at depths of 495–2523 m, a relatively years in papers about the Australian fauna or adjacent circumscribed region off the coast of northeastern Cape York regions (Cairns, 1989a, 1994, 1995, 1998, 1999a; Cairns Peninsula. & Parker, 1992; Cairns & Zibrowius, 1997). Instead, the The SEM was done by the author using an AMRAY 1810 primary purpose is to document and verify all previously scanning electron microscope. published records of the 237 Australian azooxanthellate Zoogeography. The zoogeographic affinities of the species in the form of annotated synonymies, to augment Australian azooxanthellate Scleractinia were analyzed based these reports with additional records from the various on their recorded presence in the seven states, but with Australian museums, and to list deposition of types and type Western Australia divided into a tropical and temperate zone localities for all species. This exhaustive compilation then at the Houtman Abrolhos Islands, and with three added provides the basis for a meaningful zoogeographic analysis categories of New Zealand, Indo-Pacific, and (east coast) of this fauna, the second goal of the paper. Seamounts to facilitate subsequent comparisons (Tables 2 Material and 3). The methodology used follows that of Cairns & Chapman (2002), i.e., first constructing a cluster analysis This study was based on an examination of approximately of the regions using the UPGMA method, followed by a 5500 newly-reported specimens collected from 498 more detailed ordination, i.e., Non-Metric Multi- localities, many of which are recorded in the Station List. Dimensional Scaling (MDS). The program PC-ORD4 Over the course of the last 15 years, but primarily on a visit (McCune and Mefford, 1999) was used for both analyses. to Australia in January–February, 2002, I examined the The 30 species restricted to a single locality were eliminated azooxanthellate collections of most of the major Australian from the analyses, resulting in a data matrix of 207×11, or natural history museums, including those of: SAM, NMV, 2277 cells. In order to characterize and compare the AM, Macleay, QMB, MTQ, AIMS, NTM, and WAM. groupings resulting from the MDS—as well as each region, Specimens were collected by over 34 vessels (see Station all of Australia, and all known species—Table 3 was List), but most significant among those were the Australian constructed listing the number and percentage of species vessels Kapala, Franklin, Soela, and Cidaris. Specimens having various morphological, bathymetric, and taxonomic from the first three vessels are housed at AM, and those from characteristics. Although bathymetric ranges have been the Cidaris at MTQ. As a result, of the 237 azooxanthellate published for the azooxanthellate coral genera (Vaughan & species known from Australia, original or additional records Wells, 1943), Cairns (2001b) reported the ratio of colonial/ of 186 species (78.5%) are reported herein. solitary azooxanthellate species, and Cairns (1999b) the The study also involved the re-examination of virtually taxonomic percentages of each family, Table 3 is the first all specimens of azooxanthellates previously reported from detailed compilation at the species level for depth of Australian waters, as well as the type specimens on which occurrence and mode of attachment for all azooxanthellate they were based. In the latter case, the types of 214 of the species, as well as updating the previously reported figures. 237 (90.3%) species have been examined; many of the This analysis was based on the list of species published by remainder are considered to be lost. Cairns et al. (1999) as updated through this publication. The following abbreviations are used in the text. Methods Systematic part. Synonymies are purported to be complete Museums regarding Australian records, and augmented with extralimital references that contain useful information on AIM Auckland Institute and Museum, Auckland. AIMS Australian Institute of Marine Science, synonymy, inclusion in a key, distribution, illustration, and/ Townsville. or description. For each Australian synonymy entry the state AM Australian Museum, Sydney. from which the specimen was collected is indicated in bold BM British Museum, London (now The Natural face, or there is an indication of whether it was simply History Museum). uncritically listed in that publication. The extralimital entries MCZ Museum of Comparative Zoology, Harvard are also briefly annotated but without bold face emphasis, University, Cambridge. as these do not include Australian records. Only previously MNHN Muséum national d’Histoire naturelle, Paris. unpublished new records are listed for each species, as well MTQ Museum of Tropical Queensland, Townsville. as the type deposition and type localities of senior synonyms NMW Naturhistorisches Museum, Wien. and Australian junior synonyms. Although Norfolk, NNM Nationaal Natuurhistorisch Museum, Leiden (formerly RMNH). Macquarie, Heard, Christmas, and Cocos-Keeling Islands, NMV National Museum of Victoria, Melbourne. and part of the Antarctic continent are considered as NTM Northern Territory Museum, Darwin. Australian possessions, this study does not include those NZOI New Zealand Oceanographic Institute, Wellington regions, but does include specimens from the seamounts of (now the National Institute of Water and Atmos- the Lord Howe Seamount Chain. pheric Research). Cairns: Azooxanthellate Scleractinia of Australia 261 POLIPI Politbang Oseanologi (National Institute of (=Stephanocyathus) platypus at a depth of 750 m off Sydney. Oceanology), Jakarta. He also reported two other species from Two Fold