Biology of the Antarctic Seas XI Antarctic Research Series, Volume 34, Paper 1, Pages 1-74 ANTARCTIC AND SUBANTARCTIC SCLERACTINIA Stephen D. Cairns Department of Invertebrate Zoology, Smithsonian Institution Washington, D. C. 20560 Abstract. The 37 species of Scleractinia known from the Antarctic and Subantarctic regions are described, mapped, and illustrated, including the description of 6 new species. Two more species, one of them new, from waters closely adjacent to the Subantarctic regions are also considered, as well as 4 previously published records of Sclerac- tinia of uncertain identity. A chronological, annotated list of all papers dealing with Antarctic Scleractinia is provided. A zoogeographic analysis describes common patterns of species distributions, and the faunas of various geographic areas are discussed. Scleractinia from South Pacific sea- mounts, some of which may form deepwater coral banks, are particularly noted. Introduction This paper reviews the 37 species of Scleractinia known from the Antarctic and Subantarctic regions, as well as 4 records of uncertain identity from these regions, and 2 more species from areas closely adjacent to the Subantarctic region. De- spite 28 papers dealing exclusively or partially with Antarctic Scleractinia (Table 1), only about 1200 specimens have been reported previously from 114 localities. The material upon which this study is based (primarily United States Antarctic Re- search Program vessels. Deep Freeze vessels, and the R/V Vema) includes over 7 times the number of specimens and almost 3 times the number of locali- ties reported in all the previous papers. Squires [1969] published a preliminary synthesis of the distribution of Antarctic Scleractinia but never did write his intended Antarctic monograph, thereby leaving many questions unanswered. Al- though his paper is valuable as a preliminary note, it includes numerous undocumented range extensions (all of his mapped records are undocumented), un- explained statements about complex groups of species, omissions of previous records, and refer- ence to an undescribed new species. Most, if not all, of Squires's [1969] specimens are deposited at the United States National Museum, Washington, D. C, and at the American Museum of Natural His- tory, New York. This material, along with subse- quently collected United States Antarctic Research Program specimens and an examination of most of the previously reported specimens, has allowed a documentation, and sometimes a correction, of Squires's synthesis, a reevaluation of the system- atics of the fauna, and a more meaningful zoogeo- graphic analysis of the fauna. Material and Methods This study is based on the examination of ap- proximately 8700 specimens collected throughout the Antarctic, Subantarctic, and adjacent waters and includes new material from 482 lots collected at 310 stations. Most of the specimens were col- lected by the vessels participating in the United States Antarctic Research Program (USNS Eltanin, now the ARA Islas Orcadas, and R/V Hero) and those participating in Operation Deep Freeze III and IV. These specimens, as well as those from other vessels (e.g., USCGC Eastwind, USFCS Albatross, and R/V Anton Bruun), are deposited at the United States National Museum. Other collections examined include specimens collected by the R/V Vema (depos- ited at the American Museum of Natural History), specimens collected by the Walther Herwig (depos- ited at the Zoologisches Institut und Zoologisches Museum, Hamburg), and a small collection of USNS Eltanin corals deposited at the Museum of Compara- tive Zoology, Harvard, Cambridge. Previously reported specimens from the following museums were examined: British Museum, London; Museum National d'Histoire Naturelle, Paris; Museum fur Naturkunde an der Humboldt-Universltat Berlin; Institut Royal des Sciences Naturelles de Belgique, Brussels; the Australian Museum, Sydney; the South Australian Museum, Adelaide; Museum of Comparative Zoology, Harvard, Cambridge; American Museum of Natural History, New York; and the United States National Museum, Washington, D. C. Eguchi's [1965] specimens could not be found at the National In- stitute of Polar Etesearch, Tokyo, or the Tokyo Kasei University. Synonymies are complete unless otherwise indi- cated. In the sections on material the numbers in pa- rentheses indicate the number of specimens in each lot. The number of specimens is not indicated for colonial species. Following this information, or the station number for colonial species, is an indication of where the specimen is deposited. If the vessel's cruise number is included, it precedes the station number and is linked to it with a hy- phen. Most of the holotypes and paratypes of the new species are deposited at the United States National Museum; the remainder are at the British Museum. A confirmed depth range is used to avoid erro- neous end points resulting from bathymetrically wide-ranging trawls. The stated bathymetric range for each species extends from the deepest shallow to the shallowest deep component of the trawled depth ranges [see Cairns, 1979, p. 10]. Copyright 1982 by the American Geophysical Union. 1 BIOLOGY OF THE ANTARCTIC SEAS XI TABLE 1. Chronological Listing of Research on Antarctic and Subantarctic Scleractinia Reference Vessel and/or Expedition Remarks Milne Edwards and Haime [1848] Moseley [1876] Studer [1878] Moseley [1881] von Marenzeller [1903] von Marenzeller [1904a] Pax [1910] Gardiner [1913] Gravier [1914a] Gravier [1914b] David and Priestley [1914] Gardiner [1929a] Thomson and Rennet [1931] Gardiner [1939] Niino [1958] Du Petit-Thouars, Venus HMS Challenger SMS Gazelle HMS Challenger SY Belgica, Expedition Antarctique Beige (1897-1899) Valdivia, Deutsche Tiefsee-Expedition (1898-1899) Gauss, Deutsche Siidpolar-Expedition (1901-1903) Scotia, Scottish National Antarctic Expedition (1902-1904) Pourquoi-Pas? DeuxiSme Expedition Antarctique Franqaise (1908-1910) Pourquoi-Pas? Deuxi&me Expedition Antarctique Franijaise (1908-1910) Nimrod British Antarctic Expedition (1907-1909) Terra Nova, British Antarctic Expedition (1910) Aurora Australasian Antarctic Expedition (1911-1914) RRS Discovery, RSS William Scoresby (1926-1933) Umitaka-Maru, JARE 3 Specimens of Flabellum thouarsii from off Falkland Islands; deposited at MNHNP. Preliminary report on Challenger Scleractinia, including records of 3 species from off Tristan da Cunha Group and Prince Edward Islands; deposited at EM. 2 specimens of Flabellum thouarsii reported from off southern Argentina; specimens deposited at Museum fur Naturkunde, Berlin. Final report on Challenger Scleractinia, including 10 species (5 of them new) from various Subantarctic localities, including off eastern and western southern South America, off Tristan da Cunha Group, and off Prince Edward Islands; deposited at BM. 5 specimens of 2 species from 3 stations off Peter I Island, Antarctica; deposited at Institut Royal des Sciences Naturelles de Belgique, Brussels. 2 specimens of C. antarct ica from 1 station off Bouvet0ya; deposited at Museum fiir Naturkunde, Berlin. 7 specimens of 3 species from 5 stations off Gaussland, Antarctica; histology of Flabellum impensum. 3 specimens of CaryophyIlia profunda from 1 station off Gough Island. 3 specimens of DesmophyHum antarcticum ( < Javania antarctica) from 2 stations off Palmer Archipelago; deposited at MNHNP. 13 specimens of 4 species from 3 stations off Antarctic Peninsula; deposited at MNHNP. Fossil Gardineria antarctica from Mount Larsen; deposited at the Australian Museum, Sydney. 19 specimens of 3 species from 4 stations in Ross Sea; deposited at BM. 7+ specimens of 3 species from 2 stations off Queen Mary Coast, Antarctica. Another 2 scleractinian species from off Maria Island, Tas- mania; all but F. australe are mis- identified; deposited at the Aus- tralian Museum, Sydney. 280 specimens of 12 nominal species from 33 stations off southern South America and Antarctic Peninsula; 9 of 12 species reas- signed; deposited at BM. 4 specimens of 4 species from 2 stations off Riiser-Larsen Peninsula (Cape Cook), Antarctica; all species misidentified; deposition unknown. CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA TABLE 1. (continued) Reference Vessel and/or Expedition Remarks Wells [1958] Squires [1961] Speden [1962] Squires [1962b] Squires [1963a] Squires [1964c] Squires [1965b] Eguchi [1965] Bullivant [1967] Squires [1969] Keller [1974] Podoff [1976] Sorauf and Podoff [1977] Present study Discovery, BANZARE (1929-1931) R/V Vema (cruise 14) HMNZS Endeavour (1958-1960) United States Exploring Expedition (1838-1842) HMNZS Endeavour (Macquarie Gap Crni sp) Umitaka-Maru and Soya, JARE HMNZS Endeavour D/E Ob, Academic Kurchatov miscellaneous miscellaneous USNS Eltanin; ARA Islas Orcadas; R/V Hero (USARP program) ; USS Atka; USS Burton Island; USS Staten Island; USS Edisto (Deep Freeze expeditions); HMNZS Endeavour, Rotoiti, Viti (NZOI vessels); R/V Vema, Walther Herwig; USCGC East- wind, cruise 66 97 specimens of 5 species from 10 stations off coast of Antarctica between 40? and 80?E and 110? and 120?E; 3 of 5 species reassigned; deposited at the South Australian Museum, Adelaide. 92 specimens of 4 species (including 2 new species) from 6 stations off the Falkland Islands and Tierra del Fuego; desposited at AMNH. Review of fossil records of Gardineria antarctica on Antarctic continent. 242 specimens of 5 species (including 2 new species) from 13 stations in the Ross Sea; reviews distribution of all Antarctic Scleractinia; presumably deposited at NZOI. 2 fossilized specimens of 2 species collected at Tierra del Fuego; deposited at MCZ. Discussion of research potential of Antarctic Scleractinia; no specimens reported. Specimens of 3 species from 1 station east of the Auckland Islands; deposited at NZOI and USNM. 9 specimens of 9 nominal species (including 2 new species) from 6 stations off Riiser-Larsen Peninsula (Cape Cook), Antarctica; specimens overlap with those of Niino [1958]; less than half of species correctly identified; deposition unknown. Description of Gardineria antarctica assemblage in Ross Sea; specimens previously reported by Squires [1962b]. Reviews distribution of all Antarctic Scleractinia; provides distri- bution maps for 23 species; includes new records but does not document them (presumably based on USARP and NZOI specimens); deposited at USNM and probably NZOI. 357 specimens of 2 species from 3 stations from off southeastern South America and Falkland Islands; F. antarcticum misidentified; deposition unknown. Examination of microstructure by SEM of 6 Antarctic species; unpublished M. A. thesis; deposited at USNM. Examination of microstructure of 3 Antarctic species; deposited at USNM. 8700 specimens of 39 species (including 7 new species) from 310 stations throughout Antarctic and Sub-antarctic; primarily deposited at USNM, also at AMNH and MCZ. BIOLOGY OF THE ANTARCTIC SEAS XI Some specimens have been coated with dark dye and recoated with a fine layer of ammonium chloride in order to improve their contrast for photography. These specimens are noted in the plate legends. The following abbreviations are used in the text: Vessels EAD EW GLD NZOI PD USARP WH US Museums AMNH BM MCZ NMNH MNHNP NZOI SME USNM ZIZM ZMA Other GCD LCD CD PD H S x' SEM CS USCGC Eastwind. USCGC Eastwind, cruise 66. USS Glacier, Deep Freeze IV Expedition. collected by the New Zealand Oceanographic Institute, including the HMNZS Endeavour, Rotoiti, and Viti. pebble dredge (used in conjunction with some R/V Vema stations). United States Antarctic Research Program. Walther Herwig. RSS William Scoresby. American Museum of Natural History, New York. British Museum (Natural History), London. Museum of Comparative Zoology, Harvard, Cambridge. see USNM. Museum National d'Histoire Naturelle, Paris. New Zealand Oceanographic Institute, Wellington. Station Marine d'Endoume, Marseille (most of these specimens will be deposited at the Museum National d'Histoire Naturelle, Paris). United States National Museum, Smithsonian Institution, Washington, D. C. Zoologisches Institut und Zoologisches Museum, Hamburg. Zodlogische Museum, Amsterdam. greater calicular diameter. lesser calicular diameter. calicular diameter. pedicel diameter. height. septa, costae, pali, or costosepta of cycle designated by numerical subscript. scanning electron microscopy. Checklist of Species Known From the Antarctic and Subantarctic Regions Order SCLERACTINIA Bourne, 1900 Suborder FUNGIINA Verrill, 1865 Family FUNGIIDAE Dana, 1846 Fungiacyathus Sars, 1872 F. marenzelleri (Vaughan, 1906) F. fragilis G. 0. Sars, 1872 Family MICRABACIIDAE Vaughan, 1905 Leptopenus Moseley, 1881 L. sp. cf. L. discus Moseley, 1881 Suborder FAVIINA Vaughan and Wells, 1943 Superfamily FAVIICAE Gregory, 1900 Family RHIZANGIIDAE d'Orbigny, 1851 Astrangia Milne Edwards and Haime, 1848 A. rathbuni Vaughan, 1906 Phyllangia Milne Edwards and Haime, 1848 P. fuegoensis Squires, 1963 Family OCUHNIDAE Gray, 1847 Bathelia Moseley, 1881 B. Candida Moseley, 1881 Madrepora Linnaeus, 1758 M. oculata Linnaeus, 1758 Suborder CARYOPHYLLIINA Vaughan and Wells, 1943 Superfamily CARYOPHYLLIICAE Gray, 1847 Family CARYOPHYLLIIDAE Gray, 1847 Subfamily CARYOPHYLLIINAE Gray, 1847 CaryophyIlia Lamarck, 1801 C. antarctica Marenzeller, 1904 C. squiresi n. sp. C. profunda Moseley, 1881 C. eltaninae n. sp. C. mabahithi Gardiner and Waugh, 1938 Cyathoceras Moseley, 1881 C. irregularis n. sp. Aulocyathus Marenzeller, 1904 A. recidivus (Dennant, 1906) n. comb. Subfamily TURBINOLIINAE Milne Edwards and Haime, 1848 Sphenotrochus Milne Edwards and Haime, 1848 S^. gardineri Squires, 1961 Subfamily DESMOPHYLLIINAE Vaughan and Wells, 1943 DesmophyHum Ehrenberg, 1834 D. cristagalli Milne Edwards and Haime, 1848 forma cristagalli Milne Edwards and Haime, 1848 forma ingens Moseley, 1881 forma capense Gardiner, 1904 Lophelia Milne Edwards and Haime, 1849 L. prolifera (Pallas, 1766) Subfamily PARASMILIINAE Vaughan and Wells, 1943 Solenosmilia Duncan, 1873 ?. variabilis Duncan, 1873 Goniocorella Yabe and Eguchi, 1932 G. dumosa (Alcock, 1902) CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 15 Superfamily FLABELLICAE Bourne, 1905 Family FLABELLIDAE Bourne, 1905 Flabellum Lesson, 1831 F. thouarsii Milne Edwards and Haime, 1848 F. areum n. sp. F. curvatum Moseley, 1881 F. impensum Squires, 1962 F. flexuosum n. sp. F. gardineri n. sp. F. knoxi Ralph and Squires, 1962 F. apertum Moseley, 1876 forma apertum Moseley, 1876 forma patagonichum Moseley, 1881 F. truncum n. sp. Javania Duncan, 1876 J. cailleti (Duchassaing and Michelotti, 1864) J. anCarctica (Gravier, 1914) n. comb. Gardineria Vaughan, 1907 G. antarctica Gardineria, 1929 Family GUYNIIDAE Hickson, 1910 Stenocyathus Pourtal&s, 1871 332), pi. 11, figs. 1-5, 7.?von Marenzeller, 1904b, p. 76.?Gravier, 1920, p. 97 (part).?? Eguchi, 1965, pp. 5-7, pi. 1, figs. 4a-4c. Fungia symmetrica; Duncan, 1873, p. 334, pi. 49, figs. 16-19. Bathyactis marenzelleri Vaughan, 1906b, p. 66, pi. 4, figs. 1, la, lb. Fungiacyathus symmetricus; Wells, 1958, pp. 267, pi. 2, figs. 1, 2.?Squires, 1962b, p. 13; 1969, p. 17, pi. 6, map 2. Fungiacyathus marenzelleri; pi. 7, figs. A-K.? Cairns, 1979, pp. 35-37, pi. 2, figs. 8, 9, pi. 3, figs. 3, 8. Zibrovrtus, 1980, pp. 24, 25, pi. 6, figs. A-M. Description. Base of corallum flat, center of base slightly raised. Largest specimen known measuring 40 mm across base; largest Antarctic specimen 38.5 mm in diameter. Forty-eight thin, ridged costae radiating from center of base, be- coming more raised and sinuous toward calicular edge. Costae sometimes projecting as much as 1.5 mm from base at calicular edge. Base extremely thin and fragile, sometimes perforate, especially toward calicular edge. Five to seven synapticular plates occurring in every interseptal space, be- coming increasingly larger and more oblique toward edge of corallum. S. vermiformis (PourtalSs, 1868) Suborder DENDROPHYLLIINA, Vaughan and Wells, 1943 Family DENDROPHYLLIIDAE Gray, 1847 Balanophyllia Wood, 1844 B. malouinensis Squires, 1961 B. sp. B. chnous Squires, 1962 Enallopsammia Michelotti, 1871 E. rostrata (Pourtalfes, 1878) E. sp. cf. E. marenzelleri Zibrowius, 1973 Uncertain Records Caryophyllia clavus var. smithi sensu Moseley, 1881 Flabellum transversale conicum sensu Eguchi, 1965 Flabellum ongulense Eguchi, 1965 Desmophyllum pseudoseptatum Eguchi, 1965 Species Account Order SCLERACTINIA Bourne, 1900 Suborder FUNGIINA Verrill, 1865 Family FUNGIIDAE Dana, 1846 Genus Fungiacyathus Sars, 1872 Diagnosis. Solitary, cupolate, free. Septotheca thin; costae thin and spinose. Septa irregularly dentate, laterally braced by thin ribbons extend- ing from septotheca and by thin septa! striae. Columella feeble. Paliform lobes sometimes pres- ent. Type-species: Fungiacyathus fragilis Sars. 1872, by monotypy. 1. Fungiacyathus marenzelleri (Vaughan, 1906) Plate 1, figs. 1, 2, 8 Bathyactis symmetrica; Moseley, 1881, pp. 186-190 (part: Challenger sta. 133, 147, 157, 299, 325. Septa hexamerally arranged in four cycles. '1 largest septa and only ones reaching columella without fusion to other septa. Septa of remaining cycles progressively smaller. Each S2 reaching columella, there joined by pair of S3,all loosely fused or covered over by triangular canopy composed of thin calcareous plate. Pairs of S^ fused to S3 by similar but larger and higher canopies, these extending from S2-S3 canopies to halfway to calicular edge. These canopies often perforate. Septa laterally carinate, Sj^ possessing about 7-10 carinae, or about 1 every 1.9 mm. Carinae about 0.4 mm high and usually symmetrical on both sides of septum. Near columella, carinae more closely spaced and corresponding to high septal spines. Carinae vertical near columella, oblique midway between columella and calicular edge and almost horizontal near calicular edge. Most car- inae extending from septal edge to base; some shorter, extending only halfway to base; some branching from other carinae. Near base most car- inae curving toward columella, often degenerating into rows of granules. If still solid, carinae may be confused with synapticulae but can usually be distinguished by their more oblique orientation, often intersecting synapticulae at an acute angle. Septa bearing elongate, slender spines near colu- mella but becoming less serrate and usually lobate in profile toward calicular edge. Height of exsert lobes up to 10 mm above base. All septa but S^ bearing lobes, these lobes damaged in most speci- mens. Septa extraordinarily fragile and specimens rarely collected fully intact. Septal edges straight to irregularly sinuous. Columella vari- able in size, composed of loose fusion of inner septal spines and additional trabeculae. Discussion. F. marenzelleri was frequently reported as Bathyactis or F. symmetricus by earlier authors, probably because of Moseley's [1881] as- sumption that all small specimens were simply ju- veniles of a species with a larger adult corallum. F. symmetricus has subsequently been shown to be Indemic to the western Atlantic (183-1644 m) and to be rarely larger than 14 mm in CD [Cairns, BIOLOGY OF THE ANTARCTIC SEAS XI 1, 2, 8. 3-7. Plate 1. Fungiacycathus Fungiacyathus marenzelleri (Vaughan): 1, 2, USNM 47476, Eltanin sta. 134, CD - 32.3 mm; 8, USNM 47477, Eltanin sta. 1545, x4.3, several septa coated with ammonium chloride. Fungiacyathus fragilis Sars: 3, 6, 7, USNM 47536, Eltanin sta. 1412, CD = 24.4 nmi (3 and 7 coated with ammonium chloride); 4, holotype of F. hawaiiensis Vaughan, USNM 20834, Albatross sta. 4125, CD =*23.1 mm; 5, USIM 47537, Eltanin sta. 1846, CD = 21.3 mm, base. CAIRNS; ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 1979]. F. marenzelleri is distinguished from F. symmetricus by its much larger corallum, broad- er distribution, and greater depth range. F. symmetricus also has higher septal spines and a more solid, well-defined base. Specimens from four localities (Eltanin stations 138, 426, 993, and 1545) differ from typical F. marenzelleri by having a much more crowded ar- rangement of septal carinae (about 20 per septum, or 1 every 0.8 mm) and more spinose septal faces. These specimens, all from the South Shetland Islands and east of South Orkney Islands, may represent a new species, but until the variation of F. marenzelleri is better understood, they are assigned to this species. Eguchi's [1965] ^. symmetrica also seems to belong to this form. When the deeper records of Fungiacyathus spp. are reexamined [e.g., Moseley, 1881; Gardiner and Waugh, 1939; Keller, 1976], more synonyms for F. marenzelleri may result, and its geographic range may thus be increased. All records deeper than 1800 m should be reevaluated for this possibility. This species may eventually be found to be the most widespread and deepest-living species of scleractinian coral. Material. Eltanin sta. 13 (5), USNM 47470; sta. 18 (23), USNM 47466; sta. 20-134 (5), USNM 47476; sta. 138 (3), USNM 47475; sta. 353 (1), USNM 47471; sta. 426 (8), USNM 45673; sta. 993 (1), USNM 47473; sta. 997 (6), USNM 47474; sta. 1148 (1), USNM 47469; sta. 1545 (3), USNM 47477; sta. 1957 (1), USNM 47666. Hero sta. 721-1081 (4), USNM 47468. Glacier sta. 11 (6), USNM 47465. Albatross sta. 4397 (1), USNM 47467. Specimen identified as F. symmetricus by Wells [1958], South Australian Museum H 70; specimens listed by Cairns [1979], USNM. Types of B. marenzelleri. Types. The holotype of F. marenzelleri is de- posited at the United States National Museum (47415); three paratypes are at the Museum of Com- parative Zoology. Type-locality: 8?07.5'S, 104?10.5'W (off Peru); 3820 m. Distribution. Widely distributed throughout At- lantic Ocean as far north as Greenland; eastern Pacific; circum-Subantarctic (including off lies Crozet); off South Shetland Islands; east of South Orkney Islands; WeddeU Sea; ? off Prince Harald Coast and Enderby Land, Antarctica (Map 1). Depth range: 300-5870 m. There is a direct relationship between depth of occurrence and proximity to the Antarctic, the more southerly records being shallower. The shal- lowest records of this species (300-500 m) are represented by the four continental Antarctic records; the four records from the South Shetland Islands range from 300 to 1435 m. No other record is shallower than 1805 m. 2. Fungiacyathus fragilis G. 0. Sars, 1872 Plate 1, figs. 3-7 Fungiacyathus fragilis M. Sars, 1869, pp. 250, 265, 274 (nomen nudum).?G. 0. Sars, 1872; p. 58, pi. 5, figs. 24-32. Cairns, 1979, p. 206. ?Zibrowius, 1980, pp. 23, 24, pi. 5, figs. A-J. Bathyactis symmetrica; Verrill, 1882, p. 313; 1883, p. 65.?Gravier, 1920, p. 97 (part).?Thomson, 1931. p. 9. Bathyactis hawaiiensis Vaughan, 1907, pp. 145, 146, pi. 27, figs. 1, la. Fungiocyathus fragilis; Jungersen, 1916, (part).~Broch, 1927, p. 8.~Nordgaard, 1929, p 103. 35 Description. Base of corallum flat, very thin, sometimes irregularly perforate. Evidence of re- generation from fragments of corallum. Largest specimen reported 45 mm in basal diameter; largest specimen from Subantarctic 25 mm in diameter. Slightly ridged C1-3 radiate from center of base. Septa hexamerally arranged in five complete cy- cles, fifth cycle appearing at CD of 9-10 mm. Septal arrangement similar to previously described species: S^ independent; inner edges of septa of remaining cycles fused to one another by thin, perforate triangular lamellae (canopies). Each larger septum forming large nonserrate lobe for most of its length, with few, if any, projecting spines near columella. Height of septal lobes up to one fifth to one fourth of CD. These lobes, as well as all septa, with highly sinuous outer and upper edges, corresponding to septal corrugations. Corrugations vertical near upper edge, becoming more horizontal as they curve toward columella near base. About 12 corrugations per septum, or 1/mm, giving septa 'wrinkled' aspect. Crests of corrugations regularly spaced, usually rounded and smooth, but may bear row of low, pointed granules or may even be slightly carinate. Small, pointed granules usually on all septa near columella. Septa extraordinarily fragile. All septa joined to adjacent septa by synapticular plates, these plates increasing in size toward calicular edge. About 7-10 plates occurring per centimeter, con- tinuing to add on as corallum increases in dia- meter. Columella round and small, sometimes a solid, horizontal lamella but usually a loose fusion of inner edges of larger septa. Discussion. There are four other nominal Recent species of Fungiacyathus with five cycles of septa. F. fragilis differs from western Atlantic JF. pus- illus (Pourtal&s, 1868) in being larger and having sinuous septa, from Indian Ocean F. stephanus (Al- cock, 1893) in having a flat base and lower septa, and from Indian Ocean and eastern Pacific E|. pali- ferus (Alcock, 1902) in lacking paliform lobes. It is indistinguishable from F_. hawaiiensis (Vaughan, 1907); however, comparisons Involving only one specimen (the holotype) cannot be con- clusive. F. kikaiensis (Yabe and Eguchi, 1942), also with five cycles, is from the Pliocene- Pleistocene of Japan. Material. Eltanin sta. 1412 (5+), USNM 47536; sta. 1846 (3), USNM 47537. Specimens of Verrill [1882], (Yale Peabody Museum, New Haven) YPM 8322 and USNM 47538-47539; specimens of Zibrowius [1980], SME. Holotype of B. hawaiiensis. Types. One syntype of F. fragilis is deposited at the Oslo Museum (B626). Type-locality: ' Skraaven in Lofoten'; 549 m. The holotype of B. hawaiiensis is deposited at the United States National Museum (20834). Type-locality: between Oahu and Kauai islands, Hawaii; 1761-2056 m. Distribution. Eastern Atlantic in area bordered by Norway, Cape Verde Islands, and the Azores; off Massachusetts; off Hawaii; west of South Island, New Zealand; Macquarie Ridge (Map 1). Worldwide depth range: 285-2200 m; New Zealand-Macquarie records: 1647-1693 m. BIOLOGY OF THE ANTARCTIC SEAS XI Plate 2. Leptopenus, Astrangia, and Phyllangia 1-3. 4-6. 7-9. Leptopenus sp. cf. L. discus Moseley: 1, USNM 47481, Eltanln sta. 1545, CD = 18.2 mm; 2, USNM 47483, Eltanln sta. 2002, CD = 16.5 mm, base; 3, same specimen, calice. 6, specimen reported by Squires [1963a], MCZ USNM 10974, CD = 6-7 mm. holotypic colony and calices, MCZ 5390, CD = Astrangia rathbuni Vaughan: 4, 2520, CD = 5-6 mm; 5, holotype Phyllangia fuegoensis Squires: 7.5 X 6.5 mm. CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA Map 1. Distribution of Fungiacyathus marenzelleri (solid circles), Fungiacyathus fragilis (solid triangles), and Bathelia Candida (solid squares). Family MICRABACIIDAE Vaughan, 1905 Genus Leptopenus Moseley, 1881 Diagnosis. Solitary, discoidal, free. No wall, costae alternating in position with septa. Costae and septa united by simple synapticulae producing very porous, delicate corallum. Columella trabec- ular. Type-species: Leptopenus discus Moseley, 1881, by subsequent designation [Wells, 1936]. 3. Leptopenus sp. cf. L. discus Moseley, 1881 Plate 2, figs. 1-3 Leptopenus discus Moseley, 1881, pp. 205-208, pi. 14, figs. 1-4, pi. 16, figs. 1-7.?Not L. discus Dennant, 1906, p. 162 (? Letepsammia sp.).? Wells, 1958, p. 262.?Squires, 1965a, pp. 878, 879, fig. 1: 1967. p. 505; 1969, p. 17, pi. 6, map 2.?Keller, 1977, p. 37, fig. 1.?Cairns, 1979, pp. 37, 38, pi. 3, figs. 4-7. Leptonemus discus; Agassiz, 1888, p. 154, fig. 479 (misspelling). Description. Corallum discoidal and extremely fragile. Base of corallum flat to slightly con- cave, especially near center. Largest Antarctic specimen 18.0 mm in CD. Septa hexamerally arranged in two cycles and multiple bifurcation of septa of third cycle, un- like species from any other family of Scleractinia. (The traditional method of designating septa of higher cycles as pairs of new septa flanking each previously formed septum does not apply to Lepto- penus, and therefore a new system of terminology is introduced here (Text figure 1).) Each S3 bi- furcating about 2 mm from columella, the two resul- ting septa being referred to as S3'. Not far from this junction the S3' bifurcate, each form- ing two S3''. Sometimes, near edge of calice, pair of S3''' forming, as described by Moseley for syntypes. Branching of S3 not always symme- 10 BIOLOGY OF THE ANTARCTIC SEAS XI Fig. 1. Diagraimnatic representation of one system of Leptopenus discus. Heavy lines represent septa, light lines costae. trical within one system or consistent in systems of one corallum. (For instance, in a half system of a specimen from Eltanin station 2002, one S3' produced only two S3'', whereas the other S3' produced two S3'' and four S3'''. Also, one of the syntypes [Moseley, 1881, pi. 14, fig. 1] is illustrated as having a pair of S3' ' ' ' in the upper right system. The regular arrangement of 72 septa with one pair of S3''' per system to which Moseley alluded is probably just one variation of septal arrangement.) $1 only independent septa, extending to col- umella and bearing eight or nine long, recurved spines. S2 also extending to columella and bear- ing five or six similar spines, with pair of S3 joining each S2 close to columella S3 and all of its bifurcations bearing similar recurved spines decreasing in size toward calicular edge. Thin, triangular canopy USLilly at point of junction of S3 to S2 and at every bifurcation of S3. Septal spines of S2 just distal of S2-S3 junction usually tallest spines on corallum, rising about 1 Iran above columellar spines. Septa very low and solid, composed mainly of large spines united by thin lamella. No septal granulation. Columella a very spiny mound in center of calice, merging with inner septal spines. Base of corallum consisting of radiating network of bifurcating costae, these costae alternating- in position with overlaying septa. Costae not ar- ranged like septa, because all costae bifurcate; none independent (Text figure 1). Each costa at- tached to its two adjacent septa by thin synaptic- ular bridges; space between bridges forming el- liptical pores. Viewed from above or below, corallum appearing regularly perforate. Pores increasing in size from center to calicular edge. For inner two thirds of corallum, synapticular bridges reaching upward from costae to meet septa; in outer part of corallum, however, septa becoming rudimentary or absent and synapticular bridges becoming wider and horizontal, but pores still persist. At calicular edge, costae forming thin spines projecting up to 1.6 mm beyond synapticular zone, or 17-20% of calicular radius. Discussion. The Antarctic specimens are only provisionally identified as L. discus because (1) they are taller than the syntypes, (2) the septal arrangement is sligntly different, and (3) they sometimes lack canopies at the junctions of septa. The reasons that these specimens are taller (up to 3.8 mm) may be the slight concavity of the base or their intact septal spines. Most of the spines are broken off of the syntypes, which measure less than 2 mm in height. The septal arrangement and presence or degree of development of canopies may be a matter of individual variation. Variation in this species is very poorly known. Only 6 speci- mens of L. discus have been reported previously, 2 of which were fragments. Although the 44 specimens reported herein were all collected with tissue intact, upon cleaning, much of the corallum fell apart, especially the peripheral areas. Additional whole specimens are needed to study variation of septal arrangement and costal spines. L. discus is distinguished from L. irinae Keller, 1977, by its shorter costal and peripheral septal spines. It is easily distinguished from L. hypo- coelus Moseley, 1881, by its lesser height and smaller S2 spines and from L. solidus Keller, 1977, by its alternating septa and costae. Material. Eltanin sta. 598 (1), USNM 47480; sta. 1545 (2), USNM 47481; sta. 1926 (1), USNM 47479; sta. 2002 (3), USNM 47483; sta. 2108 (37), USNM 47482. Specimen of Agassiz (1888], USNM 46916. Syntype from Challenger sta. 147. Types. The four syntypes of L. discus, collected at Challenger stations 147, 157, and 323, are de- posited at the British Museum. The specimen from Challenger station 147 is numbered 1880.11.25.159. Type-locality: since a lectotype has not been designated, the type-localities are off Rio de la Plata, South America, and Subantarctic Indian Ocean; 2926-3566 m. Distribution. Off Cuba; off Rio de la Plata; Makassar Strait, Indonesia; off South Orkney Is- lands; off South Sandwich Islands; Subantarctic Indian Ocean (including off lies Crozet); Ross Sea (Map 2). Worldwide depth range: 2000-3566 m; Antarctic records: 2035-2384 m. Suborder FAVIINA Vaughan and Wells, 1943 Superfamily FAVIICAE Gregory, 1900 Family RHIZANGIIDAE d'Orbigny, 1851 Genus Astrangia Milne Edwards and Haime, 1848 Diagnosis. Colonial, extratentacular budding forming cerioid, plocoid, or reptoid coralla. Corallites united basally by thin coenosteura or stolons. Septa dentate; columella papillose. Type-species: Astrangia michelini Milne Edwards and Haime, 1848, by subsequent designation [Milne Edwards and Haime, 1850]. 4- Astrangia rathbuni Vaughan, 1906 Plate 2, figs. 4-6 7 Astrangia Verrill, 1869, p. 526. Astrangia rathbuni Vaughan, 1906a, pp. 849-850, p. 78, figs. 1-3.?Squires, 1963a, pp. 10, 11, figs. 3-7.?Laborel, 1971, pp. 200, 201, pi. 6, fig. 1, map 7.?Zibrowius, 1974c, pp. 165, 166.?Not A. CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA U Map 2. Distribution of Madrepora oculata (solid circles), Madrepora vitiae (open circles), and Leptopenua discus (solid squares). rathbuni; Avent et al., 1977, p. 200 (is A. astreiformis Milne Edwards and Haime, 1849). ? Astrangia sp. Pax, 1910, p. 74.?Gravier, 1914b, p. 121.?Squires, 1961, p. 20. Description. Small encrusting colonies 30-50 mm in diameter; corallites united by basal coenosteura. Cylindrical corallites 5-6 mm in diameter project- ing up to 9 mm from coenosteum. Sometimes extra- tentacular budding from other corallites. Coral- lites weakly costate, usually brownish. Septa hexamerally arranged in four systems. S]^, S2, and S3 equal in size, sloping grad- ually toward deep fossa. S^ half as large and joining adjacent S-j halfway to columella. Inner edges of all septa highly dentate (beaded), each bearing 7-10 irregular teeth. Columella indis- tinguishable from lower septal edges, consisting of mass of similarly shaped teeth. Remarks. An X ray diffraction analysis of a small fragment from specimen MCZ 2520 revealed a very high calcite peak, indicating a subfossil age for the specimen. Discussion. Verrill [1869], in a general dis- cussion of the genus Astrangia, mentioned that one species (not named) occurred in the Strait of Magellan. Pax [1910], Gravier [1914b], and Squires [1961] perpetuated this obscure record as Astrangia sp., none of whom documented its occurrence. Zibrowius [1974c] suggested that both Verrill's [1869] report of the unnamed Astrangia and Squires's [1963a] record of A. rathbuni from off Tierra del Fuego were based on the same specimens. Both authors were familiar with the Museum of Com- parative Zoology coral collection, where Squires's specimens are deposited. Despite their fossilized condition. Squires's [1963a] Tierra del Fuego spec- imens are definitely A. rathbuni. Material. Specimens of Squires [1963a], MCZ 2520; specimens (A. astreiformis) of Avent et al. 12 BIOLOGY OF THE ANTARCTIC SEAS XI 1-3. 4-6. 7-9. Plate 3. Bathelia, Madrepora, and Caryophyllia Bathelia Candida Mosely: 1, syntype branch, BM, Challenger sta. 320, xl.2; 2, USNM 47512, Vema sta. 17-14', GCD = 9.5 mm; 3, same specimen, H = 98 mm. Madrepora oculata Linnaeus: 4, USNM 47499, Eltanin sta. 1346, xO.8; 5, specimen from same lot, xl.3; 6, USNM 47515, NZOI sta. B-314, CD = 2.1 ram, coated with ammonium chloride, forma vitiae. Caryophyllia antarctica Marenzeller: 7, USNM 45677, Eltanin sta. 138, GCD 28.7 mm] - 20.0 n 8, 9, syntype. Museum fiir Naturkunde 5067, Valdivia sta. n, H = 21.6 mm. 127, GCD CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 13 [1977J, Florida Department of Natural Resources FSBC I 14492. Holotype. Types. The holotypic colony of A. rathbuni is deposited at the United States National Museum (10974). Type-locality: Paqueta, Rio de Janeiro. Distribution. Off eastern coast of South America from 22''S to 37?59'S; Tierra del Fuego fossil spec- imens collected by United States Exploring Expedi- tion (1838-1842) but unfortunately without precise locality data. Several to 90 m. Genus Phyllangia Milne Edwards and Haime, 1848 Diagnosis. Colonial, extratentacular budding forming reptoid colonies, united basally by thick coenosteum. Inner septa! edges smooth to slightly dentate. Columella rudimentary; P3 sometimes present. Type-species: Phyllangia americana Milne Edwards and Haime, 1849, by subsequent designation [Milne Edwards and Haime, 1850J. 5. Phyllangia fuegoensis Squires, 1963 Plate 2, figs. 7-9 Phyllangia fuegoensis Squires, 1963a, pp. 13, 16, figs. 1, 2. Description. In the following the holotypic colony is described. Fragmentary corallum very worn, perhaps the result of fossilization. Frag- ment about 35 X 20 X 20 mm, bearing 21 corallites. Corallites cylindrical, encrusting a bivalve shell and budding from parent corallites. Corallites up to 10-15 mm tall with elliptical calices measuring up to 7.5 X 6.5 mm in diameter. Septa hexamerally arranged in four cycles. S^ thickest and most exsert; septa of remaining cycles progressively smaller. Inner septal edges worn but appear to be straight, entire, and vertical, except for S3, these usually having small, pointed paliform lobes. Pair of lobes often meeting or at least bending toward each other across each S2. Low, pointed granules on septal faces. Columella rudimentary, crispate, surrounded by paliform lobes. Discussion. Like the preceding species, P. fue- goensis was also collected by the United States Exploring Expedition from Tierra del Fuego, without precise locality. Contrary to Squires's [1963a] comparison, it is quite distinct from Astrangia floridana (Gane, 1895) but very similar to P. americana Milne Edwards and Haime, 1849. The lat- ter species shows great morphological variation in paliform lobes, corallite size, and colony form and has been found living off Rio de Janeiro [Laborel, 1971] and in the Pliocene of Venezuela [Weisbord, 1968] and Florida [Weisbord, 1974]. In view of the variation and distribution of P. amer- icana and the unique but unfortunately vague dis- tributional record of P. fuegoensis it is possible that they will be synonymized when more specimens become available. Material. Holotype Types. The holotypic colony is deposited at the Museum of Comparative Zoology (5390). Type- locality: Tierra del Fuego (? fossil). Distribution. Known only from type-locality. Family OCULINIDAE Gray, 1847 Genus Bathelia Moseley, 1881 Diagnosis. Colonial, extratentacular budding forming dendroid coralla. Coenosteum dense. Sep- tal edges smooth. Crown of pali before S3; columella of irregular ribbons. Type-species: Bathelia Candida Moseley, 1881, by monotypy. 6. Bathelia Candida Moseley, 1881 Plate 3, figs. 1-3 Bathelia Candida Moseley, 1881, pp. 177, 178, pi. 8, figs. 1-6.?Wells, 1958, p. 262.?Cairns, 1979, p. 206. Description. Colony dendroid, corallites ar- ranged in opposite and alternating fashion on branch. Extratentacular budding most common; how- ever, intratentacular budding also occurring. May have three branches originating at one calice. Maximum size of colony unknown. Branches robust (about 1 cm in diameter) and solid; no dissepiments present. Coenosteum smooth and very finely granu- lated. Thin, shallow coenosteal striae bordering wide, flat costae corresponding to all septa. Calices round to slightly elliptical, 6-10 mm in diameter, projecting obliquely several millimeters from branch. Septa hexamerally arranged in four cycles. S^ and S2 equal in size and extending to columella; S3 and S/j progressively smaller. All septa narrow and slightly exsert, extending over thick- ened calicular edge as low ridges. Inner septal edges straight; those of Sj and S2 usually entire, those of S3 and S4 usually dentate. Septal granulation variable, ranging from abundant, small, fine granules to sparse larger, blunt granu- les. Lower inner edges of S3 bearing tall, nar- row pali, these bordering columella and terminating at same height as columellar papillae. Pali usually distinguished from papillae by their larger size and elliptical to rectangular shape in cross sec- tion. Columella composed of 5-15 tall, slender papillae, these irregularly round in cross section. Pali and columellar papillae granulated. Because of narrow septa, fossa rather wide but not very deep, being filled in with pali and columella. Discussion. Bathelia is a monotypic genus and has been reported from only one locality previ- ously. Material. Vema sta. 17-14, USNM 47512. Calypso sta. 171, USNM 47513 and SME. Following WH records (H. Zibrowius, personal communication, 1979): sta. 215/66, sta. 142/71, sta. 191/71, sta. 197/71, sta. 328/71, sta. 329/71, sta. 331/71 (all WH specimens deposited at ZIZM). Syntypes. Types. The syntype branches, from Challenger sta. 320, are deposited at the British Museum. Type-locality: 37?17'S, 53?52'W (off Rio de la Plata); 1097 m. Distribution. Off southern South America from Rio Grande, Brazil, to Cabo Tres Puntas, Argentina; off Peninsula de Taitao, Chile (Map 1). Depth range: 500-1250 m. Genus Madrepora Linnaeus, 1758 Diagnosis. Colonial, extratentacular budding forming dendroid colonies. Coenosteum dense, no costae, corallites filled internally by stereome. No pali; columella spongy or absent. Type-species: Madrepora oculata Linnaeus, 1758, by subsequent designation [Verrill, 1901]. 14 BIOLOGY OF THE ANTARCTIC SEAS XI Plate 4. Caryophyllia 1-4. Caryophyllia antarctica Marenzeller: 1, USNM 47304, Eltanin sta. 1933, GCD = 14.6 nun; 2, USNM 53414, Yelcho 2-11, GCD = 13.1 nnn; 3, specimen identified as Caryophyllia clavus by Thomson and Rennet [1931], Australian Museum G 13536, sta. 10, GCD = 17.9 mm; 4, USNM 47302, Glacier sta. 1, H = 16.0 mm. 5-9. Caryophyllia squiresi n. sp.: 5, paratype, USNM 47161, Eltanin sta. 558, GCD = 15.6 mm; 6, paratype from same lot, GCD = 17.1 mm; 7, paratype from same lot, H = 38.3 mm; 8, 9, holotype, USNM 47160, Eltanin sta. 558, GCD = 15.5 mm, H = 31.0 mm, theca coated with ammonium chloride. CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 15 7. Madrepora oculata Linnaeus, 1758 Plate 3, figs. 4-6 Madrepora oculata Linnaeus, 1758, p. 798.?von Mar- enzeller, 1904b, p. 79.?Eguchi, 1968, p. C-29, pi. C-8, figs. l-9.~Zibrowius, 1974a, pp. 762- 766, pi. 2, figs. 2-5; 1980, pp. 36-40, pi. 13, figs. A-P.?Cairns, 1979, pp. 39-42, pi. 3, fig. 2, pi. 4, fig. 5, pi. 5. figs. 1-3. Amphihelia oculata; Milne Edwards and Haime, 1837, p. 119.?von Marenzeller, 1904a, p. 308, pi. 14, figs. 1, lb. Amphihelia ramea; Duncan, 1873, p. 326, pi. 44, figs. 1-3, pi. 45, figs. 4-6, pi. 46, figs. 1-19. Lophohelia Candida Moseley, 1881, pp. 179, 180, pi. 9, figs. 6-13. ? Madrepora vitiae Squires and Keyes, 1967, p. 22, pi. 1, figs. 4-8. Description. Colony bushy or flabellate, formed by extratentacular budding. End branches having sympodial arrangement of corallites, measuring between 2.3 and 4.0 mm in diameter; diameter of attached base up to 2 cm. Calices round, 2.4-3.8 mm in diameter, exsert on end branches, recessed or flush with coenosteum toward base. Coenosteum smooth, extremely finely granulated; costae and coenosteal striae rare. Septa hexamerally arranged in three cycles. S^ equal to or larger than S2; S3 much smal- ler, sometimes rudimentary. Inner edges of septa straight, sometimes thickened near columella. Septal faces covered by granules, sometimes twice as high as septal thickness. Fossa variable in depth, usually dependent on age of corallite, older corallites having shallower fossae. Columella variable, usually papillose, sometimes absent. Discussion. More complete synonymies and de- scriptions are given by Zibrowius [1974a, 1980] and Cairns [1979]. Zibrowius [1974a] lists the nominal species of Madrepora and discusses their relation- ship to M. oculata. Madrepora oculata is a widespread and extremely variable species. Characters that are subject to variation, sometimes within the same colony, in- clude frequency of branching, intercorallite dis- tance, coenosteum texture and color, relative sep- tal sizes, septal granulation, fossa depth, and development of columella. A closely related, if not identical, species, M. vitiae Squires and Keyes, 1967, was also collected off New Zealand (Eltanin stations 1814, 1816, and 1818; NZOI station C-642) (Map 2). The only dif- ference between the two is that M. vitiae usually has paliform lobes, sometimes quite thick, before the S2. However, a branch of a topotypic speci- men of M. vitiae has corallites with and without pali, and some calices have a variable number of paliform lobes (1-6). The T-shaped inner septal edges mentioned by Squires and Keyes [1967] were not observed in specimens collected from the type- locality (NZOI station B-314) or in specimens from five other lots near the type-locality. They may have been referring to the slight thickening of the inner septal edges, which is common in M. oculata. Zibrowius's [1974a] M. oculata from off lie Saint-Paul, Indian Ocean, is similar to M. vitiae; however, he did not consider the presence of P2 as a specific difference. If the presence or ab- sence of P2 is considered to be of no specific value, then M. vitiae may be dropped to a form. Material. Eltanin sta. 254, USNM 47500; sta. 1346, USNM 47499; sta. 1403, USNM 47501; sta. 1416, USNM 47665; sta. 1422, USNM 47497; sta. 1814, USNM 47502; sta. 1816, USNM 47498; sta. 1818, USNM 47504. NZOI sta. C-642, USNM 47514; sta. D-6, USNM 47503. Specimens listed by Cairns [1979], USNM; topotypic specimens of M. vitiae from NZOI sta. B-314, type lot, USNM 47515. Syn- types of L. Candida. Types. The types of M. oculata are lost. Type- locality: off Sicily and Tyrrhenian Sea, Mediter- ranean. Syntypes of L. Candida are deposited at the British Museum (1880.11.25.95). Type-locality: off Sombrero island. Lesser Antilles; 823 m. The holotype of M. vitiae is deposited at the New Zealand Oceanographic Institute (17). Type- locality: off Cape Farewell, New Zealand; 230-251 m. Distribution. According to Zibrowius [1974a, p. 776], distribution of M. oculata worldwide outside of polar seas. Three of above-mentioned records extend the southernmost distribution of M. oculata to Subantarctic waters: Hjort Seamount, a seamount in the Subantarctic South Pacific, and a seamount in the Drake Passage (Map 2). Worldwide depth range: 80-1500 m; Subantarctic records: 549-833 m. Suborder CARYOPHYLLIINA Vaughan and Wells, 1943 Superfaraily CARYOPHYLLIICAE Gray, 1847 Family CARYOPHYLLIIDAE Gray, 1847 Subfamily CARYOPHYLLIINAE Gray, 1847 Genus Caryophyllia Lamarck, 1801 Diagnosis. Solitary; ceratoid, turbinate, or subcylindrical; fixed or free. Septotheca usually costate. Pali opposite S3 in one crown (or before second group of septa when hexameral symme- try obscured). Columella fascicular, formed of twisted ribbons. Type-species: Madrepora cyathus Ellis and Solander, 1786, by subsequent designation [Broderip, 1828]. 8. Caryophyllia antarctica Marenzeller, 1904 Plate 3, figs. 7-9; Plate 4, figs. 1-4 Caryophyllia antarctica Marenzeller, 1903, p. 1 (nomen nudum); 1904a, pp. 293, 294, pi. 16, figs. 7, 7d.~Pax, 1910, pp. 65, 66, pi. 11, fig. l.~ Gravier, 1914b, pp. 129, 130, pi. 1, figs. 7, 8. ?Wells, 1958, pp. 267, 268, pi. 2, figs. 3, 4. ?Squires, 1961, p. 20; 1962b, pp. 13, 14, 16, 17, pi. 1, figs. 11, 12; 1969, pp. 16, 17, pi. 6, map 1.?Eguchi, 1965, pp. 7, 8, pi. la, lb. Cairns, 1979, p. 206. Caryophyllia clavus; Thomson and Rennet, 1931, p. 40. Caryophyllia arcuata; Gardiner, 1939, pp. 331, 332. Description. Corallum ceratoid to trochoid, usually straight, attached. Pedicel diameter one fifth to one third of GCD, expanding only slightly at substrate. Largest corallum examined 28.6 x 26.2 mm in CD and 36.5 mm tall; however, more typ- ical coralla 10-15 mm in GCD and 15-20 mm tall. Theca usually smooth, porcelaneous, sometimes with flat, equal costae bordered by shallow intercostal striae. Costal granules rare; when present, low and rounded. Calice round to elliptical. Septa usually hexamerally arranged in four cycles. Sj and S2 equal in size and exsert- 16 BIOLOGY OF THE ANTARCTIC SEAS XI ness; S3 and S4 progressively smaller. Larger coralla with up to 90 septa, accommodated by in- crease in number of half systems and acceleration of higher-cycle septa instead of by addition of another cycle of smaller septa. Inner edges of ^1> S2, and $4 slightly sinuous, those of S3 and P3 very sinuous. Septal granulation prominent, usually arranged in widely spaced rows on septal undulations oriented parallel to septal edge. Individual granules sometimes quite tall, with rounded, clavate, bifid, or squared-off tops. Granules usually fused into low, distinct carinae, these having continuous or serrated (beaded) upper edges. Carinae especially well developed near inner septal edges. Fossa shallow. Pali of varying widths (up to width of S3) stand before S3; each separated from its corresponding septum by deep, narrow notch. Pali sometimes split into two smaller lobes. Pali sometimes present before S^ and S2. Palar granulation similar to that of septa but more prominent; carinae running obliquely across palus. Columella composed of 4-20 discrete, slender, twisted ribbons aligned in greater axis of caliCular ellipse. Discussion. C. antarctica is distinguished by its distinctive septal ornamentation of carinae and squared-off granules. Thomson and Rennet's [1931] C. clavus is a typical specimen of C. ant- arctica. Material. Eltanin sta. 138 (4), USNM 45677; sta. 416 (1), US^M 47307; sta. 428 (6), USNM 45670; sta. 678 (1), USNM 47291; sta. 992 (1), USNM 47306; sta. 1067 (1), USNM 47296; sta. 1081 (2), USNM 47301; sta. 1082 (4), USNM 47297; sta. 1084 (8), USNM 47317; sta. 1870 (11), USNM 47309; sta. 1883 (2), USNM 47289; sta. 1922 (1), USNM 47294; sta. 1930 (1), USNM 47311; sta. 1931 (1), USNM 47285; sta. 1933 (11), USNM 47304; sta. 1995 (25), USNM 47316; sta. 1966 (45), USNM 47315; sta. 2007 (l), USNM 47300; sta. 2022 (3), USNM 47205; sta. 2079 (3), USNM 47293; sta. 2104 (1), USNM 47668; sta. 2106 (1), USNM 47286; sta. 2119 (1), USNM 47288; sta. 5765 (6), USNM 47284. Is las Qrcadas sta. 876-118 (1), USNM 47303. Hero sta. 721-849 (2), USNM 47308; sta. 731-1812 (1), USNM 47318. Yelcho sta. 2-11 (5), USNM 53414. Glacier sta. 1 (1), USNM 47302. Edisto sta. 21 (1), USNM 47920; sta. 31 (2), USNM 47293; sta. 36 (1), USNM 47310. Staten Island sta. 21 (1), USNM 47298. Atka sta. 23 (42), USNM 47313. Burton Island sta. 3 (20), USNM 47305. EW sta. 4 (1), USNM 47287; sta. 35 (5), USNM 47314. GLD sta. 15 (1), USNM 47299. Specimens (3) identified as Caryophyllia clavus by Thomson and Rennet [1931], Australian Museum G 13536; specimens of Wells [1958] from the following Discovery stations: sta. 39 (12), H 43; sta. 41 (3), H 46; sta. 40 (1), H 47 (all deposited at the South Australian Museum, Adelaide). Syntypes. Types. Four syntypes collected by the Belgica (station 290 (3) and station 569 (D) are deposited at the Brussels Museum. Another two syntypes from Valdivia station 127 are deposited at the Museum fiir Naturkunde, Berlin (5067). Type-locality: near Peter I Island, Antarctica, and off BouvetjSya; 567 m. Distribution. Endemic to the Antarctic region, probably circumpolar. Squires's [1969] records from Subantarctic South America undocumented (Map 3). Depth range: 87-1435 m. Caryophyllia squiresi n. sp. Plate 4, figs. 5-9 Caryophyllia sp. A Squires, 1969, p. 17 (part: 3 of 4 South American records only), pi. 5, map 1.?Cairns, 1979, p. 206. Description. Corallum ceratoid, sometimes be- coming cylindrical, often bent near base but rarely by more than 40?. Attached by narrow pedicel usually 2.9-3.4 mm in diameter (18-25% GCD of adult corallum). Holotype 15.6 x 14.5 mm in CD, 3.0 mm in PD, and 31.0 mm tall. Largest specimen 18.6 mm in GCD. Costae equal, smooth, porcela- neous, and bordered by very thin, intercostal striae. Calice slightly elliptical. Septa hexamerally arranged in four cycles. Sj^ and S2 equal in size; S3 and S4 progressively smaller. Pairs of smaller 85 present in larger coralla, flanking S4; one specimen has 10 S5, or 58 septa. These S4 then enlarged to almost size of an S3. Septa not exsert. Inner septal edges straight, except tor those of S3, these sometimes sinuous. Septal granulation sparse and usually nonlinear. Granules usually low and blunt, rarely squared off, and never fused into carinae. Fossa moderately deep. Tall, narrow pali present before S3, each separated from its corresponding septum by deep, narrow notch. Palar margins usually sinuous and granulation usually more prom- inent than that of septa. Pali (usually 12) often forming distinct crown but may merge indistin- guishably with columella. Columella composed of 3-10 discrete, twisted ribbons aligned in plane of greater axis of calicular ellipse. Discussion. This species corresponds to at least one lot of specimens identified by Squires (Eltanin station 558) and referred to by him [Squires, 1969] as Caryophyllia sp. A. Two more of his records are consistent with specimens at the United States National Museum; however, the other 17 records [Squires, 1969, pi. 6, map 1] from off western South America, Antarctica, and New Zealand are undocumented. Until these specimens are found and verified, the distribution of C. squiresi will remain as follows: off Tierra del Fuego and off the Falkland Islands (Islas Malvinas). C. squiresi is similar to C. antarctica but can be distinguished by its lack of septal carinae, deeper fossa, less sinuous inner septal edges, and less exsert septa. Etymology. This species is named in honor of D. F. Squires, who has done much to advance the knowl- edge of Antarctic Scleractinia and who first recognized this species. Material. Eltanin sta. 339 (4), USNM 47516. Vema sta. 15-PD9 (1), USNM 47517, and (1), AMNH. Types. Types. The holotype, collected at Eltanin sta- tion 558, is deposited at the United States Na- tional Museum (47160). Twenty-two paratypes, col- lected at Eltanin station 558, are deposited at the United States National Museum (47161), and another specimen from the same station is deposited at the British Museum (1979.11.1.1). Type- locality: 51?58'S, 56?38'W (off East Falkland island); 646-845 m. Distribution. See discussion (Map 3). Depth range: 406-659 m. CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 17 Map 3. Distribution of Caryophyllia antarctica (solid circles), Caryophyllia squiresi (solid squares), and Caryophyllia clavus var. smithi (solid triangle). 10. Caryophyllia profunda Moseley, 1881 Plate 5, figs. 1-5 Caryophyllia profunda Moseley, 1881, pp. 138, 139 (part: specimen from Cape Verde Islands is C. cyathus), pi. 1, figs. 6, 6b.?Not C. profunda; Jourdan, 1895, pp. 10, 11 (is C. cyathus Ellis and Soiander, 1786).?von Marenzeller, 1904a, p. 298.?Gardiner, 1913, pp. 688, 689.?Not C. pro- funda ; Gravier, 1920, p. 28 (is Caryophyllia foresti Zibrowius, 1980).?Gardiner, 1929a, p. 126; 1939, p. 331.?Ralph, 1948, p. 108, fig. 2 (top).?Squires, 1958, p. 44; 1960, pp. 196, 198-200, pi. 34, figs. 5-7, pi. 35, figs. 9-11; 1962b, pp. 13-15, pi. 1, figs. 13, 14; 1964a, p. 11; 1969, pp. 16, 17, pi. 6, map 1.?Ralph and Squires, 1962, pp. 3, 6, 7, pi. 1, figs. 8-11.? Squires and Keyes, 1967, pp. 15, 17, 23, pi. 2, figs. 1-4.?Zibrowius, 1974a, pp. 751-755, pi. 1, figs. 1-10.?Beurois, 1975, p. 46, photo 13.?Cairns, 1979, p. 206. Caryophyllia cyathus; von Marenzeller, 1904a, p. 295, pi. 16, figs. 6, 6a.?Hoffmeister, 1933, p. 14, pi. 4, figs. 4, 5.?Gardiner, 1939, pp. 330, 331.?Squires, 1961, p. 17. Caryophyllia planilamellata Dennant, 1906, pp. 157, 158, pi. 6, figs. 4a, 4b.?Squires, 1961, p. 18. Caryophyllia clavus; Wells, 1958, p. 265, pi. 1, figs. 12, 13. Caryophyllia cf. C. maculata; Ralph, 1948, p. 108, fig. 2 (bottom, right).?Ralph and Squires, 1962, pp. 3, 7, pi. 2, figs. 1, 2.?Squires and Keyes, 1967, pp. 15, 17, 23, pi. 2, figs. 4, 5. Description. Corallum trochoid to cylindrical, straight to slightly bent; strongly attached by broad, encrusting base. Pedicel variable in diam- eter, ranging from 20 to 70% of GCD. Pedicel usually greatly increased in diameter by concentric layers of external stereome. Large specimens up to 41 ram in GCD and 50 mm tall. Individual coralla 18 BIOLOGY OF THE ANTARCTIC SEAS XI Plate 5. Caryophyllia 1-5. 6-9. Caryophyllia profand a Moseley: 1, USNM 47519, Eltanin sta. 1718, H = 48.7 mm; 2, specimen from same lot, GCD = 28.2 mm; 3, specimen reported by Gardiner [1939], BM 1939.7.20.207, Discovery sta. 1187, GCD = 25.1 mm; 4, specimen identified as C. cyathus by Gardiner [1939], BM 1939.7.20.210, Discovery sta. 190, GCD = 16.2 mm; 5, syntype, BM 1880.11.25.36, Challenger sta. 135, GCD = 25.5 mm. Caryophyllia eltaninae n. sp.: 6, 7, holotype, USNM 47162, Eltanin sta. 671, H = 35.6 ram, GCD = 26.8 mm; 8, USNM 47163, Eltanin sta. 671, GCD = 25.8 mm; 9, specimen from same lot, H = 28.0 ram. CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 19 sometimes clumping into pseudocolonial arrangement. Calice round in young specimens, becoming ellip- tical in larger specimens. Theca porcelaneous, finely granulated, and often brownish. Costae usually flat and equal, although C\-j sometimes moderately ridged. Septa hexamerally arranged in five cycles. Sj and S2 equal in size and highly exsert; higher-cycle septa progressively smaller and less exsert. Calices with more than 96 septa rare, but those with less than 96 septa and 24 pali common, roughly a function of smaller calicular diameter. Inner edges of all septa straight, except those of S4, these sometimes slightly sinuous. Septal granulation variable, usually consisting of extremely fine, low granules, but sometimes larger, blunt granules; never arrang- ed in carinae. Fossa moderately deep. Narrow pali occurring before S4; each separated from its corresponding septum by a deep and narrow to shallow and broad notch. Pali sometimes bilobed or trilobed. Colu- mella variable, composed of several linearly arranged, twisted ribbons; or a fused mass of twisted ribbons generally aligned in greater calic- ular axis; or a labyrinthiform arrangement of modi- fied twisted ribbons. Remarks. Only Moseley [1881] recorded observa- tions of the living coral. He stated that the ground color of the polyp was transparent blue, encircled by a sulphur-yellow margin at the calic- ular edge. The stomadeum was white or vermillion, and the short tentacles were red knobbed. In one of the few papers that document growth rates for deepwater corals, Squires [1960] esti- mated the growth rate for this species as 0.88-2.02 mm/year in height. He also hypothesized on fea- tures characteristic of cessation of growth (or maximum size), such as lobation of pali and septa; increased thickness of septa and pedicel; and coarsened septal, palar, and costal ornamentation. Discussion. Gardiner's [1939] record of C. pro- fund a from Discovery station 190 is the only con- tinental Antarctic record for the species and pro- duces an unusual distribution pattern, which in- cludes predominantly cold temperate records, two marginal Subantarctic records (Tristan and Gough islands), and Gardiner's single Antarctic record. Zibrowius [1974a, p. 754] distinguished Gardiner's Antarctic specimen from typical C. profunda by its narrower and deeper notches between the septa and pali and the more vertical edges of its pali. Among the specimens that I have examined, I find these characters to be within the range of varia- tion for the species and, in general, not of specific value. Assuming that no labeling errors were made, the somewhat anomalous distribution of C. profunda must stand. C. profunda is easily distinguished from other Antarctic CaryophyIlia by its greater size and the presence of five cycles of septa with pali before the fourth cycle. Material. Eltanin sta. 1403 (1), USNM 47518; sta. 1718 (37), USNM 47519; sta. 1814 (1), USNM 47520. Specimens (8) identified as C. profunda and C. cyathus by Gardiner [19391, BM 1939.7.20.202-203, 207-213; some specimens of Squires and Keyes [1967], i.e., B-489 (5), C-690 (4), C-703 (5), all at USNM; some species of Zibrowius [1974a], i.e., AMS-66 (1), AMS-1474, off ile Amsterdam, 80 m, Jan. 1972, all at USNM. Ele- ven syntypes. Types. Approximately 20 syntypes of C. profunda, collected at Challenger station 135, are deposited at the British Museum (1880.11.25.36, 1880.11.25.241, 1889.7.8.1-5). The syntype frag- ment from Cape Verde Islands is C. cyathus [see Zibrowius, 1974a]. Type-locality: 37?0r50"S, 12?19'10"W (off Nightingale Island, Tristan da Cunha Group); 183-274 m. At least one syntype of C. planilamellata Dennant is deposited at the Aus- tralian Museum (G 12057). Distribution. Circumpolar in southern temperate waters: off South Africa, lie Saint-Paul and lie Amsterdam, South Australia, New Zealand, and Chat- ham Island; Subantarctic islands of Tristan and Gough; off Hugo Island, Palmer Archipelago. Squires's [1969] Subantarctic records from off South America and the Macquarie Ridge are unsub- stantiated (Map 4). Most common between 80 and 250 m; confirmed range: 35-1116 m. 11. Caryophyllia eltaninae n. sp. Plate 5, figs. 6-9 Gardineria lilliei; Gardiner, 1939, pp. 328, 329 (part: two specimens from Discovery sta. 160). Description. Corallum ceratoid to trochoid, attached or free. If attached, corallum usually straight, with reinforced pedicel up to 38% of GCD; if free, corallum often slightly bent, with an eroded base as small as lOX of GCD. Holotype (largest specimen) 26.8 x 23.5 ram in CD, 6.1 ram in PD, and 35.6 mm tall. Costae usually nongranulated and porcelaneous, bordered by thin intercostal striae. ^1^-3 slightly ridged in some speci- mens. Calice round to elliptical. Septa hexamerally arranged in five cycles. Sj^ and 83 equal in size and exsertness; septa of higher cycles progressively smaller. Full fifth cycle attained at about 13.5 mm CD. Sj and S2 extending to columella; S5 rudimentary, with ir- regularly dentate inner edges. Inner edges of all septa and pali straight. Septal granules low and blunt, never arranged in carinae. Pali (12) occurring before S3 and variable in shape; usually tall and narrow, but sometimes tri- angular or twisted like a columellar ribbon. In about one fourth of specimens examined, paliform lobes also present on S/^, often in form of small, horizontally projecting lobe directed at, and sometimes merging with, adjacent P3. Palar granulation more prominent than that of septa. Columella variable, composed of 4-30 discrete, twisted ribbons or fused mass of twisted elements aligned in greater axis of calicular ellipse. Discussion. Gardiner's [1939] misidentification of this species as G. lilliei is a result of the small size of his species (CD = 9.8 x 9.8, 9.0 x 10.0 mm), which at this stage could be confused with Gardineria. His specimens were just beginning to form pali and S5; both specimens had only one palus, corresponding to the half systems where S5 had formed. C* eltaninae is unusual in that its pali occur before the antipenultimate septal cycle, not the penultimate as is common in most Caryophyllia. This is a character shared with the Caribbean C. paucipalata Moseley, 1881. It is further distin- guished from C. antarctica and C. squiresi by its straight inner septal edges. Etymology. This species is named after the R/V 20. BIOLOGY OF THE ANTARCTIC SEAS XI Map 4. Distribution of Caryophyllia profunda (solid circles), Caryophyllia eltaninae (solid squares), Caryophyllia mabahithi (solid triangle), Cyathoceras A (open circle), and Cyathoceras irregularis (open square). Eltanin, from which many of the specimens used in this study were collected, including the holotype of this species. Material. Eltanin sta. 678 (2), US^M 47486; sta. 1535 (1), USNM 47485. Is las Orcadas . sta. 575-8 (1), USNM 47487; sta. 575-10 (3), USNM ^7490; sta. 575-12 (3), USNM 47489; sta. 575-14 (2), USNM 47488; sta. 575-17 (5), USNM 47484; sta. 575-93 (2), USNM 47491. Specimens (2) identified as G. lilliei by Gardiner [1939J, BM 1939.7.20.286-287. Types. Types. The holotype, collected at Eltanin sta- tion 671, is deposited at the United States National Museum (47162). Thirty-nine paratypes from Eltanin station 671 (number 47163) and 11 paratypes from Islas Orcadas station 575-11 (number 47164) are also deposited at the United States National Museum. One paratype from Eltanin station 671 is also deposited at the British Muse- um (1979.71.2.1.). Type-locality: 54?41'S, 38?38'W (off southwest South Georgia); 220-320 m. Distribution. Known only from the shelf and slope off the western half of South Georgia and off Shag Rocks (Map 4). Depth range: 101-261 m, except for one record at 778-814 m. 12. Caryophyllia mabahithi Gardiner and Waugh, 1938 Plate 6, figs. 1-5 Caryophyllia mabahithi Gardiner and Waugh, 1938, pp. 178, 179, text fig. 1, pi. 3, fig. 6.?Gardiner, 1939, p. 332.?Wells, 1958, p. 262.?Squires, 1961, p. 21; 1962b, pp. 14, 16, 17, pi. 1, figs. 15, 16. Description. Corallum free, trochoid, generally curved between 45? and 90?. Base 1.2-1.5 ram in CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 21 1-5. 6-9. Plate 6. Caryophy ILi-a and Cyathoceras Caryophyllia mabahithi Gardiner: 1-3, specimens reported by Gardiner (1939), BM 1939.7.20.247-248, Discovery sta. 182, GCD = 9.0 mm, 8.6 mm; 4, 5, syn- type, BM 1950.1.9.561-586, John Murray sta. 34, GCD = 9.8 mm, H = 9.3 mm. Cyathoceras irregularis n. sp.: 6, 7, paratype, USNM 47166, Eltanin sta. 1346, H = 18.5 mm, GCD = 15.8 mm; 8, 9, holotype, USNM 47165, Eltanin sta. 1346, H = 14.9 mm, GCD = 13.7 mm. 22 BIOLOGY OF THE ANTARCTIC SEAS XI diameter, usually showing hexameral partitioning of basal plate caused by first-cycle septa. An average-sized specimen is 9.3 x 8.3 ram in CD and 9.6 mm tall; however, Gardiner reported specimens 12 X 10 mm in CD and 13 mm tall. Costae flat or slightly ridged, covered by very low, rounded gran- ules arranged three or four across width of a costa near calice. Septa octamerally arranged in three cycles; addi- tional septa rare. S]^ highly exsert and extend- ing almost to columella; S2 and S3 progres- sively smaller and much less exsert. Sj and S3 with slightly sinuous inner edges, those of S2, however, more sinuous. Crown of eight pali occuring before S21 each palus separated from its corresponding septum by a deep, narrow notch. Paiar margins very sinuous. Septal granulation variable, consisting of pointed granules ranging from 0.5 to 2.0 times septal width in height; palar granulation usually even more prominent. Columella composed of one to seven broad, twisted ribbons fused among themselves and to inner edges of pali. Discussion. The three specimens reported by Gardiner [1939] from off the Palmer Archipelago are, by direct comparison, indistinguishable from syntypes of C. mabahithi from the Gulf of Aden. This disjunct distribution, both geographic and bathymetric, is very unusual and inexplicable. Two other species of octameral Caryophyllia are known: C. octopali Vaughan, 1907 (Hawaii), and C. barbadensis Cairns, 1979 (Barbados;. They are both readily distinguished by their subcylindrical, firmly attached coralla and smaller calices. Material. Specimens (3) of Gardiner [1939], BM 1939.7.20.246-248. Syntypes from John Murray sta. 34 (26 specimens). Types. Sixty-four syntypes of C. mabahithi are deposited at the British Museum. Those from sta- tion 34 are numbered 1950.1.9.561-586. Two syn- types from this lot have been permanently trans- ferred to the United States National Museum (48299). Type-locality: Gulf of Aden and Chagos Archipelago; 655-1022 m. Distribution. Gulf of Aden; Chagos Archipelago; off Anvers Island, Palmer Archipelago (Map 4). Depth range: 278-1022 m. Genus Cyathoceras Moseley, 1881 Diagnosis. Solitary, ceratoid to turbinate, fixed. Septotheca usually costate. No pali. Columella fascicular, composed of several twisted ribbons. Type-species: Cyathoceras cornu Moseley, 1881, by subsequent designation [Faustino, 1927]. 13. Cyathoceras irregularis n. sp. Plate 6, figs. 6-9 Description. Corallum ceratoid, straight to irregularly bent, attached by thin, encrusting, slightly expanded base. Pedicel one fourth to one third of GCD. Uolotype 13.7 x 10.5 mm in CD and 14.9 mm tall; largest specimen 15.4 x 14.6 mm in CD and 18.4 ram tall. Theca smooth and procela- neous, covered by very small, low granules. Costae occurring near calicular edge and base, if at all. Calice round to elliptical. Septa hexamerally arranged in four or five cycles; however, fifth cycle never complete; lar- gest specimen with 72 septa. Up to 48 septa stage. Sj and S2 equal in size, moderately exsert, and extending to columella. S3 and S4 pro- gressively narrower and less exsert. With addition of S5, septal arrangement becomes irregular. May have one or two pairs of S5 in each system. If only one pair present, enclosed (flanked) S4 invariably larger than unflanked S4. If both pairs of S5 present, both S^ enlarged, reaching almost as far toward columella as S3. Occurrence of all degrees of S5 development in one corallum possible, i.e., systems with 0, 1, or 2 pairs of S5, making interpretation of septal cycles con- fusing. Inner edges of S]^_3 broadly sinuous, corresponding to transverse septal undulations, but inner edges of higher-cycle septa straight. Large, blunt granules present on septal faces, usually arranged in lines along crests of septal undulations. Columella large, composed of numerous slender, twisted ribbons usually fused into solid mass. Remarks. All specimens examined were attached to dead coral, usually Solenosmilia variabilis. The type-locality is a seamount or ridge, which supports a presumed deepwater coral bank composed primarily of S. variabilis. C. irregularis is similar to Cyathoceras squiresi Cairns, 1979, in its association with the framework coral of deep- water banks. C. squiresi is often attached to Enallopsammia profunda, a common constituent of western Atlantic deepwater banks. Discussion. For the purposes of this comparison the following ten species are considered valid Cyathoceras: C. cornu Moseley, 1881; C. rubescens Moseley, 1881; C. tydemani Alcock, 1902; C. diomedeae Vaughan, 1907; C. niinoi Yabe and Eguchi, 1942; C. foxi Durham and Barnard, 1952; C. woodsi Wells, 1964; C. squiresi Cairns, 1979; C. avis (Durham and Barnard, 1952); and C. hoodens is (Dur- ham and Barnard, 1952). C. quay lei Durham, 1947, is herein transferred to Labyrinthocyathus, on the basis of an examination of the paratypes (USNM 547417), which have columellas composed of inter- connected lamellae. C. irregularis can be dis- tinguished from all of these species by its fused columella, composed of closely united, poorly de- fined, twisted ribbons; other Cyathoceras have well-defined columellar elements. It is also dis- tinguished by the irregularity of development of the fifth-cycle septa. Etymology. The specific name irregularis refers to the irregular manner in which the S5 are added. Material. Types. Types. The holotype, collected at Eltanin sta- tion 1346, is deposited at the United States Na- tional Museum (47165). Seven paratypes, also col- lected at Eltanin station 1346, are deposited at the United States National Museum (47166), and one paratype from this station is deposited at the British Museum (1979.11.3.1). Type-locality: 54?49'S, 129?48'W (seamount or ridge on Heezen fracture zone of Eltanin fracture zone system); 549 m. Distribution. Known only from type-locality (Map 4). 14. Cyathoceras sp. A Plate 7, figs. 1, 2 Description. Corallum ceratoid, straight, firmly attached. This specimen 8.9 x 8.6 mm in CD, 5.5 mm CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 23 Plate 7. Cyathoceras, Stephanocyathus, and Aulocyathus 1, 2. 3-6. 7-9. Cyathoceras sp. A: USNM 47521, ELtanin sta. 17-6, GCD = 8.9 ram, H = 16.5 mm. Stephanocyathus platypus (Moseley): 3, 4, syntype, BM 1880.11.25.57, Challenger sta. 164, CD = 48.4 mm; 5, 6, USNM 47522, Eltanin sta. 1718, CD = 53.0 mm. Aulocyathus reeidivus (Dennant): 7, USNM 47524, NZOI sta. C-734, CD = 8.8 mm; 8, 9, specimen from same lot, CD = 10.3 mm, H = 17.8 mm, theca coated with ammonium chloride. 24 BIOLOGY OF THE ANTARCTIC SEAS XI in PD, and 16.5 mm tall. Theca thin, porcelaneous, not granulated or costate. Thin, dull white longitudinal striae, corresponding to interseptal spaces, faintly distinguishable on parts of theca. Septa decamerally arranged in three septal sizes. Ten primaries are the largest septa, highly exsert (1.5 mm above calicular edge), and extending to columella. Ten secondaries are slightly exsert and extend two thirds of distance to columella. Twenty tertiaries barely exsert and extending one fourth of distance to columella. Inner edges of primaries and secondaries broadly sinuous (ampli- tude of undulations high and period long); ter- tiaries less corrugated. Long, prominent carinae on septal faces of primary and secondary septa, corresponding to crests of the broad septal cor- rugations. Tertiaries bearing only sparse septal granules. Columella composed of three broad, non- granulated, twisted ribbons, typical in shape for Cyathoceras. Carinae, septa, and columella all translucent. Discussion. The description above is based on only one specimen, which probably represents a new species; however, because of the lack of additional specimens to provide some indication of variation, it is not named here. This specimen was collected just north of the Subantarctic region as defined by Hedgpeth [1969] and is included here only because of its proximity to the Subantarctic region. Four of the eleven previously listed Cyathoceras have decameral symmetry: C. avis (Durham and Bar- nard, 1952); C. hoodensis (Durham and Barnard, 1952); C. woods! Wells, 1964; and C. squiresi Cairns, 1979. Cyathoceras A is distinguished from the first two eastern Pacific species by its firm attachment to the substrate and its lack of cos- tae. It is easily distinguished from C. woodsi by its larger size and septal carinae; however, it is similar to C. squiresi, especially in size, septal granulation, and septal sinuosity. The main points of difference are that Cyathoceras A has a thinner theca, exsert septa, and no costae. Furthermore, it is found far deeper than any other species of Cyathoceras. Material. Eltanin sta. 17-6 (1), USNM 47521. Distribution. Known from 52?10'S, 142?10'W (Tharp fracture zone of Eltanin fracture zone sys- tem) (Map 4). Depth range: 2305-2329 m. Genus Stephanocyathus Seguenza, 1864 Diagnosis. Solitary, patellate, free. Costae usually present. Paliform lobes usually present on all septa. Columella trabecular, papillose, or fused on surface. Type-species: Stephanocyathus elegans Seguenza, 1864, by subsequent designation [Wells, 1936]. 15. Stephanocyathus platypus (Moseley, 1876) Plate 7, figs. 3-6 Ceratotrochus platypus Moseley, 1876, p. 554. Stephanotrochus platypus; Moseley, 1881, p. 154, pi. 3, figs. 4a, 4b, 5a-5c.?Not S. platypus; Jourdan, 1895, pp. 19, 20 (is S. nobilis (Mose- ley, 1873)). Not Stephanotrochus diadema var. platypus; Gravier, 1920, pp. 46, 47 (is S^. moseleyanus (Sclater, 1886)). Stephanocyathus sp. Squires and Ralph, 1965, pp. 262, 263, figs. 3, 4. Stephanocyathus (S^. ) S?. Squires and Keyes, 1967, p. 24, pi. 2, figs. 11, 12. Stephanocyathus platypus; Zibrowius, 1980, p. 97. Description. Corallum free, bowl shaped, up to 75 mm in CD and 30 mm tall. Theca initially flat to slightly concave up to CD of 35-40 mm, then ca- licular edges turn upward rather abruptly and con- tinue to grow at an angle of 60?-70? from horizon- tal. Costae not prominent on horizontal section, but Ci and C2 usually sharply ridged on up- turned peripheral theca. Theca, except for Ci and C2, covered by very fine, low, rounded granules. Septa hexamerally arranged in five cycles; Sg common in larger coralla of up to 115 septa. S^ extraordinarily exsert in form of rounded lobe projecting up to 16 mm beyond theca. S2 also highly exsert; remaining septa barely exsert, only those flanking Sj^ and S2 rising higher than S3. Calicular margin scalloped, apices corre- sponding to Si and S2. Sj extending to center of calice, there considerably thickened and fused into rudimentary columella. S2 reaching almost to center and joining in fusion. S3 falling just short of fusion and terminating in slightly lobed free end. S^ slightly smaller than S3 and also bearing small, broad paliform lobe, this lobe bending toward adjacent S3 but rarely fus- ing with it. Where pairs of Sg present, enclosed S5 enlarged to almost size of S4 and also bearing small lobe bending toward adjacent S^. Normally, S5 and all Sg short, extending only one third of distance to center. S3, S4, and enlarged S5. bearing broad, low paliform lobes not separated by notches; S]^ and S2, however, without lobes and usually uniformly concave below level of theca. Septa straight with smooth inner edges. Septal granules low, blunt, and arranged in poorly defined lines. Discussion. There is little doubt that Squires and Ralph's [1965] Stephanocyathus sp. and Moseley's S^. platypus are identical. Moseley's [1876] original description was based on two small specimens with flat bases and calicular edges that had not yet turned upward; Squires and Ralph's very large specimen had an originally flat base that had subsequently become deeply bowl shaped. The three Eltanin specimens confirm the continuity of the ontogeny. S. diadema (Moseley, 1876) also has a flat base as a juvenile, which, like that of S. platypus, curves upward with greater size. Likewise, the base of S. laevifundus Cairns, 1979, is usually flat but is sometimes gently bowl shaped. S. platypus is most similar to S. moseleyanus (Sclater, 1886) from the northeast Atlantic, es- pecially in shape and septal exsertness. The lat- ter is distinguished by its papillose columella, Py and P2, and septal junctions near the columella. Material. Eltanin sta. 1718 (2), USNM 47522; sta. 1818 (1), USNM 47423. Syntypes. Types. Two syntypes of S. platypus, collected at Challenger station 164, are deposited at the British Museum (1880.11.25.57). Type-locality: 34?13'S, 151''38'E (off Sydney, Australia); 750 m. Distribution. Known from only four records from off Sydney, Australia; off New Zealand; and from a seamount (Eltanin station 1718) east of New Zealand (Map 5). Depth range: 622-913 m. Like the pre- CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 25 Map 5. Distribution of Sphenotrochus gardineri (solid circles), Aulocyathus recidivus (solid squares), Stephanocyathus platypus (solid triangles), and Lophelia prolifera (open circle). vious species, it does not occur in the Subantarc- tic region as defined by Hedgpeth [1969] but is included here because of its proximity to the Sub- antarctic region. Genus Aulocyathus Marenzeller, 1904 Diagnosis. Solitary, ceratoid, free. Longitu- dinal parricidal budding common. Columella trabec- ular. Type-species: Aulocyathus juvenescens Maren- zeller, 1904a, by monotypy. 16. Aulocyathus recidivus (Dennant, 1906) n. comb. Plate 7, figs. 7-9; Plate 8, fig. 1 Ceratotrochus recidivus Dennant, 1906, pp. 159, 160, pi. 6, figs. 2a-2c.?Squires, 1961, p. 18; 1969, p. 16.?Zibrowius, 1980, p. 107. Ceratotrochsu [sic] (Conotrochus) typus; Wells, 1958, pp. 265, 266, pi. 1, figs. 14, 15. ? Paracyathus conceptus; Squires and Keyes, 1967, p. 23 (part: C-648, pi. 2, figs. 7, 8). Description. The following description is based primarily on the largest specimen from NZOl sta- tion C-734. Corallum ceratoid, straight with round calice. Corallum 10.3 mm in CD and 17.8 mm tall. Most specimens originally attached to internal sur- face of fragment of parent specimen, from which they probably asexually budded. Some specimens originating from calice of unbroken parent speci- men, Theca glossy, granular, sometimes marked by shallow striae. Costal granulation often indis- tinct and irregular. Calice usually round but may be elliptical. Septa hexamerally arranged in four cycles. Sj^ larger than S2, these only slightly larger than S3. S4 smallest septa and never present as 26 BIOLOGY OF THE ANTARCTIC SEAS XI full cycle. Hexameral symmetry of younger speci- mens often not present; seven, eight, or nine groups of 4 or 6 septa often found. Large speci- men with 40 septa includes two complete systems and four systems missing one pair of S^ each. Septa not exsert with straight, vertical inner edges. Septal granules small and blunt, uniformly distributed. Fossa deep. Columella composed of 11 individual, irregularly shaped rods. Remarks. It is uncertain whether the corallum splits before the bud forms or whether the growing bud causes the corallum to fracture. Dennant [1906] implies the latter, whereas Marenzeller [1904aJ implies the former for a related species, A. juvenescens. Discussion. There are three other nominal species of Aulocyathus. A. recidivus differs from the two species known from off Japan, A. mactrici- dum (Kent, 1871) and A. conotrochoides (Yabe and Eguchi, 1932), by having a large, distinct columella. It differs from A. juvenescens Mar- enzeller, 1904 (off East Africa, 400-463), in being less slender and having fewer septa at a corres- ponding calicular diameter. Wells's [1958] speci- men measures 11.6 ram in CD, is 25.6 mm tall, and has 60 septa. Squires's [1969] reference to C. recidivus from the Macquarie Ridge was undoubtedly from NZOI station C-734. Material. NZOI sta. C-734 (9), USNM 47524. Golden Hind sta. 35 (1), MCZ. Specimen of Wells [1938] identified as C. (C.) typus from Discovery sta. 113, South Australian Museum U 31. Types. The 'numerous' syntypes of C. recidivus are not at the Australian Museum (Zibrowius, 1980] and have not been traced further. Distribution. Off southeastern Australia; off Tasmania; Macquarie Ridge (Map 128-732 m. 3.) Depth range: Subfamily TURBINOLIINAE Milne Edwards and Haime, 1848 Genus Sphenotrochus Milne Edwards and Haime, 1848 Diagnosis. Solitary, cuneiform, free; corallum small. Theca imperforate. Costae deeply ridged or reduced to aligned granules. Columella lamellar or papillose. Type-species: Turbinola crispa Lamarck, 1816, by subsequent designation [Milne Edwards and Haime, 1850]. 17. Sphenotrochus gardineri Squires, 1961 Plate 8, figs. 2-8 Sphenotrochus intermedius; Gardiner, 1939, p. 333 (part: Discovery sta. 388). Sphenotrochus gardineri Squires, 1961, pp. 26-28, 29, text figs. 6-8; 1969, p. 17, pi. 6, map 1.?Cairns, 1979, p. 206. Description. Corallum attached in younger stage. Base usually evenly rounded, but lateral edges of some specimens may form an apical angle of 30?-60?. Corallum cuneiform, highly compressed, with range of GCD/LCD of 1.5-2.2. Largest specimen known (Vema sta. 13-109) 9.3 x 4.4 mm in CD and 10.1 mm tall. Prominent, vertical costae correspond to all septa, with infrequent branching. The two principal (directive) costae, occurring on lateral edges, continuous to base, as are some costae on lateral faces; however, most costae originating at intercostal grooves flanking principal costae (Plate 8, fig. 3). Costae separated by deep, wide intercostal grooves. Row of low, rounded granules on each costa, sometimes becoming two granules wide near calice. Septa hexamerally arranged in four cycles. Si, ^2, and S3 equal in size and exsertness, except for the 2 principal septa, these aligned with columella and considerably larger than other four S|. S4 considerably smaller; S5 sometimes present in half systems adjacent to principals. Coralla with less than 48 septa, 10 or 11 half systems, common. S]^_3 having coarsely dentate upper and inner edges, these merging with columel- la. Inner edges of S4 finely dentate. Septal granules large and blunt, arranged in distinct lines originating from an axis of divergence well inside theca. Fossa shallow. Columella composed of five to eight linearly arranged rods, these fused basally; never lamellar. Remarks. Although adult coralla are always free, young specimens are initially attached to a sub- strate, such as sand particles of 0.1-0.2 mm in diameter. At a GCD of 2.3 mm, coralla have usually attained a full third cycle of septa, but costae and columella have not yet developed. A glossy epitheca is usually present. At a greater size, costae begin to develop and eventually overgrow the epitheca and original attachment, including the sand particle, at which point the coral becomes free. Coralla may also bud asexually from frag- ments of a parent specimen- Discussion. S. gardineri is distinguished from other Recent Sphenotrochus, all of which are fairly localized in distribution. The following species are distinguished on the basis of their lamellar columella: S. hancocki Durham and Barnard, 1952 (Galapagos Islands; off California); ^. auritus Pourtales, 1874 (tropical western Atlantic); S. gilchristi Gardiner, 1904; S. aurantiacus Marenzeller, 1904 (both off South Africa); and S. excavatus T. Hoods, 1878 (off Australia). Most of these species also have only three cycles of sep- ta. S. andrewianus Milne Edwards and Haime, 1848. (northeast Atlantic), has a lamellar or papillose columella but only three cycles of septa. Finally, S. ralphae Squires, 1964 (off New Zealand) [Squires, 1964b], has a similar columella, but its corallum is smaller and narrower and has fewer sep- ta. Material. Eltanin sta. 980 (2), USNM 47430. Hero sta. 712-636 (3), USNM 47432; sta. 713-694 (1), USNM 47434; sta. 715-902 (5), USNM 47433; sta. 715- 903 (3), USNM 47428. Vema sta. 14-14 (4), USNM 47435; sta. 14-16 (1), USNM 47431; sta. 15-109 (2), USNM 47429; sta. 17-11 (2), USNM 47427. Specimens of Gardiner [1939] from Discovery sta. 388 (2), MCZ; specimens of Squires [1961] from Vema sta, 14-14 (5), AMNH. Holotype. Types. The holotype is deposited at the American Museum of Natural History (3367). It was collected at Vema station 14-14. Type-locality: 54?23'S, 62?25'W (Burdwood Bank); 75 m. Distribution. Endemic to Magellanic region from Tierra del Fuego to Chiloe, Chile (Map 5). Depth range: 9-403 m. CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 27 Plate 8. Aulocyathus , Sphenotrochus, and Desmophy Hum 1. Aulocyathus recidivus (Dennant): USNM 47524, NZOI sta. C-734, H = 20.1 mm. 2-8. Sphenotrochus gardineri Squires: 2-4, USNM 47429, Vema sta. 15-109, H = 10.6 mm, GCD =9.7 mm; 5, 8, USNM 47428, Hero sta. 715-903, H = 3.5 mm, 4.4 mm. young stages with poorly developed costae; 6, 7, USNM 47432, Hero sta. 712-656, H = 3.5 mm, 3.6 mm, young attached stages showing budding from septal fragments. 9-12. DesmophyHum cristagalli Milne Edwards and Haime: 9-11, USNM 36367, Alba- tross sta. 2785, series of 3 specimens from same station illustrating formae cristagalli (GCD = 64.0 mm), capense (GCD = 50.4 mm), and ingens (GCD = 59.2 mm), respectively; 12, specimen identified as D. capense by Gardiner [1939], BM 1939.7.20.220, WS sta. 99, GCD = 66.4 28 BIOLOGY OF THE ANTARCTIC SEAS XI Plate 9. Desmophyllum, Solenosmilia, Lophelia, and Goniocorella 1-3. Desmophyllum cristagalll Milne Edwards and Haime: 1, specimen identified as D. capense by Gardiner [1939], BM 1939.7.20.220, WS sta. 99, xl.7; 2, USNM 36367, Albatross 2785, xO.52, long slender corallum; 3, specimen from same lot illustrating pseudocoloniality produced by settlement of successive solitary coralla, xO.33. 4, 5. Solenosmilia variabilis Duncan: 4, USNM 47426, Eltanin sta. 1346, xl.4, illustrating dichotomous extratentacular budding; 5, specimen from same lot, xO.54. 6. Lophelia prolifera (Pallas); USNM 47525, Eltanin sta. 1411, x3.2. 7-9. Goniocorella dumosa (Alcock): 7, USNM 47505, NZOI sta. D-175, x3.2, xO.46 and x9.8. endothecal dissepiments; respectively. 9, specimens from same lot. CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 29 Subfamily DESMOPHYLLIINAE Vaughan and Wells, 1943 Genus Desmophyllum Ehrenberg, 1834 Diagnosis. Solitary, trochoid, fixed. No pali. Columella absent or very small. Sparse endothecal dissepiments. Type-species: Desmophy- llum dianthus Ehrenberg, 1834, by subsequent des- ignation [Milne Edwards and Haime, 1850]. 18. Desmophyllum eristagalli Milne Edwards and Haime, 1848 Plate 8, figs. 9-12; Plate 9, figs. 1-3 The synonymy is complete for southern records only. Desmophyllum eristagalli Milne Edwards and Haime, 1848, p. 253, pi. 7, figs. 10, 10a.?von Maren- zeller. 1904a, pp. 267, 268, pi. 15, figs. 2a, 2b.?Gardiner, 1929a, pp. 125, 126.~Hof fmeis- ter, 1933, pp. 8, 9, pi. 2, figs. 1-4.?Ralph, 1948, p. 108, fig. 2 (bottom left) .?Wells, 1958, p. 262.?Squires, 1958, p. 91; 1961, pp. 18, 19; 1965, p. 785; 1969, p. 17, pi. 6, map 1. ?Ralph and Squires, 1962, pp. 9, 10, pi. 3, figs, 1-10.?Squires and Keyes, 1967, p. 25, pi. 3, figs. 12-14.?Zibrowius, 1974a, pp. 758-761, pi. 3, figs. 1-10; 1980, pp. 117-121, pi. 61, figs. A-0, pi. 62, figs. A-M.?Beurois, 1975, p. 46, photo 13.?Cairns, 1979, pp. 117-119, pi. 21, figs. 7, 8, pi. 22, fig. 8. Desmophyllum ingens Moseley, 1881, pp. 160-162, pi. 4, figs. 1-6, pi. 5, figs. l-4a.?Squires, 1969, p. 17, pi. 6, map 1. Desmophyllum capense; Gardiner, 1939, pp. 329, 330.?Wells, 1958, p. 262.?Cairns, 1979, p. 206. Desmophyllum capensis; Squires, 1961, p. 23, fig. 5. Description. The typical form of D. eristagalli has been adequately described elsewhere[Cairns, 1979; Zibrowius, 1980]; only a brief diagnosis is given here. Corallum variable in shape, from cylindrical to eeratoid, often greatly flared. Firmly attached by thick pedicel. Up to 80 x 50 mm in CD but averaging about 45 x 35 mm. Theca thick, covered by low, fine, rounded granules; ridged costae sometimes corresponding to S]^_3. Septa hexamerally arranged in five cycles, rarely with additional Sg. Septa thick and widely spaced, about 6-12/cm. Sj^ and 82 equal in size and exsert; septa of remaining cycles pro- gressively smaller and less exsert. Inner septal edges straight; septal faces smooth, covered by low, rounded granules. Fossa deep, endothecal dissepiments sometimes present in elongate speci- mens. Columella rare, usually absent. Forma ingens: Usually larger than typical form and often producing pseudocolonial clumps of spec- imens. Sg common, up to 192 septa. Septa usually thinner and more crowded, about 14-18/em. (Although Moseley cited only five cycles of septa in his original description of D. ingens, his il- lustrations delineate a specimen with about 136 septa, and syntypes from Challenger station 307 have up to 184 septa.) Forma capense: Similar to ingens in size and septal arrangement. Differing from it in possess- ing distinct and often deep longitudinal grooves in theca; grooves partly partitioning calice into numerous scalloped lobes. Lobes increasing perim- eter of calice, thus allowing space for more septa, up to 324 in largest specimen examined. Remarks. Squires [1965b] cited pseudocolonial D. eristagalli as the framework coral for a deep- water (334 m) coral bank on the Campbell Plateau, New Zealand. The colonial coral Goniocorella dumosa is also associated with these banks. Judg- ing by the quantity of D. eristagalli dredged off Chile, it may also form deepwater coral banks there at depths of 300-800 m. Large clumps of specimens, exhibiting four or five successive settlements, are common in the Albatross material (Plate 9, fig. 3). No associated colonial ahermatype was found, although L. prolifera, M. oculata, E. pro- funda, and S, variabilis are usually found on deepwater banks in the Atlantic. The only asso- ciated corals at the Chilean stations were Caryophyllia sp. and Javania cailleti. Approxi- mately 80% of the coralla were dead when collected. Discussion. Von Marenzeller [1904a], Hoffmeister [1933], and Zibrowius [1974a, 1980] considered D. ingens to be a junior synonym of D. eristagalli. Gardiner [1929a] considered D. ingens to be dis- tinct, and Ralph and Squires [1962] and Squires [1969] were inconclusive. D. capense has generally been acknowledged as a distinct species [Gardiner, 1904, 1939; Squires, 1961; Zibrowius, 1974a, 1980]. On the basis of a large suite of approximately 690 specimens from Albatross stations 2781 and 2785 (very near the type-locality of D. ingens) I have synonymized both D. ingens and D. capense sensu Gardiner, 1939, with D. eristagalli. Although I have examined only one syntype of D. capense Gardiner, 1904, this species is probably also a synonym of D, eristagalli. The Albatross stations contain typical specimens of D. eristagalli, forma ingens, and forma capense, as well as intermediates in size and morphology between eristagalli and Ingens and between ingens and capense. D. eris- tagalli is known to be an extremely variable species [Cairns, 1979; Zibrowius, 1980] and one with few diagnostic characters (e.g., no columella, pali, or budding). My synonymy is based on this capacity for variation and the continuous suite from the Albatross stations. Their retention as formae is an artificial separation and is retained here only to aid future revisers. Material. Forma eristagalli: Eltanin sta. 255 (2), USNM 47396; sta. 1345 (4), USNM 47406; sta. 1346 (18), USNM 47407; sta. 1403 (4), USNM 45668; sta. 1411 (5), USNM 47394; sta. 1422 (3), USNM 47395; sta. 1605 (1), USNM 47400; sta. 1691 (3), USNM 47402; sta. 1718 (5), USNM 47293; sta. 1851 (12), USNM 47412. Albatross sta. 2782 (1), USNM 36352. NZOI sta. C-734 (1), USNM 47404; sta. C-618 (3), USNM 47411; sta. D-145 (2), USNM 53381; sta. D-149 (2), USNM 47401; sta. D-159 (2), USNM 47408; sta. D-160 (1), USNM 47410; sta. D-166 (1), USNM 47403; sta. D-175 (40), USNM 47413; sta. D-176 (9), USNM 53377. Forma ingens: Eltanin sta. 214 (1), USNM 47398; sta. 25-326 (9), USNM 47405; sta. 369 (1), USNM 47399; sta. 740 (3), USNM 45669; sta. 958 (1), USNM 47397; sta. 1536 (4), USNM 47409. Vema sta. 17-39 (1), USNM 47414. WH sta. 311/66 (2), ZIZM; sta. 361/66 (1), ZIZM. Forma capense: WH sta. 361/66 (1), ZIZM. Mixture of all three formae: Albatross sta. 2781 (180), USNM 19167; sta. 2785 (510), USNM 36367. Specimens (3) of D. capense [Gardiner, 1939], BM 1939.7.20.220-223; specimens listed by Cairns [1979]. Holotype of D. eristagalli; 3 syntypes of D. ingens (BM 1880.11. 25.67); syntype of D. capense (MCZ). Types. The holotype of D. eristagalli is de- posited at the Museum National d'Histoire Naturelle, Paris, and is illustrated by Cairns 30 BIOLOGY OF THE ANTARCTIC SEAS XI Map 6. Distribution of Desmophyllum cristagalli. [1979]. Type-locality: Gulf of Gascony; depth unknown. The syntypes of D. ingens are deposited at the British Museum. Type-locality: fjords of southern Chile from 48?27'S to 52?45'S; 256-631 m. The syntypes of D. eapense are deposited at the British Museum and the Museum of Comparative Zoology (3885). Type-locality: Cape Hangklip, South Africa; 81 m. Distribution. One of the few cosmopolitan species of Scleractinia, widely distributed in the Atlantic, Pacific, and Indian oceans. In southern seas, found off southern South America, Falkland Islands, South Georgia, ? South Africa, lie Saint-Paul and lie Amsterdam, southeastern Aus- tralia, New Zealand, Auckland Island, Macquarie Island, Hjort Seamount, and several Subantarctic seamounts in South Pacific. Not present off con- tinental Antarctica. Forma ingens restricted to off southern tip of South America and off South Georgia. Forma eapense known from off southern Chile, off Falkland Islands, and off South Africa (Map 6). Worldwide depth range: 35-2460 m; Sub- antarctic records: 91-1463 m. Genus Lophelia Milne Edwards and Haime, 1849 Diagnosis. Colonial, forming large dendroid colonies by intratentacular budding. Coenosteum dense. Costae and columella poorly developed. Pali absent. Sparse tabular endothecal dissepi- ments. Type-species: Madrepora prolifera Pallas, 1766, by subsequent designation [Milne Edwards and Haime, 1850J. 19. Lophelia prolifera (Pallas, 1766) Plate 9, fig. 6 Discussion. A small branch fragment, containing only five damaged corallites, was collected at Eltanin station 1411 (USNM 47525) from a seamount on the north Macquarie Ridge just within the Sub- antarctic region. Full descriptions and synonymy CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 31 for this species can be found in the works by Cairns [1979] and Zibrowius [1980]. L. prolifera is a widely distributed species, found throughout the Atlantic Ocean, including off Tristan Island; off South Africa; off northern Madagascar; off lie Saint-Paul and lie Amsterdam; and probably off California. The Eltanin record is the southernmost report of L. prolifera and the second for the Subantarctic region, the other being Moseley's [1881] record off Tristan Island (Map 5). Worldwide bathymetric range: 60-2170 m. Subfamily PARASMILIINAE Vaughan and Wells, 1943 Genus Solenosmilia Duncan, 1873 Diagnosis. Colonial, dendroid, or subplaceloid colonies formed by intratentacular budding. Stereome granular, costae sometimes corresponding to first cycle. Tabular endothecal dissepiments. Columella small. Type-species: Solenosmilia var- iabilis Duncan, 1873, by monotypy. 20. Solenosmilia variabilis Duncan, 1873 Plate 9, figs. 4, 5 The synonymy is complete for southern records only. Solenosmilia variabilis Duncan, 1873, p. 328, pi. 42, figs. 11-18.?Moseley, 1881, p. 181, pi. 9, figs. 1-5.?von Marenzeller, 1904a, pp. 310, 311, pi. 15, figs. 4, 4a.?Hoffraeister 1933, p. 14, pi. 14, fig. 7.?Wells, 1958, p. 262.?Squires, 1969, pp. 16, 18, pi. 6, map 2.?Zibrowius, 1974a, pp. 768, 769; 1980, pp. 143-145, pi. 75, figs. A-N.?Cairns, 1979, pp. 136-138, pi. 26, figs. 2-4. Description. This species has recently been described elsewhere by Cairns [1979] and Zibrowius [1980]; a brief description follows. Colonies bushy with frequent anastomosis of branches; in- tratentacular budding. Terminal branches and calices about 6 mm in diameter. Coenosteum smooth, granular, or costate; white or grayish. Septa hexamerally arranged but very irregular in devel- opment. S4 sometimes present, but rarely as complete cycle. Septal granulation sometimes very prominent with slender granules as high as 3-4 times septal width. Tabular endothecal dissepi- ments. Columella usually absent but may be small and spongy. Discussion. Because of its distinctive branching pattern, S^. variabilis is easily distinguished from the four other colonial genera that occur in Subantarctic waters: Madrepora, Enallopsammia, Bathelia, and Goniocorella. Both ^. variabilis and M. oculata were collected at Eltanin station 1081 (east of the South Orkney Islands), which is the southernmost record for colonial Scleractihia. S. variabilis was taken in great quantity at Eltanin station 1346 (a seamount or ridge on the Heezen fracture zone of the Eltanin fracture zone system) in the South Pacific, indicating the pos- sible presence of a deepwater coral bank. Other similar deepwater structures have been found in the North Atlantic and on the Campbell Plateau, south of New Zealand. The framework corals of the northeastern Atlantic banks are L. prolifera and Madrepora oculata (?. variabilis is present to a minor extent); those in the northeast Atlantic are Enallopsammia profunda and L. prolifera (again S^. variabilis is present but not common); those on the Campbell Plateau are Goniocorella dumosa and pseudocolonial DesmophyHum cristagalli. The coral composition of the South Pacific bank is apparently about 98% S. variabilis with a small amount of M. oculata. Other associated solitary corals are Desmophyllum cristagalli, Cyathoceras irregularis, and Caryophyllia sp. Other invertebrates found at the same station include Porifera, Hydroida, Stylasterina, Gorgonacea, Actiniaria, Neraatoda, Bryozoa, Brachiopoda, Ophiuroidea, Asteroidea, Echinoidea, Holothuroidea, Pterobranchia, Poly- chaeta. Gastropoda, Polyplacophora, Bivalvia, Pycnogonida, and Crustacea. Without a sediment sample, seismic profile, and photographic documentation it is difficult to be conclusive, but in all likelihood a deepwater coral bank exists in this area. Material. Eltanin sta. 254, USNM 47423; sta. 1081, USNM 47422; sta. 1344, USNM 47424; sta. 1345, USNM 47425; sta. 1346, USNM 47426; sta, 1403, USNM 47419; sta. 1414, USNM 47420; sta. 1416, USNM 47664; sta. 1422, USNM 47421; Specimens listed by Cairns [1979], USNM. Syntypes. Types. The syntypes of S. variabilis, collected on the second cruise of the Porcupine, are depos- ited at the British Museum. Type-locality: off southwestern Spain; 1190-2003 m. Distribution. Widespread in the Atlantic and Indian oceans. Circumpolar in southern seas: off South Africa; off Prince Edward Island; off He Saint-Paul; off southeastern Australia; Hjort Sea- mount; Macquarie Ridge; off New Zealand; seamounts in South Pacific and Drake Passage; east of South Orkney Islands; off Tristan Island (Map 7). Not found off continental Antarctica. Squires's [1969] record off Chile is unsubstantiated. Worldwide depth range: 220-2165 m; Subantarctic records: 500-1830 m. Genus Goniocorella Yabe and Eguchi, 1932 Diagnosis. Colonial, extratentacular budding forming bushy, dendroid colonies. Branches anas- tomose and also joined by slender extensions of coenosteum. No columella or pali. Tabular en- dothecal dissepiments widely spaced. Type-species: Pourtalosmilia dumosa Alcock, 1902, by original designation. 21. Goniocorella dumosa (Alcock, 1902) Plate 9, figs. 7-9; Plate 10, figs. 1, 2 Pourtalosmilia dumosa Alcock, 1902, pp. 36, 37, pi. 5, figs. 33, 33a. Goniocorella dumosa; Yabe and Eguchi, 1932, pp. 389, 390; 1936, p. 167; 1941b, pp. 162, 163; 1943, pp. 495, 496, figs. 1, 2.?Squires, 1960, pp. 197, 198, pi. 33, figs. 1-4; 1964a, p. 11; 1965b, pp. 785-787; 1969, p. 17, pi. 6, map 2. ?Ralph and Squires, 1962, p. 11, pi. 4, fig. 1. ?Squires and Keyes, 1967, p. 25, pi. 3, figs. 15, 16, text fig. 4.?Eguchi, 1968, C-43, pi. C-9, figs. 11, 12.?Podoff, 1976, pp. 27, 28, pi. 1, figs. 5, 6. Description. Colony bushy, achieved by irregular extratentacular budding often at right angles to parent branch. Parent branch continuing to grow after budding; each budded branch elongating and also producing buds. Strength of colony reinforced by numerous slender (2 mm in diameter) extensions of coenosteum, these uniting adjacent branches. 32 BIOLOGY OF THE ANTARCTIC SEAS XI Plate 10. Goniocorella and Flabellum 1, 2. Goniocorella dumosa (Alcock): 1, USNM A7505, NZOI sta. D-175, x2.9, illus- trating coenosteal processes; 2, specimen from same lot, CD = 4.1 mm, illustrating dissepiment forming. 3-5. Flabellum thouarsii Milne Edwards and Haime: 3, 4, syntype, MNHNP 1028, H = 21.8 mm, GCD = 24.9 mm; 5, specimen reported by Studer [1878], Museum fiir Naturkunde, Berlin, number 1737, Gazelle sta. 54, CD = 22.7 x 15.1 mm. 6-11. Series of 3 species of Flabellum of about the same GCD: 6, 7, Flabellum thouarsii, USNM 47222, Eltanin sta. 976, GCD = 21.2 mm; 8, 9, Flabellum impensum, USNM 45629, EW sta. 9, GCD = 23.9 mm; 10, 11, Flabellum curvatum, USNM 47254, Eltanin sta. 558, GCD = 22.6 mm. CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 33 Map 7. Distribution of Solenosmilia variabilis (solid squares), Goniocorella dumosa (solid triangles), and Flabellum thouarsii (solid circles). sometimes in scalariform arrangement [Squires, I960]. Branches cylindrical, often straight, 3-5 mm in diameter, each bearing round terminal calice. Colonies up to 1 m in diameter. Corallum light gray or white; polyps and tentacles orange. Coenosteum bearing low, rounded granules. Terminal corallites often with slightly ridged Cj and C2. Septa hexamerally arranged in three cycles. S]^ very slightly exsert, with straight, vertical inner edges. Upper region of septa usually nar- rower than lower region; lower region almost reach- ing center of corallite. S2 and S3 progres- sively smaller; S3 rudimentary, with dentate inner edges. Septal faces usually smooth with fine granulation, but sometimes covered with tall, pointed granules. Fossa deep and vacuous. No columella or pali. Thin, tabular endothecal dissepiments occurring every 2-10 mm, giving dried corallum a light weight. Remarks. On the basis of a specimen attached to an underwater cable. Squires [1960] calculated the growth rate to be at least 1.67-2.94 mm/year in height. Squires [1965bJ also suggested that G. dumosa is the primary sediment-forming coral of a deepwater coral bank (coppice) on the Campbell Plateau, off New Zealand. Two other locations of Goniocorella-Desmophyllum coppices are given by Squires [1965b]. Discussion. Goniocorella is monotypic and dis- tinguished from other southern colonial corals by its distinctive branching pattern, coenosteal ex- tensions, and lack of columella. Material. Eltanin sta. 1816, USNM 47509; sta. 1848, USNM 47667. NZOI sta. A-706, USNM 47511 sta. B-319, USNM 47506; sta. C-410, USNM 47510 sta. C-618, USNM 47507; sta. C-633, USNM 47508 34 BIOLOGY OF THE ANTARCTIC SEAS XI sta. D-145, USNM 53382; sta. D-175, USNM 47505. Some of these records first reported by Ralph and Squires [1960], Squires U965b], and Squires and Keyes [1967]. Types. The syntypes of G. dumosa are at the Indian Museum (Calcutta). Type-locality: Banda Sea; 469-487 m. Distribution. Off Japan; Banda Sea; off Norfolk Island; off Bounty Islands; off New Zealand; Chatham Rise; Campbell Plateau (Map 7). Depth range: 100-638 m. Superfamily FLABELLICAE Bourne, 1905 Family FLABELLIDAE Bourne, 1905 Genus Flabellum Lesson, 1831 Diagnosis. Solitary, cuneiform to compressed turbinate, free. Wall epithecal. Base not thick- ened by stereome; no roots. Calicular edge jagged or entire. Pali absent. Columella rudimentary or absent. Type-species: Flabellum pavoninum Lesson, 1831, by subsequent designation [Milne Edwards and Haime, 1850]. 22. Flabellum thouarsii Milne Edwards and Haime, 1848 Plate 10, figs. 3-7 Flabellum thouarsii Milne Edwards and Haime, 1848, p. 265, pi. 8, fig. 5.?Studer, 1878, p. 630.?Not F. thouarsii; Gravier, 1914b, pp. 125-128 (is F. impensum Squires, 1962).?Not F. thouarsii; Wells, 1958, p. 268 (is F. impensum Squires, 1962, and F. flexuosum, n. sp.?Squires, 1961, pp. 29-38, figs"; 5^ 14-19, 21, 23, 27; 1962b, pp. 14, 18; 1969, p. 18, pi. 6, map 4.?Keller, 1974, pp. 200-203, pi. 1, figs. 1-4, pi. 2, figs. 1-7, pi. 3, figs. 1-4, pi. 4, figs. 1-7.?Cairns, 1979, p. 206. Flabellum thouarsi; Milne Edwards and Haime, 1857, pp. 89, 90. Flabellum curvatu-m; Gardiner, 1939, pp. 327, 328, pi. 20, figs. 1, 2 (part: all stations but Dis- covery sta. 182, WS sta. 839).?Squires, 1961, p. 38 (part: Vema sta. 14-18); 1962a, pp. 1-11, figs. 1-3. Description. Corallum ceratoid to flabellate, attached only in young stage. Pedicel short, about 2.5-3.2 ram in diameter and usually worn away in older specimens. Pedicel expanding into straight corallum with angle of lateral edges varying from 30? (ceratoid) to 90? (flabellate); inclination of lateral faces rarely exceeding 20?. Largest spec- imen examined 33 x 22 mm in CD and 34 mm tall; an average-size specimen, however, 21 x 14 ram in CD and 23 mm tall. Theca usually worn or encrusted with calcareous invertebrates and ploychaete sand tubes. Calice elliptical, with GCD/LCD ratios of 1.3-1.7; calicular profile arched. Septa hexamerally arranged in five cycles; how- ever, only largest specimens with 96 septa. Most specimens with 20-22 major septa (81-3) enclosing triads of septa, or 80-88 total septa. Ten or eleven Sj 2 equal in size and reaching center of fossa, sometimes fusing with those on opposite side. Ten or eleven S3 smaller, 20-22 S,;, one third to one fourth of size of S3, and S5 ru- dimentary. Septa not exsert; each larger septum having shallow, concave notch near calicular edge, but never dentate. Inner edges of larger septa straight to slightly sinuous and thickened deeper in fossa. Septal granulation coarse and irregular, granules sometimes arranged in rows parallel to septal edge. Fossa relatively shallow. Rudimentary columella formed by fusion of thickened inner edges of Sj^ and S2 and additional irregular extensions from these septa. Internal stereome present in older specimens. Remarks. Judging from substrate of attachment and attached worm tubes, this species seems to inhabit bottoms composed of small pebbles and coarse sand. Squires [1962a] discusses planula- tion, size of larva, and lack of periodicity for this species. Specimens from the northern range seem to attain a larger size and are therefore more easily confused with F. curvatum. Discussion. As Squires [1961, p. 31] implied, Gardiner [1939] was probably not aware of F. thouarsii; most of his records of F. curvatum are average- to large-size specimens of F. thouarsii from its characteristic depth range, and there is one record each of F. impensum and F. curvatum. Authors that have relied on Gardiner's identifica- tions [e.g., Wells, 1958; Keller, 1974] have been misled. Although Squires [1961] differentiated F. thouarsii from F. curvatum, he included one suite of uncharacteristically large specimens of F. thouarsii as F. curvatum (Vema station 14-18). Furthermore, his paper on the larvae of F. curvatum [Squires, 1962a] is also based on large F. thouar- sii. Keller [1974] recognized two forms of F. thouarsii, one similar to Squires's [1961] F. thouarsii and the other similar to his F. curvatum. She did not agree with Squires's [1961] reasons for separating the two species and therefore synonymized them. Keller's specimens all seem to be typical F. thouarsii. Ironically, her next species account, which describes F. antarcticum, is typical F_. curvatum. Comparisons of F. thouarsii to the closely re- lated F. curvatum and F. areum are discussed with those species. Material. Eltanin sta. 217 (1), USNM 45650; sta. 370 (22), USNM 45664; sta. 974 (232), USNM 45649; sta. 976 (184), USNM 47222; sta. 977 (309), USNM 45648. Edisto sta. 49 (1), USNM 45643; sta. 50 (1), USNM 47218. Vema sta. 14-6 (10), AMNH; sta. 14-16 (18), USNM 45654, and (13), AMNH; sta. 14-18 (1), USNM 45655; sta. 15-PD3 (6), AMNH; sta. 15-99 (14), USNM 47212, and (1), AMNH; sta. 15-102 (135), USNM 47220; sta. 15-103 (2), USNM 45651, and (3), AMNH; sta. 15-108 (6), USNM 47216, and (6), AMNH; sta. 15-110 (15), USNM 47221, and (19), AMNH; sta. 16-39 (8), USNM 47214, and (100), AMNH; sta. 17-74 (19), USNM 45617, and (8), AMNH; sta. 17-76 (40), USNM 47211, and (81), AMNH; sta. 17-88 (10), USNM 45618, and (10), AMHH; sta. 17-89 (13), USNM 47217, and (21), AMNH; sta. 17-90 (1), AMNH; sta. 17-97 (100), AMNH; sta. 18-13 (32), USNM 47219; sta. 18-14 (3), USNM 47215; sta. 18-16 (4), USNM 47213. BR sta. 25149 (2), USNM 47210. Spec- imens (2) of Studer [1878], Museum fiir Naturkunde, Berlin (number 1737); some Vema records first reported by Squires [1961, 1962a, b] as F. thouar- sii and F. curvatum; some specimens identified as F. curvatum by Gardiner [1939): WS sta. 76 (1), BM 1939.7.20.169; sta. 216 (19), BM 1939.7.20.252- 264; sta. 244 (11), BM 1939.7.20.154-164; sta. 247 (1), BM 1939.7.20.170; sta. 792 (1), BM 1939.7.20.177 and Discovery sta. 652 (7), MCZ 3589. Syntypes. CAIRNS: ANTARCTIC AND SOBANTARCTIC SCLERACTINIA 35 Types. Milne Edwards and Haime's original de- scription gave measurements for only one specimen, implying a holotype. However, at the Museum Na- tional d'Histoire Naturelle, Paris, there are four specimens labeled F. thouarsi from the 'lies Ma- louines' in the Milne Edwards Collection, two num- bered 1028 and two numbered 1029. Three are F. thouarsii; a fourth from lot 1029 appears to be an Indo-Pacific hermatype of similar shape. One of the specimens from lot 1028 has measurements simi- lar to those of the specimen in the original de- scription and may be the holotype. Gravier [1914b, pp. 127, 128] reported two types (syntypes ?) of F. thouarsii at the Museum National d'Histoire Naturelle and two additional specimens identified as this species, one from 'lies Malouines' and the other 'trouv^ dans une eponge. ' This may explain the presence of four identified specimens of F. thouarsii but does not help in determining the type. Because of this uncertainty a lectotype is not chosen. Type-locality: Malouine Islands (Falkland Islands). Distribution. Off southeastern South America from Rio de la Plata, Uruguay, to Cape Horn; off Falkand Islands. Squires's [1969] record from the Scotia Ridge undocumented (Map 7). Depth range: 71-305 m. Remarks The maximum size of this species prob- between a GCD of 25-30 mm, judging from 23. Flabellum areum n. sp. Plate 11, figs. 1-5 ably lies the extreme development of stereome and worn pedi- cels of specimens in this size range. Discussion. F. areum is most similar to F. thouarsii, particularly in size, septal arrange- ment, and geographic distribution. It can be dis- tinguished by its deeper fossa, larger pedicel diameter, lesser number of septa per centimeter (about 12 for F. areum, 18 for F. thouarsii), and much deeper bathymetric range. Etymology. The specific name areum (Latin: open or vacant space) refers to the spacious fossa. Material. Vema sta. 15-132 (25), AMNH; sta. (3) (15), AMNH: sta. 17-57 Types. Types. The holotype 17-61 , USNM 47913. collected at Eltanin sta- Description. Corallum trochoid to turbinate, sometimes campanulate; usually straight. Weakly attached by short pedicel 3.2-5.1 mm in diameter; pedicel often detached from substrate (usually a small pebble), allowing base and pedicel to erode gradually. Largest specimen 27.7 x 20.0 mm in CD and 21.2 mm tall. Theca usually worn, bearing thin, longitudinal striae corresponding to all septa, characteristic of most flabellids. Calice elliptical, with GCD/LCD ratio usually between 1.3 and 1.4; lateral edges rounded. Calicular margin entire, calicular profile straight to slightly arched. Septa hexamerally arranged in five cycles; how- ever, fifth cycle never complete. One large spec- imen with 86 septa; holotype (adult specimen) with 80 septa. Sj^ and S2 equal in size and meeting in center of calice. Sj^ and S2 not exsert and forming near calicular edge shallow, concave notch; notch may or may not be dentate. Upper thecal edge forming lip rising slightly above upper septal insertions. Size of S3 depending on presence of paired S5 in half system. If S5 present (which is more common in half systems adjacent to lateral edges), then S3 one half to three fourths of size of $1 2' when S5 absent, S3 about one third of size of S]^ 2" ^4 ^"'^ ^5 Pi^ogres- sively smaller, S4 extending only about halfway to base. Inner edges of septa usually straight but may be slightly sinuous. Septal granules low to moderately tall and arranged in lines parallel, or rows almost perpendicular, to septal edge, the lat- ter corresponding to fine sinuosity of septal edge. Fossa appearing spacious because of widely spaced septa and campanulate corallum shape. Lower, inner edges of S^ and S2 greatly thickened and fused in center, forming rudimentary columella. Theca within calice often increased substantially by deposits of stereome, sometimes obscuring smaller septa. Stereome also filling in base of more elon- gate coraiia. tion 1592, is deposited at the United States Na- tional Museum (47167). Three paratypes from Elta- nin station 1592 (number 47168), 14 from Eltanin station 973 (number 45639), 8 from Vema station 17-57 (number 47169), and 15 from Vema station 15-132 (number 45616) are deposited at the United States National Museum. Type-locality: 54?43'S, 55?30'W (Scotia Ridge east of Burdwood Bank); 1647- 2044 m. Distribution. Off Mar del Plata, Argentina; off Cape Horn, Tierra del Fuego; Scotia Ridge southeast of Falkland Islands (Map 8). Unconfirmed depth range: 1647-2229 m. 24. Flabellum curvatum Moseley, 1881 Plate 10, figs. 10, 11; Plate 11, figs. 6-9 Flabellum curvatum Moseley, 1881, pp. 174, 175, pi. 6, figs. 3a-3d.?Gardiner, 1939, pp. 327, 328 (part: WS sta. 839 only) .?Squires, 1961 (part: not Vema sta. 14-18), pp. 7, 9, 29, 38, 39, figs. 5, 11-13, 20, 22, 30.?Not F. curva- tum; Squires, 1962a, pp. 1-11, figs. 1-3 (is F. thouarsii Milne Edwards and Haine, 1848).?Squi- res, 1962b, p. 14; 1964a, p. 13; 1964c, pi. 1, fig. 1; 1969, p. 18, pi. 6, map 3.?Cairns, 1979, p. 206. Flabellum antarcticum; Keller, 1974, pp. 203-205, pi. 5, figs. 1-7. Description. Corallum ceratoid to trochoid, rarely attached above GCD of 12 mm. Pedicel long, slender (2.5-3.3 mm in diameter), and usually bent; base of pedicel often worn to a point in older specimens. Corallum usually curved, enlarging gradually from pedicel to calice. Largest specimen examined 44 x 30 mm in CD and 47 mm tall. Theca usually worn or encrusted with bryozoans, serpu- lids, or other corals; sometimes thin, incised cos- tal lines present, one corresponding to each sep- tum. Calice elliptical, with GCD/LCD ratios of 1.4-1.6; calicular profile arched. Septa hexamerally arranged in five cycles. S]^ and S2 equal in size and usually slightly larger than S3. S4 about half size of S3; S5 rudimentary and sometimes fenestrate because of weakly calcified trabeculae. Septa not exsert; upper edge of each larger septum forming shallow, concave notch near calicular edge. This notch often dentate but not always. Inner edges of lar- ger septa straight to slightly sinuous and thick- ened lower in fossa. This thickening, along with loose fusion of irregular processes from lower inner edges of larger septa, forming rudimentary columella. Septal granulation variable, ranging 36 BIOLOGY OF THE ANTARCTIC SEAS XI Plate 11. Flabellum 1-5. Flabellum areum n. sp.: 1-3, holotype, USNM 47167, Eltanin sta. 1592, H ? 21.9 mm, GCD = 25.8 nun; 4, paratype, USNM 45639, Eltanin sta. 973, H = 32.5 mm, corallura broken, revealing columella; 5, specimen (paratype) from same lot, H = 21.7 ram. Flabellum curvatum Moseley: 6, 7, syntype, BM 1974.8.510, Challenger sta. 320, GCD = 40.7 mm; 8, syntype, BM 1880.11.25.85, Challenger sta. 320, GCD = 29.7 mm; 9, USNM 47253, Eltanin sta. 340, x3.2, illustrating dentate septal notch. Flabellum impensum Squires: USNM 45666, EW sta. 37, H = 65.1 ram. 6-9. 10. CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 37 Map 8. Distribution of Flabellum impensum (solid circles), and Flabellum areum (solid squares). from low and rounded to high, slender, pointed granules, usually arranged in rows subparallel to septal edges. Fossa elongate and relatively shal- low. Stereome present in elongate specimens. Remarks. F^. curvatum usually settles on pebbles 3-4 times the diameter of its original attachment. Its bent pedicel and curved corallum probably re- flect a reorientation of the polyp after it de- taches from its substrate or when it becomes so heavy that it topples sideways. It also attaches to coralla of its own species, echinoid spines, and branching bryozoans. In turn, the theca of the living coral provides a substrate for numerous species of Bryozoa, serpulid polychaetes, barn- acles, hydrocorals, and other scleractinians. JF. curvatum probably occurs in fairly high den- sity off East Falkland island, indicated by the recovery of over 2500 specimens from a 68-min trawl. Discussion. As is indicated by the synonymy, F. curvatum has often been confused with F. thouarsiT, a closely related species. In fact7 both Wells [1958] and Keller [1974] have synonymized these species. After thorough reexamination of this species complex I find that F. curvatum can be distinguished by a combination of the following characters: (1) the pedicel is usually bent and the corallum is usually curved; (2) the pedicel is longer; (3) the maximum size of the corallum is larger; (4) the septal notch is sometimes dentate; (5) the S4 are relatively larger than those of F. thouarsii; and (6) the fossa is usually shal- lower, sometimes partially occupied by a crispate columella. Characteristics of attachment and ped- icel diameter mentioned by Squires [1961] are of no diagnostic value. Furthermore, although their depth ranges overlap, F. curvatum is usually found deeper than F. thouarsii. 38 BIOLOGY OF THE ANTARCTIC SEAS XI Map 9. Distribution of Flabellum curvatum (solid circles), Flabellum knoxi (solid squares), and Flabellum gardineri (solid triangle). F. curvatum is distinguished from F. impensum by its coarser septal granulation, smaller PD, usually curved corallum, shallower fossa, and geographical distribution. Material. Eltanin sta. 339 (73), USNM 47238; sta. 340 (139), USNM 47253; sta. 556 (8).- USNM 47243; sta. 558 (about 2500), USNM 47254; sta. 740 (6), USNM 47239; sta. 1536 (11), USNM 47242. Hero sta. 715-875 (126), USNM 47251; sta. 715-885 (1), USNM 47245; sta. 715-895 (35), USNM 47252. Vema sta. 14-12 (4), USNM 45652, and (8), AMNH; sta. 15-PD3 (5), USNM 47240; sta. 15-PD4 (39), USNM 47250; sta. 15-PD9 (3), USNM 47241; sta. 15-PDlO (5), USNM 45626, and (11), AMNH; sta. 17-59 (2), USNM 45653; sta. 17-100 (24), USNM 45621, and (37), AMNH; sta. 17-101 (9), USNM 45624, and (32), AMNH; sta. 18-8 (7), USNM 45620; sta.. 18-12 (16), USNM 45622. Calypso sta. 171 (2), USNM 47246; sta. 172 (4), SME. BR sta.. 25149 (5), USNM 47244. Specimens of Squires [1961, 1962a], USNM; specimens of Gardiner [1939] from WS sta. 839, BM 1939.7.20.129. Two syn- types (BM 1880.11.25.85 and 1974.8.5.10). Types. The eight syntypes of F. curvatum are deposited at the British Museum. Type-locality: 37?17'S, 53?52'W (off Rio de la Plata, Uruguay); 1097 m. Distribution. Off southeastern South America from Rio de la Plata, Uruguay, to Cape Horn; Burd- wood Bank; off Falkland Islands; off South Georgia (Map 9). Depth range: 115-1137 m; however, most common between 400 and 800 m. 25. Flabellum impensum Squires, 1962 Plate 10, figs. 8, 9; Plate 11, fig. 10; Plate 12, figs. 1-8 Flabellum inconstans; Pax, 1910, pp. 66-72, pi. 11, figs. 3-9, pi. 12, figs. 1-6. Flabellum thouarsii; Gravier, 1914b, pp. 125-128, pi. 1, figs. 5, 6.?Wells, 1958, p. 268 (part : CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 39 116.0 mm Eltanin sta Plate 12. Flabellum 1-8. Flabellum impensum Squires: 1, 2, USNM 45629, EW sta. 9. H = 78.5 mm, GOD = 3, USNM 47356, Eltanin sta. 1878, H = 10.1 mm; 4, USNM'47368, 1084, x0.51, muricid gastropod egg cases; 5, USNM 45637, ^'?'^^"^" ?'^- *37, xO.92. volutid gastropod egg case; 6, USNM 45635, Eltanin sta. 426, xl.7, broken corallum revealing columella; 7, USNM 45666, EW sta. 37, xl.9, dentate and fenestrate septal edges; 8, specimen identified as F. transversale by Thomson and Rennet [1931], Australian Museum G 13538 sta 10, H = 43.8 mm. 9-12. Flabellum flexuosum n. sp.: 9, USNM 47265. Eltanin sta. 418, H = 36.0 mm- 10-12, holotype, USNM 47170, Eltanin sta. 1536, GCD = 20.8 mm, H = 28.7 mm.' 40 BIOLOGY OF THE ANTARCTIC SEAS XI Discovery sta. 34, 39), pi. 2, figs. 5, 6. Flabellum transversale: Thomson and Rennet, 1931, p. 41. Flabellum harmeri; Gardiner, 1939, pp. 326, 327. Flabellum curvatum; Gardiner, 1939, pp. 327, 328 (part: Discovery sta. 182). Frabellum marmeri; Niino, 1958, p. 257, pi. 2, fig. 1 (misspellings). Flabellum sp. Squires, 1961, pp. 29, 38. Flabellum impensum Squires, 1962b, pp. 14, ~~^. 27 figs. 4-7, text fig. 3; 1969, filter feeders (usually Bryozoa) on both sides of a specimen was interpreted as an indication of an upright position, filter feeders on one side and borers on the other as a prone position, and ab- sence of organisms from one or both sides as inconclusive. In an examination of 96 specimens which were alive when they were collected, 71 were found to be inconclusive, 24 indicated an erect and 1 indicated a prone posture. The posture, 17-19, p. 18 31-38, implication is that after F. imge nsum becomes free of its original attachment, it maintains itself in an upright position, perhaps by submergence of the pedicel in the substrate. Consistent absence of filter feeders from near the base and of erosion of the base supports this theory. Two gastropods, believed to be of the families Volutidae and Muricidae (J. Houbrick, personal communication, 1979), have deposited large egg cases near the calicular edges of living specimens Description. CoraUum variable in shape, ranging (Plate 12, figs. 4 and 5). Bryozoans and serpulids (part), pi. 6, map 3.?Podoff, 1976, pp pi. 2, figs. 9, 10.~Sorauf and Podoff, 1977, p. 4 (part: pi. 1, figs. 1, 2, not pi. 3, figs. 1, 2)?Cairns, 1979. p. 206. Flabellum distinctum; Eguchi, 1965, pp.. 10, 11, text fig. 3. also commonly encrust the theca. Pax [1910J describes and illustrates some his- tology of a tentacle, including nematocysts. Discussion. The extreme forms of F. impensum might easily be mistaken for separate species: one a large flabellate corallum with almost seven cycles of septa, the other a ceratoid to trochoid corallum with less than six cycles of septa. Fur- thermore, each form usually occurs homogeneously when it is collected. However, several suites show a continuous gradation of corallum shape. No other morphological differences are apparent. from flabellate to almost conical (ceratoid to tro- choid). Largest flabellate corallum 128 x 45 mm in CD and 80.2 mm tall (Plate 12, fig. 4). Ceratoid to trochoid coralla usually smaller, with GCD/LCD ratio as low as 1.1. Angle of lateral edges of flabellate coralla often about 40?-50? for first 10 mm, then lateral edges flaring out- ward up to 150?, maintaining constant inclination of lateral faces of 30?-40?. In ceratoid to trochoid coralla, angle of lateral edges remaining constant with growth. All intergrades occurring between extreme flabellate and ceratoid shapes. Round pedicel never reinforced by external stereome Even the type-specimens show this corallum varia- and 3.5-6.0 mm in diameter. Small coralla usually tion: the holotype is flabellate and the il attached to pebbles or gastropod shells; when over lustrated paratype is ceratoid. Finally, there 30 mm tall, coralla usually becoming free of orig- are no geographic or bathymetric differences^ ex- slightly carinate. Calicular profiles of flabel- late coralla strongly arched; those of trochoid coralla more even. Septa hexamerally arranged in smaller coralla u). to 96 septa stage. Additional septa irregularly added in groups of three, up to at least 300 septa (Sy). Full fourth cycle achieved at GCD of about type or current pattern. Early records of F. impensum were usually as- signed to more northern temperate species. F. inconstans Marenzeller, 1904, reported by Pax [1910], IS a South African species with a basal fracture. F. thouarsii Milne Edwards and Haime, 1848, is a distinct species known from relatively sizes may be either S^ = S2 - S3 - S4 > S5 > S6 or Si = S2 = S3 2 S4 > S5 > S5. Septa not exsert and very thin. Larger septa usually sloping concavely away from calicular edge, producing shallow notch, this notch usually finely dentate. Toward center of calice, septum forming shoulder before dropping vertically into fossa. Inner edges of larger septa slightly sin- uous and thickened; deep within fossa, lower inner edges fused into rudimentary columella. Septal granules fine and pointed, often arranged in widely spaced lines parallel to septal edge. Fossa very deep, elongate in flabellate coralla. Small amounts of internal stereome sometimes pres- ent in base. Remarks. An analysis of the living orientation of F. impensum similar to that of Squires [1964a] for~other flabellids was made. The presence of species. Gardiner's [1939] single Antarctic record of F. curvatum is F. impensum. >ta'terial. Eltanin sta. 272 (6), USNM 45636; sta. sta sta sta sta 410 (4), USNM 45634; sta. 426 (3), USNM 45635; 428 (5), USNM 45632; sta. 437 (5), USNM 45637; 444 (1), USNM 47330; sta. 499 (1), USNM 47347; 992 (7), USNM 47335; sta. 993 (2), USNM 47323; 997 (10), USNM 47341; sta. 1002 (1), USNM 47327; sta. 1079 (7), USNM 45640; sta. 1083 (9), USNM 47331; sta. 1084 (3), USNM 47368; sta. 1089 (1), USNM 47542; sta. 1870 (8), USNM 47346; sta. 1871 (33), USNM 47361, and (1), MCZ; sta. 1878 (12), USNM 47356; sta. 1880 (6), USNM 47349; sta. 1885 (2), USNM 47344; sta. 1898 (1), USNM 47367; sta. 1916 (1), USNM 47362; sta. 1922 (1), USNM 47545; sta. 1930 (2), USNM 47339; sta. 1933 (3), USNM 47355; sta. 1996 (1), USNM 47343; sta. 2005 (2), USNM 47371; sta. 2006 (1), MCZ; sta. CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 41 U.?i^ 'lh"''^..V'l?: -li^>- ^''?' -- 2018 (3). USNM 47375, and (1), 47369; sta. 2025 (2),' USNM 47372; sta. 2045 (4), USNM 47353; sta. 2068 (1), USNM 47351; sta. 2085 (2), USNM MCZ; sta. 2021 (1), uSNM USNM 47357; sta. 2031 (1) (3), USNM 47354; sta. 2063 2065 (2), USNM 47352; sta. sta. 2070 (1), USNM 47540; 47348; sta. 2088 (4), USNM 47350; sta. 2097 (1), uSNM 47342; sta. 2099 (6) USNM 47358; sta. 2115 (1). USNM 47365; sta. 2117 (9). USNM 47374; sta. 2124 (9). USNM 47376; sta. 2143 (9). USNM 47373; sta. 5761 (1), USNM 47366- sta. 5765 (4), USNM 47326. Islas Orcadas sta.' 575-53 (11), USNM 47383; sta. 575-65 (9), USNM 47336; sta. 575-66 (4), USNM 47379; sta. '575-67 (1), USNM 47380; sta. 575-70 (2). USNM 47384; sta. 876-107 (3), USNM 47334; sta. 876-108 (6), USNM 47378; sta. 876-110 (2), USNM 47332; sta. 876-113 (2), USNM 47381; sta. 876-114 (3), USNM 47337; 876-126 (4), USNM 47382. Hero sta. 702-465 (1), USNM 47324; 876-124 (1), 47340; sta. 691-20 (15), 47543; sta. 721-1102 (1), USNM USNM 47363; sta. sta. 16 (1), Atka sta. 23 sta. USNM sta. USNM sta. 876-118 (6), USNM 47329; sta. 876-127 (1), USNM 47333; 721-1084 (1), USNM 47325; st. 47390; sta. 721-1110 (20), 731-1842 (1), USNM 47328. Edisto USNM 47391; sta. 28 (2), USNM 45641. (2), USNM 47345. EW sta. 9 (86), USNM"~45629; sta. 16 (1), USNM 47385; sta. 23 (73). USNM 45627; sta, 28 (3). USNM 45644; sta. 32 (1), USNM 47387; sta. 35 (1), USNM 47386; sta. 36 (2). USNM 45667; sta. 37 (8), USNM 45666; sta. 38 (13), USNM 45630; sta. 39 (4), USNM 45628. Westwind sta. 4 (1), USNM 47389. Vema sta. 18-32 (2), USNM 47338. NZOI sta. A-537 (1), USNM 47364; sta. A-625 (20), USNM 47392. Specimen identified as F. transversale from station 10 (Thomson and Rennet, 1931J, Australian Museum G 13538; specimen identified as F. harmeri [Gardiner, 1939], BM 1939.7.20.128; specimens (2) identified as F. curvatum from station 182 [Gardiner, 1939) BM 1939.7.20.271-272; specimens identified as F! thouarsii from Wells (1958) from stations 34 (2) and 39 (2), South Australian Museum H 63, H 65. Tjf?e?. The holotype and 61 paratypes of F. im- pensum are deposited at the New Zealand Oceano- graphic Institute. Type-locality: 73?20'S, 174?00'E (Ross Sea); 369-384 m. Distribution. Circumpolar continental Antarctic, including off South Shetland Islands, South Orkney Islands, and South Sandwich Islands, and one dis- junct record off the Antipodes Islands (Map 8). Depth range: 46-2260 m; however, temperature range probably very slight. Most records between 100 and 1000 m; one of deepest records (2010 m) at northernmost range (Eltanin sta. 2143, off Antip- odes Islands). 26. Plabellum flexuosum n. sp. Plate 12, figs. 9-12 Desmophyllum sp. Marenzeller, 1903, p. 1. Desmophyllum antarcticum; Gravier, 1914b, p. 122 (part: pi. 1, fig. 4). Gardineria lilliei; Gardiner, 1939, pp. 328, 329 (part: Discovery sta. 140, 160 (part), 181. 190). Flabellum antarcticum; Wells, 1958, p. 269, pi. 2, figs. 11-15.?Squires, 1962b, pp. 13, 14, 19^ 20; 1969, p. 18, pi. 6, map 3.?Bullivant, 1967, p. 65.~Zibrowius, 1974b, p. 18.?Not F. antarc- ticum; Keller, 1974, p. 203 (is F. curvatum Moseley, 1881). ~ Flabellum thouarsii: Wells, 1958, p. 268 (part- Discovery sta. 41, 93), pi. 2, figs. 7-10. Flabellum ongulense Eguchi, 1965, pp. 11, 12 pi 2, figs. 2a-2d. ' Description. Corallum ceratoid, tall; straight, bent, curved, or scolecoid. Pedicel 2.7-4.5 mm in diameter, expanding slightly (up to 5.5 mm) at attachment to substrate. Coralla usually remaining attached. Holotype 20.8 x 17.7 mm in CD and 28.7 mm tall: PD at break 3.0 mm. Largest specimen lEltanin station 1933, USNM 47172) 24.0 x 20.7 mm in CD, 4.0 mm in PD, and 67 mm tall. Theca very thm and porcelaneous, usually without encrusting organisms; however, bryozoans sometimes colonizing theca of living specimens. Calice elliptical, not compressed; ratio of GCD/LCD about 1.25. Septa hexamerally arranged in five cycles; how- ever, tall, slender coralla and younger specimens often with less septa. S^ and So equal in size and slightly exsert (because the septa are so delicate, their upper septal edges are invariably broken off when they are collected). Remaining septa! cycles progressively smaller, S3 sometimes 3-4 times larger than S^; S5 rudimentary. Inner edges of all septa sinuous, corresponding to shallow, transverse undulations on septa, producing wrinkled or corrugated appearance. Fine, pointed septal granules, up to 2 times septal thickness in height, aligned on crests of soptal undulations. Lower inner edges of Si and S2 usually fused, forming rudimentary columella. Discussion. Both Wells [1958] and Squires [1962b, 1969] identified this species as Flabellum antarcticum (Gravier, 1914). Gravier's (1914a] species, although it is very similar to F. flexuo- sum, is a rarely collected Javania, which always has a thick stereome-reinforced pedicel; F. flexuosum always has a typical nonreinforced Flabellum-type pedicel. However, Gravier's [1914b, pi. 1, fig. 4] third specimen from dredge VIII, which he doubtfully assigned to Desmophyllum ant- arcticum, is probably F. flexuosum. F. flexuosum has basically the same distribution as F. impensum, the only other circum-Antarctic species of Flabellum. It can be distinguished by its more slender, often bent corallum, exsert septa, and more prominent septal granulation. The single specimen from the Weddell Sea (Edisto station 20) and Wells's [1958] specimens from off eastern Antarctica differ from typical F. flexuosum in that they are large specimens and yet have only four cycles of septa. It would be helpful to have more specimens from the eastern Antarctic for com- parison. Flabellum ongulense Eguchi, 1965, may be the same species and would therefore have nomenclatural priority. However, none of Eguchi's Antarctic specimens are available for study. F. gracile (Studer. 1978), known only from off N"^W Zealand (95-196 m), is also very similar to F. flexuosum. Thomson and Rennet's [1931] Caryophyllia vermifo7- mis seems to be ?. gracile. Etymology. The specific name flexuosum (Latin: full of bends, crooked) refers to the bent and often scolecoid shape of the corallum. Material. Eltanin sta. 418 (1), USNM 47265; sta. 671 (42), USNM 47275; sta. 684 (5), USNM 45656; sta. 993 (2), USNM 53427; sta. 1535 (7), USNM 47270; sta. 1870 (3), USNM 47259; sta. 1878 (1), USNM 47261; sta. 1995 (22), USNM 47276; sta. 42 BIOLOGY OF THE ANTARCTIC SEAS XI 1996 (26), USNM 47282; sta. 1997 (10), USNM 47279; sta 2092 (1). USNM 47268; sta. 2097 (1). USNM 47258- sta. 2119 (1), USNM 47257; sta. 2120 (1), USNM 47256. Islas Orcadas sta. 575-8 (D. "SNM 47277- sta. 575-12 (4), USNM 47283; sta. 575-13 (1), USNM 47273; sta. 575-17 (3), USNM 47267; sta. 575-30 (1), USNM 47262; sta. 575-52 (1), USNM 47273; sta. 575-90 (2), USNM 47281; sta. 575-91 (6), USNM 47271. Hero sta. 691-20 (1), USNM 47280; sta 731-1812 (1), USNM 47266. Edlato sta. 20 (2)' USNM 47272. Atka sta. 23 (12), USNM 47269. Burton Island sta. ~3~r3), USNM 47264. EW sta. 6 12), USNM 47255; sta. 28 (3), USNM 47274; sta. 35 (1), USNM 47263. At bb'hO'S, 139?51'E, 220-240 m (1), SME. Specimen identified as Desmophyllum sp. by Marenzeller [1903], Brussels Museum; specimen identified as Caryophyllia vermiformis by Thomson and Rennet [1931], Australian Museum G 13535; specimens (5) identified as Gardineria UUiei by Gardiner [1939], BM 1939.7.20.238-240, and (1), MCZ- specimens identified as F. thouarsii from stations 41 (5) and 93 (3) [Wells, 1958], South Australian Museum H 66, H 69. Types. Types. The holotype, collected at Eltanin sta- tion 1536, is deposited at the United States Na- tional Museum (47170). Nine paratypes from Eltanin station 1536 (number 47171), 1 from Eltanin station 1933 (number 47172), and 21 from Islas Orcadas station 575-93 (number 47173) are deposited Ji. Ehi United States National Museum. Type- locality: 54?29'S, 39?22'W (west of South Georgia); 659-686 m. Distribution. Off Antarctic Peninsula; off South Shetland Islands; off South Orkney Islands; off South Sandwich Islands; off South Georgia and Shag Rocks; Weddell Sea; off Enderby Land; Ross Sea; Bellingshausen Sea (probably circumpolar) (Map 10). Depth range: 101-659 m. neither of which is Gardineria: Flabellum flexuosum (Discovery stations 140, 160 [part), 181, and 190); Flabellum gardineri (Discovery station 160 (part)); Flabellum sp. (Discovery station 160 (part)); and Caryophyllia eltaninae (Discovery station 160 (part)). The 2 specimens from Discovery station 160 referred to above as Flabellum sp. differ from Flabellum gardineri in having a wider pedicel, a more open calice, and a fifth cycle of septa; otherwise they are very similar and may represent a growth form. Flabellum sp. is also represented by Gardiner's [ 1929aJ record of G. antarctica from Discovery station 152. More specimens are required before this problem can be solved. Flabellum gardineri most closely resembles Flabellum flexuosum but can be distinguished by its fewer septa, more massive columella, straight inner septal edges, noncorrugated septa, and smal- ler size. Etymology. This species is named in honor of J. S. Gardiner, who contributed greatly to our know- ledge of Scleractinia, including corals of the Subantarctic region. Material. Discovery sta. 160 (4), MCZ-3574. Types. Types. The holotype, collected at Discovery station 160, is deposited at the British Museum (1939.7.20.305). Thirty-three paratypes from Dis- covery station 160 (1939.7.20.288-304, 306-314) are deposited at the British Museum. Two specimens from this lot have been permanently deposited at the United States National Museum (48300). Type- locality: 53?43'40"S, 40?57'00"W (off Shag Rocks); 17 7 m. Distribution. Known only from type-locality 27. Flabellum gardineri n. Plate 13, figs. 1-3 sp. Gardineria lilliei; Gardiner, 1939, pp. 328, 329 (part: 40 specimens from Discovery sta. 160). Description. Corallum ceratoid, straight, elongate. Pedicel diameter about 2.5 mm, expanding slightly at attachment to substrate. Holotype 8.9 X 8.3 mm in CD and 22.1 mm tall. Tallest specimen 30.5 mm. Theca dull white with thin, incised ver- tical striae, one corresponding to each septum. Calice round to elliptical. Septa hexamerally arranged in four cycles. S^ and S2 equal in size and 3-4 times larger than S3 and Sl^, these about equal in size. Septa not exsert; larger septa bearing nondentate, shal- low notch near calicular edge. Inner septal edges of Si and S2 straight and entire, fusing into a solid columella about one third to one half of distance to base. S3 and S4 very low in relief with irregular inner edges. Septal granules sparse, small, and pointed- Discussion. Gardiner [1939] reported 40 speci- mens of Gardineria lilliei from four stations and implied that 4 environmentally controlled forms were present. Forty-nine specimens, deposited at the British Museum and the Museum of Comparative Zoology, have been examined from these stations, all bearing Gardiner's identification of G. lilliei; however, the 4 implied growth forms are in fact separate species belonging to two genera. (Map 9). 28. Flabellum knoxi Ralph and Squires, 1962 Plate 13, figs. 4-7 Flabellum knoxi Ralph and Squires, 1962, pp. 14, 15, pi. 7, figs. 1, 2.?Squires, 1964a, pp. 11, 12, 19, 20, pi. 1, figs. 4-6, pi. 2, fig. 7, pi. 3, figs. 3-5, pi. 4, figs. 1-4; 1969, p. 18, pi. 6, map 4.?Squires and Keyes, 1967, pp. 26, 27, pi. 5, figs. 1, 2.~Zibrowius, 1974b, p. 18. Description. Corallum flabellate, compressed; angle of lateral edges typically 135?-180?, in- clination of lateral faces 30?-35''. Lateral edges usually rounded, not carinate. Base of pedicel small, 2.5-3.0 mm in diameter; height about 5-10 mm. One of largest specimens (holotype) 112 x 55 mm in CD and 65 nim tall. Theca very thin and fra- gile, bearing thin, incised striae, one corres- ponding to each septum. Closely spaced, transverse growth lines form chevrons, peaking at each stria. Theca uniformly reddish-brown, entirely white, or bearing reddish-brown stripes corresponding to each septum. Darker, broader stripes corresponding to major septa. Calice entire, not lacerate, and strongly arched. Up to 348 thin, fragile septa per calice, ar- ranged in three size groups. Largest septa (pri- maries) extending to columella and having very sinuous lower inner edges. Between each primary a secondary, usually smaller (three fourths of a size of primary) but in larger specimens almost reaching columella. A much smaller tertiary sep- tum, occurring between each primary and secondary, rarely extending more than halfway to pedicel. In CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 43 1-3. 4-7. 8-11. Plate 13. Flabellum Flabellum gardineri n. sp.: holotype, BM 1939.7.20.305, Discovery sta. 160 H = 22.1 mm, GCD = 8.9 mm. Flabellum knoxi Ralph and Squires: 4, USNM 53378, NZOI sta. D-177 GCD = 44.0 ram; 5, USNM 47492, NZOI sta. D-175, H = 22.7 mm, stripes on theca; 6. USNM 53380, NZOI sta. D-179, H = 42.7 mm, coated with ammonium chloride; 7. USNM 47492, NZOI sta. D-175, xl.7, corallum broken, revealing columella. Flabellum apertum Moseley: 8, 9, lectotype, BM 1880.11.25.74, Challenger sta. 145, GCD = 32.2 mm; 10. U, USNM 47444, Eltanin sta. 1412, GCD = 57.3 mm. 44 BIOLOGY OF THE ANTARCTIC SEAS XI larger specimens, rudimentary quaternaries present near calicular edge. Upper septal margins invari- ably broken, but septa do not appear to be exsert. Very small, pointed granules arranged in widely spaced rows parallel to septal margins. Columella long and slender (1.5-2.2 mm wide) but very sturdy, often remaining intact after surrounding septa have been broken away (Plate 13, fig. 7). Columella composed of loose fusion of convoluted lower inner edges of primary septa. Remarks. The living coral appears to remain in the upright position, as is concluded from the presence of attached filter feeders on both lateral faces. Squires [1964a] suggested that it maintains this orientation by sinking its pedicel into soft mud in areas of low-velocity current. Coralla are rarely attached, but if they are, to sand or peb- bles, which are usually incorporated into the ped- icel. According to Squires [1974a], specimens that are accidentally knocked to the prone posi- tion will produce recurved (angle of lateral edges up to 250?) and reflexed coralla in an effort to right the upper half of the polyp relative to the substrate. The variation in thecal striping and color of F. knoxi is similar to that found in F. pavoninum atlanticum Cairns, 1979. In general, the intensity of striping correlates with age [Squires and Keyes, 1967], the younger specimens having the more pro- nounced stripes. Discussion. F. knoxi is extremely similar and may be identical to F. magnificum Marenzeller, 1904, from off Sumatra. The type of the latter species was not examined. Material. Eltanin sta. 1398 (2), USNM 47496; sta. 1989 (7), USNM 47493. NZOI sta. A-898 (5), USNM 47495; sta. D-6 (2), USNM 47494; sta. D-175 (4), USNM 47492; sta. D-176 (15), USNM 53379; sta. D-177 (6), USNM 53378; sta. D-179 (2), USNM 53380; sta. D-207 (8), USNM 53376. Types. The holotype is deposited at the Canter- bury Museum, Christchurch, New Zealand. Type- locality: Chatham Rise; 402-512 m. Distribution. Chatham Rise; Campbell Plateau; off Macquarie Island (Map 9). Depth range: 201- 914 m. 29. Flabellum apertum Moseley, 1876 Plate 13, figs. 8-11; Plate 14, figs. 1-4 Flabellum apertum Moseley, 1876, p. 556 (part: off Prince Edward Islands); 1881, pp. 167, 168 (part: Challenger sta. 145), pi. 6, figs. 7a-7c.?Not F. apertum; Marion, 1906, pp. 120, 121, pi. 11, figs. 9, 9a (is F. angulare Moseley, 1876).?Wells, 1958, p. 262.?Not F. apertum Squires, 1958, p. 68.?Squires and Keyes, 1967, p. 26, pi. 4, figs. 4, 5.?Squires 1969, ppi 16, 18, pi. 6, map 4.?Zibrowius, 1980, p. 154. ? Flabellum patagonichum Moseley, 1881, pp. 166, 167, pi. 15, figs. 1-7.?Fowler, 1885, pp. 585-590, figs. 1-12.?Wells, 1958, p. 262. ?Squires, 1961, p. 30; 1969, pp. 17, 18, pi. 6, map 4.?Squires and Keyes, 1967, p. 27.?Cairns, 1979, p. 206. Description. Corallum campanulate and distinctly compressed. Pedicel short and cylindrical (2.0-2.5 mm in diameter); originally attached to small ob- ject, becoming free early in ontogeny. Largest specimen examined (USNM 47444) 57.2 x 39.3 mm in CD and 37.2 mm tall. The two principal costae ridged and continuous from pedicel to calice. They diverge from pedicel at an apical angle between 130? and 170? until a GCD of about 30 mm, whereupon epitheca turns upward to continue almost vertical growth. At point of inflection, the four lateral C^ usually well developed, sometimes forming spurs, and may continue as ridges to calice. C2 sometimes ridged from point of in- flection to calice, but much less than Cx. Epi- theca porcelaneous, with chevron-shaped growth lines extending between each septum. Calicular profile scalloped, a large apex corresponding to each Si and S2 and a smaller peak corresponding to every S3. Septa hexamerally arranged in four cycles, with rudiments of fifth cycle only in larger specimens. S5 first appearing in end half systems, only rarely in lateral half systems; largest specimen with 68 septa. Sx and S2 equal in size and slightly exsert. Their inner edges thickened and fusing in center of fossa, forming elongate, solid or trabecular columella. S3 and S4 progres- sively smaller and do not reach columella. S5, if present, rudimentary. All septa having straight inner edges and bearing numerous, small, pointed granules. Stereome infilling sometimes at bottom of fossa. F. apert'"" forma patagonichum differing from typical form primarily in its smaller size (maximum size reported, 28 x 21 mm in CD) and more slender shape. Ridged principal costae diverging from pedicel at an apical angle between 90? and 110? and turning upward at GCD of about 15 mm, producing smaller, more slender corallum. Other four Cx not developed. Theca porcelaneous only in region 5-10 mm from calice. Remainder of corallum usually worn and white but may also be uniformly reddish- brown or white with diffuse reddish-brown stripes corresponding to Sx and S2. S4 often missing from half systems and stereome infilling more c oramon. Discussion. F. patagonlchuni Is treated as a forma of F. apertum because both patagonichum and typiral apertum as well as a continuous series of morphological intermediates were present in two lots. The series of 246 specimens from Eltanin station 283 was particularly helpful in tracing the morphological variation possible in one popu- lation. Four other closely related species have been linked to F. apertum: F. angulare Moseley, 1876; F. conuis Moseley, 1881; F. japonicum Moseley, T881; and F. raukawaensis Squires and Keyes, 1967. Gardiner [r929b] synonymized F. angulare, F. ager- tum, and F. conuis as F. japonicum, the incorrect ii^ior synonym. He stated that F. patagonichum might prove to be a form of F. iaponicum also. Keller [1974] synonymized F. iaponicum and F. raukawaensis as F. apertum, which she reported from off South Africa and western India. Zibrowius [1980] distinguished F. angulare from F. apertum and discussed the nominal species. I have examined the type-specimens of all of the above species except F. raukawaensis, of which I have seen a specimen from very near the type- locality (Eltanin station 1403), and have made the following observations. F. iaponicum can be distinguished from F. apertum by its possession of a full fifth cycle of septa at a calicular diameter at which F. apertum has only few S3. Yabe and CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 45 Map 10. Distribution of Flabellum flexuosum (solid circles), Flabellum ongulense (open circle), and Flabellum apertum (solid squares). Eguchi [1941aJ have used this as a key character to differentiate the two species. F. japonicum has been widely reported from the northern Indian Ocean, off Indonesia, Japan, Philippine Islands, and Tasmania. F. angulare is known only from the north Atlantic (1647-3186 m) and can be distin- guished by its less compressed corallum and S3 that extend to the columella. F. conuis, known only from off the Admiralty Islands, is very similar to F. apertum forma patagonichum in shape but differs in having a deeper fossa, a more deli- cate corallum, slightly corrugated septa, and S3 that reach the columella. It is probably a separ- ate species, but obviously more specimens are needed before this can be determined with cer- tainty. F. raukawaensis, known only from four specimens from off North Island, New Zealand, is very similar to typical F. apertum but differs in that it has more S5, a larger columella, and S3 that almost reach the columella. Other records of F. apertum that I have not veri- fied include von Marenzeller [1904a], off Tanzania, East Africa, 863 m; Yabe and Eguchi [1941bJ, off Japan, 307 m; Keller [1974], off South Africa, 1400 ra, and off western India, depth unknown; Keller [1975], Caribbean, depth unknown. Material. Forma apertum: Eltanin sta. 338 (2), USNM 45675; sta. 558 (1), USNM 47442; sta. 1412 (4), USNM 47444; sta. 1414 (2), USNM 47439; sta. 1422 (5), USNM 47441. Is las Orcadas sta. 575-6 (12), USNM 47445. Edisto sta. 7 (1), USNM 4736. HH sta. 64/68 (3), ZIZM. NZOI sta. D-166 (2), USNM 47437. Mixtures of formae apertum and patagonichum: Eltanin sta. 216 (28), USNM 45674; sta. 21-283 (246), USNM 47443. Specimen of Squires and Keyes [1967], USNM 47438. Syntypes of F. apertum and F. patagonichum. ~ Types. The eight syntypes of F. apertum, two from Challenger station 3 (number 1880.11.25.73) and six from Challenger station 145 (number 46 BIOLOGY OF THE ANTARCTIC SEAS XI 1880.11.25.74), are deposited at the British Mu- seum. Because these specimens represent a mixed lot [Zibrowius, 19801, a lectotype is chosen from Challenger station 145 (Plate 13, figs. 8, 9). The remaining specimens are considered paralecto- types, those from Challenger station 3 being F. angulare [see Zibrowius, 1980]. Type-locality: 46?40'S, 37?50'E (off Prince Edward Island); 567 m. The' syntypes of F. patagonichum are also de- posited at the British Museum. Type-locality: 47?48'30"S, 74?47'W (off Isla Penguin, Chile); 220 m. Distribution. Circum-Subantarctic, including off Prince Edward Islands; Hjort Seamount; Mac- quarie Ridge; off Chile; off Falkland Islands; Falkland Plateau; off southern Brazil; off southern New Zealand (Map 10). Depth range: 220-1500 m. 30. Flabellum truncum n. sp. inconstans Marenzeller, 1904, known from off South Africa at 100 m. F. truncum is distinguished by its smaller size, lesser number of septa, and deeper bathymetric range. Etymology. The specific name truncum (Latin: piece cut off, tip) refers to the detached distal anthocyathus stage of this species. Material. Eltanin sta. 21-282 (1), USNM 47526; 47527. Islas Orcadas sta. Anton Bruun sta. 11-88 (2), Plate 14, figs. 5-8 Description. Corallum ceratoid to trochoid, compressed. Angle of lateral edges 45?-70?; incli- nation of lateral faces 22?-38?; ratio of GCD/LCD 1.4-2.2. Corallum (anthocyathus) proximally trun- cated, resulting from transverse fission from a presumably attached base (the anthocaulus). Scar of attachment from 9 to 14 mm long and from 5 to 9 mm wide; usually worn. Largest specimen 38 x 23 mm in CD; tallest specimen 39.4 mm in height. Theca thin, sometimes bearing low, rounded, longi- tudinal ridges, one corresponding to each Sj^_3. Lateral edges rounded, never carinate or spinose. Calicular margin entire; profile of margin arched. Septa hexamerally arranged in five cycles, rarely with additional Sg. Si, S2, and S3 equal in size and extending to columella. S4 about half of size of S^; S5 about one fourth of size of S4. Sometimes lower inner edges of S4 bending toward and fusing with S3. Septa not exsert and bearing large, pointed or blunt granules measuring as high as septal thickness. Lower inner edges of larger septa thickened and slightly sinuous; sinuosity corresponding to shal- low undulations of currugated septa. Columella variable in structure, but usually slender, elon- gate, loose fusion of inner edges of Si_3; may sometimes be a slightly wider, flat, solid mass or a very wide (up to 25% of LCD), loose, spongy structure. Remarks. Zibrowius [1974b] suggested that the truncated flabellids may reproduce asexually by transverse division ('strobilation') with one basal part (the anthocaulus stage [see Wells, 1966, p. 226]), producing more than one corallum (the an- thocyathus stage). Unfortunately, no attached specimens or specimens in the process of dividing were found among the lots of F. truncum. When the attached and free stages of the truncated flabel- lids are known, some species will probably be synonymized. Discussion. F. truncum belongs to the flabella truncata section of the genus Flabellum [see Milne Edwards and Hairae, 1848, p. 259], which is equiva- lent to Zibrowius's [1974b] 'second group.' Zib- rowius listed 19 species from this group, which are all characterized by a transverse division, but none of these are known from the eastern Pacific or the Subantarctic. Many are distin- guished by prominent costal spines or crests. Out of this group, F. truncum is most similar to F. sta. 338 (1), US^^M 575-5 (1), USNM 47528. USNM 47529. Types. Types. The holotype, collected at Eltanin sta- tion 21-283, is deposited at the United States National Museum (47174). Six paratypes from El- tanin station 21-283 (number 47175) and 19 from AKt3^ Bruun station 18-714 (number 47176) are de- posited at the United States National Museum. One paratype from Anton Bruun station 18-714 is depos- ited at the BTitlih Museum (1979.11.4.1). Type- locality: 53?13'S, 75?41'W (off Isla Desolacion, Chile); 1500-1666 m. Distribution. Off western coast of South America Peru to off southern Chile; south of Islands; Falkland Plateau (Map 11). 595-1896 m. from off Falkland Depth range: Genus Javania Duncan, 1876 Diagnosis. Solitary, ceratoid to trochoid, fixed. Wall epithecal. Base reinforced by layers of stereome. No pali. Calicular edge jagged. Co- lumella rudimentary. Type-species: Javania in- signia Duncan, 1876, by monotypy. 31. Javania cailleti (Duchassaing and Michelotti, 1864) Plate 14, figs. 9-12 Desmophyllum cailleti Duchassaing and Michelotti, 1864, p. 66, pi. 8, fig. U. Desmophyllum eburneum Moseley, 1881, p. 162, pi. 6, figs, la, lb. Desmophyllum nobile Verrill, 1885, pp. 150, 151. Desmophyllum vitreum Alcock, 1898, p. 20, pi. 2, figs. 2a, 2b. Flabellum sp. Marenzeller, 1904b, pi. 81. Desmophyllum galapagense Vaughan, 1906b, p. 63, pi. 1, figs. 1, lb. Javania eburnea; Zibrowius, 1974b, pp. 12, 13, pi. 3, figs. 13-17; 1980, pp. 157-159, pi. 82, figs. A-L. Javania cf. eburnea; Zibrowius, 1974b, pp. 13-16, pi. 4, figs. 22-29, pi. 5, figs. 31-34. Javania vitrea; Zibrowius, 1974b, pp. 16, 17, pi. 5, figs. 18-21. Javania cailleti; Cairns, 1979, pp. 153-156, pi. 28, figs. 8-12, pi. 30, figs. 1, 4. Description. J. cailleti has been fully des- cribed and illustrated elsewhere [Zibrowius, 1974b, 1980; Cairns, 1979]; only a brief diagnosis is given here. Corallum ceratoid, often flared dis- tally. Pedicel thick, reinforced by concentric layers of stereome up to PD of one fourth to one half of CD. Typical specimen 18 x 14 mm in CD and 35 mm tall. Theca usually smooth and porcelaneous but may be ridged with costae near calice. Septa usually hexamerally arranged in four cycles: Si S2 > S3 > S4 in size. Si and S2 highly "exsert;" S4 rudimentary. Inner septal edges straight. Septal granules low, rounded. CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 47 Plate 14. Flabellum and Javania 1-4. Flabellum apertum Moseley: 1, USNM 47443, Eltanin sta. 283, H = 24.0 mm; 2, specimen from same lot, GOD = 26.8 mm; 3, 4, syntype of F. patagonichum Moseley, BM, Challenger sta. 305, GCD about 28 mm. 5-8. Flabellum truncum n. sp.: 5, 6, holotype, USNM 47174, Eltanin sta. 283, H = 31.7 mm, GCD = 27.7 mm; 7, paratype, USNM 47175, Eltanin sta. 283, GCD = 25.6 mm, large spongy columella; 8, USNM 47529, Anton Bruun sta. 11-88, x2.2, view of basal fracture. 9-12. Javania cailleti (Duchassaing and Michelotti): 9, 10, USNM 19173, Albatross sta. 2785, H = 22.4 mm, GCD = 20.2 mm; 11, 12, USNM 47530, Eltanin sta. 1592, H = 20.4 mm, GCD = 10.2 mm. 48 BIOLOGY OF THE ANTARCTIC SEAS XI Map U. Distribution of Gardineria antarctica (solid circles), fossil Gardineria ant- arctica (open circle), and Flabellum truncum (solid squares). Fossa deep and narrow; rudimentary columella formed by solid fusion of lower inner edges of S^ and Discussion. Of the two new specimens reported, the one from Albatross station 2785, about 10 km from the type-locality of D. eburneum Moseley, is identical to the syntypes of that species.- The small specimen from Eltanin station 1592, however, has prominent costal ridges and highly exsert septa, similar to those of the holotype of D. nobile Verrill. Material. Eltanin sta. 1592 (1), USNM 47530; USNM 19173. Specimens USNM. Syntypes of D. nobile and D. ealapa- Albatross sta. 2785 (1), listed by Cairns [1979], eburneum; holotypes of D _ gense. Types. See Cairns [1979]. Type-locality: ser Antilles. Distribution. Widespread in Northern Hemisphere Les- off India; off Morocco; northwest Mediterranean; Celtic Sea; off Madeira; off Azores; off Nova Scotia; off Georgia, United States, to off Surinarae (including Gulf of Mexico and Caribbean). In Southern Hemisphere known from off Uruguay; off Burdwood Bank; off Chile (Map 12). Depth range: 86-2165 m. 32. Javania antarctica (Gravier, 1914) n. comb. Plate 15, figs. 1-4 Desmophyllum antarcticum Gravier, 1914a, pp. 236-238; 1914b, pp. 122-125 (part: not dredge 8), pi. 1, figs. 1-3. Not Flabellum antarcticum; Wells, 1958, p. 269 (is F. flexuosum tT sp.).~Not F. antarcticum; Squires, 1962b, pp. 13, 14, 19, 20 (is F. flex- uosum n. sp.) and not 1969, p. 18 (is F. flexuo- sum n. sp.).?Not F. antarcticum; Keller, 1974, in all oceans: off Galapagos Islands; off Japan; p. 203 (is F. curvatum). CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 49 !" Plate 15. Javania and Gardineria 1-4. Javania antarctlca (Gravier): 1, MNHNP (no number), locality unknown H = 64.0 mm; 2, 3, USNM 47464, Hefo sta. 731-1865, GCD = 24.7 mm; 4, USNM 53407 Ed isto sta. 15, H = 71.7 mm. ' 5-11. Gardineria antarctica Gardiner: 5, USNM 47249, Eltanin sta. 2119, H = 22.6 mm; 6, specimen from same lot, CD = 27.4 mm, columella labyrinthiform; 7, USNM 47247, Eltanin sta. 5762, CD = 25.5 mm, coated with ammonium chloride, paliforra lobes present; 8, USNM 47248, Eltanin sta. 2082, CD = 28.1 mm; 9, USNM 47205, Eltanin sta. 2117, CD = 26.1 mm, massive columella; 10, lecto- type of G. lilliei Gardiner, BM 1929.10.22.9, Terra Nova sta. 194, GCD = 12.5 mm; 11, syntype of G. antarctica, BM Terra Nova sta. 349, CD = 19.0 mm. 50 BIOLOGY OF THE ANTARCTIC SEAS XI ? Desmophyllum delicatum; Niino, 1958, p. 257, pi. 2> fig- 3. ? Desmophyllum pseudoseptata Eguchi, 1965, p. 9, pi. 2, figs. 3a-3c. Description. Corallum ceratoid to trochoid, tall, straight to slightly curved. Pedicel rein- forced by numerous concentric layers of stereome, producing PD ranging from 5.7 to 12.5 mm. Pedicel more expanded at base of attachment. Largest specimen (Gravier's [1914b] illustrated syntype) 44 X 38 mm in C0, about 8.0 in PD, and 65 mm tall. Theca very thin and porcelaneous; encrusting or ganisms may settle on theca, these organisms per- iodically covered over with stereome. Chevron- shaped growth lines peak at insertion lines of major septa. Insertion lines sometimes slightly grooved, resembling costae. Calice elliptical, ratio of GCD/LCD between 1.15 and 1.60. Septa hexamerally arranged in five cycles. Sj^ and S2 equal in size and slightly exsert. S3 slightly smaller but much larger than 84; S5 rudimentary. Septa very thin and delicate; speci- mens rarely collected with intact upper septal edges. Inner septal edges slightly sinuous, those of Si and S2 thickened lower in fossa, there loosely fused into rudimentary collumella. Septal granulation fine and pointed. Fossa deep. Discussion. J. antarctica is very similar to F. flexuosum but can be distinguished by its thicker, reinforced pedicel and its later development of a full fifth cycle of septa. Wells [1958] and Squires [1962b, 1969], perhaps not realizing the importance of the pedicel (Zibrowius resurrected Javania only as recently as 1974), lumped the two together. Eguchi's [1965] Desmophyllum pseudosep- tata is probably a Javania and may be J. antarc- tica; however, Eguchi's holotype could not be lo- cated. Niino's [1958] D. delicatum is based on the same specimen. Material. Eltanin sta. 499 (1), USNM 47462; sta. 1054 (1), USNM 47455; sta. 1081 (1), USNM 47461; sta. 1089 (2), USNM 47458. Is las Orcadas sta. 575-34 (1), USNM 47463; sta. 575-89 (1), USNM 47459. Hero sta. 731-1865 (1), USNM 47464; sta. 731-1940 (1), USNM 47456; sta. 731-1947 (3), USNM W460. Edisto sta. 15 (3), USNM 53407. EW sta. 4 (1), USNM 47457; one specimen without locality data (MNHNP). Xypes. One syntype from the Pourquoi-Pas? sta- tion 4 (illustrated by Gravier [1914b, pi. 1, figs- 1, 2]), is deposited at the Museum National d'His- toire Naturelle, Paris. The other syntype from the same station and the doubtfully assigned spec- imen from Pourquoi-Pas? station 8 (the latter probably F. flexuosum) could not be found at the Museum National d'Histoire Naturelle; however, another large, typical specimen without locality data is present. Type-locality: 64?50'S, 63?30'W (off Anvers Island, Palmer Archipelago); 53 m. Distribution. Off western Antarctic Peninsula; Scotia Ridge trora South Shetland Islands to South Georgia; off Cape Norvegia, Queen Maud Land; ? off Riiser-Larsen Peninsula (Cape Cook), Prince Harald Coast (Map 12). Depth range: 53-1280 m. Genus Gardineria Vaughan, 1907 Diagnosis. Solitary, turbinate to cylindrical, fixed. Wall epithecal but thickened internally by stereome. Septa not always arranged hexamerally. Paliform lobes opposite larger septa. Columella well developed, papillose. Type-species: Gardi- neria hawaiiensis Vaughan, 1907, by original des- ignation. 33. Gardineria antarctica Gardiner, 1929 Plate 15, figs. 5-11 Flabellum sp. Pax, 1910, p. 73, pi. 11, fig- 2. Caryophyllia sp. ? Gravier, 1914b, pp. 130, 131, pi. 1, figs. 9, 10. Gardineria antarctica Gardiner, 1929a, pp. 124, 125, 128-130, pi. 1, figs. 11-18; 1939 (part: not Discovery sta. 152), p. 328.?Wells, 1958, pp. 269, 270, pi. 2, figs. 16-18.?Squires, 1961, p. 20; 1962b, pp. 11, 13, 15, 20, 21, pi. 1, figs. 1-10; 1969, pp. 17, 18, pi. 6, map 2.?Speden, 1962, p. 756, figs, lla-llc?Zibrow- ius, 1974b, p. 24.~Podoff, 1976, pp. 45, 46, pi. 3, figs. 12, 13. Gardineria lilliei Gardiner, 1929a, p. 125, pi. 1, figs. 3-10.?Wells, 1958, p. 262. Caryophyllia inskipi; Thomson and Rennet, 1931, p. 40, pi. 10, fig. 6. Ceratorochus (Convtorochus) parphis; Niino, 1958, p. 257, pi. 2, fig. 5 (misspellings). Gardineria lillei; Squires, 1961, p. 20; 1962b, p. 13; 1969, pp. 17, 18, pi. 6, map 2.?Zibrow- ius, 1974b, p. 24. Ceratotrochus ? sp. Eguchi, 1965, p. 8, pi. 2,figs, la, lb. Description. Corallum usually straight, regular cone with round calice; trochoid to turbinate, sometimes ceratoid. Pedicel diameter 2.8-7.7 mm, expanding up to 12 mm at base of attachment. Largest specimen examined 32.8 mm in CD and 32.2 mm tali. Theca thick and not porcelaneous; usually smooth, but sometimes bearing low granulated ridges, one corresponding to each septum. Upper calicular margin horizontal and entire. Septa hexamerally arranged in five cycles. Sj^ slightly larger than $2, these slightly larger than S3. Septa of first three cycles not exsert and extending to columella. Each of these septa bearing shallow, nondentate notch near its upper junction with theca. Short thecal lip sometimes present, extending slightly above septal inser- tions. S4 about half size of S]^, sometimes bending slightly toward S3; S5 rudimentary, extending about one fourth of way to base. All septa with straight inner edges and covered by very low, rounded granules, usually arranged in rows parallel to septal edge. Fossa shallow, containing prominent columella of variable structure. S]^_3 often bearing one to three long paliform lobes, these sometimes round or flattened in cross section. Four to thirty of these lobes may be present, forming columella. Sometimes columellar lamellae labyrinthine in arrangement (Plate 15, fig. 6) and sometimes greatly thickened (Plate 15, fig. 9). Remarks. BuUivant [1967] reported a deep slope cobble assemblage on the Pennell Bank, Ross Sea (461-583 m), characterized by G. antarctica, various echinoderms, sponges, bryozoans, and stylasterine coral. Squires [1962b] had earlier reported G. antarctica, Flabellum antarcticum (actually F. flexuosum), and Caryophyllia CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 51 Map 12. Distribution of Javania antarctica pseudoseptatum (open circle), Javania cailleti vermiformis (solid squares), and Balanophyllia sp. (open square). (solid circles), (solid triangles), Desmophyllum Stenocyathus antarctica from the same locality. These three species have also been collected together several times off Cape Hallett, Ross Sea, at 342-433 m (Eltanin stations 1870, 1995, 1996; Atka station 23; and Burton Island station 3), often attached to the stylasterine coral Errina fissurata (Gray). In the assemblage from off Cape Hallett, both F. f lexuosum and C. antarctica are small and often scolecoid; the C. antarctica are often missing pali, resembling Cyathoceras. One of the specimens from Eltanin station 2097 had a full gastrovascular cavity of disarticulated and partially digested, small, shrimplike crusta- ceans. Judging from the size of the antennule scales, two species are represented, the larger one probably measuring 20 mm (B. Kensley, personal communication, 1979). Discussion. The specimens reported as G. lilliei by Gardiner [1929a] are typical small G. antarctica. His redescription of G. lilliei [Gardiner, 1939] is based on a suite of three or four species, as he suggested may have been the case: Flabellum flexuosum, Caryophyllia eltaninae, Flabellum gardineri, and Flabellum sp. Gardiner's [1929a] Gardineria sp., from off New Zealand, resembles Caryophyllia antarctica in its septa! ornamentation and columella but lacks the distinc- tive pali. G. antarctica superficially resembles G. capensis (Gardiner, 1904) from off South Africa but can be distinguished by its extra cycle of septa, in- equality in size of 81-3, and lack of exsert septa. G. antarctica is the only scleractinian fossil known from the Antarctic continent [Squires, 1962b, p. 15], reported from various localities around McMurdo Sound, Ross Sea, from the Pliocene-Lower Pleistocene to the Pleistocene-Subrecent. Speden 52 BIOLOGY OF THE ANTARCTIC SEAS XI [1962] noted that the fossil specimens have a much thicker theca. One of the fossil specimens men- tioned by Gardiner [1929a] is illustrated by David and Priestley [1914, pi. 88, figs. 4, 5]. Material. Eltanin sta. 1081 (17), USNM 47194; sta. 1082 (1), USNM 47183; sta. 1088 (2), USNM 47186; sta. 1089 (4), USNM 47206; sta. 1870 (125), USNM 47209; sta. 1871 (1), USNM 47181; sta. 1922 (2), USNM 47540; sta. 1924 (1), USNM 47200; sta. 1931 (4), USNM 47182; sta. 1933 (2), USNM 47203; sta. 1944 (3), USNM 47188; sta. 2021 (6), USNM 47189; sta. 2022 (1), USNM 47204; sta. 2045 (1), USNM 47195; sta. 2072 (2), USNM 47198; sta. 2075 (1), USNM 47191; sta. 2082 (4), USNM 47248; sta. 2097 (1), USNM 47202; sta. 2099 (1), USNM 47192; sta. 2117 (30), USNM 47205; sta. 2119 (38), USNM 47249; sta. 2120 (3), USNM 47180; sta. 2124 (3), USNM 47190; sta. 2125 (1), USNM 47185; sta. 5761 (4), USNM 47187; sta. 5762 (1), USNM 47247; sta. 5765 (9), USNM 47208. Hero sta. 691-20 (1), USNM 47196; sta. 731-1844 (1), USNM 47199. Edisto sta. 20 (1), USNM 53406. Glacier sta. 1 (1), USNM 47206. Staten Island sta. 21 (15), USNM 47193. EW sta. 9 (5), USNM 45663; sta. 32 (10), USNM 47197. NZOI sta. A-625 (4), USNM 47201. EAD sta. 2 (8), USNM 53399; sta. 3 (4), USNM 47184. Speci- mens (3) identified as Caryophyllia inskipi by Thomson and Rennet [1931], Australian Museum G 13537; specimens (2) identified as Gardineria ant- arctica from Discovery sta. 152 by Gardiner [1939], BM 1939.7.22.242-243. Syntypes of G. antarctica and G. lilliei. Types. The seven syntypes of G. antarctica, collected at Terra Nova stations 314 and 349, are deposited at the British Museum. Type-locality: The six syntypes of G. lilliei are a mixed lot, represented by five specimens of G. antarctica and one basal fragment of a Flabellum, perhaps F. flexuosum. Therefore, one specimen (Plate 15, fig. 10) is designated as lectotype. All specimens are deposited at the British Museum (1929.10.22.9- 14). Type-locality: 69?43'S, 163?24'E (off Oates Coast, Antarctica); 329-366 m. Distribution. Circumpolar: off Antarctica; off South Shetland Islands; east of South Orkney Islands; off Scott Island (Map 11). Depth range: 87-728 m. Family GUYNIIDAE Hickson, 1910 Genus Stenocyathus PourtalSs, 1871 Diagnosis. Solitary, ceratoid to cylindrical, free or attached. Wall epithecal; rows of thecal spots flank each S3. Pali, when present, oppo- site S2. Columella formed of one to four twisted, crispate ribbons. Type-species: Coeno- cyathus vermiformis PourtalSs, 1868, by monotypy. 34. Stenocyathus vermiformis (PourtalSs, 1868) Plate 16, figs. 8-11 Coenocyathus vermiformis PourtalSs, 1868, pp. 133, 134. Stenocyathus vermiformis; von Marenzeller, 1904a, pp. 298-300, pi. 18, fig. 16.?Zibrowius, 1974a, pp. 769, 770; 1980, pp. 163-165, pi. 84, figs. A-Q.~Cairns, 1979, pp. 168-170, pi. 32, figs. 8-10, pi. 33, figs. 1, 2. Stenocyathus decamera Ralph and Squires, 1962, pp. 11, 12, pi. 4, figs. 2-6.?Squires and Keyes, 1967, p. 28, pi. 6, figs. 3-5.?Squires, 1969, p. 17, pi. 6, map 2. Description. This species has been fully de- .scribed and illustrated elsewhere [Cairns, 1979; Zibrowius, 1980]; only a brief description follows. Corallum cylindrical, elongate, vermiform, up to 50 mm long but rarely over 5 mm in CD. Free or attached; when attached, reinforced basally by layers of granular stereome. Epitheca thin, por- celaneous; usually marked by 24 longitudinal rows of white spots, 1 row corresponding to each inter- septal space. Septa hexamerally arranged in three systems. Si largest septa; S2 usually larger than S3 but may be of same size or smaller. Septa usually not exsert. Inner edges of septa sinuous. Thick pali before S2; columella formed of one to four twisted ribbons. Septa and pali bear large granules. Discussion. Additional variations noted in the Subantarctic specimens include the following: (1) the Si are sometimes exsert, (2) the S3 are sometimes equal to or larger than the S2, and (3) the columella may be composed of up to four elements. The white spots on the theca are solid, smooth structures but probably represent areas of lesser calcification, as is evidenced by their earlier erosion to pores after death of the coral. Material. Eltanin sta. 1284 (8), USNM 47450; sta. 1411 (1), USNM 47448; sta. 1691 (1), USNM 47454; sta. 1851 (1), USNM 47446. NZOI sta. A-740 (1), USNM 47449; sta. D-159 (9). USNM 47452; sta. D-160 (1), USNM 47451; sta. D-175 (1), USNM 47453. Specimens (9) identified as S. decamera by Squires and Keyes [1967] from NZOI sta. B-319, USNM 47447; specimens listed by Cairns [1979], USNM. Syntypes of C. vermiformis. T^pes. The types of C. vermiformis are at the Museum of Comparative Zoology [see Cairns, 1979]. Type-locality: Florida Keys; 274-329 m. The holotype of S. decamera is deposited at the New Zealand Geological Survey, Wellington. Type- locality: off New Zealand; 110-220 m. Distribution. Widely distributed in Atlantic Ocean: Mediterranean Sea; area bordered by Celtic Sea, Azores, and Madeira; western Atlantic from off Georgia, United States, to off Rio de Janeiro, Brazil; off Penedos de SSo Pedro e Sao Paulo (St. Peter and Paul Rocks). ^ From more southern latitudes known from off lie Saint-Paul and He Amsterdam, Indian Ocean; off New Zealand; Campbell Plateau; off Antipodes Islands; several searaounts in South Pacific (Map 12). Depth range: 80-1229 m. Suborder DENDROPHYLLIINA Vaughan and Wells, 1943 Family DENDROPHYLLIIDAE Gray, 1847 Genus Balanophyllia Wood, 1844 Diagnosis. Solitary, turbinate to trochoid, fixed or free. Costae well developed. Synapti- culotheca porous, especially near calicular edge. Septa follow Pourtal&s plan. Pali present or ab- sent. Columella spongy. Type-species: Balano- phyllia calyculus Wood, 1844, by monotypy. 35. Balanophyllia malouinensis Squires, 1961 Plate 16, figs. 4-7; Plate 17, figs. 1-3 Plate 18, fig. 7 Balanophyllia cornu; Gardiner, 1939, pp. 335, 336. Balanophyllia malouinensis Squires, 1961, pp. 15, CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 53 Plate 16. Balanophyllia and Stenocyathus 1-3. Balanophyllia chnous Squires: holotype, BM 1929.10.22.25, Terra Nova sta. 191, H = 30.4 mm, GCD = 12.1 mm. ^-7. Balanophyllia malouinenais Squires: 4, specimen identified as B. cornu by Gardiner [1939], BM 1939.7.20.234, WS sta. 839, H = 34.4 mm; 5, 6, USNM 45672, Eltanin sta. 558, GCD = 22.8 mm; 7, specimen from same lot, xl.8, coated with ammonium chloride to illustrate columella, stereotheca, and synapticulotheca. 8-11. Stenocyathus vermifonnis (PourtalSs): 8, USNM 47447, NZOI sta. B-319, H = 14.4 mm; 9, USNM 47454, Eltanin sta. 1691, H = 13.9 mm; 10, USNM 47466, Eltanin sta. 1851, CD = 4.0 mm; 11, USNM 47448, Eltanin sta. 1411, H = 7.4 mm, illustrating exothecal deposits. S4 BIOLOGY OF THE ANTARCTIC SEAS XI 39, 40, 46, figs. 5, 24-26; 1969, pp. 17, 18, pi. 6, map 2.?Sorauf and Podoff, 1977, pp. 4-6, pi. 2, figs. 5, 6, pi. 3, fig. 6, pi. 4, figs. 2-5.?Cairns, 1979, p. 206. Description. Corallum ceratoid to subcylindri- cal, straight to slightly curved, usually free when adult. Basal disc diameter about 4.2 mm, narrowing slightly to PD of about 3.5 mm, then gradually expanding into ceratoid corallum. Some coralla, however, subcylindrical and remaining firmly attached by strengthening base with layers of stereome, up to 12 mm in diameter. Young cor- alla usually weakly attached to epitheca of same species, F. curvatum, small gastropods, bivalves, or pebbles. Largest specimen examined 23.1 x 21.2 mm in CD and 57.5 mm tall. Synapticulotheca thick, very porous, and spinose. Costal spines usually randomly arranged, but in some specimens aligned longitudinally and separated by striae, resembling costae. Between 60 and 100% of synapticulotheca covered by thin, irregularly banded epitheca, of- ten leaving only small ring of synapticulotheca visible at calicular edge. According to Sorauf and Podoff [1977], synapticulotheca gradually in- filled by stereome, forming more solid 'stereo- theca' (Plate 16, fig. 7). Septa hexamerally arranged in five cycles but only largest specimens with complete fifth cycle. Si and S2 equal in size and extending to coluraella. Remaining septa arranged in PourtalSs plan: S, smallest septa, S. adjacent to S^ and S2 larger than S3 and extending to columella. Septa not exsert and with straight inner edges. Septal granulation variable, from very sparse (smooth septal faces) to crowded arrangement of tall, blunt granules. Columella discrete, massive, elongate structure, resting in shallow, elliptical fossa; either spongy or composed of numerous twisted ribbons, swirled in clockwise direction. Columella may be gran- ulated. Discussion. Contrary to Squires's [1961] origi- nal description, based on six worn specimens, this species does not have exsert septa and does some- times have costae. The presence or absence of costae is the basic difference between Balanophyl- lia and Thecopsammia, the latter lacking costae. The variable nature of this character in B. ma- louinensis implies that Thecopsammia may be a jun- ior synonym of Balanophyllia. Contrary to Wells [1956J and Squires [1961], Thecopsammia socialis (type-species of Thecopsammia) has a distinct PourtalSs plan in the adult stage [see Cairns, 1979]. Material. Eltanin sta. 339 (255), USNM 47179; sta. 340 (15), USNM 45671; sta. 346 (1), USNM 47154; sta. 369 (6), USNM 47149; sta. 558 (221), USNM 45672; sta. 740 (103), USNM 47153; sta. 970 (21), USNM 47146; sta. 977 (205), USNM 47148; sta. 1521 (6), USNM 47147; sta. 1536 (12), USNM 47178; sta. 1596 (1), USNM 47155. Hero sta. 715-895 (7), USNM 47177. Vema sta. 14-12 (5), AMNH; sta. 14-18 (1), AMNH; sta. 15-PD3 (9), USNM 47150; sta. 15-PD4 (78), USNM 47151; sta. 15-PD9 (6), AMNH; sta. 15- PDIO (9), USNM 53408, and (24), AMNH; sta. 17-59 (6), USNM 53409, and (11), AMNH. Following WH records (H. Zibrowius, personal communication, 1979): sta. 324/66 (2), sta. 325/66 (1), sta. 330/66 (1), sta. 336/66 (3), sta. 357/66 (2), sta. 359/66 (2), sta. 360/66 (1), sta. 269/71 (4), sta. 270/71 (1) (all WH specimens desposited at ZIZM), Specimens (5) identified as B. cornu by Gardiner [1939], BM 1939.7.20.227-228, 234. Holotype. Types. The holotype is deposited at the American Museum of Natural History (3368). Type-locality: 52?32'S, 61?15'W (south of East Falkland island); 358 m. Distribution. Off Tierra del Fuego; off Falkland Islands; Scotia Ridge between Burdwood Bank and South Georgia (Map 13). Squires's [1969] record from off Gough Island unsubstantiated. Depth range: 75-1137 m. 36. Balanophyllia sp. Plate 17, figs. 4-8 Dendrophyllia oahensis; Gardiner, 1939, pp. 334, 335.?Squires, 1961, p. 21. Description. In the following, William Scoresby station 244 specimens are described. Ceratoid to cylindrical coralla, attached to dead coralla of same species in pseudocolonial arrangement. One 'corallite' 34.4 mm long and cylindrical (CD = 7.7 x 5.7 mm); others ceratoid, shorter, with larger calices (GCD up to 14.0 mm). Thin epitheca covers most of porous synapticulotheca; synapticulotheca longitudinally striate. Septa hexamerally arranged in four complete cycles with no S5. Sy and 52 equal in size and extending to columella. 53 half as large and enclosed by pairs of larger 54, these sometimes fusing near inner edge of S3 and extending to columella. Septa not exsert, slightly porous, especially toward theca, and bearing large, pointed granules. Columella large and spongy. Discussion. These specimens were collected in the geographic and depth range of B. malouinensis but are distinguished by their cylindrical shape, tendency toward pseudocoloniality, and lack of 55, even at a relatively large calicular dia- meter. They are, however, similar to small col- onies of the northern Atlantic Dendrophyllia corn- ucopia Pourtal&s, 1871. D. cornucopia sometimes has extratentacular budding similar to that of the pseudocolonial Balanophyllia sp., and the septal arrangement is identical when small corallites of D. cornucopia are considered. Tne holotype of D. oahensis Vaughan, 1907, known only from off Hawaii, differs from these specimens by having sinuous septal edges and a truly colonial growth form. Material. WS sta. 244 (3 pseudocolonies), BM, and (1 pseudocolony), MCZ 3571. Distribution. Known only from 52?00'S, 62?40'W (east of East Falkland island) (Map 12). Depth range: 247-253 m. 37. Balanophyllia chnous Squires, 1962 Plate 16, figs. 1-3 Thecopsammia sp. Gardiner, 1929a, pp. 126, 127. Balanophyllia sp. Gardiner, 1929a, pp. 126, 127. Balanophyllia chnous Squires, 1962b, pp. 13, 21, 22, pi. 1, fig. 17, pi. 2, figs. 1-3; 1969, pp. 17, 18, pi. 6, map 2. Description. In the following the holotype is redescribed. Corallum ceratoid, becoming cylin- drical toward calice, firmly attached to substrate by pedicel 6.2 mm in diameter. Calice elliptical, CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 55 f Plate 17. Balanophyllia 1-3. Balanophyllia malouinensis Squires: 1, USNM 47179, Eltanin sta. 339, x2.4; 2, USNM 47153, Eltanin sta. 740, H = 19.1 mm, coated with ammonium chloride; 3, USNM 47179, Eltanin sta. 339, xl.9, 3 young, attached specimens. 4-8. Balanophyllia sp. (specimens identified as B. oahensis by Gardiner [1939], BM (no number), WS sta. 244): 4, GCD = 11.1 mm; 5, GOD = 13.2 mm; 6, GOD = 7.7 mm; 7, x2.0; 8, x2.2 m. BIOLOGY OF THE ANTARCTIC SEAS XI Map 13. Distribution of Balanophyllia malouinensis (solid circles), Balanophyllia chnous (open circle), Enallopsammia rostrata (solid squares), and Enallopsammia maren- zelleri (solid triangleTi 12.1 X 9.3 ram in diameter; corallum 30.4 mm tall. Epitheca covering porous synapticulotheca to within 3 mm of calice. Epitheca heavily encrusted with Bryozoa, Foraminifera, serpulids, and sponges. Costae not apparent but may have been present before epitheca formed. Sixty-six septa hexam- erally arranged in five incomplete cycles. S]^ largest septa, slightly exsert, and extending to columella. S2 slightly smaller, not reaching columella. Remaining septa arranged in PourtalSs plan, those adjacent to Sj^ and S2 large and usually extending to columella. Development of S5 irregular within systems: the two lateral systems without S5, but the four systems adjacent to lateral edges with variable number of S5. Three half systems with all four 85. Inner edges of S]^ 2 straight and entire; those of 33-5 irregular to laciniate. Septal granules prominent, as large as septal thickness in height. Short, Low carinae present on inner edges of larger septa. Columella elongate and spongy. Discussion. B. chnous is the only dendrophylliid known from off continental Antarctica. The only specimens known were reported by Gardiner [i929a] as Thecopsammia sp. and Balanophyllia sp. He stated that the specimens were collected at Terra Nova station 91 (off New Zealand) at the beginning of his species account and at Terra Nova station 191 (Bay of Whales, Ross Sea) at the end of the species account. The original label reads 191. However, Gardiner also validly reported specimens of other species from Terra Nova station 91 in the same paper. If there was a labeling error, it might explain the subsequent lack of records of this species in the Ross Sea, even though several Eltanin stations have been made in the proximity of the type-locality. B. chnous differs from B. malouinensis by its CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 57 exsert septa; smaller size at maturity; and longer, more slender corallum. Material. Holotype. Types. The holotype (number 1929.10.22.25) and five paratypes (numbers 1929.10.22.22-24, 26-27) are deposited at the British Museum. They were presumably collected at Terra Nova station 191. Type-locality: Bay of Whales, Ross Sea; 355-457 m. Distribution. Known only from type-locality (Map 131^ Genus Enallopsammia Michelotti, 1871 Diagnosis. Dendroid (often uniplanar) colonies formed by extratentacular budding. Coenosteum compact, synapticulothecate, porous only near cal- ices. Septa arranged normally. Columella small. Type-species: Coenopsammia scillae Seguenza. 1864, by monotypy. Columella 38. Enallopsammia rostrata (Pourtales, Plate 18, figs. 1-4 1878) Amphihelia rostrata PourtalSs, 1878, p. 204, pi. 1, figs. 4, 5. Stereopsammia rostrata; PourtalSs, 1880, pp. 97, 110, 111, Dendrophyllia (Coenopsammia) amphelioides Alcock, 1902, pp. 42, 43, pi. 5, figs. 37, 37a.?Not D. (C.) amphelioides; Gardiner and Waugh, 1939, p. 238. Amphihelia adminicularis Rehberg, 1892, p. 10, pi. 4, fig. 1. Anisopsammia rostrata; von Marenzeller, 1904a, pp. 314, 315, pi. 18, fig. 23.?Gravier, 1920, pp. 102-104, pi. 12, figs. 181-185. Anisopsammia amphelioides; Vaughan, 1907, pp. 156, 157, pi. 47, figs. 1, 2. Dendrophyllia amphelioides var. cucullata Vaughan, 1907, p. 157, pi. 47, fig. 3, pi. 48, figs. 1-4. Madrepora ramea; Gardiner and Waugh, 1939, pp. 226, 227. Dendrophyllia minuscula; Gardiner and Waugh, 1939, p. 237 (part). Enallopsammia rostrata; Squires, 1959, p. 40.?La- borel, 1970, p. 156.?Zibrowius, 1973, pp. 44, 45, pi. 2, figs. 14, 15; 1980, pp. 201-203, pi. 105, figs. A-K, pi. 106, figs. A-C?Cairns, 1979, pp. 186-188, pi. 37, figs. 2, 3, 6. Enallopsammia amphelioides; Zibrowius, 1973, pp. 45-48, pi. 3, figs. 16-20; 1980, pp. 203, 204, pi. 106, figs. D-I. Description. Colony flabellate, dendroid; extra- tentacular branching occurring at every or every second or third corallite. Base of colony massive, up to 3 cm in diameter. Calices 3-5 mm in diameter, round to teardrop shaped, occurring on only one side of colony. Calices projecting upward costoseptal rostrum, formed by enlargement of one CSj^ and sometimes adjacent septa. Rostrum vari- able in development and sometimes absent. Both sides of branches costate, coenosteum solid. Septa hexamerally arranged in three cycles, rarely with additional S4. S]^ largest septa, septa of other two cycles progressively smaller; S3 some- times fused to S2 halfway to columella. Septa not exsert except for Sj forming rostrum. Inner edges of septa variable, from smooth to laciniate, sometimes bearing wide paliform lobes. All septa narrow, thick near calice and thinner toward columella, and bearing spiny granules, rudimentary, spongy. Discussion. More complete descriptions, addi- tional illustrations, and an explanation of the synonymy can be found in works by Zibrowius [1973 1980] and Cairns [1979]. According to Zibrowius [1973] the main difference between E. rostrata and E. amphelioides is the of development of the exsert Si the former has the latter has degree rostrum whereas at all. 1> or prominent rostrum, reduced rostrum or none After reexamination of Atlantic specimens previously reported [Cairns, 1979] and additional specimens from the eastern Atlantic, off Hawaii, and off New Zealand, I am led to agree with Vaughan [1907] that there is a continuous intergradation between the extreme rostrate and nonrostrate forms, not only off Hawaii but also in the western Atlan- tic and off New Zealand. In fact, a single branch from an Albatross station 3827 specimen bears ex- tremely rostrate calices, nonrostrate calices, and some that are intermediate in development with only a slightly developed Sj. The development of the rostrum seems to be influenced by microen- vironmental changes and may be a reaction to a poor feeding area. The most extreme rostra are often those of calices which are adjacent to other corals or in close proximity to other corallites of the same colony. I agree with Vaughan [1907] that these differences should be designated as varietal or formae, not as separate species. It may be useful to refer to all specimens with a rostrum, or enlarged S^, as the typical form and to those without enlarged Sj^ as forma amphe- lioides, because some colonies have calices that uniformly lack rostra, whereas calices of other colonies all have rostra. E. rostra is differentiated from other Enallop- sammia by its unifacial calices. Material. Eltanin sta. 1411, USNM 47531; sta. 1981, USNM 47532; sta. 1983, USNM 47533. NZOI sta. C-527, USNM 47534. Specimens listed by Cairns [1979], USNM. Syntypes of A. rostrata; syntypes of Dendrophyllia amphelioides var. cucullata. Types. The syntypes of A. rostrata are deposited at the Museum of Comparative Zoology. Type- locality: 23?14'N, 82?25'W (Straits of Florida); 1472 m. The syntypes of D. (C.) amphelioides are deposited at the Zoologische Museum, Amsterdam. Type-locality: off Pulau Waigeo and Pulau Misool, Indonesia; 469-1633 m. The syntypes of D. amphelioides var. cucullata are deposited at the United States National Museum. Type-locality: off Hawaii; 426-679 m. Distribution. Widely distributed except for eastern Pacific: western Atlantic (San Pablo Sea- mount to Rio de Janeiro, Brazil), eastern Atlantic (area bounded by Celtic Sea, Azores, and Gulf of Guinea), Indian Ocean (off Maldive, off Nicobar Islands), western Pacific (off Japan, Indonesia), central Pacific (off Hawaii, lies Tuamotu), South Tasmanian Rise, Macquarie Ridge, Kermadec Ridge (Map 13). Depth range: 229-2165 m. 39. Enallopsammia sp. cf. marenzeller! Zibrowius, 1973 Plate 18, figs. 5, 6 Enallopsammia marenzelleri Zibrowius, 1973, pp. 49-51, pi. 1, figs^ 1-7, pi. 2, figs. 8-11; 1980, pp. 204, 205, pi. 106, figs. J-M. 58 BIOLOGY OF THE ANTARCTIC SEAS XI Plate 18. Enallopsammia, Balanophyllia, and GaryophyIlia jpsammia rostrata (PourtalSs): 1, 2, USNM 47532, Eltanin sta. 1981, dyed red; 3, 4, calices from other branches in same lot, both x5.1, 1-4. Enallop xO.85, dyed coated with ammonium chloride. 5, 6. Enallopsammia marenzelleri Zibrowius: 5, USNM 47535, Eltanin sta. 1411, x5.5, dyed red; 6, same specimen, xl.8. 7. Balanophyllia malouinensis Squires: USNM 47179, Eltanin sta. 339, H = 57.7 8, 9. Specimen identified as Caryophyllia clavus var. smithi by Moseley [1881]: BM 1880.11.25.27, Challenger sta. 308, GCD = 8.7 mm. CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 59 Discussion. Thirteen branch fragments, the largest bearing only 13 corallites, are tentatively assigned to this species. Branching bushy, calices on alternate sides of branches. Distinct costae present. Calices round, about 4-5 mm in diameter, without enlarged S^, or rostra. Otherwise, as- pects of septa and columella similar to those of previously described species. E. marenzelleri is a poorly known species, origi- nally described from only five colonies, two of which are equal to or smaller than these fragments. On the basis of the original description these fragments seem to belong to E. marenzelleri, but not enough material is present to be conclusive. This species is distinguished from the other two Recent species with alternately placed calices (E. pusilla, E. profunda) by its possession of costaeT Material. Eltanin sta. 1411 (13 fragments), USNM 47535. Types. The holotype and paratype of E. maren- zelleri are deposited at the Zoologische Museum, Amsterdam (Coel. 6902, 588). Type-locality: 5?56.5'S, 132''47.7'E (off Kepulauan Kai (Kei Islands)); 595 m. Distribution. Meteor Seamount; off Azores; off Nicobar Islands; off Kepulauan Kai (Kei Islands); Macquarie Ridge (Map 13). Depth range: 371-815 m. Uncertain Records Caryophyllia clavus var. smithi Broderip, 1828 Plate 18, figs. 8, 9 Caryophyllia clavus var. smithi; Moseley, 1881, p. 134 (part: Challenger sta. 308). Moseley reported this species from off southern Chile (Map 3) on the basis of three specimens, two of which were damaged (BM 1880.11.25.27). Complete specimen firmly attached to gorgonian stem and 8.7 X 7.6 mm in CD and 18.7 mm tall. Theca porcelan- eous, marked by thin, vertical striae, which delimit costae. Costal granules low and rounded. Septa hexamerally arranged in four cycles but missing two S4 for 46 septa. Sj^ and S2 equal in size and moderately exsert; S3 and S4 pro- gressively smaller and less exsert. Septal edges straight except for those of S3. Crown of eleven P3 enclosing elongate columella composed of eight narrow, twisted ribbons. These species are clearly not C. clavus var. smithi sensu Duncan, 1876 (actually C. smithi Broderip, 1828), thus far known only from the eastern Atlantic from Norway to the Congo [Zibrow- ius, 1980]. They also do not belong to any of the Caryophyllia discussed in this text. They might represent an undescribed species, but more speci- mens are needed for a complete description. Flabellum transversale conicum of F. flexuosum or F. impensum, both of which are known from this region. Flabellum ongulense Eguchi, 1965 Frabellum lurvatum; Niino, 1958, pi. 2, fig. 2 (misspellings). Flabellum ongulense Eguchi, 1965, pp. 11, 12, pi. 2, figs. 2a-2d. Eguchi described this species on the basis of one specimen from off Riiser-Larsen Peninsula (Cape Cook), Antarctica, at 750 m (Map 10). Because the hol?type is not available for study, it is not possible to add anything to his original descrip- tion. It is very similar to F. flexuosum but dif- ficult to verify from Eguchi's account. Desmophyllum pseudoseptatum Eguchi, 1965 Desmophyllum delicatum; Niino, 1958, pi. 2, fig. 3. Desmophyllum pseudoseptata [sic] Eguchi, 1965, p. 9, pi. 2, figs. 3a-3c. Eguchi described this species on the basis of one specimen from off Riiser-Larsen Peninsula (Cape Cook), Antarctica, at 630-680 m (Map 12). The holotype is not available for study, but Eguchi's illustrations and description very much resemble those of Javania antarctica (Gravier, 1914). Eguchi apparently intended to name this species D. pseudocostatum. ~ Yabe and Eguchi, 1942 Flabellum transversale conicum; Eguchi, 1965, p. 11, pi. 1, figs. 3a, 3b. Zoogeographic Analysis The Antarctic and Subantarctic regions have not been uniformly and thoroughly sampled, especially not the region from the Antarctic Peninsula to the Ross Sea and from the Ross Sea to the Weddell Sea. Furthermore, several scleractinian species from this area are known from only one or two records, some are doubtfully assigned, and records of sev- eral unique specimens have not been discussed in this paper pending the collection of additional specimens. Nonetheless, it is possible to make some generalizations about patterns of distribution and regional affinities on the basis of the USARP specimens and a reevaluation of previously reported specimens. For conformity, the zoogeographic divisions and terminology of Hedgpeth [1969] will be used in this paper (Map 14). The Antarctic region is the area inside the Antarctic convergence, including South Georgia, Bouvetrfya, and Heard Island. The Subantarctic boundary follows the subtropical con- vergence only partially and includes Magellanic South America, Falkland Islands, Tristan da Cunha Group, Gough Island, Prince Edward Islands, lies Crozet, lies Kerguelen, Macquarie Ridge, Campbell and Auckland islands, and several unnamed seamounts in the South Pacific. Areas directly to the north of the Subantarctic are referred to as cold tem- perate. Eguchi reported one specimen from off Riiser- Patterns of Distribution Larsen Peninsula (Cape Cook), Antarctica, at 920 m. Unfortunately, the specimen is not available Among the 37 species of Scleractinia known from for study. Judging from the illustration provided the Antarctic-Subantarctic region, certain patterns by Eguchi, the specimen could be a basal fragment of distribution occur (Table 2, last column), most BIOLOGY OF THE ANTARCTIC SEAS XI Map 14. Locator map indicating boundaries of Antarctic and Subantarctic regions [after Hedgpeth, 1969]. of which are shared with one other group or more of benthic invertebrates. The patterns are as follows: Antarctic region species A. Circumpolar or widespread in region, usually including Scotia Ridge but not crossing Antarctic convergence. B. Endemic to single localities. Subantarctic region species A. Primarily Magellanic. Campbell Plateau, Macquarie Ridge, and usually New Zealand and/or Australia. C. Circum-Subantarctic. D. Endemic to a seamount. Cosmopolitan or widespread species (with southern limit as given below) A. Subantarctic, but only marginally. B. Antarctic region, insular or seamounts. C. Antarctic region, continental. II. III. 6 Eight species are endemic to the Antarctic region, a pattern found in most other benthic invertebrate groups. Two of these, Caryophyllia eltaninae and Flabellum gardineri, are only known from the South Georgia-Shag Rocks area, and Balanophyllia chnous is known from only one record in the Ross Sea (pattern IB). Of the remaining five (pattern lA), four are circumpolar, including Caryophyllia antarctica, representing the ideal Antarctic distribution, including Bouvet0ya. The fifth species, Javania antarctica, is found primarily off western Antarctica and the Scotia Ridge. One of the circumpolar species, Flabellum impensum, was collected once north of the Antarctic convergence off the Antipodes Islands (Eltanin station 2143) at its second greatest recorded depth. This record is not so unexpected if one considers that the distribution of benthic shelf and slope animals does not necessarily follow the CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA .61 zoogeographic boundaries imposed by surface cur- rents [Squires, 1946c, p. 454; Kussakin as cited by Deli, 1972, p. U]. Instead, the charac- teristics of the water mass must be considered, which could be Antarctic bottom water or Antarctic intermediate water off the Antipodes Islands, depending on depth [Menzies et al., 1973, p. 201]. The temperatures for the Antarctic records of F. impensum are probably at or below 0?C, fully within the Antarctic bottom water mass. Ridgway [1968] indicated a bottom temperature of 3?-4?C for the Antipodes Islands locality, but it is an area of highly compressed benthic isotherms over short distances. Upwelling may have brought colder Ant- arctic bottom water closer to the surface here, or the species may have adapted to slightly warmer water at this latitude. Dawson [1970] has reported similar distributions for primarily Antarctic Asteroidea and Ophiuroidea, hypothesizing a pos- sible migration route from the Ross Sea to the Balleny Islands, then along the ridge system from the Balleny Islands to Macquarie Island, and then to the Campbell Plateau and New Zealand. Dawson concluded that this migration route is probably not a common one but that it may explain some of the echinoderm distributions; the Antipodes Islands record of F. impensum may result from a similar migration. Nine species are characteristic of the Magellanic subregion (pattern IIA), two ol which are known from only single records (Phyllangia fuegoensis and Balanophyllia sp.). Of the remainder, five are restricted to the eastern coast of South America. Of these, two cross over the Antarctic convergence to South Georgia, and three extend northward as far as Rio de la Plata. Sphenotrochus gardineri occurs on both sides of South America. The ninth species, Flabellum truncum, has been collected in the Magellanic subregion but also extends up the western coast as far as Peru. Two other species should be mentioned here: Bathelia Candida is more characteristic of the cold temper- ate region to the north but does extend into the Magellanic subregion, off both the east and the west coast of South America. A. rathbuni is known from the warm temperate and subtropical coasts of eastern South America and is known from the Sub- antarctic only as a subfossil at Tierra del Fuego. Three species (pattern IIB) are known from the Subantarctic islands of the Campbell Plateau and/ or off Macquarie Island as southernmost records of species normally found in more northerly, cold temperate regions, such as off New Zealand, the Chatham Rise, and off southern Australia. One species, Flabellum apertum (pattern IIC), is essen- tially circum-Subantarctic, and Cyathoceras irreg- ularis is endemic to a Subantarctic seamount (pat- tern IID). Another large component, including 11 species, does not fit any pattern but represents southern- most extensions of cosmopolitan or widespread species. Six of these species penetrate the Sub- antarctic but do not cross the Antarctic conver- gence (pattern IIIA). Of these, 5 occur off the Subantarctic islands south of New Zealand, and 1 (J^. cailleti) is found in the western Magellanic subregion^ Five cosmopolitan species cross the Antarctic convergence, 3 of them extending as far south as the Antarctic islands and seamounts (pattern IIIB) and 2 reaching the coast of Ant- arctica (pattern IIIC). One of the last group (Lcptopenus discus), however, might be considered an abyssal instead of a continental Antarctic species, its shallowest record being 2000 m. The remaining two species have unusual and dis- junct distributions. C. profunda is known from the southern cold temperate regions; from Sub- antarctic Tristan and Gough islands; and from one record from off Hugo Island, Palmer Archipelago. C. mabahithi is known from the northern Indian Ocean and, like C. prof unda, from one record off the Palmer Archipelago. A peculiarity of the Antarctic scleractinian distributions is the absence of the pattern common to most benthic invertebrate groups; that is, that species are distributed throughout, but endemic to, the Antarctic-Subantarctic region [see Dell, 1972, fig. 3]. The Antarctic convergence, despite the fact that intermediate and bottom waters cross freely and are little changed below it, does seem to form a boundary for many scleractinian distri- butions. The only exceptions are (1) C. profunda, (2) C. mabahithi, (3) four cosmopolitan species, (4) one Antarctic species (F. impensum) which crosses to the north, and (5) two Magellanic species which cross over to South Georgia. None- theless, the typical Antarctic-Subantarctic benthic invertebrate pattern is not achieved by any scler- actinian. Another characteristic of the Antarctic Scler- actinia is their unusually high percentage of cosmopolitan species. Even if the abyssal Leptopenus discus is not included, 3 cosmopolitan species (8% of the Antarctic-Subantarctic species) occur in the Antarctic region, and 11 (3U) occur as far as the Subantarctic region. This is consid- erably more than is the case for other benthic invertebrate groups. The group of Antarctic benthic invertebrates that most closely resembles the Scleractinia in distributional patterns is the Echinoidea [see Pawson, 1969]. This class has most of the distri- butional patterns described for Scleractinia and is one of the few groups with a limited number of species that cross the Antarctic convergence. The position of South Georgia as a transitional area is similar for both groups. Geographic Affinities Antarctic. Seventeen species of Scleractinia are known from the Antarctic region: eight ende- mic, five cosmopolitan, and two primarily Magel- lanic ones (which cross over to South Georgia) and two species with anomalous distributions (Caryo- phyllia profunda and C. mabahithi). The percent of endemic species is therefore 47%, close to Squires's [1969, p. 17] value of 50%, despite num- erous additions and synonymies. (For this compar- ison, Squires's Caryophyllia A is included as a nonendemic Antarctic species.) If the percent of endemic species is calculated in accordance with Cailleux's [1961] method, which would exclude the two Magellanic crossovers, this figure would in- crease to 53%. The scleractinian fauna of South Georgia and Shag Rocks (eight species) is a faunal mixture of Antarctic, Magellanic, endemic, and cosmopolitan species, typical of most other groups of benthic invertebrates [Dell, 1972, p. 7] and indicative of the mixed hydrologic conditions at these islands. 62 BIOLOGY OF THE ANTARCTIC SEAS XI e < fujaiqea uoTinq-rJisTQ (apTApxJOfi) qqdaa pB3adsapiM BTUBmsBi 'BT^BJjsnv il^nog mBpjaqsrav ail P"^ tnej-quiBS 311 BOfajv qjnos BXiqaAiiog ^Bxnsuiuad oxqDiBjuy uaaqsBa puB Bas liappaM ?Bas Tiapp3? oq B3S SSO^ gBas ssoa jBas ssoa oq Binsuxuaj OfqojBquv ujaqsafj spuBXSi puBxqaqs qqnos spuBxsi XauiiJO yqnos spuE-[SI qotnpuBS qqnos s>iooa SBIIS ' BTSaoao qqnos asT^ mBqqBqo puB puBXSl raBqqBqo puB^Baz ?aN ouojqBXd Xqunog pHBaqBxd XT3<1>'?3 gSqunoraBag oxqoJBquBqng _a8pTa aijBnbDBW spuBXSi paB?pa aouTjj puBXSj q3no3 puB dnoa3 Bqung Bp UBqsx^I g5]UBa poonpang 'spuBxsi puBx^ixBj BCSOSS-OS*/) BDTjauiv qqnos uaaqsBa B(S?e5-?6V) BDTjaniv qqnos uaaqsan (SoSV-oSe) B^^Jl^IlIV qqnos uaaqsBa (S?e*7-?5C) BOTjaray qqnos uaaqsan Xpnqs STqi UT passnasfa saToadg o <; u pa M M M <; M xSj Q MMM M I-H<;M pa M (O -(P^O CMi-(CO vO^* I loo t^.-ll.-lll inllOCNCN OlnOO^ Ol I'DI^OOC^OtNQO^tl I oooo I oor^omot^JlNc^?c^?mQOao^m^-^CNm^sl^O^HC7^cn^O M M X X K X M X X X ?H en .p^ 01 ? to , .--< < , -^ .?1 ?H tJ 0) ?H u CO r-l Cd CO ?H m ?iH ?H c8 Cd s: to CO CO g ki Cd u 0) ?n< c C cd ? 4-1 CO 13 c 4-1 ^ Cd 3 ?^ ?iH 4J ?f-4 0) C>0 o x: bO -D CO ?H M-4 Cd CO cu u 4J ?H T3 CO f-( M fO to 4J J ? ? ? ? ? fa fa eJ < ft* rt S o u u o o u o V3 <: CO o ^\ .-* 0(N.-4CN\D 0^.-t.-^.-^esICNor^oCNjcri'XJcnr^om lU X T3 X u c s O (U ?-l a 0) l-< o jcNc>jcnc<)cnrncoromncncn H -u ^o in e ^o tX) CO .-< ^ 1 fo a to c o j= o> ?r^ 4J to 4J 3 -H CO O 1 4J m Q ? 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