Notes on Indo-Pacific Scleractinian Corals. Part 10.1 Late Pleistocene

Notes on Indo-Pacific Scleractinian Corals. Part 10. 1 Late Pleistocene Ahermatypic Corals from Vanuatu2

JOHN W. WELLS 3

THE OCCURRENCE OF A DEEP-WATER inver Pertinent to this briefstudy is the collection tebrate fauna of late Pleistocene age of recent Australian ahermatypes in the Aus (25,280 ± 400 yrs. B.P.) on the island ofSanto, tralian Institute of Marine Science (AIMS), Vanuatu (formerly New Hebrides), has been briefly examined by the writer in 1982. In it briefly described by H. S. Ladd (1975, 1976, were found, inter alia, specimens of a new 1982). The richly fossiliferous unlithified genus, Bourneotrochus, identical to some puz sands and silts on the Kere and Navaka rivers zling fossil examples from Vanuatu. Thanks were first made known by the geologists ofthe are due to J. E. N. Veron of AIMS for per New Hebrides Geological Survey (Mallick mission to describe these. 1971; Mallick and Greenbaum 1975). Subse Types and figured specimens are deposited quent collections were made by members of in the National Museum of Natural History the United States Geological Survey (USGs) (USNM), Washington. and the National Museum ofNatural History, Washington (USNM). The rich fauna of non reefcorals was made available to the writer for FAMILY FUNGIIDAE DANA study by the late H. S. Ladd. GENUS Diaseris MILNE EDWARDS & HAIME The ahermatypic coral fauna of 16 genera and 19 species is one typical of sandy or silty Diaseris distorta (Michelin, 1842) bottoms in moderately deep water (200 m) and consists mainly of small free-living Fungia distorta Michelin 1842, p. 316; 1843, caryophyllids and flabellids. Only five species pI. 5. are attached forms, fixed to shells, bryozoans, Diaseris distorta M. E. & H. 1851, p. 118; or octocoral axes. The faunal facies is that of 1860, p. 35, pI. D12, fig. 4. contemporary calcareous sands and silts such ?Diaseris distorta Umbgrove 1946, p. 533, as off the eastern and southern Australian pI. 81, figs. 3, 4. coasts, Hawaii, Philippines, and southern Diaseris distorta Veron & Pichon 1980, Japan, and that of the Pleistocene deposits of p. 121, figs. 194-196 (synonymy). the Ryukyu Islands (Yabe and Eguchi 1932b), MATERIAL: Two worn specimens, disasso Shortland Islands (Guppy 1885), Philippines ciated fragments. (Eguchi 1941), and Java (Umbgrove 1950). All of the 19 species except two (Deltocyathus OCCURRENCE: Kere River: USGS Sta. heteroclitus and Flabellum vanuatu) are extant 25718. Recent: Red Sea eastward through forms of the western Pacific. central Pacific to Hawaii; Japan; Galapagos (?). lO-IOOm. Pliocene, Java (Umbgrove).

1 Numbers 1-9 ofthese notes were published in Pacific Science as nos. I and 2, vol. 13, pp. 286-290, 1955; no. 3, GENUS Fungiacyathus SARS vol. 15, pp. 189-191, 1961; no. 4, vol. 20, pp. 203-205, 1966; nos. 5 and 6, vol. 22, pp. 274-276, 1968; no. 7, vol. 25, pp. 368-371, 1971; no. 8, vol. 26, pp. 183-190, 1972; Fungiacyathus fragilis Sars no. 9, vol. 36, pp. 211-219, 1982. Figure 1, 1, 2 2 Manuscript accepted 1 March 1984. 3 Cornell University, Department of Geological Fungiacyathus fragilis G. O. Sars 1872, Sciences, Ithaca, New York 14853. p. 58, pI. 5, figs. 24-32. 205 206 PACIFIC SCIENCE, Volume 38, July 1984

5 3 4

9 6 7

FIGURE 1. 1,2, Fungiacyathus jragilis, USNM 71837, x 8, x 8, Navaka R., USGS Sta. 24918; 3, 4, F. sp. cf. F. stephanus, USNM 71838, x 4, x 4, Kere R., USGS Sta. 25715; 5, 6, Stephanophyllia japonica, USNM 71839, x 4, x 4, Navaka R., USGS Sta. 24918; 7,8, Madrepora porcellana, USNM 71840, x 4, x 8, Kere R., USGS Sta. 25715; 9, M. porcellana, USNM 71841, x 4, Navaka R., USGS Sta. 24918. Indo-Pacific Scleractinian Corals. Part IO-WELLs 207

Fungiacyathus fragilis Zibrowius 1980, Stephanophyllia japonica Yabe & Eguchi p. 23, pI. 5. 1934, p. 281, figs. 1-3; 1942a, p. 156, pI. Fungiacyathus fragilis Cairns 1982, p. 9, 13, figs. 8a-c. pI. 1, figs. 3-7 (synonymy). MATERIAL: Two specimens, the larger Bathyactis kikaiensis Yabe & Eguchi 1942a, p. 155, pI. 12, figs. 6, 7. (d = 8 mm) well preserved, with five complete septal cycles, the base nearly flat with equal MATERIAL: Twenty-four examples, ranging beaded costae. in diameter from 2 to 11 mm. The septa in OCCURRENCE: Navaka River, USGS Sta. specimens up to cd = 8.5 mm are relatively 24918. Plio-Pleistocene: Ryukyu Islands; thick and strongly spinose laterally, the smallest with only three cycles. One, Kagoshima, Japan. cd = 2.5 mm, has four cycles with primary septa strongly crested. At cd = 4 mm the FAMILY OCULINIDAE GRAY "canopied" condition of the junctions of the GENUS Madrepora LINNAEUS second and third cycles has developed. At cd = 6 mm the 48 fifth cycle septa are present. Madrepora porcellana (Moseley) At cd = l4mm the septa are proportionally thinner but thickly spinose laterally. Figure 1, 7,8,9 OCCURRENCE: Navaka River: USGS Sta. Neohelia porcellana Moseley 1881, p. 176, 24918 (17 specimens); Kere River: USGS Sta. pI. 10, figs. 7, 7a. 25718 (7 specimens). Plio-Pleistocene: Japan. Neohelia porcellana Pratt 1900, p. 591, pIs. Recent: Atlantic, Hawaii, New Zealand, 62,63. Antarctic (Macquarie Ridge). 285-2200m. Neohelia porcellana Hickson 1903, p. 344. MATERIAL: Several more or less fragmen Fungiacyathus sp. cf. F. stephanus (Alcock) tary pieces of small colonies of this species seem to be distinct from the cosmopolitan Figurel, 3, 4 Madrepora oculata (Linnaeus). The calices are Bathyactis stephana Alcock 1893, p. 14, very small (1-1.5 mm) with generally fewer pI. 5, figs. 12, 12a. septa. Bathyactis stephana Alcock 1898, p. 28, pI. 3, figs. 5, 5a. OCCURRENCE: Kere River: USGS Stas. Bathyactis stephana Alcock 1902, p. 38. 25718 and 25715; Navaka River: USGS Sta. Fungiacyathus stephanus Gardiner & 24918. Recent: off Api Island, Vanuatu (Chal Waugh 1939, p. 252. lenger Sta. 177, 115m [type]); Lifu, Loyalty Island (Pratt gives no locality but most of MATERIAL: A fragment representing one Willey's corals came from Sandal Bay, Lifu); quarter of a flat-based corallum (cd = 12 mm northwestern Australia, 140-141 m, 20.7°C. est.) with five complete cycles of septa, of which those ofthe first two or three cycles rise as tall, smooth-margined folia, is referred to FAMILY CARYOPHYLLIIDAE DANA this species. GENUS Caryophyllia LAMARCK OCCURRENCE: Kere River: USGS Sta. 25715. Recent: Bay of Bengal, Macassar Caryophyllia scobinosa Alcock 1902 Straits, Natal. (34)-1240 m. Figure 2, 1-4 Caryophyllia scobinosa Alcock 1902, p. 8, FAMILY MICRABACIIDAE VAUGHAN pI. 1, figs. 1, 2a. Caryophyllia scobinosa Yabe & Eguchi GENUS Stephanophyllia MICHELIN 1942a, p. 119, pI. 10, figs. 4, 5 (syn onymy). Stephanophyllia japonica Yabe & Eguchi Caryophyllia scobinosa Gardiner & Waugh Figure 1,5,6 1938, p. 177, pI. 3, fig. 7. 208 PACIFIC SCIENCE, Volume 38, July 1984

8 7

FIGURE 2. 1,2, Caryophyllia scobinosa, USNM 71842, x 1, xl, Kere R., USGS Sta. 25715; 3, 4, C. scobinosa, USNM 71843, x 1.5, x 1.5, Kere R., USGS Sta. 25715; 5, 6, Acanihocyathus grayi, USNM 71844, x I, x 1.5, Kere R. USGS Sta. 25715; 7,8, A. grayi, USNM 71845, x I, x 1.5, Kere R., Sta. SM242; 9, A. grayi, USNM 71846, x I, Kere R., USGS Sta. 25715; 10, 11, Stephanocyathus (Acinocyathus) spiniger, USNM 71847, x 1.5, x 1.5, Navaka R., USGS Sta. 24918; 12, 13, A. (C.) spiniger, USNM 71848, x 4, x 4, Navaka R., USGS Sta. 24918. Indo-Pacific Scleractinian Corals. Part 10-WELLS 209

MATERIAL: Fifteen conical to cornute This subgenus is proposed to include coralla attached to bits of shell or sand, rang species of Stephanocyathus with six elongate, ing in size from cd = 8 x 9mm, h = 15mm, slender basal spines corresponding to the cos to cd = 15 x 19m, h = 25mm. Costae low, tae of the six primary septa. Stephanotrochus rounded, finely granulate. Specimens less than (Stephanocyathus) lacks basal spines; S. cd = 9 x 10mm have only four complete (Odontocyathus) has 12 short, blunt or ragged cycles of septa, regularly arranged. In larger spines corresponding to the 12 primary and examples fifth cycle septa occur irregularly in secondary costae. Other species of S. (Acin many of the systems. Columella of curled ocyathus), some of which are probably syn laths. onyms of S. (A.) spiniger are: Trochocyathus? mantelli Milne Edwards & Haime 1857 OCCURRENCE: Kere River, USGS Sta. (Oligocene, New Zealand), Stephanotrochus 25715, Sta. SM 242. Recent: western Indian tatei Dennant 1899 (0. Oligocene, Victoria, Ocean, 786-805 m; Indonesia, 535-794 m; Australia), Odontocyathus coloradus Smith Japan, 119-522m. 1913 (Miocene/Pliocene, Nasbate 1., Philip pines), Odontocyathus japonicus Yabe & GENUS Acanthocyathus MILNE EDWARDS & Eguchi 1932 (Neogene, Japan), Odonto HAIME cyathus stella and O. sexradiis Alcock 1902 Acanthocyathus grayi MILNE EDWARDS & (Recent, Indonesia). HAIME ETYMOLOGY: akaina (Greek = spine). Figure 2, 5-9 Acanthocyathus grayi M. E. & H. 1848, Stephanocyathus (Acinocyathus) spiniger p. 293, pI. 9, figs. 2, 2a. Marenzeller Acanthocyathus grayi Yabe & Eguchi 1941, p. 211,figs. 3a, b. Figure 2,10-13 Acanthoc,y(Jthus grayi Umbgrove 1950, Stephanotrochus spiniger Marenzeller 1888, p. 641,'figs. 27-32 (synonymy). p.20. Acanthocyathus grayi Eguchi & Miyawaki Odontocyathus spiniger Eguchi 1968, p. 1975, p. 57. C39, pI. C23, figs. 1, 2; pI. C26, figs. 12-14 (synonymy). MATERIAL: Twenty-one typical specimens Stephanocyathus (Odontocyathus) spiniger with calices ranging from 6 x 10 mm to Eguchi & Miyawaki 1975, p. 57. 16 x 23 mm. In larger calices there are from 58 to 65 septa in four complete cycles plus MATERIAL: One worn specimen (cd = 10-17 septa of the fifth. One example has two 20 mm, h = 10 mm) with the tips of the six short spines on one flank, and five on one end horizontal spines broken off. A very small and three on the other. Most specimens are individual (Figure 2, 12, 13; cd = 3 mm) tinted a pale red brown, deeper in shade to from the same locality strongly resembles ward the calice. Aplocyathus armatus (Michelotti, 1838) ofthe Oligocene and Oligocene ofEurope, examples OCCURRENCE: Kere River: USGS Sta. of which are characterized by an apparent 25715; Sta. SM 242. Recent: off coast of pentameral symmetry due to reduction ofone Burma; Andamans; Indonesia; Sulu Sea; ofthe six normal systems, with suppression of Japan. 37-490m. one spine. In a recent specimen from the Philippines (cd = 20mm) the spines are very long, gently GENUS Stephanocyathus SEGUENZA tapering, with a spread of about 50 mm. SUBGENUS Acinocyathus N.S.GEN. OCCURRENCE: Navaka River, USGS Sta. TYPE SPECIES: Stephanotrochus spiniger 24918. Neogene: Philippines, Japan. Recent: Marenzeller 1888. Recent, Japan. Philippines, Indonesia, Japan. 120-560 m. 210 PACIFIC SCIENCE, Volume 38, July 1984

GENUS Heterocyathus MILNE EDWARDS & S4 is the largest septum in each system, thick HAIME ening at the margin where it continues as one of the six blunt spines. The other S3 in each Heterocyathus aequicostatus Milne Edwards system lacks a pair of S4. The columella is & Haime formed by the fused inner ends of Sl and S2. Figure 4,1 MATERIAL: Six specimens, all from USGS Heterocyathus aequicostatus M. E. & H. Sta. 24918: five paratypes with cd's 2.5 mm 1848, p. 321, pI. 10, fig. 8. (circular, no spines), 2.5 mm (small spines), Heterocyathus aequicostatus Eguchi 1968, 3.0mm, 3.7mm, 3.8mm, and holotype 4mm. p. C30 (but not pI. C28, fig. 1; pI. C29, figs. REMARKS: The curious disposition of the 8, 9) (synonymy). septa in each of the systems distinguishes this Heterocyathus aequicostatus Folkeson from other recent and fossil species of Delto 1919, p. 8, pI. 1, figs. 4-9. cyathus. Two other calcarate species are (1) MATERIAL: One specimen, cd = 7 mm, D. calcar Pourtales of the western Atlantic covering part ofthe shell ofa small Conus, and (Cairns 1979) with spines extending from the two worn examples. Sl septa, and (2) D. ornatus Gardiner 1899 from the Loyalty Islands, which bears 12 OCCURRENCE: Kere River, USGS Sta. spines extending from the unthickened S3 25718. Navaka River: USGS Sta. 24918. Neo septa. Specimens of the latter from off the gene: Japan, Ryukyus, Taiwan, Philippines, southern Great Barrier Reefs, Queensland, in Indonesia. Recent: western Pacific, Indian the collections at the Australian Institute of Ocean, Persian Gulf, Red Sea, South Africa. Marine Science show clearly 24 S4 attached in 10-658m. pairs to the 12 spiniferous S3. In the present form the Sl are small, the S2 are not much larger, but one of the S3 in each system is GENUS Deltocyathus MILNE EDWARDS & thickened beyond the junction with the pair of HAIME S4 and continues into a prominent short, blunt spine. Deltocyathus heteroclitus n. sp. ETYMOLOGY: heteroclitus, Latin, differing Figure 3, 1-6 from the norm. DESCRIPTION: Corallum solitary, free, and HOLOTYPE: USNM 71849, Navaka River, discoid, with a slightly convex base. Costae USGS Sta. 24918. corresponding to all septa, rounded, granu lated, scarcely distinguishable toward center PARATYPES (5): USNM 71850, 71851, Navaka of base. Six short, blunt spines project hori River, USGS Sta. 24918. zontally from six of the S3 septa, giving the OCCURRENCE: Navaka River, USGS Sta. corallum a roughly hexagonal outline. Septa 24918. hexamerally arranged in four cycles, the last incomplete. The holotype and three of the paratypes have 36 septa, the other two para GENUS Bourneotrochus N.G. types have 34. The arrangement of the septa TYPE SPECIES: Bourneotrochus veroni n. sp. in each system is consistent and peculiar: Sl Recent, off southern septa are thin, thickening at the wall where Great Barrier Reefs, Queensland, they continue as costae, with a slender palus Australia, 348-531 m. near the junction with the columella. Each S2 is joined near the columella by a pair ofS3 . At DIAGNOSIS: Solitary, free, discoidal del the junction of an S2 with one of the S3 bear tocyathids, increasing asexually by transverse ing a pair of S4 a pair of thick pali form a tall division (strobilation). Calice shallow, with chevron. The palus-bearing S3 with its pair of pali before all but last cycle ofsepta. Base flat, 16

15 17 5

13 7

10 8

18 " 9

FIGURE 3. 1,2, De/tocyathus heteroclitus n. sp., holotype USNM 71849, X 8, x 8, Navaka R., USGS Sta. 24918; 3, 4, D. heterochtus, paratype USNM 71850, X 4, x 4. Sta. 24918; 5, 6, D. heteroclitus, paratype USNM 71851, X 4, x 4, USGS Sta. 24918; 7,8,9, Bourneotrochus veroni n. g., n. sp., holotype USNM 75852, calice x 4, lateral ofanthocaulus-anthocyathus x 4, base ofanthocaulus x 4, N ofFraser Island, Queensland, 476-531 m; 10, 11, calice and lateral ofsame x 10, x 10; 12, B. veroni, paratype USNM 71853, calice x 4, same locality; 13, B. veroni, USNM 71854, x 8, 25 mi E ofLady Musgrave Island, Queensland, 348-357 m; 14, base of same, x 4; 15, 16, 17, B. veroni USNM 71855, x 4, x 8, x 4, Navaka R., USGS Sta. 24918; 18, B. veroni, base, x 16, same locality. 3

4 9

8 "

FIGURE 4. 1, Heterocyathus aequicostatus USNM 71856, X 4, Kere R., USGS Sta. 25718; 2,3 Citharocyathus conicus USNM 71857, X 4, x 4, Navaka R., USGS Sta. 24918; 4, 5, C. conicus USNM 71857, X 4, x 8, same locality; 6,7, Flabellum paripavoninumusNM71858, x 1.5, x 1.5, Kere R., USGSSta.25718;8,9,10,F.sp.cf.F.deludens,usNM71859, x 1, xl, x 1.5, Navaka R., USGS Sta. 24918; 11, 12, F. vanuatu n. sp., holDtype USNM 71860, x 1.5, x 1.5, Kere R., USGS Sta. 25715.

.lliB el'S", ., Indo-Pacific Scleractinian Corals. Part IO-WELLS 213 marked by scar healed after division. The disc spines, Stephanocyathus (Acinocyathus) has oidal form distinguishes this genus from other tatus (1903, pI. 5, figs. 2-5); the other (pI. 6, caryophylliid genera exibiting transyerse figs. 8-11), represented by three very small division. deltocyathids with basal scars, here desig nated as Bourneotrochus veroni, which Bourne ETYMOLOGY: The genus is named for Gil "safely identified as the young forms ofT. has bert C. Bourne (1861-1933), British tatus. They are of considerable interest for zoologist. they show that the free forms of this species are derived by strobilation from a ... tropho zoid." Although Bourne's identification ofthe Bourneotrochus veron; n. sp. small forms with T. hastatus was in error, this Figure 3, 7-18 is the first recognition oftransverse fission in a caryophylliid, previously known only in the Trochocyathus hastatus Bourne 1903 pars, fungiids and flabellids. By coincidence in the p. 32, pI. 6, figs. 8-11, not pI. 5, figs. 2-5.. same year, Dennant (1903: 214) suggested DESCRIPTION: Corallum small, discoidal, strobilation in his new Eocene (Austra and free, with flat or slightly concave base lia) genus Pleuropodia, another very small with scar of formar attachment, or attached coral of uncertain affinities. The occurrence with impending transverse division to cali of anthocyathi still attached to parent cular surface of a similar parent corallum anthocauli of nearly equal size in the speci ("anthocyathus"). Corallum contracting mens from Queensland described above indi slightly from base to calice rim, in lateral aspect cates that the adult stage represented by the resembling a squat, truncated cone. Costae anthocaulus, itself an anthocyathus from a equal, subacute finely granulated or subspi previous division, is the full size attained in B. nose. Associated with each of the six Sl is a veroni, and that T. hastatus, a much larger blunt spine up to 2 mm long, projecting out form, as restricted here, with unscarred base, ward,and slightly down from near the basal is quite a different coral. Further examination margin. Septa in four complete cycles, each of of the Queensland dredged material may re the six S3 lacking a pair of S4' Septa scarcely veal the Deltocyathus-like initial sexual exsert, decreasing in length with increasing generation. cycle number. Pali present before Sl' larger Specimens of Dunocyathus parasiticus ones before S2 and S3' except that they are Woods 1878, ofwhich Deltocyathus rotaefor present only before S3 that are flanked by mis Woods 1878 is a synonym (Wells 1958), S4-a total of 18 pali. Septa and pali granu from the same localities as Bourneotrochus lose or spinose. Fossa shallow. veroni are also examples of strobilation: D. rotaeformis is a discoidal anthocyathus with DIMENSIONS basal scar which in some individuals is cd bd h covered by poststrobilation extensions of the Australia flat costae over the scar; D. parasiticus, com Holotype monly immersed in Bipora, is the turbinate ("anthocyathus") 2.2 3.5 1.5mm anthocaulus. ("anthocaulus") 3.0 3.2 1.0 Paratype ETYMOLOGY: The genus is named for Gil 1 2.5 3.0 0.9 bert C. Bourne (1861-1933), British marine 2 3.0 3.6 1.5 (incipient. zoologist. The species is named for J. E. N. division) Veron, Australian scleractinian systematist 3 2.8 3. 1.0 and ecologist. Vanuatu 3.5 3.75 0.75 HOLOTYPE: USNM 71852, 43 mi. N of Fraser REMARKS: Bourne's Trochocyathus hasta Island, Queensland, 476-531 m. tus included two different forms: a free steph anocyathid bearing six long horizontal basal PARATYPES: (1). USNM 71853, as above. (2). 214 PACIFIC SCIENCE, Volume 38, July 1984

USNM 71854, 25 mi. E of Lady Musgrave Citharocyathus conicus Yabe & Eguchi Island, Queensland, 348-357 m. 1932b, p. 443; 1942a, p. 122, pI. 10, figs. 17, 18. OCCURRENCE: Pleistocene: Vanuatu, USGS Citharocyathus conicus Yabe & Eguchi Sta. 24918. Recent: offsouthern Great Barrier 1941, p. 212, figs. 4a, 4b. Reefs, Queensland (holotype and paratypes), Citharocyathus conicus Eguchi 1941b, p. 62. 348-531 m; off Tutanga Island, Funafuti, Notocyathus conicus Yabe & Eguchi 1946, 766?m, Hawaii, 274-336m. p. 6, figs. I, 2. MATERIAL: Nine specimens of the slender GENUS Peponocyathus GRAVIER venustus form (conical angle = 27°), ranging in size from cd = 2.72, h = 3.5 mm, to Peponocyathus orientalis (Duncan) cd = 4.5, h = 6.25. OCCURRENCE: Navaka River, USGS Sta. Deltocyathus orientalis Duncan 1876, p. 24918. Pliocene, Mindoro, Philippines; Plio 431, pI. 38, figs. 4-7. Pleistocene: Ryukyus. Recent: Indonesia, Peponocyathus orientalis Yabe & Eguchi China Sea, Philippines, Japan. 62-522m, 1932b, p. 444, figs. 12°-26°C. Deltocyathus (Paradeltocyathus) orientalis Yabe & Eguchi 1937, p. 131, pI. 20, figs. 1-10; I942a, p. 123. Deltocyathus (Paradeltocyathus) orientalis FAMILY FLABELLIDAE BOURNE Squires & Keyes 1967, pp. 13, 24, pI. 3, GENUS Flabellum LESSON figs. 1-7. Deltocyathus (Paradeltocyathus) orientalis Flabellum paripavoninum Alcock Wells 1976, p. G9, pI. 3, figs. 1-3. Peponocyathus orientalis Cairns 1979, p. Figure 4, 6, 7 115, pI. 50, figs. 8, 9. Flabellum paripavoninum Alcock 1894, p. 187. MATERIAL: Twelve typical bowl-shaped Flabellum paripavoninum Alcock 1898, coralla of this common western Pacific turbi p. 21, pI. 2, figs. 2, 3, 3a, b. nolid, well illustrated by Yabe & Eguchi and Flabellum pavoninum paripavoninum Cairns. Vaughan 1907, p. 59, pI. 3, figs. 1-4. Flabellum paripavoninum Faustino 1927, OCCURRENCE: Navaka River: USGS Sta. p. 46, pI. 2, figs. 1-4. 24918. Eocene-Pleistocene: Tonga (Eua), New Zealand, Japan, Taiwan, Ceram. Re MATERIAL: Two specimens, one a large cent: New Zealand, Japan, Philippines, Indo corallum (cd = II x 27 mm, h = 22 mm, nesia, Indian Ocean. 75-550m. lateral angle 63°) with basal cicatrix, the other an immature corallum (cd = 6 x 12, h = 12 mm, lateral angle 82°) with basal GENUS Citharocyathus ALCOCK 1902 pedicel. The basal scar of the larger corallum is like those characteristically found in coralla Citharocyathus conicus Alcock 1902 of the F. stokesi species group, the effect of Figure 4, 2-5 strobilation. Vaughan and Faustino both Citharocyathus conicus Alcock 1902, p. 22, thought the fracture was accidental, although pI. 3, figs. 18, 18a. Faustino noted that many specimens from the Citharocyathus venustus Alcock 1902, p. 22, Philippines were similarly broken and figured pI. 3, figs. 19, 19a. one ofthem (1927, pI. 2, figs. 3, 4). It appears Citharocyathus conicus Faustino 1927, p. that transverse fission, supposedly not a char 78, pI. 6, figs. 6, 7. acter of the F. pavoninum group, does occur. Indo-Pacific Scleractinian Corals. Part to-WELLS 215

Corallum colored a faded purple-brown, as DIMENSIONS noted by Alcock in his specimen ("pale h (cicatrix to cd madder"). calice rim) OCCURRENCE: Kere River: USGS Stas. Holotype 37mm 11 x 19mm 25715 and 25718. Recent: Hawaii, Philippines, Paratype Laccadive Sea. 198-1160m. 1 20 6 x 8 2 26 9 x 13 3 28 10 x 18 4 33 11 x 18 Flabellum sp. cf. F. deludens Marenzeller 5 39 11 x 20 Figure 3,8-10 6 46 13 x 23 Flabellum deludens Marenzeller 1904, p. MATERIAL: Thirteen specimens (holotype 209, pI. 17, fig. 10. and 12 paratypes), from Kere River, USGS Flabellum deludens Vaughan 1907, p. 63, pI. Sta. 25715; one specimen from Kere River, 5, figs. 5a, 5b. USGS Sta. 25718. Flabellum deludens Yabe & Eguchi 1942a, p. 135, pI. 12, fig. 1. REMARKS: A readily distinguished species Flabellum deludens Yabe & Eguchi 1942b, marked by (1) the very low lateral angle, about p. 101, pI. 5, figs. 9, 10, 11. 10° less than the least lateral angle of any Flabellum deludens Eguchi 1968, p. C44, pI. other taxon of the Flabellum stokesi species C22, figs. 4, 5; pI. C25, figs. 3, 4. group (30° in F. irregulare Semper), (2) more lateral spines (from three to five on each side), MATERIAL: One specimen, well preserved and (3) proportionally higher corallum and except for an incomplete margin ofthe calice; smaller basal scar. four worn, small fragmentary juveniles. A re lated species is Flabellum aotearoa Squires ETYMOLOGY: vanuatu, noun in apposition (1964, Squires & Keyes 1967) from New (Vanuatu, olim New Hebrides). Zealand. HOLOTYPE: USNM 71860, Kere River, USGS OCCURRENCE: Navaka River, USGS Sta. Sta. 25715. 24918. Recent: Indian Ocean; South Pacific; PARATYPES (12): USNM 71861, Kere River, Japan; Hawaii. 82-1200m. New Zealand USGS Sta. 25715. (Flabellum aotearoa), 130-l84m. OCCURRENCE: Kere River, USGS Sta. 25715 and 25718. Flabellum vanuatu n. sp. Figure 4, 11, 12; Figure 5, 1 FAMILY DENDROPHYLLIIDAE GRAY DESCRIPTION: Corallum compressed elon GENUS Dendrophyllia BLAINVILLE gate conical, lateral edges subacute or nar rowly rounded, bearing hollow spines (lateral Dendrophyllia subcornigera Eguchi angle 20°-22°). Calice regularly oval, very deep. Base truncate with cicatrix. Septa non Figure 5, 4, 5 exsert, lateral faces with discrete acute gra Dendrophyllia subcornigera Eguchi 1968, p. nulations, in three groups diminishing in C64, pI. C32, figs. 3, 4. thickness and length: 16/16/32 = 64 septa, Dendrophyllia subcornigera cylindrica in the holotype. In the largest paratype: Eguchi 1968, p. C64, pI. C32, figs. 1,2. 20/20/40. Columella deep in the calice, com ?Dendrophyllia ijimai Eguchi 1968, p. C65, posed of a few septal trabeculae from inner pI. C16, figs. 1,2; pI. C22, fig. 1; pI. C30, edges of the larger septa. figs. 4,5. 216 PACIFIC SCIENCE, Volume 38, July 1984

4 6

FIGURE 5. 1, Flabellum vanuatu n. sp., holotype USNM 71860 (top center) and 5 paratypes USNM 71861, x I, Kere R., USGS Sta. 25715; 2, 3, Balanophyllia gigas, USNM 71862, x I, x 2, Kere R., USGS Sta. 25715; 4, 5, Dendrophyllia sub cornigera USNM 71863, x 1, x 4, Kere R., USGS Sta. 25715; 6,7, Distichopora nitida, USNM 71864, x 0.5, x 4, Kere R., Sta. SM 242. Indo-Pacific Scleractinian Corals. Part IO-WELLS 217

MATERIAL: A number of fragments of this Distichopora nitida Boschma 1959, p. 151, arborescen-t form, marked by axial corallites pI. 19, figs. 4-7; pI. 10, fig. 4; pI. 9, figs. 10 mm in diameter, tapering to 5mm with 4-7; pis. Il-13 (synonymy). scattered slender lateral corallites, and calices Distichopora nitida Eguchi 1968, p. 41, pI. 4, with 4 septal cycles, are referred to this species fig. 4, 5; pI. 25, figs. 1,2.; pI. 26, figs. 6,7, which probably subsumes Eguchi's Dendro 9; pI. 27, figs. 1-3. phyllia ijimai. MATERIAL: One large piece of corallum OCCURRENCE: Kere River: USGS Sta. from Kere River, faded ochre in color ("Disti 25715. Recent: Seto, Japan (Dendrophyllia chopora ochracea" Quelch). Several small, subcornigera, no depth data); Sagami Bay, worn fragments from the same locality. Japan (D. ijimai), 36-360m. OCCURRENCE: Kere River, USGS Stas. 25715, 242. Recent: central tropical Pacific. GENUS Balanophyllia WOOD 90-180m.

Balanophyllia gigas v.d. Horst LITERATURE CITED Figure 5, 2, 3 Balanophyllia gigas v. d. Horst 1922, p. 58, ALCOCK, A. 1893. Some newly-recorded pI. 8, fig. 22. corals from the Indian seas. Jour. Asiatic Balanophyllia gigas Eguchi 1941b, p. 63. Soc. Bengal 62: 138-149. Balanophyllia gigas Yabe & Eguchi 1942a, ---. 1894. On some new and rare corals p. 139. from the deep waters ofIndia. Jour. Asiatic Balanophyllia gigas Eguchi 1968, p. C51, pI. Soc. Bengal 63: 186-188. C21, figs. 1, 2; pI. C31, figs. 5, 6. ---. 1898. Account ofdeep-sea madrepo Balanophyllia gigas Eguchi & Mori 1973, rarian corals collected by the Royal Indian p.47. Survey Ship "Investigator." Calcutta. 29 pp. MATERIAL: Two large, worn cornute exam ---. 1902. Report on deep-sea Madrepo ples, cd = 14,x 18mm, h = 57-62mm, raria of the Siboga Expedition. Siboga measured along convex side. Wall solid, im Expeditie Monog. 16a. 55 pp. perforate, faintly costate. BOSCHMA, H. 1959. Revision of the Indo Balanophyllia gigas was a manuscript name Pacific species of the genus Distichopora. of Brueggemann's (BMNH 76.10.11.23) men Bijdragen tot de Dierkunde, Afl. 29: tioned by Moseley (1881, p. 193), finally de 121-171. scribed and figured by van der Horst in 1922. BOURNE, G. C. 1903. Some new and rare OCCURRENCE: Kere River, USGS Sta. corals from Funafuti. Jour. Linn. Soc. Lon 25715. Pliocene: Mindoro, Philippines; Pleis don, Zooi. 29:26-37. tocene: Honshu, Japan; Recent: Japan, Indo CAIRNS, S. D. 1979. The deep-water Scleract nesia. 90-500m. inia of the Caribbean Sea and adjacent waters. Studies on the fauna of Curac;ao and other Caribbean islands. 57(186). 341 pp. ---. 1982. Antarctic and Subantarctic ORDER STYLASTERINA HICKSON & ENGLAND Scleractinia.Amer. Geophys. Union, F AMILY STYLASTERIDAE GRAY Antarctic Res. Ser. 34. 74 pp. GENUS Distichopora LAMARCK DENNANT, J. 1899. Descriptions ofnew species Distichopora nitida Verrill of corals from the Australian tertiaries. Trans. R. Soc. South Australia 23 : Figure 5, 6, 7 Il2-122, 281-287. Distichopora nitida Verrill 1864, p. 46. ---. 1903. Descriptions of new species of 218 PACIFIC SCIENCE, Volume 38, July 1984

corals from the Australian tertiaries. Trans. poda from the New Hebrides. Nautilus R. Soc. South Australia 27: 208-215. 90: 127-138. DUNCAN, P. M. 1876. Notices ofsome deep-sea ---. 1982. Cenozoic fossil mollusks from and littoral corais from the Atlantic Ocean, Western Pacific Islands: Gastropods. U.S. Caribbean, Indian, New Zealand, Persian Geol. Surv. Prof. Paper 1171. 100 pp. Gulf and Japanese etc. seas. Proc. ZooI. MALLICK, D. 1. J. 1971. Annual Rept. New Soc. London 1876: 428-442. Hebrides Condominium GeoI. Surv. EGUCHI, M. 1941. [On some fossil simple 1970: 16-20. corals of Mindoro 1., Philippines]. Jour. MALLICK, D. 1. J., and D. GREENBAUM. 1975. Geol. Japan 48: 414-417. (In Japanese.) The Navaka fossiliferous sands and the ---. 1968. The hydrocorals and scleract Kere shell bed. New Hebrides Condo inian corals of Sagami Bay. BioI. Lab. minium Geol. Surv. Regional Rept. Imperial Household, Tokyo 15: 1-53, 1973: 8-12. CI-C80. MICHELIN, H. 1842. Description d'une nou EGUCHI, M., and R. MORL 1973. A study of velle espece du genre Fongie. Rev. Zool. fossil corals from Tateyama City and its en 5: 316; Magasin de ZooI. 5: plate 5 (1843). virons and recent coral fauna ofChiba Pre MICHELOTTI, J. 1838. Specimen Zoophyto fecture, Central Japan. Bull. Tokyo College logiae Diluvianae. Turin 227 pp. of Domestic Sci. 13 :41-57. MILNE EDWARDS, H., and J. HAIME. 1848. Re EGUCHI, M., and T. MIYAWAKI. 1975. Sys cherches surles polypes. Monogr. des turbi tematic study of the scleractinian corals of nolides. Ann. Sci. Nat. (3), 9: 211-344. Kushimoto and its vicinity. Marine Park ---. 1851. Recherches sur les polypes. Res. Sta. Bull. 1:47-62. Monogr. des fongides. Ann. Sci. Nat. (3), FAUSTINO, L. A. 1927. Recent Madreporaria 15: 73-144. of the Philippine Islands. Philippine Dept. ---. 1857-1860. Histoire naturelle des Agr. Nat. Resources, Bur. Sci. Mon. 22. coralliaires. Paris. Vol. 1: 326 pp. [1857]; 310 pp. Vol. 2: 633 pp. [1857]; Vol. 3: 560 pp. [1860]; FOLKESON, F. 1919. Results of Dr. E. Mjo atlas [1857]. bergs Swedish Sci. Exped. to Australia, MOSELEY, H. N. 1881. On the deep-sea Mad 1910-1913. XXII. Madreporaria. Kungl. reporaria. Rept. Sci. Results Voy. H.M.S. Svenska Vetens. Akad. Handl. 59: 1-23. "Challenger." ZooI. 2: 127-208. GARDINER, J. S. 1899. On the solitary corals PRATT, E. M. 1900. Anatomy ofNeoheliapor collected by Dr. A. Willey. Zool. Results eel/ana Moseley. Willey's Zool. Results (Willey) (Part 2): 161-170. 5: 591-602. GARDINER, J. S., and P. WAUGH. 1938. The SARS, G. O. 1872. On some remarkable forms flabellid and turbinolid corals. Sci. Repts. of animal life from the great deeps off the John Murray Exped., Brit. Mus. Norwegian coast. Christiania. 82 pp. 5: 167-202. SMITH, W. D. 1913. Contributions to the ---. 1939. Madreporaria excluding Flab stratigraphy and fossil invertebrate fauna ellidae and Turbinolidae. Sci. Repts. John of the Philippine Islands. Philippine Jour. Murray Exped., Brit. Mus. 6: 225-242. Sci. 8: 235-300. Guppy, H. B. 1885. Observations on the recent SQUIRES, D. F. 1964. New stony corals (Scler calcareous formations of the Solomon actinia) from northeastern New Zealand. Group. Trans. R. Soc. Edinburgh Records Auckland Inst. Mus. 6: 1-9. 32: 545-581. SQUIRES, D. F., and 1. W. KEYES. 1967. The HICKSON, S. J. 1903. The horny membrane of marine fauna ofNew Zealand: Scleractinia. Neohelia poreel/ana. Nature 344. N.Z. Oceanogr. Inst. Mem. 43. 46 pp. LADD, H. S. 1975. Two Pleistocene volutes UMBGROVE, J. H. F. 1946. Corals from a lower from the New Hebrides. Veliger 18: Pliocene patch reef in central Java. Jour. 134-138. PaleontoI. 20: 521-542. ---. 1976. New Pleistocene Neogastro- ---. 1950. Corals from the Putjangan beds Indo-Pacific Scleractinian Corals. Part lO-WELLS 219

(Lower Pleistocene) of Java. Jour. corals of Australia. Linn. Soc. N.S. Wales Paleontol. 24: 637-651. Proc. 2: 292-341. VAN DER HORST, C. J. 1922. Madreporaria of YABE, H., and M. EGUCHI. 1932a. A study of the Siboga Expedition. Part 3. Eupsam the recent deep-water coral fauna ofJapan. midae. Siboga Exped. Repts., Mon. Proc. Imperial Acad. Japan 8: 387-390. l6c:99-l27. ---. 1932b. Deep-water corals from the VAUGHAN, T. W. 1907. Recent Madreporaria Riukiu limestone of Kikai-Jima, Riukiu of the Hawaiian Islands and Laysan. Bull. Islands. Proc. Imperial Acad. Japan 8: U.S. Nat. Mus. 59.427 pp. 442-445. VERON, J. E. N., and M. PICHON. 1980. Scler ---. 1934. Probable generic identity of actinia of Eastern Australia. Part 3. Fami Stephanophyllia Michelin and Micrabacia lies Agaricidae, Siderastreidae, Fungiidae, E. & H. Proc. Imperial Acad. Japan Oculinidae, Merulinidae, Mussidae, Pecti 10:278-281. nidae, Caryophylliidae, Dendrophylliidae. ---. 1937. Notes on Deltocyathus and Dis Mem. Austral. Inst. Marine Sci. 4. 422 pp. cotrochus from Japan. Sci. Repts. Tohoku VERRILL, A. E. 1864. List of the polyps and Imperial Dniv. (2) 19: 127-147. corals sent by the Museum ofComparative ---. 1941. Simple corals from the Sumagui Zoology ... in exchange, with annotations. formation, Philippine Islands. Proc. Imper Bull. Mus. Compo Zool. Harvard 1:29-60. ial Acad. Japan 17: 210-215. VON MARENZELLER, E. 1888. Ueber einige ---. 1942a. Fossil and recent simple corals japanische Turbinoliiden. Ann. naturhist. from Japan. Sci. Repts. Tohoku Imperial Hofmuseum, Wien 3: 12-22. Dniv. (2) 22: 105-178. ---.1904. Steinkorallen. Wiss. Ergebnisse ---. 1942b. Fossil and recent Flabellum Deutschen Tiefsee-Expeditie aufden Damp from Japan. Sci. Repts. Tohoku Imperial fer "Valdivia" 1888-18997:263-318. Dniv. (2) 22:87-103. WELLS, J. W. 1958. Scleractinian corals. ---.1946. Generic identity ofNotocyathus B.A.N.Z. Antarctic Res, Exped., 1929 Tenison Woods and Citherocyathus Al 1931, Repts. Ser. B. 6:257-275. cock. Proc. Imperial Acad. Japan 22: 6-8. ---. 1976. Eocene corals from Eua, ZIBROWIUS, H. 1980. Les scleractiniaires de la Tonga. U.S. Geol. Surv. Prof. Paper 640-G. Mediterranee et de l'Atlantique nord 12 pp. central. Inst. oceanogr. Monaco Mem. 11. WOODS, J. E. T. 1878. On the extratropical 284 pp.