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 Material and Methods from the Antarctic and Subantarctic regions are described, mapped, and illustrated, including the This study is based on the examination of ap- description of 6 new species. Two more species, proximately 8700 specimens collected throughout one of them new, from waters closely adjacent to the Antarctic, Subantarctic, and adjacent waters the Subantarctic regions are also considered, as and includes new material from 482 lots collected well as 4 previously published records of Sclerac- at 310 stations. Most of the specimens were col- tinia of uncertain identity. A chronological, lected by the vessels participating in the United annotated list of all papers dealing with Antarctic States Antarctic Research Program (USNS Eltanin, Scleractinia is provided. A zoogeographic analysis now the ARA Islas Orcadas, and R/V Hero) and those describes common patterns of species distributions, participating in Operation Deep Freeze III and and the faunas of various geographic areas are IV. These specimens, as well as those from other discussed. Scleractinia from South Pacific sea- vessels (e.g., USCGC Eastwind, USFCS Albatross, and mounts, some of which may form deepwater coral R/V Anton Bruun), are deposited at the United banks, are particularly noted. 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- Introduction ited at the Zoologisches Institut und Zoologisches Museum, Hamburg), and a small collection of USNS This paper reviews the 37 species of Scleractinia Eltanin corals deposited at the Museum of Compara- known from the Antarctic and Subantarctic regions, tive Zoology, Harvard, Cambridge. as well as 4 records of uncertain identity from Previously reported specimens from the following these regions, and 2 more species from areas museums were examined: British Museum, London; closely adjacent to the Subantarctic region. De- Museum National d'Histoire Naturelle, Paris; Museum spite 28 papers dealing exclusively or partially fur Naturkunde an der Humboldt-Universltat Berlin; with Antarctic Scleractinia (Table 1), only about Institut Royal des Sciences Naturelles de Belgique, 1200 specimens have been reported previously from Brussels; the Australian Museum, Sydney; the South 114 localities. The material upon which this study Australian Museum, Adelaide; Museum of Comparative is based (primarily United States Antarctic Re- Zoology, Harvard, Cambridge; American Museum of search Program vessels. Deep Freeze vessels, and Natural History, New York; and the United States the R/V Vema) includes over 7 times the number of National Museum, Washington, D. C. Eguchi's [1965] specimens and almost 3 times the number of locali- specimens could not be found at the National In- ties reported in all the previous papers. stitute of Polar Etesearch, Tokyo, or the Tokyo Squires [1969] published a preliminary synthesis Kasei University. of the distribution of Antarctic Scleractinia but Synonymies are complete unless otherwise indi- never did write his intended Antarctic monograph, cated. thereby leaving many questions unanswered. Al- In the sections on material the numbers in pa- though his paper is valuable as a preliminary note, rentheses indicate the number of specimens in each it includes numerous undocumented range extensions lot. The number of specimens is not indicated for (all of his mapped records are undocumented), un- colonial species. Following this information, or explained statements about complex groups of the station number for colonial species, is an species, omissions of previous records, and refer- indication of where the specimen is deposited. If ence to an undescribed new species. Most, if not the vessel's cruise number is included, it precedes all, of Squires's [1969] specimens are deposited at the station number and is linked to it with a hy- the United States National Museum, Washington, D. phen. C, and at the American Museum of Natural His- Most of the holotypes and paratypes of the new tory, New York. This material, along with subse- species are deposited at the United States National quently collected United States Antarctic Research Museum; the remainder are at the British Museum. Program specimens and an examination of most of A confirmed depth range is used to avoid erro- the previously reported specimens, has allowed a neous end points resulting from bathymetrically documentation, and sometimes a correction, of wide-ranging trawls. The stated bathymetric range Squires's synthesis, a reevaluation of the system- for each species extends from the deepest shallow atics of the fauna, and a more meaningful zoogeo- to the shallowest deep component of the trawled graphic analysis of the fauna. 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 Vessel and/or Expedition Remarks Reference 3 Specimens of Flabellum thouarsii Milne Edwards and Du Petit-Thouars, from off Falkland Islands; Venus Haime [1848] deposited at MNHNP. HMS Challenger Preliminary report on Challenger Moseley [1876] Scleractinia, including records of 3 species from off Tristan da Cunha Group and Prince Edward Islands; deposited at EM. 2 specimens of Flabellum thouarsii SMS Gazelle Studer [1878] reported from off southern Argentina; specimens deposited at Museum fur Naturkunde, Berlin. Final report on Challenger HMS Challenger Moseley [1881] 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 SY Belgica, von Marenzeller stations off Peter I Island, Expedition Antarctique [1903] Antarctica; deposited at Institut Beige (1897-1899) Royal des Sciences Naturelles de Belgique, Brussels. 2 specimens of C. antarct ica from 1 Valdivia, Deutsche von Marenzeller station off Bouvet0ya; deposited Tiefsee-Expedition [1904a] at Museum fiir Naturkunde, Berlin. (1898-1899) 7 specimens of 3 species from 5 Gauss, Deutsche Pax [1910] stations off Gaussland, Antarctica; Siidpolar-Expedition histology of Flabellum impensum. (1901-1903) Scotia, Scottish Gardiner [1913] 3 specimens of CaryophyIlia profunda National Antarctic from 1 station off Gough Island. Expedition (1902-1904) Pourquoi-Pas? Gravier [1914a] 3 specimens of DesmophyHum DeuxiSme Expedition antarcticum ( < Javania antarctica) Antarctique Franqaise from 2 stations off Palmer Archipelago; deposited at MNHNP. (1908-1910) Pourquoi-Pas? Gravier [1914b] 13 specimens of 4 species from 3 Deuxi&me Expedition stations off Antarctic Peninsula; Antarctique Franijaise deposited at MNHNP. (1908-1910) Nimrod British Fossil Gardineria antarctica from David and Priestley Antarctic Expedition Mount Larsen; deposited at the [1914] (1907-1909) Australian Museum, Sydney. Terra Nova, British Gardiner [1929a] 19 specimens of 3 species from 4 Antarctic Expedition stations in Ross Sea; deposited at (1910) BM. Aurora Thomson and Rennet 7+ specimens of 3 species from 2 Australasian stations off Queen Mary Coast, [1931] Antarctic Expedition Antarctica. Another 2 scleractinian species from off Maria Island, Tas- (1911-1914) mania; all but F. australe are mis- identified; deposited at the Aus- tralian Museum, Sydney. RRS Discovery, RSS 280 specimens of 12 nominal species Gardiner [1939] William Scoresby from 33 stations off southern South America and Antarctic (1926-1933) Peninsula; 9 of 12 species reas- signed; deposited at BM. Umitaka-Maru, JARE 4 specimens of 4 species from 2 Niino [1958] stations off Riiser-Larsen Peninsula (Cape Cook), Antarctica; all species misidentified; deposition unknown. CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA TABLE 1. (continued) Vessel and/or Expedition Reference Remarks 97 specimens of 5 species from 10 Discovery, BANZARE Wells [1958] stations off coast of Antarctica (1929-1931) 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 R/V Vema (cruise 14) Squires [1961] 2 new species) from 6 stations off the Falkland Islands and Tierra del Fuego; desposited at AMNH. Review of fossil records of Gardineria Speden [1962] antarctica on Antarctic continent. 242 specimens of 5 species (including HMNZS Endeavour Squires [1962b] 2 new species) from 13 stations in (1958-1960) the Ross Sea; reviews distribution of all Antarctic Scleractinia; presumably deposited at NZOI. 2 fossilized specimens of 2 species United States Squires [1963a] collected at Tierra del Fuego; Exploring Expedition deposited at MCZ. (1838-1842) Discussion of research potential of Antarctic Squires [1964c] Scleractinia; no specimens reported. Specimens of 3 species from 1 station HMNZS Endeavour Squires [1965b] east of the Auckland Islands; (Macquarie Gap Crni sp) deposited at NZOI and USNM. 9 specimens of 9 nominal species Umitaka-Maru and Eguchi [1965] (including 2 new species) from 6 Soya, JARE 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 HMNZS Endeavour Bullivant [1967] assemblage in Ross Sea; specimens previously reported by Squires [1962b]. Reviews distribution of all Antarctic Squires [1969] 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 Ob, Academic Keller [1974] D/E stations from off southeastern South America Kurchatov and Falkland Islands; F. antarcticum misidentified; deposition unknown. Examination of microstructure by SEM miscellaneous Podoff [1976] of 6 Antarctic species; unpublished M. A. thesis; deposited at USNM. Sorauf and Podoff miscellaneous Examination of microstructure of 3 [1977] Antarctic species; deposited at USNM. 8700 specimens of 39 species USNS Eltanin; ARA Present study (including 7 new species) from 310 Islas Orcadas; R/V stations throughout Antarctic and Hero (USARP program) ; Sub-antarctic; primarily deposited USS Atka; USS Burton at USNM, also at AMNH and MCZ. 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 BIOLOGY OF THE ANTARCTIC SEAS XI Family MICRABACIIDAE Vaughan, 1905 Some specimens have been coated with dark dye and Leptopenus Moseley, 1881 recoated with a fine layer of ammonium chloride in order to improve their contrast for photography. L. sp. cf. L. discus Moseley, 1881 These specimens are noted in the plate legends. The following abbreviations are used in the text: Suborder FAVIINA Vaughan and Wells, 1943 Superfamily FAVIICAE Gregory, 1900 Vessels Family RHIZANGIIDAE d'Orbigny, 1851 USCGC Eastwind. EAD Astrangia Milne Edwards and Haime, 1848 EW USCGC Eastwind, cruise 66. USS Glacier, Deep Freeze IV GLD A. rathbuni Vaughan, 1906 Expedition. collected by the New Zealand NZOI Phyllangia Milne Edwards and Haime, 1848 Oceanographic Institute, including the HMNZS Endeavour, Rotoiti, and Viti. P. fuegoensis Squires, 1963 PD pebble dredge (used in conjunction with some R/V Family OCUHNIDAE Gray, 1847 Vema stations). Bathelia Moseley, 1881 United States Antarctic USARP Research Program. B. Candida Moseley, 1881 WH Walther Herwig. US RSS William Scoresby. Madrepora Linnaeus, 1758 Museums M. oculata Linnaeus, 1758 American Museum of Natural AMNH History, New York. Suborder CARYOPHYLLIINA Vaughan and Wells, 1943 BM British Museum (Natural Superfamily CARYOPHYLLIICAE Gray, 1847 History), London. Family CARYOPHYLLIIDAE Gray, 1847 MCZ Museum of Comparative Subfamily CARYOPHYLLIINAE Gray, 1847 Zoology, Harvard, Cambridge. CaryophyIlia Lamarck, 1801 NMNH see USNM. MNHNP Museum National d'Histoire C. antarctica Marenzeller, 1904 Naturelle, Paris. C. squiresi n. sp. NZOI New Zealand Oceanographic profunda Moseley, 1881 C. Institute, Wellington. C. eltaninae n. sp. SME Station Marine d'Endoume, C. mabahithi Gardiner and Waugh, 1938 Marseille (most of these specimens will be deposited Cyathoceras Moseley, 1881 at the Museum National d'Histoire Naturelle, Paris). C. irregularis n. sp. USNM United States National Museum, Smithsonian Aulocyathus Marenzeller, 1904 Institution, Washington, D. C. A. recidivus (Dennant, 1906) n. comb. ZIZM Zoologisches Institut und Zoologisches Museum, Hamburg. Subfamily TURBINOLIINAE Milne Edwards ZMA Zodlogische Museum, Amsterdam. and Haime, 1848 Other Sphenotrochus Milne Edwards and Haime, 1848 GCD greater calicular diameter. LCD lesser calicular diameter. S^. gardineri Squires, 1961 CD calicular diameter. PD pedicel diameter. Subfamily DESMOPHYLLIINAE Vaughan and Wells, 1943 H height. DesmophyHum Ehrenberg, 1834 S septa, costae, pali, or CS x' costosepta of cycle D. cristagalli Milne Edwards and Haime, 1848 designated by numerical forma cristagalli Milne Edwards and Haime, 1848 subscript. forma ingens Moseley, 1881 SEM scanning electron microscopy. forma capense Gardiner, 1904 Lophelia Milne Edwards and Haime, 1849 Checklist of Species Known From the Antarctic and Subantarctic L. prolifera (Pallas, 1766) Regions Subfamily PARASMILIINAE Vaughan and Wells, 1943 Order SCLERACTINIA Bourne, 1900 Solenosmilia Duncan, 1873 Suborder FUNGIINA Verrill, 1865 Family FUNGIIDAE Dana, 1846 £. variabilis Duncan, 1873 Fungiacyathus Sars, 1872 Goniocorella Yabe and Eguchi, 1932 F. marenzelleri (Vaughan, 1906) F. fragilis G. 0. Sars, 1872 G. dumosa (Alcock, 1902) 15 CAIRNS: ANTARCTIC AND SUBANTARCTIC SCLERACTINIA 332), pi. 11, figs. 1-5, 7.—von Marenzeller, Superfamily FLABELLICAE Bourne, 1905 1904b, p. 76.—Gravier, 1920, p. 97 (part).—? Family FLABELLIDAE Bourne, 1905 Eguchi, 1965, pp. 5-7, pi. 1, figs. 4a-4c. Flabellum Lesson, 1831 Fungia symmetrica; Duncan, 1873, p. 334, pi. 49, figs. 16-19. F. thouarsii Milne Edwards and Haime, 1848 Bathyactis marenzelleri Vaughan, 1906b, p. 66, F. areum n. sp. pi. 4, figs. 1, la, lb. F. curvatum Moseley, 1881 Fungiacyathus symmetricus; Wells, 1958, pp. 267, F. impensum Squires, 1962 pi. 2, figs. 1, 2.—Squires, 1962b, p. 13; 1969, F. flexuosum n. sp. p. 17, pi. 6, map 2. F. gardineri n. sp. Fungiacyathus marenzelleri; pi. 7, figs. A-K.— F. knoxi Ralph and Squires, 1962 F. apertum Moseley, 1876 Cairns, 1979, pp. 35-37, pi. 2, figs. 8, 9, pi. forma apertum Moseley, 1876 3, figs. 3, 8. Zibrovrtus, 1980, pp. 24, 25, pi. forma patagonichum Moseley, 1881 6, figs. A-M. F. truncum n. sp. Description. Base of corallum flat, center of Javania Duncan, 1876 base slightly raised. Largest specimen known measuring 40 mm across base; largest Antarctic J. cailleti (Duchassaing and Michelotti, 1864) specimen 38.5 mm in diameter. Forty-eight thin, J. anCarctica (Gravier, 1914) n. comb. ridged costae radiating from center of base, be- coming more raised and sinuous toward calicular Gardineria Vaughan, 1907 edge. Costae sometimes projecting as much as 1.5 mm from base at calicular edge. Base extremely G. antarctica Gardineria, 1929 thin and fragile, sometimes perforate, especially toward calicular edge. Five to seven synapticular plates occurring in every interseptal space, be- Family GUYNIIDAE Hickson, 1910 coming increasingly larger and more oblique toward Stenocyathus Pourtal&s, 1871 edge of corallum. Septa hexamerally arranged in four cycles. '1 largest septa and only ones reaching columella S. vermiformis (PourtalSs, 1868) without fusion to other septa. Septa of remaining cycles progressively smaller. Each S2 reaching Suborder DENDROPHYLLIINA, Vaughan and Wells, 1943 columella, there joined by pair of S3,all loosely Family DENDROPHYLLIIDAE Gray, 1847 fused or covered over by triangular canopy composed Balanophyllia Wood, 1844 of thin calcareous plate. Pairs of S^ fused to S3 by similar but larger and higher canopies, B. malouinensis Squires, 1961 these extending from S2-S3 canopies to halfway B. sp. to calicular edge. These canopies often perforate. B. chnous Squires, 1962 Septa laterally carinate, Sj^ possessing about 7-10 carinae, or about 1 every 1.9 mm. Carinae Enallopsammia Michelotti, 1871 about 0.4 mm high and usually symmetrical on both sides of septum. Near columella, carinae more E. rostrata (Pourtalfes, 1878) closely spaced and corresponding to high septal E. sp. cf. E. marenzelleri Zibrowius, 1973 spines. Carinae vertical near columella, oblique midway between columella and calicular edge and Uncertain Records almost horizontal near calicular edge. Most car- inae extending from septal edge to base; some Caryophyllia clavus var. smithi sensu Moseley, 1881 shorter, extending only halfway to base; some Flabellum transversale conicum sensu Eguchi, 1965 branching from other carinae. Near base most car- Flabellum ongulense Eguchi, 1965 inae curving toward columella, often degenerating Desmophyllum pseudoseptatum Eguchi, 1965 into rows of granules. If still solid, carinae may be confused with synapticulae but can usually be distinguished by their more oblique orientation, Species Account often intersecting synapticulae at an acute angle. Septa bearing elongate, slender spines near colu- Order SCLERACTINIA Bourne, 1900 mella but becoming less serrate and usually lobate Suborder FUNGIINA Verrill, 1865 in profile toward calicular edge. Height of exsert Family FUNGIIDAE Dana, 1846 lobes up to 10 mm above base. All septa but S^ Genus Fungiacyathus Sars, 1872 bearing lobes, these lobes damaged in most speci- mens. Septa extraordinarily fragile and specimens Diagnosis. Solitary, cupolate, free. Septotheca rarely collected fully intact. Septal edges thin; costae thin and spinose. Septa irregularly straight to irregularly sinuous. Columella vari- dentate, laterally braced by thin ribbons extend- able in size, composed of loose fusion of inner ing from septotheca and by thin septa! striae. septal spines and additional trabeculae. Columella feeble. Paliform lobes sometimes pres- Discussion. F. marenzelleri was frequently ent. Type-species: Fungiacyathus fragilis Sars. reported as Bathyactis or F. symmetricus by earlier 1872, by monotypy. authors, probably because of Moseley's [1881] as- sumption that all small specimens were simply ju- 1. Fungiacyathus marenzelleri (Vaughan, 1906) veniles of a species with a larger adult corallum. Plate 1, figs. 1, 2, 8 F. symmetricus has subsequently been shown to be Indemic to the western Atlantic (183-1644 m) and Bathyactis symmetrica; Moseley, 1881, pp. 186-190 to be rarely larger than 14 mm in CD [Cairns, (part: Challenger sta. 133, 147, 157, 299, 325. BIOLOGY OF THE ANTARCTIC SEAS XI Plate 1. Fungiacycathus 1, 2, 8. 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. 3-7. 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 35 1916, fragilis; Jungersen, Fungiocyathus cya 1979]. F. marenzelleri is distinguished from (part).~Broch, 1927, p. 8.~Nordgaard, 1929, p F. symmetricus by its much larger corallum, broad- 103. er distribution, and greater depth range. F. symmetricus also has higher septal spines and a more solid, well-defined base. Description. Base of corallum flat, very thin, Specimens from four localities (Eltanin stations sometimes irregularly perforate. Evidence of re- 138, 426, 993, and 1545) differ from typical F. generation from fragments of corallum. Largest marenzelleri by having a much more crowded ar- specimen reported 45 mm in basal diameter; largest rangement of septal carinae (about 20 per septum, specimen from Subantarctic 25 mm in diameter. or 1 every 0.8 mm) and more spinose septal faces. Slightly ridged C1-3 radiate from center of base. These specimens, all from the South Shetland Septa hexamerally arranged in five complete cy- Islands and east of South Orkney Islands, may cles, fifth cycle appearing at CD of 9-10 mm. represent a new species, but until the variation Septal arrangement similar to previously described of F. marenzelleri is better understood, they are species: independent; inner edges of septa S^ assigned to this species. Eguchi's [1965] ^. of remaining cycles fused to one another by thin, symmetrica also seems to belong to this form. perforate triangular lamellae (canopies). Each When the deeper records of Fungiacyathus spp. larger septum forming large nonserrate lobe for are reexamined [e.g., Moseley, 1881; Gardiner and most of its length, with few, if any, projecting Waugh, 1939; Keller, 1976], more synonyms for F. spines near columella. Height of septal lobes up marenzelleri may result, and its geographic range to one fifth to one fourth of CD. These lobes, as may thus be increased. All records deeper than well as all septa, with highly sinuous outer and 1800 m should be reevaluated for this possibility. upper edges, corresponding to septal corrugations. This species may eventually be found to be the most widespread and deepest-living species of Corrugations vertical near upper edge, becoming more horizontal as they curve toward columella scleractinian coral. near base. About 12 corrugations per septum, or Material. Eltanin sta. 13 (5), USNM 47470; sta. 1/mm, giving septa 'wrinkled' aspect. Crests of 18 (23), USNM 47466; sta. 20-134 (5), USNM 47476; corrugations regularly spaced, usually rounded and sta. 138 (3), USNM 47475; sta. 353 (1), USNM 47471; smooth, but may bear row of low, pointed granules sta. 426 (8), USNM 45673; sta. 993 (1), USNM 47473; or may even be slightly carinate. Small, pointed sta. 997 (6), USNM 47474; sta. 1148 (1), USNM granules usually on all septa near columella. 47469; sta. 1545 (3), USNM 47477; sta. 1957 (1), Septa extraordinarily fragile. All septa joined USNM 47666. Hero sta. 721-1081 (4), USNM 47468. to adjacent septa by synapticular plates, these Glacier sta. 11 (6), USNM 47465. Albatross sta. plates increasing in size toward calicular edge. 4397 (1), USNM 47467. Specimen identified as F. About 7-10 plates occurring per centimeter, con- symmetricus by Wells [1958], South Australian tinuing to add on as corallum increases in dia- Museum H 70; specimens listed by Cairns [1979], meter. USNM. Types of B. marenzelleri. Columella round and small, sometimes a solid, Types. The holotype of F. marenzelleri is de- horizontal lamella but usually a loose fusion of posited at the United States National Museum inner edges of larger septa. (47415); three paratypes are at the Museum of Com- Discussion. There are four other nominal Recent parative Zoology. Type-locality: 8°07.5'S, species of Fungiacyathus with five cycles of septa. 104°10.5'W (off Peru); 3820 m. F. fragilis differs from western Atlantic JF. pus- Distribution. Widely distributed throughout At- illus (Pourtal&s, 1868) in being larger and having lantic Ocean as far north as Greenland; eastern sinuous septa, from Indian Ocean F. stephanus (Al- Pacific; circum-Subantarctic (including off lies cock, 1893) in having a flat base and lower septa, Crozet); off South Shetland Islands; east of South and from Indian Ocean and eastern Pacific E|. pali- Orkney Islands; WeddeU Sea; ? off Prince Harald ferus (Alcock, 1902) in lacking paliform lobes. Coast and Enderby Land, Antarctica (Map 1). Depth It is indistinguishable from hawaiiensis F_. range: 300-5870 m. (Vaughan, 1907); however, comparisons Involving There is a direct relationship between depth of only one specimen (the holotype) cannot be con- occurrence and proximity to the Antarctic, the clusive. F. kikaiensis (Yabe and Eguchi, 1942), more southerly records being shallower. The shal- also with five cycles, is from the Pliocene- lowest records of this species (300-500 m) are Pleistocene of Japan. represented by the four continental Antarctic Material. Eltanin sta. 1412 (5+), USNM 47536; records; the four records from the South Shetland sta. 1846 (3), USNM 47537. Specimens of Verrill Islands range from 300 to 1435 m. No other record [1882], (Yale Peabody Museum, New Haven) YPM 8322 is shallower than 1805 m. and USNM 47538-47539; specimens of Zibrowius [1980], SME. Holotype of B. hawaiiensis. 2. Fungiacyathus fragilis G. 0. Sars, 1872 Types. One syntype of F. fragilis is deposited Plate 1, figs. 3-7 at the Oslo Museum (B626). Type-locality: ' Skraaven in Lofoten'; 549 m. The holotype of B. Fungiacyathus fragilis M. Sars, 1869, pp. 250, hawaiiensis is deposited at the United States 265, 274 (nomen nudum).—G. 0. Sars, 1872; p. National Museum (20834). Type-locality: between 58, pi. 5, figs. 24-32. Cairns, 1979, p. 206. Oahu and Kauai islands, Hawaii; 1761-2056 m. Distribution. Eastern Atlantic in area bordered —Zibrowius, 1980, pp. 23, 24, pi. 5, figs. A-J. by Norway, Cape Verde Islands, and the Azores; off Bathyactis symmetrica; Verrill, 1882, p. 313; 1883, Massachusetts; off Hawaii; west of South Island, p. 65.—Gravier, 1920, p. 97 (part).—Thomson, New Zealand; Macquarie Ridge (Map 1). Worldwide 1931. p. 9. depth range: 285-2200 m; New Zealand-Macquarie Bathyactis hawaiiensis Vaughan, 1907, pp. 145, 146, records: 1647-1693 m. pi. 27, figs. 1, la. BIOLOGY OF THE ANTARCTIC SEAS XI Plate 2. Leptopenus, Astrangia, and Phyllangia Leptopenus sp. cf. L. discus Moseley: 1, USNM 47481, Eltanln sta. 1545, CD 1-3. = 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 Astrangia rathbuni Vaughan: 4, 4-6. USNM 10974, CD = 6-7 mm. 2520, CD = 5-6 mm; 5, holotype holotypic colony and calices, MCZ 5390, CD = Phyllangia fuegoensis Squires: 7-9. 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). Leptonemus discus; Agassiz, 1888, p. 154, fig. 479 Family MICRABACIIDAE Vaughan, 1905 (misspelling). Genus Leptopenus Moseley, 1881 Description. Corallum discoidal and extremely Diagnosis. Solitary, discoidal, free. No wall, fragile. Base of corallum flat to slightly con- costae alternating in position with septa. Costae cave, especially near center. Largest Antarctic and septa united by simple synapticulae producing specimen 18.0 mm in CD. very porous, delicate corallum. Columella trabec- Septa hexamerally arranged in two cycles and ular. Type-species: Leptopenus discus Moseley, multiple bifurcation of septa of third cycle, un- 1881, by subsequent designation [Wells, 1936]. like species from any other family of Scleractinia. (The traditional method of designating septa of 3. Leptopenus sp. cf. L. discus Moseley, 1881 higher cycles as pairs of new septa flanking each Plate 2, figs. 1-3 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- Leptopenus discus Moseley, 1881, pp. 205-208, pi. furcating about 2 mm from columella, the two resul- 14, figs. 1-4, pi. 16, figs. 1-7.—Not L. discus ting septa being referred to as S3'. Not far Dennant, 1906, p. 162 (? Letepsammia sp.).— from this junction the S3' bifurcate, each form- Wells, 1958, p. 262.—Squires, 1965a, pp. 878, ing two S3''. Sometimes, near edge of calice, 879, fig. 1: 1967. p. 505; 1969, p. 17, pi. pair of S3''' forming, as described by Moseley 6, map 2.—Keller, 1977, p. 37, fig. 1.—Cairns, for syntypes. Branching of S3 not always symme- 1979, pp. 37, 38, pi. 3, figs. 4-7. 10 BIOLOGY OF THE ANTARCTIC SEAS XI 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 Fig. 1. Diagraimnatic representation of one system spines. It is easily distinguished from L. hypo- of Leptopenus discus. Heavy lines represent septa, coelus Moseley, 1881, by its lesser height and light lines costae. smaller S2 spines and from L. solidus Keller, 1977, by its alternating septa and costae. Material. Eltanin sta. 598 (1), USNM 47480; trical within one system or consistent in systems sta. 1545 (2), USNM 47481; sta. 1926 (1), USNM of one corallum. (For instance, in a half system 47479; sta. 2002 (3), USNM 47483; sta. 2108 (37), of a specimen from Eltanin station 2002, one S3' USNM 47482. Specimen of Agassiz (1888], USNM 46916. produced only two S3'', whereas the other S3' Syntype from Challenger sta. 147. produced two S3'' and four S3'''. Also, one Types. The four syntypes of L. discus, collected of the syntypes [Moseley, 1881, pi. 14, fig. 1] is at Challenger stations 147, 157, and 323, are de- illustrated as having a pair of S3' ' ' ' in the posited at the British Museum. The specimen from upper right system. The regular arrangement of 72 Challenger station 147 is numbered 1880.11.25.159. septa with one pair of S3''' per system to which Type-locality: since a lectotype has not been Moseley alluded is probably just one variation of designated, the type-localities are off Rio de la septal arrangement.) only independent septa, extending to col- Plata, South America, and Subantarctic Indian $1 Ocean; 2926-3566 m. umella and bearing eight or nine long, recurved spines. S2 also extending to columella and bear- Distribution. Off Cuba; off Rio de la Plata; Makassar Strait, Indonesia; off South Orkney Is- ing five or six similar spines, with pair of S3 joining each S2 close to columella S3 and all lands; off South Sandwich Islands; Subantarctic of its bifurcations bearing similar recurved spines Indian Ocean (including off lies Crozet); Ross Sea decreasing in size toward calicular edge. Thin, (Map 2). Worldwide depth range: 2000-3566 m; triangular canopy illy at point of junction of Antarctic records: 2035-2384 m. USL S3 to S2 and at every bifurcation of S3. Septal spines of S2 just distal of S2-S3 Suborder FAVIINA Vaughan and Wells, 1943 junction usually tallest spines on corallum, rising Superfamily FAVIICAE Gregory, 1900 about 1 Iran above columellar spines. Septa very Family RHIZANGIIDAE d'Orbigny, 1851 low and solid, composed mainly of large spines Genus Astrangia Milne Edwards and Haime, 1848 united by thin lamella. No septal granulation. Columella a very spiny mound in center of calice, Diagnosis. Colonial, extratentacular budding merging with inner septal spines. forming cerioid, plocoid, or reptoid coralla. Base of corallum consisting of radiating network Corallites united basally by thin coenosteura or of bifurcating costae, these costae alternating- in position with overlaying septa. Costae not ar- stolons. Septa dentate; columella papillose. ranged like septa, because all costae bifurcate; Type-species: Astrangia michelini Milne Edwards none independent (Text figure 1). Each costa at- and Haime, 1848, by subsequent designation [Milne tached to its two adjacent septa by thin synaptic- Edwards and Haime, 1850]. ular bridges; space between bridges forming el- liptical pores. Viewed from above or below, 4- Astrangia rathbuni Vaughan, 1906 corallum appearing regularly perforate. Pores Plate 2, figs. 4-6 increasing in size from center to calicular edge. For inner two thirds of corallum, synapticular 7 Astrangia Verrill, 1869, p. 526. bridges reaching upward from costae to meet septa; Astrangia rathbuni Vaughan, 1906a, pp. 849-850, p. in outer part of corallum, however, septa becoming 78, figs. 1-3.—Squires, 1963a, pp. 10, 11, figs. rudimentary or absent and synapticular bridges 3-7.—Laborel, 1971, pp. 200, 201, pi. 6, fig. 1, becoming wider and horizontal, but pores still map 7.—Zibrowius, 1974c, pp. 165, 166.—Not A.