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Scleractinia of the Temperate 

North Pacific 

STEPHEN D. CAIRNS 

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 557

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S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y • N U M B E R 5 5 7 


North Pacific 

Stephen D. Cairns 

SMITHSONIAN INSTITUTION PRESS 

Washington, D.C. 

1994

ABSTRACT 

Cairns, Stephen D. Scleractinia of the Temperate North Pacific. Smithsonian Contributions to 

Zoology, number 557, 150 pages, 3 figures, 42 plates, 5 tables, 1994.—The 119 species of 

temperate North Pacific azooxanthellate Scleractinia are described or diagnosed in two separate 

accounts: the 25 species known to occur in the temperate northeastern Pacific and the 102 

species known to occur in the temperate northwestern Pacific, there being only eight species in 

common. Three genera, six species, and one subspecies are described as new; nine new 

combinations are suggested; and 28 new records for the northern temperate regions are reported. 

A neotype for Desmophyllum dianthus (Esper, 1794) is designated. A dichotomous key to the 

temperate northeastern Pacific Scleractinia is provided, as well as a key to the seven species of 

northwestern Pacific Truncatoflabellum. Separate historical resumes are discussed and 

tabularized for the northeastern and northwestern temperate faunas. The study is based on new 

material from a variety of sources, but primarily from the cruises of the Albatross (USNM) and 

the R/V Tansei Mam (ORI), and the collections of the California Academy of Sciences, Royal 

British Columbia Museum, and the Scripps Institution of Oceanography. 

The region covered in this account extends from Bahia Magdalena on the Pacific coast of Baja 

California to Formosa Strait, off China, including two warm temperate provinces (California 

and Japan) and four cold temperate provinces (Oregon, Aleutian, Oriental, and Kurile). The 

most species-rich temperate region is the warm temperate northwestern Pacific Japan Province, 

which contains 92 azooxanthellate species. This province is directly adjacent to the even more 

species-rich Indo-West Pacific tropical region from which it receives many tropical and 

eurythermic tropical species via the warm, northerly flowing Kuroshio Current. Azooxanthellate 

species diversity decreases with higher latitude: 106 species occur in the combined warm 

temperate North Pacific provinces, 37 in the lower boreal provinces (Oregon and Oriental), and 

only 14 species occur in the combined upper boreal provinces (Aleutian and Kurile). The 

northernmost record of a scleractinian coral in the North Pacific is Caryophyllia arnoldi from 

Prince William Sound (60°48'N) in the Aleutian Province. This province also holds the record 

for the deepest living scleractinian coral, Fungiacyathus marenzelleri, at 6328 m in the Aleutian 

Trench. 

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is 

recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral 

Montastrea cavernosa (Linnaeus). 

Library of Congress Cataloging-in-Publication Dala 

Cairns, Stephen 

Scleractinia of the temperate North Pacific / Stephen D. Cairns 

p. cm.—(Smithsonian contributions to zoology ; no. 557 

Includes bibliographical references (p. ). 

1. Scleractinia—North Pacific Ocean. I. Series. 

QLl.S54no. 557 [QL377.C7] 591 s-dc20 [59.V.5] 94-32147 CIP 

'S> The paper used in this publication meets the minimum requirements of the American 

National Standard for Permanence of Paper for Printed Library Materials Z39.48—1984.

Contents 

Page 

Introduction 1 

Abbreviations 1 

Acknowledgments 2 

Historical Resumes 2 

Temperate Northeastern Pacific 2 

Temperate Northwestern Pacific 3 

Material 6 

Methods 6 

Zoogeography 8 

Temperate Northeastern Pacific 8 

Temperate Northwestern Pacific 12 

Comparisons 12 

Systematic Account 13 

Part 1: Temperate Northeastern Pacific 13 

Key to the 25 Species of Temperate Northeastern Pacific Scleractinia 13 

Suborder FUNGIINA 14 

Superfamily FUNGIOIDEA 14 

Family FUNGIACYATHIDAE 14 

Fungiacyathus Sars, 1872 14 

Subgenus Fungiacyathus (Bathyactis) Moseley, 1881 14 

Fungiacyathus (B.) marenzelleri (Vaughan, 1906) 15 

Family MlCRABACHDAE 16 

Leptopenus Moseley, 1881 16 

Leptopenus discus Moseley, 1881 16 

Suborder FAVIINA 17 

Superfamily FAVIOIDEA 17 

Family RHIZANGIIDAE 17 

Astrangia Milne Edwards and Haime, 1848 17 

Astrangia haimei Verrill, 1866 17 

Family OCULINIDAE 18 

Oculina Lamarck, 1816 18 

Oculina profunda Cairns, 1991 18 

Madrepora Linnaeus, 1758 18 

Madrepora oculata Linnaeus, 1758 18 

Suborder CARYOPHYLLIINA 19 

Superfamily CARYOPHYLLIOIDEA 19 

Family CARYOPHYLLIIDAE 19 

Caryophyllia Lamarck, 1816 19 

Subgenus Caryophyllia (Caryophyllia) Lamarck, 1816 19 

Caryophyllia (C.) arnoldi Vaughan, 1900 19 

Caryophyllia (C.) alaskensis Vaughan, 1941 20 

Caryophyllia (C.) sp. A 21 

Labyrinthocyathus Caims, 1979 21 

Labyrinthocyathus quaylei (Durham, 1947) 21 

Crispatotrochus Tenison Woods, 1879 22 

Crispatotrochus foxi (Durham and Barnard, 1952) 22 

in

IV SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 

Paracyathus Milne Edwards and Haime, 1848 22 

Paracyathus stearnsii Verriil, 1869 22 

Paracyathus montereyensis Durham, 1947 24 

Coenocyathus Milne Edwards and Haime, 1848 24 

Coenocyathus bowersi Vaughan, 1906 25 

Nomlandia Durham and Barnard, 1952 25 

Nomlandia californica Durham and Barnard, 1952 26 

Desmophyllum Ehrenberg, 1834 26 

Desmophyllum dianthus (Esper, 1794) 26 

Lophelia Milne Edwards and Haime, 1849 27 

Lophelia pertusa (Linnaeus, 1758) 27 

Superfamily FLABELLOIDEA 28 

Family FLABELLIDAE 28 

Flabellum Lesson, 1831 28 

Subgenus Flabellum (Flabellum) Lesson, 1831 28 

Flabellum (F.) sp. A 28 

Javania Duncan, 1876 29 

Javania cailletti (Duchassaing and Michelotti, 1864) 29 

Javania borealis, sp. nov 30 

Javania californica, sp. nov 30 

Polymyces Caims, 1979 31 

Polymyces montereyensis (Durham, 1947) 31 

Suborder DENDROPHYLLIINA 32 

Family DENDROPHYLLIIDAE 32 

Balanophyllia Wood, 1844 32 

Balanophyllia elegans Verriil, 1864 32 

Balanophyllia cedrosensis Durham, 1947 34 

Dendrophyllia Blainville, 1830 34 

Dendrophyllia oldroydae Oldroyd, 1924 34 

Dendrophyllia californica Durham, 1947 35 

Part 2: Temperate Northwestern Pacific 36 

Suborder ASTROCOENMNA 36 

Family POCILLOPORIDAE 36 

Madracis Milne Edwards and Haime, 1849 36 

Madracis sp. A 36 

Suborder FUNGIINA 37 

Superfamily FUNGIOIDEA 37 

Family FUNGIACYATHIDAE 37 

Fungiacyathus Sars, 1872 37 

Subgenus Fungiacyathus (Fungiacyathus) Sars, 1872 37 

Fungiacyathus (F.) stephanus (Alcock, 1893) 37 

Fungiacyathus (F.) paliferus (Alcock, 1902) 37 

Fungiacyathus (F.) sp. A 38 

Subgenus Fungiacyathus (Bathyactis) Moseley, 1881 38 

Fungiacyathus (B.) marenzelleri (Vaughan, 1906) 38 

Fungiacyathus (B.) variegatus Caims, 1989 38 

Fungiacyathus (B.) granulosus Caims, 1989 39 

Family MICRABACHDAE 39 

Leptopenus Moseley, 1881 39 

Leptopenus discus Moseley, 1881 39 

Leptopenus solidus Keller, 1977 39 

Letepsammia Yabe and Eguchi, 1932 40 

Letepsammia formosissima (Moseley', 1876) 40 

Rhomhopsammia Owens, 1986 41 

Rhombopsammia niphada Owens, 1986 41

NUMBER 557 

Stephanophyllia Michelin, 1841 41 

Stephanophyllia fungulus Alcock, 1902 41 

Suborder FAVIINA 42 

Superfamily FAVIOIDEA 42 

Family RHIZANGIIDAE 42 

Culicia Dana, 1846 42 

Culicia japonica Yabe and Eguchi, 1936 42 

Oulangia Milne Edwards and Haime, 1848 42 

Oulangia stokesiana miltoni Yabe and Eguchi, 1932 42 

Family OCULINIDAE 43 

Madrepora Linnaeus, 1758 43 

Madrepora oculata Linnaeus, 1758 43 

Cyathelia Milne Edwards and Haime, 1849 43 

Cyathelia axillaris (Ellis and Solander, 1786) 43 

Family ANTHEMIPHYLLIIDAE 44 

Anthemiphyllia Pourtales, 1878 44 

Anthemiphyllia dentata (Alcock, 1902) 44 

Anthemiphyllia frustum, sp. nov 44 

Suborder CARYOPHYLLIINA 45 

Superfamily CARYOPHYLLIOIDEA 45 

Family CARYOPHYLLHDAE 45 

Caryophyllia Lamarck, 1816 45 

Subgenus Caryophyllia (Caryophyllia) Lamarck, 1816 45 

Caryophyllia (C.) japonica Marenzeller, 1888 45 

Caryophyllia (C.) alaskensis Vaughan, 1941 45 

Caryophyllia (C.) sp. cf. C. scobinosa Alcock, 1902 45 

Caryophyllia (C.) quadragenaria Alcock, 1902 46 

Caryophyllia (C.) rugosa Moseley, 1881 47 

Caryophyllia (C.) jogashimaensis Eguchi, 1968 47 

Caryophyllia (C.) ambrosia ambrosia Alcock, 1898 48 

Caryophyllia (C.) laevicostata Moseley, 1881 48 

Subgenus Caryophyllia (Acanthocyathus) Milne Edwards and Haime, 

1848 49 

Caryophyllia (A.) grayi Milne Edwards and Haime, 1848 49 

Caryophyllia (A.) spiniger Kent, 1871 49 

Subgenus Caryophyllia (Premocyathus) Yabe and Euchi, 1942 50 

Caryophyllia (P.) compressa Yabe and Eguchi, 1942 50 

Crispatotrochus Tenison Woods, 1879 51 

Crispatotrochus rubescens (Moseley, 1881) 51 

Crispatotrochus niinoi (Yabe and Eguchi, 1942) 51 

Paracyathus Milne Edwards and Haime, 1848 51 

Paracyathus pruinosus Alcock, 1902 52 

Trochocyathus Milne Edwards and Haime, 1848 52 

Trochocyathus caryophylloides Alcock, 1902 52 

Trochocyathus japonicus Eguchi, 1968 53 

Trochocyathus decamera, sp. nov 53 

Trochocyathus cooperi (Gardiner, 1905), comb, nov 54 

Deltocyathus Milne Edwards and Haime, 1848 54 

Deltocyathus vaughani Yabe and Eguchi, 1932 54 

Deltocyathus rotulus (Alcock, 1898) 55 

Deltocyathus magnificus Moseley, 1876 56 

Stephanocyathus Seguenza, 1864 56 

Subgenus Stephanocyathus (Acinocyathus) Wells, 1984 57 

Stephanocyathus (Acinocyanthus) spiniger (Marenzeller, 1888) . . . . 57 

Subgenus Stephanocyathus (Odontocyathus) Moseley, 1881 57

Vi SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 

Stephanocyathus (Odontocyathus) weberianus (Alcock, 1902), comb. 

nov 57 

Conotrochus Seguenza, 1864 58 

Conotochus funicolumna (Alcock, 1902) 58 

Aulocyathus Marenzeller, 1904 59 

Aulocyathus recidivus (Dennant, 1906) 59 

Aulocyathus matricidus (Kent, 1871) 60 

Desmophyllum Ehrenberg, 1834 60 

Desmophyllum dianthus (Esper, 1794) 61 

Lophelia Milne Edwards and Haime, 1849 61 

Lophelia pertusa (Linnaeus, 1758) 61 

Anomocora Studer, 1878 61 

Anomocora sp 61 

Coenosmilia Pourtales, 1874 61 

Coenosmilia sp. cf. C. arbuscula Pourtales, 1874 61 

Phyllangia Milne Edwards and Haime, 1848 61 

Phyllangia hayamaensis (Eguchi, 1968) 62 

Rhizosmilia Cairns, 1978 62 

Rhizosmilia sagamiensis (Eguchi, 1968), comb, nov 62 

Dasmosmilia Pourtales, 1880 63 

Dasmosmilia pacifica (Yabe and Eguchi, 1932), comb, nov 63 

Goniocorella Yabe and Eguchi, 1932 63 

Goniocorella dumosa (Alcock, 1902) 63 

Family TURBINOLHDAE 64 

Notocyathus Tenison-Woods, 1880 64 

Notocyathus venustus (Alcock, 1902) 64 

Notocyathus conicus (Alcock, 1902) 64 

Peponocyathus Gravier, 1915 65 

Peponocyathus australiensis (Duncan, 1870) 65 

Peponocyathus folliculus (Pourtales, 1868) 66 

Tropidocyathus Milne Edwards and Haime, 1848 67 

Tropidocyathus lessoni (Michelin, 1842) 67 

Tropidocyathus pileus (Alcock, 1902) 68 

Alatotrochus, gen. nov 68 

Alatotrochus rubescens (Moseley, 1876), comb, nov 68 

Idiotrochus Wells, 1935 69 

Idiotrochus kikutii (Yabe and Eguchi, 1941) 69 

Superfamily FLABELLOIDEA 69 

Family GUYNIIDAE 69 

Stenocyathus Pourtales, 1871 69 

Stenocyathus vermiformis (Pourtales, 1868) 69 

Truncatoguynia Cairns, 1989 70 

Truncatoguynia irregularis Cairns, 1989 70 

Family FLABELLIDAE 70 

Flabellum Lesson, 1831 70 

Subgenus Flabellum (Flabellum) Lesson, 1831 70 

Flabellum (F.) pavoninum Lesson, 1831 70 

Flabellum (F.) patens Moseley, 1881 71 

Flabellum (F.) magnificum Marenzeller, 1904 72 

Flabellum (F.) angustum Yabe and Eguchi, 1942 72 

Flabellum (F.) politum Cairns, 1989 73 

Subgenus Flabellum (Ulocyathus) Sars, 1851 73 

Flabellum (U.) deludens Marenzeller, 1904 73 

Flabellum (U.) japonicum Moseley, 1881 73

NUMBER 557 vii 

Flabellum (U.) apertum borealis, subsp. nov 74 

Truncatoflabellum Caims, 1989 75 

Key to the Temperate Northwest Pacific Truncatoflabellum 75 

Truncatoflabellum spheniscus (Dana, 1846) 76 

Truncatoflabellum candeanum (Milne Edwards and Haime, 1848)... 76 

Truncatoflabellum formosum Caims, 1989 77 

Truncatoflabellum carinatum Cairns, 1989 77 

Truncatoflabellum gardineri Cairns, 1993 78 

Truncatoflabellum sp. A 79 

Truncatoflabellum sp. B 79 

Placotrochides Alcock, 1902 79 

Placotrochides scaphula Alcock, 1902 79 

Javania Duncan, 1876 80 

Javania insignis Duncan, 1876 80 

Javania cailleti (Duchassaing and Michelotti, 1864) 80 

Javania borealis, sp. nov 80 

Rhizotrochus Milne Edwards and Haime, 1848 80 

Rhizotrochus typus Milne Edwards and Haime, 1848 81 

Suborder DENDROPHYLLIINA 81 

Family DENDROPHYLLIIDAE 81 

Balanophyllia Wood, 1844 81 

Balanophyllia cumingii Milne Edwards and Haime, 1848 81 

Balanophyllia cornu Moseley, 1881 82 

Balanophyllia gigas Moseley, 1881 83 

Balanophyllia ponderosa Van der Horst, 1926 83 

Balanophyllia sp. A 84 

Balanophyllia teres, sp. nov 84 

Endopachys Lonsdale, 1845 84 

Endopachys grayi Milne Edwards and Haime, 1848 84 

Eguchipsammia, gen. nov 85 

Eguchipsammia gaditana (Duncan, 1873), comb, nov 85 

Eguchipsammia wellsi (Eguchi, 1968), comb, nov 86 

Rhizopsammia Verrill, 1870 87 

Rhizopsammia minuta mutsuensis Yabe and Eguchi, 1932 87 

Cladopsammia Lacaze-Duthiers, 1897 87 

Cladopsammia gracilis (Milne Edwards and Haime, 1848), comb. 

nov 88 

Cladopsammia eguchii (Wells, 1982), comb, nov 88 

Dendrophyllia Blainville, 1830 89 

Dendrophyllia ijimai Yabe and Eguchi, 1934 89 

Dendrophyllia cribrosa Milne Edwards and Haime, 1851 89 

Dendrophyllia japonica Rehberg, 1892 90 

Dendrophyllia arbuscula Van der Horst, 1922 90 

Dendrophyllia boschmai Van der Horst, 1926 91 

Dendrophyllia florulenta Alcock, 1902 91 

Enallopsammia Michelotti, 1871 92 

Enallopsammia rostrata (Pourtales, 1878) 92 

Tubastraea Lesson, 1829 93 

Tubastraea coccinea Lesson, 1829 93 

Schizopsammia, gen. nov 94 

Schizopsammia songae, sp. nov 94 

Appendix: Station List 95 

Literture Cited 100 

Plates 1-42 108

Introduction 

The temperate North Pacific coral region, as defined in 

greater detail in the Zoogeography section, extends from Bahia 

Magdelena on the Pacific coast of Baja California to Formosa 

Strait, China, and spans six zoogeographic provinces. The 

taxonomic and zoogeographic accounts are separated into 

northeastern and northwestern Pacific sections, the dividing 

point being the channel between the Aleutian and Commander 

islands, such that each section could be used as a regional 

faunistic guide. This separation is supported by an almost 

completely separate body of literature on the corals from these 

two regions (see Tables 1, 2), as well as an essentially separate 

fauna in each of the two regions. Of the 119 azooxanthellate 

species known from the temperate North Pacific, only eight 

species occur in both regions, most of those being cosmopolitan 

species. 

Only seven new species or subspecies are described in this 

account, a surprisingly low 6% of the fauna, which is a tribute 

to the earlier efforts of M. Eguchi, H. Yabe, J.W. Durham, and 

J.L. Barnard between the years 1932-1973. 

The most interesting zoogeographic result of this study was 

to show the strong influence of the Indo-West Pacific tropical 

fauna on the warm temperate northwestern Pacific Japan 

Province. As more Indo-West Pacific azooxanthellates are 

found in the Japan Province and as more Japanese "endemics" 

are synonymized with previously described species common in 

the Indo-West Pacific realm, it becomes increasingly apparent 

that the Japan Province is the northern border/transition zone of 

the dominant Indo-West Pacific fauna, a conclusion also 

reached by Veron (1992:3) regarding hermatypic (zooxanthellate) 

corals. As a result, an increasing number of species are 

now known to have distributions extending from the southwestern 

Indian Ocean to the southern coasts of Kyushu and 

Honshu. 

Stephen D. Cairns, Department of Invertebrate Zoology, National 

Museum of Natural History, Smithsonian Institution, Washington, D. 

C. 20560. 


North Pacific 

Stephen D. Cairns 

Abbreviations 

Museums and Collecting Institutions 

AHF Allan Hancock Foundation, University of Southern California, 

Los Angeles 

BM British Museum (Natural History), London 

CAS California Academy of Sciences, San Francisco 

IOM Institute of Oceanology, Moscow 

MCZ Museum of Comparative Zoology, Harvard University. 

Cambridge 

MNHNP Museum National d'Histoire Naturelle, Paris 

NMCIC National Museum of Canada (Canadian Museum of Nature), 

Invertebrate Collection, Ottawa 

NMW Naturhistorisches Museum, Wien 

ORI Ocean Research Institute, University of Tbkyo, Tokyo 

RBCM Royal British Columbia Museum, Victoria, Canada 

RMNH Rijksmuseum van Natuurlijke Historic Leiden 

SBMNH Santa Barbara Museum of Natural History, Santa Barbara 

SIO Scripps Institution of Oceanography, University of California, 

La Jolla 

TIUS Institute of Geology and Paleontology, Tohoku (Imperial) 

University, Sendai. Japan 

UA University of Alaska Museum, Fairbanks 

UCMP University of California, Berkeley, Museum of Paleontology 

USNM former United States National Museum, collection now in 

National Museum of Natural History, Washington, D. C. 

YPM Yale Peabody Museum of Natural History, New Haven 

ZMA Zoologisch Museum, Amsterdam 

ZMB Zoologisches Museum, Berlin 

ZMC Zoologisk Museum, Copenhagen 

Vessels and Expeditions 

Alb U.S.F.W.S. Albatross 

OCSEAP Outer Continental Shelf Environmental Assessment Program 

SEPBOP South East Pacific Biological Oceanographic Program 

TM R/V Tansei Maru (operated by the ORI) 

Morphological Terms 

GCD Greater Calicular Diameter 

GCD:LCD Ratio of greater calicular diameter to lesser calicular 

diameter

D:H Ratio of calicular diameter to height of a corallum 

H:D Ratio of height to diameter of a corallum 

LCD Lesser Calicular Diameter 

PD:GCD Ratio of pedicel diameter to greater calicular diameter 

Sx, Cx, Px, CSX Septa, costae, pali, or costosepta (respectively) of cycle 

designated by numerical subscript 

Sx > S Septa of cycle x more broad than those of 

cycle y 

ACKNOWLEDGMENTS.—I dedicate this publication to the 

memory of my friend and fellow Smithsonian curator, J. 

Laurens Barnard (1928-1991). Much better known as an 

amphipod taxonomist and even as an amateur ornithologist, 

Jerry Barnard co-authored one paper on scleractinian corals 

early in his career (Durham and Barnard, 1952), which has 

served as the basis for coral taxonomy of the eastern Pacific. 

I especially thank Yoshihisa Shirayama (ORI) for allowing 

me to study the diverse deep-water coral collections made off 

southern Japan by the R/V Tansei Maru. I am also very grateful 

to Helmut Zibrowius (Station Marine d'Endoume) and Harry 

Filkorn (Kent State University) for their meticulous efforts in 

reviewing this manuscript. 

I would like to thank the following people who have 

generously loaned me specimens used in this study: N. Foster 

(UA), J.A. Fournier (NMCIC), B.W. Hoeksema (RMNH), C. 

Hussey (BM), A. Johnston (MCZ), N.B. Keller (IOM), E. 

Kools and G. Williams (CAS), E. Kritscher (NMW), D.H.H. 

Kiihlmann (ZMB), P. Lambert and J.A. Cosgrove (RBCM), 

E.A. Lazo-Wasem (YPM), D. Lindberg (UCPM), S.R. Luke 

(SIO), P.H. Scott (SBMNH), O.S. Tendal (ZMC), and R.W.M. 

van Soest (ZMA). 

The scanning electron photomicrographs were taken in the 

SEM Laboratory of the National Museum of Natural History, 

Smithsonian Instituion. Figure 2 was drawn by staff illustrator 

Molly Ryan. 

Historical Resumes 

TEMPERATE NORTHEASTERN PACIFIC (Table 1).—The earliest 

report of a scleractinian from the temperate northeastern 

Pacific appears to have been the original description of 

Balanophyllia elegans Verrill, 1864, from off northern California. 

That this should have been the first species reported is not 

unexpected because it is one of only three shallow-water 

species known from this region and it has a very colorful polyp. 

Shortly thereafter, Verrill (1869) described the second of the 

three shallow-water species, Paracyathus stearnsii, from off 

Monterey, California. 

All subsequent records from this region are listed in Table 1, 

only the larger taxonomic papers or series of papers being 

discussed below. The first comprehensive review of the eastern 

Pacific Scleractinia was that of Durham (1947), who reported 

the corals from the E. W. Scripps Expedition to the Gulf of 

California made in 1940. Durham expanded his revision to the 

coral fauna north of the Gulf as well, including 15 records of 

species from the temperate northeastern Pacific, including 

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 

seven new species: Astrangia lajollaensis (= A. haimei), 

Cyathoceras (= Labyrinthocyathus) quay lei, Paracyathus 

montereyensis, Lophelia californica (= L. pertusa), Flabellum 

(= Polymyces) montereyensis, Dendrophyllia californica, and 

Balanophyllia cedrosensis. Type and nontype material reported 

in this paper are deposited primarily at the University of 

California Museum of Paleontology (UCMP), but some are 

also represented in the collections of the CAS and USNM. 

The most comprehensive review of the eastern Pacific 

Scleractinia was that of Durham and Barnard (1952), who 

based their new records on the collections of the Velero III and 

IV. Although they listed 98 species from the eastern Pacific, 

only 13 of these species pertain to the temperate northeastern 

Pacific as defined herein, including the description of four new 

species: Cyathoceras (= Crispatotrochus) foxi, Dendrosmilia 

nomlandi (= Lophelia pertusa), Nomlandia californica, and 

Flabellum tannerense (= Polymyces montereyensis). Their type 

and nontype specimens were transferred from the Allan 

Hancock Foundation Museum to the Santa Barbara Museum of 

Natural History in 1990. 

In a series of three papers, Keller (1976, 1977, 1981a) 

reported new records of Fungiacyathus, Leptopenus, and 

Caryophyllia, respectively, from bathyal and abyssal depths (to 

6328 m) along the northern rim of the North Pacific from off 

Oregon to off Japan. Both Fungiacyathus marenzelleri and 

Leptopenus discus were reported for the first time from the 

temperate northeastern Pacific, but under different names. Her 

reports of Caryophyllia ambrosia and C. alaskensis are 

discussed in the text. Many of her Vityaz specimens were 

examined on loan from the IOM in 1991, but some voucher 

specimens are also deposited at the USNM. 

Following the natural history observations of Gerrodette 

(1979, 1981) on the distribution and planular dispersal of 

Balanophyllia elegans, Fadlallah produced an excellent series 

of papers on the natural history of the three common, 

shallow-water temperate northeastern Pacific species: Astrangia 

lajollaensis (= A. haimei), Balanophyllia elegans, and 

Paracyathus stearnsii (see Fadlallah, 1982, 1983b; Fadlallah 

and Pearse, 1982a,b). Included in these papers are fascinating 

observations on reproductive ecology, sex ratios, periodicity of 

sexual cycles, planular sizes, growth rates, longevity, and 

population density of adult coralla. 

Bythell (1986) published a fine identification guide to the 17 

species occurring off Southern California (32°-35°N), his new 

records coming from the Scripps Institution of Oceanography 

Invertebrate Collection. Bythell included a key to the genera of 

this region and many useful illustrations, including some of the 

polyps and nematocysts. 

Checklists and fieldguides too numerous to mention have 

included Scleractinia from the northeastern Pacific, all invariably 

including a reference to B. elegans. Although not listed in 

Table 1, some of these references are Ricketts and Calvin 

(1952), Guberlet (1962), Johnson and Snook (1967), Hand 

(1975), Brusca and Brusca (1978), Lewbel et al. (1981),

NUMBER 557 

TABLE 1.—Chronology of Temperate Northeastern Pacific Scleractinian Publications (* denotes significant 

taxonomic publications). 

Year Author(s) Comments 

1864 

1869 

1870a 

1870b 

1886 

1900 

1903 

1906a 

1917 

1924 

1931 

1936 

1941 

1943 

1947 

1949 

1952 

1956 

1961 

1966 

1972 

1976 

1977 

1979 

1981a 

1981 

1982 

1982a 

1982b 

1983b 

1986 

1991a 

1991 

in press 

Verrill 

Verrill 

Verrill 

Verrill 

Whiteaves 

Vaughan 

Vaughan 

Vaughan 

Hickman 

Oldroyd 

Faustino 

Williams 

Vaughan 

Durham 

Durham 

Durham 

Durham and Barnard 

Emerson 

Addicott 

Durham 

Talmadge 

Keller 

Keller 

Gerrodette 

Keller 

Gerrodette 

Fadlallah 

Fadlallah and Pearse 

Fadlallah and Pearse 

Fadlallah 

Bythell 

Cairns 

Hellberg 

Wilson 

Kozloff (1983, 1987), and Austin (1985). Other useful regional 

references, but concerned primarily with tropical eastern 

Pacific species, are Marenzeller (1904b), Vaughan (1906b), 

Squires (1959), Parker (1964), and Wilson (1990). 

In this paper 25 species are reported from the northeastern 

temperate Pacific (Table 4), including six new records for the 

region and two new species. One species not included in the 

species account, however, is Solenosmilia variabilis Duncan, 

1873, which was reported from shallow water (0-200 m) off 

Knight Island, British Columbia by Austin (1985) in his 

checklist of invertebrates from the northeastern Pacific coast. 

Austin's specimen could not be obtained for verification. 

Solenosmilia variabilis is a widespread deep-water species that 

Balanophyllia elegans from off northern California 

Paracyathus stearnsii and P. caltha n. spp. from off Monterey, 

California 

Additional records of B. elegans and original description of Astrangia 

conferta from off Mexico 

B. elegans from off Puget Sound, Washington 

Paracyathus caltha and B. elegans from off British Columbia 

Caryophyllia arnoldi n. sp. from Pleistocene of San Pedro, California 

C. pedroensis, C. californica and Paracyathus pedroensis n. spp. from 

Pleistocene and Pliocene of San Pedro, California 

Coenocyathus bowersi n. sp. from Channel Ids., California 

Reiteration of British Columbia records of B. elegans and P. caltha 

Dendrophyllia oldroydi n. sp. from off California 

Dendrophyllia oldroydi n. sp. from off California 

Another report of D. oldroydi from off southern California 

Caryophyllia alaskensis n. sp. from Alexander Archipelago, Alaska 

Dendrophyllia oldroydae from Pleistocene of California 

15 species, including 7 new species, from temperate Northeast Pacific 

6 species figured and discussed in context of description of polycyclic 

development 

13 species, including 4 new species, reported from temperate Northeast 

Pacific 

B. elegans from Pacific, Baja California 

B. elegans from Pleistocene of central California 

Noncritical listing of 103 species of Scleractinia from East Pacific 

B. elegans from Washington and British Columbia; C. alaskensis from 

Alaska; Desmophyllum cristagalli from northern California 

First record for region of Fungiacyathus marenzelleri (as F. symmetricus 

aleuticus) from off Alaska 

Leptopenus irinae (=L. discus) from off Alaska and Washington 

Discussion of equatorial submergence of B. elegans 

New records of two species of Caryophyllia from off Oregon and 

Alaska 

Discussion of planula dispersal of B. elegans 

Reproductive ecology of Astrangia lajollaensis 

Discussion of sexual reproduction in B. elegans 

Discussion of sexual reproduction in P. stearnsii 

Discussion of population dynamics of B. elegans 

Identification guide to 17 species occurring off southern California 

Oculina profunda n. sp. from off central California 

Discussion of larval dispersal in B. elegans (Abstract) 

Madracis sp. cf. M. pharensis and Phyllangia consagensis from off 

Rocas Alijos, off Baja California 

occurs throughout the Atlantic and Indian Oceans and 

Subantarctic region at depths of 220-2165 m (Cairns, 1982). 

Heretofore, it has not been reported elsewhere in the Pacific 

Ocean or at depths shallower than 200 m, both of which cast 

doubt on the British Columbia record. 

TEMPERATE NORTHWESTERN PACIFIC (Table 2).—The earliest 

record of azooxanthellate corals from the temperate 

northwestern Pacific appears to be that of Gray (1849), who 

listed three previously described species from "off Japan," all 

based on specimens deposited at the BM: Flabellwn distinction 

Milne Edwards and Haime, 1848a (= F. pavoninum), F. bairdi 

Milne Edwards and Haime, 1848a (= Truncatoflabellum 

bairdi), and F. elegans Milne Edwards and Haime, 1848a


TABLE 2.—Abbreviated Chronology of Temperate Northwestern Pacific Scleractinian Publications (* denotes 

significant taxonomic works). 

Year Authors) Comments 

1849 

1850b 

1857 

1865 

1871 

1876 

1881 

1888 

1888a 

1888b 

1892 

1900 

*1922 

•1932g 

*1937 

*1942a 

*1942b 

*1968 

1976 

1977 

1977 

1980 

1981 

1981a 

1982 

1982 

1983 

1986 

1986a 

1987 

1989a 

M991 Song 

Gray 

Milne Edwards 

and Haime 

Milne Edwards 

and Haime 

Verrill 

Kent 

Duncan 

Moseley 

Ortmann 

Marenzeller 

Marenzeller 

Rehberg 

Vaughan 

Van der Horst 

Yabe and Eguchi 




Eguchi 

Keller 

Keller 

Cheng 

Mori and Minoura 

Zou 

Keller 

Keller 

Song 

Mori and Minoura 

Yajima 

Owens 

Mori 

Cairns 

(= Truncatoflabellum candeanum). His report of F. distinction 

from off Japan was a correction of the type locality of the Red 

Sea given in the original account, but his Japanese assignations 

for F. bairdi and F. elegans also contradict (supplement?) the 

reports of an unknown type locality given by the authors of 

those species (Milne Edwards and Haime, 1848a, 1857). Milne 

Edwards and Haime (1850b. 1857, respectively) did specifically 

report two azooxanthellate species from off Japan: 

Cyathelia axillaris (Ellis and Solander, 1786) and Desmophyllum 

dianthus (Esper, 1794). 

Yabe and Eguchi (1942b) credited Verrill (1869) with the 

report of Eupsammia stimpsoniana (= Balanophyllia stimpsonii) 

from Kagosima Bay; however, Verrill's original and 

subsequent descriptions (Verrill, 1865, 1866) refer only to the 

"North China Sea," not a specific Japanese or temperate 

locality. 

Shortly thereafter, Saville Kent (1871) described two new 

species from unspecified localities off Japan: Acanthocyathus 

3 species of Flabellum reported off Japan 

Cyathelia axillaris listed for Japan 

Desmophyllum dianthus reported from off Japan 

Eupsammia stimpsoniana (= Balanophyllia stimpsonii) from "North China 

Sea" 

2 new species described from off Japan 

2 new species described from off Honshu, Japan 

Flabellum japonicum, new species, described from Sagami Bay 

Stephanophyllia superstes, new species, described from Sagami Bay 

3 species of Flabellum reported from off Japan, one new 

3 species reported from Sagami Bay, including 2 new species 

3 new species reported off Japan 

Levipalifer orientalis, new species, described off Honshu 

5 dendrophylliid species reported from off Japan, including 1 new species 

Revision of fossil and Recent Stephanophyllia from Japan 

Revision of Deltocyathus from Japanese region 

Revision of the 13 species of the genus Flabellum from off Japan 

84 fossil and Recent species of solitary corals reported from off Japan, 

including 14 new species 

62 Recent species reported from Sagami Bay, including 12 new species 

Deep-water Fungiacyathus reported from off Hokkaido 

Abyssal Leptopenus reported from off Kurile Islands 

Flabellum apertum reported off Kyushu 

Ontogeny of Flabellum coalitum (= F. pavoninum) discussed 

Checklist of azooxanthellates from East China Sea 

Deep-water Caryophyllia reported from off Kurile Islands 

Deep-water Deltocyathus reported from off Honshu 

First report of azooxanthellate corals from off South Korea 

Discussion of genetic control of septal number in Cylindrophyllia orientalis 

Distribution of Rhizopsammia minuta mutsuensis discussed 

Rhombopsammia squiresi, new species, descibed from off Kyushu 

Discussion of genetic control of septal number in Caryophyllia compressa 

Several azooxanthellate corals reported from off Pratas Islands, northern 

South China Sea 

19 azooxanthellate corals listed for waters off South Korea 

spiniger and Flabellum (= Aulocyathus) matricidum, the types 

of which are deposited at the British Museum. 

Also based on British Museum specimens, Duncan (1876) 

described two new species from off Owase, Mie-ken, Honshu: 

Deltocyathus orientalis (= Peponocyathus australiensis) and 

Javania insignis. 

Only one species from off Japan was reported in the 

Challenger Expedition Reports: the new species Flabellum 

japonicum Moseley, 1881, from Sagami Bay, Honshu. The 

type of this species is deposited at the British Museum. The 

new species Stephanophyllia superstes (= Letepsammia formosissima) 

was also described from Sagami Bay by Ortmann 

(1888), the type deposited at the Strasbourg Zoological 

Museum (H. Zibrowius, pers. comm.). 

Marenzeller (1888a) reported three species of Flabellum 

from off Japan (probably from Sagami Bay), including one new 

species, F. coalitum (= F. pavoninum); these specimens are 

deposited at the NMW. In the same year Marenzeller (1888b)

NUMBER 557 

described two new species from Sagami Bay, Caryophyllia 

japonica and Stephanotrochus (= Stephanocyathus) spiniger, 

and extended the range of Cyathoceras (= Crispatotrochus) 

rubescens Moseley, 1881; these specimens are also deposited 

in the NMW. 

Rehberg (1892) reported the following three new species 

from off Japan: Amphehelia adminicularis (= Enallopsammia 

rostrata), Dendrophyllia japonica, and Coenopsammia 

ramiculosa (= ?Tubastraea micrantha); and an additional 

Japanese record of Dendrophyllia conferta Quelch, 1866 (= 

?D. arbuscula). Some of Rehberg's specimens are still 

deposited at the ZMB. 

Vaughan (1900) described the commonly collected Levipalifer 

orientalis (later renamed Deltocyathus vaughani) from 

off Honshu, but in the next 32 years only one paper was 

published that included azooxanthellate corals from the 

Japanese region: Van der Horst's (1922) report on the 

Indonesian Siboga dendrophylliids. From off Japan he reported: 

Dendrophyllia jponica (later renamed D. boschmai Van 

der Horst, 1926); D. cribrosa Milne Edwards and Haime, 1851; 

D. coccinea (Ehrenberg, 1834) (= ?Cladopsammia gracilis); D. 

willeyi (Gardiner, 1899) (= ?Cladopsammia gracilis); and 

Balanophyllia gigas Moseley, 1881; as well as many other 

species from Indonesia now known to occur in Japanese waters. 

His specimens are deposited at the ZMA. 

Beginning in 1932 and continuing for 45 years until 1977 

(one year before his death) Motoki Eguchi authored 95 papers 

on fossil and Recent corals of the Japanese region, many of his 

earlier papers coauthored with Hisakatsu Yabe. The excellent 

biography by Mori (1980) lists all of Eguchi's papers and 

includes a cross reference to every mention of a species name 

made in his papers. Only five of his most significant papers are 

listed in Table 2 and discussed herein. From the standpoint of 

azooxanthellate Scleractinia, the most significant contribution 

was: "Fossil and Recent Simple Corals from Japan" (Yabe and 

Eguchi, 1942b), which must be considered as the starting point 

for serious work on northwestern Pacific azooxanthallate 

corals. It was the first attempt at a synthesis of knowledge of the 

corals from this region, including a treatment of 84 species, 14 

of them new species. Their report was based primarily on the 

collections of the vessel Soyo-Maru (1926-1930) and to a 

lesser extent from the vessels Husa-Maru, Sitito-Maru, and 

Hukui-Maru, as well as numerous fossil localities. Most of 

these specimens are deposited at the TIUS. Yabe and Eguchi's 

(1942a) revision of the 13 species of fossil and Recent 

flabellids from Japan is also significant in that it laid the 

foundation for their contemporaneous paper on the fossil and 

Recent simple corals from the same region. 

Much later in his career, Eguchi (1968) wrote "The 

Scleractinian Corals of Sagami Bay," in which he described 

and illustrated 62 azooxanthellate species and subspecies, 

including 12 new species or subspecies and one new subgenus, 

Alcockia. Unlike the previously discussed paper, this book also 

included colonial species, but was restricted to the fauna of 

Sagami Bay. The specimens that he studied were primarily 

from the collection of Emperor Hirohito and are deposited in 

the Biological Laboratory of the Imperial Household, Tokyo. 

Two other papers by Yabe and Eguchi (1932g, 1937) deserve 

mention for their comprehensive revision of the micrabaciids 

and the genus Deltocyathus, respectively, in the Japanese 

region. 

In a series of three papers, Mori and Minoura (1980, 1983) 

and Mori (1987) discussed genetic control of septal number 

using Japanese specimens of Cylindrophyllia orientalis and 

Caryophyllia compressa. Cheng (1977) reported Flabellum 

apertum (= F. apertum borealis) from several stations off 

Kyushu, and Yajima (1986) summarized all records of the Sea 

of Japan endemic Rhizopsammia minuta mutsuensis. Finally, 

Owens (1986a) described a new genus and species of 

micrabaciid from off Koshiki Island, Kyushu: Rhombopsammia 

squiresi. 

Various checklists and brief species descriptions of corals 

from the Japanese region were published by: Utinomi (1956, 

1965, 1971), Mori (1964), Kikuchi (1968), Hamada (1969), 

Okutani (1969), and Tribble and Randall (1986). These are not 

listed in Table 2 but are referenced in the Literature Cited and 

in species synonymies. Grygier (1983) and Zibrowius and 

Grygier (1985) also published records of Japanese azooxanthellates 

in the context of their being hosts for ascothoracid 

parasites. Most recently, Ogawa and Matsuzaki (1992a,b) 

published a listing of the 871 nominal species of Scleractinia 

reported from Japanese waters. 

Off northern Japan, including Hokkaido and the Kurile 

Islands, Keller (1976, 1977, 1981a, 1982) reported various 

species of Fungiacyathus, Leptopenus, Caryophyllia, and 

Deltocyathus collected by the Vityaz from bathyal depths. To 

the south of Japan, but still in the warm temperate region 

(western East China Sea and Formosa Strait), azooxanthellate 

species were reported by Zou (1981) and Cairns (1989a). To 

the west of Japan, the only reports of azooxanthellate corals 

from off South Korea - including the eastern Yellow Sea, 

Chejo Do, and the western Sea of Japan - have been the papers 

of Song (1982, 1988, 1991), who reported a total of 19 

azooxanthellate species from this region. 

In this account 102 species are reported from the temperate 

northwestern Pacific (Table 5), including 22 new records for 

the region, three new genera, 5 new species, one new 

subspecies, and 9 new combinations. Caryophyllia laevicostata 

is mentioned in the species account but not counted as a species 

record. 

For the sake of completeness, it should be noted that several 

nonima nuda have been published from the Japanese region, 

especially by Yabe and Eguchi (1932a,b) and Eguchi (1934), 

but most of these names were made available by the same 

authors in later publications. But, three that remain as nomina 

nuda are: Balanophyllia niinoi, Eupsammia miyasakiensis, and 

Letepsammia marukawai, all listed by Eguchi (1934:368). 

Another nomen nudum from Japan was listed by Blainville

(1834:354) as Dendrophyllia semiramea, a name later synonymized 

with Dendrophyllia coccinea by Van der Horst 

(1922). 

Material 

This study is based on the examination of previously 

unstudied material from nine institutional sources, herein listed 

in order of decreasing size of the contribution: USNM (167 

stations of the USFWS Albatross in the northeastern and 

northwestern Pacific, 1888-1909), ORI (a large and diverse 

collection from 41 stations of the R/V Tansei Maru cruises off 

Japan, primarily cruises KT9015, KT9202, KT9309, 1974- 

1993), CAS (a large collection made primarily off California 

and the northwestern Pacific), RBCM (a large but not diverse 

collection made off British Columbia and Alaska), SIO 

(miscellaneous deep-water collections from throughout the 

northeastern Pacific, primarily off California), ZMC (Mortensen's 

Pacific Expedition of 1914, Japanese collections), IOM 

(several unreported specimens from off the Kurile Islands), 

NMCIC (several deep-water corals from off British Columbia), 

and UA (several specimens in Prince William Sound, Alaska). 

In addition to these unstudied collections, previously 

reported specimens from the following museums were examined: 

BM (Kent, 1871; Moseley, 1881; Van der Horst, 1926); 

TIUS (Yabe and Eguchi, 1942a,b); CAS (Durham, 1947; 

Faustino, 1931); IOM (Keller, 1976, 1977, 1981a); MCZ 

(Verrill, 1864); NMW (Marenzeller, 1888b); RMNH (Milne 

Edwards and Haime, 1851); SBMNH (Durham and Barnard, 

1952); SIO (Bythell, 1986); UCPM (Durham, 1947; Faustino, 

1931); USNM (Vaughan, 1900, 1903, 1906a,b, 1941; Williams, 

1936; Durham, 1947; Owens, 1986a; Cairns, 1989a, 

1991a); YPM (Verrill, 1866, 1869); ZMA (Alcock, 1902c); and 

ZMB (Marenzeller, 1904a). 

The examination of previously reported specimens combined 

with unstudied specimens provided a good base for the 

revision of the temperate North Pacific fauna, especially those 

species from the northeastern Pacific. Additional records are 

reported herein for 23 of the 25 species from the northeastern 

Pacific, only Oculina profunda and the unusual Nomlandia 

californica remaining known only from their type specimens. 

Type material of 21 of the 25 species was examined, as well as 

the types of the 10 junior synonyms; the four unexamined types 

are presumed to be lost. The coverage of the temperate 

northwestern Pacific corals was not as complete; however, 

additional records of 89 of the 102 species are reported herein 

and types of 64 of the 102 taxa were examined. For only four 

species was no material examined, in which case a diagnosis 

was abstracted from the literature, these species being: 

Crispatotrochus niinoi (Yabe and Eguchi, 1942b); Trochocyathus 

japonicus Eguchi, 1968; Phyllangia hayamaensis 

Eguchi, 1968; and Dendrophyllia hoschmai Van der Horst, 

1926. 


Methods 

It was attempted to provide complete species synonymies, at 

least regarding records from the temperate North Pacific 

region, but it is acknowledged that various checklists and 

smaller publications may have been overlooked. The original 

description and other significant references outside the North 

Pacific are also included in the synonymies, the latter often 

being a key to the extended synonymy (chresonomy) of the 

species. Efforts were made to examine as many types as 

possible and to verify as many of the previously published 

records as possible (see "Material"), but when specimens were 

not available for study and the published accounts unclear, the 

synonymy entries and corresponding distribution records are 

queried. 

All 119 species included in this revision are described or 

diagnosed, most of them based on new material (see 

"Material"). Conventional scleractinian terminology is used in 

describing the coralla (see Wells, 1956; Cairns, 1981, 1989a; 

Figures 1, 2). One new ratio is introduced in this paper: 

PD:GCD (see "Abbreviations"). 

After long deliberation, it was decided to present the species 

account in two geographic sections rather than in a continuous 

phylogenetic sequence. The reason for this was to facilitate its 

use as a regional faunistic guide for either the northeastern or 

northwestern Pacific, only eight species being found in 

common to both regions. These eight species are presented in 

detail in the first (northeastern Pacific) section of the Species 

Account, and only cross referenced in the second section. The 

order of the plates follows the same rationale, the first 12 plates 

illustrating the northeast Pacific species, plates 13-42 illustrating 

the northwest Pacific species. Specimens illustrated on 

plates 40-42 are out of order due to the inclusion of a loan 

received after the main sequence of plates was finished. 

It is important to document which specimens were actually 

examined by the author of a systematic paper and where those 

specimens are deposited. To this end, I have segregated the 

"Material Examined" sections into "New Records" and 

"Previous Records"; the former listing previously unreported 

specimens, the latter listing specimens that have been cited in 

previous publications. A third category is added for some 

species named "Reference Specimens," for specimens examined 

of closely related but not conspecific species. In each 

section each record begins with a station number followed by 

the number of specimens in the lot, and finally the catalog 

number and/or museum of deposition. 

Holotypes and paratypes of all new species reported herein 

are deposited primarily at the USNM, some paratypes also 

deposited at the ORI. An effort was made to list the museum of 

deposition and type locality for all senior and junior synonyms 

of species treated in this account. 

A detailed list of the geographic and bathymetric ranges

NUMBER 557 

INNER 

SEPTAL 

EDGE 

CALICE 

I 

EXSERT SEPTUM 

PEDICEL 

SEPTAL FACE 

PALUS 

FASCICULAR 

COLUMELLA 

BASE 

FIGURE 1.—Cutaway diagram of a species of Caryophyllia illustrating the basic morphological features of an 

attached, solitary scleractinian (.after Cairns, 1981). 

within the temperate North Pacific is given for each species, as 

well as the extended geographic and bathymetric ranges for the 

species, if any. 

The scanning electron photomicrographs were taken by the 

author on a Cambridge Stereoscan 100. In some cases 

specimens that lacked sufficient contrast for conventional 

photography were dyed dark red and coated with a thin layer of 

sublimed ammonium chloride.

Zoogeography 

I. Northeastern Pacific 

A. Tropical (PANAMANIAN PROVINCE, including "CORTEZ PROV- 

INCE") 

B. Temperate Region 

1. Warm Temperate (CALIFORNIA PROVINCE, including "SAN 

DIEGO PROVINCE") 

2. Cold Temperate 

a. Lower Boreal (OREGON PROVINCE) 

b. Upper Boreal (ALEUTIAN PROVINCE) 

II. Northwestern Pacific 

A. Tropical (INDO-WEST PACIFIC PROVINCE) 

B. Temperate Region 

1. Warm Temperate (JAPAN PROVINCE) 

2. Cold Temperate 

a. Lower Boreal (ORIENTAL PROVINCE) 

b. Upper Boreal (KURILE and OKHOTSK PROVINCES) 

TEMPERATE NORTHEASTERN PACIFIC.—In general, the 

boundaries of the various zoogeographic regions and provinces 

discussed in this analysis follow the synthesis of Briggs (1974, 

see outline above); however, an exception was made with 

respect to the southern boundary of the warm termperate 

province of the northeastern Pacific. Briggs recognized a 

distinct warm temperate province, called the Cortez Province, 

1 halfsystem 

Pourtales Plan 

system 

FIGURE 2.—Composite cross-sectional diagram of a calice illustrating various 

septal insertion patterns: upper right system with three cycles of septa, upper 

left system with four cycles, and lower two systems with various stages of 

development of the Pourtales Plan (after Cairns and Parker, 1992). Numbers 

refer to cycle to which septa belong. 


TABLE 3.—Azooxanthellate Scleractinia known from the Gulf of California 

Aslrangia haimei Verrill, 1866 

Astrangia coslata Verrill, 1866 

Astrangia conferia Verrill, 1870a 

Paracyathus stearnsii Verrill, 1869 

Coenocyalhus bowersi Vaughan, 1906 

Ceratotrochus franciscanus Durham and Barnard, 1952 

Phyllangia consagensis (Durham and Barnard, 1952) 

Desmophyllum dianthus (Esper, 1794) 

Balanophyllia cedrosensis Durham, 1947 

Endopachys grayi Milne Edwards and Haime, 1848b 

Dendrophyllia oldroydae Oldroyd, 1924 

Tubastraea coccinea Lesson, 1829 

that included the waters of the Gulf of California from La Paz, 

Baja California on the west coast of the Gulf to Topolobampo, 

Sinaloa, Mexico on the eastern coast. However, most zoogeographers 

(e.g., Ekman, 1953; Squires, 1959; Brusca, 1980) 

consider the Gulf of California to be an attenuated tropical or 

subtropical fauna. This categorization is supported by the 

affinities of the 12 azooxanthellate species found in the Gulf 

(Table 3). Aside from three species that are cosmopolitan, 

seven of the remaining nine species known from the Gulf are 

tropical species with their northern limits in the Gulf, or 

eurythermic tropical species having their northern limits 

slightly farther north in the warm temperate region. Only one 

species, Paracyathus stearnsii, is primarily temperate in 

distribution, and one species, Ceratotrochus franciscanus, is 

endemic to the Gulf. The endemic nature of the latter species 

must be viewed with doubt because it is known from only one 

specimen. 

The northern boundary of the northeastern Pacific warm 

temperate California Province is generally regarded as Point 

Conception, California, but its southern boundary on the 

Pacific coast of Baja California is less well defined. Various 

zoogeographers have placed it as far south as Cabo San Lucas 

and as far north as Bahia Sebastian Vizcaino (see Briggs, 

1974); however, the latitude of Isla Magdalena (24°40'N) is 

most often cited, and is adopted in this analysis. At least one 

tropical azooxanthellate, Astrangia conferta, has its northern 

limit at Isla Magdalena, and this latitude also coincides with the 

northern limit of zooxanthellate corals on the northeastern 

Pacific coast of Baja California (Squires, 1959; Wilson, 1988, 

1991). Seventeen scleractinians are known to occur in the 

California Province (Table 4), of which four are cosmopolitan 

and one, Crispatotrochusfoxi, has a disjunct distribution in that 

province and the Aleutian Province. Two species are endemic 

to the province (Nomlandia calif or nica and Dendrophyllia 

californica), but both of these are known from very few records. 

One species, Leptopenus discus, appears to be bitemperate, 

occurring in the northern and southern temperate regions. The 

remaining nine species are an almost equal mixture of five 

eurythermic tropical species and 4 eurythermic temperate 

species, consistent with Briggs' (1974) characterization of this 

province as a transition zone between the tropical and 

temperate regions. Rocas Alijos, a small group of rocks 343 km 

west of Isla MagdaJena at 24°57'N, was included in the warm

NUMBER 557 

TABLE 4.—Distribution of the 25 species of Temperate Northeastern Pacific Azooxanthellate Scleractinia. 

Species 

Fungiacyathus marenzelleri 

Leptopenus discus 

Astrangia haimei 

Ocutina profunda 

*Madrepora oculata 

Caryophyllia arnoldi 

Caryophyllia alaskensis 

*Caryophyllia sp. A 

Labyrinthocyathus quaylei 

Crispatotrochus foxi 

Paracyathus stearnsii 

Paracyathus montereyensis 

Coenocyathus bowersi 

Nomlandia californica 

Desmophyllum dianthus 

Lophelia pertusa 

*Flabellum sp. A 

*Javania cailleti 

*Javania borealis 

*Javania californica 

Polymyces montereyensis 

Balanophyllia elegans 

Balanophyllia cedrosensis 

Dendrophyllia oldroydae 

Dendrophyllia californica 

Totals 

Tropical 

Eastern 

Pacific 

X 

X 

X 

X* 

X 

X 

X 

X 

X 

X 

- 

Warm 

California 

Province 

* New record for temperate northeastern Pacific. 

1 Depth range in meters for records in northeastern Pacific. 

2 Presence based on very few records. 

3 Gulf of California only. 

temperate province by Briggs (1974), but both of the coral 

species that occur there (Madracis sp. cf. M. pharensis and 

Phyllangia consagensis, fide Wilson, in press, 1994) are 

characteristic of the tropical region, and thus are not considered 

in this account. Three species are known from Isla Guadelupe 

260 km off the coast of Baja California at about 29°N: 

Coenocyathus bowersi, Desmophyllum dianthus, and Lophelia 

pertusa (one eurythermic tropical and two cosmoplitan species, 

respectively). 

The cold temperate region of the northeastern Pacific is 

divided into two provinces: the lower boreal Oregon Province 

and the upper boreal Aleutian Province. According to Briggs, 

the Oregon Province extends from Point Conception, California 

to Dixon Entrance, just north of the Queen Charlotte 

Islands, British Columbia. Sixteen species are known to occur 

in this province (Table 4), including four cosmopolitan and 

three endemic species: Javania californica, Paracyathus 

montereyensis, and Oculina profunda; the latter two are known 

from very few records. The bitemperate Leptopenus discus is 

also known from this province. Of the remaining eight species, 

three are considered as eurythermic tropical species having 

X 

X 

X 

X 

X 

X 

X 2 

X 

X 

X 2 

X 

X 

X 

X 

X 

X 

X 2 

17 

Temperate 

Oregon 

Province 

X 

X 

X 

X 2 

X 

X 

X 

X 

X 2 

X 

X 

X 

X 

X 

X 

X 

16 

Cold 

Aleutian 

Province 

X 

X 

X 

X 

X 2 

X 

X 2 

X 

X 

9 

Temperate 

Northwest 

Pacific 

X 

X 

X 

X 

X 

X 

X 

X 

8 

Elsewhere 

Cosmopolitan 

Subantarctic 

Cosmopolitan 

Cosmopolitan 

Cosmopolitan 

? Antarctic 

Cosmopolitan 

Depth 

2999-6328 

3599-5000 

1-53 

119-742 

84-488 

40-656 

102-399 

247 

37-293 

82-274 

20-134 

75-146 

9-302 

82 

33-1097 

82-488 

55-507 

1280-1371 

247-348 

62-170 

69-212 

0-293 

66-119 

40-576 

42-93 

their northern limit at Monterey Bay, California, and five 

species are characteristic of the northeastern Pacific temperate 

region. 

The upper boreal Aleutian Province is considered to extend 

from Dixon Entrance to the western tip of the Aleutian Islands 

(Briggs, 1974:280). This province contains only nine coral 

species (Table 4): one cosmopolitan, one endemic (Caryophyllia 

sp. A), one disjunct {Crispatotrochus foxi), two bipolar or 

bitemperate species (Leptopenus discus and Flabellum sp. A), 

and four species exclusively northern temperate in distribution. 

In the latter category, Javania borealis also occurs in the 

northwestern Pacific upper boreal Kurile Province; Caryophyllia 

alaskensis also occurs in the lower boreal (Oregon 

Province); and two species (Caryophyllia arnoldi and Balanophyllia 

elegans) have distributions encompassing all three 

provinces in the northeastern Pacific temperate region. 

The northernmost record of a scleractinian species in the 

Pacific occurs in the Aleutian Province, consisting of several 

records of Caryophyllia arnoldi from Prince William Sound, 

Alaska (60°48'N). Several species occur throughout the 

Aleutian and Commander Islands, but none are known to occur

10 


TABLE 5.—Distribution of the 102 species of Temperate Northwestern Pacific Azooxanthellate Scleractinia. 

Species 

Madracis sp. A 

*Fungiacyathus stephanus 

Fungiacyathus paliferus 

Fungiacyathus sp. A 2 

Fungiacyathus marenzelleri 

*Fungiacyathus variegatus 

* Fungiacyathus granulosus 

Leptopenus discus 

Ltptopenus solidus 2 

Letepsammia formosissima 

Rhombopsammia niphada 2 

Stephanophyllia fungulus 

Culicia japonica 

Oulangia stokesiana miltoni 

Madrepora oculata 

Cyathelia axillaris 

Anthemiphyllia dentata 

*Anthemiphyllia frustum 2 

Caryophyllia japonica 

Caryophyllia alaskensis 

Caryophyllia cf. scobinosa 

Caryophyllia quadagenaria 

Caryophyllia rugosa 

Caryophyllia jogashimaensis 2 

Caryophyllia a. ambrosia 

Caryophyllia (A.) grayi 

Caryophyllia (A.) spinigera 2 

Caryophyllia (P.) compressa 

Crispatotrochus rubescens 

Crispatotrochus niinoi 2 

Paracyathus pruinosus 

Trochocyathus caryophylloides 

Trochocyathus japonicus 2 

*Trochocyathus decamera 2 

'Trochocyathus cooperi 2 

Deltocyathus vaughani 

*Deltocyathus rotulus 

Deltocyathus magnificus 

Stephanocyathus (A.) spiniger 

*Stephanocyathus (O.) weberianus 

Conotrochus funicolumna 

'Aulocyathus recidivus 2 

Aulocyathus matricidus 

Desmophyllum dianthus 

*Lophelia pertusa 

Anomocora sp. 2 

*Coenosmilia cf. arbuscula 2 

Phyllangia hayamaensis 2 

Rhizosmilia sagamiensis 

Dasmosmilia pacifwa 

Goniocorella dumosa 

*Notocyathus venustus 2 

Notocyathus conicus 

Peponocyathus australiensis 

Peponocyathus folliculus 

Tropidocyathus lessoni 

Tropidocyathus pileus 

Tropical Warm 

Temperate 

Cold Temperate 

ndo-West Japan Oriental Kurile Northeast 

Pacific Province Province Province Pacific Elsewhere Depth 1 

Cosmopolitan 

S. Australia 

x Cosmopolitan 

x Subantarctic 


Atlantic 

Maldives 

S. Australia 

S. Australia 

S. Australia 

S. Australia 



? Atlantic 

New Zealand 

Cosmopolitan 

Atlantic 

S. Australia 

55-110 

446-1317 

70-364 

3175-4110 

4690-5450 

422-715 

402-410 

3175-3250 

3175-3250 

77-307 

660-783 

15-256 

5-100 

0-135 

72-549 

15-366 

50-499 

237-241 

77-1680 

223 

119-805 

70-422 

71-240 

52-98 

311-2450 

108-191 

9 

115-422 

110-344 

104 

176-198 

115-344 

150-450 

70-88 

80-88 

88-1097 

799-1187 

88-422 

106-783 

715-1302 

88-600 

741 

84-207 

77-715 

150-340 

55-100 

238-240 

5.5 

60-98 

168-355 

100-366 

193-422 

70-110 

59-494 

30-402 

98-155 

123-422

NUMBER 557 11 

Species 

*Alatotrochus rubescens 2 

Idiotrochus kikulii 

Stenocyathus vermiformis 2 

Truncatoguynia irregularis 

Flabellum pavoninum 

Flabellum patens 

Flabellum magnificum 

Flabellum angustum 2 

Flabellum politum 

Flabellum deludens 

Flabellum japonicum 

Flabellum apertum borealis 

Truncatoflabellum spheniscus 

Truncatoflabellum candeanum 

Truncatoflabellum formosum 

Truncatoflabellum carinatum 

*Truncatoflabellum gardineri 

*Truncatoflabellum sp. A 2 

^Truncateflabellum sp. B 2 

*Placotrochides scaphula 

Javania insignis 

Javania cailleti 

*Javania borealis 

Rhizotrochus typus 

Balanophyllia cumingii 

Balanophyllia cornu 

Balanophyllia gigas 

Balanophyllia ponderosa 

Balanophyllia sp. A 2 

*Balanophyllia teres 

Endopachys grayi 

*Eguchipsammia gaditana 

Eguchipsammia wellsi 

Rhizopsammia minuta mutsuensis 

Cladopsammia gracilis 

Cladopsammia eguchii 

Dendrophyllia ijimai 

Dendrophyllia cribrosa 2 

Dendrophyllia japonica 

Dendrophyllia arbuscula 

Dendrophyllia boschmai 

Dendrophyllia florulenta 

Enallopsammia rostrata 

Tubastraea coccinea 

*Schizopsammia songae 2 

Totals 

Tropical 

Indo-West 

Pacific 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

_ 

TABLE 5.—Continued. 

Warm 

Japan 

Province 

* New record for the temperate northwestern Pacific. 

1 Depth range in meters for records in northwestern Pacific. 

2 Presence based on very few records. 

in the Bering Sea proper. 

Eight of the 25 temperate northeastern Pacific coral species 

are common to both the northeastern and northwestern Pacific 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

92 

Temperate 

Cold 

Oriental Kurile 

Province Province 

X 

X 

X 

X 

X 

X X 

X 

X 

X 

X 

X X 

X 

24 8 

Temperate 

Northeast 

Pacific Elsewhere 

Atlantic 

? E. Atlantic 


X 

8 

E. Pac. tropics 

Atlantic 

E. Pac. tropics 

E. Atlantic 

New Zealand 

Widespread 

Atlantic, E. 

Pac. tropics 

Depth 1 

193-422 

? 

300 

80-161 

73-658 

223-402 

307-700 

7 

70-402 

115-783 

119-631 

307-1141 

66-106 

88-223 

106-210 

30-274 

100-144 

964-1031 

80-88 

80-457 

46-249 

539-785 

348 

20-104 

65-307 

60-274 

115-245 

14-16 

56-100 

154-237 

70-274 

110-188 

110-1% 

0-2 

3-95 

7 

10-200 

7-40 

115-250 

40-240 

40-165 

70-110 

270-331 

0-15 

(Tables 4 and 5), but six are cosmoploitan; only two species 

appear to be characteristic of the North Pacific temperate 

region: Caryophyllia alaskensis and Javania borealis. But, of 

7

12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 

the 18 scleractinian genera known from the temperate 

northeastern Pacific, 12 (67%) are also represented in the 

northwestern Pacific. 

TEMPERATE NORTHWESTERN PACIFIC.—The warm temperate 

northwestern Pacific Japan Province is dominated by the 

northerly flowing tropical Kuroshio Current. Its boundaries are 

quite complicated because of the complex geography of the 

region and the variations in the path of the Kuroshio (Veron, 

1992). According to Briggs (1974), the southern limit of the 

warm temperate province is Hong Kong, China and includes 

the Chinese mainland to Wenling, the west coast of Taiwan, 

Formosa Strait, and most of the East China Sea, but not the 

southern Ryukyu Islands. The northern boundaries of the Japan 

Province include the southern tip of South Korea (Haenam to 

P'ohang), Korea Strait (including Tsushima and Mishima), the 

southwestern tip of Honshu to about the latitude of Hamada, the 

northern Ryukyu Islands from Tokara Retto northward, and the 

east coast of Honshu to Cape Inubo. Unlike the warm temperate 

northeastern Pacific in which zooxanthellate corals are absent, 

zooxanthellate corals are present in the Japan Province to its 

northern limits (Eguchi, 1968; Tribble and Randall, 1986; 

Veron, 1992). The 92 azooxanthellate species known from this 

province (Table 5) consist of six cosmopolitan species, 21 

(23%) endemic species, 58 (63%) eurythermic tropical species, 

six species characteristic of the boreal temperate region, and 

one species that remains unclassified {Stenocyathus vermiformis). 

Of the 21 endemic species, 13 are poor indicators of 

endemism as they are known from only one or two records, but 

the remaining eight species appear to be characteristic of this 

province: Deltocyathus vaughani, Rhizosmilia sagamiensis, 

Dasmosmilia pacifica, Phyllangia hayamaensis, Flabellum 

apertum borealis, Truncatoguynia irregularis, Balanophyllia 

teres, Eguchipsammia wellsi, and Dendrophyllia boschmai. 

Among the eurythermic tropical species, many conform to the 

previously stated northern border of the warm temperate 

province (i.e., southern South Korea, Mishima, Cape Inubo), 

including: Letepsammia formosissima, Culicia japonica, 

Cyathelia axillaris, Caryophyllia quadragenaria, Javania 

insignis, and Endopachys grayi; however, the range of at least 

six species extend slightly farther north into the cold temperate 

Sea of Japan coast of Honshu as far as Oki Channel (e.g., 

Stephanophyllia fungulus, Fungiacyathus paliferus, Anthemiphyllia 

dentata, Flabellum deludens, Truncatoflabellum 

formosum, and T. gardineri), and two more species penetrate to 

Wakasa Bay {Rhizotrochus typus and Balanophyllia cumingii). 

Several other species reported from this province are primarily 

tropical with only marginal occurrences in the southern warm 

temperate province, such as: Tropidocyathus lessoni, Stenocyathus 

vermiformis, Truncatoflabellum sp. A, T. sp. B, and 

Enallopsammia rostrata. The six cold temperate species found 

in the Japan Province are: Oulangia stokesiana miltoni, 

Caryophyllia compressa, C. japonica, Aulocyathus matricidus, 

Balanophyllia sp. A, and Rhizopsammia minuta mutsuensis. 

According to Briggs (1974:267), the lower boreal cold 

temperate Oriental Province includes the Yellow Sea and much 

of the Sea of Japan, from P'chang, South Korea to Oh'ongjin, 

North Korea to the west, Hamada to Tsugaru Strait on the west 

coast of Honshu, and from Cape Inubo to Tsugaru Strait on the 

east coast of Honshu. Of the 24 scleractinian species known to 

occur in this province (Table 5), the largest component (11 

species, 46%) is that of eurythermic tropical species, most of 

which are found only a short distance into the province along 

the west coast of Honshu (e.g., Oki Channel, Wakasa Bay, see 

above). Of the remaining 13 species, five are cosmopolitan, 

none are endemic, six are characteristic of the northwestern 

temperate region, and two are unclassified {Caryophyllia 

alaskensis and Dendrophyllia japonica). Although no endemics 

are known, Rhizopsammia minuta mutsuensis typifies this 

province, being found only slightly north and south of the 

stated borders of the province. Of the six temperate species, 

four are found in the Japan and Oriental Provinces {Oulangia 

stokesiana miltoni, Carophyllia compressa, Balanophyllia sp. 

A, and Aulocyathus matricidus), the other two being characteristic 

of all three provinces of the northwestern Pacific region: 

Caryophyllia japonica and Rhizopsammia minuta mutsuensis. 

Only eight species of Scleractinia are known from the upper 

boreal cold temperate Kurile Province (Table 5), defined as the 

northern Sea of Japan, off Hokkaido, and the eastern coasts of 

the Kurile Islands and Kamchatka Peninsula. The affinities of 

this fauna are quite similar to the upper boreal of the 

northeastern Pacific Aleutian Province, consisting of: two 

cosmopolitan species, one endemic {Fungiacyathus sp. A), one 

bitemperate species {Leptopenus discus), two northwestern 

temperate species {Caryophyllia japonica and Rhizopsammia 

minuta mutsuensis), one upper boreal endemic (Javania 

borealis), and one species having a disjunct distribution in the 

tropical and boreal regions {Leptopenus solidus). 

A second, upper boreal cold temperate province is recognized 

by Briggs, the Okhotsk Province, which includes the Sea 

of Okhotsk, but no Scleractinia have been reported from this 

province. 

COMPARISONS.—According to the zoogeographic synthesis 

of Briggs (1974:196), warm temperate marine faunas are 

usually strongly influenced by the adjacent eurythermic 

tropical species or simply by dispersal of tropical species that 

happen to be carried to higher latitudes by strong currents. This 

would be especially true of northwestern boundary currents, 

wherein a warm northerly flowing current (e.g., Gulf Stream, 

Kuroshio) brings tropical species into northern latitudes. This 

phenomenon is clearly seen in the northwestern Pacific warm 

temperate Japan Province wherein 58 (63%) of the 92 species 

are eurythermic tropical species, but less evident in the 

California warm temperate province wherein only 5 (29%) of 

the 17 species are considered as eurythermic tropical. The 

much higher number of azooxanthellate species in the Japan 

Province (92) compared to that of the California Province (17) 

is undoubtedly also due to their proximity to tropical faunas of 

vastly different sizes, i.e., the estimated number of tropical 

eastern Pacific azooxanthellates (Durham and Barnard, 1952; 

Squires, 1959; Cairns, 1991a) is about 55; that of the Indo-West

NUMBER 557 13 

Pacific region, about 400. On the other hand, the influence of 16, difference = 8). 

temperate species is larger in the northeastern Pacific warm The northernmost, adjacent upper boreal provinces (Aleutian 

temperate (4 species, 23.5%) than in the northwestern warm and Kurile) have virtually the same number of species and 

temperate (6 species, 6.5%), no doubt caused by the cool, similar ZOOgeographic affinities, although they have only one 

southerly flowing California Current coming from higher noncosmopolitan species in commori) Javania horealis. Both 

latitudes. 

„, . , . . , , . . . 

The number of species in the northwestern lower boreal 

~ . . , „ . j , t l. ... j . i . 

Onental Province drops dramatically with regard to the 

adjacent Japan Province, primarily due to the great reduction of 

. . „ .. . . . .. . 

provinces have small cosmopolitan, endemic, and bipolar 

. . . , . 

components, and are characterized by species that are restricted 

. 

t0 their "SJ** cold temperate regions. It was also noted that 

the eurythermic tropical species from 58 to 11 and the complete most of tne "P^ 1 " ^real s P ec[es nave relatively deep 

absence of endemic species. The paucity of collections from bathymetric ranges, often exceeding 500 m. In fact, the deepest 

this region probably also contributes to its low known diversity. record of a scleractinian coral occurs in the Aleutian Province: 

The sister Oregon Province in the northeastern Pacific has only Fungiacyathus marenzelleri (reported as F. symmetricus 

one fewer species than its adjacent warm temperate province to aleuticus by Keller, 1976) at 6296-6328 m in the western 

the south (16 vs 17) and shows a greater affinity with cold Aleutian Trench (53°37'N, 159°40'W). 

temperate regions (5 species, 31%) than with the tropical region When amphi-Pacific provinces are combined and species 

(3 species, 18%). The lesser influence of eurythermic tropical held in common accounted for, a total of 106 azooxanthellate 

species in the Oregon Province compared to the Oriental species are known from the warm temperate North Pacific 

Province (11 vs 3, difference = 8) essentially accounts for the provinces, 37 in the lower boreal cold temperate provinces, and 

difference in number of species of these two provinces (24 vs only 14 in the upper boreal cold temperate provinces. 

SYSTEMATIC ACCOUNT 

Part 1: Temperate Northeastern Pacific 

Key to the 25 Species of Temperate Northeastern Pacific Scleractinia 

1. Corallum solitary 2 

Corallum colonial 19 

2. Corallum attached (fixed) 3 

Corallum unattached (free) 17 

3. Columella present 4 

Columella absent, or only a rudimentary basal fusion of lower, inner septal edges 

12 

4. Corallum discoidal, without a thecal wall Nomlandia californica 

Corallum conical (trochoid, ceratoid), with a thecal wall (septo- or synapticulothecate) 

5 

5. Theca rough and porous; septa arranged in a Pourtales Plan 6 

Theca costate or smooth, not porous; septa normally inserted 7 

6. Columella flat to slightly concave; distinctive stellate pattern of septal arrangement; 

Alaska to Baja California (29°44'N) Balanophyllia elegans 

Columella convex; Pourtales Plan not so pronounced, stellate pattern not obvious; 

Cedros Island, Baja California southward Balanophyllia cedrosensis 

7. Pali or paliform lobes present 8 

Pali and paliform lobes absent 11 

8. Paliform lobes present before all but last cycle of septa (e.g., P1_3), often divided 

and indistinguishable from columella 9 

Pali occur only before penultimate cycle of septa (e.g., usually 12 P3) 10 

9. Septa hexamerally arranged in 5 of more cycles (>96 septa); septa crowded; 3 or 4 

palar crowns Paracyathus stearnsii 

Septa decamerally arranged in 3 to 4 cycles (42-46 septa); septa well spaced; 2 

palar crowns Paracyathus montereyensis 

10. Pedicel thick (up to 65% GCD); costae granular, calicular septal apices acute; San 

Diego to Gulf of Alaska Caryophyllia arnoldi 

Pedicel thinner (18%-33% GCD); theca porcellaneous; calicular septal apices 

equilateral; British Columbia to Korea Strait Caryophyllia alaskensis


11. Columella labyrinthiform, composed of short interconnected lamellae; inner septal 

edges straight; theca smooth Labyrinthocyathus quaylei 

Columella fascicular, composed of twisted laths; inner septal edges slightly sinuous 

Crispatotrochus foxi 

12. Theca costate and granular, often brown to grey Desmophyllum dianthus 

Theca smooth and porcellaneous, white 13 

13. Pedicel reinforced by 12-24 hollow rootlets, best seen in cross section of lower 

pedicel Polymyces montereyensis 

Pedicel does not contain rootlets: either slender or stereome-reinforced . . . . 14 

14. Pedicel not reinforced with stereome and therefore relatively narrow (25% 

GCD) 15 

15. Corallum relatively small (

NUMBER 557 15 

septa (see Cairns, 1991a). 

TYPE SPECIES.—Fungia symmetrica Pourtales, 1871, by 

monotypy. 

Fungiacyathus (Bathyactis) marenzelleri (Vaughan, 1906) 

PLATE la-/ 

Bathyactis symmetrica.—Moseley, 1881:189 [in part: Challenger-24\, 244].— 

Marenzeller, 1904b:76 [in part: only Alb-3376, specimens from the other 

three stations being too fragmentary to identify].—Not Yabe and Eguchi, 

I942b:\31 [=F.paliferus]. 

Bathyactis marenzelleri Vaughan. 1906b:66-67, pi. 4: figs. 1-lb. 

Fungiacyathus symmetricus aleuticus Keller, 1976:39-41, pi. 2: figs. 1-5; pi. 

3: figs. 1-19. 

Fungiacyathus symmetricus fragilis Keller, 1976:41-43 [in part: W/yaz-4158, 

4191]. 

Fungiacyathus marenzelleri.—Cairns, 1979:35-37, pi. 2: figs. 8, 9; pi. 3: figs. 

3, 8 [synonymy]; 1982:5-7, pi. 1: figs. 1, 2, 8 [synonymy].—Zibrowius, 

1980:24-25, pi. 6: figs. A-M; pi. 7: figs. A-K [synonymy]. 

Fungiacyathus sp. "a".—Zibrowius and Grygier, 1985:119 [Challenger-24\]. 

DESCRIPTION.—Corallum fragile, light, and discoidal, having 

a very thin (0.15 mm), flat to slightly convex, circular base 

up to 27 mm in diameter in North Pacific specimens. Costae 

thin, finely serrate, slightly sinuous ridges up to 1 mm in height 

near calicular edge. Length and height of costae progressively 

decrease in size according to formula: C1_2>C3>C4. Corallum 

white or light brown. 

SU2 extend from center of calice, whereas inner edges of 

each pair of S3 fuse with the adjacent S2 about one-fourth 

radius from center of calice in a short triangular canopy. 

Likewise, each pair of S4 fuse with their adjacent S3 about half 

radius distance from calicular center through more elongate 

canopies. S1 largest and only independent septa, composed of 

7-10 trabeculae. Innermost 3 or 4 trabeculae of S1 project as 

inwardly curved carinate spines, the outermost 5 or 6 trabeculae 

forming a tall (up to 11 mm in height) solid peripheral lobe. 

Septal faces carinate, each trabeculum producing a curved, 

slightly serrate ridge up to 0.4 mm in height and symmetrically 

arranged on each side of septum. Trabecular carinae on St 

widely spaced (up to 1.2 mm apart) but occasionally doubled, 

two being directly adjacent. S2 consist of about 8 trabeculae, 

have a much lower peripheral lobe, and have only 5 or 6 inner 

spines. S3 consist of 4 or 5 trabeculae in a single peripheral 

lobe; S4 consist of 3 or 4 trabeculae in a low peripheral 

lamellae. The third trabecular spines from caliclular center of 

the 12 S^ are the most exsert, projecting up to 2.5 mm above 

upper septal edge. Depending on diameter of corallum, 6-11 

synapticulae occur per S1t becoming progressively larger 

toward calicular edge, where they intersect the curved 

trabecular carinae at an oblique angle (Plate \f), the outermost 

synapticulae rising well above the S4. Columella variable in 

development and sometimes even absent, but usually consists 

of a thin, horizontal circular plate up to 6 mm in diameter, 

through which project the innermost S:_2 trabecular spines. 

DISCUSSION.—Most of Keller's (1976) types off. symmetricus 

aleuticus were examined and found to be conspecific with 

F. marenzelleri, a species of which she was apparently 

unaware. Her (Keller, 1976) two Pacific records of F. 

symmetricus fragilis were examined and also found to be 

conspecific. The latter two records were inadvertently symbolized 

as F. symmetricus aleuticus on her distributional map 

(Keller, 1976:32). Her new subspecies F. s. fragilis is a junior 

primary homonym of F. fragilis Sars, 1872, but may not 

require a replacement name since her type series includes 

several previously described species. 

Approximately 14 species comprise the subgenus Bathyactis 

(see Cairns, 1989a). Fungiacyathus marenzelleri is distinguished 

from other species by its large corallum size and by 

having straight septa with relatively few, widely-spaced 

trabecular ridges. Only one other species of Fungiacyathus is 

known from the temperate North Pacific, a species reported as 

F. paliferus by Keller (see Fungiacyathus sp. A in northwest 

Pacific account), which is easily distinguished from F. 

marenzelleri by having five cycles of septa. 

Over 100 specimens of F. marenzelleri were collected by the 

PULSE cruises of SIO, which repeatedly sampled one abyssal 

locality about 220 km off San Luis Obispo Bay, California at 

4100 m. F. marenzelleri is one of the few scleractinians known 

from that depth in the North Pacific and holds the depth record 

for a scleractinian coral at 6328 m (Wryaz-4120) in the Aleutian 

Trench (Keller, 1976). 

MATERIAL EXAMINED.—New Records: Alb-4397, 1, 

USNM 47467; Ve/ero-4-7228-60, 2, USNM 80119; Vityaz- 

5626, 2, USNM 92439; PULSE 1-124, 3, USNM 85790; 

PULSE 2-216, 25, USNM 87616; PULSE 2-224, 9, USNM 

87615; PULSE 3-312, 1, USNM 92440; PULSE 3-314, 15, 

USNM 92441; PULSE 4-426,8, USNM 92442; PULSE 5-505, 

10, USNM 92443; PULSE 6-606, 3, USNM 92444; PULSE 

7-721, 5, USNM 92445; PULSE 8-803. 30, USNM 92446; 

PULSE 9-907, 15, USNM 92447; PULSE 9-910, 8, USNM 

92448; PULSE 10-1007, 10, USNM 92449; PULSE 10-1017, 

6, USNM 92450; MV 66-II-4, 1, SIO Co 1272; Washington 

MV 67-111-22, 15, SIO Co 1269; Melville MV69-VI-9, 1, SIO 

Co 1288; Melville 70-22, 15, SIO Co 1270; Melville 

MV70-III-6, 10, SIO Co 1268; Horizon MET-133, 5, SIO Co 

944. Previous Records: Alb-3376, 1, USNM 22078 (Marenzeller, 

1904b); types of F. marenzelleri Vaughan, 1906b, 

Alb-4721 (USNM), Alb-4670 (MCZ); types of/ 7 , symmetricus 

aleuticus Keller, 1976: Vityaz-2074, 3166 (5 at USNM 92436), 

4120, 5634, 5605 (Plate \e), 5623, 5621, 5622, 5624 (10 at 

USNM 92437), 6136, 6143 (Plate \c,d), 6142, 6088 (7 at 

USNM 92438), unless otherwise indicated deposited at IOM; 

types of F. symmetricus fragilis Keller, 1976: V/fyaz-4158, 

deposited at IOM; specimens listed by Cairns (1979,1982) and 

Zibrowius (1980). 

TYPES.—The holotype of F. marenzelleri is deposited at the 

USNM (47415). Three paratypes from Alb-4670 are deposited 

at the MCZ. Type Locality: Alb-4721: 80°7.5'S, 104°10.5'W 

(off Peru), 3820 m. 

The holotype and most paratypes (Plate \c-e) of F. 

symmetricus aleuticus Keller, 1976 are deposited at the IOM,


some being present at the USNM (see Material Examined). 

Type Locality: Vityaz-6136: 53°25'N, 163°23'W (Aleutian 

Trench off the Fox Ids.), 4620 m. 

DISTRIBUTION.—Widespread in Pacific from off Peru (Vaughan, 

1906b) to Colombia (Marenzeller, 1904b); off Pacific 

coast of Baja California, California, and Washington (Keller, 

1976); Aleutian, Kurile-Kamchatka, and Japan Trenches 

(Keller, 1976); Shatskiy Rise (Moseley, 1881). Elsewhere: 

Bahamas and off Cuba (Cairns, 1979); eastern 

Atlantic (Zibrowius, 1980); Subantarctic and off continental 

Antarctica (Cairns, 1982); 300-6328 m, but most records from 

North Pacific fall between 3800-5000 m. 

Family MICRABACIIDAE 

Leptopenus Moseley, 1881 

DIAGNOSIS.—Corallum solitary, discoidal, and free; extremely 

fragile. Synapticulotheca horizontal and extremely 

porous. Costae and septa alternate in position. Septa rudimentary, 

composed primarily of a series of tall spines. Pali absent; 

columella spiny. Polyp completely invests corallum. Exclusively 

azooxanthellae and deep water in habit. 

TYPE SPECIES.—Leptopenus discus Moseley, 1881, by 

subsequent designation (Wells, 1936). 

Leptopenus discus Moseley, 1881 

PLATE la-d 

leptopenus discus Moseley, 1881:205-208, pi. 14: figs. 1-4; pi. 16: figs. 

1-7.—Caims, 1979:37-38, pi. 3: figs. 4-7 [synonymy].—Not Leptopenus 

discus.—Squires, 1965:878. fig. 1 [=L. solidus Keller, 1977]; 1967:505. 

Leptopenus irinae Keller, 1977:38-40, pi. 1: figs. 1-4, 6, text-fig. 2. 

Not Leptopenus sp. cf. L. discus.— Caims, 1982:9 [= L. antarcticus Caims, 

1989a]. 

DESCRIPTION.—Corallum discoidal and extremely fragile, 

the peripheral costal spines invariably broken during collection. 

Largest specimen 25 mm in diameter, the height varying from 

1-4 mm, resulting in the relatively great range of D:H ratio of 

5-17. Base flat, the costae alternating in position with septa. 

Septa and adjacent costae joined to one another by thin 

synapticular bridges, each separated by large pores up to 0.5 

mm long and 0.25-0.30 mm wide. Six major costae radiate 

from a circular central region about 0.5 mm in diameter, each 

costa soon dividing twice resulting in 4 costae in each system. 

These 24 costae further subdivide to match the number of septa 

in the corallum and ultimately extend up to one-third the 

calicular radius beyond the synapticular region (Keller, 1977). 

Septa hexamerally arranged in typical micrabaciid fashion 

(see Cairns, 1982, text-fig. 1), usually resulting in 72 septa. S! 

only independent septa, bearing 4 or 5 slender, cylindrical (0.1 

mm in diameter) spines up to 0.9 mm in height, united by a low 

septal webbing. S2 also originate at center of calice and bear 3 

tall robust spines, the one closest to center quite large: up to 0.2 

mm in diameter and over 1 mm in height, constituting the 

greatest height of the corallum. A high web unites the S2 

spines. A pair of S3 diverge from each of the 6 S2 relatively 

near columella, their union with the S2 covered by a low 

canopy. Each S3 bifurcates one to three times, depending on the 

ontogenetic stage, resulting in 48-72 septa. Examined paratype 

of L. irinae from Vityaz-4\5S (USNM 92435) relatively small 

(calicular diameter = 11.0 mm), having only 48 septa. All septal 

spines are obliquely oriented towards calicular edge, never 

curved inward toward columella, and restricted to central 

portion of corallum. Columella a spinose central mound, 

penetrated by 10-15 trabecular spines, each spine about 0.1 

mm in diameter. 

DISCUSSION.—Keller (1977) distinguished L. irinae from L. 

discus by its long costal ribs that extend one-third the radius 

beyond the synapticular region. Although these peripheral 

costae are usually lost during collection, Moseley (1881, pi. 14: 

figs. 1,2) clearly illustrated these costal extensions forL. discus 

extending at least 20% of the radius beyond the synapticular 

zone. The paratype from Vityaz-4158 appears to be identical to 

the syntypes of L. discus previously examined from the British 

Museum. 

Four other Recent species of Leptopenus are known. 

Leptopenus hypocoelus Moseley, 1881 is easily distinguished 

by its very tall corallum (D:H = about 2), its enormous S2 

spines, and by having only 48 septa. Leptopenus antarcticus 

Caims, 1989a also differs in having a low D:H ratio (about 5), 

a concave base, inwardly curved septal spines, and rudimentary 

or absent septal canopies. Likewise, Leptopenus sp. A Cairns, 

1989a, differs in having 96 septa, rudimentary or absent septal 

canopies, and inwardly curved septal spines. Finally, L. solidus 

Keller, 1977, differs in having smaller thecal pores, a much 

more solid base, and by having incurved spines. 

The seven specimens collected by Scripps vessels from four 

abyssal locations west of the Patton Escarpment, off northern 

Baja California (3599-3950 m) are only tentatively assigned to 

this species. All specimens were fragmentary, held together 

only by abundant tissue; bleaching of these specimens would 

have led to disarticulation of the broken fragments. 

MATERIAL EXAMINED.—New Records: Melville 70-22, 3, 

SIO Co 1270; Melville 70-III-1, 1, SIO Co 1271; Melville 

70-III-6, 1, SIO Co 1268; Horizon MET-123, 2, SIO Co 927. 

Previous Records: Syntype of L. discus from Challenger- 

147, BM 1880.11.25.159; paratype of L. irinae from Vityaz- 

4158, USNM 92435; specimens reported by Cairns (1979). 

TYPES.—Four syntypes of L. discus collected from three 

Challenger stations (147, 157, and 323) are deposited at the 

BM, the specimen from Challenger-\41 cataloged as: BM 

1880.11.25.159. Type Locality: Southern Indian Ocean and 

southwest Atlantic; 2926-3566 m. 

The holotype and 12 paratypes (Plate Id) of L. irinae Keller, 

1977 are deposited at the IOM. The holotype was collected 

from V7ryaz-6143, the paratypes from Vityaz stations 4158, 

5603, and 6142. One paratype from Vityaz-4\5& is also 

deposited at the USNM (92435). Type Locality: S

NUMBER 557 17 

163°00'W (Aleutian Trench), 4820 m. 

DISTRIBUTION.—?Off Baja California (west of Patton Escarpment); 

off Washington (Keller, 1977); Aleutian and Kurile 

Trenches (Keller, 1977); 3599-500 m. Elsewhere: Northeast 

of Cuba (Cairns, 1979); off Rio de la Plata, Argentina 

(Moseley, 1881); near Crozet Island (Moseley, 1881); 2842- 

5000 m. 

Suborder FAVIINA 

Superfamily FAVIOIDEA 

Family RHIZANGHDAE 

Astrangia Milne Edwards and Haime, 1848c 

DIAGNOSIS.—Corallum colonial and attached, having extratentacular 

budding originating from stolons or a common basal 

coenosteum. Corallum base often polycyclic. Septa dentate. 

Pali absent; columella papillose. Primarily azooxanthellate and 

invariable found in shallow water. 

TYPE SPECIES.—Astrangia michelinii Milne Edwards and 

Haime, 1848d (= A. poculata Ellis and Solander, 1786), by 

subsequent designation (Milne Edwards and Haime, 1850: 

xliv). 

Astrangia haimei Verrill, 1866 

PLATES 2e-g, 3a-c 

Astrangia haimei Verrill, 1866:330-331; 1870a:526-527, pi. 9: figs. 6, 

6a.—Marenzeller, 19O4b:75— Durham and Barnard, 1952:71-72, pi. 6: fig. 

30.—Squires, 1959:415-418 [synonymy]. 

Astrangia sp.—Johnson and Snook, 1927:108; 1967:108-109, fig. 86. 

Astrangia insignifica.—Ricketts and Calvin, 1939:39, 264. 

Astrangia lajollaensis Durham, 1947:28-29, pi. 2: figs. 14, 15, 18, 20, 21; 

1949: 151, pi. 5: figs. 9, 14, text-figs. 5. 6.—Durham and Barnard, 

1952:74-75, pi. 7: fig. 34.—Ricketts and Calvin, 1952:37.—Fadlallah, 

1982:379-387, figs. 3, 6; 1983a:139.—Bythell, 1986:13, pi. 2: fig. B; pi. 4: 

figs. A-B.—Cairns et al., 1991:46.—Chad wick, 1991:42-47. 

DESCRIPTION.—Colonies encrust rocks and bivalve shells, 

the largest colony reported (Fadlallah, 1982) 1.75 m 2 in surface 

area, constituting approximately 14,000 corallites. Smaller 

colonies circular in outline (e.g., Plate 2/: 25 cm 2 and 90 

corallites) and reptoid colonies are more common. Corallites 

cylindrical, rarely more than 6 mm in height and 3-6 mm in 

calicular diameter, the larger corallites typical of the northern 

(e.g., Monterey Bay) range of the species. Corallites at edge of 

colony invariably strongly inclined outward, their outer edges 

often flush with common coenosteum. Corallite base polycyclic, 

often consisting of 5 thecal rings (Durham, 1949). Costae 

well defined: broad (0.35-0.40 mm), slightly convex, and 

granular, separated by very thin intercostal striae. Corallum 

light brown, the color more intense at the calice. Polyps orange 

or coral-red, with white-tipped tentacles (Johnson and Snook, 

1967). 

Septal complement variable, depending on size and location 

of corallite within colony. Large, centrally-placed corallites 

have septa hexamerally arranged in four complete cycles (48 

septa) according to the formula: S1>S2.3»S4. However, most 

corallites lack pairs of S4 in several half-systems, and smaller 

peripherally located corallites usually have only 24-36 septa. 

S, slightly exsert and only about 1 mm wide, their vertical to 

slightly inclined inner edges divided into 4 or 5 narrow, 

granular lobes or teeth that merge imperceptibly with the 

columella. S2.3 only slightly less exsert and as wide as the Sv 

having similarly dentate inner edges. Each pair of S3 bend 

toward their adjacent S2, with a tendency to resemble the 

Pourtales Plan, as mentioned by Durham and Barnard (1952). 

S4 rudimentary, much smaller than S2.3, and have laciniate 

inner edges. All septal faces highly granular, the largest 

granules occurring on septal teeth. Fossa relatively deep and 

large. Columella consists of a flat to slightly concave field of 

fine (80-100 |im in diameter) papillae similar in size and 

shape to lower septal teeth. 

DISCUSSION.—It is not my goal to resolve the "Astrangia 

problem," as Squires (1959:416) phrased it, i.e., the validity of 

the 21 nominal Astrangia species described from the eastern 

Pacific. Fadlallah (1982) is probably correct in assuming that 

there is only one species of Astrangia north of Baja California 

and that it is A. lajollaensis (= A. haimei). The species problem 

more accurately pertains to those species occurring in the 

tropical eastern Pacific, particularly those described from the 

Gulf of California. However, a direct comparison of the 

lectotype of A. haimei to the holotype of A. lajollaensis shows 

no significant differences and thus A. lajollaensis is considered 

a junior synonym of A. haimei, as first suggested by Squires 

(1959). Nonetheless, the northern populations of A. haimei, 

previously referred to as A. lajollaensis, are in general, more 

robust than those to the south. 

In a detailed study of the reproduction of A. lajollaensis (= 

A. haimei), Fadlallah (1982) reported that it had separate sexes 

that broadcast large numbers of gametes in an annual cycle, as 

well as extratentacular budding of contiguous corallites to form 

colonies. Furthermore, he predicted that a colony 1 m 2 would 

be 46-383 years old, the range depending on differing 

assumptions for the average rate of budding per year, which he 

estimated to vary from 0.12-1.0 bud/corallite/year. 

MATERIAL EXAMINED.—New Records: Monterey wharf, 1 

colony, CAS 74819; Anacapal., 1 colony, CAS 74812; various 

Channel Islands, USNM 62489-92 and SIO Co 1167; Monterey 

breakwater, 1 colony, USNM 89300; Ensenada, 4 corallites, 

USNM 19198; off La Jolla, 10 colonies, SIO Co 1174, 1186; 

off San Onofre, CA, 4 colonies, SIO Co 980; off Coronado I., 

Gulf of California, 1 colony, SIO Co 1170. Previous 

Records: Lectotype of A. haimei; 1 colony from Zorritos, 

Peru, USNM 92451 (Verrill, 1870a); Bay of Panama, 1, USNM 

22087 (Marenzeller, 1904b); holotype of A. lajollaensis 

Durham, 1947. 

TYPES.—The lectotype of A. haimei, designated by Durham 

and Barnard (1952:71), is deposited at the YPM (598a) (Plate 

3d). About 15 paralectotypes are also deposited at the YPM,


one cataloged as 593 (Lazo-Wasem, pers. comm.). The 

lectotype colony measures 27.5 mm in diameter and consists of 

33-36 corallites, the typical calicular diameter being about 3.6 

mm. Type Locality: Bay of Panama; depth unknown. 

The holotype and paratypes of A. lajollaensis are deposited 

at the UCMP, cataloged as 30282, and 30308 and 30296, 

respectively. Type Locality: ?Ensenada, Baja California; 

depth unknown. 

DISTRIBUTION.—Known from off Peru (Verrill, 1870a) to 

Monterey Bay, California, including the Gulf of California; 

1-53 m. 

Family OCULINIDAE 

Oculina Lamarck, 1816 

DIAGNOSIS.—Corallum branches through extratentacular 

sympodial budding; axial corallites absent. Coenosteum dense 

and costate. Pali present before first two cycles of septa; 

columella papillose. Zooxanthellate and azooxanthellate species 

occur, primarily in shallow water. 

TYPE SPECIES.—Madrepora virginea Lamarck, 1816 (= 

Oculina diffusa Lamarck, 1816), by subsequent designation 

(Milne Edwards and Haime, 185Oa:xix). 

Oculina profunda Cairns, 1991 

PLATE 3d,e 

Oculina profunda Cairns. 1991a: 10-11, pi. 3: figs, e.g; pi. 4: figs. a,b 

[synonymy]. 

DIAGNOSIS.—Colonies arborescent and presumably attached. 

Corallites uniformly distributed on branches, each 

2.6-3.7 mm in diameter, and only slightly exsert. Conoesteum 

costate and white. Septa arranged in three complete cycles. P1 

crown composed of small pali; P2 crown much more 

prominent. Columella papillose. 

DISCUSSION.—Oculina profunda has been so recently 

described and illustrated (Cairns, 1991a) that it is unnecessary 

to recast the original description, especially as no additional 

specimens have been taken of this rarely collected species. 

MATERIAL EXAMINED.—Type series. 

TYPES.—The holotype (Plate 3d) and paratypes (Plate 3e) of 

O. profunda are all deposited at the USNM (see Caims 1991a, 

b). Type Locality: Alb-3170: 38° 1VN, 123°29'W (off Bodega 

Bay, California), 305 m. 

DISTRIBUTION.—Off St. Lucia Range and Bodega Bay, 

California; Galapagos; 119-742 m. ?Pleistocene of Mexico 

(Palmer, 1928). 

Madrepora Linnaeus, 1758 

DIAGNOSIS.—Colonial, extratentacular sympodial budding 

forming dendroid colonies. Coenosteum dense; costae absent. 

Pali absent; columella papillose or absent. Exclusively 

azooxanthellate. 

TYPE SPECIES.—Madrepora oculata Linnaeus, 1758, by 

subsequent designation (Verrill, 1901). 

Madrepora oculata Linnaeus, 1758 

PLATE 3/-/I 

Madrepora oculata Linnaeus, 1758:798.—Marenzeller, 1904b:79.—Durham 

and Barnard, 1952:11.—Eguchi, 1968:C29, pi. C8: figs 1-9.—Zibrowius, 

1974a:762-766, pi. 2: figs. 3-5 [synonymy].—Cairns, 1979:39-42, pi. 3: 

fig. 2; pi. 4: fig. 5; pi. 5: figs. 1-3 [synonymy]; 1982:15, pi. 3: figs. 4-6 

[synonymy]; 1984:10, pi. 1: fig. H; 1991a:9-10, pi. 2: fig. j; pi. 3: figs, 

a-d.—Zibrowius, 1980:36-40, pi. 13: figs. A-P [synonymy]. 

Madrepora galapagensis Vaughan, 1906b:63-64, pi. 1: fig. 2; pi. 2: figs. 1, 

lb.—Durham and Barnard, 1952:11.—Wells, 1983:234, pi. 13: figs. 1, 2. 

Madrepora (Sclerhelia) sp.—Eguchi, 1938, table 2. 

Madrepora (Amphelia) sp.—Yabe and Eguchi, 1941b:102. 

Madrepora cf. oculata.—Eguchi, 1942:136-137, pi. 6: fig. 1. 

DESCRIPTION OF NORTHEAST PACIFIC SPECIMENS.— 

Colonies uniplanar, formed by closely spaced extratentacular 

sympodial budding. Calices circular and 3.0-3.7 mm in 

diameter, exsert on end branches but flush or recessed into 

coenosteum on larger diameter branches. Coenosteum faintly 

striate and finely granular; light brown in color. 

Septa hexamerally arranged in 3 complete cycles (24 septa) 

according to the formula: S1_2»S3. S^ not exsert and have 

slightly sinuous inner edges that fuse with the columella. S3 

rudimentary. Fossa deep and relatively broad. Columella a 

bolus of trabeculae, interconnected to lower, inner edges of 

Si-2- 

DISCUSSION.—Madrepora oculata is a widespread and 

highly variable species that has been described under a dozen 

different names in regional accounts (see Zibrowius, 1974a, for 

a discussion of extended synonymy). Of the four forms of the 

species described by Cairns (1991a) from the Galapagos, the 

Fieberling Seamount specimens most clearly resemble forma 

galapagensis. The Japanese specimens differ from those in the 

eastern Pacific in having well-developed S3, almost as large as 

the S2. 

MATERIAL EXAMINED.—New Records: R/V Washington 

PPTU-II, 10 branches, USNM 83581; Alb-2978, 1 branch, 

USNM 92454; TM (KT7414, B2), 1 branch, ORI; off 

Enoshima, Sagami Bay, 1 branch, USNM 92658. Previous 

Records: Alb-3401, USNM 22085 (Marenzeller, 1904b); 

syntypes of M. galapagensis Vaughan, 1906b, USNM; 

specimens reported by Zibrowius (1974a, 1980) and Cairns 

(1979, 1982, 1984, 1991a). 

TYPES.—The types of M. oculata are lost (see Zibrowius, 

1980). Type Locality: Tyrrhenian Sea and Sicily, Mediterranean; 


Two syntypes of M. galapagensis are deposited at the 

USNM (68276). Type Locality: Alb-4642:l°30.5'S, 

89°35.0'W (south of Espanola, Galapagos), 549 m. 

DISTRIBUTION.—Madrepora oculata is cosmopolitan in 

distribution outside Antarctic Seas, including the Atlantic 

(Cairns, 1979; Zibrowius, 1980), Subantarctic (Cairns, 1982), 

Indian Ocean (Zibrowius, 1974a), New Zealand (Squires and

NUMBER 557 19 

Keyes, 1967), Hawaiian Islands (Cairns, 1984), Galapagos 

(Marenzeller, 1904b; Vaughan, 1906b; Cairns, 1991a), and off 

Japan (Eguchi, 1968) (off Honshu from the northwestern tip to 

Toyama Bay; Sagami and Suruga Bays; Korea Strait south of 

Tshushima) at depths of 15-1500 m. However, the two records 

reported herein from Anacapa Island (84 m) and Fieberling 

Seamount (440-488 m) are the first known from the northeast 

Pacific Basin. 

Suborder CARYOPHYLLIINA 

Superfamily CARYOPHYLLIOIDEA 

Family CARYOPHYLLIIDAE 

Caryophyllia Lamarck, 1816 

DIAGNOSIS.—Corallum solitary, attached or free: if attached, 

corallum cylindrical, trochoid, or ceratoid; if free, corallum 

usually cornute. Calice circular, elliptical, or compressed; 

thecal edge spines present on species having compressed 

coral la. Septal symmetry variable, but hexameral symmetry 

with four cycles of septa most common (Cairns, 1991a). One 

crown of pali present before penultimate or antipenultimate 

cycle of septa, which is usually the S3. Columella fascicular, 

composed of several twisted laths. Exclusively azooxanthellate 

and common in deep water. 

Subgenus Caryophyllia (Caryophyllia) Lamarck, 1816 

DIAGNOSIS.—Caryophyllia in which the calice is circular to 

elliptical (not compressed), and which do not have thecal edge 

spines or crests. 

TYPE SPECIES.—Madrepora cyathus Ellis and Solander, 

1786, by subsequent designation (Broderip, 1828). 

Caryophyllia (C.) arnoldi Vaughan, 1900 

PLATE 4a-e 

Caryophyllia arnoldi Vaughan, 1900:199-200, pi. 16: figs. 1,2; 1903:86. pi. 3: 

figs. 4, 4a.—Durham, 1947:33-34, pi. 2: figs. 3, 7, 16, 17.—Durham and 

Barnard, 1952:81-82, pi. 9: fig. 42a,b.—Keller, 1981:19, fig. 4, table 

4.—Austin, 1985:81.—Bythell. 1986:13-14.—Cairns etal.. 1991:47. 

Caryophyllia alaskensis: Durham, 1947:33 [in part: specimens from California, 

including pi. 2: figs. 4, 8, 9, 12, 13); 1949:152-153, pi. 4: fig. 9.—Durham 

and Barnard, 1952:81, pi. 9: fig. 41a,b—Talmadge, 1972:81 [in part: 

California specimens].—Keller, 1981:21 [in part: Vifyaz-4139, 4179].— 

Bythell, 1986:14 [in part: pi. 4: figs. C-F]. 

Caryophyllia ambrosia.—Keller, 1981a:15 [in part: V/0>ar-6127]. 

DESCRIPTION.—Corallum robust, ceratoid to trochoid, always 

attached through a relatively thick pedicel up to 65% of 

GCD. Calice circular to slightly elliptical; calicular edge 

serrate, each septum producing an acute triangular apex. 

Largest specimen known (Alb-4463) 16.0 x 14.5 mm in 

calicular diameter and 17.0 mm in height, with a pedicel 

diameter of 8.8 mm. Theca thick; costae usually have rounded 

edges, especially near calicular edge, and are separated by 

narrow and shallow intercostal striae. Costae usually bear very 

small, rounded granules: 4 or 5 occurring across the width of a 

costa. Occasionally the theca is smooth, lacking granulation. 

Corallum white. 

Above a calicular diameter of 8 mm septa are hexamerally 

arranged in 4 complete cycles (48 septa) according to the 

formula: S1_2>S3>S4. Su2 moderately exsert (1.4-2.3 mm) 

and have straight inner edges that extend about three-quarters 

distance to columella. S3 about three-quarters width of S^_2, 

0.9-1.3 mm exsert, and have slightly sinuous inner edges. S4 

80%-100% width of S3 (the wider S3 occurring in larger 

coralla), have straight inner edges, and are equally exsert as S3. 

Twelve broad, extremely sinuous P3 form a discrete circular to 

slightly elliptical palar crown. Pali bear tall granules, sometimes 

fused into oblique carinae. Viewed from above, the 

sinuous pali give the appearance of being thicker than the septa, 

but both septa and pali are equally thick. Fossa relatively 

shallow, containing a robust, fascicular columella composed of 

a circular to slightly elliptical field of 5-25 twisted laths, 

occasionally fused together into a massive structure. 

DISCUSSION.—Caryophyllia arnoldi and C. alaskensis are 

very similar, so much so that Durham (1947) identified some of 

his specimens of C. alaskensis as "pathologically" robust, 

herein reidentified as C. arnoldi. Durham and Barnard (1952) 

expressed doubt about their identification of C. arnoldi, and 

Bythell (1986:14) stated that the differences between the two 

species "appear to be slight and without further ecological 

evidence they cannot be designated as separate species with 

certainty." It is admitted that they are very similar and may 

even be sister species or even subspecies, but examination of 

the types of both species and a large suite of specimens from 

California to Alaska show several consistent morphological 

differences between the two as well as a geographic division. 

Caryophyllia arnoldi has a more robust and denser corallum, 

with thicker walls and a wider pedicel diameter. It has better 

defined costae that are usually granular, not porcellaneous. C. 

arnoldi consistently has 48 septa and 12 pali, whereas C. 

alaskensis occasionally has 14-16 pali and correspondingly 56 

or 64 septa. The septa of C. arnoldi are more exsert, their S^ 

usually well over 1 mm above the calicular edge, whereas those 

of C. alaskensis are invariably less than 1 mm exsert. The 

calicular apices of C. arnoldi are acute; those of C. alaskansis 

are usually equilateral. Finally, the S4 of C. arnoldi are almost 

as wide as their S3; the S4 of C. alskensis are only about 

three-quarters the width of their S3. Geographically, C. 

alaskensis is known only from the boreal temperate region, 

whereas C. arnoldi has a range from San Diego to the Gulf of 

Alaska, their ranges overlapping only off British Columbia and 

the Gulf of Alaska. Perhaps no single character will definitively 

separate the two species, but, when taken together, the 

characters listed above usually are diagnostic. 

Two other species of Caryophyllia are known from the 

Pleistocene of San Pedro, California, the type-locality of C. 

arnoldi: C. californica Vaughan, 1903. and C. pedroensis

20 

Vaughan, 1903. Caryophyllia californica is easily distinguished 

by having five complete cycles of septa at a GCD of 

only 10.7 mm (Plate 5d,e, USNM 92479). The holotype of C. 

pedroensis (Plate 5c,f, USNM Ml64736) resembles C. arnoldi 

in being free and having 48 septa, but its poor preservation does 

not allow a definite identification, none of its pali being intact. 

Durham (1947:33) suggested that C. pedroensis Vaughan, 

1903 might be the senior synonym of C. alaskensis Vaughan, 

1941, but it is more likely to be a junior synonym of C. arnoldi 

Vaughan, 1900. 

The holotype of C. arnoldi (USNM Ml57509) is unattached, 

whereas all subsequently collected Recent specimens are firmly 

attached and have a stout pedicel. This being the only apparent 

difference between the Pleistocene type and Recent species, I 

follow Durham's (1947) reasoning in using this name for the 

Recent species. 


USNM 19264; Alb-2864, 6, USNM 19211; Alb-2886, 6, 

USNM 92467; Alb-2893, 4, USNM 19283; AIb-2935, 1, 





USNM 92468; Alb-4328, 1, USNM M547401; Alb-4332, 1, 

USNM M547399; Alb-4359, 1, USNM 92470; Alb-4361, 2, 

USNM 77418; Alb-4377, 1, USNM M547402; Alb-4431, 2, 


USNM M547400; Lower Inlet, B.C., 2, USNM M547319 and 

M547321; Departure Bay, B. C, 4, USNM M547323; Work 

Canal, B. C, 1, USNM M547322; 60°22TSf, 147°42'W, 70-73 

m, 2, USNM 91421; Treadwell Mine, Alaska, Pleistocene, 2, 

USNM Ml23169; Snittishan, Juneau, Alaska, Pleistocene, 3, 

USNM 92466; Eshany Bay, P.W.S., 149 m, 7, UA; Ressurection 

Bay, Alaska, 58 m, 1, UA; 60°48TSf, 148° M'W, 1, UA; 

Point Valdez, Alaska, 1, UA; 45°01.5'N, 124°43.2'W, 273 m, 

13, UA; Channel Islands, 10, CAS 74811, 74803, 74909, 

74805, 74908, 80909, 80899; off Cordell Bank, 8, CAS 74831, 

74832; OCSEAP 20136, 2, CAS; OCSEAP 26502, 1, CAS; 

OCSEAP 20134, 1, CAS; OCSEAP 26499, 1, CAS; off 

Bodega, California, 1, CAS 80911; 32°46.5TSf, 118°2O.5 / W, 

505 m, 1, SIO Co 1248; 32°00'N, 117°56.2'W, 183 m, 1, SIO 

Co 404; 79 specimens from off British Columbia from 40-377 

m, all deposited at the RBCM and cataloged as: 974-389-5, 

974-224-6, 976-1030-1, 976-1051-1, 976-1033-2, 976-111-50, 

976-17-3, 980-257-40, 980-245-32, 980-268-16, 980-264-24, 

980-256-18, 981-202-3, 981-204-2, 981-208-1, 986-275-3, 

986-278-6, 988-2-12, 988-15-6, 988-2-13. Previous Records: 

Holotype of C. arnoldi (USNM); Albatross specimens 

reported by Durham, 1947 (USNM); specimen of Talmadge, 

1972 (CAS); Vityaz (stations 4139, 4179, 6127) specimens of 

Keller, 1981 (IOM). Reference Specimens: Holotypes of C. 

californica and C. pedroensis (both USNM). 

TYPES.—The holotype (Plate 4a,b) of C. arnoldi is deposited 

at the USNM (M164736). Type Locality: San Pedro, California 

(Pleistocene). 


DISTRIBUTION.—Known from San Diego to Prince William 

Sound, Gulf of Alaska, including Queen Charlotte Islands, 

Vancouver Island, San Juan Island, off British Columbia, and 

the Channel Islands and Banks; 40-656 m.-Pleistocene of San 

Pedro, California and Juneau, Alaska. 

Caryophyllia (C.) alaskensis Vaughan, 1941 

PLATE 4f-h.j 

Caryophyllia alaskensis Vaughan, 1941:280-281, pi. 40: figs. 1-4.—Durham, 

1947:33 [in part: only specimen from Alaska].—Not Durham, 1949:152- 

153 [= C. arnoldi].—Not Durham and Barnard, 1952:81 [= C. arnoldi].— 

Talmadge, 1972:81 [in part: Alaskan specimen].—?Ricketts and Calvin, 

1952:37.—Not Keller, 1981a:21 [= C. arnoldi and C. japonica].—Austin, 

1985:81 [in part: northern records].—Bythell, 1986:14 [in part: not pi. 4: 

figs. C-F, only northern latitude range].—Cairns et al., 1991:47. 

DESCRIPTION.—Corallum ceratoid, always attached through 

a relatively slender pedicel (18%-33% GCD). Calice circular 

to elliptical; calicular edge finely serrate, each septum forming 

a broad equilateral apex. Largest specimen examined (Alb- 

3317) 21.1 x 16.6 mm in diameter and 20.3 mm in height. 

Theca smooth, porcellaneous (not granular), and thin (translucent); 

milky white. Cu3 occasionally slightly ridged near 

calicular edge; otherwise, costae not expressed. Thin, chalky 

white intercostal striae delimit costae but there is no relief to 

theca. 

Septa often hexamerally arranged in four complete cycles 

(48 septa and 12 pali) according to formula: S1_2>S3>S4, but 

large coralla have 56 septa and 14 pali or 64 septa and 16 pali, 

the largest specimen having an almost complete fifth cycle (94 

septa). Instead of a fourth size class of septa to accommodate 

the fifth cycle, larger calices maintain only three size classes by 

adding equivalent half-systems of septa (i.e., 1 S2, 1 S3, and 2 

S4), rather than two pairs of S5. Su2 very slightly exsert (less 

than 1 mm) and have straight to slightly sinuous inner edges 

that merge with the columella deep within fossa. S3 only half 

width of S.|_2, having slightly sinuous inner edges and bordered 

internally by very sinuous, large pali, each 1-3 times width of 

an S3. P3 prominently granulated and extend into columella. S4 

about three-quarters width of S3 and have sinuous inner edges. 

Fossa moderate to shallow, containing a fascicular columella 

composed of 6-17 narrow twisted laths, which are usually 

independent of one another. 

DISCUSSION.—Comparisons to the closely related C. arnoldi 

and C. japonica are found in the accounts of those species. 



USNM 92460; Alb-4782, 1, USNM 92461; Alb-4788, I, 



USNM 92463; Alpha Helix-30, 1, UA; Lets Go-80, 1, UA; 2, 

RBCM 980-238-15. Previous Records: Syntypes of C. 

alaskensis; specimens reported by Durham, 1947 (USNM). 

Reference Specimens: Specimens reported as C. alaskensis 

by Keller (1981a) (IOM).

NUMBER 557 21 

TYPES.—The syntypes (Plate 4 f,h) of C. alaskensis are 

deposited at the USNM: two from Alb-4231 (M547317) and 

one from Alb-4300 (M547318). Type Locality: Behm Canal 

and Summer Strait, Alexander Archipelago, Alaska; 150- 

399 m. 

DISTRIBUTION.—Strait of Georgia, British Columbia to the 

Commander Islands, off Kamchatka (Keller, 1981a), including 

Alexander Archipelago, Gulf of Alaska, Aleutian Islands, and 

one record from the northern Korea Strait off South Korea; 

102-399 m. 

Caryophyllia (C.) sp. A 

PLATE 5a,b 

DESCRIPTION OF SPECIMEN FROM Alb-4784.—Corallum 

base and calice damaged: approximately 34 mm in GCD and 27 

mm in height. Corallum apparently cornute and thus probably 

free. Costae completely covered with an encrusting bryozoan. 

Corallum white. Septa hexamerally arranged in 5 complete 

cycles (96 septa) according to formula: S1_2»S3>S4>S5. S^ 

only moderately exsert (2.0-2.5 mm) and have slightly sinuous 

inner edges that border on the columella. S3 only about 

two-thirds width of S^, slightly less exsert, and also have 

slightly sinuous inner edges. S4_5 progressively smaller, each 

about three-quarters width of next lower cycle of septa, and less 

exsert, except for those S5 adjacent to S^_2, which are more 

exsert than the S4. A crown of small P3 occurs adjacent to the 

columella, well separated from their corresponding septa. Pairs 

of much broader P4 also present, forming a more prominent 

palar crown slightly recessed from the columella and arranged 

in chevrons with the P3 within each half-system. Fossa 6 mm 

deep, containing a massive fascicular columella composed of 

about 23 slender, laterally fused, twisted laths. 

DISCUSSION.—Among the 56 Recent species of Caryophyllia 

listed by Cairns (1991a), only seven have a full fifth 

cycle of septa, and of those seven, only two are cornute: C. 

grandis Gardiner and Waugh, 1938, and C. planilamellata 

Dennant, 1906. The North Pacific Caryophyllia differs from 

these two species in having pali before not only the penultimate 

septal cycle (P4) but also before the antipenultimate cycle (P3) 

and in having a larger, more open calice. Although this 

specimen may represent an undescribed species endemic to the 

northern boreal Pacific, the examination of one damaged 

specimen is not considered adequate to establish a new 

species. 

The specimens reported by Keller (1981a) as C. ambrosia 

from the Gulf of Alaska and the Kurile Islands are C. arnoldi 

and C. alaskensis, respectively. The Gulf of Alaska record 

(Vityaz-6121) may have been the basis for Austin's (1985) 

listing of C. ambrosia for off British Columbia. 

MATERIAL EXAMINED.—New Record: Alb-4784, 1, 

USNM 92478. Reference Specimens: Wryaz-5638 and 6127, 

C. ambrosia of Keller, 1981 (IOM). 

DISTRIBUTION.—Off Attu I., Aleutian Islands; 247 m. 

Labyrinthocyathus Cairns, 1979 

DIAGNOSIS.—Corallum solitary, ceratoid to trochoid, attached. 

Costae usually absent. Endotheca and pali absent; 

columella composed of an interconnected maze of lamellar 

plates. Exclusively azooxanthellate. 

TYPE SPECIES.—Labyrinthocyathus langae Cairns, 1979, by 

original designation. 

Labyrinthocyathus quay lei (Durham, 1947) 

PLATES 5g-i, 6aJ> 

Cyalhoceras quay lei Durham, 1947:32, pi. 1: figs. 1-4; 1949:153, text-fig. 

8-2.—Austin, 1985:81.—Kozloff, 1987:72.—Cairns et al., 1991:47. 

Labyrinthocyathus quaylei.—Cairns, 1982:22.—Bythell, 1986:15-16, pi. 2: 

fig. A; pi. 6: figs. CD; pi. 7: figs. A-E. 

DESCRIPTION.—Corallum ceratoid to cylindrical, often several 

attached to one another and thus resembling a small 

colony. Largest specimen examined (CAS 16405) 15.8 x 14.1 

mm in calicular diameter and 21.0 mm in height. Calice usually 

circular, occasionally elliptical. Pedicel relatively slender: 

PD:GCD = 0.35-0.50. Costae usually not discemable, the 

theca being uniformly covered with very small, rounded 

granules giving a smooth aspect to theca. Occasionally costae 

are visible as broad flat strips delineated by narrow and shallow 

intercostal striae. Corallum white. 

Septa hexamerally arranged in 5 cycles according to the 

formula: S1.2>S3»S4>S5. A full fourth cycle (48 septa) is 

attained at a GCD of 4-5 mm, and the fifth cycle (96 septa) at 

12-13 mm GCD, although the holotype (which is 14.0 x 12.7 

mm in calicular diameter) has only 90 septa. S^_2 on W slightly 

exsert (1.0-1.5 mm) and have sinuous inner edges that border 

the columella. S3 about 80% width of Su2 and also have 

sinuous edges. S4 much smaller, only 40%-50% width of S3, 

and have straight inner edges. S5 rudimentary, only about 

one-third size of S4. Fossa moderate in depth, containing a 

large columella consisting of short, interconnected lamellae, 

the lamellae sometimes vertically ridged. 

DISCUSSION.—Other than L. quaylei, seven species are 

recognized in Labyrinthocyathus: L. delicatus (Marenzeller, 

1904a) (southwest Indian Ocean); L. mentaldoensis (Chevalier, 

1961) (Miocene of Italy); L. taurinensis (Chevalier, 1961) 

(Miocene of Italy); L. limatulus (Squires, 1964) (New 

Zealand); L. kondoi (Wells, 1977) (Eocene of Tonga); L. 

facetus Cairns, 1979; and L. langae Cairns, 1979 (western 

Atlantic). Labyrinthocyathus quaylei, the only species known 

from the North Pacific, is distinguished from the others by 

having five, not four, cycles of septa. In the North Pacific, L. 

quaylei is most similar to Crispatotrochus foxi (Durham and 

Barnard, 1952), as discussed in the account of that species. 

Subsequent to its original description, L. quaylei was 

reported only once from Scripps Submarine Canyon, La Jolla 

by Bythell (1986), who illustrated corallum variation and 

several living specimens. Ten additional records are reported 

herein.

22 



USNM 92458; Alb-4551, 4, USNM M547418 (topotypic); 

Cordell Bank, 5, USNM 92455; Monterey Bay, 50-60 m, 10, 

CAS 16405, 16315, 16289; Zaca-42, 1, CAS 80932; off Santa 

Barbara, 64-80 m, 4, CAS 74905. Previous Records: 

Holotype and paratype of C. quaylei (UCMP and 

USNM); BythelFs (1986) specimens, 5, SIO Co 1176. 

TYPES.—The holotype (Plate 5h,i) of C. quaylei is deposited 

at the UCMP (30245). Two paratypes (Plate 5g) from the same 

station (Alb-4551) are deposited at the UCMP (30244) and the 

USNM (M547417). Type Locality: 36°45'N, 121°55'W 

(Point Pinos, Monterey Bay), 84-102 m. 

DISTRIBUTION.—Point Loma, San Diego to Cordell Bank, 

California, including the Channel Islands; 37-293 m. The 

record from Jarvis Inlet, B. C. (Austin 1985) has not been 

verified and is doubted. 

Crispatotrochus Tenison Woods, 1878 

DIAGNOSIS.—Corallum solitary, ceratoid to turbinate, and 

attached through a robust pedicel. Septotheca costate. Pali 

absent; columella fascicular, composed of twisted laths. 

Exclusively azooxanthellate. 

TYPE SPECIES.—Crispatotrochus inornatus Tenison Woods, 

1878, by monotypy. 

Crispatotrochus foxi (Durham and Barnard, 1952) 

PLATE 6C -e 

Cyathoceras foxi Durham and Barnard, 1952:84-85, pi. 10: fig. 46a,b.— 

Bythell, 1986:15, pi. 6: figs. A3.—Cairns et al., 1991:47. 

Crispatotrochus foxi.—Cairns, 1991a: 15. 

DESCRIPTION.—Corallum trochoid, firmly attached through 

a pedicel about half calicular diameter. Holotype 14.6 x 12.8 

mm in calicular diameter and 14.5 mm in height; Alaskan 

specimen (Alb-3324) slightly larger: 14.7 mm in calicular 

diameter and 18.5 mm in height, with a pedicel diameter of 6.6 

mm. Calice essentially circular but slightly irregular in outline 

due to small outpocketings of the theca. All costae (C^_5) 

slightly ridged, but only in upper one-third to one-half of 

corallum. Small, low granules cover costae. Corallum white. 


formula: S1.2>S3»S4>S5. Holotype lacks 1 pair of S5 (94 

septa), whereas Alaskan specimen has a full 5 cycles (96 septa). 

Su2 slightly exsert (1.3-2.5 mm) and have straight inner edges 

that attain the columella. S3 less exsert, about 80% width of 

Su2, and also have straight inner edges. S4 half or less width of 

S3; S5 rudimentary, only one-quarter to one-third width of S4. 

Septal faces sparsely granulated but upper faces of Su3 finely 

pleated, bearing very low, fine trabecular ridges oriented 

perpendicular to septal edges. Fossa of moderate depth, 

containing a relatively small fascicular columella consisting of 

4-7 broad, twisted laths that are laterally fused to one another. 


DISCUSSION.—Twelve species (Cairns, 1991a) are known in 

Crispatotrochus, the senior synonym of Cyathoceras, only 

three of which have five cycles of hexamerally arranged septa: 

C. rubescens (Moseley, 1881), C. niinoi (Yabe and Eguchi, 

1942b), and C.foxi (Durham and Barnard, 1952). C. rubescens, 

which is known from the central and western Pacific (Cairns, 

1984), differs from C. foxi in having a larger, more robust 

corallum with more exsert Su2 and sinuous inner septal edges. 

Crispatotrochus niinoi, known only from off Japan, is similar 

to C. foxi in size and septal formula but appears to differ in 

having a narrower pedicel (PD:GCD = 0.13 vs 0.41 -0.45 for C. 

foxi), a thicker theca, and a smooth, noncostate theca. 

In the northeastern Pacific, C. foxi could easily be confused 

with Labyrinthocyathus quaylei, both species being about the 

same size and shape, and having the same septal formula. 

Crispatotrochus foxi differs by having a fascicular columella 

(not interconnected plates), slightly ridged upper costae, 

straight inner septal edges, pleated septal faces, and, in general, 

a more delicate corallum. 

Crispatotrochus foxi is known from only three specimens: 

the holotype, a non-type juvenile also reported in the original 

description, and a third specimen reported herein, which is 

almost identical to the holotype. The species was originally 

named for the collector of the type specimen, Earl Fox, and thus 

it is coincidental that the specimen reported herein was 

collected from the Fox Islands, Alaska. 


USNM 19210. Previous Records: Holotype. 

TYPES.—The holotype (Plate 6c) is now deposited at the 

SBMNH (ex AHF 112). Type Locality: Richardson Point, 

San Miguel I., Channel Islands, California, 82 m. 

DISTRIBUTION.—Known only from the Channel Islands (San 

Miguel and Santa Catalina) and off Unalaska, Fox Islands, 

Aleutian Chain; 82-274 m. 

Paracyathus Milne Edwards and Haime, 1848a 

DIAGNOSIS.—Corallum solitary, turbinate, fixed or free. 

Septotheca costate. Paliform lobes usually bi- or trilobed, 

occurring before all but last cycle. Columella papillose, often 

indistinguishable from lower paliform lobes. 

TYPE SPECIES.—Paracyathus procumbens Milne Edwards 

and Haime, 1848a, by subsequent designation (Milne Edwards 

and Haime, 185Oa:xv). 

Paracyathus stearnsii Verrill, 1869 

PLATES 6f-k, la-f 

Paracyathus stearnsii Verrill, 1869:393-394; 1870a:537-538; 1870b:560 — 

Durham, 1947:35, pi. 2: figs. 1, 2, 5, 6; 1949:153-156.161, pi. 4: fig. 12; pi. 

5: fig. 1, text-figs. 8-1, 9, 17-9.—Durham and Barnard, 1952:92-93, pi. 13: 

fig. 55a-e.—?Hertlein and Grant, 1960:80-81, pi. 19: figs. 8-13.—Lewbel 

et al., 1981:165.—Fadlallah and Pearse, 1982b:233-238,6 figs.—Fadlallah, 

1983a: 132.—Bythell, 1986:17, pi. 3: fig. C; pi. 9: figs. A-F — Kozloff, 

1987:72.—Cairns et al., 1991:47.—Chadwick, 1991:42-47. 

Paracyathus caltha Verrill, 1869:394; 1870a:537, pi. 9: figs. 9, 9a; 1870b:

NUMBER 557 23 

560.—Whiteaves, 1886:115.—Hickson, 1917:24.—Durham, 1947:34.— 

Austin, 1985:81. 

Paracyathus tiburonensis Durham, 1947:35-36, pi. 3: figs. 5, 6 [new 

synonym]; 1949:156.—Durham and Barnard, 1952:94, pi. 13: fig. 56.— 

Squires, 1959:423. 

Paracyathus stearnsi.—Austin, 1985:81.—Kozloff, 1987:72. 

Paracyathus calthus.—Cairns et al., 1991:47. 

DESCRIPTION.—Corallum trochoid, firmly attached through 

a robust pedicel 40%-65% that of GCD, and firmly anchored 

to gastropod or bivalve shells or small pebbles by a thin 

expansive base up to twice calicular diameter in size. Young 

coralla (Plate la-c) demonstrate a polycyclic basal development, 

older coralla having up to 6 concentric basal thecal rings 

(Durham, 1949). Largest corallum examined (USNM 92603) 

26.5 x 17.6 mm in calicular diameter and 18.3 mm in height. 

Calice elliptical; smaller coralla occasionally circular. Costae 

equal in width, low, and rounded, separated by relatively 

narrow and shallow intercostal striae about one-quarter width 

of costae. Costae covered with low, rounded granules, 2-3 

occurring across the width of a costa. In some coralla, costae 

are slightly ridged near the calice, in which case the granules 

are less apparent and the intercostal striae are wider. Upper 

quarter to third of well-preserved coralla, including all calicular 

elements, pigmented a light to dark brown. 

Septa hexamerally arranged in 5 cycles accordingly: Sl- 

2>S3>S4>S5. A full fourth cycle is obtained at a GCD as small 

as 3.5 mm; the fifth cycle is usually complete at a GCD of 

12-14 mm; and additional pairs of S6 occur in larger coralla, up 

to 146 septa in the largest specimen examined (USNM 92603). 

All septa exsert, but only moderately so, their inner edges 

straight and vertical (not sinuous). S^ easily distinguished 

from others by their much larger size, each bordered internally 

by an undivided, rounded paliform lobe that sits low in the 

fossa directly adjacent to the columella. P3 located higher in 

fossa, slightly recessed from columella, and often dissected into 

2 or 3 lobes, the lowermost indistinguishable from columellar 

elements. P4 located highest in fossa, most recessed from 

columella, and also frequently dissected into several lobes. If 

not fragmented, P4 are the broadest paliform lobes, and, each 

pair, together with the enclosed P3, form a chevron in each 

half-system. Septal faces bear prominent, pointed granules; 

palar faces often bear short carinae and/or granules. Paliform 

lobes thick as septa. 

Fossa of variable depth but usually deep, containing multiple 

paliform crowns and a large, elliptical columella. Columella 

composed of a field of 2-40 small cylindrical, irregularlyshaped 

pillars, all pillars terminating at the same height several 

mm below the P^.2- 

DISCUSSION.—Examination of the types of P. caltha and P. 

tiburonensis indicate that they fall well within the range of 

variation for P. stearnsii, the former synonymy having been 

implied by Durham (1947:34). The distinguishing characters of 

P. tiburonensis listed by Durham (1947) (i.e., little-exsert, thin 

septa; straight septal edges; equal costae) are all typical of P. 

stearnsii. Durham and Barnard (1952) also suggested that the 

Pleistocene P. pedroensis Vaughan, 1903 was a junior 

synonym of P. stearnsii, but examination of the holotype (Plate 

8/i) revealed that although it is very similar, its septa are thicker 

and less crowded—only 66 septa present in a corallum with a 

GCD of 11.4 mm. A specimen of P. stearnsii of corresponding 

size would have 90-96 septa and well-developed pairs of P4. 

For this reason I consider P. pedroensis to be a distinct species. 

Hertlein and Grant's (1960) Pliocene records of P. stearnsii 

rely on the synonymy of P. pedroensis and are therefore 

suspect. Comparisons of P. stearnsii to P. montereyensis are 

made in the account of the latter species. Paracyathus stearnsii 

is also remarkably similar to the amphi-Atlantic P. pulchellus 

(Philippi, 1842), but the Atlantic species does not attain the 

large size or full fifth cycle of closely spaced septa characteristic 

of P. stearnsii. 

Being a common, relatively shallow-water species, P. 

stearnsii is one of the most frequently collected corals in the 

Northeast Pacific. Variations in hydrographic conditions 

associated with near-shore habitats probably contribute to the 

morphologic variation seen in the corallum. The most variable 

character of this species is the size and nature of the paliform 

lobes, particularly whether each lobe is intact or divided into 

several smaller lobes. Fossa depth is also variable, usually 

moderately deep but occasionally quite shallow, some specimens 

having paliform lobes rising even above the calicular 

edge. Nonetheless, the species is characterized by having 

crowded septa with a tendency toward five full cyles of septa, 

having 12 S^_2 that evenly divide the calice into 12 

half-systems, and a characteristic P3-P4 chevron arrangement 

in each half-system. 

In a study of the reproduction of P. stearnsii, Fadlallah and 

Pearse (1982a) found the species to have separate sexes in a 1:1 

ratio and to broadcast its gametes in a synchronized cycle. 

External fertilization results in very small (about 160 (im) 

planktonic planulae, which suffer a high rate of mortality. 

Densities of P. stearnsii were found to average 24.5 coralla/m 2 

and longevity was estimated to be about 40 years. 





USNM 19203 and 19239; Alb-2913, 11, USNM 19230 and 

19281; Alb-2922, 20, USNM 19213 and 19225; Alb-2932, 1, 

USNM 92495; Alb-2939, 10, USNM 19240 and 19254; 

Alb-2940, 1, USNM 19231; Alb-2943, 13, USNM 19219; 

Alb-2944, 5, USNM 92596; Alb-2945, 5, USNM 19204 and 

19265; Alb-2958,5, USNM 19218; Alb-2961,4, USNM 19234 

and 19253; Alb-2965, 8, USNM 19201; Alb-2968, 5, USNM 

19238; Alb-2969, 10, USNM 19242; Alb-2974, 5, USNM 


19243; Alb-2977, 5, USNM 19214 and 36447; Alb-2978, 4, 


USNM 19263; Alb-3088, 7, USNM 19216; Alb-3102, 23,

24 




USNM 92602; approximately 230 specimens from the Channel 

Islands deposited at the CAS and cataloged as: 74827, 74911, 

74800, 74826, 74824, 74825, 74804, 74801, 69608; 45 

specimens from Monterey Bay: CAS 74813,74824,74822; off 

Pacific Grove, 11, CAS 74821; Cordell Bank, 22, CAS 74830, 

74835; off Huntington Beach, 2, CAS 74906; south of Carmel, 

2, CAS 80904; 25 specimens from Channel Islands deposited at 

SIO, cataloged as: Co 441, 1167, 1169; southwest of Cedros I., 

27 m, 1, SIO Co 1172; off La Jolla, 13, SIO Co 1164, 1186; 

north side of Middle I., Coronado I., 15 m, SIO Co 1170; 

27°0lX 114°16.5'W, 99-108 m, 1, SIO Co 1273; 72 

specimens from off B. C. deposited at RBCM, cataloged as: 

973-234-4, 973-235-5, 973-246-2, 973-231-2, 976-1043-3, 

976-1078-2, 976-1163-60, 976-1077-3, 976-1037-1, 977-156- 

14, 978-96-2, 980-329-25, 980-343-18, 984-417-2, 985-469- 

73. Previous Records: Holotypes of P. stearnsii, P. caltha, 

and P. tiburonensis; Skidgate, B. C, 25-90 m, 1, USNM 78624 

(Durham, 1947). Reference Specimens: Holotype of P. 

pedroensis, USNM Ml64738. 

TYPES.—The holotype (Plate Id-f) of P. stearnsii is 

deposited at the YPM (2187). It measures 17.9 x 12.9 mm in 

calicular diameter and has 104 septa. Type Locality: 

Monterey Bay, California, depth unknown. 

The holotype (Plate 6/) of P. caltha is also deposited at the 

YPM (965). It measures 11.5 x 8.2 mm in calicular diameter 

and has 96 septa. Type Locality: Monterey Bay, California, 


The holotype (Plate 6h) of P. tiburonensis is deposited at the 

UCMP (30485). It measures 9.3 x 8.0 mm in calicular diameter 

and has 82 septa. Type Locality: Off Tiburon I., Gulf of 

California, 73 m. 

DISTRIBUTION.—From Skidgate, Queen Charlotte Islands 

(Durham, 1947) to Bahfa Asuncion (27°01'N), Baja California 

and Tiburon Island, Gulf of California (as P. tiburonensis); 

20-134 m. 

Paracyathus montereyensis Durham, 1947 

PLATE lg-i 

Paracyathus montereyensis Durham, 1947:34-35. pi. 2: figs. 10, 19.—Durham 

and Barnard, 1952:11.—Austin, 1985.—Bythell, 1986:18.—Cairns et al., 

1991:47. 

DESCRIPTION.—Holotype 6.03 mm in circular calicular 

diameter and 8.4 mm in height, attached by a thick pedicel 4.6 

mm in diameter that expands into a broad, thin base. Costae 

coarsely granular and light brown in color. Septa decamerally 

arranged in three cycles plus 3 pairs of quaternaries: 10:10:20:6 

= 46 septa. Primary septa about 1 mm exsert and 1.6 mm wide, 

with straight to slightly sinuous, vertical inner edges. Inner 

edges of primaries separated by a narrow notch from slender 

pali, each about 0.4 mm wide. Secondary septa less exsert and 


about two-thirds width of primaries, each bordered internally 

by a robust palus about 0.6 mm wide and greatly thickened on 

its edge adjacent to the notch separating it from its corresponding 

septum. In addition to being wider, these pali reach higher 

in the fossa than the PI and are slightly recessed from the 

columella, the 20 paliform lobes together forming two crowns 

distinguished by size, height, thickness, and position. Three 

sectors bear pairs of rudimentary quaternary septa. Within the 

crowns of pali is a papillose columella composed of 12 slender 

elements. 

The larger specimen (Alb-2888, USNM 92604, Plate lg,h) 

from off Oregon measures 11.1 x 9.7 mm in calicular diameter 

and 12.8 mm im height and also is brownish in color. Its septa 

are decamerally arranged in 3 cycles, with 1 additional pair of 

quaternaries, for a total of only 42 septa. 

DISCUSSION.—This rarely collected species was previously 

known from only the three syntypes; one additional specimen 

is reported herein. Paracyathus montereyensis differs from the 

much more commonly collected P. stearnsii primarily in its 

septal symmetry and its relatively fewer septa at a corresponding 

calicular diameter. The septal symmetry of P. montereyensis 

appears to be decameral, attaining and maintaining 40-46 

septa at a GCD of 6 mm and above. The symmetry of P. 

stearnsii is hexameral, a corallum 6 mm in calicular diameter 

already having about 60 septa and a specimen 11 mm in 

calicular diameter having a full fifth cycle (96 septa). 

Associated with the lower number of septa, P. montereyensis 

has only two palar crowns, not three or more as in P. stearnsii, 

and its septa are thicker, yet less crowded than those of P. 

stearnsii. 

The Albatross specimen from off Oregon is quite similar to 

the type of P. pedroensis Vaughan, 1903, known from the 

Pleistocene of San Pedro, California, except that the symmetry 

of P. pedroensis is hexameral, the holotype measuring 11.3 x 

10.0 mm in calicular diameter and having 66 septa (Plate %h). 


USNM 92604. Previous Records: Holotype of P. montereyensis. 

Reference Specimen: Holotype (Plate 8/i) of P. pedroensis 

Vaughan, 1903 (USNM Ml64738). 

TYPES.—The holotype (Plate 7i) and two paratypes are 

deposited at the UCMP (30341 and 30342-43, respectively). 

Type Locality: Off Point Lopez, 17 mile drive, Monterey 

Peninsula, California; 146 m. 

DISTRIBUTION.—Known only from off Monterey, California 

and off Oregon Dunes, Oregon; 75-146 m. 

Coenocyathus Milne Edwards and Haime, 1848a 

DIAGNOSIS.—Like Caryophyllia (Caryophyllia) but forming 

colonies through extratentacular budding from a thick basal 

coenosteum and occasionally from the theca of other corallites. 

Exclusively azooxanthellate, occurring in relatively shallow 

water. 

TYPE SPECIES.—Coenocyathus cylindricus Milne Edwards

NUMBER 557 25 

and Haime, 1848a, by subsequent designation (Milne Edwards 

and Haime, 1850a:xii). 

Coenocyathus bowersi Vaughan, 1906 

PLATE 8a-/ 

Coenocyathus bowersi Vaughan, 1906a:847, pi. 77: figs. 1-3.—Durham, 

1947:34.—Durham and Barnard, 1952:83-84, pi. 10: fig. 45a-d— Wells, 

1956:F422, fig. 323,8.—Squires, 1959:422.—Parker, 1964:150.— 

Zibrowius, 1980:73.—Austin, 1985:81.—Bythell, 1986:14-15, pi. 3: fig. A; 

pi. 5: figs. A-F, color cover illustration.—Cairns et al., 1991:47. 

DESCRIPTION.—Corallum colonial, the most common habit 

being a thick mounded coenosteal encrustation from which 

cylindrical corallites arise by extratentacular budding, each 

coral lite 5-8 mm in height. However, some colonies have 

widely spaced and taller (up to 20 mm) corallites (Plate %b) and 

others have closely spaced (cerioid) and very short corallites 

(Plate 8a). One corallum (Plate 8c) even adopted a branching 

mode, some of its corallites budding from the theca of parent 

corallites. Holotypic colony only 4 cm in width; however, 

Durham (1947) reported a colony 1 foot (30.5 cm) in diameter, 

and a specimen at the CAS (77797) measures 37 cm across. 

Colonies usually encrust bivalves or rocks. Calices usually 

circular, but may be elliptical or somewhat irregular in shape. 

Calice size quite variable, some colonies (e.g., the holotype) 

having corallites 3.5-7.5 mm in calicular diameter, others 

averaging up to 13-14 mm in diameter. In addition to the 

typical method of extratentacular budding, examples of 

intratentacular budding are found in most coralla, in which a 

corallite is in the process of or has split into two or as many as 

10 smaller corallites (Plate Sd,e), a process termed multiple 

fission by Durham and Barnard (1952). Costae broad and flat to 

slightly convex, equal in width, and separated by very narrow, 

slender striae. Costae sometimes extend over coenosteal base. 


Septal symmetry extremely irregular. Each calice has 6-14 

primary septa, which are larger than all others and extend to the 

columella or center of calice, dividing the corallite into 6-14 

sectors. Each sector encloses 3, 5, or 7 septa. The 3-septasectors 

consist of 1 secondary and 2 smaller tertiary septa plus 

1 P2. The 5-septa-sector consists of 1 secondary, 2 tertiary, and 

a pair of small quaternary septa, plus 1 P2. The 7-septa-sector 

consists of 1 secondary, 2 tertiary, and two pairs of quaternary 

septa, accompanied by 2 P3. Some calices have a typical 

hexameral symmetry of 12 primary, 12 secondary, and 24 

tertiary septa with 12 P2 (48 septa, Plate 8/), but most calices 

have a mixture of developmental stages, producing a very 

irregular septal insertion pattern, but no corallite examined had 

over 56 septa. All septa little exsert and have slightly sinuous 

inner edges. Pali (P2 or P3) relatively slender and extremely 

sinuous. In some calices they form a well-defined crown (Plate 

8/) as in Caryophyllia, but in others pali are poorly formed or 

even missing in various sectors. 

Fossa of moderate depth, containing a small fascicular 

columella composed of 1-4 twisted laths. Occasionally the 

columella is absent, the inner edges of the primary septa being 

slightly expanded and almost meeting in center of calice. 

DISCUSSION.—Five Recent species of Coenocyathus are 

known: C. cylindricus Milne Edwards and Haime, 1848a 

(northeastern Atlantic); C. anthophyllites Milne Edwards and 

Haime, 1848a (northeastern Atlantic); C. bowersi Vaughan, 

1906a; C. sagamiensis Eguchi, 1968 (Japan); and C. goreaui 

Wells, 1972 (Bermuda). Although Zibrowius (1980) expressed 

some reservation about placing C. bowersi in this genus, 

comparison to the type species does appear to justify its 

placement. The only other known Pacific species, C. sagamiensis, 

differs in having a papillose columella, hexamerally 

arranged septa, and in apparently lacking pali. 

Both Durham and Barnard (1952) and Bythell (1986) 

commented on the extreme variability of this species, with 

which I agree. Characters showing variation include: colony 

shape; corallite diameter and height; method of budding; septal 

insertion pattern; palar development; and columellar development. 

Nonetheless, C. bowersi is a distinctive species known 

from a circumscribed area, not easily confused with any other 

species in the northeastern Pacific. According the Bythell 

(1986), the peculiar intratentacular budding may be caused by 

the mechanical stress of sediment burial. 

MATERIAL EXAMINED.—New Records: Alb-2895,6 colonies, 




USNM 92607; off southern California, 20-45 m, 1, USNM 

78634; off Santa Cruz, 2, USNM 78631 and 78643; off Santa 

Catalina, 1, USNM 92608; Monterey Bay, 1, USNM 78613; 

Pillsbury-5\2, USNM 92609; east of Point Conception, 64-80 

m, 2, CAS 74904; Zaca-23, 2, CAS 80913; Zaca-42, 2, CAS 

80916; Zaca-43, 1, CAS 80921; off Bird Rock, Santa Catalina, 

13-15 m, 3, CAS 77797, 77799 and SIO Co 1168; off Catalina 

I., 1, CAS 80903; off Palos Verde, California, 13-15 m, CAS 

80939; Agassiz MV71-I-1, 1, SIO Co 1309. Previous Records: 

Holotype of C. bowersi; Alb-4463, 1, USNM 

M547379 (Durham, 1947); off La Jolla, 30 fms (= 55 m), 1, 

USNM 92920 (Durham, 1947). 

TYPES.—The holotype colony (Plate Sb) of C. bowersi is 

deposited at the USNM (21138). Type Locality: Off San 

Miguel Island, Channel Islands, California; depth unknown. 

DISTRIBUTION.—Colonet, Pacific coast of Baja California 

(Parker, 1964) to Monterey Bay, including Channel Islands, 

Cortes Bank, and Isla Guadelupe; Gulf of California; Gulf of 

Panama (reported herein); 9-302 m. 

Nomlandia Durham and Barnard, 1952 

DIAGNOSIS.—Corallum solitary, discoidal, and attached. 

Thecal wall lacking. Pali present (?); columella fascicular. No 

dissepiments or synapticulae.

26 

TYPE SPECIES.—Nomlandia californica Durham and Barnard, 

1952, by original designation. 

Nomlandia californica Durham and Barnard, 1952 

PLATE %g 

Nomlandia californica Durham and Barnard, 1952:91, pi. 12: fig. 53.—Wells, 

1956:F423.—Bythell, 1986, 19.—Cairns et al., 1991:47. 

REDESCRIPTION OF HOLOTYPE.—Coral lum discoidal and 

firmly attached (encrusting): 9.7 x 7.0 mm in calicular diameter 

and only 2.4 mm in height. Thecal wall and costae absent; outer 

edges of septa attenuate in height toward perimeter of basal 

plate. Septa hexamerally arranged in 4 complete cycles (48 

septa) according to formula: S1>S2>S3>S4. S1 extend to 

columella, the higher cycle septa progressively less wide. All 

septa semi-circular in shape (arched), the S, being the highest 

of the septa. Septal edges finely pleated corresponding to 

underlying trabeculae that are oriented perpendicular to septal 

edges. Closer to basal plate and toward columella the septa bear 

tall, elongate carinae oriented parallel to septal edges. A 

sinuous palus (P3) occurs in only one half-system, being absent 

from all other half-systems. Columella a single low, twisted 

lath, which is attached to the inner edges of the 6 Sv 

DISCUSSION.—This unusual species is known from only the 

holotype, collected from a sunken bouy off San Miguel Island, 

California. It does not resemble any North Pacific or any other 

previously described coral, justifying Durham and Barnard's 

creation of a new genus. Durham and Barnard (1952) suggested 

that Nomlandia was related to Bathycyathus, but its inconsistent 

presence of pali and lack of a thecal wall argue against that 

relation. Wells (1956) questioningly placed the genus in the 

Caryophylliinae. The holotype appears to be a mature 

specimen, not a juvenile stage of a larger, better-known species, 

but, until more specimens are collected, its range of variation 

and phylogenetic affinities remain enigmatic. Its encrustation 

of a sunken bouy suggests that it might have been transported 

a great distance, either alive or after death. 

MATERIAL EXAMINED.—New Records: None. Previous 

Records: Holotype. 

TYPE.—The holotype (Plate 8g) of N. californica is 

deposited at the SBMNH (35560) ex AHF 19. Type Locality: 

1.1 km off Richardson Point, San Miguel Island, Channel 

Islands, California; 82 m on a sunken bouy. 

DISTRIBUTION.—Known only from the type locality. 

DesmophyUum Ehrenberg, 1834 

DIAGNOSIS.—Solitary, trochoid, fixed. Septothecate. Pali 

absent; columella absent or quite rudimentary. Sparse endothecal 

dissepiments. Azooxanthellate. 

TYPE SPECIES.—Madrepora dianthus Esper, 1794, here 

designated. Milne Edwards and Haime (185Oa:xvii) have been 

cited as the authors to have subsequently designated DesmophyUum 

cristagalli Milne Edwards and Haime, 1848a, as the 


type species of DesmophyUum, however this cannot be valid 

since D. cristagalli was not included as a species in the original 

description of the genus. In Ehrenberg's (1834:299) original 

description of DesmophyUum, he listed two species: "D. 

dianthus (= Madrepora dianthus Esper)" and D. stellaria sp. 

nov. Since the type species of a genus must originate from the 

species originally placed in that genus when it was established 

(ICZN Articles 67g and 69a), D. cristagalli cannot be 

considered as the type species, even if it is considered to be a 

junior synonym of the type species. In 1857, Milne Edwards 

and Haime made a distinction between the Red Sea specimens 

Ehrenberg called D. dianthus (having five cycles of septa) and 

the "East Indian" specimen (having six cycles of septa) that 

Esper called D. dianthus. This undoubtedly led Wells 

(1956:F426) to designate Ehrenberg's "D. dianthus (non 

Madrepora dianthus Esper)" as the type species of the genus. 

But, regardless of the specimens Ehrenberg had in hand at the 

Muse"e de Berlin (see Zibrowius, 1980:117), he clearly equated 

his D. dianthus to Esper's species, and thus the type of the 

genus cannot be D. dianthus sensu Ehrenberg, as implied by 

Wells (1956), but must be the original D. dianthus (Esper, 

1794). According to Zibrowius (1980:117), Ehrenberg's D. 

stellaria is a junior synonym of Balanophyllia europaea (Risso, 

1826). It is therefore appropriate (ICZN recommendation 

69B(3): choice by elimination) to designate the other species 

listed by Ehrenberg (1834) in his original generic account, M. 

dianthus Esper, 1794, as the type species of the genus. 

Esper's type of D. dianthus is lost (Scheer, 1990:406), and 

his description and illustrations leave room for doubt as to its 

identity, but it is known (Cairns, 1979, 1982) that D. cristagalli 

is a widespread and quite variable species, having five, six, or 

even more cycles of septa. It is quite likely that Esper's 

six-cycle D. dianthus from the "East Indies" was the same as 

Ehrenberg's five-cycle D. dianthus from the Red Sea, a species 

better known as D. cristagalli Milne Edwards and Haime, 

1848a. Even though the type of Esper's D. dianthus is lost, this 

name has nomenclatural priority as well as being the type 

species of the genus, and thus a neotype is chosen for this 

species: a specimen having six cycles of septa from Sagami 

Bay (Plate 9a,/?). 

DesmophyUum dianthus (Esper, 1794) 

PLATE 9a-d 

Madrepora dianthus Esper, 1794, pi. 69: figs. 1-3; 1795:85-86.—Scheer, 

1990:406. 

DesmophyUum dianthus.—Ehrenberg, 1834:299-300.—Milne Edwards and 

Haime, 1848a:254-255; 1857:77-78.—Yabe and Eguchi, 1942b:l 13-114, 

pi. 9: figs. 1-3.—Eguchi, 1965:290, 2 figs.; 1968:C41, pi. C33: fig. 6. 

DesmophyUum cristagalli Milne Edwards and Haime, 1848a:253, pi. 7: figs. 

10, 10a.—Marenzeller, 1904b:81.—Durham, 1947:36-37, pi. I: figs. 6, 10, 

15, 17; 1949:158-159, pi. 4: figs. 2, 4, 7, 8.—Durham and Barnard, 1952: 

86-87, pi. 11: fig. 48 [not Cartego Bay specimen].—Parker, 1964:150.— 

Talmadge, 1972:81, 2 figs.—Zibrowius, 1974a:758-761, pi. 3: figs. 1-10 

[synonymy]; 1980:117-121, pi. 61: figs. A-O; pi. 62: figs. A-M.—Cairns, 

1979:117-119, pi. 21: figs. 7, 8; pi. 22: fig. 8; 1982:29-30, pi. 8: figs. 9-12;

NUMBER 557 27 

pi. 9: figs. 1-3 [synonymy]; 199la: 17, pi. 6: figs, g-i.—Austin, 1985:81.— 

Bythell, 1986:16-17, pi. 8: figs. A-D.—Kozloff, 1987:72. 

Desmophyllum cumingii Milne Edwards and Haime, 1848a:254, pi. 7: fig. 11. 

DESCRIPTION.—Corallum ceratoid, often flaring at calice 

(trumpet-shaped), attached through a robust pedicel 20%-40% 

that of GCD. Largest North Pacific specimen (USNM 83583) 

60 x 40 mm in calicular diameter and 50 mm in height, with a 

pedicel diameter of 20 mm. Calice circular, elliptical, or 

scalloped. Theca uniformly covered with small low granules; 

costae rarely expressed, but occasionally C:_3 are present in 

upper half of corallum as thin ridges. Corallum light brown or 

grey. 

Septa hexamerally arranged in 5 to 6 cycles according to the 

formula: S1_2>S3»S4>S5>S6. Fourth cycle (48 septa) attained 

at a calicular diameter of about 7 mm and fifth cycle at a 

calicular diameter of about 18 mm; a complete sixth cycle (192 

septa) is often present in Japanese specimens. S:_2 extremely 

robust, up to 2 mm thick at thecal edge, and up to 11 mm exsert. 

S^_2 have straight, vertical inner edges that define a deep, 

narrow fossa, the inner edges of opposing SU2 sometimes 

almost touching in center of fossa. S3 also highly exsert, but 

narrower—only 80%-90% width of S^_2. S4 much smaller 

than S3 (50%-70% width) and least exsert of the septal cycles. 

S5 only about half width of S4, but highly exsert, rising well 

above S4 and often becoming incorporated into adjacent S^_3 

in large coralla. Septal faces smooth, covered with small, fine, 

rounded granules. Fossa deep and slender. Columella usually 

absent but may consist of up to 5 slender fascicular or papillose 

elements, usually hidden from view in an intact corallum. 

DISCUSSION.—Specimens from the northeastern Pacific tend 

to have five cycles of thick septa, occasionally with pairs of S6. 

Japanese specimens tend to have a full sixth cycle, typifying the 

typical form originally described by Esper and the form 

identified as D. dianthus by Yabe and Eguchi (1942b). 



USNM 92477; Alb-2987, 4, USNM 19202 and 19250; 


Alb-4912, 3, USNM M547422; R/V Washington, 32°25.78'N, 

127°47.4'W (Fieberling Seamount), 440-488 m, 4, USNM 

83583; Cobb Seamount, 312 m, 2, USNM 78630; off Santa 

Barbara, 64-80 m, 2, CAS 74903; Eel Canyon, 366 m, 5, CAS 

74910; Carmel Bay, 183 m, 1, CAS 80926; Monterey Bay, 

91-110 m, 2, CAS 15647 and 16309; La Jolla Canyon, 33 m, 

1, SIO Co 1265; 32°46'N, 117°22.8'W, 183 m, 8, SIO Co 1193; 

32°46.5'N, 1 lS^O.S'W, 505 m, 2, SIO Co 1276; east of South 

Point, Guadelupe, 274 m, 3, SIO Co 1333; off Point Loma, 229 

m, 10, SIO Co 945; TM (KT7802,Z61), 1, IOM; Enoshima, 

Sagami Bay, 9, USNM 92612; 32°21'N, 128°41'E, 179-201 m, 

2, ZMC; 32°21'N, 128°39'E, 274-366 m, 5, ZMC. Previous 

Records: Neotype of D. dianthus; holotype of D. cristagalli; 

specimens reported by Marenzeller (1904b) from Alb-3384 and 

3401 (USNM); specimens reported by Durham (1947) from 

Alb-4370,4373, 4376,4377 (USNM). 

TYPES.—Although we may never know the identity of 

Esper's D. dianthus from the "East Indies" because the type is 

lost and the description is brief, it nonetheless must be 

considered as the type species of the genus (see above). 

Because it is the type species of Desmophyllum, it is 

appropriate to designate a neotype (Plate 9a,b) for the species, 

herein deignated as a specimen from Sagami Bay, depth 

unknown (USNM 92475). 

The holotype of D. cristagalli is deposited at the MNHNP 

and illustrated by Cairns (1979, pi. 21: figs. 7, 8). Type 

Locality: Gulf of Gascogne, depth unknown. 

The type of D. cumingii has not been traced. Type 

Locality: "Pacific coast of South America," depth unknown. 

DISTRIBUTION.—Northeast Pacific: Vancouver Island, British 

Columbia (Austin, 1985); Cobb Seamont, Washington to 

off San Diego, including Channel Isalnds, Cordell Bank, and 

Fieberling Seamount; Isla de Guadelupe, Mexico; Gulf of 

California (Parker, 1964); Gulf of Panama (Marenzeller, 

1904b); Cocos and Galapagos Islands (Cairns, 1991a); 33- 

1097 m. Off Japan: Sagami Bay to Kii Strait, Honshu; off 

Shikoku; off Koshiki Island, southwest Kyushu; 77-715 m. 

Elsewhere: Cosmopolitan except for off continental Antarctica 

and northern boreal Pacific; 35-2460 m (Cairns, 1982). 

Common on seamounts, guyots, and deep-water coral banks 

associated with framework building species (Cairns and 

Stanley, 1982). 

Lophelia Milne Edwards and Haime, 1849 

DIAGNOSIS.—Colonial, forming large dendroid colonies by 

intratentacular budding. Coenosteum dense; costae poorly 

developed. Pali absent; columella rudimentary. Sparse tabular 

endothecal dissepiments present. Azooxanthellate. 

TYPE SPECIES.—Madrepora prolifera Pallas, 1766 (= L. 

pertusa L., 1758), by subsequent designation (Milne Edwards 

and Haime, 185Oa:xx). 

Lophelia pertusa (Linnaeus, 1758) 

PLATE 9e-i 

Madrepora pertusa Linnaeus, 1758:797. 

Madrepora prolifera Pallas, 1766:307. 

Lophelia californica Durham, 1947:36, pi. 1: figs. 13, 16; pi. 2: fig. 

11.—Bythell. 1986:18, pi. 10: figs. A-E — Kozloff, 1987:72.—Cairns, 

1991a:17. 

Dendrosmilia nomlandi Durham and Barnard, 1952:85, pi. 10: fig. 47.— 

Caims, 1979:126.—Bythell, 1986:16, pi. 10: fig. F; 1991a:18. 

Lophelia prolifera.—Cairns, 1979:125-127, pi. 24: figs. 1-5 [synonymy]; 

1982:30-31, pi. 9: fig. 6; 1991a:17-18, pi. 6: fig. j. 

Lophelia pertusa.—Zibrowius, 1980:126-130, pi. 66: figs. A-L [synonymy]. 

Description of northeastern Pacific Specimens.-Colonies up 

to 25 x 12 cm in size (Bythell, 1986), achieved by profuse 

intratentacular budding. Up to 8 corallites may bud from the 

perimeter of a single corallite (Plate 9/), which ultimately 

produces a very dense, sometimes anastomotic, colony.

28 

Corallites ceratoid, often slightly curved, long, and slender: up 

to 25 mm long and usually 4-6 mm in diameter, but some large 

corallites up to 11 mm in diameter. Calices circular to slightly 

elliptical. Costae usually not present, the theca covered with 

low, closely spaced, rounded granules. Occasionally C^_2 

expressed as low ridges near calice. Corallum white. 

Septa irregularly arranged in 3 or 4 size classes. Six to 12 

primary septa define an equal number of sectors, each sector 

bisected by a smaller secondary septum. Another cycle of 

ternaries follows and a variable number of quaternaries, 32-66 

septa per calice being the most common totals for those 

examined. Primary septa fairly highly exsert (up to 1.2 mm), 

quite thin, and relatively narrow, their straight, vertical inner 

edges reaching halfway to calice center. Secondary septa about 

half width of primaries and correspondingly less exsert. 

Tertiaries quite small, but slightly enlarged if flanked by a pair 

of quaternaries. Lower, inner edges of all septa slightly sinuous. 

Septal faces smooth, covered by very small, low, rounded 

granules. Tabular endothecal dissepiments present low in fossa. 

Fossa deep and often curved, such that base cannot be seen. 

Columella usually absent, but occasionally expressed as a small 

crispate lath. 

DISCUSSION.—As discussed by Cairns (1979, 1991a), both 

L. californica and Dendrosmilia nomlandi are considered to be 

junior synonyms of L. prolifera (= L. pertusa). Lophelia 

californica represents the gracilis variation (sensu Duncan, 

1873), characterized by delicate, slender, elongate corallites. 

Also typical of this form is extremely profuse budding. D. 

nomlandi represents the more robust brachycephala variation 

(sensu Moseley, 1881), characterized by short, stout corallites 

and, in this case, a rudimentary columella. 

MATERIAL EXAMINED.—New Records: Alb-2946, 10 

colonies, USNM 92606; Alb-2948, 1 branch, USNM 19215; 

Alb-2984, 4 colonies, USNM 19237; Alb-2987, 10 colonies, 

USNM 19247; R/V Washington, 32°55'N, 127°47'W (Fieberling 

Seamount), 440-488 m, 1 branch, USNM 83582; Cobb 

Seamount, 415 m, branches, USNM 78616; Cobb Seamount, 

312 m, 2 branches, USNM 78605; Saucer Dive 345, off Bird 

Rock, La Jolla, 110-200 m, many fragments, SIO Co 532; TM 

(KT7414, B2), 1 branch, USNM 92611, 1 branch, ORI. 

Previous Records: Holotype of Dendrosmilia nomlandi. 

TYPES.—The Linnaean type of M. pertusa appears to be lost 

(Zibrowius, 1980). Type Locality: Not stated, but probably 

the fjords of Norway. 

The holotype of L. californica, stated to be deposited at the 

Los Angeles County Museum (2001), is no longer there and is 

presumed to be lost (G. Hendler, pers. comm.). Type 

Locality: Off "Southern California," depth unknown. 

The holotype of Dendrosmilia nomlandi (Plate 9h,i) is 

deposited at the SBMNH (35559) ex. AHF 15. Type Locality: 

VeleroA 172-40: 8.8 km southeast of Santa Catalina, 82- 

274 m. 

DISTRIBUTION.—Northeastern Pacific: known primarily 

from offshore seamounts and islands, including: Cobb Seamount, 

Washington; Channel Islands; off La Jolla (Bird Rock); 


Fieberling Seamount; and Isla de Guadelupe, off Baja 

California, Mexico; 82-488 m. Northwestern Pacific: Suruga 

Bay, Honshu; 150-340 m (first record for western Pacific). 

Elsewhere: Amphi-Atlantic (Cairns, 1979); northeast of 

Madagascar (Cairns and Keller, 1993) and off South Africa; 

Saint Paul and Amsterdam Islands (Zibrowius, 1974a); and 

Macquarie Ridge (Cairns, 1982); 60-2170 m. Lophelia pertusa 

is a framework species of deep-water coral banks (Cairns and 

Stanley, 1982), often providing the substrate for attachment by 

Desmophyllum dianthus. 

Superfamily FLABELLOIDAE 

Family FLA BELLI DAK 

Flabellum Lesson, 1831 

DIAGNOSIS.—Corallum solitary, fixed or free. Corallum 

ceratoid, campanulate, or compressed; base not reinforced with 

stereome. Wall epithecal, usually lacking costae. Transverse 

division lacking. Pali, dissepiments, and synapticulae absent. 

Columella rudimentary: a simple fusion of lower inner edges of 

larger septa. Exclusively azooxanthellate. 

Subgenus Flabellum (Flabellum) Lesson, 1831 

DIAGNOSIS.—Flabellum with a smooth (not serrate) calicular 

edge. 

TYPE SPECIES.—Flabellum pavoninum Lesson, 1831, by 

monotypy. 

Flabellum (F.) sp. A 

PLATE \0a,b 

DESCRIPTION.—Corallum ceratoid, largest specimen examined 

(Plate \0a,b) 25.8 x 21.5 mm in calicular diameter and 

38.5 mm in height, with a pedicel diameter of 4.1 mm and a 

basal plate diameter of 5.3 mm. Three of the 7 specimens 

reported herein attached to epitheca of conspecific specimens. 

Thin, shallow, longitudinal intercostal striae visible on worn 

specimens, otherwise, well-preserved coralla display transverse 

epithecal bands encircling the theca. Calice slightly elliptical 

(GCD:LCD = about 1.2); calicular edge very slightly scalloped, 

the outer edges of the S^ rising about 0.8 mm as apices. 

Septa hexamerally arranged in 5 complete cycles according 

to the formula: S1_2>S3»S4_5. A juvenile corallum only 4.8 

mm in calicular diameter has only 2 cycles of septa (12), 

whereas at a calicular diameter of 5.2 mm a full third cycle (24) 

is present and at a calicular diameter of 11.6 mm a full fourth 

plus 3 pairs of S5 are present (54 septa). The illustrated 

specimen (GCD = 25.8 mm) has a perfectly symmetrical 

arrangement of 5 cycles (96 septa). All septa nonexsert and 

quite thin, their faces covered with small, pointed granules. 

Su2 have straight inner edges, which fuse to one another deep 

in fossa. S3 one-half to two-thirds width of Su2, their inner 

edges reaching deep into fossa but not joining to Su2 fusion. 

S4 one-tenth to one-third width of S3; S5 rudimentary, less than

NUMBER 557 29 

1 mm wide. S4 extend deep into fossa but S5 extend only 

one-third distance down inner thecal wall. 

DISCUSSION.—None of the 28 Recent species in the 

nominate subgenus of Flabellum (see Cairns, 1989a:46) occur 

in the eastern Pacific or boreal North Pacific; however, the 

Alaskan specimens are most similar to several Antarctic- 

Subantarctic species that have conical (not flabellate) coralla, 

especially F. impensum Squires, 1962. Comparison of the 

Alaskan specimens to ceratoid specimens of F. impensum of 

equivalent size show them to be similar in shape, pedicel 

diameter, theca, septal shape, and septal formula. The only 

difference noted between the two was the slightly scalloped 

calicular margin of the Alaskan specimens, the calice of F. 

impensum being perfectly smooth. Flabellum impensum is 

known only from the Antarctic and Antipodes Islands at a 

depth range of 46-2260 m (Cairns, 1982). 

Several specimens of an unnamed species belonging to this 

subgenus (F. (F.) sp. A) were reported from the Galapagos 

Islands at 441-717 m (Cairns, 1991a), but are easily 

distinguished from the Alaskan specimens by their much 

smaller size, smaller pedicel diameter, and fewer septa. 

MATERIAL EXAMINED.—Alb-3315, 4, USNM 19226; Station 

69RD3, 3, NMCIC 1982-1492. Reference Specimens: 

Numerous specimens of F. impensum reported by 

Cairns (1982). 

DISTRIBUTION.—Aleutian Islands: Unalaska Bay, Fox Islands 

and off Amchitka, Rat Islands; 55-507 m. 

Javania Duncan, 1876 

DIAGNOSIS.—Corallum solitary, ceratoid to trochoid, and 

firmly attached by a stereome-reinforced pedicel and base; 

transverse division lacking. Theca smooth. Septa usually 

highly exsert; calicular edge scalloped. Fossa deep; pali 

lacking, columella absent or rudimentary. Azooxanthellate. 

TYPE SPECIES.—Javania insignis Duncan, 1876, by monotypy. 

Javania cailleti (Duchassaing and Michelotti, 1864) 

PLATE \0g-i 

Desmophyllum cailleti Duchassaing and Michelotti, 1864:66, pi. 8: fig. 11. 

Desmophyllum eburneum Moseley, 1881:162, pi. 6: figs. 1, la,b. 

Desmophyllum nobile Verrill, 1885:150-151. 

Desmophyllum vitreum Alcock, 1898:20, pi. 2: figs. 2, 2a,b. 

Desmophyllum galapagense Vaughan, 19O6b:63, pi. 1: figs. 1-lb.—Durham 

and Barnard, 1952:11.—Durham. 1962:46; 1966:125. 

Desmophyllum delicatum Yabe and Eguchi, 1942b: 115, 144, pi. 9: fig. 7a,b. 

Javania delicata.—Zibrowius, 1974b: 18. 

Javania cailleti—Cairns, 1979:153-156, pi. 28: figs. 8-12; pi. 30: figs. 1, 4 

[synonymy and discussion]).—Zibrowius. 1980:157-159, pi. 82: figs. A-L 

[synonymy and discussion].—Cairns, 1982:46-48, pi. 14: figs. 9-12; 

1991a:21, pi. 8: figs, c-e.—Not Wells, 1983:238. 

DESCRIPTION.—Corallum of large specimens ceratoid, 

firmly attached, and distally flared (trumpet-shaped). One 

figured specimen (Plate lOg) is 31 x 23 mm in calicular 

diameter (estimated) and 29 mm in height, with a thick pedicel 

7.5 mm in diameter. Pedicel diameter of a damaged specimen 

from British Columbia 13.8 mm. Theca white, porcellaneous, 

and smooth, except near calice where the theca overlaying the 

S3 is flared outwards. 


according to the formula: S1_2>S3>S4. S1_2 usually highly 

exsert (up to 4.5 mm in the British Columbia specimens) and 

have straight inner edges that fuse lower in fossa as a solid, 

rudimentary columella. S3 less exsert (about 2 mm above 

calicular edge), about three-quarters width of S^g, and extend 

deep into fossa but do not merge with columella. S4 nonexsert, 

about one-third width of S3, and extend only about halfway 

down inner thecal wall. 

DISCUSSION.—Javania cailleti is distinguished from all but 

one of the other nine species known in this genus (Cairns, 

1989a) by having four, not three or five, cycles of septa. /. 

pseudoalabastra Zibrowius, 1974b, known only from several 

records in the Atlantic (Zibrowius, 1980), also has four cycles 

of septa, but is easily distinguished by its elongate calice and 

red-brown coenosteum. At least six names have been applied to 

this species, which is not unusual for widely distributed species 

that are collected and described from the various ocean basins. 

Javania cailleti has been collected only rarely in the North 

Pacific. It is known from only three specimens from two 

localities in the Queen Charlotte Islands in the northeastern 

Pacific, and likewise from only three specimens from two 

localities off Japan (two of these specimens reported as 

Desmophyllum delicatum by Yabe and Eguchi, 1942b). 

MATERIAL EXAMINED.—-New Records: LM49,2, NMCIC 

1900-8362; 67-41 GBR, 1, NMCIC 1982-1491; Alb-4982, 1, 

USNM 92747. Previous Records: Types of D. eburneum, D. 

nobile, and D. galapagense. 

TYPES.—The holotype of D. cailleti appears to be lost 

(Cairns 1979). Type Locality: Lesser Antilles, depth unknown. 

Five syntypes of D. eburneum are deposited at the BM 

(1880. 11.25.65). Type Locality: Challenger-3O6A: 48°27'S, 

74^0^ (off southern Chile), 627 m. 

The holotype of D. nobile is deposited at the USNM (7964). 

Type Locality: 44°28'N, 57° 13*W (off Nova Scotia), 549 m. 

Most of the syntypes of D. vitreum are deposited at the 

Indian Museum, Calcutta, but five syntypes are also deposited 

at the MNHNP, two at the ZMA (Coel. 1198), and two at the 

MCZ. Type Locality: Investigator-232: 7°17'3O"N, 

76°54'30"E (Laccadive Sea), 787 m. 

The holotype of D. galapagense is deposited at the USNM 

(68275). Type Locality: Alb-4642: 1°3O.5'S, 89°35'W 

(Galapagos), 549 m. 

The holotype of D. delicatum is deposited at the TIUS 

(59131). Type Locality: Soyo Maru-22: 36°46'N, 141°30'E, 

off Hitachi, Honshu), 539 m. 

DISTRIBUTION.—Widespread (perhaps cosmopolitan): previously 

reported from throughout Atlantic Ocean from Nova

30 

Scotia to Burdwood Bank (Cairns, 1979; Zibrowius, 1980); 

southern Chile (Moseley, 1881; Cairns, 1982); Galapagos 

(Cairns, 1991a); off Japan (off Hitachi, Honshu and Sea of 

Japan, off southwestern Hokkaido); Arabian Sea (Alcock, 

1898); 86-2165 m. The two records reported from off British 

Columbia are the first for the northeastern Pacific. 

Javania borealis, sp. nov. 

PLATE 10cd 

?Javania cailleti — Wells, 1983:238 [in part: Alb-2816]. 

Javania n. sp.—Cairns, 1989a:76. 

?Javania sp. A.—Cairns, 1991a:21, pi. 8: figs. g,h. 

DESCRIPTION.—Corallum massive and trochoid, the thecal 

walls growing upward at a constant angle (not flared). Holotype 

36.4 x 27.8 mm in calicular diameter and 34.4 mm in height, 

firmly attached through a robust stereome-reinforced pedicel 

10.6 mm in diameter. A broken cross section of a pedicel of a 

paratype revealed 11 concentric layers of solid stereome. Calice 

elliptical; calicular edge slightly serrate, each S1_4 rising to a 

small triangular apex around calicular perimeter. Costae 

glisteny white. Theca and septa thin and fragile, easily broken, 

both about 0.2 mm thick. 


according to the formula: S1_2>S3>S4»S5. All septa have 

slightly sinuous inner edges and faces sparsely covered with 

low, pointed granules. Upper, inner edges of opposing St_2 

almost meet in center of fossa, and fuse into a rudimentary 

columella deeper in fossa. S3 about 80% width of Su2 and 

nonexsert, their lower inner edges also fused with the Su2 even 

deeper within fossa. S4 about half width of S3; S5 rudimentary, 

only about one-fourth width of S4. Fossa elongate and quite 

deep, defined by inner edges of S^. 

DISCUSSION.—Of the seven Recent species of Javania (see 

Cairns, 1989a:76), four have five cycles of septa: J. insignis 

Duncan, 1876; J. lamprotichum (Moseley, 1880); J. antarcticum 

(Gravier, 1914); and/, borealis. Javania borealis is quite 

similar to J. lamprotichum, which is known only from the 

Hawaiian Islands and off Johnston Atoll at 244-322 m (Cairns, 

1984). The Hawaiian species differs in having a flared calice, a 

reddish brown corallum, and more sinuous septal edges. 

Javania borealis differs from J. insignis by having: much less 

exsert septa; thinner, and thus more delicate, theca and septa; 

and more widely spaced septa. Each septum of J. borealis is 

separated from its adjacent septa by 0.5-0.6 mm, whereas in J. 

insignis the distance is 0.3 mm or less, their Su2 often so thick 

that adjacent S5 become fused to them near thecal edge. An 

unidentified species of Javania collected from off the Galapagos 

Islands at 143 m was also compared to /. borealis by Cairns 

(1991a), but the Galapagan specimen was not adequately 

preserved to make a definite identification. 

ETYMOLOGY.—The species name borealis (Latin borealis, 

northern) refers to the high northern latitude distribution of this 

species. 


MATERIAL EXAMINED/TYPES.—Holotype: Alb-4784, 

USNM 82019 (Plate \0c,d). Paratypes: Alb-4784, 3, USNM 

92618; ALB-4994, 1, USNM 92619. Type Locality: 

52°55'40"N, 173°26'E (northeast of Attu Island, Near Islands, 

Aleutian Chain, Alaska), 247 m. Reference Specimen: Javania 

sp. A of Caims (1991a), Alb-2816, USNM 19142. 

DISTRIBUTION.—Known only from off Attu Island (western 

Aleutian Chain) and off Rebun Island (Sea of Japan off 

northwest tip of Hokkaido); 247-348 m. 

Javania californica, sp. nov. 

PLATE \0e,f 

Flabellum montereyense Durham, 1947:37 (in part: paratype and 3 nontype 

specimens from Alb-4551; specimen from Alb-4550; 1 specimen from 

Alb-4553). 

DESCRIPTION.—Corallum trochoid and relatively small, 

attached by a thick, stereome-reinforced pedicel. Largest 

specimen (holotype) only 12.6 x 10.7 mm in calicular diameter 

and 12.6 mm in height, with a pedicel diameter of 3.6 mm. 

Calice elliptical; calicular edge highly serrate. Theca white, 

porcellaneous, and relatively thin in upper corallum. 

Septa hexamerally arranged in 3 complete cycles (only 24 

septa) according to the formula: S1>S2»S3. Each S1 forms a 

triangular apex about 1.2 mm high at calicular edge and has a 

straight inner edge that thickens lower in fossa. S2 also form 

triangular calicular apices of equal size and are about 80% 

width of Sv their lower, inner edges also thickened and fused 

with those of the S1 into a rudimentary columella. S3 much 

smaller, only about 20%-25% width of S2, have slightly 

sinuous inner edges, and form smaller calicular apices of about 

0.5 mm height. Septa thin (about 0.2 mm thick) and widely 

spaced, separated by approximately 1.1-1.2 mm from one 

another. Fossa deep and elongate. 

DISCUSSION.—Javania californica is distinguished from 

other Recent species in the genus by having only three cycles of 

widely spaced septa. Only one other species in the genus, J. 

duncani Wells, 1977 (Eocene of Tonga), has three septal 

cycles, but its generic placement is questionable and furthermore, 

the Eocene species has a much more slender corallum 

and thicker septa. 

Durham (1947) included several specimens of 7. californica 

in his description of Polymyces montereyensis. Although 

similar in young stages, and sometimes found together, /. 

californica is distinguished by having a solid, stereomereinforced 

pedicel (no rootlets); only three cycles of septa (S4 

are present in P. montereyensis even in the protothecal stage at 

6 mm calicular diameter); septa more exsert; a more robust 

columella; and a smaller adult size. 

Etymology.—The species name alludes to the region of 

collection of the type material. 

MATERIAL EXAMINED/TYPES.—Holotype: Alb-4550, 

USNM 92613 (ex. USNM M547411), Plate 10e,/. Paratypes: 

Alb-3168, 1, USNM 92615; Alb-4550, 4, USNM

NUMBER 557 31 

M547411, reported as F. montereyensis by Durham (1947); 

Alb-4551,4, USNM M547407, M547410, reported as paratype 

and nontypes by Durham (1947); Alb-4543, 1, USNM 92614 

(ex. USNM M547409) reported as P. montereyensis by 

Durham (1947). Reference Specimen: Holotype of J. duncani 

Wells, 1977 (USNM 1208332). Type Locality of J. californica: 

Alb-4550: 36°45'N, 121°55'W (off Point Pinos, 

Monterey Bay, California), 91-104 m. 

DISTRIBUTION.—Known only from Monterey Bay and 

Cordell Bank; 62-170 m. 

Polymyces Cairns, 1979 

DIAGNOSIS.—Corallum solitary, ceratoid to trochoid, fixed. 

Wall epithecal, reinforced basally by symmetrically or asymmetrically 

arranged, contiguous hollow rootlets. Calicular edge 

lacerate to serrate. Pali absent; columella rudimentary. 

TYPE SPECIES.—Rhizotrochus fragilis Pourtales, 1871, by 


Polymyces montereyensis (Durham, 1947) 

PLATE lla-/ 

Flabellum (?) montereyense Durham, 1947:37, pi. 1: figs. 5, 9 [in part: not 

paratype from Alb-4551; only 1 of 2 specimens from Alb-4543; only 1 of 3 

nontypes from Alb-4551; not Alb-4550: all Javania californica].—Durham 

and Barnard, 1952:97, pi. 14: fig. 59a-c—Austin, 1985:81. 

Flabellum tannerense Durham and Barnard, 1952:97-98, pi. 14: fig. 60a-c 

[new synonym].—Bythell, 1986:19. 

Polymyces montereyensis.—Cairns, 1979:158; 1991a:22.—Cairns et al., 

1991:48. 

Polymyces montereyense.—Bythell, 1986:19-20. 

Polymyces tannerensis.—Cairns, 1991a:22.—Cairns etal., 1991:48. 

DESCRIPTION.—Corallum trochoid, the thecal walls increasing 

in diameter at a constant angle. Largest known specimen 

(SEPBOP 18B-764, Plate 11/) 39.1 x 30.7 mm in calicular 

diameter and 34.4 mm in height, with a pedicel diameter of 

10.8 mm. Calice elliptical and only slightly serrate, each S1-4 

forming a small triangular apex. Corallum pedicel enlarged by 

a ring of contiguous rootlets, characteristic of the genus (see 

Discussion for more detail). Corallum white. 

Septa hexamerally arranged in five incomplete cycles 

according to the formula: S1_2>S3>S4»S5. At a GCD of 11 

mm 36-48 septa are present; the largest two coralla examined 

have only 90 or 92 septa, each lacking several pairs of S5. All 

septa have slightly sinuous inner edges and sparsely granular 

septal faces. S^_2 slightly exsert: the upper, inner edges of 

opposing lateral septa almost meeting in center of fossa, their 

lower inner edges fusing into a rudimentary columella deep in 

fossa. S3 about 80% width of Su2, nonexsert, and do not fuse 

with columella. S4 50%-80% width of S3; S5 rudimentary, 

about one-quarter width of S4 and extending only 4-5 mm 

down inner theca. Fossa deep and elongate, defined by inner 

edges of S^_2. 

DISCUSSION.—Polymyces montereyensis is easily distin- 

guished from the two other species in the genus (Cairns, 1991a) 

by having five cycles of septa, the other species having only 

four cycles. It is more difficult to distinguish it from the genus 

Javania, especially J. borealis, the basic difference between the 

two species being that J. borealis has a solid, stereomereinforced 

pedicel, whereas that of P. montereyensis is 

reinforced by hollow rootlets and secondarily by stereome, 

which gives it a less dense corallum. Because rootlets are hard 

to detect in adult coralla, a cross section of the pedicel 

sometimes must be made to verify the generic placement. 

Polymyces montereyensis also differs from J. borealis in 

having less wide S3 that do not fuse with the S^_2. The broad 

pedicel of P. montereyensis led Durham (1947) to correctly 

doubt its placement in Flabellum; the inclusion of specimens of 

Javania californica in the type series of F. montereyense added 

to that uncertainty. 

Polymyces tannerensis is herein synonymized with P. 

montereyensis. It was previously thought that it could be 

distinguished from P. montereyensis by its lack of a fifth cycle 

and its highly exsert S^; however, small specimens of P. 

montereyensis of the same size as the types of P. tannerensis 

(holotype: 11.2 x 8.9 mm in calicular diameter; paratype: 9.9 x 

8.6 mm in calicular diameter) have only four septal cycles. The 

highly exsert S^ of P. tannerense may be explained by the 

poor preservation of the two types, their epitheca entirely worn 

away at the calicular edge between major septa. In all other 

characters, P. tannerensis resembles P. montereyensis of an 

equivalent size. Furthermore, both "species" were collected at 

the same station (Velero-1348-41) and no additional specimens 

of the P. tannerensis growth form have been collected 

subsequently. 

The rootlets appear to form in the following manner. Each 

corallum forms a prototheca about 5-6 mm in height and 

diameter. A secondary epitheca develops from the basal plate 

and encircles the prototheca at a distance of 1.0-1.5 mm, the 

secondary epitheca eventually growing upward to form the 

adult wall. In the basalmost 1-2 mm of the pedicel, the ring 

formed by the encirclement of the prototheca by the epitheca is 

divided into 12 compartments (rootlets) by outward extentions 

of the S3 (Figure 3A; Plate 1 lc). Slightly higher in the pedicel 

(4-6 mm above basal plate) outward extensions of the S^ 

further subdivide the ring into 24 compartments (rootlets). 

These 24 chambers communicate with the polyp through a 

series of 12 pairs of pores, each pair flanking an S3 about 6-8 

mm above calicular base, essentially at the top of the 

protothecal ring (Figure 3B). Rudimentary S4 are also found 

within the prototheca, these septa being continuous with the S4 

of the adult corallum, only higher in the corallum. S5 are not 

found within the prototheca, these septa formed later and thus 

higher in the corallum. The space between each S4 and adjacent 

S1 and S2 within the prototheca does not result in an additional 

pore, but is sealed with a dissepiment-like structure. Sometimes 

rootlet development is irregular, several pairs missing from one 

side of a corallum. Rootlets can usually be distinguished

32 

2 prototheca 

secondary 

eDitheca 

FIGURE 3.—Cross-section diagram of two stages of rootlet development in 

Polymyces montereyensis. A (top). Early stage of rootlet development 

characterized by relatively wide outer ring (= 12 rootlets) and only three septal 

cycles (see Plate lie). B (bottom). Later development characterized by a 

narrower outer ring (24 rootlets) and four septal cycles. Location of pores 

through which rootlets communicate to polyp indicated by ellipses. 

externally as slightly convex, longitudinal strips near the 

coral lum base that often alter the architecture of the epitheca; 

however, in larger coralla the pedicel is secondarily reinforced 


with peripheral stereome and thus rootlets are seen only in cross 

section of the pedicel. Rootlets can also be seen in a calicular 

view of juvenile coralla, in which the paired pores leading to 

the rootlets flanking each S3 are visible within the corallum. 



USNM 83528; SEPBOP 18B-764, 1, USNM 83521; Zaca-42, 

1, CAS 80931. Previous Records: Holotype, paratypes, and 

nontypes of/ 7 , montereyense Durham, 1947 [Alb-4543 (USNM 

M547409), 4550 (= /. californica, USNM M547411), 4551 (ex 

USNM M547410, now USNM 92617)]; holotype of F. 

tannerense. 

TYPES.—The holotype (Plate 1 \a,b) of F. montereyense is 

deposited at the USNM (M547406). Two paratypes also exist: 

one deposited at the USNM (Alb-4551, USNM M547407, = J. 

californica) and one at the CAS (loc. 21810, CAS 7852). Type 

Locality: Alb-4543: 36°45'N, 121 "SS'W (west of Point Pinos 

Lighthouse, Monterey Bay), 97-170 m. 

The holotype (Plate 1 \d,e) and paratype of F. tannerense are 

both deposited at the SBMNH, the holotype numbered 35562 

(ex AHF 22). Type Locality: Velero 1348-41: Tanner Bank, 

off Southern California, 82-84 m; however, Velero 1348-41 is 

given as 33° 16X 118°15'W (off southeastern Santa Catalina) 

at 212 m in the station log. 

DISTRIBUTION.—Channel Islands, including Cortes and 

Tanner Banks, to Monterey Bay; off Cape Blanco, Peru; 

69-212 m. 

Suborder DENDROPHYLLIINA 

Family DENDROPHYLLIIDAE 

Balanophyllia Wood, 1844 

DIAGNOSIS.—Corallum solitary, turbinate to trochoid, fixed 

or free. Costae usually well developed. Synapticulotheca 

especially well developed near calice. Septa arranged in a 

Pourtales Plan. Pali may or may not be present; columella 

invariably spongy. Azooxanthellate; variable in depth range. 

TYPE SPECIES.—Balanophyllia calyculus Wood 1844, by 

monotypy. 

Balanophyllia elegans Verrill, 1864 

PLATE 12a-/ 

Balanophyllia elegans Verrill, 1864:44; 1870a:511-512, pi. 10: fig. 3; 

1870b:560.—Whiteaves, 1886:115.—Hickson, 1917:24.—Durham, 

1947:41, pi. 1: figs. 7, 8, 11, 12; pi. 10: figs. 3, 4, text-fig. 2A; 1949: 

142-145, 161, text-figs. 3, 17-7, pi. 5: figs. 2, 3, 6, 7, 10, 12, 15.—Durham 

and Barnard, 1952:99-100, pi. 14: fig. 62a-c.—Ricketts and Calvin, 

1952:37, pi. 6: fig. 5.—Emerson, 1956:394.—Hertlein and Emerson, 

1960:84, pi. 19: figs. 24, 19-21, pi. 24: figs. 4, 5, 8, 13, 15-17.—Guberlet, 

1962:53, figure.—Addicott, 1966, pi. 1: figs. 25, 26.—Johnson and Snook, 

1967:109, pi. 3: fig. 87.—Talmadge, 1972:81, fig.—Hand, 1975:93.— 

Brusca and Brusca, 1978:54, fig. 30.—Gerrodette, 1979:227-234; 

1981:611-618.—Lewbel et al., 1981:165.—Fadlallah and Pearse, 

1982a:223-230. 9 figs.—Fadlallah, 1983a: 132; 1983b:20O-2O6, 7 figs — 

Austin. 1985: 81.—Bythell, 1986:20, pi. 3: fig. B; pi. 11: figs. A-F — 

Kozloff, 1987:72.—Hellberg, 1991:30.—Cairns et al., 1991:48.— 

Chadwick, 1991:42-47.

NUMBER 557 33 

DESCRIPTION.—Corallum cylindrical to trochoid, often short 

and squat, but occasionally taller and more vasiform. Pedicel 

robust: PD:GCD = 0.50-0.80; base polycyclic (Durham, 

1949). Largest specimen examined (Alb-4552, Plate \2b,c) 

16.8 x 13.7 mm in calicular diameter and 17.3 mm in height. 

Coralla invariably secondarily epithecate, the epitheca covering 

at least the lower third to as much as the entire underlying 

synapticulothecate wall. Epitheca often encrusted with coralline 

algae, bryozoans, and serpulids. Synapticulotheca porous; 

costae usually not well developed, but when present are low 

and rounded, separated by porous intercostal furrows about 

one-quarter width of a costa. Costae covered with small fine 

granules. Corallum white; polyp a vivid red-orange or bright 

red. 

Septa hexamerally arranged in four or five incomplete 

cycles. Fourth cycle (48 septa) complete at a GCD of about 7-8 

mm, half the fifth cycle (72 septa) at a GCD of 11-12 mm, and 

up to 88 septa in the largest coralla examined; a full fifth cycle 

was not observed. S^_2 equal in size, only slightly exsert, and 

have straight, entire inner edges that attain the columella. S3 

one-third to half width of Su2 and have dentate lower inner 

edges. Remaining septa arranged in a Pourtales Plan (Figure 2). 

In small coralla having no S5, each pair of S4 meet before its 

adjacent S3 and extends to the columella as a single septum. 

When pairs of S5 occur, they first occur flanking S4 that are 

adjacent to S.,, not those adjacent to S2 (Figure 2, lower right 

half-system adjacent to medial S^. At this stage the pair of S5 

in each half-system unites before its adjacent S3 and also joins 

with the singleton S4 near the columella. After 12 pairs of S5 

are inserted in this manner (72 septa), additional pairs of S5 

form flanking the S4 adjacent to the S2 (Figure 2, lower right 

half-system adjacent to lateral S^. Inner edges of S4_5 

laciniate. All septal faces finely granular. Fossa of moderate 

depth, containing an elongate, spongy columella, which may 

have a flat of concave surface. 

DISCUSSION.—Balanophyllia elegans is without doubt the 

most commonly collected and highly visible scleractinian in 

the northeastern Pacific and therefore has been cited in many 

fieldguides (see synonymy) and is one of the few azooxanthellates 

for which life history information is known. Durham 

(1947) used it as an example to explain his discovery of 

polycyclic development, by which he illustrated that B. elegans 

originally possesses an epithecal prototheca, surrounded by 

two concentric synapticulothecate walls, the outermost usually 

partially covered by another epitheca. Gerrodette (1981) 

published a study of larval dispersal of the species, showing it 

to have demersal planulae with extremely limited dispersal 

ability. In a study of its reproductive habits, Fadlallah and 

Pearse (1982a) discovered that it reproduced only sexually, had 

populations of equal sex ratio, and brooded large embryos to an 

advanced planula stage in large calicular cavities structured by 

its Pourtales Plan septal arrangement. Production of sperm was 

found to be seasonal (late summer) but oocytes were found year 

round in females. Population densities of 530/m 2 were 

reported. In a follow-up study on growth, population dynamics, 

and demography of B. elegans, Fadlallah (1983b) estimated 

coralla to have a longevity of 6-11 years, based on mortality 

curves and growth rates, the latter averaging 1.7 mm/year in 

calicular diameter, 1.8 mm/year in height, and 175 mm 3 /year in 

volume. Population densities ranged from 385-816 coralla/m 2 

in his study area. Individuals reached reproductive maturity 

relatively early (less than a year) and planula mortality was 

relatively low. 

Comparisons to its neighboring species B. cedrosensis are 

made in the account of that species. 








USNM 92626; Alb-4552, 2, USNM 92627; off Cordell Bank, 

73 m, 8, USNM 92628 and CAS 74802, CAS 74834, CAS 

80940, CAS 74836; Monterey Bay, 42, USNM 78640 and CAS 

74810, CAS 16325, CAS 74809, CAS 74814; Moss Beach, 15, 

USNM 78636 and CAS 7602; off Nanaimo, B.C., 1, USNM 

92629; off Banks I., B.C., 1, USNM 92630; Pleistocene of San 

Pedro, 1, USNM 92631; off Pigeon Point, 1, CAS 6149; Neah 

Bay, B.C., 3, CAS 69606; off Santa Barbara, 1, CAS 74817; 

36°3TSf, m^W, 183 m, 3, CAS 74808; Snipe Bay (west 

coast of Baranof I.), Alaska, 3, UA; Quast Rock, La Jolla, 21 m, 

15, SIO Co 1164 and 1186; Ben's Rock, Isla San Martin, 

Pacific coast of northern Baja California, 27 m, 8, SIO Co 

1171; Islas Coronados, Pacific coast of northern Baja California, 

15 m, 9, SIO Co 1170; Point Loma, 13 m, 7, SIO Co 1173; 

127 specimens collected off British Columbia deposited at the 

RBCM, cataloged as: 973-180-2, 973-138-3, 973-251-10, 

973-154-2, 973-166-2, 973-139-23, 974-230-8, 974-234-2, 

974-239-2, 975-753-23, 975-772-26, 976-1040-3, 976-1238- 

59, 976-111-39, 976-111-38, 977-156-10, 980-350-19, 980- 

327-24, 980-381-5, 980-338-24, 980-267-29, 980-343-17, 

981-212-7. Previous Records: Syntypes of B. elegans; 

specimen reported by Durham (1947) [Alb-4534, 1, USNM 

M547412; Alb-4543, 2, USNM M547414-15; Alb-4555, 1, 

USNMM547413]. 

TYPES.—Forty-two syntypes of B. elegans are deposited at 

the MCZ in three lots: Crescent City (old MCZ 1085, new 

MCZ 5472, Plate \2e,J), 3 dry syntypes; Crescent City (MCZ 

223), 23 syntypes preserved in alcohol; Mendacino (MCZ 

222), 16 syntypes preserved in alcohol. All syntypes were 

received and cataloged in the early 1860s and bear Verrill's 

handwriting regarding type status. Type Locality: Off Crescent 

City and Mendacino, California, depth unknown. 

DISTRIBUTION.—Pliocene: San Diego (Hertlein and Grant, 

1960). Pleistocene: San Pedro, California (Addicott, 1966). 

Recent: Snipe Bay, Alaska to Sacramento Reef, Pacific coast of 

northern Baja California (29°44'N) (Gerrodette, 1979), including 

Queen Charlotte Islands, Vancouver Island, and Cortes, 

Tanner, and Cordell Banks. Bathymetrically known from

34 

0-293 m (Durham, 1947), with one outlying record at 587 m 

(Durham and Barnard, 1952). According to Gerrodette (1979), 

specimens south of Point Conception are always found deeper 

than 10 m, which he suggests is an example of equatorial 

submergence that occurs at the cold-warm temperate zoogeographic 

boundary. 

Balanophyllia cedrosensis Durham, 1947 

PLATE 1 \g-j 

Balanophyllia cedrosensis Durham. 1947:40-41, pi. 11: figs. 3, 5.—Durham 

and Barnard, 1952:99, pi. 14: fig. 61a,b. 

Balanophyllia tiburonensis Durham, 1947:41-42, pi. 10: figs. 5, 7 [new 

synonym].—Squires, 1959:423-426. 

REDESCRIPTION OF HOLOTYPE.—Corallum poorly preserved, 

obviously dead when collected. Corallum ceratoid: 13.7 

x 10.4 mm in calicular diameter and 16.4 mm in height, with a 

pedicel diameter of 6.2 mm. Costal granulation worn; 

intercostal striae almost as wide as costae and quite porous. A 

thin epitheca covers lower two-thirds of corallum. Septa 

hexamerally arranged in 5 cycles, the last cycle incomplete, the 

corallum having approximately 64 septa. S^_2 equal in size, 

porous near the theca, but solid on edge toward fossa. Inner 

edges of S^ vertical and straight, reaching the columella. S3 

and S4 bear large pores, each pair of S4 fused before its adjacent 

S3 and continuing to the columella. Pairs of S5 present in some 

half-systems, each pair fused to their adjacent S4. Depth of 

fossa difficult to determine because of damage to corallum. 

Columella discrete, elongate, and spongy, the numerous 

columellar elements quite fine and fused to one another. 

DISCUSSION.—Examination of nontype specimens show that 

the species can attain a calicular diameter of 19 x 15 mm and a 

height of 23 mm (Pillsbury-521) and have a full fifth cycle of 

septa (96). With greater age and size, the porosity of the septa 

decreases until they are solid. No other specimens examined 

displayed an epitheca, the costae being distinct from calice to 

base. The inner edges of the fused S4 are usually coarsely 

dentate, not smooth as is the case in the worn type. Finally, the 

columella may consist of a rounded, elliptical field of slightly 

swirled elements. 

Balanophyllia cedrosensis is distinguished from B. elegans 

by having a discrete, convex columella; porous septa in early 

ontogeny; and less dentate inner S4 margins. It lacks the stellate 

septal pattern characteristic of the strong Pourtales Plan of B. 

elegans. The species are also geographically distinct, B. elegans 

known only north of Sacremento Reef, northern Baja California, 

and B. cedrosensis from south of Isla Cedros to the Bay of 

Panama, including the Gulf of California. No differences were 

found between the types of B. cedrosensis and B. tiburonensis, 

notwithstanding the variation discussed by Durham (1947, 

text-fig. 2B.C). 

MATERIAL EXAMINED.—New Records: Pillsbury-52\, 5, 

USNM 83531; Pillsbury-530, 20, USNM 83532. Previous 

Records: Holotype of B. cedrosensis; holotype and paratype 


of B. tiburonensis; Velero 1259-41, 1, SBMNH (Durham and 

Barnard, 1952). 

TYPES.—The holotype (Plate 11/-;) of B. cedrosensis is 

deposited at the CAS (29961). Type Locality: "Near Cedros 

Island on the way to San Benito," depth unknown. 

The holotype (Plate 1\g) and paratype of B. tiburonensis are 

deposited at the UCMP (14833, 14834, respectively). Type 

Locality: 28°44.3'N, 112°34.6'W (off Tiburon I., Gulf of 

California), 73 m. 

DISTRIBUTION.—Known only from a small region from 

Cedros Island to Punta Abreojos, Pacific coast of Baja 

California Sur, off Tiburon Island, Gulf of California; and the 

Bay of Panama; 66- 119m. 

Dendrophyllia Blainville, 1830 

DIAGNOSIS (emended).—Extratentacular budding forms 

colonies of three general forms: arborescent colonies with axial 

corallites, small bushy colonies with sparse branching, or 

dendroid colonies with sympodial branching. All three forms 

originate from a single basal stem. Synapticulothecate; costae 

usually well defined. Septa arranged in a Pourtales Plan. Pali 

may be present or absent; columella spongy. Tabular endothecal 

dissepiments may be present. Azooxanthellate, often found 

in shallow water. 

TYPE SPECIES.—Madrepora ramea Linnaeus, 1758, by 

subsequent designation (Milne Edwards and Haime, 

Dendrophyllia oldroydae Oldroyd 1924, nom. correct 

PLATES \2jjc, \3a,b 

Dendrophyllia oldroydi Oldroyd, 1924, pi. 49: fig. 7.—Williams, 1936:27-28, 

1 fig- 

Dendrophyllia oldroydi Faustino, 1931:286-287, pi. 1: fig. 2.—?Durham, 

1943:201, pi. 32: fig. 1; 1947:38, pi. 10: figs. 1,9.—Bythell, 1986:21, pi. 12: 

figs, a-d.—Prahl, 1987:230, figs. 6, 7.—Prahl and Erhardt, 1989:549-550, 

fig. 9.—Cairns et al., 1991:48. 

Dendrophyllia californica.—Durham and Barnard, 1952:101 [in part: pi. 15: 

fig. 65a,b].—Cairns, 1991a:23-24, pi. 10: figs. c-e. 

Dendrophyllia cortezi Durham and Barnard, 1952:102-103, pi. 16: fig. 

66a,b.—Squires, 1959:426. 

Dendrophyllia cf. D. oldroydi Faustino.—Hertlein and Grant, 1960:82-83, pi. 

19: figs. 5, 6, 15. 

DESCRIPTION.—Corallum sympodially branched, forming 

large bushy colonies up to 1 m in height (Cairns, 1991a) and 

having robust basal branches up to 5 cm in diameter; largest 

colonies examined from off California only 35 cm in height 

with a basal branch diameter of 2.2 cm. Corallites short and 

stout on large-diameter basal branches, projecting only 4-7 

mm perpendicular to main branch (e.g., Durham and Barnard, 

1952, pi. 15: fig. 65a), but distal corallites, especially those of 

fast growing coralla, are inclined distally and elongate, up to 23 

mm long (e.g., Williams, 1936, fig. 1). Calices elliptical and 

usually less than 10 mm in GCD, but may reach up to 15 x 13 

mm in GCD x LCD (USNM 80453). Costae equal in width and 

slightly convex, separated by narrow intercostal striae and

NUMBER 557 35 

covered with small (0.10 mm diameter) granules. However, 

within 5-7 mm of the calice, intercostal striae are often 

perforate, and C^_2 in this region are slightly narrower than the 

others and slightly ridged. Corallum white; coenosarc yellow, 

drying to a dark brown. 

Septa hexamerally arranged in 5 cycles, the last cycle always 

incomplete. Each half-system usually contains either no S5, 

sometimes 1 pair of S5, or rarely 2 pairs of S5, for a total of only 

48 septa if no S5 are present, and up to 72 if 12 pairs of S5 are 

present, as in the case of the largest calice. Most corallites have 

48-58 septa (0-6 pairs of S5). S^ equal in size, only about 

1.5 mm exsert, relatively narrow (only about 1.5 mm wide), 

and have straight inner edges that attain the columella. S3 little 

exsert and are the smallest septa, only about 0.5 mm wide and 

merging into the paliform lobe formed by flanking S4. S4 

paired and equal in exsertness to S3, but about 1.2 mm in width, 

the inner edges of each pair solidly fused before its adjacent S3 

to form a tall, rounded paliform lobe, which is thicker than an 

S3 and of equal width to an S4. In most corallites these 12 P3 

form a crown of lobes, but in larger corallites having pairs of 

S5, 2 P4 are formed in each half-system instead of 1 P3. Inner 

edges of paliform lobes reach columella and appear to be 

continuous with columellar structure. Fossa shallow, containing 

a columella of variable size and structure. Columella 

usually composed of an elongate to elliptical mass of swirled 

elements, sometimes appearing fascicular, other times as 

coarsely papillose. Columella width varies from rudimentary, 

only 12% LCD, to quite large (Plate 12/), up to 37% LCD. All 

septal faces, paliform lobes, and columellar elements covered 

with a fine granulation. 

DISCUSSION.—Dendrophyllia oldroydae was first published 

as a figure and its caption by Oldroyd (1924) without 

description, but with a reference to an unpublished manuscript 

by Faustina (sic). According to Article 12b7 of the International 

Code of Zoological Nomenclature (1985), because this 

illustration and figure caption were published before 1931, they 

satisfy the criteria of availability, and thus Oldroyd (1924) must 

be considered as the author of the species. When Faustino's 

(1931) D. oldroydi was finally published as a new species 

account, based on topotypic but different specimens, it thus 

must be considered as a junior primary homonym as well as a 

junior subjective synonym, subjective because different holotypes 

were chosen. The intent of the name in both cases was 

obviously to honor Ida S. Oldroyd, and thus the spelling of the 

name should be changed to the feminine oldroydae. 

I (Cairns, 1991 a) was in error to have identified specimens of 

D. oldroydae from the Galapagos Islands as D. californica. At 

that time I had not examined the types of D. oldroydi nor did I 

appreciate the great variability of its columella and growth 

form. I was also influenced by Durham and Barnard's (1952, 

pi. 15: fig. 65b) misidentification of their D. californica. 

However, I still support the synonymy of D. cortezi and D. 

oldroydae, the former having poorly formed paliform lobes 

(dentate inner margins), slightly porous S4 fusions, and 

elongate corallites, all characteristic of a rapidly growing 

terminal branchlet of D. oldroydae. 

Dendrophyllia oldroydae is compared to D. californica in 

the account of the latter species. 

MATERIAL EXAMINED.—New Records: 6.4 km off Del 

Mar, California, 99-125 m, 1 colony, USNM 80453; San 

Pedro Bay, California (topotypic), 366 m, 10 colonies, USNM 

38114; 27°28.3'N, 114°51'W, 110 m, 2 branches, SIO Co 1317. 

Previous Records: Holotype and paratypes of D. oldroydi 

Faustino, 1931; off Huntington Beach, 183 m, 1 colony, 

USNM 92610 (Williams, 1936); holotype of D. cortezi; Velero 

1254-41, 1 colony, SBMNH 35563 (D. californica of Durham 

and Barnard, 1952); off Gorgona I., Colombia, 40 m, 1 colony, 

USNM 78795 (Prahl, 1987); off La Jolla, 100-180 m, 1 

colony, SIO Co 1179 (Bythell, 1986). 

TYPES.—The holotype of D. oldroydi Oldroyd, 1924, 

originally deposited in the Paleontology Department of 

Stanford University, appears to be lost (Hertlein and Grant, 

1960). Type Locality: Off San Pedro, California, 366 m. 

Faustino (1931) mentioned two specimens in the description 

of his D. oldroydi, the illustrated specimen (holotype (Plate 

\2k), cataloged as CAS 36397) being part of a larger colony 

having two more pieces. The identification of his "second" 

specimen, which must be considered a paratype (Plate 13b), is 

confused, since the CAS has eight other colonies of the same 

species from the same locality also numbered 36397. There is 

also a specimen of D. oldroydi deposited at the UCMP (12200) 

designated as a paratype, mentioned by Durham (1947). 

Finally, there are 10 colonies from the same collection 

deposited at the USNM (38114), which are topotypic but not 

considered to be type specimens. Type Locality: "Sunken 

Valley, between San Pedro and Redonda, California, 200 

fathoms [= 366 m]; deep water off San Diego, California." 

The holotype (Plate 13a) of D. cortezi is deposited at the 

SBMNH (35564) ex AHF 24. Type Locality: Velero 561-36: 

south of Isla Partida, Gulf of California, 128 m. 

DISTRIBUTION.—Dendrophyllia oldroydae is the most common 

Dendrophyllia found off California and Baja California, 

occasionally being snagged by deep-sea fishermen. It is known 

from off Redondo (type locality) to halfway down the Pacific 

coast of Baja California (27°28'N), 99-366 m; also off Angel 

de la Guardia and Isla Partida, Gulf of California. Middle 

Pliocene San Diego (Hertlein and Grant, 1960). ?Plio- 

Pleistocene of Humboldt Co., California (Durham, 1943). 

Elsewhere: Off Pacific coast of Colombia (Prahl, 1987) and 

off Galdpagos and Cocos Islands (Cairns, 1991a); 40-576 m. 

Dendrophyllia californica Durham, 1947 

PLATE ]2g-i 

Dendrophyllia californica Durham, 1947:37-38. pi. 10: figs. 2. 6.—Bythell, 

1986:21, pi. 12:figs.e.f. 

Not Dendrophyllia californica.—Durham and Barnard, 1952:101-102, pi. 15: 

fig. 65a.b [= D. oldroydae].—Caims. 199la:23-24. pi. 10: figs, c-e [= D 

oldroydae]. 

DESCRIPTION.—Colonies small, none known over 5 cm in

36 

height or diameter. Extratentacular budding sparse, colonies 

examined have no more than 6 or 7 corallites and lack third 

generation buds. Corallites elongate and ceratoid, up to 19.9 x 

14.4 mm in calicular diameter and 32 mm long. Pedicel 

diameter of holotype 14.8 mm. Costae slightly convex and 

separated by narrow, porous intercostal striae. C^_2 slightly 

narrower than other costae, but not ridged. Costae finely 

granular, producing a rough texture. Corallum white. 

Septa hexamerally arranged in 5 cycles, the last cycle 

complete only in largest corallites. Sy_2 only about 0.7 mm 

exsert and quite slender (only about 1.5 mm wide), with 

straight, vertical inner edges that merge with columella low in 

fossa. Remaining septa barely exsert, the S3 being the smallest 

septa, only about 0.9 mm wide. Inner edges of each pair of S4 

form a rudimentary, porous fusion before the S3 and extend to 

the columella but do not contribute to it. Occasionally pairs of 

S4 do not quite fuse near the columella, especially if S5 are 

present. Inner edges of the 12 S^ and 12 combined S4 extend 

same distance into calice. If present, inner edges of S5 fuse 

before their adjacent S4. Paliform lobes absent. Fossa deep. 

Columella similar to that of D. oldroydae: an elongate, swirled 

mass of elements, but modest in size, only 20%-25% LCD in 

width. 

DISCUSSION.—Dendrophyllia californica is distinguished 

from D. oldroydae by having a small colony with very sparse 

branching; longer, larger corallites with more septa; and a much 

deeper fossa. It also differs in lacking paliform lobes, lacking 

ridged costae, and in having rudimentary, porous S4 fusions. It 

also is known from a more limited geographic and depth range. 

MATERIAL EXAMINED.—New Record: Alb-2832, 2 

branches, USNM 38113. Previous Records: Holotype of D. 

californica; 26°12TSf, 112°35'W, 51-55 m, 1 colony, SIO Co 

437 (Bythell, 1986). Reference Specimens: Specimens reported 

as D. californica by Durham and Barnard (1952) and 

Cairns (1991a). 

TYPES.—The holotype (Plate \2g,h) of D. californica is 

deposited at the CAS (29960). Type Locality: 27°52'3O"N, 

114°54'45"W (Bahia Sebastian Vizcaino, near Cedros I., 

Pacific coast of Baja California), 42 m. 

DISTRIBUTION.—Known from only three localities from the 

Pacific coast of Baja California, from Bahia Sebastian Vizcaino 

to Bahia Magdalena; 42-93 m. 

Part 2: Temperate Northwestern Pacific 

Suborder ASTROCOENIINA 

Family POCILLOPORIDAE 

Madracis Milne Edwards and Haime, 1849 

DIAGNOSIS.—Colonial, extratentacular budding producing 

massive, encrusting, or branching coralla. Coenosteum solid. 

Septa arranged in groups of 6, 8, or 10, but rarely in more than 

two cycles (i.e., 12, 16, or 20 septa). Columella styliform; 

paliform lobes sometimes present. 


TYPE SPECIES.—Madracis asperula Milne Edwards and 

Haime, 1849, by monotypy. 

Madracis sp. A 

PLATE 13C-/ 

?Madracis sp.: Wells. 1954:414, pi. 99: fig. 5. 

Madracis asanoi.—Eguchi, 1968:C11, pi. C5: figs. 3-7. 

DIAGNOSIS.—Colonies delicately branched, terminal branchlets 

slender (2-3 mm in diameter), basal branches somewhat 

compressed and up to 5 mm in diameter. Largest colony 

examined (ZMC, Sagami Bay) only 26 mm in height. 

Corallites circular to elliptical in shape and closely spaced, 

some directly adjacent to one another, and rarely separated by 

more than 1 mm from adjacent corallites. Corallites 1.7-2.5 

mm in diameter. Coenosteum faintly striate and covered with 

tall (up to 0.4 mm), slender (0.12-0.18 mm) coenosteal spines. 


Septa decamerally arranged in 2 cycles (10:10 = 20 septa). 

Primary septa highly exsert but rather slender, each bearing a 

small paliform lobe, the 10 lobes forming a crown encircling 

the columella. Secondary septa rudimentary, each composed of 

a row of small, slender teeth. Fossa relatively shallow, its 

central portion a circular horizontal platform on which the 

columella and paliform lobes project. Columella a prominent, 

plate-like style up to 0.4 mm in height and 0.14 mm in 

diameter, usually reaching above the calicular edge. Longer 

axis of columellar plate aligned parallel to branch axis. 

DISCUSSION.—The geographically closest known species of 

Madracis to Japan is M. asanoi Yabe and Sugiyama, 1936, 

described from the Pelau Islands. Madracis asanoi is similar to 

the Japanese species in calicular diameter and corallite spacing, 

but differs in having only 10 (not 20) septa; thicker, more 

robust branches; and in lacking paliform lobes. The Japanese 

specimens are quite similar to the unnamed species of 

Madracis reported by Wells (1954) from Bikini Atoll, Marshall 

Islands; however, the growth form of that small specimen was 

encrusting. It is not possible to determine if the encrusting 

mode was the permanent growth form of this specimen or just 

the foundation of a larger branching colony. Madracis 

kauaiensis Vaughan, 1907, (Hawaiian and Johnston Islands) 

differs in having robust branches; smaller, more widely spaced 

corallites; only one cycle of 10 septa; no paliform lobes; and a 

much smaller columella. Madracis sp. cf. M. asperula, reported 

by Durham and Barnard (1952) and Cairns (1991a) from the 

Galapagos Islands, differs in having smaller, more widely 

spaced corallites; only 10 septa per corallite; and no paliform 

lobes. Thus, the specimens from Sagami Bay appear to 

represent an undescribed species, but not enough material is 

available to justify the proposal of a new species. 

MATERIAL EXAMINED.^MW Records: Off Okinose, Sagami 

Bay, 110 m, Mortensen's Pacific Expedition of 1914, 3 

colonies, ZMC; 26°30'N, 127°50'54"E (off Okinawa), 64 m, 1 

colony, USNM 88378. Previous Records: Madracis sp. of 

Wells (1954), USNM 44407. Reference Specimens: Types of

NUMBER 557 37 

M. kauaiensis, USNM; M. sp. cf. M. asperula of Durham and 

Barnard (1952), AHF 1.1. 

DISTRIBUTION.—Sagami Bay, Honshu; off Okinawa, 

Ryukyu Islands; ?Bikini Atoll, Marshall Islands; ?46-55- 

110 m. 

DIAGNOSIS.—See Part 1. 

Suborder FUNGIINA 

Superfamily FUNGIOIDEA 

Family FUNGIACYATHIDAE 

Fungiacyathus Sars, 1872 

Subgenus Fungiacyathus (Fungiacyathus) Sars, 1872 

DIAGNOSIS.—Fungiacyathus having five cycles of septa. 

Fungiacyathus (F.) stephanus (Alcock, 1893) 

PLATE \3g-i 

Bathyactis stephanus Alcock, 1893:149, pi. 5: figs. 12, 12a. 

Bathyactis Stephana.—Alcock, 1898:28-29, pi. 3: figs. 5, 5a; 1902c:38. 

Fungiacyathus stephanus.—Caims, 1989a:7-9, pi. 1: figs, a-k; pi. 2: figs. a,b 

[synonymy].—Cairns and Keller, 1993:230: 

DESCRIPTION.—Corallum circular and quite fragile, the 

more typical concave-base form up to 39 mm in diameter and 

20 mm in height (D.H = 1.95), whereas the flat-base form 

attains 45 mm in diameter but only 16-17 mm in height (D:H 

= 2.7). Costal ridges thin, finely dentate, and straight to slightly 

sinuous. Corallum white. 


to the formula: S1>S2>S3>S4>S5. S1 composed of an 

extremely tall inner lobe bearing 12-14 vertical, finely dentate 

ridges (trabeculae) and a lower outer marginal shelf about 5 

mm wide and only about 1.5-2.0 mm in height. Ten to 13 

synapticulae occur along an S, of a large specimen, the largest 

synapticulae being 6th or 7th from columella, which rise about 

to level of adjacent S4. Each S2 bears a prominent paliform lobe 

internally, followed by a tall septal lobe (but smaller than that 

of the Sj) composed of 10-12 trabeculae, and a similar outer 

marginal shelf. Each S3 also has an inner paliform lobe but 

more recessed from the columella, a smaller and lower septal 

lobe, and a peripheral marginal shelf. S4 joined to adjacent S3 

in a canopied structure, lack paliform lobes, and have only a 

small septal lobe and low marginal shelf. S5 quite small, 

lacking both paliform and septal lobes, and merge with 

adjacent S4 via a long, low canopied structure; their outer shelf 

region is only about 0.5 mm in height. All septal faces 

corrugated, but upper septal edges are often straight. Columella 

a low, circular trabecular mass 4-5 mm in diameter. 

DISCUSSION.—A more complete synonymy and description 

of this species is given by Cairns (1989a) in the context of a 

revision of the Philippine deep-water Scleractinia. The records 

reported herein constitute a northern range extension for the 

species and the first records from the Japanese region. 

Fungiacyathus stephanus is distinguished from other species 

in the subgenus by having extremely tall septal lobes, a broad 

marginal shelf, and corrugated septa. Bathymetrically, it is 

intermediate between the shallower F. paliferus and the much 

deeper F. sp. A. 



USNM 80853; Alb-4911, 1, USNM 80123; Alb-4912, 1, CAS 

1160; Alb-4915, 2, CAS 74999; Alb-4916, 1, USNM 80124; 


AIb-4958, 1, USNM 80127; Alb-4959, 1, USNM 92633; 


Alb-4968, 3, CAS 80929; Alb-4970, 2, USNM 80129; 

Alb-5086, 6, USNM 80130; TM (KT9202, YT6), 1, ORI. 

TYPES.—The holotype of B. stephanus is presumed to be 

deposited at the Indian Museum, Calcutta. Type Locality: 

Investigator-133:15°43'30"N, 81 ° 19'30"E (off Kistna 

Delta, Bay of Bengal), 1240 m. 

DISTRIBUTION.—Japan: Off southeastern Honshu from Sagami 

Bay to Wakayama Prefecture; Bungo Strait; off Koshiki 

Island, Kyushu; East China Sea off northern Ryukyu Islands; 

446-1317 m. Elsewhere: Throughout the Philippines, 

Indonesia, and western Indian Ocean (Caims, 1989a); 245- 

1977 m. 

Fungiacyathus (F.) paliferus (Alcock, 1902) 

PLATE \4a-e 

Bathyactis palifera Alcock, 1902a:108; 19O2c:38. pi. 5: figs. 34, 34a.—Yabe 

and Eguchi, 1942b: 137-138. pi. 12: fig. 5. 

Fungia (Diaseris form).—Yabe and Eguchi, I932e: 387. 

Bathyactis symmetrica.—Yabe and Eguchi, 1942b: 137 [in part: Syo Afaru-283, 

Soyu Maru-259]. 

Bathyactis kikaiensis Yabe and Eguchi. 1932b:443 [nomen nudum]; 

1942b: 138, 155-156, pi. 12: figs. 6, 7. 

? Fungiacyathus symmetricus.—Utinomi, 1965:248-249. 

Not Fungiacytahus palifera.—Keller, 1976:33-34 [= Fungiacyathus sp. A]. 

Fungiacyathus paliferus.—Cairns, 1989a:9-10, pi. 2: figs, c-i; pi. 3: figs, a-c 

[synonymy].—Caims and Parker, 1992:6-7, fig. la,b. 

DESCRIPTION.—Coralla occur in two shapes: (1) circular 

(Plate 14a), up to 16.2 mm in calicular diameter and 7.0 mm in 

height (D:H = 2.3-3.45), and (2) semi-circular (Plate 14fr), 

rarely greater than 10 mm in diameter, the latter form 

apparently the result of fragmentation and referred to as the 

Diaseris form by Yabe and Eguchi (1932a, 1942b) because of 

its resemblance to that genus. Base flat to only slightly 

concave, bearing low, rounded, granulated costae. Corallum 

white. 


to the formula: S1>S2>S3>S4>S5 (98 septa). The semi-circular 

Diaseris form usually consists of 2 full systems flanked by 2 

half-systems, including the S2, for a total of 48 septa, but 

paliform lobes and columellas are usually absent in these 

forms. S1 consist of 4 or 5 inner trabecular spines; an 

intermediate septal lobe bearing up to 15 coarsely dentate,

38 

closely spaced carinae; and a low, arched peripheral marginal 

shelf about 1.5 mm wide. Seven or eight synapticulae occur 

along an S1f the largest being the fourth or fifth from the 

columella. S2 consist of 2 or 3 inner spines, a broad paliform 

lobe, an intermediate septal lobe of 11 or 12 trabeculae, and a 

peripheral marginal shelf. S3 have a broad internal paliform 

lobe, a small septal lobe composed of 5-8 trabeculae, and a 

marginal shelf. S4 consist of only a small septal lobe of 4 or 5 

trabeculae and a small marginal shelf. S5 exist only in the 

marginal shelf region and are not lobate or spinose. All septa 

planar, with straight septal edges. Columella rudimentary to 

absent. 

DISCUSSION.—Fungiacyathus paliferus is easily distinguished 

from the two other Japanese congeners having five 

cycles of septa by having granular costae; a small, relatively 

robust corallum; and a tendency to asexually fragment into 

semi-circular fragments. Bathymetrically, it is the shallowest of 

the three species. 

As noted in a previous publication (Cairns, 1989a), 

Bathyactis kikaiensis is undoubtedly a junior synonym of F. 

paliferus, but the types of this species were not available for the 

confirmation of the synonymy. 

MATERIAL EXAMINED.—New Records: TM (KT9015, 

BS1), 1, USNM 92634; TM (9015, BS2), 4, USNM 92635; TM 

(KT9015, CB1-2), 4, USNM 92636; TM (9015, CB1-1), 1, 

USNM 92637; TM (KT9015, HK5), 4, ORI; TM (KT9015, 

OKI), 2, USNM 92638; TM (KT9015, OK2), 1, USNM 92639; 

TM (KT9202, OS2), fragments, USNM 92640; TM (KT9202, 

OS3), 1, USNM 92641; TM (KT9202, YS1), 4, USNM 92642; 

TM (KT9209, AM6), 1, ORI. Previous Records: Soyu 

Maru-412, 1, TIUS 58913 (B. palifera of Yabe and Eguchi, 

1942b); Soyu Maru-259, 1, TIUS 58948 (F. symmetrica of 

Yabe and Eguchi, 1942b); Soyu Afaru-283, 1, TIUS 58950 (F. 

symmetrica of Yabe and Eguchi, 1942b); syntypes of B. 

palifera, 3, ZMA. 

TYPES.—Three syntypes of Bathyactis palifera are deposited 

at the ZMA, collected at Siboga-153 (Coel. 1171) (van Soest, 

1979). Type Localities: Siboga stations 98 and 153: Sulu Sea 

and off the Moluccas, 143-350 m. 

One hundred ten syntypes of B. kikaiensis are deposited at 

the TIUS, the measurements of three of which are given in the 

text. Two catalog numbers were given to this lot by Yabe and 

Eguchi: 50236 (pages 155-156 and plate caption) and 50097 

(pages 138, 173). The earlier number was also assigned to the 

type of Stephanophyllia japonica, which might imply that 

50097 is the correct number. Type Locality: Plio-Pleistocene 

of the Ryukyu Limestone of Kakai-jima, Kagoshima-ken. 

DISTRIBUTION.—Japan: Suruga Bay, Honshu; off Shikoku 

and Bungo Strait; off southern Kyushu; East China Sea off 

Danjo Gunto; off Mi Shima, Korea Strait; Oki Strait, Sea of 

Japan; northern Tokara Retto and off Amami Oshima, Ryukyu 

Islands; 70-364 m. Elsewhere: Pleistocene of Vanuatu and 

Ryukyu Islands; Recent: Philippines, Indonesia, Great Australian 

Bight, Reunion; 75-522 m (Cairns, 1989a: 10). 


Fungiacyathus (F.) sp. A 

PLATE 14/-/I 

Fungiacyathus palifera [sic].—Keller, 1976:33-34, pi. 1: figs. 1, 2. 

DIAGNOSIS.—Two specimens of Keller's (1976) F. palifera 

were examined, both from Vityaz-2209, including her figured 

specimen, which is 35.7 mm in calicular diameter and 6.0 mm 

in height and missing 2 systems of its corallum. Its base is flat 

and thin, bearing 5 cycles of sinuous, nongranular costal ridges. 

Five full cycles of septa are present (96 septa). All septa planar 

and have straight inner edges, each Su2 bearing 20-23 low, 

closely spaced trabecular carinae. Large, porous canopies occur 

at S3_4 and S4_5 junctions. Eight to nine synapticulae occur per 

Sv the outermost being the most highly developed. Columella 

4.5 mm in diameter, circular, and quite spongy. 

DISCUSSION.—The specimen described above is distinguished 

from F. paliferus by having a much larger corallum, 

lacking paliform lobes, and having ridged (not rounded or 

granular) costae. It is actually more similar to F. stephanus than 

F. paliferus, especially in corallum size and fragility, but differs 

in having relatively low and straight (noncorrugated) septa, no 

marginal shelf or paliform lobes, and very spongy columellar 

and canopy regions. It differs from F. fragilis Sars, 1872, in 

having planar septa and spongy canopies and columella. 

Fungiacyathus sp. A is bathymetrically and geographically 

distinct from the two other Japanese species that have five 

cycles of septa in being by far the deepest of the three and the 

most northerly in distribution. 

Despite the distinctness of Fungiacyathus sp. A, until 

additional specimens are collected and examined, a formal 

description is postponed. 

MATERIAL EXAMINED.—Vityaz-2209, 1, IOM, and 1 central 

disc, USNM 92643 (Plate 14/-/i). 

DISTRIBUTION.—Off eastern Kurile Islands bordering Kurile 

Trench; 3175-4110 m. Keller's (1976) mapping of Vityaz- 

5603 and V7ryaz-5635 off the Aleutian Islands and off Honshu, 

respectively, are mistaken, the coordinates of all three of her 

stations being off the Kurile Islands. 

Subgenus Fungiacyathus (Bathyactis) Moseley, 1881 


Fungiacyathus (B.) marenzelleri (Vaughan, 1906b) 

Account.—See Part 1. 

Fungiacyathus (B.) variegatus Cairns, 1989 

PLATE I5a,b 

Fungiacyathus (B.) variegatus Cairns, 1989a: II-12, pi. 5: figs, a-h 

[synonymy]. 

DIAGNOSIS.—The largest Japanese specimen (Alb-4911) is a 

slightly worn corallum 9.0 mm in calicular diameter, probably

NUMBER 557 39 

dead when collected. Base slightly concave, circular in outline, 

but with a finely serrate edge, each septum projecting slightly 

beyond calicular perimeter. C^_2 more highly ridged than C3_4, 

each costa bearing a linear row of coarse granules. Corallum 

white. Septa hexamerally arranged in 4 complete cycles. S1 

consist of 4 or 5 inner trabecular spines and an outer lobe 

bearing 10-12 coarsely dentate ridges. S2 also have 5 or 6 inner 

spines, the third or fourth from the columella being quite robust 

and curved inward toward the columella. A pair of S3 joins to 

each S2 through a robust, solid canopy, each S3 consisting of 11 

or 12 trabeculae, which, like those of the lower cycle septa, 

extend above the upper septal edges as narrow spines. 

Likewise, each pair of S4 joins to an adjacent S3 through a 

robust canopy, each S4 consisting of 8 or 9 small trabecular 

spines. All septa planar, with straight inner edges. No marginal 

shelf present, the outer edges of all septa extending around 

calicular edge and thus continuous with underlying costae. 

Synapticulae sparse, only 3 or 4 along length of an Sv the 

innermost 2 linking the S, to its adjacent S4 in the region of the 

S^ canopy, and the outermost synapticulae linked directly to 

the S4. Columella rudimentary. 

DISCUSSION.—The Japanese specimens differ from the type 

series only in lacking the characteristic pigmentation of the 

species, but it is noted that some of the wom paratypes from 

Alb-5278 and Alb-5314 also lack pigmentation. Fungiacyathus 

variegatus is easily distinguished from other species in this 

subgenus in this region by usually having a pigmented 

corallum, having very few synapticulae, and quite welldeveloped 

septal canopies. 


USNM 82018; TM (KT9309, AM8), 3, USNM 93167, 3, ORI. 

Previous Records: Type series. 

TYPES.—The holotype and paratypes of F. variegatus are all 

deposited at the USNM (Cairns, 1991b). Type Locality: Alb- 

5113: H ^ H 120°51'E (off Luzon, Philippines), 291 m. 

DISTRIBUTION.—Japan: Off Koshiki Island, southwestern 

Kyushu; off Amami Oshima, Ryukyu Islands; 422-715 m. 

Elsewhere: Pleistocene of Vanuatu; Recent: Philippines; 

South China Sea off Hong Kong; 187-333 m (Cairns, 1989a). 

Fungiacyathus (B.) granulosus Cairns, 1989 

PLATE \5d,e 

Fungiacyathus granulosus Cairns, 1989a: 11, pi. 4: figs, d-i [synonymy]. 

DISCUSSION.—The single, worn specimen (now in three 

pieces) reported herein serves only to verify the existence of F. 

granulosus in Japanese waters, which is the northernmost 

record of the species. The specimen is 15.8 mm in calicular 

diameter and has a slightly concave base with coarsely 

granular, rounded costae. Each S, has approximately 22 closely 

spaced trabecular ridges and about 8 synapticulae, but the worn 

condition of the specimen does not allow remarks on the upper 

septal edges. All septa are planar with straight edges. The 

columella is circular and 2.6 mm in diameter. 

MATERIAL EXAMINED.—New Record: TM (KT9202, 

YT4), 1, USNM 92645. Previous Records: Type series. 

TYPES.—The holotype and paratypes are all deposited at the 

USNM. Type Locality: Alb-5590: 4°10'50"N, 118°39'35 / ^ 

(off Sabah, Celebes Sea), 567 m. 

DISTRIBUTION.—Off Japan: Osumi Shoto, northern Ryukyu 

Islands; 402-410 m. Elsewhere: Off Philippine Islands; 

Celebes Sea off Sabah; 390-567 m. 


Family MICRABACIIDAE 

Leptopenus Moseley, 1881 

Leptopenus discus Moseley, 1881 

ACCOUNT.—See Part 1. 

Leptopenus solidus Keller, 1977 

PLATE \Sg,h 

Leptopenus discus.—Squires. 1965:878-879, fig. 1; 1967:505. 

Leptopenus solidus Keller, 1977:40-41, pi. 1: fig. 5a,b, text-fig. 3. 

Leptopenus sp.—Cairns, 1989a: 15, pi. 6: figs. h,i. 

DESCRIPTION OF HOLOTYPE.—Corallum discoidal and very 

fragile, having only 2 peripheral costal spines intact. Corallum 

12.6 mm in maximum calicular diameter (including costal 

spines) and 3.1 mm in height, resulting in a D:H of 4.1. Costae 

alternate in position with septa, joined together only by thin 

synapticular bridges about 0.08 mm in diameter. Thecal pores 

up to 0.21 x 0.10 mm in diameter only in circular region 

0.35-2.9 mm from center of corallum, the epicenter and 

outermost 1.5 mm perimeter being solid. As in L. discus, 6 

major costae radiate from a solid, circular epicenter, each costa 

dividing twice to form 24 broad (0.30 mm wide), flat costae, 

which further subdivide into 68 costae in the holotype. 

Septa arranged in micrabaciid fashion; however, S, are not 

independent, but are usually united to an adjacent S2 near the 

center. S, consist of 5 or 6 tall, slender (about 0.15 mm in 

diameter), outwardly inclined trabecular spines that are united 

basally by a low web. The third and fourth trabecular spines 

from the center are the highest, their tips slightly recurved. S2 

are the most robust septa, also composed of 5 or 6 spines, the 

third of which is quite thick (about 0.35 mm in diameter), 

constituting the maximum corallum height. S2 trabecular 

spines also outwardly inclined but recurved at tips. Higher 

cycle septa composed of 1-3 spines of similar size to those of 

the S^ Septal canopies occur at S2-S3 and S3-S3' junctions. 

Columella consists of 9 tall, slender spines similar in shape to 

the S, trabecular spines. 

DISCUSSION.—Among the four other species of Leptopenus, 

L. solidus is most similar to L. discus, both having massive S2 

trabecular spines, but L. solidus differs in having a lower D:H

40 

ratio and a more solid base, with much smaller and fewer thecal 

pores. 

Keller (1977) reported the diameter of the holotype to be 24 

mm, but the actual diameter (including spines) is only about 

half this size. Likewise, she described the thecal pores to be 

small, with a diameter of 0.1 cm or less, but undoubtedly she 

meant to say 0.1 mm or less. She also stated that the septa were 

pierced with very large pores, but essentially they are solid. 

Finally, she reported five cycles of septa, implying 96 septa, 

whereas there are only about 54 septa (and 68 costae) present in 

the damaged holotype. The development of higher order septa 

appears to lag that of the costae. 

MATERIAL EXAMINED.—No new records.-Previous Records: 

Holotype of L. solidus; Galathea-453,1, ZMC (L. discus 

of Squires, 1965). 

TYPE.—The holotype (Plate \5g,h) is deposited at the IOM. 

Type Locality: Vityaz-5603: 46°22'N, 153°03'E (Kurile 

Trench), 3175-3250 m. 

DISTRIBUTION.—Kurile Trench and Makassar Strait, Indonesia; 

2000-3250 m. 

Letepsammia Yabe and Eguchi, 1932g 

DIAGNOSIS.—Corallum solitary, discoidal, and free. Synapticulothecate; 

marginal shelf present. Costae thin dentate 

ridges, the intercostal spaces much broader than costae and 

penetrated by large pores. Septa also highly porous, with 

complex dentition. Septa alternate in position with costae. 

Septa arranged in typical micrabaciid pattern, having multiple 

bifurcations of the S3; number of septa a function of calicular 

diameter, but 120 is the common adult number. Columella 

spongy. 

TYPE SPECIES.—Stephanophyllia formosissima Moseley, 


Letepsammia formosissima (Moseley, 1876) 

PLATE 15C./ 

Stephanophyllia formosissima Moseley, 1876:561-562; 1881: 201-204.pl. 4: 

fig. 11; pi. 13: figs. 6. 7; pi. 16: figs. 8, 9.—Eguchi. 1934:368; 1938. table 

2.—Utinomi. 1965:249. 

Stephanophyllia superstes Ortmann, 1888:160-161. pi. 6: Fig. 5.—Yabe and 

Eguchi. 1932g:58; 1942b: 112.—Owens, 1986b:487. 

Stephanophyllia (Letepsammia) formosissima.—Yabe and Eguchi, 1932b:443; 

1932g:61-63, pi. 8: figs. 7, 8; 1942b: 138-139.—Eguchi. 1968:C16-17. pi. 

C6: figs. 7-11; pi. C25: figs. 10-13; pi. C27: figs. 2, 3.—Eguchi and 

Miyawaki. 1975: 58. 

Stephanophyllia (Letepsammia) japonica Yabe and Eguchi, 1932b:443 [nomen 

nudum]; 1934b:281, figs. 1-3; 1942b: 139, 156-157.pl. 12: fig. 8. 

Micrabacia japonica.—Omura, 1983:119. 

Stephanophyllia japonica.—Zou. 1988:75, pi. 5: fig. 7. 

Letepsammia formosissima.—Owens, 1986b:486-487.—Cairns, 1989a: 15- 

18, pi. 6: fig. j; pi. 7: figs, g-i; pi. 8: figs, a-d [synonymy].—Caims and 

Parker. 1992:8-9, fig. If.h.—Caims and Keller. 1993:230-231. fig. 3E. 

DESCRIPTION.—Corallum circular, Japanese specimens up 

to 20 mm in diameter, and extremely porous, resulting in a very 


low density corallum. Base slightly convex, composed of thin 

costae 0.10-0.15 mm wide separated by wide intercostal 

spaces 3-4 times that width. Intercostal spaces bridged at 

regular intervals by thin synapticulae, which produce a series of 

large, elliptical pores adjacent to each costa. Corallum white. 

Septa arranged in micrabaciid fashion: S1 independent 

(nonbifurcating) and straight, extending from columella to 

calicular edge, but appearing somewhat isolated because their 

adjacent septa are always inclined away from them. S2 also 

straight (nonbifurcate), and extend from columella to calicular 

edge, but are usually taller than the S1 and are joined by 

adjacent S3 near columella. Each S3 usually bifurcates into 7 

(sometimes 6) septa, resulting in 96 or 84 septa, respectively. 

S3 tallest septa, gradually decreasing in height as they 

repeatedly bifurcate toward calicular edge. All septa highly 

porous. A small marginal shelf occurs peripherally. Columella 

elliptical in shape and papillose. 

DISCUSSION.—Letepsammia formosissima is a commonly 

collected deep-water coral in the Japanese region, representing 

the northern range of a widely distributed Indo-West Pacific 

species. The Japanese specimens are uniformly smaller (

NUMBER 557 41 

DISTRIBUTION.—Japan: Pleistocene of Kakai-jima, Ryukyu 

Islands. Recent: Southeastern Honshu from Sagami Bay to 

the Inland Sea; Bungo Strait; Osumi Shoto, northern Ryukyu 

Islands; western Kyushu; Korea Strait south of Tsushima and 

northeast of Cheju Do, South Korea; off Hong Kong; 77-307 

m. Elsewhere: Philippines, Indonesia, New Zealand, Australia, 

Tasmania, Hawaiian Islands, southwest Indian Ocean; 

97-470 m. 

Rhombopsammia Owens, 1986a 


broad marginal shelf present. Costae thin dentate 

ridges; intercostal spaces broader than costae, penetrated by 

large pores. Septa alternate in position with costae. Septa 

imperforate, numbering 96-144. Columella spongy. 

TYPE SPECIES.—Rhombopsammia squiresi Owens, 1986a, 

by original designation. 

Rhombopsammia niphada Owens, 1986 

PLATES 15/-*, 16e 

Rhombopsammia niphada Owens, 1986a:252-255, figs. 2B, 3A-D.—Cairns, 

1989a: 19-20, text-fig. 2, pi. 9: figs, d-i; pi. 10: figs. a,b [synonymy]. 

DIAGNOSIS.—Discoidal corallum up to 41.6 mm in diameter, 

base usually flat. Costae thin (0.06-0.07 mm) ridges. Intercostal 

regions much wider (about 0.45 mm) than costae, traversed 

by thin synapticulae, which produce a series of pores in each 

elongate space. Septa arranged in typical micrabaciid fashion, 

both costae and septa attaining 144 elements. Marginal shelf 

present but not wide and often damaged. Columella elongate 

and spongy. 

DISCUSSION.—Only two additional specimens are reported 

herein, the larger (TM (KT9202, YT5)) measuring 41.6 mm in 

diameter. R. niphada has been adequately described and 

figured by Owens (1986a) and Caims (1989a); nothing more 

can be added to its characterization from these specimens. This 

species is distinguished from other micrabaciids in the North 

Pacific by having 144 imperforate septa and a relatively large 

calicular diameter. 

MATERIAL EXAMINED.—New Records: Alb-4912,1, CAS 

74999; TM (KT9202, YT5), 1, USNM 92647. Previous 

Records: Type series, USNM; specimen from Misake reported 

by Cairns (1989a). 

TYPES.—The holotype (Plate 15*) and paratypes are all 

deposited at the USNM. Type Locality: Alb-4911: 31°38'N, 

129°19'E (East China Sea off southwestern Kyushu), 715 m. 

DISTRIBUTION.—East China Sea off Kyushu; Osumi Shoto, 

northern Ryukyu Islands; off Misaki, Honshu; 660-783. 

Elsewhere: South China Sea off Philippines; 512-686 m. 

Stephanophyllia Michelin, 1841 


narrow maginal shelf usually present. Costae 

granular, equal to or thicker than intercostal spaces; base thick. 

Septa alternate in position with costae. Septa primarily 

imperforate, being perforate only near base; 96 septa present in 

a mature specimen. Columella solid, compact, and often 

lenticular. 

TYPE SPECIES.—Fungia elegans Bronn, 1837, by original 

designation. 

Stephanophyllia fungulus Alcock, 1902 

PLATE \6a-d,f, g 

Stephanophyllia fungulus Alcock, 1902b: 122; 1902c:40, pi. 5: fig. 35a,b — 

Yabe and Eguchi, 1932b:443; 1932g:58, 60-61. pi. 8: figs. 1-6; pi. 9: figs. 

1-8; 1942b: 138.—Eguchi, 1934:368.—Uchida, 1963:17-21.—Cairns, 

1989a:2l-23, pi. 10: figs, c-k; pi. 11: figs. a,b [synonymy].—Caims and 

Keller. 1993:231. 

DESCRIPTION.—Corallum circular, with a flat base and 

evenly rounded convex calice having a D:H of 1.9-2.1. Largest 

northwestern Pacific specimen (Alb-5311) 12.9 mm in diameter. 

Base quite thick (up to 0.6 mm) and usually upturned at 

calicular edge as much as 1 mm. Costae flat, about 0.15 mm 

wide, and separated by narrow intercostal spaces 70-100 \im 

wide. Each costa bordered on both edges by a row of very small 

(0.02 mm in diameter) granules (Plate 16/). Costae solidly 

fused via rings (about 13 in figured specimen) of massive mural 

synapticulae termed fulturae by Gill (1979) (Plate \6c,d), 

altogether producing a sturdy, robust corallum. Mural fulturae 

closely spaced and parallel in orientation, each about 0.2 mm 

thick but of variable height, increasing in size from center of 

calice to peripheral edge (0.2 to 0.8 mm). Corallum white. 

Septa! arrangement quite conservative, all specimens above 

a calicular diameter of about 6.9 mm having 96 septa arranged 

in typical micrabaciid fashion as illustrated by Caims (1989a, 

text-fig. 3, but having 96 septa, not 98 as I mistakenly indicated 

in the caption). S,_2 straight, extending from columella to 

calicular edge, nonbifurcate, and composed of 20-22 trabeculae, 

each trabeculum projecting about 0.3 mm above septal 

edge as a transversely flattened spine. A pair of S3 originate 

from each S2 very close to the columella: the S3i closest to the 

S1 invariably divide twice to form 3 terminal septa, the S3' 

closest to the S2 divide 3 times to form 4 terminal septa. Septa 

fairly solid, perforate only near base and at points of S3 

bifurcation. All septa structurally reinforced by massive septal 

fulturae, which are irregular in distribution and parallel to 

septal trabeculae. Columella usually consists of a massive 

lamella, lenticular in cross section, and aligned with the 2 

principal septa. Often additional papillae surround the central 

structure. Occasionally, especially in small specimens, the 

columella is formed of a massive fusion of aligned tubercles. 

DISCUSSION.—Two other Recent species of Stephanophyllia 

are known, both occurring in the Indonesian region: S. neglecta 

Boschma, 1923 and S. complicata Moseley, 1876. Stephanophyllia 

fungulus is distinguished by its massive columella and 

thick, upturned base (also see Cairns, 1989a, table 2).

42 

MATERIAL EXAMINED.^WW Records: TM (KT9015, 

BS2), 88, USNM 92652, 22, ORI; TM (9015, HK2), 1, USNM 

92653; TM (KT9015, OK2), 1, USNM 92654; TM (KT9202, 

OS2), 1, USNM 92655; USGS 17633 (Pleistocene of Okinawa), 

1, USNM 88438. Previous Records: 5 syntypes, 

ZMA; Alb-5311 and Alb-5312 (reported by Cairns, 1989a); 

Pleistocene of Ryukyu Islands, 1, USNM 81858 (reported by 

Yabe and Eguchi, 1932b). 

TYPES.—Five syntypes of 5. fungulus are deposited at the 

ZMA (1321). Type Locality: Siboga-100: 6° 11X 120°37.5'E 

(Sulu Archipelago), 450 m. 

DISTRIBUTION.—Pleistocene of Kazusa, Honshu; Okinawa 

and Kakai-shima, Ryukyu Islands. Recent: Tokyo Bay, 

Honshu; Bungo Strait; Van Dienem Strait, Kyushu; East China 

Sea off Goto Retto; Eastern Channel of Korea Strait; Oki Strait, 

Sea of Japan, western Honshu; off Hong Kong; 73-256 m. 

Elsewhere: Philippines, Celebes Sea, Maldives, South Africa, 

Mozambique; 98-635 m (Cairns, 1989a). 

Suborder FAVIINA 

Superfamily FAVIOIDEA 

Family RHIZANGIIDAE 

Culicia Dana, 1846 

DIAGNOSIS.—Corallum colonial (reptoid), consisting of low, 

cylindrical corallites linked together by stolons. Corallites 

epithecate. S1 weakly dentate or lobate; higher cycle septa 

finely dentate. Pali absent; columella rudimentary (papillose). 

Shallow water cryptic azooxanthellates. 

TYPE SPECIES.—Culicia stellata Dana, 1846, by subsequent 

designation (Wells, 1936). 

Culicia japonica Yabe and Eguchi, 1936 

PLATE \la-e 

Culicia japonica Yabe and Eguchi, 1936:167-168, figs. 1-3.—Eguchi, 

1968:C26-27. pi. C9: figs. 1-3.—Kikuchi, 1968:7, pi. 4: fig. 9a,b.—Song, 

1982:133-134, pi. 1: figs. 7-9; 1988:26-27, pi. 1: figs. 7-10; 1991:131.— 

Tribbie and Randall. 1986:108. 

Culitia I sic] japonica.—Yabe and Eguchi, 1942b: 128. 

DIAGNOSIS.—Corallites cylindrical, 3.1-3.7 mm in calicular 

diameter and up to 7 mm in height, although most are only 2-3 

in height. Corallites encrusting and well spaced, their placement 

ranging from directly adjacent to 5-6 mm from one 

another, connected to their parent corallite by a thin, flat stolon. 

Corallites epithecate and white, the epitheca often rising well 

above upper septal margins. Polyps yellowish orange. Septa 

hexamerally arranged in 4 cycles, the last cycle always 

incomplete, corallites usually having 34-44 septa. S, composed 

of a tall, broad septal lobe bordered internally by 2 or 3 

smaller lobes. S2.3 about equal in size and composed of 3 or 4 

tall, slender, bluntly tipped septal lobes, their inner edges fused 

near columella. S4 rudimentary and irregular in occurrence. 


Columella circular and papillose, the elements indistinguishable 

from the Su3 inner septal lobes. 

DISCUSSION.—The differences among C. japonica, C. 

truncata Dana, 1846, and C. stellata Dana, 1846, are slight, if 

any. A revision of the 15-18 nominal species of this genus is 

badly needed. 

MATERIAL EXAMINED.—New Records: Alb-4925, 1 colony, 

USNM 92657; TM (KT9015, CB2-2), 1 colony, ORI; off 

Sesoko Island, Okinawa, 1, USNM 86826; off Horseshoe 

Cliffs, Onna Village, Okinawa, 1, USNM 88385; off Ogasawara 

Island, Bonin Islands, 1 colony, USNM 92656. 

Reference Specimens: Types of C. stellata and C. truncata, 

USNM. 

TYPES.—The holotype of C. japonica is deposited at the 

TIUS (59328). Type Locality: Off Owasi, Mie Prefecture, 

Honshu, 100 m. 

DISTRIBUTION.—Southeastern Honshu from Sagami Bay to 

Owasi; Tokara Retto and Okinawa, Ryukyu Islands; off 

Amakusa Island, Kyushu; Eastern Channel, Korea Strait; off 

southern Cheju Do, Korea; western Channel, Korea Strait off 

southeastern Korea; Bonin Islands; 5-100 m. 

Oulangia Milne Edwards and Haime, 1848c 

DIAGNOSIS.—Corallum colonial (reptoid), corallites connected 

by thin stolons. Corallites shaped as low, wide 

cylinders. Theca costate. Septa exsert, usually in five cycles; 

inner septal edges finely dentate to laciniate. Pali absent; 

columella papillose, merging with lower, inner septal processes. 

Shallow water azooxanthellates. 

TYPE SPECIES.—Oulangia stokesiana Milne Edwards and 

Haime, 1848d, by subsequent designation (Milne Edwards and 

Haime, 185Oa:xlv). 

Oulangia stokesiana miltoni Yabe and Eguchi, 1932 

PLATES 17/, 4\a,b 

Oulangia stokesiana var. miltoni Yabe and Eguchi, I932e:29-31, pi. 4: fip. 

1-9; 1933:83-85, figs. 1,2. 

Oulangia stokesiana miltoni.—Eguchi, 1968:C27, pi. C4: figs. 2, 3 [color].— 

Hamada, 1969:253, pi. 2: fig. 6a-c.—Song, 1991:132, pi. 1: fig. 2; pi. 2: 

figs. 2, 3. 

?0ulangia cf. O. stokesiana.—Tribbie and Randall, 1986:158. 

DIAGNOSIS (based primarily on syntypes).—Corallites short, 

squat cylinders, up to 10 mm in GCD and 4 mm in height, but 

invariably wider than tall. Propagation by asexual budding 

from a thin layer of common basal coenosteum, the connection 

among polyps usually lost or obscured after budding. Base 

polycyclic, up to three concentric thecal walls visible in worn 

corallites. Costae broad, flat to slightly convex, and equal in 

width (about 0.5 mm), separated by narrow (0.15 mm), shallow 

intercostal striae, which become wider near calice as costae 

become correspondingly narrower. Costae finely granular, 2 or 

3 granules occurring across a costal width. Corallum blackish 

brown. Septa hexamerally arranged in 5 cycles according to the

NUMBER 557 43 

formula: S1_2>S3>S4>S5. Last (fifth) cycle rarely present and 

never complete, the largest number of septa reported being 68, 

corallites with only 48 septa being most common. S^_2 only 

slightly exsert, their inner edges attaining the columella, their 

upper edges bearing 5-7 coarse teeth. S3 shortest septa, bearing 

3 or 4 small, blunt teeth. Each S3 flanked by a pair of long S4, 

each bearing 7 or 8 blunt teeth and meeting in a V-shaped 

junction near inner edge of common S3 and continuing to 

columella to meet the inner edge of the S2. Fossa of variable 

depth, but usually quite shallow. Columella papillose, consisting 

of papillae indistinguishable from teeth on inner edges of 

S.,_2, and S4. 

DISCUSSION.—Oulangia stokesiana miltoni is distinguished 

from the nominate subspecies and two other Recent species (O. 

bradleyi Verrill, 1866, and O. cyathiformis Chevalier, 1971) by 

having a relatively small corallum with correspondingly fewer 

septa: 48-66 instead of 96. Previously known only from off 

northeastern Honshu, and South Korea, this subspecies is 

herein reported from the Sea of Japan in the Oki Channel, off 

southwestern Honshu at 100-135 m, the greatest recorded 

depth for this subspecies. Like Letepsammia formosissima, 

specimens from the northern range (off Japan) of this 

widespread Indo-West Pacific species have a smaller corallum, 

and thus has been distinguished as a subspecies. 


CB-6), 3 corallites, USNM 92646. Previous Records: 75 

syntypes, TIUS. 

TYPES.—Approximately 150 syntypes (Plate 41 a,b) are 

deposited at the TIUS (41051). Type Locality: Shiogamamachi 

(east of Teizan Canal), Matsushima Bay, near Sendai, 

Honshu, depth unknown. 

DISTRIBUTION.—Pleistocene of Sagami Bay region. Recent: 

Northeastern coast of Honshu from Shiogama to 

Sagami Bay; ?Miyake-jima; Oki Channel, off southwestern 

Honshu; coast of South Korea from Pusan to Tokchokto, 

Yellow Sea; 0-135 m. 


Family OCULINIDAE 

Madrepora Linnaeus, 1758 

Madrepora oculata Linnaeus, 1758 


Cyathelia Milne Edwards and Haime, 1849 

DIAGNOSIS.—Colonies arborescent, formed by regular, 

alternate, extratentacular budding. Coenosteum dense and 

faintly costate. Well-developed pali occur before first three 

septal cycles; columella papillose. 

TYPE SPECIES.—Madrepora axillaris Ellis and Solander, 


Cyathelia axillaris (Ellis and Solander, 1786) 

PLATE \%a-c 

Madrepora axillaris Ellis and Solander. 1786:153, pi. 13: fig. 5. 

Cyathelia axillaris.—Milne Edwards and Haime. 185Ob:72-73. 

Cyathohetia axillaris.—Milne Edwards and Haime, 1857:110-111.—Duncan, 

1876:438.—Alcock, 1898:28.—Bedot, 1907:145, pi. 15: fig. 103. 

Cyathelia cf. axillaris.—Eguchi, 1942:138 (142), pi. 6: figs. 2, 3 [synonymy]. 

Cyathelia axillaris.—Eguchi, 1968:C28, pi. C20: figs. 5-7; pi. C24: figs. 4. 5 

[synonymy].—Kikuchi, 1968:8, pi. 5: fig. 1. 

DESCRIPTION.—Coralla sparsely branched, resulting in 

small, robust, bushy colonies, the largest known about 7.5 cm 

in height, supporting approximately 100 corallites (Eguchi, 

1968, pi. C24: fig. 4). Branching is essentially sympodial, but 

two buds often originate on opposite sides of a terminal 

corallite, the parent corallite ultimately becoming immersed in 

thick coenosteum within the branch axil. Corallites circular 

when small, often becoming elliptical or even medially 

constricted if located at a branch axil. Corallites relatively 

large, up to 11 mm in GCD. Branch coenosteum dense, 

granular, light brown to tan in color, and usually faintly costate. 

Corallites often pigmented a darker shade of brown. 

Septa usually hexamerally arranged in 4 cycles; however, 

small corallites (i.e., 10 mm GCD) sometimes have an extra (13th) half-system for 

a total of 52 septa. Septal formula: S1_2>S3>S4. Su2 up to 1.5 

mm exsert, quite thick, and have straight inner edges reaching 

about one-fourth distance into fossa. S3 equally exsert but only 

about two-thirds as wide as S:_2- S4 less exsert and about 80% 

width of the S3. Twelve thick pali, each about 1.5 mm wide, 

form a palar crown encircling columella. Another crown of 12 

P3, each palus of equal width but slightly thicker than P1-2, 

occurs slightly recessed from the columella and slightly higher 

in fossa. Pali and septal faces highly granular. Columella 

papillose. 

DISCUSSION.—Cyathelia is a montypic genus distinguished 

from other oculinid genera by having two crowns of pali before 

its first three septal cycles. It has a distinctive colony shape, 

easily recognized by its sparse branching and large corallites in 

relation to its branch diameter. 

MATERIAL EXAMINED.—New Records: Alb-4944, 1 colony, 

USNM 92660; TM (KT9015, CB2-2), 1 colony, ORI; TM 

(KT9015, CB1-2), 2 colonies, USNM 92661; TM (KT9202, 

YS1), 1, USNM 92662; TM (KT9202, YT2), 1, USNM 92663; 

off Yenoshima, Honshu, 4 colonies, USNM 92664; Monoiso, 

Misaki, Sagami Bay, 1 colony, USNM 92665; Ashima, Sagami 

Bay, 1 colony, USNM 92666. Previous Records: 

Challenger-\96, 1 colony, BM 1880.11.25.88 (reported by 

Moseley, 1881). 

TYPES.— The types of M. axillaris have not been traced, 

although Ellis and Solander's (1786) original illustration of the 

species is diagnostic for this distinctive species. Type Locality: 

Eastern Indian Ocean, depth unknown.

44 

DISTRIBUTION.—Japan: Sagami Bay to Izu Bay, Honshu; 

Osumi Shoto, northern Ryukyu Islands; Kagoshima Bay and 

off Amakusa Island, Kyushu; off Mishima, Eastern Channel, 

Korea Strait; 15-366 m. Elsewhere: Moluccas; Arabian Sea 

and Bay of Bengal; 161-71509 m. 

Moseley's (1881) specimen from 1509 m is correctly 

identified but clearly was dead when dredged. It seems unlikely 

that C. axillaris would occur at such a great depth. Likewise, 

although the identity of Alcock's (1898) record from 810 m has 

not been verified, it would also seem to be too deep for this 

species based on all other known records. 

Family ANTHEMIPHYLLIIDAE 

Anthemiphyllia Pourtales, 1878 

DIAGNOSIS.—Solitary, patellate or discoidal, and free. 

Septotheca thick: porcellaneous or costate. Septal edges lobate 

to laciniate. Pali absent; colurnella papillose. 

TYPE SPECIES.—Anthemiphyllia patera Pourtales, 1878, by 

monotypy. 

Anthemiphyllia dentata (Alcock, 1902) 

PLATE \id-f 

Discotrochus dentatus Alcock, 1902a: 104; 19O2c:27, pi. 4: figs. 26, 26a.— 

Yabe and Eguchi, 1932b:443; 1937a: 143-145, pi. 20: fig. 15a-c. 

?Discothrochus [sic] sp.—Yabe and Eguchi, 1937:145-146. 

Anthemiphyllia dentata.—Yabe and Eguchi, 1942b: 128-129.—Eguchi, 

1968:C29-30, pi. C6: figs. 12-21.—Not Cairns, 1984:11 [= undescribed 

species].—Best and Hoeksema, 1987:398-399, fig. 9a-c.—Cairns and 

Parker, 1992:16-17. fig. 4e,f [synonymy].—Caims and Keller, 1993:233, 

fig. 3E. 

Anthemiphyllia dentatus.—Eguchi. 1965:285. 2 figs. 

?Anthemiphyllia sp..—Eguchi, 1968:C30-31, pi. C9: figs. 13. 14. 

Deltocyathus andamanicus.—Keller, 1982:52 [in part: pi. 1 [= 4]: figs. 3, 4). 

DISCUSSION.—Anthemiphyllia dentata is a widespread species 

known from the western Indian Ocean to off Japan, the 

latter being the northernmost range for the species. It has been 

widely reported (Yabe and Eguchi, 1937, 1942b) from off 

southern Japan from moderate depths (50-499 m). The three 

specimens reported herein (1 fossil, 1 badly preserved, and 1 

juvenile) do not add to the previous descriptions of the species 

given by Alcock (1902a) and Cairns and Parker (1992), and the 

fine illustrations published by Yabe and Eguchi (1937) and 

Eguchi (1968). Anthemiphyllia dentata is distinguished from 

the three other Recent species in the genus by attaining a fifth 

cycle of septa at a relatively small calicular diameter, ultimately 

attaining a large (up to 27 mm) calicular diameter, and in 

having finely dentate to laciniate septal margins. 

MATERIAL EXAMINED.—New Records: Pleistocene: 

USGS 17455 (south of Tomari, Okinawa, Shimijin marl), 2 

fragments, USNM 88435; Recent: Alb-5094, 1, USNM 

92667; TM (KT9202, OS3), 1, ORI. 

TYPES.—Seven syntypes of D. dentatus are deposited at the 

ZMA (Coel. 716-718) (van Soest, 1979). Type Local- 


ity: Sibof>a stations 95, 98, 100 (Sulu Sea), 350-522 m. 

DISTRIBUTION.—Pleistocene of Ryukyu Islands off Kikaijima 

and Okinawa. Recent: Japan: southeastern Honshu from 

Sagami Bay to Kii Strait; off eastern Shikoku and southern 

Kyushu; Sea of Japan, Oki Strait; 50-499 m. Elsewhere: 

Sulu Sea (Alcock, 1902a), Philippines (Keller, 1982), 

Banda Sea (Best and Hoeksema, 1987), southern Australia 

(Cairns and Parker, 1992), Maldives, Arabian Sea, Saya de 

Malha; 75-560 m. 

Anthemiphyllia frustum, sp. nov. 

PLATES l&g-i, \9a,b 

DESCRIPTION.—Corallum tympanoid or frustum-shaped, 

with a flat base and a low, slightly inwardly curved theca 

resulting in a basal diameter slightly larger than that of calice. 

Holotype 9.94 mm in basal diameter, 8.96 mm in calicular 

diameter, and 4.71 mm in height. Costae equal in width 

(0.5-0.7 mm at edge of base), relatively flat, and separated by 

quite narrow (0.02-0.05 mm), shallow intercostal striae. 

Costae finely granular, 5 or 6 small (0.05 mm in diameter) 

rounded granules occurring across the width of a costae near 

corallum edge. Corallum white or reddish brown. 


in all specimens examined. S1 bear 5 or 6 massive triangular 

teeth, each tooth up to 1 mm in height and 0.9 mm in width, the 

innermost tooth being directly adjacent to columella. S2 similar 

to S1 in construction, but their teeth not quite as tall. Both S1 

and S2 quite robust and closely spaced, measuring about 0.5 

mm in thickness at calicular edge and separated by only about 

0.1 mm. S3 support 4 or 5 tall, triangular teeth, each tooth up to 

0.5 mm wide, but not quite as tall as those of the S-i_2- S4 bear 

5 or 6 triangular teeth, which decrease in width toward 

columella. S3_4 about 0.35 mm in thickness. Each pair of S4 

within a half-system unites before its adjacent S3 near the 

columella in a common triangular septal tooth about 0.2 mm 

wide. Orientation of faces of septal teeth is parallel to that of 

septal faces. All septal faces coarsely granular, except for tips 

of large septal teeth, which are smooth. Columella papillose, 

composed of 10-15 massive, rounded, finely granular papillae. 

DISCUSSION.—Anthemiphyllia frustum is readily distinguished 

from A. patera Pourtales, 1878; A. dentata (Alcock, 

1902); and A. pacifica Vaughan, 1907, by: its frustum-shaped 

corallum (the other three species are bowl-shaped); its 

limitation to 48 septa (the other species have some or all of the 

fifth cycle); and its massive septal teeth, the faces of which are 

parallel to the septal plane, not perpendicular as in the other 

three species. Anthemiphyllia frustum is most similar to A. 

pacifica Vaughan, 1907, particularly in having the same 

number of septa, the same orientation and number of septal 

teeth per septum, and approximately the same corallum size. 

Anthemiphyllia pacifica differs significantly, however, in 

corallum shape (bowl-shaped), having much smaller S4 in 

relation to its S3, having broader intercostal and interseptal

NUMBER 557 45 

spaces, and in having clavate, not triangular, septal teeth. 

ETYMOLOGY.—The species name frustum (Latin frustum, 

meaning "a bit," or "part"), refers to the corallum shape. In 

geometry, a frustum is the solid figure formed when the top of 

a cone is cut off by a plane parallel to the base, which is the 

shape of the corallum of this species. 

MATERIAL EXAMINED (Types).—Holotype: TM (KT9202, 

OS2), USNM 92668 (Plate \9a,b). Paratypes: TM (KT9202, 

OS2), 4, USNM 92669 (Plate \Sg-i), 1, ORI. Type Locality: 

3O°59TM, 13O°32"E (Osumi Strait, southern Kyushu), 

237-241 m. 


Suborder CARYOPHYLLIINA 

Superfamily CARYOPHYLLIOIDEA 


Family CARYOPHYLLIIDAE 

Caryophyllia Lamarck, 1816 

Subgenus Caryophyllia (Caryophyllia) Lamarck, 1816 


Caryophyllia (C.) japonica Marenzeller, 1888 

PLATES 4/, 19c-/ 

Caryophyllia (C. japonica Marenzeller, 1888b: 16.—Yabe and Eguchi, 

1941b:102; 1942b:119, pi. 10: figs. 1-3.—Eguchi, 1965:285, 2 figs.; 

1968:C31-32, pi. Cl 1: figs. 4-6, 10-29; pi. C23: figs. 7-9; pi. C25: figs. 5, 

6; pi. C29: figs. 6, 7.—?Kikuchi, 1968:8, pi. 5: fig. 8.—Eguchi and 

Miyawaki, 1975:56.—Song, 1982:134, pi. 2: figs. 3-5; 1988:27, pi. 3: figs. 

9-11; 1991:132-133. 

Caryophyllia ephyala.—Yabe and Eguchi, 1932a:388, 389. 

Caryophyllia arcuata.—Yabe and Eguchi, 1936:167.—Eguchi, 1938, table 2. 

Caryophyllia alaskensis.—Keller, 1981a:21 [in part: Vifyaz-2078, 3353, 5592, 

5640], pi. 2: fig. 1. 

Caryophyllia ambrosia.—Keller, 198la: 15 [in part: V/ryaz-5638]. 

DESCRIPTION.—Corallum robust, ceratoid to trochoid, and 

firmly attached by a relatively slender pedicel: PD:GCD = 

0.25-0.39. Calice circular to elliptical. Largest corallum 

examined (Alb-4982) 17.8 x 16.0 mm in calicular diameter and 

18.4 mm in height, with a pedicel diameter of only 6.7 mm. 

Theca thick. Cl-3 sometimes slightly ridged near calice; 

otherwise, costae are flat with low, rounded granules or 

porcellaneous in texture. Corallum white. 


to the formula: S1_2>S3>S4. S^ moderately exsert (1.5-2.1 

mm) and have straight, vertical inner edges that merge with the 

columella low in fossa. S3 70%-75% width of Su2, less exsert 

(about 1 mm), and have sinuous inner edges. S4 less exsert and 

slightly less wide than S3. A crown of 12 slender, lamellar but 

sinuous pali occurs before the S3, each palus 1.5-2.0 mm in 

width. 

Fossa of moderate depth. Columella well developed, 

composed of a large elliptical field of 10-20 fused, twisted 

elements. The previous description was based primarily on 

specimens from Alb-4982 and the two syntypes. 

DISCUSSION.—Three species—Caryophillia japonica, C. 

alaskensis and C. arnoldi—form a closely related group that 

might well be considered as geographic subspecies of one 

species: C. japonica, known from Japan to the Commander 

Islands; C. alaskensis, from the Commander Islands (at 

shallower depths) to British Columbia; and C. arnoldi, from 

Prince William Sound, Alaska to the California Channel 

Islands. Caryophyllia japonica differs from the geographically 

adjacent C. alaskensis in much the same ways that C. arnoldi 

differs from C. alaskensis. Caryophyllia japonica has a more 

robust corallum; consistently 48, more highly exsert septa; and 

S4 that are only slightly less wide than the S3. Caryophyllia 

japonica is almost indistinguishable from its trans-Pacific 

cognate, C. arnoldi, differing only in having a narrower 

pedicel. 


USNM 82158, 2, ORI; Alb-5088, 1, USNM 92676; Alb-5093, 

1, USNM 92677; Academik Keldish-23\2, 4, IOM. Previous 

Records: 2 syntypes of C. japonica; specimens reported as C. 

alaskensis by Keller (1981a) from: Vityaz-2QH% (IOM), 3353 

(USNM 92678, Plate 19c,/), and 5640 (IOM); specimen 

reported as C. ambrosia by Keller (1981a) from V/fyaz-5638, 

IOM (Plate 40. 

TYPES.—Two syntypes (Plate \9d,g,i) of C. japonica are 

deposited at the NMW (8168). Type Locality: "Enosima," 

Japan, depth unknown. Although there are several Enosima's 

(= Enoshima) listed in a Japanese gazetteer, the most likely is 

the one in Sagami Bay. 

DISTRIBUTION.—Throughout Japanese waters from Ishikari 

Bay, Hokkaido to southern Kyushu, including both Pacific and 

Sea of Japan coasts; off Cheju Island, Korea Strait and Sea of 

Japan off South Korea; Pacific coast of Kurile Islands; 

Commander Islands, Bering Sea; 77-1680 m, with a tendency 

to occur deeper in the northern range. 

Caryophyllia (C.) alaskensis Vaughan, 1941 


Caryophyllia (C.) sp. cf. C. scobinosa Alcock, 1902 

PLATES 19/-/, 20a.* 

Caryophyllia scobinosa Alcock, 1902a:90; 19O2c:8, pi. 1: figs. 2. 2a.—Yabe 

and Eguchi. 1942b:119-120 [in part; not pi. 10: fig. 5].—Utinomi, 1965: 

254.—Eguchi, 1965:285. fig.—Cairns and Keller, 1993:235 [synonymy]. 

Caryophyllia cf. scobinosa.—Utinomi, 1956:42. 

DESCRIPTION OF SPECIMENS FROM TM (KT7911, OT4).— 

Corallum ceratoid, free, and curved about 45. Corallum 10.4 

mm in calicular diameter, 17.9 mm in height, and 1.7 mm in 

pedicel diameter. Costae flat and granular, separated by narrow 

intercostal striae. Corallum white. Septa hexamerally arranged

46 

in 4 complete cycles according to the formula: S1_2>S3»S4. 

S^ only slightly exsert and have straight, vertical inner edges 

that do not attain the columella. S3 about three-fourths width of 

S^ and have slightly sinuous inner edges. S4 rudimentary, 

only about one-third width of S3, and have irregular to lacerate 

inner edges. Twelve broad lamellar pali occur in a crown before 

the S3, each palus almost as wide as its adjacent S3 and having 

only moderately sinuous inner edges. Fossa of moderate depth, 

containing a columella of 5 slender, twisted elements. 

DISCUSSION.—Although the specimens reported herein, as 

well as those reported by Yabe and Eguchi (1942b) and 

Utinomi (1965), are similar to typical C. scobinosa, they are 

consistently different in several characters. The Japanese 

specimens have much smaller S4 in relation to their S3; in 

typical C. scobinosa the S4 are equal to or wider than the S3. 

Also, the Japanese specimens have less exsert septa and a 

narrower, less curved corallum. They may represent an extreme 

variation for the species at its northern limit, or, perhaps, a 

separate species. 


74994; TM (KT7911, OT4), 3, USNM 92680. Previous 

Records: Syntypes of C. scobinosa, ZMA. 

TYPES.—Six syntypes (Plate 20a,b) of C. scobinosa are 

deposited at the ZMA (Coel. 574, 575Xsee van Soest, 1979). 

Type Localities: Siboga stations 45 and 102: Celebes and 

Sulu Seas, 535-794 m. 

DISTRIBUTION.—Japan: Suruga Bay and Kii Strait, Honshu; 

off eastern Shikoku; East China Sea off southwestern Kyushu; 

Tokara Retto, northern Ryukyu Islands; 119-805 m. Elsewhere 

(as C. scobinosa): Philippines, Sulu Sea, Celebes, 

southwestern Indian Ocean; 535-960 m. 

Caryophyllia (C.) quadragenaria Alcock, 1902 

PLATES 20C -h,4\c4 

Caryophyllia quadragenaria Alcock, 1902a:9I-92; 1902c:10, pi. 1: figs. 4, 

4a.—Cairns, 1991a:12. 

Caryophyllia scobinosa.—Yabe and Eguchi, 1942b: 119 [in part: pi. 10: fig. 

5a,b]. 

Caryophyllia scobinosa decapali Yabe and Eguchi, 1942b: 120, 149, pi. 10: 

figs. 6. 7.—Eguchi. 1968:C33-34.—Eguchi and Miyawaki, 1975:56 — 

Cairns. 1991a:12. 

Caryophyllia decapali.—Grygier, 1983:420.—Zibrowius and Grygier, 

1985:120. figs. 10, 11. 

DESCRIPTION.—Corallum ceratoid to subcylindrical, straight 

to slightly curved, and attached by a rather narrow pedicel 

(PD:GCD = 0.16-0.33). Largest specimen examined (paratype 

of C. scobinosa decapali, Soyo Maru-222) 12.2 x 10.3 mm in 

calicular diameter and 21.8 mm in height, with a slender 

pedicel diameter of 1.39 mm. Costae equal in width (about 0.5 

mm), flat, and covered with large, low, rounded granules, 2-3 

across the width of a costa. Intercostal striae shallow and 

narrow. Corallum white to light brown. Calicular margin 

serrate, each primary septum along with its fused adjacent pair 

of tertiary septa projecting as a thecal extension. 


Septa decamerally arranged in 3 size classes: 10:10:20 (40 

septa). Primary septa highly exsert (up to 1.8 mm) and have 

vertical, slightly sinuous inner edges that do not reach the 

columella. Secondary septa least exsert (about 0.7 mm), about 

three-quarters width of primaries, and have extremely sinuous 

inner edges. Tertiary septa about 1.0 mm exsert and have 

moderately sinuous inner edges, the upper outer edge of each 

S3 fused to its adjacent primary septum. Width of tertiary septa 

variable, usually equal to or wider than the secondaries, but in 

some specimens only one-third to one-half width of secondaries. 

A crown of 10 broad (1.3 mm wide), very sinuous P2 

tightly encircle the elongate columella, the latter composed of 

4-8 broad, twisted elements. 

DISCUSSION.—Among the approximately 60 species of 

Recent Caryophyllia, only seven have decameral septal 

symmetry (Cairns, 1991a, table 3). In their original diagnosis of 

C. scobinosa decapali, Yabe and Eguchi (1942b) noted the 

extreme resemblance of their Japanese subspecies to C. 

quadragenaria, according to them the latter differing only in its 

attachment by a "cylindrical stalk." In fact, both the Japanese 

C. decapali and C. quadragenaria are attached by narrow 

pedicels, and comparison of syntypes of C. quadragenaria to 

Japanese specimens show no differences, and therefore C. 

decapali is synonymized with C. quadragenaria. 

The character of the relative widths of the last (tertiaries) and 

penultimate cycle (secondaries) of septa can usually be used as 

a species level character, but in C. quadragenaria the tertiaries 

may be wider or considerably less wide than the secondaries. 

For instance, the smallest and largest syntypes of C. quadragenaria 

(Siboga-90, 289) have tertiaries larger than secondaries, 

whereas the tertiaries of the syntype from Siboga-251 are only 

one-third the width of a secondary. And, in the lot of 8 

specimens from TM (KT9202, OS2), most specimens have 

broad tertiaries, but several have tertiaries less broad than the 

secondaries. 

The specimen from Soyo Maru-220, identified by Yabe and 

Eguchi (1942b) as C. scobinosa, is identical to specimens in the 

type series of C. decapali, but has 48 septa and 12 pali. It is 

either an aberrant specimen or indicative that the species may 

have either 10 or 12 pali (40 or 48 septa). 


USNM 92670; TM (KT7811, OT6), 1, USNM 92671; TM 

(KT9015, CB1-2), 4, ORI; TM (KT9202, OS2), 8, USNM 

92672; TM (KT9309, AM8), 1, USNM 93162,2, ORI; Sagami 

Bay, 110 m, 11 June 1914, Mortensen's Pacific Expedition, 1, 

ZMC; 32°10'N, 128°20'E, 183 m, 23 May 1898, 1, ZMC. 

Previous Records: Two syntypes of C. quadragenaria 

{Siboga-90, 251); holotype and 6 paratypes of C. scobinosa 

decapali, Soyo Maru-222, TIUS; C. scobinosa of Yabe and 

Eguchi (1942b), Soyo Maru-220, TIUS 53634. 

TYPES.—Two of the three syntypes of C. quadragenaria are 

deposited at the ZMA: Siboga-90 (Coel. 5534) and Siboga-251 

(Coel. 5529, Plate 20d,g). The third syntype from Siboga-2%9 

is missing (Van Soest, 1979). The missing syntype was the

NUMBER 557 47 

largest and figured specimen. Type Localities: Indonesia 

(Makassar Strait, Banda and Timor seas), 54-281 m. 

The holotype of C. scobinosa decapali (53640), as limited 

by Yabe and Eguchi by the designation of a type locality, and 

paratypes are deposited at the TIUS (Plate 4\c,d). Type 

Locality: Soyo Maru-210: 33°29'N, HS^'E (Kii Strait, off 

southeastern Honshu), 165 m. 

DISTRIBUTION.—Japan: Sagami and Suruga Bays, Honshu; 

Kii Strait; off Shikoku; Osumi Shoto, northern Ryukyu Islands; 

East China Sea off Danjo Gunto; Eastern Channel of Korea 

Strait off Mishima; 70-422 m. Elsewhere: Makassar Strait, 

Banda and Timor Seas; 54-281 m. 

Caryophyllia (C.) rugosa Moseley, 1881 

PLATES 20/, 21a 

Caryophyllia rugosa Moseley, 1881:141-143, pi. 1: fig. 8.—Wells, 1954:469, 

pi. 177: figs. 5, 6.—Cairns. 1984:11-13, pi. 2: figs. A.B; pi. 4: fig. I 

[synonymy].—Cairns and Keller, 1993:236, fig. 3i. 

Caryophyllia paraoctopali Yabe and Eguchi, 1942b:120. 150, pi. 10: fig. 12. 

DESCRIPTION.—Corallum ceratoid to cylindrical, often 

firmly attached to a bivalve or barnacle through a broad 

pedicel. Largest specimen examined (ZMC) 8.5 x 7.5 mm in 

calicular diameter and 14.3 mm in height. Costae equal in 

width (about 0.6 mm wide in large coralla), flat, and covered 

with fine transverse rugae (Plate 21a). Corallum white. 

Septa of most coralla (i.e., calicular diameter < 5.5 mm) 

octamerally arranged in 3 cycles: 8:8:16 (= 32 septa). Primary 

septa only slightly exsert (about 0.8 mm), extend about 

three-quarters distance to columella, and have very sinuous 

inner edges. Secondary septa less exsert (about 0.5 mm), about 

three-quarters width of a primary, and also have very sinuous 

inner edges. Tertiary septa about 0.4 mm exsert, three-quarters 

width of a secondary, and have moderately sinuous inner edges. 

A crown of 8 slender, quite sinuous pali occur before the 

secondary septa. In larger coralla (i.e., calicular diameter > 5.5 

mm), additional septa occur but octameral symmetry is 

maintained. In these larger coraila, pairs of tertiary septa within 

various sectors are accelerated to an equivalent width of a 

secondary and are flanked by smaller quaternary septa. In such 

a sector, a palus occurs before each tertiary septa, not the 

secondary, leading to coralla with 52 septa and 9 pali, or 56 

septa and 10 pali. If only 1 tertiary septum is enlarged within a 

sector, the palus remains single before the secondary septum. 

Fossa quite shallow, containing 3-6 slender, twisted elements. 

DISCUSSION.—There are four species of octamerally symmetrical 

Caryophyllia with 32 septa (Cairns, 1991a); C. rugosa 

is distinguished from the other three by having transversely 

sculptured costae and a very small corallum. In their 

description of C. paraoctopali, Yabe and Eguchi (1942b) 

distinguished it from C. rugosa by its having octameral 

symmetry; however, C. rugosa also has octameral symmetry 

and is clearly the senior synonym. 


USNM 92673; TM (KT9202, YS1), 3, USNM 92674; TM 

(KT9202, YS2), 1, ORI; 32°15'N, 128°20'E, 181 m, 17 April 

1926, 30 specimens, ZMC; 32°21'N, 128°49'E, 179-291 m, 2 

specimens, ZMC. 

TYPES.—The syntypes of C. rugosa are deposited at the BM. 

Type Localities: Challenger stations 192 and 201 (Indonesia 

and Philippine Islands), 187-230 m. 

Four syntypes of C. paraoctopali are deposited at the TIUS 

(53645-53648). Type Locality: "Pacific coast of Honshu, 

Shikoku, and Kyushu," 71-183 m. 

DISTRIBUTION.—Japan: Suruga Bay, Honshu; off southeastern 

Kyushu and northern Ryukyu Islands (Colnett Strait and 

Tokara Retto); East China Sea off Danjo Gunto; 71-240 m. 

Elsewhere: Off Taiwan, Philippines, Ceram Sea, Bikini, 

Hawaiian Islands, and southwest Indian Ocean; 100-439 m. 

Caryophyllia (C.) jogashimaensis Eguchi, 1968 

PLATE21A,C 

Caryophyllia jogashimaensis Eguchi, 1968:C33, pi. C18: figs. 4-8.—Cairns, 

1991a:12. 

Description of Specimen from TM (KT9202, YT2).— 

Corallum trochoid: 12.1 x 9.7 mm in calicular diameter, 15.7 

mm in height, and 4.7 mm in pedicel diameter. Theca worn and 

heavily encrusted with serpulids and foraminifera; however, 

theca near base appears to bear fine transverse rugae. Upper 

outer (near theca) septal edges light red in color. 

Septa arranged in 4 size classes accordingly: 9:9:18:34 (= 70 

septa), 1 of the 18 half-sectors missing its pair of quaternaries. 

Primary septa moderately exsert (about 1.6 mm), extend about 

three-quarters distance to columella, and have slightly sinuous 

inner edges. Secondary septa less exsert (about 0.9 mm), about 

the same width, and also have slightly sinuous inner edges. 

Tertiary septa least exsert (about 0.5 mm), about three-quarters 

width of a secondary, and have very sinuous inner edges. 

Quaternary septa slightly more exsert than tertiary septa 

because they are fused at calicular edge to their adjacent S1 and 

S2 in short, rectangular projections. Quaternaries quite broad, 

almost as wide as tertiaries, and have slightly sinuous inner 

edges. A crown of 17 well-developed (1.2 mm wide), highly 

granular, sinuous pali occur before the tertiary septa. Fossa of 

moderate depth, containing an elongate columella consisting of 

4 or 5 twisted and partially fused elements. 

DISCUSSION.—Eguchi (1968) described the larger (GCD = 

19-20 mm) syntype (1968, pi. C18: figs. 4,5) of this species as 

having 72 septa arranged 9:9:18:36 and 18 pali in 18 

half-sectors; however, it easily can be seen from his illustration 

that there are either 17 or 19 half-sectors (17:17:34:4 or 

19:19:34, respectively) and 19 pali, two end half-sectors being 

in the process of expanding. The smaller syntype (pi. C18: figs. 

6-8; GCD = 14 mm) appears to have 15 half-sectors arranged: 

15:15:30 (60 septa), with 1 palus per half-sector. The number of 

sectors, septa, and pali (i.e., 15-19 half-sectors, 60-72 septa,

48 

and 15-19 pali) thus appears to a function of size, additional 

sectors and pali first forming near the end-sectors. 

MATERIAL EXAMINED.—TM (KT9202, YT2), 1, USNM 

93675, 1, ORI. Previous Records: The type specimens were 

unavailable for study. 

TYPES.—Two syntypes are presumed to be deposited at the 

Biological Laboratory of the Imperial Household, Tokyo 

(#852). Type Locality: Off Jogashima, 1.4 km west of the 

Lighthouse, Sagami Bay, 52-56 m. 

DISTRIBUTION.—Sagami Bay; Colnett Strait, Osumi Shoto, 

northern Ryukyu Islands; 52-98 m. 

Caryophyllia (C.) ambrosia ambrosia Alcock, 1898 

PLATE 2\d-h 

?Caryophyllia scillaeomorpha Alcock. 1394:186; 1898:13. pi. 1: figs. 3, 3a. 

Caryophyllia ambrosia Alcock, 1898:12, pi. 1: Tigs. 1, la.—Zibrowius, 

1980:63-65, pi. 25: figs. A-K [synonymy].—Keller, 1981a:15-16 [in part: 

Akademik Kurchatov station 441 ]. 

Caryophyllia cf. alcocki.—Yabe and Eguchi, 1942b: 120, pi. 10: fig. 8. 

"Unidentified solitary coral."—Okutani, 1969:12, pi. 1: fig. 7. 

Caryophyllia ambrosia ambrosia.—Cairns, 1979:59.—Cairns and Keller, 

1993:234, fig. 3H. 

DESCRIPTION.—Corallum cornute and free, pedicel invariably 

curved about 90° and terminating in a narrow point or 

worn nub. Largest specimen examined (Alb-4960) 31.4 x 26.1 

mm in calicular diameter and 26 mm in height. Costae equal, 

broad, and flat; separated by thin, shallow intercostal striae; and 

covered with low, rounded granules. Primary costae slightly 

ridged at calicular margin. Lower half to two-thirds of theca 

usually discolored or eroded, the costae being well preserved 

only within 5-7 mm of calicular edge. Corallum otherwise 

white. 

Septa arranged in 3 size classes, with a tendency of most 

specimens examined to have 18-20 primary, 18-20 secondary, 

and 36-40 tertiary septa (72-80 total). Smaller specimens 

(e.g., those of Yabe and Eguchi, 1942b) of only 19 mm 

GCD have only 12 primary septa (48 septa total); specimens of 

24-26 mm GCD (e.g., the syntypes) have 14-16 primary septa 

(56-64 total septa). Primary septa highly exsert (up to 6 mm) 

and have straight, vertical inner edges that do not quite reach 

the columella. Secondary septa are the smallest septa, only 1-2 

mm exsert, three-quarters width of a primary, and have a 

sinuous inner edge. Tertiary septa 2-3 mm exsert (pairs fused 

to each adjacent primary septa at calicular edge) and equal to or 

slightly wider than the flanked secondary septum. Each 

secondary septum bordered internally by a high, granular, 

sinuous palus, usually wider (about 5 mm) than the secondary 

septum it borders. Total number of pali in palar crown 

corresponds to the number of secondary (or primary) septa in 

the calice, which is, in turn, correlated to the GCD. Fossa 

moderate in depth, containing a well-developed, elongate 

fascicular columella, composed of broad, closely adjacent 

elements. 

DISCUSSION.—Caryophyllia ambrosia belongs to a group of 

five species (see Cairns, 1991a, table 3) characterized by 


having relatively large, cornute, unattached coralla and three 

size classes of nonhexamerally arranged septa, the total usually 

exceeding 48: C. seguenzae Duncan, 1873; C. ambrosia 

Alcock, 1898 (?= C. scillaeomorpha Alcock, 1894); C. 

planilamellata Dennant, 1906; C. grandis Gardiner and Waugh, 

1938; and C. valdiviae Zibrowius and Gili, 1990. Caryophyllia 

ambrosia is distinguished from the other four species by having 

tertiary septa that are wider than their secondary septa and very 

broad columellar elements. There is little doubt that Yabe and 

Eguchi's (1942b) Caryophyllia cf. alcocki is simply an 

immature specimen of this species. 

Caryophyllia ambrosia occurs from the western Atlantic 

eastward to Honshu, Japan (Cairns, 1979; Zibrowius, 1980; 

Cairns and Keller, 1993); however, a separate subspecies, C. 

ambrosia caribbeana Cairns, 1979, was distinguished for the 

tropical western Atlantic populations. Likewise, North Pacific 

specimens differ from typical Indian Ocean specimens in 

having a tendency to have more septa and pali, i.e., usually 

72-80 septa and 18-20 pali. The typical form is not known to 

have more than 72 septa and 18 pali (Zibrowius, 1980). This 

may ultimately warrant the naming of another subspecies for 

the North Pacific specimens. 

Caryophyllia scillaeomorpha Alcock, 1894 antedates C. 

ambrosia by four years and appears to be conspecific, but until 

the types are examined I hesitate to make this synonymy, 

especially because of its unusually shallow depth of occurrence 

(196 m). 


USNM 92791; Alb-4960, 1, USNM 92792; Alb-4969, 1, CAS 

15259; TM (KT8916, T3-2), 2, USNM 92793; TM (KT9202, 

YT6), 17, USNM 92794, 3, ORI. Previous Records: 2 

syntypes of C. ambrosia from Laccadive Sea, 1070 fms, 

USNM 18157; specimens reported from eastern Atlantic by 

Zibrowius (1980), USNM and MNHNP; C. alcocki of Yabe 

and Eguchi (1942b), Soyo Maru-223, TIUS 53625 (Plate 21 e). 

TYPES.—Two hundred syntypes of C. ambrosia are presumably 

deposited at the Indian Museum, Calcutta; two are also 

deposited at the USNM (18157, Plate 2\d,g); and several 

specimens are at the MNHNP. Type Localities: Investigator 

station 104: 11 ° 12'47"N, 74°25'30"E, 1829 m (Elicapeni Bank, 

Laccadive Sea, Arabian Sea) and Investigator station 176: 

1 l°47'06"N, 73°57'3O"E (Laccadive Sea), 1957 m. 

The holotype of C. scillaeomorpha is presumed to be 

deposited at the Indian Museum, Calcutta. Type Locality: 

Investigator station 170: 13°01'06"N, 8O°36'56"E (off 

Madras, Bay of Bengal), 196 m. 

DISTRIBUTION.—Japan: Pleistocene of Okinawa, Naha limestone, 

2, USNM 88678. Recent: off Sagami Bay and 

southeastern Honshu; Miyaki-jima; mouth of Bungo Strait; 

Osumi Shoto, northern Ryukyu Islands; 311-2450 m. Elsewhere: 

Amphi-Atlantic, Indian Ocean; 430-2670 m. 

Caryophyllia (C.) laevicostata Moseley, 1881 

Caryophyllia laevicostata.—Yabe and Eguchi, 1932a:388. 

Caryophyllia sp.—Yabe and Eguchi, 1942b: 162.

NUMBER 557 49 

DISCUSSION.—Yabe and Eguchi (1932a) listed a specimen 

of C. laevicostata from 604 m "off Japan" but did not describe 

or illustrate it. The only Soyo Maru station made at that depth 

was station 262, from Suruga Bay. The specimen was later 

(Yabe and Eguchi, 1942b) referred to as Caryophyllia sp. 

Because the specimen has never been described, illustrated, or 

definitively identified, it is not considered as a legitimate record 

for the North Pacific. Caryophyllia laevicostata was originally 

described from off Ascension Island, Atlantic at 776 m and 

later synonymized by Zibrowius (1980) with C. atlantica 

(Duncan, 1873). C. atlantica has a distribution including the 

eastern Atlantic and Hawaiian Islands at depths of 776-2165 

m. 

Subgenus Caryophyllia (Acanthocyathus) 

Milne Edwards and Haime, 1848 

DIAGNOSIS.—Caryophyllia having coralla with edge spines. 

TYPE SPECIES.—Acanthocyathus grayi Milne Edwards and 

Haime, 1848a, by subsequent designation (Milne Edwards and 

Haime, 1850a:xiii). 

Caryophyllia (A.) grayi Milne Edwards 

and Haime, 1848 

PLATE 2\i-k 

Acanthocyathus grayi Milne Edwards and Haime, 1848a:293, pi. 9: figs. 2, 

2a.—Yabe and Eguchi, 1942b:121-122 [synonymy, no figure].— 

Umbgrove, 1950:641-642, pi. 81: figs. 27-32 [synonymy].—Wells, 

1984:209, pi. 2: figs. 5-9 [synonymy].—Eguchi and Miyawaki, 1975:57.— 

Zou, 1988:76, pi. 5: figs. 8, 9. 

DESCRIPTION.—Corallum ceratoid, compressed (GCD:LCD 

= 1.3-1.5), and usually slightly curved in plane of GCD. 

Largest specimen examined (USNM 71846, Pleistocene of 

Vanuatu) 21.7 x 16.9 mm in calicular diameter, but Yabe and 

Eguchi (1942b) reported a specimen 28 x 20 mm in calicular 

diameter, and the holotype is reported to be even larger at 30 x 

20 mm in calicular diameter and 40 mm in height. Coralla 

unattached, the pedicel narrowing to 1.5-1.8 mm in diameter. 

Each mature specimen bears up to 8 lateral spines: 2 or 3 on the 

concave edge and 3-5 on the convex edge. Spines up to 8 mm 

in length (but commonly broken) and circular to slightly 

elliptical in cross section. Costae equal in width, slightly 

convex, and finely granular. Corallum light reddish brown. 

Septa arranged in 14 to 18 sectors, the most common 

arrangement being: 14:14:28 (= 56 septa); however, end half 

systems often have extra pairs of quaternaries, i.e., 14:14:28:6 

(= 62 septa) and coralla with 18 sectors have up to 72 septa. 

Primary septa up to 2.8 mm exsert, extend about three-quarters 

distance to columella, and have straight inner edges. Secondary 

septa about 1.8 mm exsert, three-quarters width of primaries, 

and have sinuous inner edges. Tertiary septa equally exsert as 

secondaries but only about three-quarters their width, and have 

slightly sinuous inner edges. Quaternary septa usually restricted 

to end half-systems. A distinct crown of 14-18 wide 

(about 2 mm) pali occur before the secondary septa, each palus 

planar but with slightly sinuous edges. Fossa of moderate 

depth, containing an elongate columella composed of 7-10 

highly fused, slender twisted elements. 

DISCUSSION.—The only specimens of this species previously 

known from off Japan were those reported by Yabe and 

Eguchi (1942b) off Seto, Wakayama-ken, and those reported 

by Eguchi and Miyawaki (1975) off Kushimoto, Honshu. The 

two large specimens reported by Yabe and Eguchi (1942b), 

stated to be deposited at the Seto Marine Biological Laboratory 

of Kyoto University, are no longer there (Grygier, pers. 

comm.). Comparisons of this species to C. (A.) spiniger are 

made in the account of that species. 


AM6), 3, ORI; TM (KT9309, AM7), 5, USNM 93161, 8, ORI. 

Previous Records: Vanuatu Pleistocene specimens reported 

by Wells (1984), USNM. 

TYPES.—Holotype not traced. Type Locality: Unknown. 

DISTRIBUTION.—Japan: Off Seto and Kushimoto, Honshu; 

off Amami Oshima, Ryukyu Islands; 108-191 m. Elsewhere: 

Pliocene of Java. Plio-Pleistocene of Java, Japan, 

Ceram, Talaud, and Vanuatu. Recent: South China Sea; 

Philippines 04/6-5381, 5593); Andamans; off Burma; 37- 

490 m. 

Caryophyllia (A.) spiniger Kent, 1871 

PLATES 21/, 22a-d 

Acanthocyathus spiniger Kent, 1871:275-276, pi. 23: fig. la-c—Yabe and 

Eguchi, 1941c:212; 1942b: 112, 122. 

DESCRIPTION.—Corallum ceratoid to trochoid, compressed 

(GCD:LCD = 1.3-1.4), and straight. Largest specimen 

examined (Alb-5371) 18.2 x 14.1 mm in calicular diameter and 

17.6 mm in height. Corallum unattached, the pedicel only 

0.8-1.3 mm in diameter. C1-2 ridged, the two lateral C, giving 

the corallum alate edges. A pair of short (up to 5 mm), slender 

(about 0.7 mm in diameter) edge spines occurs close to the 

pedicel. A much larger pair of spatulate spines occurs slightly 

higher on calicular edges, each measuring up to 13 mm in 

length and up to 3.1 x 0.8 mm in cross section, the greater 

diameter of the spine being in plane of greater axis of corallum. 

In some large coralla, a third pair of spines similar to the second 

pair occur higher on thecal edge. In about one-quarter of the 

specimens examined, 4 large additional thecal spines occur 

(Plate 22d), similar to the second edge pair in size, one midway 

on each lateral Cl. Corallum reddish brown to white. 

Septa hexamerally arranged in four complete cycles according 

to the formula: S1>S2>S3>S4. S, highly exsert (up to 4.5 

mm), extend about four-fifths distance to columella, and have 

straight inner edges. S2 only about 2.5 mm exsert, only 

four-fifths width of an S,, and also have straight inner edges. S3 

least exsert (about 1.5 mm), about two-thirds width of an S2, 

and have sinuous inner edges. Each pair of S4 adjacent to an S, 

are highly exsert (about 2.3 mm) and fused to its adjacent S^ 

likewise, each pair of S4 adjacent to an S2 are as exsert as an S3 

and also fused to its adjacent S2. S4 about two-thirds width of

50 

an S3 and have slightly sinuous inner edges. Septa are well 

spaced. A crown of 12 planar (2 mm wide) P3 occurs within the 

fossa encircling a linear columella composed of 3-6 broad, 

twisted elements. 

DISCUSSION.—The only Japanese record of this species was 

Kent's original description from an unspecified location and 

depth off Japan. No additional North Pacific specimens are 

reported herein, and the description above is based on three 

syntypes and a suite of specimens from off the Philippines 

(Alb-5369,5371). 

Caryophyllia (A.) spiniger is distinguished from C. (A.) 

grayi by several characters. Caryophyllia spiniger always has 

48 septa and 12 pali; C. grayi has 56-72 septa and 14-18 pali. 

Also, C. spiniger has: ridged Cuz* not ' ow > slightly convex 

costae; highly exsert S, that are larger than its S2; and a 

straight, not curved, corallum and thus a symmetrical arrangement 

of edge spines, not unequal as in C. grayi. Furthermore, 

the edge spines of C. spiniger are spatulate, those of C. grayi 

are circular to elliptical in cross section. Finally, there is a 

tendency for lateral thecal spines to occur in C. spiniger, but not 

in C. grayi. 


USNM 92689; Alb-5371, 28, USNM 92690, 2, ORI. Previous 

Records: 3 syntypes, BM. 

TYPES.—At least 3 syntypes (Plates 21/, 22c) of A. spiniger 

are deposited at the BM (uncataloged). Type Locality: 

"Japan," depth unknown. 

DISTRIBUTION.—Off Japan, depth range unknown. Elsewhere: 

Philippines; 152-194 m. 

Subgenus Caryophyllia (Premocyathus) Yabe 

and Eguchi, 1942b 

DIAGNOSIS.—Caryophyllia having compressed coralla with 

alate to carinate lateral edges. 

TYPE SPECIES.—Premocyathus compressus Yabe and 

Eguchi, 1942b, by original designation. 

DISCUSSION.—Yabe and Eguchi (1942b) established Premocyathus 

as a separate genus but Wells (1956) later 

considered it to be a subgenus of Caryophyllia. I (Cairns, 

1991a) treated it as separate genus, but now agree with Wells, 

that it should be a subgenus of Caryophyllia. Its variation from 

typical Caryophyllia is no more than that of the other subgenus 

Acanthocyathus, both subgenera differing only in the degree of 

septa! edge modification. 

Caryophyllia (P.) compressa Yabe and Eguchi, 1942 

PLATE 22*,/ 

Caryophyllia compressa Yabe and Eguchi, 1932b:443 [nomen nudum]. 

Premocyathus compressus Yabe and Eguchi, 1942b: 121, 151-152, pi. 10: figs. 

13. 14.—Eguchi, 1965:285, fig.—Eguchi and Miyawaki. 1975:57. 

Not Caryophyllia compressa Gardiner and Waugh. 1938:180 (junior homonym, 

replacement name C. zanzibarensis Zou. 1984]. 

Not Caryophyllia (Premocyathus) compressus.—Wells. I956:F422, fig. 323, 3. 


Not Premocyathus compressus.—Cairns, 1984:14. 

Caryophyllia {Premocyathus) compressa.—Mori, 1987:21-30,9 figs. 

Description of Recent Specimens.—Corallum compressed, 

free, and evenly curved up to 90° in plane of GCD. Largest 

specimen examined (TM (KT9015, BS2)) 8.3 x 5.4 mm in 

calicular diameter (GCD:LCD = 1.53) and 18.5 mm in height. 

Base of corallum invariably broken, revealing an irregularly 

circular scar 1.5-2.0 mm in diameter. Outer (convex) side of 

corallum slightly ridged; inner (concave) edge evenly rounded. 

Costae well developed, slightly convex, and coarsely granular. 


Septa usually decamerally arranged in three size classes, the 

tertiary cycle often incomplete. Common septal/palar arrangements 

include: 10:10:20, 10; 10:10:18, 9; 10:10:10, 5; and 

10:10:8, 4 (notation follows that of Mori, 1987, in which the 

number of pali follows the comma). There is in general a direct 

correlation between number of septa and GCD, the first pairs of 

tertiary septa (and P2) occurring in sectors adjacent to the 

principal septa (termed "spaces iv and v" by Mori, 1987). 

Primary septa moderately exsert (up to 0.7 mm), and have 

sinuous, vertical inner edges that attain the columella. 

Secondary septa about three-quarters width of primaries and 

also have quite sinuous inner edges. Tertiary septa equal to or 

only slightly less wide than secondaries, and have almost 

straight inner edges. Slender, sinuous pali occur only before 

those secondary septa flanked by tertiary septa, the palar crown 

thus ranging from 1-10 elements. Fossa of moderate depth, 

containing a fascicular columella composed of 8-15 slender, 

twisted elements. 

DISCUSSION.—In a thorough mophological analysis of over 

a thousand Pleistocene specimens of C. compressa from the 

type locality, Mori (1987, fig. 3) graphed a direct relationship 

between calicular size and number of septa, showing that when 

tertiary septa are present, they are most likely to first occur in 

the sectors adjacent to the principal septa and least likely to 

occur in lateral sectors. Nonetheless, he concluded that septal 

number was not a function of size, but rather genetically 

determined. Mori also found an enormous range of variation of 

septal arrangements (57) and septal symmetries (9), ranging in 

symmetry from hexameral (6:6:12, 6) to decatetrameral 

(14:14:12,6), the first and last symmetries being quite rare. The 

most common symmetry was decameral (68% of the coralla) 

and the most common septal arrangement was: 10:10:14, 7 

(i.e., 34 septa, 7 pali); the maximum number of septa and pali 

never exceeding 48 and 12, respectively. 

Specimens reported as P. compressus by Wells (1956) and 

Caims (1984) from the Philippines are now thought to be 

misidentified for the following reasons. The Philippine 

specimens have: papillose (not fascicular) columellas; less 

compressed calices; straight-edged septa; and consistently 

dodecamerally arranged septa (xl2). Notwithstanding the fact 

that Mori found some (3%) dodecameral C. (P.) compressa, 

virtually all of the Philippine specimens have that symmetry.

NUMBER 557 51 

MATERIAL EXAMINED.—New Records: USGS 17450, near 

Iwa, Okinawa, Shimijiri marl (Pleistocene), 1, USNM 88652; 

TM (KT9015, BS2), 8, USNM 92686, 3, ORI; TM (KT9202, 

OS2), 1, USNM 92687; TM (KT9202, OS3), 4, USNM 92688; 

TM (KT9309, AM8), 1, ORI; off Misaki, Japan, Mortensen 

Expedition, 30 June 1914, 366 m, ZMC. Previous Records: 

Specimen illustrated by Wells (1956), USNM. 

TYPES.—Syntypes of P. compressus are deposited at the 

TIUS (see Yabe and Eguchi, 1942b). Type Locality: Pleistocene, 

Ryukyu limestone of Kikai-jima. 

DISTRIBUTION.—Pleistocene of Ryukyu Islands. Recent: 

Kii and Bungo Straits; van Dienem Strait, southern 

Kyushu; Wakasa Bay, Honshu, Sea of Japan; off Amami 

Oshima, Ryukyu Islands; 115-422 m. 

Crispatotrochus Tenison Woods, 1878 


Crispatotrochus rube see ns (Moseley, 1881) 

PLATE 22gji 

Cyathoceras rubescens Moseley, 1881:157, pi. 2: fig. 8a-c.—Marenzeller, 

1888b:21-22.—Yabe and Eguchi, 1942b: 117.—Cairns. 1984:15. 

Cyathoceras diomedeae Vaughan, 1907:77-78, pi. 7: figs. 1, 2. 

Cyathoceras diomedae [sic].—Yabe and Eguchi, 1942b: 116-117, pi. 9: fig. 8. 

Crispatotrochus rubescens.—Cairns, 199la: 15. 

DESCRIPTION.—Corallum ceratoid, with a flared calice and a 

firmly attached pedicel about one-quarter diameter of calice. 

Illustrated specimen (Soyo Afaru-331) 24.3 x 20.6 mm in 

calicular diameter and 30.4 mm in height, with a pedicel 

diameter of 6.6 mm. Costae ridged only near calice, otherwise 

consisting of broad, flat, finely granulated strips. Corallum 

white or reddish brown. 

Septa hexamerally arranged in 5 cycles: S1.2>S3>S4>S5. 

S1-2 highly exsert (as much as 7 mm), and have sinuous, 

vertical inner edges that join the columella. S3 considerably 

less exsert (about 4 mm), but as much as 80% as wide as S^ 

and also have sinuous inner edges. S4.5 equally exsert (about 

2.5 mm), the S4 about half size of S3 and the S5 about half size 

of S4. Fossa moderately deep, containing a crispate, fascicular 

columella composed of 15-20 slender, twisted elements. 

DISCUSSION.—Of the 12 valid species of Crispatotrochus 

(see Cairns, 1991a), only three have five cycles of hexamerally 

arranged septa: C. rubescens (Moseley, 1881); C. niinoi (Yabe 

and Eguchi, 1942b); and C.foxi (Durham and Barnard, 1952), 

all of which occur in the temperate North Pacific and included 

in this account. C. rubescens is compared to the other two 

species in the discussions of those species. 

Both new records reported herein are small specimens with 

a GCD less than 8 mm and have only four cycles of septa. 

MATERIAL EXAMINED.—New Records: TM (CR79-1), 1, 

USNM 92685; Okinose, Sagami Bay, 110 m, 11 June 1914, 1, 

ZMC. Previous Records: Soyo Maru-425,2, TIUS 53695 (C 

rubescens of Yabe and Eguchi, 1942b); Soyo Afaru-331, 1, 

TIUS 53694 (C. diomedae of Yabe and Eguchi, 1942b), Plate 

22/i. 

TYPES.—The holotype of Cyathoceras rubescens is lost (see 

Caims, 1984). Type Locality: Challenger-192: 5°49 / 15"S, 

132°14'15"E(Kei I., Banda Sea, Indonesia), 236 m. 

The types of Cyathoceras diomedeae are deposited at the 

USNM (see Cairns, 1991b). Type Locality: Alb-3835: 

21°00'10"N, 157°5(rw (south of Molokai, Hawaiian Islands), 

309-333 m. 

DISTRIBUTION.—Japan: Sagami Bay and off Kushimoto, 

Honshu; off southeastern Shikoku; off Koshiki I., southwestern 

Kyushu; 110-344 m. Elsewhere: Hawaiian Islands, Christmas 

I., Philippines, Banda Sea; 183-634 m. 

Crispatotrochus niinoi (Yabe and Eguchi, 1942) 

Cyathoceras niinoi Yabe and Eguchi, 1942b:117, 145-146, pi. 9: fig. 

9a,b.—?Song, 1982:134-135, pi. 2: figs. 1, 2; 1991:133. 

Crispatotrochus niinoi.—Cairns, 1991 a: 15. 

REDIAGNOSIS OF HOLOTYPE.—Corallum ceratoid: 15 x 10.5 

mm in calicular diameter and 20 mm in height, with a pedicel 

diameter of 2 mm (PD:GCD = 0.13). Theca thick and smooth; 

noncostate. Upper theca white, lower three-quarters of theca 

eroded and encrusted. Septa hexamerally arranged in 5 

complete cycles (96 septa) according to the formula: 

S1.2>S3>S4>S5. S,_2 slightly exsert (about 1.1 mm) and 

extend to the columella. S3 about three-quarters width of an 

S^; S4 about three-quarters width of an S3; S5 rudimentary. 

Columella fascicular, composed of 7 fused twisted elements. 

DISCUSSION.—Crispatotrochus niinoi is known from only 

one, perhaps two (if Song's specimen is included), specimens, 

neither of which is available for study. The dignosis above 

comes from Yabe and Eguchi's (1942b) original description. 

Only two other species of Crispatotrochus are known to 

have five cycles of septa: C. rubescens (Moseley, 1881) and C. 

foxi (Durham and Barnard, 1952). Crispatotrochus niinoi 

differs from C. rubescens in having a smaller corallum and 

pedicel (PD:GCD = 0.13 vs 0.22-0.27 for C. rubescens); a 

thicker, noncostate theca; and less exsert septa. Crispatotrochus 

niinoi is compared to C.foxi in the account of that species. 

Song's (1982) record from off South Korea is queried because 

the PD:GCD is 0.80. 

MATERIAL EXAMINED.—None. 

TYPES.—The holotype of Cyathoceras niinoi is deposited at 

the TIUS (59070). Type Locality: Taito-zaki, Chiba-ken (off 

Boso Peninsula, Honshu), 104 m. 

DISTRIBUTION.—Off Boso Peninsula, Honshu; ? Western 

Channel of Korea Strait off Pusan, South Korea, depth 

unknown (Song, 1982); 104 m. 

Paracyathus Milne Edwards and Haime, 1848a 

DIAGNOSIS.—See Part 1.

52 

Paracyathus pruinosus Alcock, 1902 

PLATES 22/,;, 42a 

Paracyathus pruinosus Alcock, 1902c: 18, 19, pi. 3: figs. 17, 17a.—Van der 

Hoist, 1931:9, pi. 1: figs. 9,11; pi. 2: fig. 6.—Yabe and Eguchi. 1942b: 126. 

pi. 11: fig. 1.—Eguchi, 1965:287, fig.—NotZou, 1988:76, pi. 5: figs. 10, 11. 

REDESCRIPTION OF FIGURED SYNTYPE (ZMA Coel. 1307).— 

Corallum robust and dense, 20.1 x 15.6 mm in calicular 

diameter, and 21.9 mm in height, with a pedicel diameter of 

11.8 mm (PD:GCD = 0.59). Theca worn and encrusted, but 

near calice costae are flat and equal in width (about 1 mm), 

covered with very small granules 5-6 across the width of a 

costa. Intercostal striae thin and shallow. Corallum light brown. 

Septa arranged in 13 half-systems accordingly: 13:13:26:4 (56 

septa). The other, slightly larger (GCD = 20.6 mm) syntype 

(ZMA 1089) has 82 septa, but its symmetry is obscured by the 

polyp. Eleven of the half-systems are composed of two thick 

bordering primary septa, a secondary septum, and a pair of 

tertiary septa, whereas 2 of the half-systems also have a pair of 

quateraries. Primary septa only slightly exsert (about 1.8 mm) 

and have straight, entire inner edges, each bordered by a thick 

palus 1-2 mm wide that is adjacent to the columella. 

Secondary and tertiary septa only slightly less exsert than 

primary septa (about 1.6 mm), the secondaries about 80% 

width of a primary and also bordered by a palar crown, each P2 

standing taller in the fossa and frequently divided into 2-4 

subelements. Tertiary septa of about the same width as 

secondaries but much less thick. All septa closely spaced and 

covered with prominent ridge-like granules, the orientation of 

the ridges parallel to the septal edges. Palar faces also 

prominently ridged. Fossa shallow, containing a large elliptical 

and slightly convex papillose columella composed of about 25 

elements. 

DISCUSSION.—The two Japanese specimens examined (Soyo 

Maru-235) are 18.0 and 14.8 mm in GCD and appear to be 

conspecific. Although the septal symmetry is difficult to 

distinguish, both specimens seem to have hexamerally arranged 

septa in five cycles, the last incomplete. The larger 

specimen contains 66 septa: 6 half-systems having no S5, 3 

half-systems having 1 pair of S5, and 3 half-systems having 2 

pairs of S5. The smaller specimen has 70 septa: 4 half-systems 

without S5, 5 with 1 pair of S5, and 3 with 2 pair of S5. The 

half-systems that lack pairs of S5 are invariably those on the 

lateral faces of the calicular ellipse; those having 2 pairs 

invariably in the end half-systems. The most similar species to 

P. pruinosus in the North Pacific is Trochocyathus caryophylloides, 

both species having a similar corallum shape, size, 

columella, and palar configuration. Paracyathus pruinosus is 

distinguished by having divided paliform lobes (P3); little 

exsert, very thick septa; highly granular septal and palar faces; 

and a shallower fossa. 


Records: Soyo Maru-235, 2, TIUS 53681 (Yabe and Eguchi, 

1942b) (Plate 42a); syntypes of P. pruinosus. 


TYPES.—Two syntypes (Plate 22/J) of P. pruinosus are 

deposited at the ZMA (Coel. 1307, 1089). Type Locality: 

Siboga-96: Southeast of Pearl Bank, Sulu Archipelago; 

?15 m. 

DISTRIBUTION.—Japan: Off Hochijo Jima (off Sagami Bay); 

176-198 m. Elsewhere: Sulu Archipelago; Saya de Malha, 

Indian Ocean; [15]-150 m. 

Trochocyathus Milne Edwards and Haime, 1848a 

DIAGNOSIS.—Corallum solitary, turbinate to ceratoid, fixed 

or free. Septotheca costate, sometimes covered with a thin 

epitheca. Pali before all but last cycle in two crowns; columella 

papillose. 

TYPE SPECIES.—Turbinolia mitrata Goldfuss, 1827, by 

subsequent designation (Milne Edwards and Haime, 

1850a:xiv). 

Trochocyathus caryophylloides Alcock, 1902 

PLATE 23a-c,h 

Trochocyathus caryophylloides Alcock, 1902a:94; 1902c:14-15,pl. 2: figs. 10, 

10a.—Yabe and Eguchi, 1942b: 123-124, pi. 10: fig. 21.—Not Zou. 

1988:76, pi. 5: fig. 5, 5a. 

DESCRIPTION OF SPECIMEN FROM Alb-4891.—Corallum 

trochoid: 15.6 x 13.9 mm in calicular diameter, 15.2 mm in 

height, and 6.8 mm in pedicel diameter (PD.GCD = 0.44). 

Costae broad (0.6-0.7 mm), flat, and equal in width, covered 

with small (0.10 mm in diameter) granules up to 4 across a 

costal width near the calice; intercostal striae quite thin (0.05 

mm) and shallow. Corallum white. 

Septa arranged in 15 sectors accordingly: 15:15:30:2 (62 

septa). Primary septa only slightly exsert (about 1.7 mm) and 

extend about two-thirds distance to columella, where each is 

bordered by a small (0.7 mm width), low palus, together 

forming a palar crown tightly encircling the columella. 

Secondary septa only slightly less exsert (about 1.5 mm), half 

width of primaries, and have sinuous inner edges. Each 

secondary septum bordered by a broad (about 1.8 mm wide), 

sinuous, highly granular palus, together forming a second palar 

crown that is much more prominent than the first. Tertiary septa 

only slightly less exsert and less wide than secondaries. In one 

(developing?) sector, a pair of quaternaries encloses the tertiary 

septum, the latter bearing a palus, thus resulting in 31 pali in the 

corallum. Fossa of moderate depth, containing an columella 

composed of an elliptical field of 15 discrete, papillose 

elements. 

DISCUSSION.—The Albatross specimen differs from the 

syntypes (GCD up to 21 mm) in having a smaller corallum, two 

less septa and one less palus, and a wider pedicel. The lesser 

number of septa and pali are probably due to the smaller size of 

the Albatross specimen, one of its sectors apparently being in 

the process of enlarging. Alcock (1902c) illustrated only one of 

the six syntypes, one having a very narrow pedicel (PD:GCD =

NUMBER 557 53 

0.11); however the other syntypes have much more robust 

pedicels, up to a PD:GCD of 0.38 (Siboga-253). Therefore, the 

PD:GCD of 0.44 of the Albatross specimen is not considered 

unusual. Yabe and Eguchi's (1942b) specimen of 21.0 mm 

GCD has the typical 64 septa in 16 sectors and a PD:GCD of 

0.29. T. caryophylloides is easily distinguished from the other 

North Pacific Trochocyathus by its decahexameral (xl6) septal 

symmmetry. For this reason, Zou's (1988) identification 

probably is not correct, his specimen having four hexamerally 

arranged cycles of septa (48 septa). 

MATERIAL EXAMINED.—New Record: Alb-4891, 1, CAS 

74940. Previous Records: Syntypes of T. caryophylloides 

from Siboga-96 (1) ZMA Coel. 1325, Siboga-253 (2) ZMA 

Coel. 1324. 

TYPES.—Six syntypes (Plate 23b,c) of T. caryophylloides 

were discussed by Alcock, five of which are deposited at the 

ZMA (Coel. 1324, 1325, 5436, 5437) (see van Soest, 1979). 

Type Localities: Siboga-96, 251, 253: Celebes and Banda 

seas, Indonesia, ?15, 115-304 m. 

DISTRIBUTION.—Japan: Off Boso Hanto, Honshu; Bungo 

Strait; East China Sea south of Fukue Jima; 115-344m. 

Elsewhere: Celebes and Banda Seas; [15]-304 m. 

Trochocyathus japonicus Eguchi, 1968, nom. correct 

Trochocyathus japonka Eguchi, 1968:C34-35, pi. C6: figs. 4-6. 

Ceratotrochus japonicus Eguchi, 1968:C38, pi. C6: figs. 1-3 [not pi. Cl 1: figs. 

1-3; pi. 20: figs. 1,2). 

?Ceratorochus [sic] jogashimaensis Eguchi, 1968:C38, pi. Cl 1: figs. 7-9. 

DISCUSSION.—Eguchi (1968) described three remarkably 

similar species from virtually the same locality, the holotypes 

of the first two species coming from the same haul: 

Trochocyathus japonicus, Ceratotrochus japonicus, and Ceratotrochus 

jogashimaensis. Each species was described from a 

single specimen. 

From Eguchi's (1968) account, Trochocyathus japonicus 

can be characterized as having a small (GCD = 12 mm), 

trochoid corallum; hexamerally arranged septa in four complete 

cycles (S1_2>S3>S4); two crowns of pali, one before the S^ 

and another before the S3; a papillose columella; and a costate 

theca. Ceratotrochus japonicus, illustrated on the same plate as 

T. japonicus (plate C6) and collected in the same haul, is 10 

mm in GCD and also has four hexamerally arranged cycles of 

septa, a papillose columella, and costate theca. Eguchi was 

equivocal about the presence of pali, stating that lobes were 

present before some of the S3 as well as having 6 "pali-like 

rods" radiating from the columella. By definition (see 

Chevalier, 1961), the genus Ceratotrochus is distinguished 

from Trochocyathus by having lobes before only the first two 

cycles, whereas in Trochocyathus, they occur before all but the 

last cycle (i.e., in this case, the first three cycles) in two crowns. 

The preservation of the holotype of C. japonicus apparently 

does not allow this observation, but, if some P3 are present in 

this specimen, they would indicate an affinity to Trocho- 

cyathus, not Ceratotrochus, and probably conspecificity with 

T. japonicus. The additional specimen of C. japonicus 

illustrated by Eguchi (Specimen #796b: Plate Cl 1: figs. 1-3; 

PI. 20: figs. 1, 2) is not mentioned in the text and appears to be 

a juvenile Conotrochus; it may have been confused or 

mislabelled in his text. 

The holotype of Ceratotrochus jogashimaensis (GCD = 11 

mm) was also collected from virtually the same locality but 

from slightly deeper water. It is identical to the other two 

species except that it appears to lack pali. 

MATERIAL EXAMINED.—None. The types of these three 

species are not available for study and, unfortunately, no 

additional specimens are available to further characterize and 

illustrate this taxon. 

TYPES.—The holotypes of Trochocyathus japonica and 

Ceratotrochus japonicus are presumed to be deposited at the 


(#785, 782, respectively). Type Locality: 5 km WSW of 

Jogashima, Sagami Bay, 150-250 m. 

The holotype of Ceratotrochus jogashimaensis is also 

presumed to be deposited at the Biological Laboratory of the 

Imperial Household, Tokyo (#779). Type Locality: 5 km SW 

of Jogashima, Sagami Bay, 300-450 m. 

DISTRIBUTION.—Known only from 5 km WSW of Jogashima, 

Sagami Bay, Honshu, Japan; 150-450 m. 

Trochocyathus decamera, sp. nov. 

PLATE 23d.e 

DESCRIPTION.—Corallum trochoid to subcylindrical and 

firmly attached through a wide pedicel. Holotype 6.4 x 6.3 mm 

in calicular diameter, 7.9 mm in height, and 5.0 mm in pedicel 

diameter. Theca faintly costate, overlain with very thin, 

transversely banded epitheca through which underlying granules 

can be seen. Corallum white. 

Septa decamerally arranged in three cycles, the holotype 

having an extra pair of quaternary septa (10:10:20:2, 42 septa). 

Primary septa little exsert (about 0.7 mm), extend about 80% 

distance to the columella, and have slightly sinuous inner 

edges. Each primary septum bordered internally by a small 

palus about 0.5 mm in width, which together form a crown 

adjacent to and rising just above the columella. Secondary 

septa about 0.5 mm exsert, extend about half distance to 

columella, and are bordered by much larger, sinuous pali. Each 

palus about 0.9 mm in width, together forming another palar 

crown that is higher in the fossa than the first but reaching the 

same distance into the fossa. Tertiary septa equally exsert and 

slightly more wide than secondary septa. The pair of 

quaternaries in the holotype reach only one-third distance to the 

columella and cause a palus to form before the tertiary septum 

in this sector, thus resulting in 21 pali in the holotype. The 

unique tertiary palus is larger than the others and extends even 

higher in the fossa than those before secondary septa. Fossa

54 

shallow, containing an elliptical field of 12-15 small, 

granulated pillars. 

DISCUSSION.—Trochocyathus decamera is superficially 

similar to T. japonicus but can be distinguished by having 

decameral symmetry, a thin epitheca, and tertiary septa wider 

than its secondary septa. In T. japonicus, the S4 (equivalent to 

tertiaries of T. decamera) are less wide than the S3 (equivalent 

to secondaries of T. japonica). 

The genus Tethocyathus differs from Trochocyathus in 

having a thick epitheca, Trochocyathus having only a thin or no 

epitheca (Cairns, 1979:76-77). Therefore, T. decamera is 

placed in Trochocyathus. 

ETYMOLOGY.—The species name decamera (Greek: deka, 

meaning 10, and meros, meaning division or parts) refers to the 

decamerally arranged septa in 10 septal sectors. 

MATERIAL EXAMINED/TYPES.—Holotype: TM (KT9202, 

YT1), USNM 92693.-Paratypes: TM (KT9202, YT1), 1, 

USNM 92694; TM (KT9015, CB1-2), 1, ORI. Type Locality: 

30°15'N, 130°46'E (Osumi Shoto, northern Ryukyu 

Islands), 80-88 m. 

DISTRIBUTION.—Osumi Shoto, northern Ryukyu Islands and 

off Mishima, Eastern Channel, Korea Strait; 70-88 m. 

Trochocyathus cooperi (Gardiner, 1905), comb. nov. 

PLATE 23f,g 

Tropidocyathus cooperi Gardiner, 1905:955, pi. 93: fig. 30. 

Trochocyathus.—Vaughan and Wells, 1943:48, fig. 20b. 

DESCRIPTION.—Corallum (anthocyathus) a highly compressed 

(GCD:LCD = 1.4-2.3), straight cylinder, with parallel 

thecal edges and only slightly divergent thecal faces. Largest 

Japanese specimen 13.4 x 8.6 mm in calicular diameter and 

18.5 mm in height. Base of each thecal edge bears a downward 

projecting crest up to 4.5 mm in length and flattened in plane of 

GCD. Between these 2 crests is the basal scar of attachment, 

which is elliptical to almost circular: 3.9-4.3 x 3.3-3.4 mm in 

diameter. Only one small specimen is fully intact, still having 

its anthocyathus attached to the fixed anthocaulus. It measures 

7.3 x 3.9 mm in calicular diameter and 8.0 mm in height, with 

2 edge crests, but it is assumed that the eventual transverse 

division would have taken place about 3.3 mm from the basal 

disc, just under the edge crests, where the cross sectional 

diameter is about 4x3 mm. The holotype also has a partial 

anthocaulus remaining attached to its base. Costae equal in 

width (0.5 mm), flat, and granular (about 2 granules across a 

costa), becoming slightly convex and separated by deeper 

furrows near calicular edge. Corallum reddish brown, with 

slightly more intense color and dark costal speckling in a 

narrow band about 1.5 mm from the calicular edge. 

Septa hexamerally arranged in 4 complete cycles: 

S1_2>S3>S4 (48 septa); however, larger coralla often have pairs 

of S5 in the end half-systems comprising up to 56-62 septa. 

S,_2 little exsert (about 0.7 mm), extend about two-thirds 

distance to columella, and have slightly sinuous inner edges. A 


slender (about 0.6 mm wide) palus sometimes occurs on the 

inner edge of each S^, forming a deep-seated palar crown 

directly adjacent to the columella, but often these pali are 

missing or indistinguishable from columellar elements. S3 only 

slightly less exsert (0.6 mm), about two-thirds width of S,_2, 

and have sinuous inner edges. Each S3 bordered by a wide (up 

to 1.1 mm) palus, together forming a prominent palar ring 

encircling columella. S4 0.5 mm exsert and about threequarters 

width of an S3. When present, pairs of S5 are quite 

small. Fossa of moderate depth, containing an elongate 

columella consisting of several papillae, some of which may be 

indistinguishable in size and position with P1-2. 

DISCUSSION.—Several caryophylliid genera reproduce 

asexually by transverse division (Cairns, 1989b), Trochocyathus 

being one of them. The original reference to transverse 

division in this genus was Vaughan and Wells (1943:47), who 

cited the process in two undescribed species from the 

Philippines and in Trochocyathus hastatus Boume, 1903. 

Bourne's species subsequently has been divided (Wells, 1984) 

into two species: Stephanocyathus (A.) hastatus and 

Bourneotrochus veroni Wells, 1984 (= Deltocyathus stellulatus 

Cairns, 1984), the latter established for a spinose, discoidal, 

deltocyathid-like species having transverse division. Therefore, 

T. cooperi is the only described species of Trochocyathus that 

is known to have transverse division. 

The two syntypes reported by Gardiner (1905) are larger than 

the Japanese specimens, having up to 10 pairs of S5 (68 septa) 

and a better developed columella. The Japanese specimens 

appear to be the first report of this species subsequent to its 

description. 


YT1), 23, USNM 92691, 3, ORI. 

TYPES.—Two syntypes of T. cooperi are presumed to be 

deposited at the BM. Type Locality: Kolumadulu and 

Suvadiva, Maldive Islands, 64-70 m. 

DISTRIBUTION.—Off Japan: Osumi Shoto, northern Ryukyu 

Islands; 80-88 m. Elsewhere: Maldive Islands; Philippines; 

64-70 m. 

Deltocyathus Milne Edwards and Haime, 1848a 

DIAGNOSIS.—Solitary, discoidal to patellate, free in adult 

stage. Well-developed costae present. Pali present before all 

but last cycle; within each system the inner edges of each pair 

of P3 fuse to the P2 near the columella, forming characteristic 

chevrons (deltas). Paliform lobes may also be present before 

fourth septal cycle. Columella papillose. 

TYPE SPECIES.—Turbinolia italica Michelotti, 1838, by 

monotypy. 

Deltocyathus vaughani Yabe and Eguchi, 1932 

PLATES 23/,./, 24a-c,f 

Not Deltocyathus orientalis Duncan, 1876:431 [= Peponocyathus). 

Levipalifer orientalis Vaughan, 1900:201-202, pi. 16: figs. 3-7 [junior

NUMBER 557 55 

secondary homonym of D. orientalis Duncan, 1876]. 

Deltocyathus vaughani Yabe and Eguchi, 1932a:388-389 [replacement name 

for D. orientalis Vaughan, 1900]; 1937:130, 135-138, pi. 20: fig. lla-c, 

12a-c; 1942b:113, 126.—Eguchi, 1965:287, 3 figs.; 1968:C35.—Eguchi 

and Miyawaki, 1975:57.—Not Keller, 1982:51, pi. 1 [= 4, plates 

miscaptioned]: figs. 1, 2, 7.—Zibrowius and Grygier, 1985:121, fig. 12. 

Deltocyathus (Levipalifer) vaughani.—Wells, 1956:F423-424, figs. 325, 4a,b. 

DESCRIPTION.—Corallum patellate, having a convex to 

almost flat base with a basal angle ranging from 130°-170°. 

Largest corallum examined (Alb-4973) 22.9 mm in calicular 

diameter and 8.9 mm in height, but Yabe and Eguchi (1937) 

reported an even larger specimen of 24 mm calicular diameter. 

Holotype 19.7 mm in calicular diameter and 7.8 mm in height. 

Costae ridged, becoming increasingly prominent toward 

calicular edge where they are up to 1.5 mm in height. C1-2 

originate at epicenter, C3, about 0.5 mm from epicenter; and C4, 

about 2 mm from epicenter. Each costa bears a linear row of 

tall, blunt to clavate granules, giving costal edges a coarsely 

serrate aspect. Costae also laterally spinose. Intercostal space 

up to twice width of a costa and usually flat; however, near 

calicular edges of large specimens there is sometimes a low 

ridge bisecting this space. Corallum white. 

Septa hexamerally arranged in 4 complete cycles, all 

specimens examined (as small as 9 mm in calicular diameter) 

having 48 septa. S! highly exsert (up to 3.5 mm), each bearing 

a prominent palus 1.3-1.6 mm wide, which joins the 

columella. S2 less exsert (about 3.0 mm), about three-quarters 

width of an Sv and also bear a similar-sized palus, but 

positioned slightly higher in fossa and more recessed from 

columella than the P^. S3^ equally exsert (about 2.5 mm), the 

S3 being about two-thirds width of an S2, but S4 are slightly 

wider than S3, about three-quarters width of an S2. Each S3 

bears a very wide palus equal to or wider than the septum it 

borders, these lobes forming a crown higher in the fossa and 

more recessed than P2 crown. S4 often bear narrow paliform 

lobes, each pair of which fuse to its enclosed P3. If P4 are 

absent, S4 inner edges fuse to P3. Inner edges of P2 and P3 

loosely connected near columella. Fossa shallow. Columella an 

elongate field of small interconnected papillae. 

DISCUSSION.—Deltocyathus vaughani is a relatively common 

and very distinctive deep-water species in the Japanese 

region. It resembles the Atlantic D. conicus Zibrowius, 1980 (= 

D. sp. cf. D. italicus sensu Cairns, 1979) in conical corallum 

shape and costal morphology, but differs in having P4. In fact, 

the character of having P4 distinguishes D. vaughani from all 

other species in the genus that have four cycles of septa and was 

used as the basis for the creation of the genus Levipalifer 

Vaughan, 1900. However, I agree with Yabe and Eguchi 

(1937), not Vaughan (1900) or Wells (1956), that the variable 

presence of P4 does not justify a separate genus or even 

subgenus. 

Even though Duncan's (1876) Deltocyathus orientalis was 

transferred to the genus Peponocyathus, it was necessary to 

provide a replacement name for Vaughan's (1900) Levipalifer 

orientalis, being a junior secondary homonym of Duncan's 

species. Yabe and Eguchi (1932a, 1937) provided the name 

Deltocyathus vaughani as this replacement name. 

The specimens reported by Keller (1982) from the South 

Pacific (425-1640 m) are similar to D. vaughani in corallum 

shape and in having P4 lobes, but their costae appear to be very 

different and are therefore not included in the synonymy of this 

species. 

One specimen (TM (KT7811, OT4), Plate 24/) has a 

well-developed ascothoracid gall, similar to those previously 

described by Zibrowius and Grygier (1985) for this species. 




USNM 92716; Alb-4915, 32, CAS 74942; A1M958, 1, USNM 

92717; Alb-4966, 1, USNM 82152; Alb-4967, 6, CAS 16326; 

Alb-4968, 3, CAS 80906; Alb-4972, 11, CAS 80942; Alb- 

4973, 1, USNM 92718; Alb-5054, 57, USNM 92719; 


Alb-5093, 4, USNM 92722; TM (KT7811, OT4), 4, USNM 

92723, 26, ORI; TM (KT7818, OT10), 2, USNM 92724; TM 

(KT7911, OT4), 7, USNM 92725. Previous Records: Holotype 

of L. orientalis, USNM. 

TYPES.—The holotype (Plate 23/,/) of Levipalifer orientalis 

is deposited at the USNM (19391). Type Locality: "Bosyu (= 

Awa), Japan," depth unknown. 

DISTRIBUTION.—Japan: from Sagami Bay to Bungo Strait, 

Honshu; off southwestern Kyushu; 88-1097 m, but most 

records are deeper than 400 m. 

Deltocyathus rotulus (Alcock, 1898) 

PLATE 24/,* 

Trochocyathus rotulus Alcock, 1898:16, pi. 2: figs. 1, la. 

Deltocyathus fragilis Alcock, 1902a:99-100; 1902c:21, pi. 2: figs. 15, 15a. 

Deltocyathus rotulus.—Cairns and Keller, 1993:245, fig. 5l [synonymy]. 

DESCRIPTION.—Corallum shaped like a shallow bowl, with 

a flat to gently rounded base and upturned edges. Largest 

Japanese specimen examined (Alb-5079) 31.1 mm in diameter 

and 7.3 mm in height; however, a specimen from the Celebes 

Sea (Alb-5582) is larger, measuring 36.4 mm in diameter. 

Circular region 7-12 mm in diameter at center of base of large 

coralla invariably worn and without costae. Otherwise, peripheral 

region of base crenulated, the theca associated with each C4 

and flanking C5 forming slightly raised radial strips about 2 

mm wide that project up to 1.5 mm beyond the calicular edge 

formed by the C^_3. Cu3 extremely narrow (about 0.2 mm), 

finely serrate ridges that are slightly recessed in valleys formed 

by ridged C4_5. Corallum white. 

Septa hexamerally arranged in 5 cycles, the fifth cycle 

usually completed at a calicular diameter of 30-32 mm. Small 

coralla often lack all 4 S5 within one or more half-systems, a 

specimen 20-30 mm in diameter having anywhere from 72-92 

septa. S1.2 moderately exsert (about 3 mm) and are independent 

septa, i.e., not fused to any adjacent septa. Each S^g 

usually bears a small palus (about 1 mm wide), which is

56 

separated from its septum by a relatively wide notch. S3 

slightly less exsert and usually do not bear pali, but, if present, 

pali are irregular in occurrence and usually quite small. S4 

equally exsert as S3, each bearing a very wide (up to 4 mm), tall 

palus, together producing a prominent and distinct palar crown 

of up to 24 lobes. In a half-system lacking all 4 S5, the S3 

within that half-system resembles an S4 and has a correspondingly 

large palus. Inner edges of paired P4 merge with inner 

edges of adjacent S3 near columella. S5 rudimentary, each 

connected to its adjacent S4 through a porous lamella in the P4 

region, but close to basal theca. Fossa shallow, containing a 

circular to elliptical columella in the form of an undercut, 

horizontal platform, through which irregularly shaped papillae 

penetrate. 

DISCUSSION.—Only two species of Deltocyathus are known 

to attain five cycles of septa in the adult state: D. magnificus 

and D. rotulus. The latter is distinguished from D. magnificus, 

as well as all other congeners, by its prominent P4 crown and 

distinctively serrate calicular margin. Its bowl-shaped corallum 

and unique columella also serve to distinguish it from D. 

magnificus. 


16331 and 80934; Alb-4970, 3, USNM 82157; Alb-4972, 18, 

CAS 80942; Alb-4973, 8, USNM 82155, 1, ORI; Alb-5079,1, 

USNM 92726; Alb-5582, 8, USNM 92727. 

TYPES.—The holotype of Trochocyathus rotulus is presumed 

to be deposited at the Indian Museum, Calcutta. Type 

Locality: North Maldive Atoll (probably Investigator station 

216), 1408-1756 m. 

Six syntypes of Deltocyathus fragilis are deposited at the 

ZMA (Coel. 1188). Type Locality: Siboga-45: 7°24'S, 

1 lS 0 ^^ (Flores Sea), 794 m. 

DISTRIBUTION.—Off southeastern Japan from off Hamamatsu, 

Honshu to the Bungo Strait (new record for Japanese 

waters); 799-1187 m. Elsewhere: Flores and Celebes Seas; 

Indian Ocean from Durban, South Africa to Sri Lanka; 

510-1986 m. 

Deltocyathus magnificus Moseley, 1876 

PLATE 2Ad.ejJi 

Deltocyathus magnificus Moseley, 1876:552-553: 1881:147-148, pi. 4: fig. 

10; pi. 13: figs. 1. 2.—Alcock, 1902c:20.—Yabe and Eguchi, 1937:138, pi. 

20: figs. 13a-c. 14a-e.—Eguchi, 1938, table 2; 1942b:126; 1965:286, 2 

figs.—Utinomi, 1965:254.—Eguchi and Miyawaki, 1975:57.—Cairns and 

Parker, 1992:27-28, pi. 7: figs, j-l; pi. 8: fig. a. 

DESCRIPTION.—Corallum discoidal, with a flat to slightly 

concave base that is not upturned at its edges. Largest Japanese 

specimen examined (Tsuchida-102) 30.0 mm in calicular 

diameter and 8.4 mm in height; however, Yabe and Eguchi 

(1937) reported a specimen 33.5 mm in diameter, and a 

specimen from the USNM collection from the Philippines 

(Alb-5444) measures 37.5 mm in diameter. Costae thin (0.2 

mm), straight, finely serrate ridges separated by rather wide, 


flat intercostal spaces 3-4 times a costal width. All costae 

project about 1.5 mm beyond calicular edge and are continuous 

with their corresponding septa above. Corallum white; according 

to Moseley (1881), the coenosarc is ochre-yellow in color, 

somewhat red toward the margin, the polyps having white 

tentacles. 

Septa hexamerally arranged in 5 complete cycles (96 septa), 

a juvenile 6.2 mm in calicular diameter having only 4 cycles, 

but a slightly larger corallum of 8.2 mm having all 96 septa. 

One large specimen (Tsuchida-102) has 2 S6, for a total of 98 

septa. S1 are the only independent septa, joining the columella 

through a wide (about 4.5 mm) palus. S2 equal in height to S1 

but slightly less wide, due to their even wider (about 6 mm) 

palus, which reaches to the columella. S3 also equal in height 

but considerably less wide, each bearing a broad palus that is 

connected to its adjacent P2 near the columella. S4 much lower 

in height than S^_3 and only 3-4 mm wide, but bordered 

internally by a broad palus that is fused to the adjacent P3 

through a porous lamella. S5 least exsert septa but equally as 

wide as an S4, each S5 usually bordered by a narrow P5, each 

pair fused to the adjacent P4 through porous lamellae. Notch 

separating P5 from S5 sometimes quite broad or poorly formed, 

making the distinction of the P5 ambiguous. Fossa absent; 

columella an elongate fusion of irregular papillae. 

DISCUSSION.—Deltocyathus magnificus is relatively easy to 

distinguish from D. rotulus, the only other congener having 

five cycles of septa (see Discussion of D. rotulus). Furthermore, 

in the Japanese region, D. magnificus consistently is 

found at shallower depths than D. rotulus (88-393 m vs 

869-1187 m). 

Material Examined.—Mew Records: Alb-4903, 1, USNM 

92728; Alb-5444, 1, USNM 62712; TM (KT9015, BS2), 10, 

USNM 92729, 2, ORI; TM (KT9309, AM8), 2, ORI; 

Tsuchida-102, 1, USNM 92730; Tsuchida-851, 2, USNM 

92731. Previous Records: Syntype of D. magnificus, BM. 

TYPES.—The single remaining syntype (Plate 24d,e) of D. 

magnificus is deposited at the BM (uncataloged). Type 

Locality: Challenger-192: 5°49'S, 132° 14^ (off Kei Islands, 

Banda Sea), 236 m. 

DISTRIBUTION.—Off Pacific coast of Japan from Kii Strait to 

Makejima, northern Ryukyu Islands; East China Sea off 

southwestern Kyushu, including Danjo Gunto; 88-422 m. 

Elsewhere: Sulu Archipelago, Moluccas, off southern Australia, 

and Bass Strait; 118-1500 m. 

Stephanocyathus Seguenza, 1864 

DIAGNOSIS.—Solitary, patellate to bowl-shaped, and free. 

Costae usually well developed, some of which are sometimes 

highly spinose< Paliform lobes usually present on all septa. 

Columella trabecular, papillose, of fused. 

TYPE SPECIES.—Stephanocyathus elegans Seguenza, 1864, 

by subsequent designation (Wells, 1936).

NUMBER 557 57 

Subgenus Stephanocyathus (Acinocyathus) Wells, 1984 

DIAGNOSIS.—Stephanocyathus having six elongate, slender 

basal spines (C^). 

TYPE SPECIES.—Stephanotrochus spiniger Marenzeller, 


Stephanocyathus (Acinocyathus) spiniger 

(Marenzeller, 1888) 

PLATE 25a-c 

Stephanotrochus spiniger Marenzeller, 1888b:20-21. 

Odontocyathus japonicus Yabe and Eguchi, 1932c: 151-152, text-figs. 1-3, pi. 

14; 1942b: 125. 

Odontocyathus spiniger.—Yabe and Eguchi, 1942b: 124-125, pi. 10: figs. 

26-28.—Eguchi, 1968:C39-40, pi. C20: figs. 12-14; pi. C23: fig. 1. 

Stephanocyathus spiniger.—Eguchi, 1965:288, 2 figs. 

Stephanocyathus (Odontocyathus) spiniger.—Utinomi, 1965:254.—Eguchi 

and Miyawaki, 1975:57.—Song, 1982:136, pi. 4: figs. 1, 2; 1991:134. 

Stephanocyathus (Odontocyathus) spinifer [sic].—Eguchi and Miyawaki, 

1975:57. 

Stephanocyathus (Acinocyathus) spiniger.—Wells, 1984:209, pi. 2: figs. 

10-13.—Cairns and Parker, 1992:26-27, pi. 7: figs, g-i [synonymy].— 

Cairns and Keller, 1993:243. 

Description of Specimen from Tosa Bay.—Corallum bowlshaped: 

47.9 mm in diameter (exclusive of costal spines) and 

26.0 mm in height, supported by 6 massive, tapered thecal 

spines (C^), each reaching over 15 mm in length and 5 mm in 

basal diameter. Lateral theca covered with equally wide, 

slightly convex, finely granulated costae. Calicular margin 

serrate, with 6 very prominent apices corresponding to the CS1 

and 6 lesser apices corresponding to the 6 CS2. Corallum 

uniformly white. 

Septa hexamerally arranged in 5 cycles with 6 additional 

pairs of S6 (= 108 septa) according to the formula: 

S1»S2>S3>S4»S5. Each S, extremely exsert (as much as 10 

mm) and bears a small (about 1.1 mm wide) paliform lobe that 

is separated from its septum by a broad notch, the lobes located 

directly adjacent to columella. Each S2 much less exsert (about 

4 mm) and bears a paliform lobe slightly larger and positioned 

higher in fossa than P1t but, like the Pv are separated by a 

broad, shallow notch from their septa, and are also located 

directly adjacent to the columella. S3_6 all about 2 mm exsert, 

except for those Sg^ directly adjacent to an S1, which are much 

more highly exsert and laterally fused to the S1 at the calicular 

margin. Each S3 bears a paliform lobe about the same size of a 

P2, but positioned higher and recessed farther from the 

columella than the P1_2. Each S4 bears 1-3 small, broad 

paliform lobes, the innermost lobe fused to its adjacent P3. S5_g 

rudimentary and lack lobes. Fossa shallow; columella papillose. 

DISCUSSION.—Stephanocyathus spiniger is easily distinguished 

from all other North Pacific species by its 6 distinctive 

costal spines, which apparently elevate the corallum above the 

substratum or stabilize it in a soft substatum. Because of its 

unusual apperance and relatively shallow depth range, it is 

sometimes found for sale in shell and curio shops. Acinocyathus 

includes about ten nominal species (see Wells, 1984; 

Cairns and Parker, 1992), ranging from the Miocene to Recent 

and throughout the Indo-West Pacific. Both Wells (1984) and 

Cairns and Parker (1992) suggested that they may all represent 

the same species; however, at least one other species is known: 

5. (A.) explanans (Marenzeller, 1904), from the southwest 

Indian Ocean. 


74944; Alb-4933, 2, USNM 92732; Tosa Bay, Shikoku, depth 

unknown, 1, USNM 92733. 

TYPES.—The holotype of S. spiniger is deposited at the 

NMW. Type Locality: Enosima (Sagami Bay), Honshu; 


The holotype of O. japonicus is deposited at the TIUS 

(40876). Type Locality: Neogene of Segoe, near Takaokamachi, 

Miyazaka-ken, southwestern Kyushu. 

DISTRIBUTION.—Japan: Neogene of southwestern Kyushu. 

Recent: Sagami Bay to Kii Strait; Tosa Bay, Shikoku; off 

southern Kyushu; off Cheju Do, Korea Strait, South Korea; 

106-783 m. Elsewhere: Philippines, Indonesia, Great Australian 

Bight, southwest Indian Ocean; 120-695 m. 

Subgenus Stephanocyathus (Odontocyathus) Moseley, 1881 

DIAGNOSIS.—Stephanocyathus having 12-18 short basal 

spines or tubercles (C1 _3), sometimes fusing into a basal rim. 

TYPE SPECIES.—Platytrochus coronatus Pourtales, 1867, by 

monotypy. 

Stephanocyathus (Odontocyathus) weberianus 

(Alcock, 1902), comb. nov. 

PLATE 25d-f 

Stephanotrochus weberianus Alcock, 1902a:101-102; 19O2c:25, pi. 3: figs. 

22,22a. 

Stephanotrochus Sibogae Alcock, 1902a: 102-103; 19O2c:25-26, pi. 3: figs. 

23, 23a. 

Stephanotrmhus sp.—Alcock, 1902c:26. 

Stephanocyathus (Odontocyathus) ixine Squires, 1958:54 [in part: pi. 8: figs. 3, 

4]. 

Stephanocyathus nobilis.—Zou, 1988:74-75 [in part: pi. 1: figs. 1-3]. 

DESCRIPTION.—Corallum bowl-shaped, with a flat to 

slightly convex base up to 26-28 mm in diameter, beyond 

which the thecal walls are inflected upward at an angle of 

75°-8O° from the horizontal. At level of thecal inflexion there 

are often 12-18 costal tubercles, each up to 2 mm long, one 

corresponding to each C,_2 and those C3 in which that 

half-system possesses 4 C5. Usually, however, the entire base 

and several mm of the lower thecal edge are completely eroded, 

which causes the tubercles to appear quite wom or to be absent. 

Tubercles often best developed on small coralla, before basal 

erosion occurs. In some coralla, thecal region corresponding to 

costal tubercles appears 'swollen' into a thick rim encircling 

the corallum base. Largest specimen examined (Alb-4969) 41.8

58 

mm in calicular diameter and 22.2 mm in height. Costae on 

thecal perimeter slightly convex and granular, the Cu2 being 

slightly wider than other costae and ridged near the calice. 


Septa hexamerally arranged in 5 cycles, the last incomplete, 

according to the formula: S1_2>S3>S4»S5. Large specimens 

(e.g., GCD > 40 mm) have only 72 septa, whereas mid-sized 

coralla of GCD 25-37 mm have only 64-70, and smaller 

coralla 18-25 mm GCD have 50-68 septa. No specimens were 

found with only 48 septa. Half-systems within a single 

specimen quite variable in development, some having 0, 2, or 

4 S5. S^ highly exsert (about 5 mm), each bearing a low, 

oblique to almost horizontally projecting paliform lobe that 

extends into the columellar region. S3 slightly less exsert 

(3.0-3.5 mm), each also bearing a small paliform lobe but 

positioned slightly higher in fossa and slightly farther from 

columella than P^. S4 smaller still (only about 2.7 mm exsert), 

each bearing a wide paliform lobe which contributes to a palar 

crown that is located higher and farther recessed from the 

columella than the P3 crown. Inner edges of P4 usually bend 

toward and are fused to P3 near columella. S5 rudimentary, 

extending as narrow lamellae only partially down inner theca. 

When S5 are absent from a half-system, the S3 bears a wide 

paliform lobe equal in size and position to that of a P4. All septa 

have very finely sinuous inner edges and virtually smooth 

septal faces. All paliform lobes separated from their respective 

septa by broad, shallow notches. Fossa moderately deep, 

containing a papillose columella. 

DISCUSSION.—Stephanocyathus weberianus is very similar 

to the type-species of the subgenus, 5. (O.) coronatus 

(Pourtales, 1867), which is known only from the western 

Atlantic at 543-1250 m (Cairns, 1979). A detailed comparison, 

however, reveals that S. coronatus invariably has only 12 large, 

often complexly ornamented costal tubercles, whereas S. 

weberianus has 12-18 rather simple tubercles that are usually 

quite worn. The base of S. coronatus is usually convex and 

coarsely dentate (C^_2)\ that of 5. weberianus is flat and eroded. 

The corallum of 5. coronatus is, in general, higher in proportion 

to its diameter (H:D) than S. weberianus in proportion to its 

diameter, producing a more slender corallum with less septa. 

For instance, coralla containing only 48 septa are common for 

S. coronatus but quite rare for S. weberianus, the latter often 

having half-systems with a full complement of 4 S5. Finally, 

the S3_5 of S. coronatus are less exsert in relation to their S,_2 

than in S. weberianus. 

Stephanocyathus weberianus is also similar to 5. nobilis 

(Moseley, 1873), known only from the Atlantic and western 

Indian Oceans at 609-2200 m (Cairns and Keller, 1993). Zou 

(1988) synonymized the two species, considering S. nobilis to 

be a cosmopolitan species in the Atlantic, Pacific, and Indian 

Oceans. However, when closely compared, S. nobilis differs 

significantly in having very inconspicuous paliform lobes and 

in lacking costal tubercles, having only a series of large costal 

spines on the C^. Furthermore, its costal spines are limited to 


the twelve C1-2, even when a half-system has all 4 S5, whereas 

additional costal tubercles are invariably present in specimens 

of 5. weberianus having the same number of septa. Stephanocyathus 

nohilis also has an evenly rounded, convex base, and, 

like S. coronatus, much more exsert S^ in relation to the 

remaining S3.5. 

The difference between 5. weberianus and S. sibogae as 

described by Alcock (1902a) was that the former had a swollen 

ring around its base, the latter, a worn base with atrophied 

costal tubercles. It is now clear that these conditions are within 

the range of variation for one species: S. weberianus. 



USNM 92736; Alb-4957, 4, CAS 1102; Alb-4958, 1, USNM 



92740; Alb-4975, 1, USNM 92741; TM (KT9202, ATI), 2, 

USNM 92742; TM (KT9202, YT6), 10, USNM 92743,4, ORI. 

Previous Records: Holotype of 5. weberianus, ZMA; Alb- 

5445, 46, USNM 46819 (5. ixine of Squires, 1958). 

TYPES.—The holotype of S. weberianus is deposited at the 

ZMA (Coel. 1322). Type Locality: Siboga-2%A: 8°43.1'S, 

127° 16.7'E (Timor Sea), 828 m. 

The holotype of 5. Sibogae is presumed to be deposited at the 

ZMA, but is not listed by van Soest (1979). Type Locality: 

Siboga-%%: 0°34.6TSf, \\9°Q%.5 f E (Makassar Strait), 

1301 m. 

DISTRIBUTION.—First reports from off Japan: Off southeastern 

Honshu; Bungo Strait; off Koshiki I., southwestern 

Kyushu; northen Ryukyu Islands (Osumi Shoto and Tokara 

Retto); 715-1302 m. Elsewhere: South China Sea, Sulu Sea, 

Makassar Strait, Banda and Timor Seas; 206-1301 m. 

Conotrochus Seguenza, 1864 

DIAGNOSIS.—Solitary, ceratoid to trochoid, free or attached 

through a small pedicel, which is often augmented by a lateral 

thecal attachment. Theca thick, but covered with epitheca; 

costae usually obscure. Septa exsert, but upper outer septal 

edges join the theca below upper thecal edge, forming an exsert 

calicular rim. Pali absent; columella prominent, composed of 

elongate, twisted lamellar elements. 

TYPE SPECIES.—Conotrochus typus Seguenza, 1864, by 


Conotrochus funicolumna (Alcock, 1902) 

PLATES 24/, 25g-l 

Ceratotrochus (Conotrochus) funicolumna Alcock, 1902a:93; 1902c: 11-12, 

pi. 1: figs. 6, 6a.—Faustino, 1927:66, pi. 9: figs. 7, 8.—Yabe and Eguchi, 

1942b:117. pi. 9: fig. 11.—Eguchi, 1968:C38-39.—Zou, 1988:77, pi. 5: 

figs. 1, la. 

Conotrochus funicolumna.—Cairns, 1984:14, pi. 2: figs. IJ. 

Conotrochus sp. cf. C. funicolumna.—Cairns and Parker, 1992:22, pi. 6: 

figs, c, f. 

Ceratotrochus hiugaensis Yabe and Eguchi, 1942b:117, 146, pi. 9: fig.

NUMBER 557 59 

10a,b.—Eguchi, 1965:288, 2 figs. 

Ceratotrochus (Conotrochus) parahispidus Yabe and Eguchi, 1942b: 118, 

147-148, pi. 9: fig. 12a,b.—Eguchi, 1965:288, 2 figs; 1968:C39.—Eguchi 

and Miyawaki, 1975:57. 

DESCRIPTION.—Corallum trochoid to ceratoid, slightly 

curved, and usually free, but sometimes attached through a 

slender pedicel or attached along a small section of lower theca. 

One illustrated specimen (Plate 25/,/) is 13.6 x 13.0 mm in 

calicular diameter and 17.8 mm in height. Alcock's (1902c) 

figured syntype is 11.9 x 11.7 mm in calicular diameter and 

13.1 mm in height. Theca thick but invariably worn, revealing 

coarsely granular costae covered by a thin epitheca. Upper 

theca projects 0.5-0.7 mm above upper, outer septal edges as 

a continuous rim, which is characteristic of the genus. Corallum 

white. 

Septa hexamerally arranged in 4 complete cycles: 

S1_2>S3>S4. S^ only slightly exsert above encircling thecal 

rim and have straight, vertical inner edges that join the 

columella only deep within fossa. S3 about 80% width of S^ 

and also have straight inner edges that fuse with the columella 

only lower in fossa. S4 one-quarter to one-third width of S3 and 

have entire inner edges that do not attain the columella. All 

septal faces bear very low granules, giving the impression of 

smooth septal faces. Fossa shallow. Columella prominent, 

composed of an elliptical mass of slightly swirled lamellar 

elements, the uppermost 5 mm remaining free of septal fusion. 

DISCUSSION.—Conotrochus parahispidus (Yabe and 

Eguchi, 1942b) was originally distinguished from C. funicolumna 

by having a narrower (GCD = 9.5-14 mm), elongate 

corallum, all other characters being similar. Having examined 

two syntypes of C. parahispidus, I conclude that they fall 

within the range of corallum shape variation of C. funicolumna 

and therefore suggest its synonymy. Likewise, although not 

examined, the holotype of Ceratotrochus hiugaensis Yabe and 

Eguchi, 1942b (GCD = 9.0 mm) appears to be simply a juvenile 

C. funicolumna. 

At least three other species of Conotrochus are known: C. 

typus Seguenza, 1864 (Miocene of Italy); C. elongatus Yabe 

and Eguchi, 1942b (Plio-Pleistocene of Ryukyu Islands); and 

C. brunneus (Moseley, 1881) (Recent, Indo-West Pacific). 

Conotrochus funicolumna differs from the latter, the only other 

Recent species, in having a larger adult corallum with more 

septa (^48 septa vs

60 

A. matricidus by its broader and shorter corallum, its thinner 

S^, and its 1:1 correspondence of septa and costae. It is quite 

similar to A. atlanticus Zibrowius, 1980 (known only from the 

northeast Atlantic at 450-1716 m), the Pacific species differing 

primarily in having notched septa near the thecal edge. It might 

also be confused with Conotrochus funicolumna, but is usually 

easily distinguished by its residual parent fragment that 

remains attached to the base. 


USNM 92704. 

TYPES.—The "numerous" syntypes of A. recidivus reported 

by Dennant (1906) cannot be located (see Zibrowius, 1980). 

Type Localities: Off Cape Jaffa and Neptune Island, South 

Australia, 165-190 m. 

DISTRIBUTION.—Japan: Bungo Strait (Okino I.) off Shikoku 

(new record for North Pacific); 741 m. Elsewhere: South 

Australia, Tasmania, Macquarie Ridge, off Madagascar, 128— 

1000 m. 

Aulocyathus matricidus (Kent, 1871) 

PLATES 26C -g, MJb-d 

Flabellum matricidum Kent, 1871:276, pi. 23: fig. 2a-c. 

Fragilocyathus conotrochoides Yabc and Eguchi, 1932a:388, 389, fig. 1; 

1941b:101; 1942b:116, 145. pi. 9: fig. 15.—Eguchi, 1965:288. 4 figs — 

Eguchi and Miyawaki, 1975:57.—Zibrowius, 1980:105, 107. 

Aulocyathus cf. matricidum.—Yabe and Eguchi, 1941b: 101. 

Aulocyathus cf. matricidus.—Yabe and Eguchi, 1942b: 112, 116. 

DESCRIPTION.—Corallum elongate-conical, up to 31.0 mm 

in length but with a maximum calicular diameter of 8.5 mm 

(Yabe and Eguchi, 1942b). Illustrated specimen (TM (KT7802, 

Z4)) only 16.4 mm in length and 6.2 mm in diameter, tapering 

gradually to a broad pedicel 2.1 mm in diameter. Coralla appear 

to reproduce predominantly by budding from corallum fragments 

following the longitudinal fission of a parent corallum. 

Theca rough and striate, possessing twice the number of 

longitudinal ridges as septa. The true costae, those ridges 

corresponding in position to the septa, are 0.18-0.20 mm wide; 

whereas, between each costa occurs another narrower ridge 

0.10-0.11 mm in width. Both ridges bear hollow, conical, 

apically inclined granules, many of which are abraded apically 

revealing a hollow interior. Corallum light brown. 

Septa hexamerally arranged in 4 cycles, the fourth cycle 

rarely complete. Each system of a typical corallum contains 1 

S2,2 S3, and only 2 S4, resulting in a corallum total of 36 septa. 

As the corallum increases in size, additional pairs of S4 are 

added, completing various systems. Yabe and Eguchi (1942b) 

reported specimens of GCD = 6-8.5 mm to have 36-49 septa; 

the holotype of F. conotrochoides (GCD = 6.6 mm) has 44 

septa; specimens reported herein (GCD = 5.2-7.5 mm) range 

from 38-40 septa; and a syntype of F. matricidum (GCD = 7.4 

mm) has 40 septa. S1 only very slightly exsert (0.5 mm) and 

have vertical, straight to slightly sinuous inner edges, which 

become quite thick (up to 0.7 mm) and coarsely granular to 

tuberculate at their lower edges. S2 about three-quarters width 


of an S1 and also have thickened, tuberculate inner edges, 

sometimes bearing a narrow paliform lobe or columellar 

process that extends upward into the fossa. S3 that are flanked 

by a pair of S4 are about three-quarters width of an S2 and have 

thick inner edges; S3 that stand alone are only half width of an 

S2 and have coarsely dentate, narrow inner edges. S4 similar in 

shape to those S3 unflanked by S4. Upper outer edges of septa 

not notched near thecal edge. Fossa deep. Columella rudimentary, 

composed of the tuberculate, widened lower inner edges 

of the SU2 and often a single rudimentary tubercle. 

DISCUSSION.—The holotype of F. conotrochoides is a 

well-preserved specimen 30.9 mm in height and 6.6 mm in 

calicular diameter, having 44 septa. It is the only specimen 

known to me that does not evidence asexual development from 

a parent fragment, instead having a slender pedicel 0.9 mm in 

diameter including the six original protoseta. It is also 

distinctive in bearing 14 brown longitudinal stripes of various 

thickness on its theca, these stripes not associated with any 

particular septal cycle. Despite these peculiarities, F. conotrochoides 

is considered to be a junior synonym of Aulocyathus 

matricidus. 

Three other species occur in the genus: A. juvenescens 

Marenzeller, 1904a (known only from off Tanzania, 302-463 

m); A. recidivus (Dennant, 1906) (S. Australia, New Zealand, 

Madagascar, off Japan, 128-1000 m); and A. atlanticus 

Zibrowius, 1980 (northeast Atlantic, 450-1716 m). Aulocyathus 

matricidus is unique in having pseudocostae in 

addition to costae and tuberculate inner edges of S1-2. It is 

more slender and has a less well-developed columella than A. 

recidivus and A. atlanticus, but has a broader corallum than A. 

juvenescens (Plate 26h,i). Among the four species, it has the 

shallowest depth range (84-207 m). 


Z4), 1, USNM 92703; TM (KT7414, B2), 1, ORI. Previous 

Records: Syntypes of F. matricidum, BM. Reference Specimens: 

4 syntypes of A. juvenescens, ZMB 5064 and 7032; 

holotype of F. conotrochoides, TIUS; Toyama Bay, 2, TIUS 

58223 (Yabe and Eguchi, 1942b) (Plate 42c,d). 

TYPES.—Two syntypes (Plate 26c,e,f) of Flabellum matricidum 

are deposited at the BM (1862.7.16.72). Type Locality: 

"Off Japan," depth not reported, but Zibrowius (1980) 

gives 84 m, this information obtained from the label with the 

syntypes. 

The holotype (Plate 42b) and paratype of Fragilocyathus 

conotrochoides are deposited at the TIUS (50084 and 50083, 

respectively). Type Locality: Soyo Maru-352: 33 o 39'50"N, 

135°O6'3O"E (Kii Strait, Honshu), 154 m (elucidated by Yabe 

and Eguchi, 1942b: 145, footnote). 

DISTRIBUTION.—Tsugara Strait; Pacific coast of Honshu 

from Suruga Bay to Kii Strait; Sea of Japan from Toyama Bay 

to Wakasa Bay; 84-207 m. 

Desmophyllum Ehrenberg, 1834 

DIAGNOSIS.—See Part 1.

NUMBER 557 61 

Desmophyllum dianthus (Esper, 1794) 


Lophelia Milne Edwards and Haime, 1849 


Lophelia pertusa (Linnaeus, 1758) 


Anomocora Studer, 1878 

DIAGNOSIS.—Corallum subcylindrical and free, with a 

tendency to bud new corallites at random from thecal wall, the 

buds subsequently losing their organic connection. Before bud 

detachment the corallum is colonial. Wall thin. Columella 

trabecular, multiple irregularly shaped paliform lobes usually 

present on S^_3. Tabular endothecal dissepiments common and 

widely spaced. 

TYPE SPECIES.—Coelosmilia fecunda Pourtales, 1871, by 

monotypy. 

Anomocora sp. 

Parasmilia fecunda Yabe and Eguchi, 1932b:443. 

Anomocora fecunda.—Eguchi, 1965:290, 2 figs.; 1968:C42, pi. CIO: figs. 1-5; 

?pl. C20: figs. 10, 11; ?pl. C23: fig. 3.—Cairns, 199la: 19. 

DISCUSSION.—No additional specimens of Anomocora are 

reported in the present study and the two specimens of A. 

fecunda reported by Eguchi (1968) are not available for study. 

Eguchi (1968) did not describe his Japanese specimens, but his 

specimen (#701) illustrated on plate CIO does appear to be an 

Anomocora; the other specimen (#741) cannot be identified 

from the illustrations. Specimen #701, from off Chigasaki, 

Sagami Bay (100 m), has relatively large corallites 12-16 mm 

in calicular diameter, thin, ridged costae; an incomplete fifth 

cycle of septa; prominent paliform lobes; and sparse budding. 

I agree with Zibrowius (1980) that the Japanese specimens are 

not A. fecunda, which appears to be restricted to the Atlantic 

(Caims, 1979). The Japanese specimens have larger corallites, 

more septa per corallite, more prominent paliform lobes, and 

less frequent budding. Anomocora has been reported several 

times in the Indo-West Pacific by: Gardiner and Waugh (1939) 

Red Sea; Marenzeller (1904a) off Sumatra; Cairns (1984) 

Hawaiian Islands; and Caims (1991a) off the Galapagos. 


Records: Reference specimens of A. fecunda from the 

western Atlantic (Cairns, 1979). 

DISTRIBUTION.—Sagami Bay; 50-100 m. 

Coenosmilia Pourtales, 1874 

DIAGNOSIS.—Corallum colonial, small bushy colonies 

formed by extratentacular budding of ceratoid corallites just 

below calice edge or from a commom basal coenosteum. 

Corallum firmly attached. Columella a solid fusion of 

elements; paliform lobes absent. Tabular endothecal dissepiments 

present. 

DISCUSSION.—Coenosmilia and Anomocora are quite similar 

in morphology and have been synonymized by Zibrowius 

(1980). Coenosmilia, however, can be distinguished by having 

an attached corallum, more regular budding, a more solid 

columella, and the absence of paliform lobes. 

TYPE SPECIES.—Coenosmilia arbuscula Pourtales, 1874, by 

monotypy. 

Coenosmilia sp. cf. C. arbuscula Pourtales, 1874 

PLATE 27a,* 

DESCRIPTION OF SPECIMENS FROM TM (KT9202, YS2).—A 

well-preserved corallum is ceratoid: 7.6 x 6.7 mm in calicular 

diameter, 3.1 mm in pedicel diameter, and 24 mm in height, 

bearing no buds. A second poorly preserved specimen is only 

slightly smaller and has 3 buds equally spaced just below the 

caliclar edge. C[_2 low, finely serrate ridges; C3 slightly less 

prominent ridges; C4 broad and unridged. Corallum white. 

Septa hexamerally arranged in four cycles, but with one pair 

of S4 missing (= 46 septa), according to the formula: 

S1>S2»S3>S4. S1 little exsert (0.8 mm), and not wide (about 

0.8 mm wide), having vertical, slightly sinuous inner edges that 

merge with the columella. S2 of same exsertness, less wide (0.6 

mm), and have inner edges that also merge with the columella. 

S3 less exsert and only about half width of S,_2, their inner 

edges not coming close to the columella. S4 rudimentary, each 

composed of a row of small spines. Pali and paliform lobes 

absent. Fossa deep, containing a large trabecular columella. 

Tabular endothecal dissepiments present. 

DISCUSSION.—The Japanese specimens are extremely similar 

to C. arbuscula, including aspects of their budding pattern, 

size, costal shape, and number of septa. The two specimens at 

hand differ only in having a deeper fossa and smaller S3 than C. 

arbuscula. Coenosmilia arbuscula is known only from the 

North Atlantic from 109-622 m (Cairns, 1979; Zibrowius, 

1980); before a confident identification can be made, more 

North Pacific specimens will need to be examined and 

compared. 


YS2), 2, USNM 92709. Reference Specimens: Specimens 

from Atlantic reported by Cairns (1979) and Zibrowius (1980), 

including the syntypes (MCZ). 

DISTRIBUTION.—A seamount (Yaku-Shinsone) in the northern 

Tokara Retto, Ryukyu Islands; 238-240 m. 

Phyllangia Milne Edwards and Haime, 1848c 

DIAGNOSIS.—Colonial, extratentacular budding forming 

reptoid to plocoid colonies, often basally united by thick 

coenosteum. Inner edges of S, smooth, those of higher cycle 

septa smooth to finely dentate. Sparse endotheca present. 

Columella rudimentary; P2 or P3 sometimes present.

62 

TYPE SPECIES.—Phyllangia americana Milne Edwards and 

Haime, 1849, by subsequent designation (Milne Edwards and 

Haime, 185Oa:xliv). 

Phyllangia hayamaensis (Eguchi, 1968) 

Astrangia hayamaensis Eguchi, 1968:C26, pi. C27: figs. 8-10; pi. C28: fig. 

7.—Wells, 1983:233. 

Phyllangia hayamaensis.—Cairns, 1991 a: 18. 

DIAGNOSIS.—This species is known only from the holotype 

colony, which is about 7.4 cm wide and consists of an 

encrusting corallum of about 40 corallites. Unfortunately this 

specimen is not available for study and thus the following 

diagnosis is taken from the original description. Corallites 

cylindrical, up to 7 x 6 mm in diameter and 7 mm in height, and 

bud from a thick, common basal coenosteum. Theca thin and 

covered with small granules. Septal symmetry octameral, each 

coral lite having 8 primary, 8 secondary, and 2-5 pairs of 

tertiary septa, for a total of 20-26 septa. Primary septa thick, 

exsert, and have entire (smooth) vertical inner edges. Secondary 

septa only slightly less wide but have dentate inner edges. 

S3 rudimentary and also have dentate inner edges. A crown of 

8 paliform lobes (P2) occur before the secondary septa. 

Columella spongy. 

DISCUSSION.—Wells (1983) implied that A. hayamaensis 

belonged in the genus Phyllangia by stating that it was a 

"related species" to Phyllangia consagensis. I concur that the 

species belongs to Phyllangia, but suggest that it is probably 

more closely related to the eastern Pacific P. dispersa Verrill, 

1864. These two species have approximately the same calicular 

diameter, and both have highly exsert primary septa, rudimentary 

P2, and a similar growth form. Phyllangia hayamaensis 

appears to differ in having only about 20-26 octamerally 

arranged septa, whereas P. dispersa usually has 48 hexamerally 

arranged septa. Examination of additional specimens from off 

Japan is required to understand this species and to properly 

compare it with its eastern Pacific congener. 

MATERIAL EXAMINED.—New Records: None. Reference 

Specimens: P. dispersa, Paitilla Point, Canal Zone, Gulf of 

Panama, 1 colony, USNM 83526. 

TYPES.—The holotype of P. hayamaensis is deposited in the 


(#640). Type Locality: Kamegisho One, Sagami Bay, 5.5 m. 


RhizosmiUa Cairns, 1978 

DIAGNOSIS.—Phaceloid coral la formed by extratentacular 

budding from a thin common basal coenosteum. Corallite bases 

increase in diameter by adding exothecal dissepiments over 

raised costae producing concentric rings of partitioned chambers 

resembling polycyclic development in cross section. 

Vesicular endothecal dissepiments present. Paliform lobes 

present before penultimate cycle (usually P3). Columella 

variable, including papillose, lamellar, and fascicular. 


TYPE SPECIES.—RhizosmiUa gerdae Cairns, 1978, by original 

designation. 

RhizosmiUa sagamiensis (Eguchi, 1968), comb. nov. 

PLATE 27C -e 

Coenocyalhus sagamiensis Eguchi, 1968:C34, pi. C10: figs. 6, 7. 

DESCRIPTION OF SPECIMENS FROM Alb-4944.—One corallum 

consists of a phaceloid clump of 13 interconnected 

corallites (Plate 27c), the other corallum is a large solitary 

specimen (Plate 21d,e). Corallites of colony connected basally 

by a thin encrusting coenosteum. Corallites ceratoid, the largest 

measuring 13.3 x 10.3 mm in calicular diameter, 20.1 mm in 

height, and 5.2 mm in pedicel diameter. Pedicel reinforced by 

thin raised costae overlain with exothecal dissepiments, this 

process usually visible on any large specimen. Costae equal in 

width (0.5-0.6 mm) and finely granular, the C^_3 being 

slightly ridged near calice. Lower half of theca of largest 

(solitary) specimen also bears fine transverse rugae. Corallum 

white, except for brown crescent-shaped bands that parallel the 

septal edges of S^. 

Septa generally hexamerally arranged in 4 complete cycles 

according to the formula: S1>S2>S3>S4, but larger corallites 

have additional septa and paliform lobes, the largest calice 

having 68 septa and 16 paliform lobes. S1 highly exsert (up to 

3.4 mm), relatively thick, and have straight, vertical inner edges 

that almost attain the columella. S2 up to 2.5 mm exsert, 

three-quarters width of S,, and also have straight inner edges. 

S3 least exsert septa (about 1.1 mm), about three-quarters width 

of S2, and have slightly sinuous inner edges. A wide (1.1-1.3 

mm), lamellar paliform lobe occurs before each S3, separated 

from its adjacent septum by a deep, narrow notch. S4 less wide 

but more exsert (1.6-2.0 mm) than S3. Each pair of S4 flanking 

an S1 or S2 is fused with that septum at the calicular edge to 

form a 3-septum calicular extension resulting in a highly serrate 

calicular edge. Fossa of moderate depth, containing the P3 

crown of 12-16 lobes and a central fascicular columella 

consisting of several slender, loosely twisted elements. 

DISCUSSION.—The holotype, which was the only known 

specimen of this species, is unavailable for study. Eguchi's 

illustration of the holotype shows a worn or rejuvenescent 

corallum of three corallites. The specimens described above are 

consistent with Eguchi's description and depth range and thus 

believed to be conspecific. 

Three other species of RhizosmiUa are known: R. maculata 

(Pourtales, 1874); R. gerdae Cairns, 1978; and R. robusta 

Cairns in Caims and Keller, 1993. R. sagamiensis is most 

similar to the type species, R. gerdae, known only from the 

western Atlantic at 123-287 m, both species having four cycles 

of septa and approximately the same calicular diameter. 

RhizosmiUa sagamiensis differs in having ceratoid corallites, a 

fascicular columella, and highly exsert, pigmented S.^. 

MATERIAL EXAMINED.—New Records: Alb-4944, 1 col-

NUMBER 557 63 

ony and 1 corallite, USNM 92705; TM (KT9202, YT2), 1 

corallite, USNM 92706. 

Types.—The holotype is deposited in the Biological Laboratory 

of the Imperial Household, Tokyo (#657). Type Locality: 

Amadaiba, Okino-Kannonzukadashi, Sagami Bay, 60- 

80 m. 

DISTRIBUTION.—Known only from off Japan: Sagami Bay; 

Kagoshima Bay, Kyushu; Colnett Strait, Osumi Shoto, 


Dasmosmilia Pourtales, 1880 

DIAGNOSIS.—Solitary, ceratoid to turbinate, free. Parricidal 

budding common. Theca and septa very thin. Paliform lobes 

present before penultimate cycle of septa and occasionally also 

before lower cycle septa. Columella trabecular. Endotheca 

sparse or absent. 

TYPE SPECIES.—Parasmilia lymani Pourtales, 1871, by 

subsequent designation (Wells, 1933). 

Dasmosmilia pacifica (Yabe and Eguchi, 1932), comb. nov. 

PLATES 27/-I, 4 \f,g 

Goniocyathus pacificus Yabe and Eguchi, 1932a:389, text-fig. 2; 1932b:443; 

1942b: 122, 152-153, pi. 10: figs. 15, 16.—Eguchi, 1965:286, 2 figs.— 

Zibrowius, 1980:70. 

Caryophyllia pacifica—Wells, 1956:F422. 

DESCRIPTION.—Corallum fragile, due to extremely thin 

theca and septa. AH specimens examined had asexually 

generated from a wedge-shaped fragment of parent corallum, 

often resulting in somewhat irregular septal development and 

symmetry. Largest specimen examined (syntype) 14.9 mm in 

GCD. Corallum above base ceratoid, with a circular to 

irregularly shaped calice. Each primary septum, along with its 

2 adjacent higher-cycle septa, are fused at calicular edge and 

project 1.0-1.5 mm above the edge, producing a serrate 

margin. Costae slightly convex and coarsely granular, the 

12-24 primary costae ridged. Corallum white, with a slight 

brownish tint near the calice. 

Septal symmetry irregular, the number of septa apparently 

correlated with calicular diameter. Among the few specimens 

examined, there are coralla with 12-24 primary septa, 12-24 

secondary septa, 24-48 tertiary septa, one corallum (large 

syntype, TIUS 50086) also having a pair of quaternary septa, 

for a total of 98 septa. Primary septa exsert as much as 2.2 mm 

and have vertical, sinuous inner edges, several of which clearly 

bear tall twisted fascicular columella elements. Secondary 

septa least exsert and only about 40% width of a primary. Each 

secondary septum bears a wide paliform lobe (about 1.3 mm), 

together forming a crown of 12-18 lobes. Tertiary septa 

slightly more exsert but less wide than secondaries, becoming 

rudimentary lower in fossa. When pairs of quaternary septa 

occur within a half-system, they resemble tertiaries in size, and 

the flanked tertiary is slightly enlarged in width and adds a 

paliform lobe. Endothecal dissepiments not observed. 

DISCUSSION.—Vaughan and Wells (1943) and Wells (1956) 

placed the monotypic Goniocyathus Yabe and Eguchi, 1932a in 

synonymy with Caryophyllia; however, Zibrowius (1980) 

suggested that it might fall closer to Dasmosmilia. After 

comparing the Japanese specimens reported herein with typical 

D. lymani, I agree with Zibrowius that Goniocyathus is a junior 

synonym of Dasmosmilia. In fact, the differences between D. 

pacifica and D. lymani are quite minor. Dasmosmilia pacifica 

differs in having exsert primary septa, always fused with 

adjacent higher-cycle septa to form a serrate calicular margin, 

whereas the primary septa of D. lymani are not very exsert and 

not fused with adjacent septa near the calice. Also, the corallum 

of D. pacifica is more open and the septa more widely spaced 

than in D. lymani. 

The thin theca of this genus facilitates its mode of asexual 

budding. 


USNM 92710; Alb-5055, 1, USNM 92711; YO70-1009, 1, 

ORI. Previous Records: 3 syntypes, TIUS. 

TYPES.—The syntypes of G. pacificus are deposited at the 

TIUS (50086, 50097)(Plate 4\f,g). Type Locality (vide Yabe 

and Eguchi, 1942b): Soyo Maru-198: 34°17'45"N, 

137 o 04'45 / ^(off Ise Bay, Honshu), 168 m. 

DISTRIBUTION.—Known only from the Pacific coast of 

Japan from off Boso Hanto, Chiba-ken, Honshu to southeastern 

Kyushu; 168-355 m. Pleistocene of Ryukyu Islands (Yabe and 

Eguchi, 1932b). 

Goniocorella Yabe and Eguchi, 1932a 

DIAGNOSIS.—Colonial, extratentacular budding forming 

bushy colonies. Branch anastomosis common, the integrity of 

the corallum further increased by numerous slender, tubular 

coenosteal bridges. No pali or columella. Tabular endothecal 

dissepiments common and widely spaced. 

TYPE SPECIES.—Pourtalosmilia dumosa Alcock, 1902, by 


Goniocorella dumosa (Alcock, 1902) 

PLATE 27/ 

Pourtalosmilia dumosa Alcock, 1902c:36-37, pi. 5: fig. 33. 

Goniocorella dumosa.—Yabe and Eguchi, 1932a:389-390; 1936:167; 

1943:494-496, figs. 1, 2; 1942b:162, 163.—Eguchi, 1965:291, 2 figs.; 

1968:C43, pi. C9: figs. 11. 12.—Cairns, 1982:31-34, pi. 9: figs. 7-9; pi. 10: 

figs. 1. 2 [synonymy].—Cairns and Keller. 1993:250. fig. 6E.—Song, 

1991:134-135, pi. 1: fig. 3; pi. 2: figs. 4-7. 

Goniocorella sp. aff. G. dumosa.—Eguchi and Miyawaki. 1975:58. 

DESCRIPTION.—Colony bushy, budding often at right angle 

to parent branch. Colony reinforced by slender (about 2 mm in 

diameter), hollow, tubular coenosteal bridges, which unite 

adjacent branches. Branches cylindrical, 3-5 mm in diameter, 

each bearing a terminal calice of equal diameter. Costae 

inconspicuous. Corallum white or light brown.

64 


to the formula: S1»S2>S3. St not very exsert and quite narrow 

(only about 0.6 mm wide), with straight, vertical inner edges 

that extend to uppermost dissepiment. S2 much thinner (only 

about 0.2 mm wide); S3 smaller still (only about 0.1 mm wide), 

both S2 and S3 extend deep into fossa. Fossa usually deep and 

vacuous, bordered laterally by inner edges of narrow septa and 

basally by a horizontal dissepiment. Thin, tabular endothecal 

dissepiments occur every 2-10 mm, giving the corallum a very 

low density. 

DISCUSSION.—This species is well described and illustrated 

by Yabe and Eguchi (1936) and Cairns (1982). Goniocorella is 

monotypic. 

MATERIAL EXAMINED.—New Records: Sagami Bay, 

depth unknown, 1 colony, USNM 92708, 1 fragment, ORI; 

Okinose, Sagami Bay, 366 m, several colonies, ZMC; off Izu 

Peninsula, Honshu, depth unknown, 1 colony, USNM 92707. 

TYPES.—Several syntypes of P. dumosa are deposited at the 

ZMA (Coel. 1097). Another syntype is presumed to be 

deposited at the Indian Museum, Calcutta (see van Soest, 1979; 

Cairns, 1982). Type Localities: Siboga stations 156, 259, 

Banda Sea, Indonesia, 469-487 m. 

DISTRIBUTION.—Off Honshu from Sagami Bay to Owase; 

off southwestern Shikoku; off Ullung Do, South Korea, Sea of 

Japan; 100-366 m. Elsewhere: Off South Africa; Banda Sea; 

New Zealand region; 100-760 m. 

Family TURBINOLIIDAE 

Notocyathus Tenison-Woods, 1880 

DIAGNOSIS.—Corallum solitary, cylindro-conical or cuneiform, 

with a pointed, unattached base; transverse division 

absent. Theca imperforate; costae serrate and correspond to 

septa. Septa highly exsert. Pali before all but last cycle, but 

Pu2 suppressed in adult; pairs of P3 unite in V-shaped 

structures in each system. Columella papillose. 

TYPE SPECIES.—Caryophyllia viola Duncan, 1865, by 

subsequent designation (Felix, 1927). 

Notocyathus venustus (Alcock, 1902) 

PLATE 27*./ 

Citharocyathus venustus Alcock, 1902b: 119; 1902c:22, pi. 3: figs. 19, 

19a.—?Yabe and Eguchi. 1932b:443,444. 

?Citharocyathus conicus forma venustus.—Wells. 1984:214, pi. 4: figs. 2-5. 

Notocyathus venustus.—Cairns, 1989a:27-28. pi. 12: figs, c-h [synonymy]. 

DESCRIPTION.—Lower half of corallum conical (ceratoid), 

but upper half a constant-diameter cylinder. Largest Japanese 

specimen examined 7.15 x 6.95 mm in calicular diameter and 

12.6 mm in height, which is the largest reported specimen. C, 

independent C2 trifurcate very near base epicenter, resulting in 

2 C3 and a medial C2. C3 also trifurcate within 2 mm of 

epicenter, each C3 producing 2 C4 and a medial C3. C^ 

slightly broader than C^. All costae bear a uniserial row of 

teeth as well as smaller lateral granules that project into 


intercostal spaces. Intercostal furrows deep and narrow, which 

is characteristic of the family. Corallum white. 


to the formula: S1>S2>S3»S4. S1 highly exsert (about 2.2 

mm) and have straight, vertical inner edges that fuse with the 

columella deep in fossa. S2 less exsert (about 1.6 mm) and 

about two-thirds width of an S,, their inner edges fused to the 

V-shaped palar fusion of adjacent P3. S3 slightly less exsert and 

about two-thirds width of an S2, each S3 separated by a broad 

notch from a tall, lamellar palus about 0.5 mm wide. Inner 

edges of each P3 pair fused near columella, producing a 

V-shaped paliform structure in each of the 6 systems. S4 small, 

only about one-third as exsert and two-thirds as wide as an S3. 

Fossa lacking, the columellar and palar elements rising well 

above calicular edge. Columella papillose, composed of 7-12 

fused elements. 

DISCUSSION.—Yabe and Eguchi (1942b, 1946) did not 

consider N. venustus and N. conicus to be distinct species, but 

after examining the types of both species, I (Cairns, 1989a) 

listed several characters that consistently differentiate the two. 

To summarize, N. venustus has (1) a nearly circular calice 

(GCD:LCD = 1.02-1.05 vs 1.05-1.22 for N. conicus), (2) 

more exsert S1-3 and less exsert S4, and 3) no fossa. 

Furthermore, the inner septal edges of N. venustus are straight, 

whereas the inner edges of the Su3 ofN. conicus are sinuous. 

Notocyathus venustus was described and illustrated in 

greater detail by Cairns (1989a); however, his distributional 

record of Pleistocene from the Ryukyu Islands attributed to 

Yabe and Eguchi (1942b) was an error. 

MATERIAL EXAMINED.^/Vew Records: TM (KT9015, 

BS2), 12, USNM 92778; TM (KT9309, AM8), 4, ORI. 

Previous Records: 3 syntypes of C. venustus, ZMA; C. 

conicus forma venustus of Wells (1984), USNM 73968. 

TYPES.—Three of the four syntypes of C. venustus are 

deposited at the ZMA (Coel. 1244); the location of the fourth 

syntype is unknown (Van Soest, 1979). Type Locality: 

Siboga-59: 10°22.7'S, 123° 16.5^ (Savu Sea, Indonesia), 

390 m. 

DISTRIBUTION.—Japan: Bungo Strait off northeastern 

Kyushu; off Amami Oshima, Ryukyu Islands (new record for 

Japan); 193-422 m. Elsewhere: South China Sea; Philippines; 

Celebes and Savu Sea, Indonesia; 70-555 m. ?Pleistocene 

of Vanuatu (Wells, 1984) and Ryukyu Islands (Yabe and 

Eguchi, 1932b). 

Notocyathus conicus (Alcock, 1902) 

PLATE 2ia,b 

Citharocyathus conicus Alcock, 19O2b:l 18-119; 1902c:22, pi. 3: figs. 18, 

18a.—Yabe and Eguchi, 1941c:212, fig. 4a,b; 1942b:122, pi. 10: figs. 17,18. 

Sphenotrochus viola.—Gerth, 1921:393, pi. 57: figs. 10, 11. 

Notocyathus conicus.—Yabe and Eguchi, 1946:7.—Eguchi, 1965:289, 2 

figs.—Cairns, 1989a:28, pi. 13: figs, a-i [synonymy]. 

DESCRIPTION.—Corallum regularly conical, never cylindrical, 

with a slightly compressed calice having a GCD:LCD

NUMBER 557 65 

range of 1.05-1.22. Largest Japanese specimen examined (TM 

(KT9015, CB1-2)) 5.85 x 5.39 mm in calicular diameter and 

7.60 mm in height. Costal structure and corallum color as in N. 

venustus. 

Septa hexamerally arranged in 4 complete cycles, as in N. 

venustus, but the relative sizes of septa differ. S1 exsert (about 

1.2 mm) and have sinuous vertical inner edges that fuse with 

the columella deep within fossa. S2 slightly less exsert (about 

1.0 mm), two-thirds width of S,, and also have sinuous inner 

edges that join with the inner P3 fusions. S3 slightly less exsert 

than S2 (about 0.8 mm) and about two-thirds width of an S2, 

each S3 separated by a wide notch from a tall, wide (about 0.75 

mm) palus. Inner edges of each pair of P3 within a system fused 

in a V-shaped structure before the S2, as in N. venustus. S4 

slightly less exsert than S3 and equal to or slightly less wide 

than the S3. Fossa quite shallow, containing a papillose 

columella. 

DISCUSSION.—Characters differentiating N. conicus from N. 

venustus are listed by Cairns (1989a) and in the previous 

species account. A more complete description and illustration 

of this species is given by Cairns (1989a). Based on the 

GCD:LCD ratio of Yabe and Eguchi's (1941c) specimen from 

the Philippine Pleistocene (1.19), I now consider it to be N. 

conicus, not N. venustus. 


CB1-2), 1, USNM 92777; TM (KT9015, HK3), 1, ORI. 

Previous Records: Syntypes of C. conicus, ZMA. 

TYPES.—Two syntypes of C. conicus are deposited at the 

ZMA (Coel. 1185). Type Locality: Siboga-95: S^.S^, 

119°40'E(Sulu Sea), 522 m. 

DISTRIBUTION—Japan: Bungo Strait; off Mi Shimi, Eastern 

Strait, southwestern Honshu; 70-110 m. Elsewhere: 

Philippines; Makassar Strait; off Sabah, Indonesia; 

34-923 m. Pleistocene of Philippines and Ryukyu Islands; 

Miocene of Java (Gerth, 1921). 

Peponocyathus Gravier, 1915 

DIAGNOSIS.—Corallum solitary and free, but quite variable 

in shape, including bowl-shaped, cylindrical, and globose. 

Theca imperforate; costae serrate, corresponding to septa. 

Septa highly exsert. Pali present before all but last cycle in two 

crowns, but P, often suppressed. Columella papillose. 

TYPE SPECIES.—Peponocyathus variabilis Gravier, 1915 (= 

P. folliculus), by original designation. 

Peponocyathus australiensis (Duncan, 1870) 

PLATES 28c-/, 41/ 

Deltocyathus italicus var. australiensis Duncan, 1870:297, pi. 19: fig. 4. 

Deltocyathus orientalis Duncan, 1876:431. pi. 38: Figs. 4-7.—Yabe and 

Eguchi, 1932a:387, 388; 194la:418-420, text-figs. 1-3; 1941b:102; 

1942b:l 12, 125-126.—Mori, 1964:314, pi. 46: figs. 1. 2. 

Deltocyathoides japonicus Yabe and Eguchi, 1932a:389, fig. 3; 1937:140-141, 

pi. 20: fig. 23a-c.—Eguchi. 1968:C35-36 — Eguchi and Miyawaki, 

1975:57. 

Not Peponocyathus orientalis Yabe and Eguchi, 1932b:444—445 [= P. 

folliculus]. 

Deltocyathus (Paradeltocyathus) orientalis Duncan.—Yabe and Eguchi, 

1937:130. 131-135.pl. 20: figs. 1-10. 

Deltocyathus (Paradeltocyathus) australiensis.—Yabe and Eguchi, 1937:130. 

Deltocyathoides japonicum.—Yabe and Eguchi, 1942b: 126. 

Paradeltocyathus orientalis Duncan.—Eguchi, 1965:289, 2 figs.—Kikuchi, 

1968:11. 

Notocyathus (Paradeltocyathus) orientalis Duncan.—Eguchi, 1968:C40-41.— 

Hamada. 1969:253-254. pi. 2: fig. 4a-c.—Otnuraet al.. 1984: 33. fig. IB. 

Deltocyathus sp.—Eguchi, 1974:228, pi. 70: figs. 6-11. 

Peponocyathus australiensis.—Cairns, 1989a:29, 30 32, pi. 14: figs, d-j; pi. 

15: figs, a-d [synonymy].—Cairns and Parker, 1992:39-40, pi. 13: figs. 

c-d.—Cairns and Keller, 1993:259-261. 

DESCRIPTION.—Corallum bowl-shaped and small, rarely 

exceeding 8 mm in calicular diameter or 7 mm in height. 

Largest typical specimen examined (TM (KT9015, BS2)) only 

8.6 mm in calicular diameter, but most Japanese specimens 

only 5.0-6.5 mm in diameter with a relatively high H:D ratio 

of 0.70-0.83. Costae rounded and equal in width (0.18-0.22 

mm), separated by very thin (about 0.1 mm) and deep (up to 1 

mm at calicular edge) intercostal furrows, which do not afford 

a view of underlying theca. Epicenter of base 0.8-1.0 mm in 

diameter and granular, from which 6 independent C1 originate. 

Within every system the C2 and 2 C3 originate in a trifurcation 

at the border of the epicentral region. Only 0.5 mm beyond this 

region a pair of C4 split from each C3. Corallum white. 

Septa invariably hexamerally arranged in 4 complete cycles 

(48 septa). S1 easily distinguished by their highly exsert upper 

margins and greater width, each having a straight, vertical inner 

edge that almost reaches the columella. S2 less exsert and about 

three-quarters width of an Sv each usually bearing a small (0.5 

mm wide) lamellar paliform lobe, the 6 P2 forming a palar 

crown encircling columella. S3 about three-quarters width of an 

S2 and often bear 1-3 papillose paliform lobes, positioned 

slightly outward from the columella with resepect to P2 crown. 

S4 adjacent to S1 usually slightly wider than those adjacent to 

S2. All septal and palar faces highly granular. Fossa shallow. 

Columella papillose, composed of up to 12 cylindrical, 

granular elements. 

DISCUSSION.—I (Cairns, 1989a) synonymized Deltocyathoides 

japonicus with Peponocyathus australiensis, but did not 

explain why. Specimens of the D. japonicus growth form 

invariably result from asexual regeneration of a parent 

fragment, which is easily recognized by the incorporation of the 

parent fragment into the new corallum (Plate 2%d,e). Specimens 

of this form are usually larger than typical specimens and have 

more septa (e.g., 72 septa in holotype of GCD 15.1 mm); have 

taller costal spines; and have much wider intercostal spaces, 

allowing a view of the underlying theca. Their septal symmetry 

is irregular, their palar structure is often rudimentary, and the 

columella is often lacking. In general, these specimens have a 

poorly organized structure typical of a juvenile corallum, even 

though they are among the largest specimens of the species. 

Because this morphology was believed to be the result of 

asexual fragmentation of P. australiensis, it was synonymized

66 

with that species (Cairns, 1989a), but may still be referred to as 

the "japonicus" growth form. 

Good descriptions of the typical form of this species are 

given by Yabe and Eguchi (1937), Cairns (1989a), and Cairns 

and Parker (1992). A complete synonymy is given by Cairns 

(1989a). 

MATERIAL EXAMINED.—New Records of Typical 

Form: Alb-3708,9, USNM 92754; Alb-4807,2, CAS 80912; 

Alb-5071, 6, USNM 81827; TM (KT7802, B), 1, USNM 

92755; TM (KT78-11, OT1), 2, USNM 92756; TM (KT78-11, 

OT6), 43, USNM 92757; TM (CR79-1), 31, USNM 92758; TM 

(KT9015, BS1), 2, USNM 92759; TM (KT9015, BS2), 3, 

USNM 92760; TM (KT9015, CB1-1), 2, USNM 92761; TM 

(KT9015, CB1-2), 4, USNM 92762; TM (KT9015, CB6-1), 30, 

USNM 92763; TM (KT9015, HK2), 2, USNM 92764; TM 

(KT9015, HK3), 13, USNM 92765; TM (KT9015, HK5), 5, 

USNM 92766; TM (KT9O15, OK2), 142, USNM 92767; TM 

(KT9015, OKI), 63, ORI; TM (KT9202, KB3), 2, USNM 

92768; TM (KT9202, OS2), 1, USNM 92769; TM (KT9202, 

OS3), 1, USNM 92770; USGS 17445, 17447, 17450, 17451, 

17503, Pleistocene of Okinawi, USNM 88666-671. New 

Records ofjaponicus form: ALB-3738,2, USNM 81828; TM 

(KT7414, B4), 3, USNM 92917; TM (KT7818, OT10), 3, 

USNM 92915, 1, ORI; TM (KT7911, OT4), 4, USNM 92916. 

Previous Records: Holotype of D. italic us australiensis, BM; 

holotype of D. japonicus, TIUS. 

TYPES.—The holotype of Deltocyathus italicus var. australiensis 

is deposited at the BM (R29255). Type Locality: 2.5 

km west of Cape Otway, Victoria, Australia (Miocene). 

The holotype of Deltocyathus orientalis Duncan appears to 

be lost (Zibrowius, 1980). Type Locality: 34°12'N, 136°20'E 

(off Owase, southeastern Honshu), 95 m. 

The holotype (Plate 41/) of Deltocyathoides japonicus is 

deposited at the TIUS (50091). Type Locality: Soyo Maru-21: 

36°47TM, 141°14 / E (off Hitachi, Honshu), 209 m. Although 

Yabe and Eguchi (1932a) did not specify the type or type 

locality, a specimen captioned as the holotype from Soyo 

Maru-2] was illustrated by Yabe and Eguchi (1937). 

DISTRIBUTION.—Peponocyathus australiensis is the most 

commonly collected and reported deep-sea coral in the 

Japanese region, and also one of the most widely distributed 

species worldwide (Caims, 1989a). 

It is found off both the Sea of Japan and Pacific coasts of 

Honshu, from Tsugaru Strait to the Eastern Channel, Korea 

Strait, as well as off Shikoku and southwestern Kyushu; 

59-494 m. Pleistocene of Ryukyu Islands. Elsewhere: 

Widespread in Atlantic and Indo-West Pacific, including 

Southern Australia and New Zealand; 44-635 m. 

Peponocyathus foUiculus (Pourtales, 1868) 

PLATE 28J?-* 

Stephanophyllia foUiculus Pourtales, 1868:139. 

Peponocyathus orientalis Yabe and Eguchi, 1932b:444-445. 15 figs. [= junior 


secondary homonym of Deltocyathus orientalis Duncan, 1876]; 

1942b: 123.—Eguchi, 1974:228, pi. 70: figs. 17, 18. 

Discotrochus (Cylindrophyllia) minimus Yabe and Eguchi, 1937: 146-147, pi. 

20: figs. 16-22; 1942b: 118. 

Kionotrochus minimus.—Yabe and Eguchi, 1941b: 102. 

Cylindrophyllia minima.—Eguchi, 1965:289, 2 figs.—Kikuchi, 1968:8, pi. 5: 

fig. 3. 

Cylindrophyllia orientalis.—Mori and Minoura, 1983:185-191, fig. lA-O. 

Peponocyathus foUiculus.—Cairns, 1979:113-115, pi. 22: figs. 1-4; pi. 20: 

fig. 11 [synonymy]; 1989a:32-33 [in part: pi. 15: figs, e-h; not pi. 16: figs. 

a-c, not Alb-5277, 5584, 5577, synonymy]. 

DESCRIPTION.—Corallum cylindrical with a flat base that 

appears to be the result of transverse division. Corallum 

relatively small: coralla rarely exceed 5 mm in calicular 

diameter and 6.5 mm in height, adults invariably taller than 

their diameter, usually having H:D greater than 1. Large 

illustrated specimen (TM (KT9015, CB1-2)) 4.9 mm in 

calicular diameter and 5.1 mm in height. Costae equal in width 

but not uniformly wide from base to calice, being widest 

(0.32-0.37 mm) near base to mid-corallum, and narrowest near 

calice. Deep intercostal furrows correspondingly narrow near 

base and wider near calice. Costae evenly rounded and 

uniformly covered on outer and lateral edges with fine (50 urn 

diameter) granules. Corallum white. 

Septa hexamerally arranged in 4 cycles, the last cycle rarely 

complete. Many specimens have only 1 pair of S4 in each 

system for a total of 36 septa, whereas juvenile coralla usually 

have only 24 septa. S1 only slightly more exsert than other 

septa and have straight, vertical inner edges that fuse with the 

columella. S2 about three-quarters width of an S1, each bearing 

a lamellar paliform lobe about 0.4 mm wide, the 6 P2 forming 

a small palar crown encircling the columella. S3 about 

three-quarters width of an S2 and their inner edges usually 

fused to adjacent S2; P3 were not observed in the specimens 

examined. S4 about three-quarters width of an S3. Fossa 

shallow to absent. Septal faces covered with prominent (up to 

80 urn tall), blunt granules. Columella papillose but small. 

DISCUSSION.—Peponocyathus orientalis and Cylindrophyllia 

minima were names originally given and predominantly 

applied to Neogene fossil specimens from the Japanese region; 

however, some Recent specimens were reported under the latter 

name by Yabe and Eguchi (1937, 1941b) and Kikuchi (1968). 

In a detailed study of almost 800 Pleistocene specimens, Mori 

and Minoura (1983) demonstrated that P. orientalis and C. 

minima were synonymous, the former name generally applying 

to specimens having 30-48 septa, the latter to smaller 

specimens with only 24 septa. They also stated that septal 

number was controlled by intrinsic genetic factors and was thus 

not a function of size, with 24 and 36 being the bimodal 

distribution of this character. I (Cairns, 1989a) synonymized 

the Japanese species with P. foUiculus and still maintain their 

equivalence; however, the Japanese specimens are, in general, 

larger than those from the Atlantic and tend to have more S4. If 

the Japanese populations are found to be a separate species, its 

correct name would be Peponocyathus minimus (Yabe and

NUMBER 557 67 

Eguchi, 1937), since P. orientalis Yabe and Eguchi, 1932 is a 

junior secondary homonym of Deltocyathus orientalis Duncan, 

1876 (= Peponocyathus australiensis). 


HK2), 1, USNM 92772; TM (KT9015, CB2-2), 2, USNM 

92773; TM (KT9015, CB1-2), 6, USNM 92774, 2, ORI; TM 

(KT9202, YT4), 1, USNM 92775; USGS 17681, Pleistocene of 

Okinawa, 1, USNM 88670. Previous Records: Holotype of 5. 

folliculus, MCZ. 

TYPES.—The holotype of Stephanophyllia folliculus is 

deposited at the MCZ. Type Locality: Bibb-5\: 24°12'40"N, 

81°19'25"W (Straits of Florida), 433 m. 

Five syntypes of Peponocyathus orientalis Yabe and Eguchi, 

1932b are deposited at the TIUS (43423). Type Locality: 

Pleistocene ofKikai-jima, Ryukyu Islands. 

Approximately 62 syntypes of Discotrochus minimus from 

11 localities are deposited at the TIUS. Type Localities: 

Neogene of Taiwan; Chiba Prefecture, Honshu; Shimbara 

Peninsula, Kyushu. Recent of Toyama Bay, depth unknown. 

DISTRIBUTION.—Recent of Japan: Sagami and Suruga Bays, 

Honshu; off Amakusa Island, Kyushu; Eastern Strait off 

southwestern Honshu; Toyama Bay, Honshu; Osumi Shoto, 

northern Ryukyu Ids; 30-402 m. Neogene of Chiba Prefecture; 

Shimbara Peninsula, Kyushu; Okinawa, Ryukyu Islands. 

Elsewhere: South China Sea, Philippines, Celebes, Atlantic 

Ocean; 50-582 m. 

Tropidocyathus Milne Edwards 

and Haime, 1848a 

DIAGNOSIS.—Corallum solitary and cuneiform to campanulate, 

with a rounded base; transverse division absent, but 

asexual fragmentation occurs in some species. Alate lateral 

thecal crests present in some species. Theca imperforate; costae 

serrate to granular and correspond to septa. Septa highly exsert. 

Pali occur in three crowns before all but last cycle; columella 

papillose to lamellar. 

TYPE SPECIES.—Flabellum Lessonii Michelin, 1842, by 

monotypy. 

Tropidocyathus lessoni (Michelin, 1842) 

PLATE 29aJ> 

Flabellum Lessonii Michelin, 1842:119. 

Trochocyathus (Tropidocyathus) cf. lessoni.—Yabe and Eguchi, 1942b: 124. 

Trochocyathus (Tropidocyathus) wellsi Yabe and Eguchi 1942b: 124, 153, pi. 

10: fig. 22a,b. 

Tropidocyathus lessonii.—Eguchi, 1965:289, 2 figs. 

Tropidocyathus wellsi.—Eguchi, 1965:290. 2 figs. 

Tropidocyathus lessoni.—Cairns, 1989a:33-34, pi. 16: figs, d-1 [synonymy].—Caims 

and Keller. 1993:253, fig. 7c. 

DESCRIPTION.—Corallum cuneiform, with a rounded base 

and pronounced edge crests that may project laterally up to 5.2 

mm and be 0.6-1.4 mm thick. Largest specimen known, the 

holotype of T. wellsi, measures 18 x 14 mm in calicular 

diameter and is 17 mm in height. Calice elliptical to 

diamond-shaped, with GCD:LCD ratios of 1.1-1.5. Costae 

broad and flat, the Cu3 being slightly wider than the C4. CU3 

bear 3 or 4 granules across their width near the calice, whereas 

C4 are only about 2 granules wide. Intercostal furrows narrow 

(about 0.15 mm) and relatively shallow (0.4 mm), degenerating 

about two-thirds distance to corallum base, the lower third of 

the thecal faces uniformly granulated and lacking costal 

furrows. Theca of fresh specimens pale orange; septa white. 


to the formula S1>S2>S4>S3. S! highly exsert (up to 2.5 mm), 

extend about three-quarters distance to columella, and have 

thick, straight inner edges. A relatively small (0.3 mm wide), 

low P, occurs on every Sv S2 almost as exsert and as wide as 

an Sv and also have thick straight inner edges, but each usually 

bears a large palus (e.g., 0.5 mm wide) that reaches higher in 

the fossa than the Pt; P2 occasionally missing. S3 considerably 

less exsert than S1-2, about three-quarters width of an S2, and 

have relatively thin, slightly sinuous inner edges. Each S3 bears 

a rather large palus up to 1.6 mm in width, pairs of which often 

loosely fuse to its flanked P2 near the columella into a 

chevron-like structure. Inner edges of all pali (P^) border on 

columella. S4 less exsert but slightly wider than an S3. Septal 

and palar faces bear tall granules. Fossa of moderate depth, 

containing an elongate columella composed of basally fused 

papillae. 

DISCUSSION.—Tropidocyathus lessoni is relatively easily 

distinguished from T. pileus by the following characters. 

Tropidocyathus lessoni has prominent, granular thecal edge 

crests; T. pileus does not. The costae of T. lessoni degenerate 

into granules towards the base and the intercostal spaces are 

shallow; costae and well-defined intercostal spaces of T. pileus 

extend to the corallum base. Furthermore, the costae of T. 

lessoni are flat and bear 2-4 granules across their width, 

whereas those of T. pileus are rounded and bear essentially a 

single row of peripheral granules. Finally, the theca of T. 

lessoni is orange, that of T. pileus is white. 

A more complete description is given by Cairns (1989a). 


YT2), 7, USNM 92779, 2, ORI; TM (KT9309, AM6), 3, ORI; 

USGS 17451, north of Iwa, Okinawa, Pleistocene, 1, USNM 

88674. Previous Records: Specimens reported from the 

Philippines (Cairns, 1989a) and the southwest Indian Ocean 

(Cairns and Keller, 1993). 

TYPES.—The holotype of Flabellum Lessonii is deposited in 

the Michelin Collection at the MNHNP. No type locality was 

given. 

The holotype of Trochocyathus wellsi is deposited at the 

TIUS (43691). Type Locality: Soyo Maru-439: 31°52TS(, 

128°O1 'E (off Danjo Gunto, East China Sea), 155 m. 

DISTRIBUTION.—Japan: East China Sea off southwestern 

Kyushu and Danjo Gunto; off Osumi Shoto and Amami 

Oshima, northern Ryukyu Islands; 98-155 m. Pleistocene of 

Ryukyu Islands. Elsewhere: South China Sea, Philippines, 

Indonesia, southwest Indian Ocean; 62-421 m.

68 

Tropidocyathus pilots (Alcock, 1902) 

PLATE 29d,e 

Trochocyathus pile us Alcock, 1902a:96-97; 1902c:15-l6, pi. 2: figs. 11, 

1 la.—Yabe and Eguchi, 1942b:123, pi. 10: figs. 19,20.—Eguchi, 1965:286, 

2 figs. 

Trochocyathus (Tropidocyathus) intermedius Yabe and Eguchi, 1932b:443 

[notnen nudum]. 

Tropidocyathus pileus.—Caims, 1989a:34-35, pi. 17: figs, a-h [synonymy]. 

DESCRIPTION.—Looking at a lateral face, the corallum is 

trapezoidal, the lower lateral edges being either rounded or 

slightly protuberent, but never carinate (crested). Largest 

Japanese specimen examined (TM (KT9015, BS2)) 18.7 x 11.0 

mm in calicular diameter and 19.5 mm in height. Costae 

continuous from calice to base, where many are continuous 

with their counterparts from the opposite face. Edge protuberances, 

if present, vertically costate and not granular. Costae 

rounded and equal in width, each bearing an outer unilinear row 

of coarse teeth, as well as smaller granules on their lateral faces. 

Intercostal furrows deep. Corallum white. 

Septa hexamerally arranged in 4 cycles, larger specimens 

having pairs of S5 in their end half-systems, e.g., the illustrated 

specimen has 6 pairs of S5 for a total of 60 septa. S1 about 2 

mm exsert and have slightly sinuous inner edges, each S1 

bordered by a small (about 0.8 mm wide), highly granular palus 

that is positioned directly adjacent to the columella. S2 only 

slightly less exsert and about 80% width of an S^ each S2 also 

bearing a palus of equal width and position to that of the P1 but 

rising higher in the fossa. S3 least exsert septa (only about 1 

mm) and about two-thirds width of an S2, each having a broad 

(about 1.5 mm wide) palus slightly recessed from the columella 

and rising higher in the fossa than the P2. Each pair of P3 within 

a system slant toward their flanked P2, the inner edges of these 

triads (1 P2 and 2 P3) sometimes loosely fused into a chevron 

arrangement. S4 slightly more exsert and wider than S3, their 

outer edges fused to adjacent S^ at calicular edge producing 

calicular projections that result in a serrate calicular margin. P4 

present only if pairs of S5 present in a half-system. Fossa of 

moderate depth. Columella linear-papillose, the elements 

strongly fused together into a solid lamella. 

DISCUSSION.—The name Tropidocyathus intermedius (Yabe 

and Eguchi, 1932b), attributed to a Pleistocene fossil coral, was 

never described but nonetheless was synonymized by Yabe and 

Eguchi in 1942(e). A second Pleistocene specimen from the 

same general region is reported herein. 

Comparisons to T. lessoni are made in the account of that 

species. A more complete description and illustration of T. 

pileus can be found in Cairns (1989a). 


BS2), 6, USNM 92781,1, ORI; TM (KT9202, OS2), 2, USNM 

92780; TM (KT9309, AM8), 11, ORI, 5, USNM 93164; USGS 

17445, 1.6 km west of Yonabaru, Okinawa, Ryukyu Islands, 

Pleistocene, 1, USNM 88673. Previous Records: Syntypes of 

Trochocyathus pileus, ZMA. 

TYPES.—Four syntypes of T. pileus are deposited at the 


ZMA (Coel. 7352). Type Locality: Siboga-95: 5 0 43TSf, 

119°40'E (Sulu Sea), 522 m. 

DISTRIBUTION.—Off Japan: Suruga Bay, Honshu; off Shikoku; 

off southern Kyushu; off Amami Oshima, Ryukyu 

Islands; 123-422 m. Pleistocene of Ryukyu Islands. Elsewhere: 

South China Sea, Philippines, off southeastern Australia, 

?Gulf of Manaar; 123-522 m. 

Alatotrochus, gen. nov. 

DIAGNOSIS.—Corallum cuneiform, with a rounded (free) 

base and prominent, costate thecal edge crests. Theca imperforate. 

Costae serrate, extending from calice to base; costae 

number twice that of septa. Four cycles of highly exsert septa. 

Pali absent; columella linear-papillose. 

DISCUSSION.—Using my key to the turbinoliid genera 

(Cairns, 1989a:25-26), Alatotrochus would key to Platytrochus, 

the genus in which Moseley (1876) originally placed the 

type species. Alatotrochus does share several characters with 

Platytrochus, such as a cuneiform corallum with edge crests, a 

papillose columella, and the absence of pali. Alatotrochus 

differs from the type species, Platytrochus stokesi (Lea), 

however, in several significant characters. It has a much larger 

corallum, an additional cycle of septa, and twice as many costae 

as septa. Its costae are serrate and continuous from calice to 

base, whereas those of P. stokesi are discontinuous and 

granular. Finally, its columellar elements are large and linearly 

arranged, whereas those of Platytrochus are much smaller and 

grouped in an elliptical field or two rows. 

Moseley (1881) later placed Platytrochus rubescens in the 

genus Sphenotrochus, which is not surprising given the 

similarity of Platytrochus and Sphenotrochus (see Cairns, 

1989a: 38). Nonetheless, Alatotrochus differs from species of 

Sphenotrochus in having a papillose (not lamellar) columella; 

a larger corallum; serrate (not smooth) costae; and twice as 

many costae as septa. 

Alatotrochus is monotypic. 

ETYMOLOGY.—The name Alatotrochus (Latin alatus, meaning 

"winged" + trochus, a common coral suffix, literally 

meaning "wheel"), refers to the prominent edge crests of this 

genera that resemble wings. Gender: masculine. 

TYPE SPECIES.—Platytrochus rubescens Moseley, 1876, 

here designated. 

DISTRIBUTION.—Bungo Strait, off Kyushu; off Amami 

Oshima, Ryukyu Islands; 193-422; Banda Sea, 236 m; 

Pleistocene of Okinawa, Ryukyu Islands. 

Alatotrochus rubescens (Moseley, 1876), comb. nov. 

PLATE 29/J-/ 

Platytrochus ruhescens Moseley, 1876:553. 

Sphenotrochus rubescens.—Moseley, 1881:157-159, pi. 6: figs. 8, 8a. 

DESCRIPTION.—Corallum compressed, very much resembling 

the shape of Tropidocyathus lessoni, with thin, solid (not 

hollow) edge crests extending as much as 4 mm from the thecal

NUMBER 557 69 

edge. Largest Japanese specimen examined 16.1 x 10.5 mm in 

calicular diameter and 14.7 mm in height; however, the 

syntypes are larger: up to 20 x 16 mm in calicular diameter and 

17-19 mm in height. Costae thin (about 0.8 mm wide), high, 

serrate ridges, one corresponding to each septum, and another 

of equal size corresponding to each interseptal space. Near 

calice the costae that correspond to septa widen at the expense 

of the interseptal costae. Additional costae, also uncorrelated to 

septa, cover the thecal edge crests, oriented at right angles to 

face costae. Many, but not all, costae are continuous from calice 

to base. The 2 principal costae extend along the outside of each 

lateral crest and are continuous along the sharp-edged base of 

the corallum, meeting one another at base epicenter. Intercostal 

spaces broad (0.18-0.20 mm) but diminish in width near 

calice. Theca, calicular edge, and exsert portion of septa reddish 

brown. 

Septa hexamerally arranged in 4 cycles (48 septa) according 

to the formula: S1>S2>S3>S4; however, some larger syntypes 

have pairs of S5 in end half-systems, for a total of 56 septa. S1 

highly exsert (up to 4 mm) and quite thick (0.25 mm), with 

straight, vertical inner edges that fuse with the columella low in 

fossa. S2 only slightly less exsert (about 3.5 mm), but otherwise 

similar to the Sr S3 2.0-2.2 mm exsert and about two-thirds 

width of an S^g, with finely sinuous inner edges that do not 

attain the columella. S4 1.3-1.5 mm exsert and only about 

one-third width of an S3. All septal faces covered with very 

low, rounded granules, producing an apparently smooth aspect. 

Fossa shallow but commodious, caused by widely spaced septa 

and the absence of pali. Columella papillose, composed of 5 or 

6 tall, cylindrical, finely granulated, aligned pillars, each pillar 

0.4-0.7 mm in diameter and reaching almost to level of calice. 

DISCUSSION.—This is only the second report of this rare and 

beautiful species, heretofore known only from four syntypes 

from the type locality. Aside from the six specimens reported 

herein from Bungo Strait, a Pleistocene specimen (USGS 

17633) is tentatively assigned to this species. It differs only in 

that its four lateral C1 are swollen into robust protuberances 

near the calicular edge. 


BS2), 5, USNM 92776, 1, ORI; TM (KT9309, AM7), 4, ORI; 

TM (KT9309, AM8), 1 ORI; USGS 17633, 1 km southwest of 

China, Chinen Peninsula, Okinawa, Ryukyu Islands, Pleistocene, 

1, USNM 88672. Previous Records: 1 syntype of P. 

rubescens, BM. 

TYPES.—Four syntypes of Platytrochus rubescens, one 

illustrated in Plate 29h,i,l, are deposited at the BM (1 numbered 

1880.11.25.163). Type Locality: Challenger-192: 5°49' 15"S, 

132°14'15"E(off Kei Island, Banda Sea), 136 m. 

DISTRIBUTION.—Same as that for genus. 

Idiotrochus Wells, 1935 

DIAGNOSIS.—Corallum cuneiform; anthocaulus often having 

a compressed but expansive base, which may bear 2 lateral 

projections; transverse division present. Theca imperforate; 

costae smooth; intercostal striae shallow. Septa alternate in 

position with costae. Septal faces carinate and granular. Pali 

present before first 2 cycles; columella papillose to fascicular. 

TYPE SPECIES.—Sphenotrochus emarciatus Duncan, 1865, 

by original designation. 

Idiotrochus kikutii (Yabe and Eguchi, 1941) 

PLATE 30a-d 

Placotrot hides ? kikutii Yabe and Eguchi. 1941b:104, 3 figs.; 1942b:l 18.149. 

pi. 9: fig. 16a-c. 

Idiotrochus kikutii.—Cairns, 1989a:36-37, pi. 18: figs. a,b, d-h [synonymy]. 

DIAGNOSIS.—Corallum (anthocyathus) compressed, with a 

"pinched," blade-like lower edge. Specimens typically 2.3 x 1.7 

mm in calicular diameter and 4-5 mm in height Costae wide 

(0.21 mm) and smooth to only slightly granular, intercostal 

striae very narrow and shallow. Septa hexamerally arranged in 

3 cycles, most septal faces bearing elongate carinae. Ten to 12 

pali occur before S1-2, the 2 principal P1 often absent. 

Columella fascicular. 

DISCUSSION.—No additional specimens of this unusually 

shaped species are reported herein. It is known from only one 

record in Japanese waters, the type locality in Toyama Bay, 

Honshu. A full description with illustrations of this species 

based on Philippine specimens is given by Cairns (1989a). 

MATERIAL EXAMINED.—New Records: None. Reference 

Specimens: Philippine specimens reported by Cairns 

(1989a). 

TYPES.—Six syntypes of Placotrochides kikutii are deposited 

at the TIUS (63088). Type Locality: Toyama Bay, 

Honshu, depth not recorded. 

DISTRIBUTION.—Sea of Japan, Toyama Bay, Honshu; South 

China Sea; Philippines; Celebes Sea; 143-645 m. 

Superfamily FLABELLOIDEA 

Family GUYNHDAE 

Stenocyathus Pourtales, 1871 

DIAGNOSIS.—Solitary, ceratoid to cylindrical, free or attached. 

Wall epithecate; rows of thecal spots in every 

intercostal region, aligned somewhat closer to the C3 than Cu2- 

Only 3 cycles of septa. Pali opposite S2; columella formed of 

1-2 twisted, crispate elements. 

TYPE SPECIES.—Coenocyathus vermiformis Pourtales, 1868, 

by monotypy. 

Stenocyathus vermiformis (Pourtales, 1868) 

PLATES 22g, 29c,/ 

Coenocyathus vermiformis Pourtales, 1868:133. 

Stenocyathus vermiformis.—Cairns, 1979:168-170, pi. 32: figs. 8-10; pi. 33: 

figs. 1. 2 [synonymy]; 1982:52, pi. 16: figs. 8-11.—Cairns and Parker, 

1992:43. pi. 14: figs. b.c. 

DISCUSSION.—Stenocyathus is a monotypic genus, and thus 

the generic dignosis also serves as the species diagnosis. 5.

70 

vermiformis is easily distinguished from all other North Pacific 

corals by its small, slender, cylindrical corallum and aligned 

mural spots. Its small size and inconspicuous habit have 

undoubtedly caused this relatively widespread species to have 

been overlooked in the past. The specimen reported herein from 

Soyo Maru-425 is 2.3 mm in calicular diameter and 3.2 mm in 

length with large P2 and a fascicular columella. It can be 

identified based on the generic diagnosis; illustrations of this 

species may be found in the references listed in the synonymy. 

MATERIAL EXAMINED.—New Records: Soyo Maru-425,1 

specimen attached to a dead Crispatotrochus rubescens, TIUS 

53695, Plate 22g. Previous Records: Syntypes of C. vermiformis, 

MCZ; specimens reported by Cairns (1979, 1982) and 

Cairns and Parker (1992). 

TYPES.—Thirty-eight syntypes of Coenocyathus vermiformis 

are deposited at the MCZ (Cairns, 1979). Type 

Locality: Off Florida Keys, 274-329 m. 

DISTRIBUTION.—The single Japanese record from off Koshiki 

Retto, southwestern Kyushu (300 m) is the first report of 

this genus from the North Pacific. It was apparently overlooked 

by Yabe and Eguchi (1942b) in their report of Cyathoceras 

rubescens. Elsewhere: It is known to be common on both 

sides of the Atlantic; off St. Paul and Amsterdam Islands, 

southern Indian Ocean; the New Zealand region, including the 

Antipodes and Campbell Plateaus; off Tasmania; and off 

southeastern Australia; 80-1229 m. 

Truncatoguynia Cairns, 1989a 

DIAGNOSIS.—Solitary, elongate compressed-cylindrical, and 

free. Budding by transverse division. Epitheca smooth; rows of 

thecal pores occur in every intercostal space, alignment of rows 

appearing to flank the secondary costae. Septa only slightly 

exsert and variable in symmetry, arranged in 2 or 3 size classes. 

Pali and columella absent. 

TYPE SPECIES.—Truncatoguynia irregularis Cairns, 1989a, 

by monotypy. 

Truncatoguynia irregularis Cairns, 1989 

PLATE 30r/ 

Truncatoguynia irregularis Cairns. 1989a:43. pi. 22: figs. f,g; pi. 23: figs. c,f. 

DESCRIPTION.—Corallum an elongate, laterally compressed 

cylinder with rounded thecal edges. Largest Japanese specimen 

examined 44.4 mm in length, with calicular diameters up to 5.8 

x 4.3 mm. Base of specimens (anthocauli) terminate in an 

elliptical scar about 3.9 x 3.1 mm in diameter, revealing 12-24 

primary septa. Theca thin, smooth, and in ail Japanese 

specimens, somewhat wom, the coralla obviously dead when 

collected. Thecal pores irregular in appearance, more common 

toward the base of a corallum and often absent from theca near 

calice. In several weathered coralla and the older (basal) 

portions of others, the secondary septa and associated overlying 

theca are absent, resulting in a characteristic diagenetic form 


consisting of 12-18 lamella (septa) radiating from a central 

columella. Corallum white. 

Most coralla hexamerally arranged in three cycles of septa 

and an additional pair of S4 in each of the four end half-systems 

according to the formula: S1_2>S3>S4 and a total of 32 septa; 

however, one specimen had 14 primary septa and another had 

18. Su2 slightly exsert (1.1 mm) with slightly sinuous inner 

edges that are strongly fused to the columella. S3 only one-third 

to one-half width of an S1 and also have slightly sinuous inner 

edges, but they do not attain the columella. S4 rudimentary, 

consisting of a short row of spines. Fossa deep. Columella an 

elongate fusion of lower, inner edges of S^. 

DISCUSSION.—This species was previously known only 

from its type locality in the South China Sea. Although all the 

Japanese specimens were dead when collected, they do allow a 

refinement of the original description and reinforce the 

observation that the septal symmetry of this species is indeed 

irregular. 

MATERIAL EXAMINED.—-New Records: TM (KT9202, 

YT1), 1, USNM 92753, 2, ORI. Previous Records: Types of 

T. irregularis, USNM. 

TYPES.—The holotype (81890) and 10 paratypes (81891) of 

T. irregularis are deposited at the USNM. Another paratype is 

deposited at the Australian Museum (G15252). Type Locality: 

Alb-5311: 21°33'N, 1 ^tf'E (South China Sea north of 

Pratas Island), 161 m. 

DISTRIBUTION.—Osumi Shoto, northern Ryukyu Islands; 

north of Pratas Island, South China Sea; 80-161 m. 


Family FLABELLIDAE 

Flabellum Lesson, 1831 

Subgenus Flabellum (Flabellum) Lesson, 1831 


Flabellum (F.) pavoninum Lesson, 1831 

PLATES 30g-i, 3la-e 

Flabeilum pavoninum Lesson, 1831:2.—Gray, 1849:75-76.—Marenzeller, 

1888a:41-42.—Cairns, 1989a:46-50, pi. 23: figs, g-1; pi. 24: figs. a-d,g,h 

[synonymy]. 

Flabellum distinctum Milne Edwards and Haime, 1848a:262.—Marenzeller, 

I888a:42.—Yabe and Eguchi, 1932a:387, 389; 1932b:443; 1941b:101; 

l942a:93-95, pi. 5: figs. 3-6; pi. 6: figs. 3,4,9, 10; pi. 7: fig. 7; 1942b:l 12, 

130-131, pi. 11: figs. 10-12.—Eguchi, 1938, table 2; 1944:132-134, 5 figs; 

I968:C44 [in part: pi. C28: figs. 5, 6].—Utinomi, 1965:255.—Eguchi and 

Miyawaki, 1975:58.—Mori and Minoura, 1980:321-326, figs. 1-5.—Zou 

et al., 1981. pi. 1: figs. 2, 3.—Song, 1982:136-137, pi. 3: figs. 1-4; 

1991:135. 

Flabellum coalitum Marenzeller, 1888a:48-49.—Cairns, 1989a:46, 50, pi. 24: 

figs, e.f.i-1. 

Flabellum sp. cf. distinctum.—Okutani, 1969:12.—?Hamada, 1969:254. 

DESCRIPTION.—Corallum compressed, the planar faces 

meeting at acute edges and projecting 2.5-3.8 mm beyond the

NUMBER 557 71 

edge as low, discontinuous crests. Angle of thecal edges quite 

variable, ranging from 74° (coalitum form) to 139° (typical 

form); inclination of lateral faces, 27°-42°.Edge angle of lower 

5-8 mm of corallum considerably lower (e.g., 48°) than 

remainder of corallum. GCD:LCD also variable, ranging from 

1.9-2.1 {coalitum form) to 2.4 (typical form). Largest 

specimen known (lectotype of F. distinction) 60.6 x 25.1 mm in 

calicular diameter and 42.8 mm in height. Holotype of F. 

coalitum 32.3 x 14.3 mm in calicular diameter and 30.0 mm in 

height. Pedicel circular, 1.0-1.5 mm in diameter, and usually 

worn, revealing 6 protosepta. Thecal faces of most specimens 

examined discolored and worn; however, in well-preserved 

specimens there are narrow brown stripes associated with the 

theca overlaying every septum. Calicular edge a smooth arc. 

Septa hexamerally arranged in 6 cycles (S1-3>S4>S5>S6), 

the last cycle progressively appearing throughout a GCD range 

of 30-45 mm, and a full sixth cycle (192 septa) usually attained 

at a GCD of 45-47 mm. S1-3 quite wide, their inner edges 

extremely sinuous and thick, almost touching their counterparts 

from the opposite face. S4 of larger coralla as wide as S^_3, but 

have less thick and less sinuous inner edges. S4 of smaller 

coralla about 80% width of an S^_3 and thus more easily 

distinguished. S5 about half width of an S1-4 and also have 

moderately sinuous inner edges. S6 rudimentary, only onequarter 

to one-third width of an S5. Fossa very deep and 

elongate. Columella an elongate fusion of inner edges of S1_4, 

but often obscured from view by the narrow fossa. 

DISCUSSION.—Yabe and Eguchi (1942a) synonymized F. 

coalitum Marenzeller, 1888a (type locality: Japan) with F. 

distinctum Milne Edwards and Haime, 1848a (type locality: 

Japan); however, based on an examination of the type series of 

both species, I (Cairns, 1989a) maintained their distinction 

based on the considerably lower edge angle (74°) and lower 

number of septa (130°) of the holotype of F. coalitum. Despite 

its long synonymy, F. pavoninum (= F. distinctum) is known 

from remarkable few specimens, but the Japanese material 

reported herein includes one group (TM (KT8412, 14)) of 44 

specimens that displays a range of edge angles of 78°-117° and 

a range of septal number from 130°-192°, the septal number 

being directly correlated to GCD. Since all other characters of 

F. pavoninum and F. coalitum are the same, these Japanese 

specimens would appear to link the two extremes of the 

species: form coalitum having a relatively low edge angle and 

5'/2 septal cycles; the typical form having a greater edge angle 

(up to 139°) and 6 full cycles. Although only several of Yabe 

and Eguchi's (1942a,b) Japanese specimens were examined 

(see "Material Examined"), they all appear to be the coalitum 

form of the species. 


USNM 86552; Alb-5312, 3, USNM 40752; TM (KT78O2, Z2), 

1, USNM 92818; TM (KT8412, 14), 38, USNM 92819,6, ORI; 

TM (KT9015, CB2-2), 1, USNM 92820; TM (KT9202, KB3), 

1, USNM 92815; Tsuchida-45, 1, USNM 92821; Formosa 

Strait, 90 m, 2, CAS 74995; East China Sea, southeast of Cheju 

Do, 80 m, 1, CAS 74997. Previous Records: Types of F. 

pavoninum, F. distinctum, and F. coalitum; F. distinctum of 

Yabe and Eguchi (1942b): Soyo Maru-431, 1, TIUS 43448; 

Soyo Maru-429, 1, TIUS 39732 (Plate 30/); Sagami Bay, 1, 

TIUS 43409; near Seto, 2, TIUS 39236 (Plate 30g,h); F. 

distinctum of Marenzeller (1888a), off Japan, NMW 8197 

(Plate 31 b,c); F. pavoninum of Marenzeller (1888a), off China, 

NMW 8201 (Plate 31a). 

TYPES.—The lectotype and 4 paralectotypes of F. pavoninum 

are deposited at the MNHNP (#372) (Cairns, 1989a). 

Type Locality: "Sandwich Islands" (?Hawaiian Islands), 


The lectotype and 3 paralectotypes of F. distinctum are 

deposited at the BM and MNHNP (Cairns, 1989a). Type 

Locality: "Japan" (see Gray, 1849), depth unknown. 

The holotype (Plate 3ld,e) off. coalitum is deposited at the 

NMW (#8196). Type Locality: "Japan," depth unknown. 

DISTRIBUTION.—Japan: common off Honshu, Shikoku, and 

Kyushu, its northern limits at 39°36'N (Soyo Maru-624) in the 

Sea of Japan and 35° 18'N (Hasu Maru-45) on the Pacific coast 

of Japan. Also known from Korea Strait and off Cheju Do, 

Korea; Formosa Strait; South China Sea north of Pratas Island; 

73-658 m. Elsewhere: Hawaiian Islands, southwest Indian 

Ocean; 98-665 m. 

F lube Hum (F.) patens Moseley, 1881 

PLATE 3\g-i 

Flabellum patens Moseley, 1881:172 [in part: pi. 6: fig. 5].—Cairns. 

1989a:5l-52. pi. 26: figs, a-i [synonymy]. 

Flabellum pavoninum paripavoninum.—Yabe and Eguchi, 1942a:91-93 [in 

part: Soyo Maru-343. 419. pi. 5: fig. 7a-c]; 1942b:129 [in part: Soyo 

Maru-M3, 419. not pi. 11, fig. 9).—Eguchi, 1965:291, 2 figs.—Utinomi, 

1965:255. 

?Flabellum pavoninum lamellulosum.—Yabe and Eguchi, 1942b: 131. 

DESCRIPTION.—Corallum highly compressed, having virtually 

planar thecal faces that meet at a sharp angle and project as 

much as 6 mm as prominent edge crests. Angle of thecal edges 

(exclusive of edge crests) 117°-155°, but including the aliform 

crests, the thecal edges may approach 180°. Inclination of 

lateral faces 20°-32°, the faces of large specimens slightly 

concave. GCD:LCD = 2.05-2.50; GCD:H about 1.04, the 

corallum being about equally tall as broad. Largest specimen 

known (Alb-5313) 67.5 x 28.9 mm in calicular diameter and 

54.7 mm in height. Pedicel small and circular, only 1.5-2.0 

mm in diameter, and often exposing the 6 protosepta. Theca 

white to grey, relatively smooth, and regularly partitioned by 

thin intercostal striae. Theca near calice often discolored by an 

arched band about 5 mm wide than parallels the calicular edge, 

which is the indication of a former association with a 

commensal Lumbrineris polychaete (Zibrowius, Southward, 

and Day, 1975). Upper calicular edge smooth. 


formula: S,-4>S5>S6»S7. Sixth cycle complete at a GCD of

72 

38-41 mm; seventh cycle gradually added up to a GCD of 

about 65 mm at which size all 384 septa are usually present. 

S1_4 equal in size only in large coralla, the S4 being slightly 

less wide than S1-3 in smaller coralla; regardless of size, inner 

edges of all S^ attain the columella. Upper, inner edges of 

S1-4 moderately sinuous, becoming thicker and more sinuous 

deeper in fossa. S5 half to three-quarters width of S1_4, have 

moderately sinuous inner edges, and do not attain the 

columella. S6 about one-third width of S5, and S7, if present, 

rudimentary: only about 0.5 mm wide and 2-3 mm long. Fossa 

deep and narrow, formed by vertical inner edges of S1_4. 

Columella elongate, composed of a trabecular to solid fusion of 

the lower, inner edges of the S1_4, as much as 30 mm long and 

about 2 mm wide. 

DISCUSSION.—Several of the Indo-West Pacific species of 

Flabellum (Flabellum) were compared by Cairns (1989a, table 

5) and further distinguished by discriminant analysis (Cairns, 

1989b). Flabellum patens was found to be most similar to F. 

pavoninum, differing primarily in corallum shape. To quantify 

the difference in shape, F. patens has a lower face angle 

(20°-32° vs 32°-42° for F. pavoninum), a higher GCD:LCD 

(2.05-2.50 vs 1.98-2.41), a lower GCD:H (1.04 vs 1.34), 

higher edge crests (4.8-6.1 vs 2.5-3.8 mm), and a tendency to 

have S7. 

Yabe and Eguchi's (1942a) report of F. pavoninum 

lamellulosum from off Seto was not illustrated and their two 

specimens cannot be located, but based on their description, 

these specimens would appear to be F. patens. 


USNM 40757,2, ORI; Alb-5314,1, USNM 92813; YO70-770, 

1, USNM 92314; off Pengue, Taiwan, 3, CAS 74993. Previous 

Records: 4 syntypes of F. patens, BM; F. pavoninum 

paripavoninum of Yabe and Eguchi (1942a,e): Soyo Maru-4\9, 

1, TIUS 43441; 5oyo Maru-343, 1, TIUS 43445. 

TYPES.—The lectotype and 5 paralectotypes of F. patens are 

deposited at the BM, the lectotype designated by Cairns 

(1989a:52). Type Locality: Challenger-192: 5°49'S, 

132°14'E (Kei Islands, Banda Sea), 256 m. 

DISTRIBUTION.—Japan: Kii Strait, Honshu; off northeastern 

Shikoku; off southwestern Kyushu; Formosa Strait; South 

China Sea north of Pratas Island; 223-402 m. Elsewhere: 

Philippines, Banda Sea; 256-439 m. 

Flabellum (F.) magnificum Marenzeller, 1904 

PLATE 31./-/ 

Flabellum magnificum Marenzeller. 19O4a:276-277.pl. 17: fig. 13.—Utinomi, 

1965:255.—Cairns, 1989a:5O-51, pi. 25: figs, a-j [synonymy]. 

Flabellum pavoninum.—Yabe and Eguchi. I942a:90-91. pi. 5: fig. 2a-c [in 

part Soyo Marn-437]; 1942b: 129. pi. 11: fig. 7a-c. 

Flabellum pavoninum magnificum.—Yabe and Eguchi, 1942a:89-90, pi. 5: 

fig. la-c; 1942b:129, pi. 11: fig. 8a-c. 

DIAGNOSIS.—Angle of thecal edges 140°-172°; inclination 


of planar thecal faces quite open, 44°-58°. Discontinuous edge 

crests usually not present. Thecal faces bear reddish brown 

stripes associated with each costa. Pedicel narrow, only 

1.5-2.3 mm in diameter. Ratio of GCD:LCD = 1.65-1.96. 

Septa hexamerally arranged in 7 cycles, the last cycle never 

complete, the largest known specimen having 376 septa. Septal 

formula: S1_4>S5>S6>S7. Columella well developed. 

DISCUSSION.—No additional specimens of this large and 

elegant species are reported herein. It is known in Japanese 

waters from only four specimens reported by Yabe and Eguchi 

(1942a,b) and Utinomi (1965). A full description and illustrations 

are given by Cairns (1989a). It is distinguished (Cairns, 

1989a,b) from other Indo-West Pacific Flabellum by attaining 

a very large corallum size with up to 7 cycles of septa, having 

the largest face angle (44°-58°), and having very long lateral 

edges. 


Records: Soyo Maru-437, 1, TIUS 43448 (F. pavoninum of 

Yabe and Eguchi, 1942a) (Plate 31»; Soyo Marw-416, 1, TIUS 

50094 (F. p. magnificum of Yabe and Eguchi, 1942a). 

TYPES.—The holotype appears to be lost from the ZMB (see 

Cairns, 1989a). Type Locality: Valdivia-199: 0° 15.57*, 

98°04'E (off western Sumatra), 470 m. 

DISTRIBUTION.—Japan: Off Seto, Kii Strait, Honshu; off 

southeastern Shikoku; East China Sea off southwestern Kyushu 

(induing off Danjo Gunto); 307-700 m. Elsewhere: Philippines, 

Indonesia, west of Sumatra; 291-616 m. 

Flabellum (F.) angustum Yabe and Eguchi, 1942 

PLATE 31/ 

Flabellum disticlum angustum Yabe and Eguchi, 1942a:95, pi. 6: figs. 5-7; 

1942b: 131.—Eguchi, 1968:C4. 

DISCUSSION.—A new subspecies of F. distinctum was 

described by Yabe and Eguchi (1942a,b) based on 60 Pliocene 

coralla from Kochi-ken, Shikoku. Although a Recent specimen 

from Sagami Bay was also attributed to this taxon, Yabe and 

Eguchi (1942a) clearly indicated that the type locality 

corresponded to the fossil locality and illustrated only fossil 

specimens. Their rationale for the new subspecies was that the 

specimens had edge angles less than 70°. In fact, the edge angle 

of the fossil specimens is bimodal, ranging from 102°-123° 

near the pedicel to 52°-54° in upper corallum. The fossil 

specimens from Shikoku do appear to be a separate species, but 

the Recent specimen(s) from Sagami Bay were not illustrated 

and were not examined by the author. 


Records: Syntypes of F. p. angustum, TIUS. 

TYPES AND TYPE LOCALITIES.—Sixty syntypes (Plate 31/) 

from the Pliocene of Tonohama, Kochi-ken, Shikoku are 

deposited at the TIUS (43436). Additional Recent syntypes 

from Sagami Bay are also deposited at the TIUS (59333). 

DISTRIBUTION.—Known only from the type localities.

NUMBER 557 73 

Flabellum (F.) politum Cairns, 1989 

PLATE yia-c 

Flabellum pavoninum paripavoninum.—Yabe and Eguchi, 1942a:91-93 [in 

part: Soyo Afarn-419, pi. 5: fig. 8a-c]; 1942b: 129-130 [in part: pi. 11: fig. 

9a-c]. 

Flabellum politum Cairns, 1989a:53-54, pi. 28: figs, a-f [synonymy]. 

DISCUSSION.—The 10 specimens reported herein are either 

juvenile or poorly preserved coralla collected when dead. Aside 

from the distributional records they contribute nothing to our 

knowledge of this species. The reader is referred to Cairns 

(1989a) for a complete description and illustration of F. 

politum. It is distinguished from other Japanese species in the 

subgenus by having highly sinuous septa; a smooth, lustrous 

theca; and a very tall corallum, invariably taller than wide. 


USNM 40735; Alb4935, 1, USNM 40790; Alb-4936, 2, 

USNM 40736; TM (KT9015, CB1-2), 4, ORI; TM (KT9015, 

HK2), 1, USNM 92816; TM(KT9015, HK5), 1, USNM 92817. 

Previous Records: Types of F. politum, USNM. 

TYPES.—All types of F. politum are deposited at the USNM. 

Type Locality: Alb-5391: 12° 13' 15"N, 124°05'03"E (Philippines), 

216 m. 

DISTRIBUTION.—Japan: Eastern Channel of Korea Strait off 

southwestern Honshu; East China Sea off southern and 

southwestern Kyushu, including off Fukue Island and Osumi 

Retto; 70-402 m. Elsewhere: South China Sea, Philippines, 

Banda Sea; 40-717 m. 

Subgenus Flabellum (Ulocyathus) Sars, 1851 

DIAGNOSIS.—Flabellum having a serrate (jagged) calicular 

edge. 

TYPE SPECIES.—Ulocyathus arcticus Sars, 1851 (= Flabellum 

macandrewi Gray, 1849), by monotypy. 

Flabellum (U.) deludens Marenzeller, 1904 

PLATE 32d,e 

Flabellum japonicum.— Marenzeller, 1888a:45-46. 

Flabellum deludens Marenzeller, 19O4a:269-272, pi. 17: figs. 10, 10a.—Yabe 

and Eguchi, 1932a:387; 1942a:101-103, pi. 5: figs. 9a,c, 10a,c, lla,c; 

1942b: 135-136, pi. 12: fig. la-c—Eguchi, 1938: table 2; 1965:292. 2 figs; 

1968:C44~45, pi. C22: figs. 4, 5; pi. C25: figs. 3. 4.—Utinomi, 1965: 

256.—Eguchi and Miyawaki, 1975:58.—Cairns, I989a:55-56, pi. 29: figs. 

a-f [synonymy]. 

?Flabellum cf. apertum.—Yabe and Eguchi, 1942a:89. pi. 5: fig. 12a,c; 

1942b:136, pi. 12: fig. 3a-c. 

DESCRIPTION.—Corallum highly compressed, with virtually 

planar thecal faces joined at sharp, carinate thecal edges. Thecal 

edge crests up to 2 mm in height and usually extend from 

pedicel to calice. Angle of thecal edges, 115°-150°; inclination 

of lateral faces, 44°-80°. GCDrLCD = 1.3-1.5. Largest 

Japanese specimen examined (Alb-4915) 47.9 x 37.7 mm in 

calicular diameter, however, Cairns (1989a) reported a larger 

specimen from the Philippines of 53 mm GCD. Thecal faces 

flat (unridged) and bear a uniform coarse granulation in 

addition to chevron-shaped growth lines looped between all 

costae but the C5. Calicular edge deeply lacerate, a tall (up to 6 

mm), rectangular thecal extension corresponding to each Sy_2 

and adjacent pair of S4 (S5 rarely developed adjacent to S^_2). 

A smaller, triangular calicular apex up to 1.7 mm in height 

corresponds to each S3. Theca and septa fragile. Base color 

white, with reddish brown stripes corresponding to the C^_2 

and occasionally the C3; septal faces of S1 _2 also reddish brown 

near theca. 

Septa hexamerally arranged in 5 cycles, the fifth cycle never 

complete even in large specimens. Medium-sized specimens 

(about 35 mm GCD) often have only 4 cycles of septa; larger 

specimens have pairs of S5 that form adjacent to S3 up to a total 

of 72 total septa. S5 rarely form adjacent to S^. S-i_2 equal in 

size and have vertical, sinuous inner edges that define a deep, 

narrow fossa. S3 about three-quarters width of an S1 and also 

have sinuous inner edges. S4 half to three-quarters width of an 

S3 and have free inner edges, not fused to adjacent S3 or the 

columella. When present, S5 rudimentary. Lower, inner edges 

of S^ form an elongate columella; inner edges of S3 barely 

attain the columella. 

DISCUSSION.—Flabellum deludens is compared to F.japonicus 

in the account of the latter species. A more complete 

synonymy and description of this species is found in Cairns 

(1989a). 



USNM 40760; Alb-5094, 2, CAS 16310 and 74992; YO69-3, 

1, ORI; YO69-37-39, 1, USNM 92833; Tsuchida-197, 1, 

USNM 92834; TM (KT7802, Z4), 1, USNM 92835; TM 

(KT7811, OT1), 1, USNM 92836; TM (KT9015, BS2), 3, 

USNM 92837; TM (KT9202, OS3), 1, USNM 92838. Previous 

Records: Syntypes of F. deludens, ZMB; Soyo Maru-421,1, 

TIUS 43443 (F. cf. apertum of Yabe and Eguchi, 1942a,b); 

Philippine specimens (Cairns, 1989a), USNM. 

TYPES.—Two syntypes of F. deludens are deposited at the 

ZMB (#5086, 7086). Type Locality: Valdivia stations 185, 

203: west of Sumatra, 614-660 m. 

DISTRIBUTION.—Quite common off Pacific coast of Honshu 

(from 36°16'N southward), Shikoku, and Kyushu; East China 

Sea off southwestern Kyushu; Eastern Channel, Korea Strait; 

Sea of Japan off southwestern Honshu; 115-783 m. Elsewhere: 

Philippines, eastern Indian Ocean; 106-1035 m. 

Flabellum (U.) japonicum Moseley, 1881 

PLATE 32g,h 

Flabellum japonicum Moseley, 1881:168-169. pi. 7: figs. 4, 4a: pi. 16: fig. 

12.—Faustino, 1927:47-48, pi. 2: figs. 5, 6.—Yabe and Eguchi, 1942a:101. 

pi. 7: fig. lla.c; I942b:136, pi. 12: fig. 2a-c.—Eguchi, 1968:C45, pi. C28:

74 

figs. 2, 3.—Kikuchi, 1968:8—Fadlallah, 1983a: 135.—Cairns, 1989a:56- 

57, pi. 29: figs, g-i [not Alb-5083] [synonymy]. 

Not Flabellum japonicum.—Marenzeller. 1888a:45-46 [= F. deludens]. 

DESCRIPTION.—Corallum campanulate, but laterally compressed, 

with gently rounded, slightly convex thecal faces and 

a sharp, crested thecal edge. Angle of thecal edges, 90°-108°; 

inclination of lateral faces, 65°-88°. GCD:LCD = 1.3-1.4. 

Largest specimen known (YO69-37-39) 57.6 x 39.5 mm in 

calicular diameter and 37.0 mm in height. Thecal edge crests up 

to 2.0 mm in height, but rarely extend to the calice of a 

medium- to large-sized specimen. Pedicel size constant 

throughout ontogeny, 2.5 x 2.0 mm in diameter and 2.0 mm in 

height The 10 C,_2 occurring on thecal faces slightly raised as 

rounded ridges. Fine chevron-shaped thecal growth lines occur 

between all costae but C5; theca otherwise smooth, not 

granular. In well-preserved specimens, calicular edge regularly 

serrate, with an equilaterally triangular apex up to 3.5 mm in 

height corresponding to each S1-2, a smaller apex of about 1.3 

mm in height corresponding to each S3, and a very small apex 

about 0.5 mm in height corresponding to each S4. Theca and 

septa thin and fragile, theca of lower half of corallum often 

discolored. Upper, fresh theca white or reddish brown, with 

more intense pigmentation occuring in stripes overlaying the 

C1-2 and upper, inner (thecal) edges of S^. 


formula: S1_2>S3>S4»S5. S1-2 have vertical, highly sinuous 

inner edges that define a deep, elongate fossa. S3 about 

three-quarters width of an S1 and also have sinuous inner 

edges. S4 half to three-quarters width of an S3 and have sinuous 

inner edges that often loosely join to their adjacent S3 low in 

fossa. S5 rudimentary, extending only partially down inner 

theca. Lower, inner edges of S.|_2 form a rudimentary 

columella; inner edges of S3 barely attain columella. 

DISCUSSION.—Large, well-preserved specimens of F. 

japonicum are easily distinguished from F. deludens by having 

less serrate calicular margins (i.e., equilateral calicular apices 

vs rectangular). If the calicular edges are broken, which is 

common, F. japonicum can be distinguished by having less 

prominent edge crests near the calice; a less open corallum (a 

smaller edge angle); the presence of S5 adjacent to S,^; and 

smooth, nongranular theca. Small specimens (GCD < 20 mm) 

are virtually indistinguishable. Comparisons to the closley 

related F. apertum borealis are made in the account of that 

species. 

MATERIAL EXAMINED.—New Records: YO69-3, 3, 

USNM 92828; YO69-37-39, 3, USNM 92829, 2, ORI. 

Previous Records: One syntype of F. japonicum; specimens 

from off Japan and the Philippines reported by Cairns (1989a). 

TYPES.—Eight syntypes of F. japonicum are deposited at the 

BM (uncataloged). Type Locality: Challenger-232: 35° 1IX 

139°28'E(Sagami Bay), 631 m. 

DISTRIBUTION.—Japan: Off southeastern Honshu from Sagami 

Bay to off Owase; Kii Strait, Shikoku; off Amakusa 

Islands, Kyushu; 119-631 m. Elsewhere: Philippines, Indonesia; 

128-1141 m. 


Flabellum (U.) apertum borealis, subsp. nov. 

PLATE 32/,/-/ 

Not Flabellum cf. apertum.—Yabe and Eguchi, 1942a:89, pi. 5: fig. 12a-c; 

1942b: 136. pi. 12: fig. 3a-c [= IF. deludens]. 

Flabellum apertum.—Cheng, 1977:135-137,6 figs. 

Flabellum japonicum.—Cairns, 1989a:57 [in part: Alb-5083]. 

DESCRIPTION.—Corallum campanulate and laterally compressed, 

the thecal edges forming a sharp angle but not crested. 

Angle of thecal edges, 90°-112°; inclination of lateral faces, 

54°-75°. GCD:LCD = 1.17-1.32. Largest specimen examined 

(Alb-5086) 35.3 x 30.4 mm in calicular diameter and 31.6 mm 

in height. The 4 lateral C, are invariably ridged for a short 

distance about 9-14 mm above pedicel; C2 not expressed. In 

well-preserved specimens, calicular edge is serrate, having a 

tall, isosceles triangular apex up to 4 mm in height corresponding 

to every S1-2 anc * a smaller apex corresponding to each S3. 

Pedicel invariably eroded, the lower half to three-quarters of 

theca often discolored. Theca porcellaneous, without granulation; 

coralla often white or reddish brown with darker pigment 

overlaying the C1-2. 


formula: S1_2>S3>S4»S5, small specimens (GCD < 24 mm) 

have only 48 septa, but larger specimens (GCD > 30 mm) have 

a full fifth cycle. Su2 have vertical, sinuous inner edges that 

define a very narrow, elongate fossa. S3 two-thirds to 

three-quarters width of S^ and also have sinuous inner edges 

but do not fuse with the columella. S4 about half width of an S3 

and do not fuse to adjacent S3. S5 rudimentary. Lower, inner 

edges of S^_2 form the rudimentary columella. 

DISCUSSION.—This subspecies is similar to F. apertum in 

that both taxa have campanulate coralla and 6 ridged (but not 

crested) Cv The Japanese subspecies differs in its tendency to 

have an additional cycle of septa (S5) and thus more closely 

spaced septa. Flabellum apertum borealis also has a lesser 

upward inflection of its thecal faces above the ridged C^ F. 

apertum apertum having a fuller, more campanulate corallum. 

Given the similarity of this form to typical F. apertum, but the 

disjunct distribution of the two forms (F. apertum is known 

only from the Subantarctic), this taxon is described as a 

subspecies of Flabellum apertum. 

Flabellum apertum borealis could easily be confused with F. 

japonicum, but differs by (1) having protuberant (ridged) Cv 

(2) lacking edge crests, (3) having S3 that do not extend to the 

columella, (4) having a sharper calicular serration (isosceles 

apices vs equilateral apices), and 5) living at a greater depth. 

ETYMOLOGY.—The subspecies name borealis (Latin borealis, 

meaning "boreal" or "northern"), refers to the disjunct 

northern distribution of this subspecies. 

MATERIAL EXAMINED AND TYPES.—Holotype: Alb-5086,1, 

USNM 40710 (Plate 32/,/,/). Paratypes: Alb-4960, 1, USNM 

40703; Alb-4969, 1, CAS 74943; Alb-4973, 2, USNM 40705; 

Alb-5083, 1, USNM 40709, (Plate 32;,*;; off Jogashima, 

Sagami Bay, depth unknown, 1, USNM 92841, 1, ORI; TM 

(KT9202, ATI), 1, USNM 92842; TM (KT9202, YT5), 6,

NUMBER 557 75 

USNM 92843; TM (KT9202, YT6), 1, USNM 92844. 

Nontypes: Alb-5054, 4, USNM 40706. Type Locality: Alb- 

5086: Z5°Q%^, 139°20'E (Sagami Bay), 534 m. 

DISTRIBUTION.—Japan from Sagami Bay to southwestern 

Kyushu, including Bungo Strait and northern Ryukyu Islands; 

southwest of Ishigaki Islands (east of Taiwan); 307-1141 m. 

Distribution of Flabellum apertum apertum is circumsubantarctic 

at 220-1500 m (Cairns, 1982). 

Truncatoflabellum Cairns, 1989a 

DIAGNOSIS.—Like Flabellum, but also reproducing asexually 

by transverse division resulting in a distal anthocyathus 

budded from a basal anthocaulus. Upper calicular margin 

smooth to slightly serrate. Most species with one or more pairs 

of thecal edge spines or edge crests on anthocyathus and one 

pair of spines on anthocaulus. Pali absent; columella rudimentary. 

DISCUSSION.—Yabe and Eguchi (1942a,b) reported Recent 

Flabellum rubrum (Quoy and Gaimard, 1833) and three of its 

subspecies (stokesii, debile, and candeanum) from approximately 

35 localities in Japanese waters (88-353 m); however, 

Squires (1963) convincingly demonstrated that F. rubrum is 

endemic to New Zealand waters, the species later being 

transferred to the genus Monomyces (see Caims, 1989a). After 

examining about one-third of Yabe and Eguchi's (1942a,b) 

specimens off. rubrum, I (Caims, 1989a) concluded that they 

comprised at least four species: Truncatoflabellum spheniscus, 

T. formosum, T. candeanum, and T. carinatum. Other records 

of Japanese F. rubrum by Yabe and Eguchi (1932b), Eguchi 

Key to the Temperate Northwest Pacific Truncatoflabellum 

(1938), Utinomi (1965), Kikuchi (1968), Eguchi (1968), and 

Eguchi and Miyawaki (1975) may also represent a mixture of 

species. 

In similar manner, Flabellum transversale Moseley, 1881 

was widely reported from Japanese waters as the nominate 

species and two subspecies (conicum Yabe and Eguchi, 1942a; 

and triangulare Eguchi, 1968) by Yabe and Eguchi (1942a), 

Eguchi (1968) and others: Yabe and Eguchi (1932a,b, 1941b, 

1942b), Eguchi (1938, 1965), Mori (1964), and Eguchi and 

Miyawaki (1975). However, F. transversale appears to be 

endemic to the coast of Victoria, Australia (Cairns and Parker, 

1992) and the Japanese records pertain to other species. 

Although I have not examined all of the records (summarized 

by Eguchi, 1968), the specimens reported as typical F. 

transversale (38-219 m) appear to represent at least three 

species: an undetermined fossil species of Truncatoflabellum 

(Yabe and Eguchi, 1942a, pi. 7: figs. 1-5,12), an undetermined 

fossil species of F. (Flabellum) (Yabe and Eguchi, 1942a, pi. 7: 

fig. 6), and an undescribed Recent species of Truncatoflabellum 

(Yabe and Eguchi, 1942a, pi. 7: fig. 9a-c) similar to T. 

formosum. Flabellum transversale belongs to the nominate 

subgenus of Flabellum. The specimens identified as subspecies 

F. transversale conicum (70-219 m) (Plate 4\e,i) and F. 

transversale triangulare (50-60 m) all appear to be juvenile 

specimens of undetermined species of Flabellum (Flabellum) 

or perhaps the anthocauli of a Truncatoflabellum. A precise 

identification is not attempted. 

TYPE SPECIES.—Euphyllia spheniscus Dana, 1846, by 


1. Edges of corallum bear one or more pairs of spines 2 

Edges of corallum do not bear spines: edges may bear crests or lack both spines and 

crests 5 

2. Calicular edge slightly serrate; inclination of lateral faces (FAN) = 30°-41°. . . . 

T. candeanum 

Calicular edge smooth; face angle < 25° 3 

3. Corallum small (GCD < 8 mm); calices having 32 septa; thecal edges virtually 

parallel T. sp. B 

Corallum larger (GCD up to 50 mm); calices having 80-192 septa; thecal edge angle 

37°-88° 4 

4. Calice elongate (GCD:LCD = 2.3-3.4), containing 6 cycles of hexamerally arranged 

septa (192) T. sphenicus 

Calice elliptical (GCD:LCD = 1.4-1.9), containing 80 septa arranged in 3 size classes 

T. formosum 

5. Edges of coralla without crests; lives at depth greater than 900 m T. sp. A 

Edges of coralla bear 1 or more crests; lives at less than 300 m 6 

6. Coralla have five cycles of septa (96) and several pairs of edge crests; edge angle 

35-57 T. carinatum 

Coralla with fifth cycle of septa absent or incomplete (48-66 septa) and only 1 pair 

of basal edge crests; edges virtually parallel T. gardineri

76 

TruncatoflabeUum spheniscus (Dana, 1846) 

PLATE 33a-d 

Euphyllia spheniscus Dana, 1846:160-161, pi. 6: fig. la-e. 

Flabellum debile Milne Edwards and Haime, 1848a:274. 

Flabellum bairdi Milne Edwards and Haime, 1848a:274-275 [new synonymy]; 

1857:93.—Cairns, 1989a:66-67, pi. 33: fig. k; pi. 34: figs. a-c. 

Flabellum profundum Milne Edwards and Haime, 1848a:276 [new synonym]; 

1857:93-94, pi. Dl: figs. 5, 5a.—Cairns, 1989a:67, pi. 34: figs. d-h. 

Flabellum crenulatum Milne Edwards and Haime. 1848a:277 [new synonym]; 

1857:95.—Yabe and Eguchi, 1942b:134. 

?Flabellum sumatrense Milne Edwards and Haime, 1848a:27l; 1857:89. 

Flabellum affine Milne Edwards and Haime. 1848a:274. 

?Flabellum stokesi.—Yabe and Eguchi, 1932b:443. 

Flabellum rubrum stokesii.—Yabe and Eguchi, 1942a:98-99 [in part: pi. 8: fig. 

5a-c]. 

?Flabellum cf. multijore.—Yabe and Eguchi, 1942a: 103, pi. 6: fig. 8a-c; 

1942b: 136. 

Flabellum rubrum debile.—Yabe and Eguchi, 1942b: 132-133 [in part: pi. 11: 

fig. 15a-c].—NotKikuchi, 1968:8, pi. 5: fig. 13 [= TruncatoflabeUum]. 

TruncatoflabeUum spheniscus.—Cairns, 1989a:65-66, pi. 32: figs, g-k 

[synonymy]. 

DESCRIPTION.—Anthocyathus robust and highly compressed 

(GCD:LCD = 2.3-3.4), the planar thecal faces meeting 

in rounded thecal edges that bear 1 or 2 pairs of thecal spines. 

Angle of thecal edges large, 57°-88°; inclination of lateral 

faces narrow, 20°-31°. Largest specimen known (holotype of 

F. profundum) 50.0 x 21.0 mm in calicular diameter and 35.1 

mm in height. Basal scar of anthocyathus usually large: 

5.3-7.3 x 11.5-13.5 mm in diameter. Upper calicular margin 

evenly arched and smooth to very finely serrate. In wellpreserved 

specimens fine chevron-shaped thecal growth lines 

form apices at every S.,_5. In other specimens (e.g., types of F. 

profundum) growth lines are lacking but coarse longitudinal 

costae corresponding to C^_3 are present. Theca reddish brown 

to blackish brown in color, concentrating in bands parallel to 

calicular margin. 

Septa essentially hexamerally arranged in 6 cycles (192 

septa), larger specimen having several pairs of S7 in end 

half-systems, and smaller specimens having developmental 

stages of 40, 42, 44, or 46 primary septa (S^), with a 

corresponding number of S5 and S6, resulting in 160,168, 176, 

or 184 septa. Primary septa (S1_4) slightly concave (notched) 

near calicular margin but project well into fossa and have 

slightly sinuous inner edges that thicken near the columella. S4 

of small- to medium-sized specimens sometimes slightly less 

wide than the S^_3, but their lower, inner edges also attain the 

columella. Secondary septa (S5) only about half the width of a 

secondary and do not attain the columella. Tertiary septa (= S6) 

rudimentary, only about one-quarter width of a secondary. 

Fossa deep and elongate, containing a well-developed columella 

about 1.1-1.5 mm wide composed of the fusion of the 

lower, inner edges of the S1^. 

DISCUSSION.—I (Cairns, 1989a) previously included Flabellum 

debile and F. affine in the synonymy of TruncatoflabeUum 

spheniscus, but treated TruncatoflabeUum profundum and T. 

bairdi as separate species based on their lesser number of septa 


and/or thecal texture. These latter two species were known 

from very few specimens, essentially their type series, which 

led to an unavoidably typological characterization. A lot of 13 

specimens from the Formosa Strait (CAS 74990), however, 

allowed a study of variation as it related to corallum size, which 

demonstrated the developmental stages of septal insertion 

characteristic of the other nominate species. 

TruncatoflabeUum spheniscus is distinguished from the 

other Indo-West Pacific congeners by its high thecal edge angle 

(57°-88°)and its narrow, elongate calice that results in the high 

GCD:LCD of 2.3-3.4. 


CB-00-1), 2, USNM 92796; TM (KT9015, CB-02-2), 12, 

USNM 92798; TM (KT9015, HK2), 2, USNM 92799; TM 

(KT9202, CB1-2), 17, USNM 92797; TM (KT9202, YT1), 5, 

USNM 92800; TM (KT9202, YT2), 5, USNM 92801; off 

Pescadores Islands, Taiwan Strait, 30-50 m, 5 May 1972, 13, 

CAS 74990. Previous Records: 4 syntypes of F. spheniscus, 

USNM; holotypes of F. bairdi, F. profundum, F. crenulatum, 

and F. sumatrense; Soyo Maru-339, 1, TIUS 50228 (F. rubrum 

stokesi of Yabe and Eguchi, 1942a); F. cfr. multifore of Yabe 

and Eguchi (1942a), TIUS 41052. 

TYPES.—The four syntypes of Euphyllia spheniscus are 

deposited at the USNM (89, 91-93). Type Locality: Off 

Singapore, 3-6 m. 

The illustrated syntype of F. debile is deposited at the BM 

(1855.12.27.2). Type Locality: Philippines, depth unknown. 

The holotype of F. profundum is deposited at the MNHNP 

(1026). Type Locality: "Off China," depth unknown. 

The holotype of F. crenulatum is deposited at the MNHNP 

(1020). Type Locality: Unknown. 

The holotype of F. sumatrense is deposited at the MNHNP 

(374). Type Locality: Off Sumatra, depth unknown. 

The holotype of F. affine appears to be lost. Type 

Locality: Sir Charles Hardy Island, Torres Strait, depth 

unknown. 

DISTRIBUTION.—Japan: Tosa Bay, Shikoku; Eastern Channel, 

Korea Strait, Honshu; Osumi Shoto, northern Ryukyu 

Islands; 66-106 m. Elsewhere: Formosa Strait, South China 

Sea; off Singapore; Philippines; Torres Strait; ?off Sumatra; 

2-36 m. 

TruncatoflabeUum candeanum (Milne Edwards 

and Haime, 1848) 

PLATE 33

NUMBER 557 77 

inclination of convex thecal faces 30°-41°. Large specimens up 

to 32 x 16 mm in calicular diameter and 22 mm in height. Two 

or three pairs of long thecal edge spines on anthocyathus. Basal 

scar 4-6 x 3-4 mm in diameter. GCD:LCD = 1.6-1.7. Calicular 

edge slightly scalloped, the apices corresponding to the primary 

septa. Septa arranged in 3 size classes, usually 20:20:40 (80 

septa). Columella well developed. 

DISCUSSION.—Based on the six small specimens reported 

herein little can be added to my previous redescription of this 

species (Cairns, 1989a) except that the anthocyathus basal scar 

may attain a greater diameter of up to 8 mm. 

Truncatoflabellum candeanum is distinguished from its 

congeners by its very long and broad edge spines; its slightly 

scalloped calicular margin; its relatively wide thecal face angle 

(30°-41°);and its striped reddish brown corallum (see Cairns, 

1989a, table 6). 


HK3), 3 including 2 anthocauli, USNM 92802, 2, ORI; TM 

(KT9015, HK5), 1 anthocyathus, USNM 92803. Previous 

Records: Neotype of F. candeanum; holotype of F. elegans; 

F. candeanum of Marcnzeller (1888a), 1, ZMB 8194; Soyo 

Maru-422 (2) and 465 (Plate 33e)(3 of 4 mentioned), TIUS 

50233 and 50229, respectively (F. rubrum of Yabe and Eguchi, 

1942a); Philippine specimens of Cairns (1989a). 

TYPES.—The neotype of Flabellum candeanum, designated 

by Cairns (1989a:71), is deposited at the USNM (81963). Type 

Locality: Alb-5369: 13°48'N, 121°43'E (Tayabas Bay, 

Luzon, Philippines), 194 m. 

The holotype of F. elegans is deposited at the BM 

(1846.7.1.58). Type Locality: ?Japan, depth unknown. 

DISTRIBUTION.—Japan: Eastern Channel, Korea Strait, Honshu; 

northwestern Kyushu; off Kagoshima, southwestern 

Kyushu; reports off Shikoku by Yabe and Eguchi (1942b) and 

Kikuchi (1968) are not verified; north of Pratas Islands, South 

China Sea; 88-223 m. Elsewhere: Philippines, Celebes Sea; 

70-249 m. 

Truncatoflabellum formosum Cairns, 1989 

PLATE 33/?,/r 

Flabellum rubrum.—Yabe and Eguchi, 1942a:96-98 [in part: form A, pi. 8: 

figs. 16-19, 23, 24]; 1942b:131-132 [in part: Soyo Maru-107, 540, pi. 11: 

fig. 14]. 

Truncatoflabellum formosum Cairns, 1989a:69-70, pi. 35: figs, j.k; pi. 36: figs. 

a,b.—Cairns and Keller, 1993:265. figs. 101, 1 lA. 

Truncatoflabellum n. sp.: Cairns, 1989a:73, pi. 38: figs. g.h. 

DESCRIPTION.—Corallum compressed, the convex thecal 

faces meeting in rounded thecal edges. Angle of thecal edges, 

37°-48°; inclination of thecal faces, 18°-25°. Largest specimen 

known (anthocyathus from TM (CR79-18)) 26.7 x 14.5 mm in 

calicular diameter and 28.9 mm in height, having a basal scar of 

only 3.3 x 4.6 mm. Two pairs of edge spines usually present, 

the lower pair cylindrical in cross section and pointed 

downward, the upper pair much more broadly based and 

oriented perpendicular to theca. GCD:LCD = 1.42-1.85. Theca 

reddish brown to white, bearing very fine transverse (not 

chevron-shaped) epithecal corrugations. Inside calice, 1 -2 mm 

below calicular edge, theca adjacent to each of the 20 primary 

septa is lightly pigmented against a white background, giving 

the impression of a circumferential band within the theca. 

Calicular margin smooth. 

Septa arranged in 3 distinct size classes in all specimens 

examined regardless of calicular diameter, accordingly: 

20:20:40 (80 total septa). Primary septa nonexsert, their upper, 

outer margins meeting the thecal wall in a broad, graceful arch. 

Inner edges of primary septa vertical and slightly sinuous, 

merging with the columella. Secondary septa half to two-thirds 

width of primaries, their inner edges less sinuous and not 

attaining the columella. Tertiary septa only one-quarter to 

one-third width of a secondary. Fossa deep and narrow, 

containing an elongate, solid columella formed from fusion of 

lower, inner edges of 20 primary septa. 

DISCUSSION.—Truncatoflabellum formosum differs from 

other species in the genus by having a septal arrangement of 

20:20:40 and having attenuate primary septa. I (Cairns, 

1989a:69) previously indicated that one of Yabe and Eguchi's 

(1942a, pi. 8: fig. 14) specimens of F. rubrum was T. 

formosum, but, this synonymy should have read: pi. 8: fig. 

16a-c. 


USNM 40789; Alb-4937, 1, USNM 40791; TM (CR79-18), 1, 

USNM 92804. Pleistocene of Ryukyu Islands, 1, TIUS. 

Previous Records: Types of T. formosum, USNM; Flabellum 

rubrum form A of Yabe and Eguchi (1942a,b): Soyo Maru-465, 

1, TIUS 50229; Soyo Maru-\0% 1, TIUS 50231; ISoyo 

Marw-540, I, TIUS 50230. 

TYPES.—The type series of T. formosum is deposited at the 

USNM. Type Locality: Alb-5249: 7 o 06 / 06"N, 125°40 / 08"E 

(Daveo Gulf, off Mindinao, Philippines), 42 m. 

DISTRIBUTION.—Japan: Off Boso Hanto, Honshu; off Kii 

Peninsula, Honshu; Tosa Bay, Shikoku; East China Sea off 

southwestern Kyushu; Eastern Channel, Korea Strait; Sea of 

Japan, Wakasa Bay, Honshu; 106-210 m. Elsewhere: Philippines, 

Celebes Sea, southwest Indian Ocean; 37-933 m. 

Truncatoflabellum carinatum Cairns, 1989 

PLATE 33/,* 

Flabellum rubrum.—Faustino, 1927:53 [in part: Alb-5311].—Yabe and 

Eguchi. 1942a:96-98 [in part: Soyo Maru 222, pi. 8: figs. 6-12, 20]; 

1942b:131-132 [in part]. 

Flabellum rubrum stokesii.—Yabe and Eguchi, 1942a:98-99 [in part Soyo 

Maru-295. 304]. 

Truncatoflabellum carinatum Cairns, 1989a:73-74, pi. 38: figs. b-e. 

DESCRIPTION.—The following description is based on one 

well-preserved specimen from off the Pescadores Islands (ex 

CAS 74990). Corallum (anthocyathus) compressed, the evenly 

convex thecal faces meeting in an acute angle at thecal edges. 

Three or 4 pairs of low (about 1.0 mm in height) discontinuous

78 

crests or spurs (not spines) occur along each thecal edge. 

Corallum 26.7 x 14.4 mm in calicular diameter, 22.5 mm in 

height, and 3.7 x 6.2 mm in basal scar diameter, making it the 

largest specimen known. Edge angle 57°; inclination of lateral 

faces, 33°. Theca white and somewhat rough, displaying 

chevron-shaped epithecal growth lines that peak at the 24 S1-3. 

Septa hexamerally arranged in 5 complete cycles, with 4 

additional pairs of S6 in quarter-systems adjacent to the 2 

principal septa (104 septa). S^g have vertical, sinuous inner 

edges that attain the columella. Septal faces distinctively lined 

with 15-18 low ridges oriented perpendicular to the septal 

edge, each ridge bearing a series of low, triangular granules. 

Ridges alternate in position with those on opposite face of same 

septum. S4 about two-thirds width of an S^ and have poorly 

formed, finely dentate inner edges that do not fuse with the 

columella. S5 quite small, only about one-quarter width of an 

S4. The 4 pairs of S6 are rudimentary. Fossa deep and elongate, 

containing a well-formed, elongate columella about 2.1 mm 

wide. 

DISCUSSION.—All previously reported specimens of this 

species were poorly preserved, either represented as fossils or 

Recent specimens that were dead when collected. The 

specimen described above is the largest known specimen and is 

well preserved, making possible an improvement of the 

original description. Truncatoflabellum carinatum is distinguished 

from most congeners by having nonspinose but acutely 

angled and crested thecal edges; a very small face angle, 

especially near the basal scar, and a rather small basal scar. The 

edge spines attributed to this species by Cairns (1989a) are 

more accurately interpreted as low, discontinuous edge crests. 

MATERIAL EXAMINED.—New Records: AIb-5311, 6, 

USNM 40787; off Pescadores Islands, Formosa Strait, South 

China Sea, 30-50 m, 1, ex CAS 74990. Previous Records: 

Types of T. carinatum; F. rubrum of Yabe and Eguchi 

(1942a,b): Pleistocene of Tonbe, Sizuoko-ken, 2, TIUS 29233; 

Pliocene of Koti-ken, 1, TIUS 43434; F. rubrum stokesii of 

Yabe and Eguchi (1942a), Soyo Maru-295, 1, TIUS 51220; F. 

rubrum of Yabe and Eguchi (1942a), Soyo Maru-222, 3, TIUS 

50232. 

TYPES.—The holotype and most paratypes of T. carinatum 

are deposited at the USNM; 1 paratype is also at the Australian 

Museum. Type Locality: Alb-5312:21 °30 / N, 116°32'E (north 

of Pratas Island, South China Sea), 256 m. 

DISTRIBUTION.—Recent: Formosa Strait north of Pratas 

Islands, South China Sea; southeastern coast of Kyushu; Tusa 

Bay, Shikoku; 30-274 m. Pliocene-Pleistocene of Honshu and 

Shikoku, Japan. 

Truncatoflabellum gardineri Cairns, 1993 

PLATE 34ajj 

Truncatoflabellum gardineri Cairns in Cairns and Keller. 1993:266, figs. 

11B-D. 

DESCRIPTION.—Coralla elongate and compressed, having a 

relatively low thecal edge angle of 21°-39°, the thecal edges 


becoming almost parallel in upper region of some coralla. 

Inclination of thecal faces, 18°-23°. Largest specimen from 

Japan (TM (KT9202, OS3) 16.7 x 11.3 mm in calicular 

diameter and 19.8 mm in height, with a basal scar diameter of 

3.7 x 4.5 mm. Calicular edge smooth; GCDrLCD = 1.30-1.48. 

Basal scar of anthocyathus rather small and elliptical, almost 

circular, ranging from 4.5-4.7 x 3.3-3.7 mm in diameter. 

Thecal edges rounded except for region approximately 3-10 

mm above basal scar, which extends as a single narrow crest up 

to 2.5 mm in height. Theca bears narrow longitudinal striae 

corresponding to each septum as well as chevron-shaped 

growth lines that peak at all but last septal cycle. In one 

well-preserved corallum, the theca was reddish brown with an 

indication of longitudinal striping along the major septa. 

Septa hexamerally arranged in 4 complete to 5 incomplete 

cycles according to the formula: S1_2>S3>S4>S5. Coralla less 

than 11 mm GCD usually have 48 septa arranged in 4 complete 

cycles. Between a GCD of 11-14 mm pairs of S5 begin to 

appear in the quarter-systems directly adjacent to the 2 

principal septa (50-56 septa) with a corresponding increase in 

the width of the flanked S4 to the size of an S2 and the 

acceleration of the adjacent S3 to the size of an Su2. Above a 

GCD of 15 mm additional pairs of S5 may occur in the 

remainder of the end half-systems and even quarter-systems 

adjacent to those, up to a total of 66 septa. S,_2 w 'de and have 

thickened, sinuous lower inner edges that fuse with the 

columella. S3 about half width of S1-2 and do not attain the 

columella, unless it is adjacent to an S5 (see above), in which 

case it is almost as wide as an S1-2 and fuses with the 

columella. S4 small, only about one third width of an S3, larger 

if flanked by a pair of S5. S5 rudimentary. Septa widely spaced, 

separated from one another by 2-3 septal thicknesses. Fossa 

deep and narrow, containing a well-developed elongate fossa 

about 1.8 mm wide. 

DISCUSSION.—The only differences between the Japanese 

specimens of T. gardineri and the type series of that species 

from off Durban, South Africa is that the Japanese specimens 

are larger and thus have more septa. The largest South African 

specimens of 11-12 mm GCD have 56 septa, which is 

consistent with the number of septa of a Japanese specimen of 

equivalent size. 

Truncatoflabellum gardineri is most similar to T. carinatum, 

both species having acutely angled thecal edges that bear low 

crests (not spines), but T. gardineri can be distinguished by 

having only one pair of edge spines, virtually parallel thecal 

edges, and fewer septa (see Key 2). 


CB6-1), 1, USNM 92807; TM (KT9202, KB3), 3, USNM 

92808; TM (KT9202, OS3), 17, USNM 92809, 2, ORI. 

Previous Records: Types of T. gardineri. 

TYPES.—The holotype and most paratypes of T. gardineri 

are deposited at the USNM (91736, 91737); 3 paratypes are 

also deposited at the South African Museum. Type Locality: 

Anton Bruun-390S: 29°34'S, 31 ^'E (off Durban, South 

Africa), 138 m.

NUMBER 557 79 

DISTRIBUTION.—Japan: Oki Strait, Honshu, Sea of Japan; 

Osumi Strait, Kyushu; 100-144 m. Elsewhere: Off Durban, 

South Africa; 138 m. 

Truncatoflabellum sp. A 

PLATE 34C -e 

?Flabellum stabile Marenzeller, 1904a:273-274, pi. 17: fig. 12.—Zibrowius, 

1980:150. 

?Truncatoflabellum stabile.—Cairns, 1989a:61.—Zibrowius and Gili, 

1990:39. 

Truncatoflabellum sp. cf. T. stabile.—Cairns and Keller, 1993:264-265, fig. 

10C.F. 

DESCRIPTION BASED ON 2 SPECIMENS FROM TM (KT9202, 

YT6).—Angle of straight thecal edges 60°-65°; inclination of 

slightly convex thecal faces, 33°-34°. Thecal faces meet in 

acutely angled edges that are quite long but that lack spines and 

crests. Largest specimen: 42.6 x 26.5 mm in calicular diameter 

and 40.5 mm in height, with a basal scar diameter of 2.1 x 6.0 

mm. Calicular edge arched and finely serrate; GCD:LCD of 

both specimens 1.61; GCD:H = 1.05-1.07. Basal scar small in 

relation to corallum size: 2.1-2.6 x 5.0-6.0 mm. Even though 

these specimens appear to have been collected alive, the theca 

is quite worn and chalky in texture. 


to the formula: S1_3>S4>S5, both specimens having 96 septa. 

S^g wide, their lower, inner edges thickened and sinuous, 

fusing with the medial columella. S4 about three-quarters width 

of an S^_3 and do not fuse with the columella. S5 small, only 

about one-quarter width of an S4. Fossa deep and elongate, 

containing a well-developed columella about 3 mm in width. 

DISCUSSION.—Five species of Truncatoflabellum are 

characterized as having nonspinose and noncrested thecal 

edges: T. paripavoninum (Alcock, 1898); 71 stabile (Marenzeller, 

1904a); 71 inconstans (Marenzeller, 1904a); 71 trapezoideum 

(Keller, 1981b); and Truncatoflabellum sp. (Zibrowius 

and Gili, 1990). All but 71 inconstans are found exclusively in 

deep water (600-3000 m). Having five cycles of septa, the 

Japanese specimens must be compared to T. trapezoideum and 

71 stabile, the other species having either four or six cycles of 

septa. The Japanese specimens differ from 71 trapezoideum in 

having a much smaller basal scar, nonridged costae; rounded 

thecal edges; and a much taller corallum (having a lower 

GCD:H ratio, 1.05-1.07 vs about 1.20 for F. trapezoideum). 

They are, however, very similar to 71 stabile, especially to the 

specimen reported from off Mozambique by Cairns and Keller 

(1993). Because the types of F. stabile are lost and there are few 

other specimens of this species with which to compare, the 

identity of the Japanese specimens remains uncertain. 


YT6), 2, USNM 92810. Previous Records: Specimen reported 

by Cairns and Keller (1993), IOM. 

TYPES.—The two syntypes of Flabellum stabile are presumed 

to be lost. Type Locality: Valdivia-31: 16°14'01"N, 

22°38'03*W (Cape Verde Islands), 1694 m. 

DISTRIBUTION.—Japan: Off Osumi Shoto, northern Ryukyu 

Islands; 964-1031 m. Elsewhere: Off Cape Verde, Selvagems, 

and Madeira Islands; off southeastern Mozambique; 

1450-3010 m. 

Truncatoflabellum sp. B 

PLATE 33/,/ 

DESCRIPTION OF SPECIMEN FROM TM (KT9202, TY1).— 

Corallum 5.9 x 3.1 mm in calicular diameter (GCD:LCD = 1.9) 

and 17.8 mm in height, with a basal scar diameter of 3.7 x 2.3 

mm. Thecal edges rounded and straight, virtually parallel to 

one another as are the convex thecal faces. Thecal edges bear 6 

pairs of small (0.4 mm in diameter), hollow spines. Thecal 

faces porcellaneous and reddish brown in color, the pigmentation 

stonger in crescent-shaped bands parallel to upper calicular 

edge. 

Septa hexamerally arranged in 3 cycles plus 4 pairs of S4 in 

end half-systems for a total of 32 septa. S^_2 equal in size and 

have sinuous inner edges that reach the columella. S3 half 

width of Su2 and do not reach the columella unless they are 

flanked by a pair of S4, in which case they are the same width 

of an S^g. S4 small, only about one-quarter width of an S3. 

Columella robust, about 1.1 mm wide. 

DISCUSSION.—This species is distinguished from its congeners 

by its virtually parallel thecal edges and elongate corallum. 

Together with additional specimens from the Kermadec 

Islands, these specimens will form the basis of the description 

of a new species (Caims, in prep.). 

MATERIAL EXAMINED.—TM (KT9202, YT1), 1, USNM 

92811. 

DISTRIBUTION.—Japan: Osumi Shoto, northern Ryukyu 

Islands; 80-88 m. Elsewhere: Kermadec Islands. 

Placotrochides Alcock, 1902b 

DIAGNOSIS.—Corallum compressed-cylindrical; transverse 

division present, resulting in an anthocyathus with a basal scar 

almost as large at its calicular diameter. Thecal spines absent. 

Three to four cycles of nonexsert septa; calicular edge smooth. 

Columella well developed, trabecular. 

TYPE SPECIES.—Placotrochides scaphula Alcock, 1902b, 

by subsequent designation (Wells, 1936). 

Placotrochides scaphula Alcock, 1902 

PLATE 34/-* 

Placotrochides scaphula Alcock, 1902b:121-122; 1902c:34, pi. 4: figs. 32, 

32a.—Cairns, 1989a:78-79, pi. 40: fig. 1; pi. 41: figs, a-e [synonymy].— 

Caims and Keller, 1993:272-273, fig. 12D.G. 

DESCRIPTION.—Corallum shaped like a flattened cylinder, 

with virtually parallel thecal faces and parallel, rounded thecal 

edges. No edge spines. Largest Japanese specimen 10.0 x 5.4 

mm in calicular diameter and 9.4 mm in height, with a basal 

scar measuring 9.5 x 5.3 mm, almost as large as the calice.

80 

Theca marked by thin (0.05 mm wide), shallow, vertical striae 

that delimit wide (0.5-0.6 mm), flat costae. 


formula: S1_2>S3>S4, 4 pairs of S4 often lacking resulting in 

40 septa. S^g about one-third LCD in width, their vertical, 

slightly sinuous lower, inner edges fusing to the columella. S3 

unflanked by S4 only about one-quarter width of an S, and 

have finely serrate inner edges; however, if S3 are flanked by a 

pair of S4, they are about three-quaters width of an S1 and have 

smooth inner edges that fuse with the columella. S4 rudimentary, 

only about 0.3-0.4 mm wide, and have finely serrate inner 

edges. All septa relatively thin (S^_2 about 0.2 mm thick) and 

widely spaced, i.e., 0.5-0.6 mm from one another. Fossa deep 

and elongate, containing a well-developed, elongate trabecular 

columella that occupies the medial third of fossa. 

DISCUSSION.—Placotrochides is very similar to 

Truncatoflabellum, but can be distinguished by its more robust 

columella, and by having virtually parallel thecal edges and 

faces. Only one other species is known in the genus, P. frustum 

Cairns, 1979 (nom. correct), which is known only from the 

Atlantic. It is distinguished by its smaller corallum size and 

fewer (^26) number of septa. 

MATERIAL EXAMINED.—Ate* Records: TM (KT7911, 

OT4), 5,USNM 92748; TM (KT9202, YT1), 2, USNM 92749, 

1, ORI. Previous Records: Holotype of P. scaphula; Philippine 

specimens (Cairns, 1989a). 

TYPES.—The holotype of P. scaphula is deposited at the 

ZMA (Coel. 1094). Type Locality: Sihoga-2\2: 5°54.5'S, 

120°19.2'E (Flores Sea), 462 m. 

DISTRIBUTION.—Japan: Suruga Bay, Honshu; Osumi Shoto, 

northern Ryukyu Islands (first records for Japan); 80-457 m. 

Elsewhere: Philippines, Flores Sea; southwest Indian Ocean; 

462-1628 m. 


Javania Duncan, 1876 

Javania insignis Duncan, 1876 

PLATE 34/-* 

Javania insignis Duncan, 1876:435. pi. 39: figs. 11-13.—Zibrowius, 1974b: 

8-9. pi. 1: figs. 1-6.—Cairns, 1989a:77-78, pi. 40: figs. d,e,g,h.j,k 

[synonymy].—Caims and Keller, 1993:272. 

Flabellum weberi Alcock, 1902a: 107. 

Desmophyllum cf. insigne.—Yabe and Eguchi, 1942b: 115, pi. 9: figs. 5, 6. 

Desmophyllum insignis.—Eguchi. 1965:290, 2 figs.; 1968:C41-42, pi. C9: 

figs. 4-9.—Song. 1982:136, pi. 2: figs. 5, 6; 1988:27-28, pi. 3: figs. 9-11; 

1991:134. 

DESCRIPTION.—Corallum ceratoid and robust, with thick 

theca and septa, and a massive, stereome-reinforced pedicel 

24%-55% of GCD. Largest Japanese specimen examined 

(ZMC: Fukue Jima) 32.7 x 24.9 mm in calicular diameter and 

56.2 mm in height, with a pedicel 7.8 mm in diameter, although 

a slightly larger specimen was reported by Cairns (1984) from 

Christmas Island. Theca milky white and smooth, sometimes 

porcellaneous. 



to the formula: S1_2>S3>S4>S5 (96 septa). S^ exsert as much 

as 4 mm and have quite thick, straight, vertical inner edges that 

disappear into the fossa. S3 exsert as much as 2.5 mm, about 

four-fifths width of an S^ and have relatively thin inner edges. 

S4_5 nonexsert, the S4 only about one-third width of an S3, the 

S5 about half width of an S4. S1 _2 often so thick that their 

adjacent S5 are fused to them. Fossa very deep and elongate; no 

columella visible. 

DISCUSSION.—I (Cairns, 1989a:77) previously cited and 

figured Alcock's "manuscript type" of Flabellum weberi as a 

specimen of/, insignis. Because F. weberi was not reported by 

Alcock (1902c) in his Siboga report, I assumed that he had 

chosen not to describe this species. But since 1989, I have 

become aware of two obscure papers published by Alcock 

(1902a,b) in July, 1902, one month before his Siboga report 

was published, in which he briefly described most of the 

Siboga new species. However, for some reason, eight species 

described in those earlier papers (Alcock, 1902a,b) were not 

redescribed in the later Siboga report (Alcock, 1902c), F. 

weberi being one of them. Thus, F. weberi is a legitimate junior 

synonym of/, insignis, not merely a manuscript type. 

Javania insignis is compared to /. borealis in the account of 

that species. 


OT6-2), 1, USNM 92744; TM (KT9015, CB1-2), 1, USNM 

92745; off Jogashima, Sagami Bay, 1, USNM 92746, 1, ORI; 

off Misaki, Sagami Bay, 1,ZMC; 32°26'N, 128°37'E,249m, 1, 

ZMC. Previous Records: Holotype of F. weberi, ZMA; 

Philippine and Japanese specimens (Caims, 1989a). 

TYPES.—The holotype of/, insignis is deposited at the BM 

(1973.2.20.1). Type Locality: 34°13TST, 136°13'E (off Owase, 

Mie-ken, Honshu), 88 m. 

The holotype of F. weberi is deposited at the ZMA (Coel. 

1232). Type Locality: Siboga-3\0: 8°3O'S, 119°07.5'E 

(Flores Sea), 73 m. 


off Owase, Mie-ken, Honshu; East China Sea off Fukue Jima; 

Eastern Channel, Korea Strait, off southwestern Honshu; 

southern Cheju Do, Korea; 46-249 m. Elsewhere: Widespread, 

including Hawaiian Islands, Philippines, Indonesia, 

Red Sea, and southwest Indian Ocean; 73-825 m. 

Javania cailleti (Duchassaing and Michelotti, 1864) 



Javania borealis sp. nov. 

Rhiwtrochus Milne Edwards and Haime, 1848a 

DIAGNOSIS.—Corallum ceratoid to turbinate or compressed. 

Transverse division absent; pedicel small and not reinforced 

with stereome. Thecal spines absent; however, 2-20 slender

NUMBER 557 81 

hollow rootlets anchor corallum base, the rootlets tending to be 

concentrically arranged in cycles. Three to six cycles of 

nonexsert septa, the lower cycle septa being highly concave 

near calicular edge. Pali absent; columella rudimentary. 

TYPE SPECIES.—Rhizotrochus typus Milne Edwards and 

Haime, 1848a, by monotypy. 

Rhizotrochus typus Milne Edwards and Haime, 1848 

PLATES 35a-c, 40/!./ 

Rhizotrochus typus Milne Edwards and Haime, 1848a:282, pi. 8: fig. 

16.—Cairns, 1989a:79-81, pi. 41: figs, f-j [synonymy]. 

Rhizotrochus niinoi Yabe and Eguchi, 1942b: 136-137, 154-155, pi. 12: fig. 

4a,b [new synonym]. 

Monomyces niinoi.—Eguchi, 1965:292, 2 figs; 1968:C48-49.—Kikuchi, 

1968:8, pi. 5: fig. 9. 

Monomyces typica.—Eguchi, 1968:C49. 

Flabellwn transversale.—Nishimura and Suzuki, 1971:11, pi. 4: fig. 1. 

Monomyces uchiuraensis Eguchi, 1972:160, pi. 1: figs. 1-7; 1973:83-84, pi. 

1: figs. 1,2. 

DISCUSSION.—Based on the 10 small specimens reported 

herein, nothing can be added to the characterization (Caims, 

1989a) of this species. R. typus is distinguished from the other 

Pacific species, R. levidensis Gardiner, 1899 (Loyalty Islands), 

by having a rather squat (trochoid) corallum; up to six cycles of 

septa; and one to two or more cycles of rootlets (6, 18, >20). 

Rhizotrochus levidensis has a much smaller, ceratoid corallum; 

an incomplete fourth cycle of septa; and 7 or less rootlets. 

At a GCD of 10-12 mm a full four cycles of septa are 

usually present; between 12-27 mm the fifth cycle is formed; 

and above 27 to 57 mm GCD the sixth cycle is formed, but 

never fully completed (Cairns, 1989a). The holotype of R. 

niinoi has a GCD of 9.75 mm and has 48 septa (the S4 being 

rudimentary), which is consistent with the ontogeny of R. 

typus. Furthermore, at this small size it is not unusual for the S1 

to be larger and thicker than the S2. It can also be seen from this 

small specimen that the pores that lead to the first ring of six 

rootlets flank the six Sv whereas the pores that correspond to 

the second, higher ring of 6-12 rootlets flank both the S1 and 

S2. 

MATERIAL EXAMINED.—New Records: Off Misaki, Sagami 

Bay, depth unknown, 9, USNM 92750, 1, ORI; Okinose, 

Sagami Bay, 110 m, 1, ZMC. Previous Records: Holotype of 

R. niinoi, TIUS; Philippine specimens (Caims, 1989a). Reference 

Specimens: 3 syntype of R. levidensis, BM 1970.1.26.9- 

10. 

TYPES.—Two syntypes of R. typus are deposited at the 

MNHNP. Type Locality: Singapore, depth unknown. 

The holotype (Plate 40h,i) of R. niinoi is deposited at the 

TIUS (60820). Type Locality: Hukui Maru-16: Wakasa Bay, 

Sea of Japan, Honshu, 76-104 m. 

The deposition of the 3 syntypes of Monomyces uchiuraensis 

is unknown. Eguchi (1973b) designated a lectotype from the 

syntype series (specimen #45), a specimen that was originally 

illustrated by Eguchi (1972d, pi. 1: figs. 3, 4). Type Locality: 

Near Awashima, Uchiura Bay, Suruga Bay, Honshu, 20 m. 

DISTRIBUTION.—Japan: Sagami, Suruga, and Wakasa Bays; 

off Amakusa Island, Kyushu; 20-104 m. Elsewhere: Philippines, 

Singapore, Pelau, Indonesia, Andaman Islands, Persian 

Gulf, Red Sea (see Caims, 1989a); 20-1048 m. 

Suborder DENDROPHYLLIINA 

Family DENDROPHYLLIIDAE 

Balanophyllia Wood, 1844 


DISCUSSION.—The taxonomy of the genus Balanophyllia is 

quite confused, there being about 53 described Recent species, 

at least 31 of which occur in the Indo-West Pacific region. This 

number will certainly be refined and perhaps reduced as larger 

collections and more types are examined. Not all of the 

specimens available for study have been reported herein. In this 

regard I agree with Yabe and Eguchi (1942b: 142), who stated: 

"Besides the above mentioned species, we have examined 

many different forms of the genus Balanophyllia, some of them 

may probably constitute new species, but mostly represented 

by [a] single, worn specimen; their description are left for 

another occasion." 

Balanophyllia cumingii Milne Edwards 

and Haime, 1848b 

PLATE 35d.e 

Balanophyllia cumingii Milne Edwards and Haime, 1848b:87-88, pi. 1: fig. 

8.—Eguchi. 1968:C51, pi. C21: figs. 7, 8. 

Balanophyllia cumingi.—Eguchi, 1934:368. 

?Balanophyllia qffinis.—Yafoe and Eguchi, 1942b: 140-141, pi. 12: figs. 11. 

12.—Eguchi, 1968:C50-51. pi. C12: figs. 1-3, 7-9, 13, 14. 

Balanophyllia cf. cumingii.—Yabe and Eguchi, 1942b: 141 [in part: pi. 12: fig. 

13a,b. and specimen from Soyo Maru-429 cataloged as TIUS 58990, not 

58236]. 

Balanophyllia diomedae [sic].—Yabe and Eguchi, 1942b: 142. 

?Balanophyllia cf. imperialis— Eguchi, 1968:C52-53, pi. C21: figs. 9, 10. 

DESCRIPTION.—Corallum ceratoid and small, the largest 

specimen examined (TM (KT9015, CB1-2)) only 10.3 x 7.8 

mm in calicular diameter and 18.9 mm in height, with a pedicel 

diameter of 4.0 mm. Corallum firmly attached: PD:GCD = 

0.40-0.50. Costae porous and coarsely granular. Often a thin 

epitheca encircles the lower synapticulotheca, providing a 

better substratum for encrusting organisms. Corallum white. 

Septa hexamerally arranged in 4-5 cycles, the fourth cycle 

complete at a GCD of about 7.5 mm, and the fifth cycle 

remaining incomplete (up to total complement of 90 septa) 

even in the largest specimens examined. S1-2 equal in size, 

about 1.3 mm exsert, and thickened at upper, outer (thecal) 

edges such that they fuse with their adjacent lower cycle septa. 

Inner edges of S^ straight, vertical, and smooth for upper 

half, but becoming dentate to laciniate adjacent to columella. 

83.4 equally exsert (about 0.8 mm), the S3 only about half the 

width of an S1 _2. In small coralla, each pair of S4 fuse before its 

adjacent S3 and extends to the columella as a single septum, its

82 

inner edge being dentate to laciniate along its full margin. In 

larger coralla (Plate 35e) containing pairs of S5, the S3 extend 

to the columella and S4 are short septa, each S4 flanked by a 

pair of S5 arranged in the Pourtales Plan, the S5 adjacent to the 

S^_2 (not the S3) being the predominant septum. In these 

coralla the paired S5 either fuse before the S4 or extend 

independently (but closely parallel) to the columella. Their 

inner edges are highly dentate. Fossa deep, containing a large, 

elliptical columella flat to slightly convex in shape, composed 

of numerous, interconnected papillose elements, sometimes 

arranged in a swirled fashion. 

DISCUSSION.—The seven lots of specimens reported herein 

are conspecific and appear to be the four species reported by 

Yabe and Eguchi (1942b) as: B. affinis Semper, 1872; B. 

diomedae Vaughan, 1907; B. imperialis Kent, 1871; and B. 

cumingii Milne Edwards and Haime, 1848b. Balanophyllia 

affinis (junior synonym of B. stimpsonii (Verrill, 1865), see 

Zibrowius, 1985) differs in having a free, unattached corallum. 

Likewise, it is not B. diomedeae, a conclusion I reached earlier 

(Cairns, 1984) after comparing Yabe and Eguchi's specimens 

to the types of B. diomedeae. B. imperialis differs in having a 

larger, more robust corallum; having a curved corallum; and 

having a smaller pedicel. The identification with B. cumingii is 

consistent with the original description of that species; 

however, for a definitive identification the type would have to 

be examined. The most distinctive characteristics of B. 

cumingii are its regular dentition of its septal edges and its 

relatively large columella. 



USNM 92885; TM (KT9015, CB1-2), 7, USNM 92886; TM 

(KT9015, CB-2-2), 1, ORI; TM (KT9015, HK2), 1, USNM 

92887; Okinose, Sagami Bay, 183 m, Mortensen's 1914 Pacific 

Expedition, 15 June 1914, 1, ZMC. Previous Records: B. 

diomedae of Yabe and Eguchi (1942b): Soyo Maru-244, 1, 

TIUS 59027; Soyo Maru-266, 1, TIUS 29026; B. cf. cumingii 

of Yabe and Eguchi (1942b), Soyo Maru-429, 3, TIUS 58990 

and 1, TIUS 58236). 

TYPES.—The holotype of B. cumingii is presumed to be 

deposited at the MNHNP. Type Locality: Philippines, depth 

unknown. 


Eastern Channel off southwestern Honshu; off southern 

Kyushu; Wakasa Bay, Sea of Japan, Honshu; 65-307 m. 

Elsewhere: Philippines. 

Balanophyllia cornu Moseley, 1881 

PLATE 35/-/ 

Balanopkyllia cornu Moseley, 1881:192-193, pi. 12: figs. 11-15. 

?Balanophyllia cf. hawaiiensis.—Eguchi, 1934:368.—Yabe and Eguchi, 

1942b: 142, pi. 12: fig. 20. 

?Balanophyllia ponderosa.—Eguchi, 1968:C54 [in part: pi. C12: figs. 15-17]. 

DESCRIPTION (based on specimen from Formosa Strait, CAS 

74991).—Corallum trochoid: 24.0 x 18.6 mm in calicular 


diameter and 34 mm in height, with a slender pedicel 4.5 mm 

in diameter (only 18% of GCD). Corallum regularly curved. 

Costae variable in width, alternating between 0.3 and 0.6 mm; 

intercostal striae narrow, about 0.05 mm wide. Costae flat and 

very finely granular. Synapticulotheca porous and white, the 

edge zone extending only 6-9 mm from calicular edge, below 

which corallum is epithecate but free from encrustation. 

Septa hexamerally arranged in 5 complete cycles. S1 slightly 

wider and have thicker inner edges than S2. S^ 1.2-1.5 mm 

exsert and have straight, vertical, smooth inner edges that 

extend to the columella, the inner edges of the 4 lateral S! 

actually constricting the columella into 3 lobes. S3 only about 

0.8 mm exsert and two-thirds width of an S^, also having 

straight and entire inner edges. S4_5 arranged in a Pourtales 

Plan, the S4 being only slightly exsert and about two-thirds 

width of an S3. S5 adjacent to S.,_2 fused to those septa at 

calicular edge. Inner edges of S5 curve toward each other, each 

pair fusing before their adjacent S4, these 2 combined S5 within 

each half-system uniting before the S3 near the columella. S5 

adjacent to S3 equal to or only slightly wider and as exsert as 

adjacent S3, but S5 adjacent to S1-2 are the predominant S5, 

being much more exsert and extending to the columella. Inner 

edges of S4_5 also smooth, without teeth or laciniation. Fossa 

of moderate depth, containing an elongate, solid (not porous) 

columella composed of tightly fused, twisted laths. 

DISCUSSION.—The South China Sea specimen described 

above differs from the syntypes in having a larger corallum (5 

mm larger in GCD) and consequently more septa (96 vs 72 for 

the largest syntype). It also has a proportionately smaller 

pedicel, the PD:GCD being only 0.18 for the South China Sea 

specimen, but 0.41-0.46 for the syntypes. 

The specimen figured as B. cf. hawaiiensis by Yabe and 

Eguchi (1942b) (TIUS 58994) could not be found in the TIUS 

collections in 1980 and is presumed to be lost. From its 

illustration it would appear to be B. cornu (see Cairns, 

1984:26). 

Balanophyllia cornu is similar to B. imperialis Kent, 1871 

(type locality: Singapore) in growth form, size, and number and 

arrangement of septa. However, B. imperialis differs in having 

laciniate inner edges on S5, a much deeper fossa, and an even 

smaller pedicel diameter (PD:GCD of holotype = 0.09). 

MATERIAL EXAMINED.—Alb-5311, 3, USNM 92879; Alb- 

5313,11, USNM 92880; TM (KT9015, BS2), 3, USNM 92881, 

2, ORI; Formosa Strait, South China Sea, 60 m, Feb. 1972, 1, 

CAS 74991. Previous Records: Syntypes of B. cornu. 

Reference Specimens: Holotype of B. imperialis, BM 

1984.4.27.3. 

TYPES.—Four syntypes (Plate 35/,/) of B. cornu are 

deposited at the BM (1880.11.25.143). Type Locality: Challenger-\92: 

5°42'S, 132°25'E (Kei I., Banda Sea), 236 m. 

DISTRIBUTION.—North Pacific: Bungo Strait; ?Sagami Bay, 

Honshu (B. ponderosa of Eguchi, 1968); ?Amakusa Islands, off 

Kyushu (B. hawaiiensis of Yabe and Eguchi, 1942b); Formosa 

Strait and north of Pratas Islands, South China Sea; 60-274 m.

NUMBER 557 83 

Elsewhere: Philippines, Banda Sea, Hawaiian Islands; 236- 

470 m. 

Balanophyllia gigas Moseley, 1881 

PLATE 35./-/ 

Balanophyllia gigas Moseley, 1881:193.—Van der Horst, 1922:58-59, pi. 8: 

fig. 22.—Eguchi, 1934:368; 1965:292, 2 figs.; 1968:C51-52, pi. C21: figs. 

1, 2; pi. C31: figs. 5,6.—Yabe and Eguchi, 1942b:l 13,139-140.—Utinomi, 

1965:256.—Eguchi and Miyawaki, 1975:59. 

?Balanophyllia ponderosa.—Yabe and Eguchi, 1942b:140, pi. 12: fig. 

9.—Eguchi, 1965:293,2 figs. 

DESCRIPTION.—The following description is based primarily 

on the specimen from Alb-5070, which is quite similar to the 

holotype. Corallum ceratoid: 29.1 x 23.6 mm in calicular 

diameter and 76 mm in height, with a slender but sturdy pedicel 

10.0 mm in diameter (PD:GCD = 0.34). Holotype 28.9 x 20.0 

mm in calicular diameter and 58.8 mm in height, with a pedicel 

diameter of 10.5 mm (PD:GCD = 0.36). Corallum irregularly 

bent, not regularly curved. Costae slightly convex and 0.6-0.7 

mm wide, separated by thin, porous intercostal striae and 

covered with fine granules and slightly larger, coarse spines, 

altogether producing a rough texture. Synapticulotheca porous 

and white, the edge zone extending only 7-16 mm below 

calicular edge, below which the corallum is epithecate, 

discolored, and covered with encrusting organisms. 

Septa hexamerally arranged in 5 cycles, the last cycle not 

complete, both the holotype and Albatross specimen having 

only 88 septa. S1-2 equal in size, up to 4 mm exsert, and have 

straight, smooth inner edges that attain the columella. S3 half as 

exsert and about three-quarters width of an S1-2 and also have 

straight, smooth inner edges. S4_5 arranged in a Pourtales Plan. 

In quarter-systems lacking S5, the S4 is independent, extending 

to the columella. If, however, the S4 is flanked by a pair of S5, 

the S4 is short, a pair of S5 fusing before it, this combined S5 

extending to the columella. Whereas inner edges of S^ are 

entire (smooth) and their faces are solid, inner edges of the S4_5 

are corsely dentate and the septa porous. Fossa of moderate 

depth, containing a discrete, slender, elongate, columella, 

composed of many slender elements loosely fused together. 

DISCUSSION.—Several authors, including Moseley (1881), 

have attributed the authorship of B. gigas to Brueggemann 

based on an apparently unpublished manuscript in which he 

described and named this species. However, the authorship 

must be attributed to Moseley (1881) who first published the 

name and commented on the species, if only as a brief 

comparison to another species. 

Balanophyllia gigas can be distinguished from other 

congeners by its large size and the coarse dentition and porosity 

of its higher cycle septa. 


USNM 92882; TM (KT7818, OT8-1), I, USNM 92883; TM 

(KT8412, 14), 1, ORI. Previous Records: Holotype of B. 

gigas, BM. 

TYPES.—The holotype (Plate 35*) of B. gigas is deposited at 

the BM (1876.10.11.23). Type Locality: "Japan," depth 

unknown. 

DISTRIBUTION.—Japan: From Boso Honto, Honshu (including 

Sagami Bay) to southwestern Kyushu, including Shikoku; 

115-245 m. Elsewhere: Kei Islands, Banda Sea; 90 m. 

Balanophyllia ponderosa Van der Horst, 1926 

PLATE 36a,b 

Balanophyllia ponderosa Van der Horst, 1926:49-50, pi. 3: figs. 6, 7.—Not 

Yabe and Eguchi, 1942b: 140, pi. 12: fig. 9 [= IB. gigas].—Not Eguchi, 

1965:293. 2 figs.; 1968:C54 [in part: pi. C17: figs. 6-11, 13, 14).— 

?Kikuchi. 1968:9, pi. 5: fig. 1 la,b.—?Eguchi and Miyawaki. 1975:54, pi. 6: 

fig. 3.—Cairns and Keller, 1993:274. 

DISCUSSION.—No additional specimens of B. ponderosa are 

reported herein and the Japanese specimens reported by Yabe 

and Eguchi (1942b) and Eguchi (1968) are unavailable for 

study, but the holotype of B. ponderosa was examined and 

compared to the published illustrations of specimens from 

Japan. The specimens figured by Eguchi (1968, pi. C17: figs. 

6-11, 13, 14) appear to be conspecific; however, the specimen 

illustrated in pi. C12, figs. 15-17 appears to be B. cornu, and 

the specimen illustrated by Yabe and Eguchi (1942b, pi. 12: fig. 

9) appears to be B. gigas. 

The holotype of B. ponderosa is 23.7 x 17.5 mm in calicular 

diameter and 30.0 mm in height, with a thick pedicel measuring 

14.2 x 13.4 mm in diameter (PD:GCD = 0.60). A thick, 

horizontally striate epitheca covers lower half of corallum, the 

upper synapticulotheca consisting of finely granular, convex 

costae 0.4-0.6 mm wide separated by very narrow striae. Septa 

hexamerally arranged in 5 cycles, only 1 pair of S5 lacking, for 

a total of 94 septa. S,_2 equal in size and have straight, vertical, 

entire inner edges. S3 about half width of an S^ and have 

similar inner edges, all 24 S1-3 reaching the columella. S4 are 

the smallest septa. Each pair of S5 fuses before an S4 and 

extends to the columella, where it fuses to the inner edges of 

adjacent S3. Inner edges of S4_5 finely dentate. All septa 

nonexsert. Fossa deep, containing a large, elliptical columella 

10.0 x 3.8 mm in size that is virtually flat on top. The columella 

consists of numerous small, flattened elements weakly swirled 

in a clockwise direction and tightly fused to one another. 

Balanophyllia ponderosa is distinguished from other Japanese 

congeners by its relatively thick pedicel and its large, low, 

spongy columella. 


Records: Holotype of B. ponderosa, BM; 1 specimen off 

South Africa (Caims and Keller, 1993). 

TYPES.—The holotype (Plate 36a,b) and two paratypes of B. 

ponderosa are deposited at the BM, the holotype cataloged as 

1939.7.20.62. Type Locality: Off Nilandu, Maldive Islands; 


DISTRIBUTION.—Japan: Sagami Bay, Honshu; 14-16 m 

(Eguchi, 1968). Elsewhere: Southwest Indian Ocean; Maldive 

Islands; 51-59 m.

84 

Balanophyllia sp. A 

PLATE 40/ 

Balanophyllia italica.—Eguchi, 1934:368. 

Balanophyllia cf. italica.—Yabe and Eguchi. 1942b: 140, pi. 12: fig. 

10a.b.—Eguchi. 1968:C53-54. pi. C12: figs. 10-12; pi. C20: figs. 3. 4. 

DISCUSSION.—Yabe and Eguchi (1942b) and Eguchi (1968) 

reported 7 specimens from off Japan for which they suggested 

a comparison to the French Miocene species B. italica. They 

did not provide a description of their species and only one 

specimen was available for examination and illustration. From 

their illustration it would appear that these specimens differ 

from their Japanese congeners by having a crown of pali (P4) 

and a more prominent upper thecal edge along the thecal faces 

rather than the edges. No specimens in the study material has 

pali or such a corallum shape. However, the relatively small 

specimen examined (Soyo Maru-641) of GCD = 10.2 mm had 

only incipient S5 development and no pali. 


Records: Soyo Maru-(A1, 1, TIUS 59163 (Yabe and Eguchi, 

1942b) (Plate 40/). Reference Specimens: Pliocene B. italica 

from Pavona, Italy, 1, USNM Ml56368. 

DISTRIBUTION.—Sagami Bay, Honshu; Sea of Japan off 

extreme northwestern Honshu; 56-100 m. 

Balanophyllia teres, sp. nov. 

PLATE 36CM 

Balanophyllia fistula Yabe and Eguchi. 1942b:141 [in part: "first form," pi. 12: 

fig. 14a,b).—Eguchi, 1968:C63. 

Dendrophyllia fistula.—Eguchi, 1965:295 [in part: middle figure]; 1968:C62- 

63 [in part: "simple form," ?pl. 12: fig. 4-6]. 

DESCRIPTION.—Corallum terete (cylindrical to slightly conical) 

and quite elongate, straight to irregularly curved. Holotype 

4.33 mm in calicular diameter and 30.7 mm long; largest 

specimen (Alb-4903) 7.0 mm in calicular diameter and 60.0 

mm long. Basal tip of colony free: either pointed and worn or 

transversely fractured. Buds absent. Corallum obviously 

epithecate, a continuous, smooth, thick epitheca extending 

virtually to calice. Underlying noncostate synapticulotheca 

visible only on lower half of corallum or on coralla dead when 

collected. Corallum white. 

Septa arranged in a weak Pourtales Plan of 3-4 cycles. A 

corallum of 3.5-4.5 mm GCD has only 3 septal cycles (24 

septa), whereas a slightly larger specimen of GCD = 5.6 mm 

has about 40 septa, and the largest specimen of GCD = 7.0 mm 

has a full fourth cycle (48 septa). S, slightly exsert (0.3-0.6 

mm) and relatively narrow, with straight, entire inner edges that 

fuse with the columella only deep within fossa. S2 of small 

coralla (e.g., holotype) small (only about one-third width of an 

St) but in a larger corallum having pairs of S4 within their 

half-systems, S2 are three-quarters width of an S, and almost 

attain the columella. S3 of smaller coralla slightly wider than 

the S2, the inner laciniate edges of each pair of S3 within a 

system fusing before its adjacent S2 near the columella. S3 of 


larger coralla quite small, each flanked by a pair of larger S4 

that loosely fuse near the columella. Fossa moderately deep, 

containing a discrete spongy columella, that, in larger 

specimens, is constricted by the inner edges of the 4 lateral Sv 

DISCUSSION.—Yabe and Eguchi (1942b) distinguished two 

similar taxa of dendrophylliids that they referred to as forms of 

Balanophyllia fistula, but later (Eguchi, 1968) as two separate 

species: DendrophyllialBalanophyllia fistula and Alcockia 

wellsi. The former species was characterized as being simple 

and epithecate, the latter as colonial and costate (nonepithecate). 

It is clear from Yabe and Eguchi's (1942b, pi. 12: fig. 

14a,b) illustration of a specimen from Soyo Maru-3\6, later 

listed as Balanophyllia fistula by Eguchi, 1968, that it is 

conspecific with B. teres. Balanophyllia teres is distinctive 

among the other Recent species in having such a slender, 

elongate, cylindrical corallum. It is not Balanophyllia fistula 

Alcock, 1902 (Plate 36/,g), which is a colonial coral herein 

assigned to the genus Eguchipammia. The costate colonial 

form referred to as Alcockia wellsi by Eguchi (1968) is 

discussed in the account of Eguchipsammia wellsi. 

ETYMOLOGY.—The species name teres (Latin teres, meaning 

"rounded," "well turned," "cylindrical" or "terete") refers to 

the slender corallum shape of this species. 

MATERIAL EXAMINED/TYPES.—Holotype: TM (KT9202, 

OS2), USNM 92888 (Plate 36c). Paratypes: Alb-4903, 3, 

USNM 92889; Alb-4904, 2, USNM 92890; TM (KT9015, 

BS2), 7, USNM 92891; TM (KT9202, OS2), 1, ORI. Previous 

Records: B. fistula of Yabe and Eguchi (1942b), Soyo 

Maru-316, 1, TIUS 58971. Type Locality: 30°59 / N, 130°31 'E 

(mouth of Kagoshima Bay, Kyushu), 237-241 m. 

DISTRIBUTION.—Kii Strait, Honshu to Fukue Jima, East 

China Sea; 154-237 m. 

Endopachys Lonsdale, 1845 

DIAGNOSIS.—Corallum solitary, cuneiform, and free, with 

alate edge crests. Septa arranged in a Pourtales Plan. Paliform 

lobes present; columella spongy. 

TYPE SPECIES.—Turbinolia maclurii Lea, 1833, by subsequent 

designation (Milne Edwards and Haime, 1850a:lii). 

Endopachys grayi Milne Edwards and Haime, 1848 

PLATES 36eji, 37/ 

Endopachys grayi Milne Edwards and Haime, 1848b:82-83, pi. 1: figs. 2, 

2a.—Yabe and Eguchi, 1942b: 139.—Cairns, 1984:27, pi. 5: fig. E 

[synonymy].—Zibrowius and Grygier, 1985:128, figs. 39-42.—Cairns and 

Keller, 1993:276 [synonymy]. 

Endopachys japonicum Yabe and Eguchi, 1932a:388, 389 [nomen nudum]; 

1932b:443 [nomen nudum]; 1932d:14-17, pi. 2: figs. 1-6; 1942b: 139.— 

Eguchi. 1934:268; 1965:293, 3 figs.—Eguchi and Miyawaki. 1975:59. 

Endopachys vaughani Durham, 1947:39-40, pi. 11: figs. 6-8, 10, 11.— 

Durham and Barnard, 1952:103, pi. 16: fig. 67a,b.—Squires, 1959:426-427. 

DESCRIPTION.—Edge angle, exclusive of lateral crests, 

50°-55°; inclination of lateral faces changes 7-8 mm above the 

base, increasing from a rather narrow 22°-28° to a more open

NUMBER 557 85 

48°-58°. Lateral edges meet at an acute angle and project 

outward up to 3 mm as lateral edge crests. Crests usually 

confined to lower half of corallum and are about 1 mm thick. 

Up to 4 buds, none over 4 mm in GCD, have been observed to 

grow from a thecal edge (Plate 37/), above the region of the 

edge crest. Largest Japanese specimen examined (TM 

(KT9015, BS1)) 15.3 x 11.0 mm in calicular diameter and 14.6 

mm in height; however, the species can grow much larger, e.g., 

up to 39 mm in GCD (Cairns and Keller, 1993). 

Septa hexamerally arranged in 5 incomplete cycles, the 

fourth cycle usually complete at a GCD of 8-10 mm and a 

specimen of 15 mm GCD having 72 septa. Larger (non 

Japanese) specimens have a full complement of 5 cycles (96 

septa). S1-2 equal in size, moderately exsert, and fairly thick 

and porous near their upper edges; their inner edges are straight 

and vertical, extending to the columella. S3 only about 

one-third width of an S1 _2. Each pair of S4 within a half-system 

unites before its adjacent S3 to form a small paliform lobe at the 

junction. In larger coralla having a fifth cycle, pairs of S5 also 

fuse before their adjacent S4, forming another crown of 

paliform lobes. Fossa deep and narrow, containing a rudimentary, 

spongy columella composed of slender, interconnected 

papillae. 

DISCUSSION.—Only one widely distributed Recent species 

of Endopachys is acknowledged, most of the nominal species 

having been synonymized by Umbgrove (1950); however, 

there are at least five species known exclusively from fossils 

(see Wells, 1975) dating from as early as the Eocene. 

Endopachys is sometimes confused with Tropidocyathus, both 

genera having the same shape, but the porous synapticulotheca 

and Pourtales Plan easily distinguish Endopachys. Phylogenetically 

Endopachys is closest to Balanophyllia, but is 

distinguished by its compressed (cuneiform) corallum. 




USNM 92864; Alb-5315, 1, USNM 92865; TM (KT9015, 

BS1), 70, USNM 92866; TM (KT9015, BS2), 28, ORI; TM 

(KT9015, CB1-2), 1, USNM 92867; TM (KT9015, HK2), I, 

USNM 92868; TM (KT9202, OS3), 1, USNM 92869; 27.3N, 

121.3E, 80-100 m, 1, CAS 74996; TM (KT9309, AM6), 1, 

ORI; TM (KT9309, AM7), 3, ORI; USGS 17441, 17456, 

17457, 17503, Pleistocene of Okinawa, 6, USNM 88661- 

88665. 

TYPES.—The holotype of E. grayi was not traced. Type 

Locality: Unknown. 

The description of E. japonicum was based on 47 specimens 

(TIUS 41929) from the Plio-Pleistocene of Makuta-maru, Boso 

Peninsula, Honshu (type locality) and 3 specimens (TIUS 

41930) from the Pliocene Byoritsu Beds of Kanpouko, Taiwan. 

The designation of a holotype from the Honshu lot was made 

by Yabe and Eguchi (1932d) in the figure captions, and thus the 

other 49 specimens should be considered as paratypes. 

The holotype and three paratypes of E. vaughani are 

deposited in the CAS Paleontology Museum (7839-7842). Two 

additional paratypes are deposited at the UCMP (30199- 

30200). Type Locality: Loc. 27584: 23°.3'N, lW^W (mouth 

of Gulf of California), 27-402 m. 

DISTRIBUTION.—Northwest Pacific: Off Boso and Kii 

Peninsulas; off Shikoku; off eastern Kyushu; Eastern Channel, 

Korea Strait, off western Honshu; off Amami Oshima, Ryukyu 

Islands; northern Formosa Strait, East China Sea; South China 

Sea northeast of Pratas Islands. Pleistocene of Ryukyu Islands 

and Honshu. Pliocene of Taiwan; 70-274 m. Elsewhere: 

Widely distributed in Indo-Pacific from off South 

Africa to the Gulf of California, including the Hawaiian 

Islands; 37-274 m. 

Eguchipsammia, gen. nov. 

Dendrophyllia (Alcockia) Eguchi, 1968:C63 [junior homonym]. 

DIAGNOSIS.—Unattached, recumbent coralla formed 

through sparse extratentacular (rarely intratentacular) budding 

from a predominant axial corallite, only rarely with third 

generation buds present on a colony. Synapticulothecate: 

costate and/or epithecate. Septa arranged in a Pourtales Plan. 

Pali absent; columella spongy. 

TYPE SPECIES.—Dendrophyllia cornucopia Pourtales, 1871, 

here designated. 

DISCUSSION.—Six species are assigned to the genus 

Eguchipsammia: E. gaditana (Duncan, 1873); E. cornucopia 

(Pourtales, 1871); E. fistula (Alcock, 1902); E. serpentina 

(Vaughan, 1907); E. oahensis (Vaughan, 1907); and E. wellsi 

(Eguchi, 1968). The genus is characterized by having a free 

(unattached), recumbent corallum with a variable number of 

asexually generated buds attached to the theca. Because the 

buds often become free of the parent corallum, these species are 

not considered as a true Dendrophyllia, but because the buds 

are temporarily part of the corallum, these taxa cannot be 

considered as Balanophyllia. As Eguchi (1968) stated in his 

discussion of the subgenus Alcockia, it "has intermediate 

characters between Balanophyllia and Dendrophyllia." Because 

the species of Eguchipsammia have a different reproductive 

mode than either Balanophyllia of Dendrophyllia and 

because they are free-living, which would also have ecological 

consequences, a new genus is proposed. 

ETYMOLOGY.—Eguchi (1968) was the first to suggest a 

supraspecific name for those species having the abovementioned 

characteristics, i.e., Alcockia, but that name was 

preoccupied by Goode and Beane (1896) for a genus of fish. 

The genus is thus renamed in honor of Motoki Eguchi 

(1905-1978). Gender: feminine. 

DISTRIBUTION.—Miocene (Caims and Wells, 1987) to 

Recent. Living species circumtropical to warm temperate in 

western Pacific; 34-960 m. 

Eguchipsammia gaditana (Duncan, 1873), comb. nov. 

Balanophyllia gaditana Duncan, 1873:333. 

PLATE 37d-fji

86 

BalanophyIlia fistula—Yabe and Eguchi, 1942b: 141 [in part: ?pl. 12: fig. 

15a,b). 

Dendropkyllia gaditana.—Gums, 1979:181-182, pi. 36: figs. 5-10; 1984:25, 

pi. 4: fig. I [synonymy].—Zibrowius, 1980:176-178, pi. 89: figs. A-N.— 

Cairns and Keller, 1993:279-280 [synonymy]. 

DESCRIPTION.—The corallum consists of a relatively slender, 

irregularly bent, cylindrical axial coral lite from which 

smaller corallites bud at irregular intervals. Largest corallum 

(ZMC, Okinose) only 53 mm long and bears 15 secondary 

corallites or broken bases of corallites from the axial corallite 

and 1 tertiary corallite that originates from a secondary. In only 

one of the 17 coralla examined was there an example of 

intratentacular budding. All coralla examined were free, with 

no evidence of previous attachment to the substrate. Axial 

corallites (and branches) 3.0-4.9 mm in diameter, secondary 

corallites slightly smaller: 2.6-4.0 mm in diameter. Synapticulotheca 

covered by a very thin, transparent epitheca, which 

often extends to calicular edge and gives the theca a coarsely 

granular porcellaneous texture. Theca porous only near calice. 

Six Cl often slightly ridged, giving branches a polygonal cross 

section. Corallum white. 

Septa arranged in a Pourtales Plan of 3 cycles (24 septa) in 

small corallites and up to 36 septa in larger corallites. S1 

significantly exsert (0.5-0.6 mm), along with their adjacent 

pairs of higher cycle septa, together forming 6 calicular apices 

around calicular margin. S1 relatively thin and have vertical, 

entire inner edges that attain the columella. In small corallites, 

S2 are quite small, each flanked by a pair of larger S3 that fuse 

before the S2 high in the fossa and extend to the columella as a 

combined septum. In larger corallites each system contains 5, 

not 3, septa, consisting of a medial S2, 1 wide S3 and 1 small 

S3, the latter flanked by 2 wide S4 that fuse and extend to the 

columella. Inner edges of S4 and those S3 that extend to the 

columella laciniate. Fossa of moderate depth, containing a 

rudimentary, spongy, nondiscrete (edges of columella merge 

with inner septal edges) columella. 

DISCUSSION.—Yabe and Eguchi were insightful when in 

1942(e) they distinguished three forms of Balanophyllia 

fistula, some of which occured at the same stations. One form 

was solitary and heavily epithecate (their pi. 12: fig. 14), which 

is described as Balanophyllia teres herein. The other forms 

were colonial and non- or little epithecate (their pi. 12: figs. 15, 

16). The colonial forms consist of two very similar species: 

Eguchipsammia gaditana and E. wellsi, E. fistula being a much 

more robust species with four full cycles of septa (Plate 36/,g). 

E. gaditana differs from E. wellsi by having a spongy (not 

honeycomb-shaped) and nondiscrete columella (the columellar 

edges are not vertical, but merge directly with the inner septal 

edges). Secondly, in E. gaditana pairs of S4 and some S3 fuse 

high in the fossa and continue to the columella as a laciniate 

septum. In E. wellsi, the S3 and S4 paired septa rarely if ever 

fuse and they have entire inner edges. Thirdly, E. gaditana has 

a very thin epitheca covering most of its synapticulotheca and 

inconspicuous costae; E. wellsi has a costate theca, lacking an 

epitheca. Finally, the coralla of E. gaditana are more delicate 

than those of E. wellsi. 


Because the differences between E. gaditana and E. wellsi 

are so subtle, all previously published records of Balanophyl- 

HalDendrophyllia fistula from Japan should be re-examined for 

a proper identification. This includes reports by: Yabe and 

Eguchi (1932a, 1932b, 1936, 1941b, 1942b), Utinomi (1956, 

1965), Kikuchi (1968), Eguchi (1968), and Eguchi and 

Miyawaki(1975). 

MATERIAL EXAMINED.—New Records: Okinose, Sagami 

Bay, 110 m, Mortensen's 1914 Pacific Expedition, 16 coralla, 

ZMC, 1, USNM 92894. Previous Records: Specimens 

reported from Atlantic (Cairns, 1979; Zibrowius, 1980), 

Hawaiian Islands (Cairns, 1984), and southwest Indian Ocean 


TYPES.—The holotype of B. gaditana is deposited at the BM 

(1883.12.10.97). Type Locality: Porcupine-29: 36°20TSf, 

6°47'W (Iberian-Morocco Gulf), 417 m. 

DISTRIBUTION.—Off Japan: Sagami Bay, Honshu; ?Bungo 

Strait, Shikoku (Soyo Maru-23\); Osumi Strait, southern 

Kyushu; 110-188 m. Elsewhere: Widespread, including 

Atlantic, southwestern Indian Ocean, Hawaiian Islands, Australia; 

73-505 m. 

Eguchipsammia wellsi (Eguchi, 1968), comb. nov. 

PLATE 37a-c,g 

Balanophyllia fistula—Eguchi, 1934:368.—Yabe and Eguchi, 1942b:141 [in 

part: pi. 12: fig. 16a,b]. 

Dendrophylliafistula.—Eguchi, 1965:295 [in part: right fig.]. 

Dendrophyllia (Alcockia) wellsi Eguchi, 1968:C63-64. 

DESCRIPTION.—Corallum shape identical to that of E. 

gaditana, consisting of an axial corallite from which slightly 

smaller secondary corallites bud at irregular intervals, although 

E. wellsi is slightly more robust that E. gaditana, their axial 

corallites being 4.5-5.1 mm in GCD and their secondaries 

2.5-4.0 mm in GCD. Largest corallum (ZMC, Okinose) 74 

mm in length and bears 8 secondary corallites. Tertiary 

corallites and examples of intratentacular budding absent. 

Synapticulotheca costate, with no evidence of an epitheca. 

Costae equal in width (about 0.3 mm) and bear very fine, sharp 

granules. Corallum white. 

As in E. gaditana, septa hexamerally arranged in 3 to 4 

cycles (up to 36 septa) in a Pourtales Plan. S1 slightly exsert 

and have entire, vertical inner edges that attain the columella. 

S2 of small corallites are small and flanked by a pair of S3, each 

of which extends almost to the columella but do not quite fuse 

with one another. In larger corallites having systems including 

5 septa, there is a large S2, 1 even wider S3 that attains the 

columella and another very small S3 that is flanked by a pair of 

S4. Inner edges of each S4 pair within in a half-system bend 

toward each other but do not quite meet. Inner edges of all septa 

entire (i.e., smooth), not dentate or laciniate. Fossa of medium 

depth, containing a discrete (a self-contained structure with 

vertical edges, not merging imperceptibly with inner septal 

edges) columella composed of small lamellae densely fused 

into a honeycomb-like structure. 

DISCUSSION.—Eguchipsammia wellsi is one of the three

NUMBER 557 87 

forms of B. fistula reported by Yabe and Eguchi (1942b) and 

one of the two forms of D. fistula reported by Eguchi (1968). 

See the previous discussion of E. gaditana for comparisons to 

that species. 


USNM 92895; AIb-4936, 2, USNM 92896; TM (KT9202, 

YS1), 1, ORI; Okinose, Sagami Bay, 110 m, Mortensen's 1914 

Pacific Expedition, 7 coralla, ZMC, 1, USNM 92897; Soyo 

Maru-2\2, 3, TIUS 58974. Reference Material: Syntypes of 

B. fistula, ZMA(Coel 564). 

TYPES.—The original description of the new subgenus and 

species Alcockia wellsi is quite confusing and rather brief, but 

qualifies as a legitimate description under Article 13C of the 

ICZN. It can be deduced that A. wellsi was reported by Eguchi 

(1968) from 13 Soyo Maru stations, all of these specimens 

herein considered to be syntypes. Two of these specimens were 

previously illustrated by Yabe and Eguchi (1942b, pi. 12: figs. 

15, 16), but because I consider one of these specimens (Soyo 

Maru-231, pi. 12, fig. 15) to be E. gaditana, I suggest that the 

other figured specimen from Soyo Maru-2\0 (pi. 12: fig. 16a,b, 

TIUS 58969) be considered as the lectotype. Type Locality: 

Soyo Maru-210: SS^X nS^S'E (off Kii Peninsula, 

Honshu), 165 m. 

DISTRIBUTION.—Sagami Bay and Kii Peninsula, Honshu; 

south of Fukue Jima; Osumi Strait, Kyushu; Tokara Retto, 


Rhizopsammia Verrill, 1870a 

DIAGNOSIS.—Like Balanophyllia, but forming reptoid colonies 

by extratentacular stoloniferous budding. Pourtales Plan 

present. Pali absent; columella rudimentary. 

TYPE SPECIES.—Rhizopsammia pulchra Verrill, 1870a, by 

monotypy. 

DISCUSSION.—Rhizopsammia represents the first level of 

polyp organization among the Recent colonial dendrophylliids 

having a Pourtales Plan, i.e., that of stoloniferous budding, 

being just a short step from a solitary Balanophyllia. The next 

higher level of organization is characterized by Cladopsammia 

in which phaceloid colonies originate from a common basal 

coenosteum. 

Rhizopsammia minuta mutsuensis Yabe and Eguchi, 1932 

PLATE AQf.g 

Rhizopsammia minuta var. mutsuensis Yabe and Eguchi. 1932f:208-209, pi. 9: 

figs. 1-3.—Abe, 1939:175-187.—Fadlallah, 1983a: 133. 

Rhizopsammia minuta mutsuensis.—Eguchi, 1934:368; 1965:293, 2 figs.; 

1968:C72, pi. C4: fig. 4 [color]; pi. C14: figs. 1-3.—Yabe and Eguchi, 

1942b: 143.—Suzuki, 1969:17-24. figs. 6-9.—Yajima. 1986:37-40.— 

Song, 1991:138-139, pi. 1: fig. 15; pi. 3: figs. 3, 6. 

DESCRIPTION OF HOLOTYPIC COLONY.—Corallum consists 

of approximately 46 interconnected encrusting corallites 

covering an area of about 5 cm 2 . Corallites bud extratentacularly 

from reptoid stolons, the stolons maintaining their 

connection throughout colony development. Corallites elliptical 

(GCD:LCD about 1.1), up to 5.8 mm in GCD, and 4 mm in 

height. All coraliites epithecate, the synapticulotheca visible 

only at calicular edge. Corallum white; coenosarc yelloworange 

(Song, 1991). 

Septa hexamerally arranged in 4 cycles in a Pourtales Plan. 

Only the largest corallites (>5.5 mm GCD) have the full 48 

septa; most corallites lack several pairs of S4. S1 only 

independent septa, and have straight, smooth inner edges that 

attain the columella. S2 also have smooth inner edges, but do 

not quite attain the columella. S3 smallest septa and have finely 

dentate inner edges extending only about half distance to 

columella. Inner edges of S4 coarsely dentate, each pair of S4 

fused before the inner edge of its common S3 and continues 

toward columella where it is loosly fused to an S2 near 

columella. Fossa of moderate depth, containing a small 

elliptical papillose to porous columella. 

DISCUSSION.—Rhizopsammia minuta mutsuensis is stated to 

differ from the nominate subspecies, R. minuta minuta van der 

Horst, 1922, by having taller corallites, a deeper fossa, and 

costate stolons. The nominate subspecies was described from 

the Lesser Sunda Islands, Banda Sea. A third subspecies, R. 

minuta bikiniensis Wells, 1954, described from the Marshall 

Islands, differs in having smaller corallites and consequently 

fewer septa. Eleven other species are attributed to this genus 



Records: Holotype of R. minuta mutsuensis, TIUS. Reference 

Specimens: Holotype of/?, minuta bikiniensis, USNM 45106. 

TYPES.—The holotype colony of R. minuta mutsuensis, 

which was designated in a plate caption, is deposited at the 

TIUS (41391). Four more paratype colonies exist. Type 

Locality: Moura-shima (40°55.5'N), near Asamushi, Mutsu 

Bay, Honshu, Japan, 1 -2 m. 

DISTRIBUTION.—Endemic to the cold temperate northwest 

Pacific: Sea of Japan coast of Honshu from Wakasa Bay to 

Mutsu Bay; Ishikari Bay, Hokkaido; Sagami and Suruga Bays, 

Honshu; Yellow Sea off South Korea; Sea of Japan coast of 

South Korea; Ullung Do, Sea of Japan; 0-2 m. 

Cladopsammia Lacaze-Duthiers, 1897 

DIAGNOSIS.—Small phaceloid colonies formed by extratentacular 

budding from a common basal coenosteum and from 

edge zone of larger corallites. Pourtales Plan well developed. 

Pali absent; columella spongy. 

TYPE SPECIES.—Cladopsammia rolandi Lacaze-Duthiers, 


DISCUSSION.—If Dendrophyllia applies to those species 

with axial, sympodial, or bushy growth forms originating from 

a single pedicel, then Cladopsammia may be reserved for those 

species having a phaceloid growth form characterized by 

numerous corallites budding from a common basal coenosteum. 

The next lower level of corallite integration is that of 

stoloniferous, reptoid budding, designated as the genus 

Rhizopsammia: the next higher level of integration is the 

branching mode, i.e., Dendrophyllia. Four species are recognized 

in Cladopsammia: C. rolandi Lacaze-Duthiers, 1897; C.

88 

gracilis (Milne Edwards and Haime, 1848b); C. eguchii (Wells, 

1982); and C. echinata Cairns, 1984. 

Cladopsammia gracilis (Milne Edwards 

and Haime, 1848), comb. nov. 

PLATE i%d,e 

Dendrophyllia gracilis Milne Edwards and Haime, 1848b: 100-101, pi. 1: fig. 

13.—Faulkner and Chesher, 1979:305-306, pi. 179-189.—Wells, 

1983:240-241, pi. 17: figs. 1-4 [synonymy].—Cairns, 1991a:23, pi. 10: 

figs. a,b. 

?Dendrophyllia coccinea.—Van der Horst, 1922:55-56, pi. 8: fig. 21. 

?Dendrophyllia willeyi.—Van der Horst, 1922:56, pi. 8: figs. 17,18. 

Dendrophyllia coccinea titigimaensis Eguchi, 1934:367 [nomen nudum]. 

Tubastraea coccinea titijimaensis Eguchi, 1968:C71-72, pi. C17: fig. 16; pi. 

C31: figs. 1-4; pi. C33: figs. 1-4. 

Dendrophyllia cf. gracilis.—Eguchi, 1968:C60, pi. C23: figs. 4-6. 

?Dendrophyllia sp.—Eguchi, 1968:C66, pi. CI4: figs. 6.7; C67, pi. C30: figs. 

6,7. 

Dendrophyllia arbuscula Utinomi, 1971:220, pi. 13: fig. 4a,b. 

DESCRIPTION.—Illustrated colony a small (5 cm broad) 

phaceloid clump of 18 corallites, some of the corallites budding 

in close proximity from a common basal coenosteum, the rest 

budding from the lower edge zone of larger corallites. Calices 

up to 12 x 11 mm in calicular diameter. Theca thin and porous; 

no epitheca present Costae fairly well defined and about 0.3 

mm wide, each bearing a unilinear row of small teeth. Corallum 

white; coenosarc orange, pink, yellow, salmon, or green. 

Septa arranged in a pronounced Pourtales Plan in 5 

incomplete cycles, pairs of S5 often missing even from largest 

corallites. S1 slightly exsert and relatively narrow, with 

straight, vertical, entire inner edges that reach the columella. S2 

about half width of an S, and also have entire inner edges, the 

S2_5 having laciniate inner edges. S3 quite narrow, but 

independent, as are the S4; however, each pair of S5 unite 

before its adjacent S4, which in turn unites with its counterpart 

before the S3 and then curves toward the S2 within its system 

and almost meets its counterpart from the adjacent half-system 

near the columella. Fossa quite deep, containing a spongy 

columella of variable size. 

DISCUSSION.—Faulkner and Chesher (1979) published 11 

color plates of this colorful species in situ, which Faulkner 

referred to as the "Festival Tube Coral" and the "Monet Tube 

Coral." In the captions to their plates they credit J. W. Wells 

with the observation that the coenosarc color is of no 

taxonomic importance for this species. 

Cladopsammia gracilis has been previously identified in the 

genus Tubastraea and more commonly in Dendrophyllia. To 

reiterate, C. gracilis differs from Tubastraea in having a 

Pourtales Plan septal arrangement and differs from Dendrophyllia 

in having a low, phaceloid colony with corallites 

originating from a common basal coenosteum. Comparisons to 

C. eguchii are made in the account of that species. 

MATERIAL EXAMINED.—New Records: Off Seto Marine 

Lab, Shirahama, Honshu, 3 m, 1 colony, USNM 92870. 

Previous Records: Specimens reported by Faulkner and 

Chesher (1979) and Cairns (1991a), all at the USNM. 


TYPES.—The types of D. gracilis are deposited at the 

MNHNP. Type Locality: Off "China"; depth unknown. 

The holotype of Tubastraea coccinea titijimaensis is 

deposited at the TIUS (#895). Type Locality: Futami port, 

Chichijima, Bonin Islands, depth unknown. 

DISTRIBUTION.—Sagami Bay, Honshu; Shirahama, Kii 

Strait; Bonin Islands; 3-95 m. Elsewhere: Common throughout 

Indo-West Pacific and eastern Australia; 0-45 m. 

Cladopsammia eguchii (Wells, 1982), comb. nov. 

PLATE 3Sa,b 

Dendrophyllia arbuscula var. compressa Eguchi and Suzuki, 1973:84, pi. 1: 

fig- 3. 

Balanophyllia eguchii Wells, 1982:211, 213, pi. 1: figs. 4-6; 1983:239-240.— 

Cairns. 1991a:23, pi. 9: figs. h-j. 

DESCRIPTION.—Illustrated colony a small (2.5 cm in width) 

phaceloid corallum composed of only 5 corallites, 3 of them 

rooted in a common basal coenosteum, the other 2 budded from 

2 of the larger corallites. Calices of small corallites circular, but 

calices of larger corallites (up to 13x9 mm in diameter) tend 

to be elongate. Synaticulotheca thick and porous, covered by 

costae 0.35 mm wide, each bearing a unilinear row of dentate 

granules. Corallum white; polyps red (Eguchi and Suzuki, 

1973). 

Septa hexamerally arranged in a weak Pourtales Plan, the 

largest corallites having a full fifth cycle (96 septa), but most 

corallites lacking many pairs of S5. S1-3 equal in size, little 

exsert, and have straight, vertical, entire inner edges that border 

the columella. All S4 quite narrow. Pairs of S5 unite before their 

mutually adjacent S4 and either extend as a single septum to the 

columella or are fused with their counterpart within the 

half-system before the S5 and continue on to the columella. 

Thus, inner edges of 24-48 septa may border the columella of 

a large corallite. Fossa of moderate depth, containing an 

elongate, spongy columella. 

DISCUSSION.—Cladopsammia eguchii differs from C. gracilis 

primarily by having a weakly developed Pourtales Plan, its 

S2 and even S3 often attaining the columella, only its S5 

consistently fused before the S4. C. eguchii also has a thicker 

synapticulotheca and septa, more elongate calices, and more 

widely spaced corallites. In the last character, C. eguchii 

approximates the reptoid growth form of Rhizopsammia, but no 

stolons were ever observed uniting adjacent corallites, only the 

continuous basal coenosteum diagnostic of the genus. 


Records: Holotype of B. eguchii, USNM; off Seto Lab, 

Shirahama, Kii Strait, 7 m, 1 colony, USNM 78646 (Wells, 

1982). 

TYPES.—The holotype of Balanophyllia eguchii is deposited 

at the USNM (46966). Type Locality: Marchena I., Galapagos; 

6 m. 

DISTRIBUTION.—Off Japan: Off Nakagi, Minami-Izu machi, 

Shizuoka Prefecture, Honshu; off Shirahama, Kii Strait; 7 m. 

Elsewhere: Hawaiian Islands; off Queensland, Australia; 

Gulf of Panama; Malpelo Island; Galapagos Islands; 1-85 m.

NUMBER 557 89 

Dendrophyllia Blainville, 1830 


DISCUSSION.—Of the 25-30 Recent species of Dendrophyllia, 

three basic growth forms can be discerned, (1) tall, 

arborescent colonies, often flabellate, having several larger 

axial corallites from which short corallites bud in an irregular 

manner, (2) small, bushy colonies that also have axial 

corallites, each bearing relatively few, large corallites, and (3) 

large, dendroid colonies with fairly regular, sympodial branching, 

the terminal corallite of each branch being replaced by each 

successive corallite. The type-species, D. ramea (Linnaeus, 

1758), and four other species (D. cribrosa Milne Edwards and 

Haime, 1851; D. minuscula Bourne, 1905; D. ijimai Yabe and 

Eguchi, 1934; and D. indica Pillai, 1967) are characteristic of 

the first form, termed the "axial" group. Species belonging to 

the second ("bushy") group include: D. cornigera (Lamarck, 

1816); D. japonica Rehberg, 1892; D. arbuscula (Van der 

Horst, 1922); D. cladonia Van der Horst, 1927; D. horsti 

(Gardiner and Waugh, 1939), and several others. The third 

"sympodial" group consists of: D. alternata Pourtales, 1880; 

D. florulenta Alcock, 1902; D. oldroydae Oldroyd, 1924; D. 

boschmai Van der Horst, 1926; D. dilatata Van der Horst, 

1927; D. californica Durham, 1947; D. alcocki (Wells, 1954); 

and D. johnsoni Cairns, 1991a. All species of Dendrophyllia 

are herein characterized by having a single basal branch from 

which all subsequent branching occurs. Two species that were 

previously listed in Dendrophyllia, both of which bud from a 

common basal coenosteum, are herein considered to belong to 

Cladopsammia: C. gracilis (Milne Edwards and Haime, 

1848b) and C. eguchii (Wells, 1982). Likewise, several other 

species are transferred from Dendrophyllia to Eguchipsammia 

based on their quasicolonial, unattached growth form: E. 

gaditana (Duncan, 1873); E. cornucopia (Pourtales, 1871); E. 

fistula (Alcock, 1902); E. serpentine (Vaughan, 1907); and E. 

oahensis (Vaughan, 1907). 

Dendrophyllia ijimai Yabe and Eguchi, 1934 

PLATE 38C./ 

Dendrophyllia ijimai Yabe and Eguchi, 1934a:2026 — Eguchi, 1965:294. 2 

figs.; 1968:C65, pi. C16: figs. 1, 2; pi. C22: fig. 1; pi. C30: figs. 4, 5 

[synonymy].—Kikuchi, 1968:9, pi. 15: fig. 2.—Eguchi and Miyawaki, 

1975:54.—Cairns and Keller, 1993:280, fig. 13c. 

Dendrophyllia micranthus.—Eguchi, 1965:294. 1 fig; 1968:C66. pi. C24: figs. 

2, 3.—?Utinomi, 1965:256-257; 1971:219-220.—Not Kikuchi. 1968:9, pi. 

5: fig. 10.—Not Eguchi and Miyawaki, 1975:54, pi. 7: fig. 1.—Not Song. 

1991:137, pi. 1: fig. 6; pi. 3: fig. 2. 

Dendrophyllia minuscula— Utinomi, 1965:257.—?Eguchi. 1968:C60-61.— 

Tribble and Randall, 1986:159. 

DESCRIPTION.— Corallum composed of elongate, relatively 

straight axial coralites, circular in cross section, and gradually 

attenuating in diameter to terminal calices 6-7 mm in diameter. 

Numerous stout corallites bud perpendicular and in all 

directions from the axial corallite, usually only 3-9 mm in 

height and 5-6 mm in diameter. Costae well defined, 0.3-0.4 

mm wide, and separated by deep, porous intercostal furrows 

about 0.15 mm wide. Costae bear 1, sometimes 2, rows of small 

pointed granules. Corallum white. 

Septa hexamerally arranged in 4 cycles (48 septa) in a 

Pourtales Plan. S, largest septa and have vertical, straight, 

entire inner edges that reach the columella. S2 only about half 

width of an St and have laciniate inner edges. S3 rudimentary 

and also have laciniate inner edges. S4 equal in width to an S3 

near calice but lower in fossa each pair of S4 are fused before its 

adjacent S3 and extended toward columella where it unites with 

the other combined S4 within the system before the common S2 

near the columella. Inner edges of S4 highly laciniate. Axial 

corallites differ from the more numerous lateral corallites in 

being slightly larger and having correspondingly more septa 

(some S5), having a deeper fossa, and lacking the final S4 

fusion before the S2- Fossa of lateral corallites of moderate 

depth and contains a small, spongy columella. 

DISCUSSION.—As stated in the genus discussion, four other 

species are recognized in the same group (axial) of Dendrophyllia 

species as D. ijimai, only one of which occurs in the 

Japanese region: D. cribrosa. Dendrophyllia ijimai is distinguished 

from that species by have nonanastomotic branches 

and exsert corallites (not flush with the coenosteum). 

The axial corallite growth form of D. ijimai is similar to that 

of Tubastraea micranthus, with which it has been confused. 

Tubastraea micranthus can be distinguished by having 

normally arranged septa; D. ijimai has a Pourtales Plan. 

MATERIAL EXAMINED.—New Records: Moroisa, Sagami 

Bay, 55 m, 2 colonies, USNM 92872,1 colony, ORI; off Japan, 

1 colony, USNM 92874; TM (KT9202, YT1), 1 colony, USNM 

92873; Misaki, Sagami Bay, 55 m, Mortensen's 1914 Pacific 

Expedition of 1914, 5 colonies, ZMC. 

TYPES.—The types of D. ijimai have not been traced. Type 

Locality: Unknown, but presumed to be off Japan. 

DISTRIBUTION.—Japan: Sagami Bay and Izu Peninsula, 

Honshu; off Kii Peninsula; Osumi Shoto, northern Ryukyu 

Islands; Amakusa Islands; north of Cheju Do, South Korea, 

East China Sea; 10-200 m. Elsewhere: Western Indian 

Ocean (Cairns and Keller, 1993), 37-366 m. 

Dendrophyllia cribrosa Milne Edwards and Haime, 1851 

PLATE 3%g,h 

Dentipora crihrosa Blainville, 1830:348 [nomen nudum]; 1834:382 [nomen 

nudum]. 

Dentipora anastomozans Blainville, 1830:348 [nomen nudum]; 1834:382 

[nomen nudum]. 

Oculina anastomozans Blainville, 1830:348 [nomen nudum]; 1834:382 

[nomen nudum]. 

Dendrophyllia sp.—Milne Edwards and Haime, 1850b: 137. 

Dendrophyllia crihrosa Milne Edwards and Haime, 1851:137; 1860: 117- 

118.—Van der Horst, 1922:52-53, pi. 7: fig. 2.—Eguchi, 1965:295, 1 fig.; 

1968:C58-59 [but not second paragraph], pi. C2: fig. 2; pi. C21: figs. 3, 

4.—Hamada: 1969:255-257. pi. 1: fig. la-e; pi. 2: figs. 2, 3.—?Song, 

1982:139. pi. 3: figs. 9. 10; 71988:29. pi. 3: figs. 1-8; 71991:137. 

?Dendrophyllia anastomozans—Monod, 1950:60; 1954:226-230. text-figs. 

6-10, pi. I: figs. 1-3.—Chevalier, 1966:1379-1382, text-fig. 30. 

REDESCRIPTION OF LARGER SYNTYPE (RMNH 9212).— 

Corallum 15 cm wide and 12 cm in height, essential planar in

90 

shape. Colony obviously dead when collected, the coenosteum 

well worn and discolored to a light brown. Branches robust, 

basal branches about 31 mm in diameter, distal blunt tipped 

branches about 13 mm in diameter, branch anastomosis 

common. Corallites 5-6 mm in diameter and flush with 

coenosteum, perhaps due to wear of corallum. Theca vermiculatc 

as well as obscurely costate, the coenosteum penetrated by 

numerous small (0.32 mm in diameter) circular pores (?borings). 

Septa hexamerally arranged in 4 cycles in a Pourtales 

Plan, some of the larger corallites having several pairs of S5. S, 

slightly wider than S2, both cycles of septa attaining the 

columella. S3 narrowest of septa, each S3 flanked by a pair of 

S4 which fuse before the S3 and extend to the columella. Fossa 

shallow, containing a large, spongy columella. 

DISCUSSION.—Dendrophyllia cribrosa is treated as a member 

of the first group of species as defined in the generic 

discussion because it has an arborescent corallum and 

nonsympodially budded corallites. It is relatively easily 

distinguished from other North Pacific species by having thick, 

anastomotic branches; very short (flush) corallites oriented 

perpendicular to parent branches; circular corallites 4.5-8.0 

mm in diameter, and a robust columella. The proper authorship 

of the species is Milne Edwards and Haime (1851), not Milne 

Edwards and Haime (1860), as suggested by some authors. 

Blainville's earlier names published in lists must be considered 

as nomina nuda and de Haan's Oculina anastomozans was 

apparently only another unpublished manuscript name cited by 

Blainvilic (1830). 


Records: Larger syntype of D. cribrosa. RMNH. 

TYPES.—Two syntypes (Plate 38tf./i) of D. cribrosa are 

deposited at the RMNH: the larger specimen described above 

(RMNH 9212. manuscript type of Dentipora anastomozans) 

and a much smaller fragment 8.2 cm long (RMNH 9209, 

manuscript type of Dentipora cribrosa). Type Locality: 

Unknown. 

DISTRIBUTION.—Japanese region: Sagami Bay. Honshu; 

?Sea of Japan off eastern and southern South Korea (Song, 

1982); Pleistocene of Kanagawa-ken; 7-40 m. Elsewhere: 

Off Banka. southeastern Sumatra; ?off Angola (Cabinda) 

and the Congo; 20-30 m. 

Dtndrophyllia japonic a Rehberg, 1892 

DendropMvllui faponuu Rehberg. IR92.28-29. pi. 4: fig. 4.—Van der Horst. 

1926 44-45. p| V figs. 4. V-Yabe and Eguchi. 1932a: 388; 1941b: 102 — 

Eguchi. 1934:367; I965295. 2 figs.. I968C6I-62. p». C13: figs. 3-5 

[synonymy].—Squires and Key**. 1967:28. pi 6: Tigs. 6-8 (synonymy). 

Not DrmJrvpMvllui japnmua Van der Honl. I922:.M. pi. 7: fig. 6 [junior 

homonym: replacement name D hnuhmai Van der Horst. 1926). 

DIAGNOSIS.—Colonies consist of a primary axial corallite up 

to 9 cm in height and up to 27 mm in GCD from which several 

slightly smaller, but still quite robust, corallites bud at irregular 

intervals. All colonies examined as well as those reported in the 

literature arc broken basally. but it would appear from the larger 

diameter of the basal fractures that the normal condition of the 


species is attached, not free as in Eguchipsammia. Costae are 

flat and porous; epitheca absent. Septa hexamerally arranged in 

a Pourtales Plan, larger corallites having a full fifth cycle of 

septa (96), smaller corallites usually having numerous pairs of 

S5. Su2 equal in size, slightly exsert, and quite wide, their 

smooth inner edges bordering the columella. S3 and S4 

progressively smaller. Pairs of S5 fuse before their adjacent S4 

and again before the S3 in their half-system before reaching the 

columella. Fossa deep, containing an elongate narrow columella. 

DISCUSSION.—Dendrophyllia japonica belongs to the "second 

group" of Dendrophyllia species, those characterized as 

bushy, having small colonies with sparsely budded large 

corallites. It is compared to D. arbuscula in the account of that 

species. 


Records: Specimens reported by Squires and Keyes (1967) 

from off New Zealand, NZOI. 

TYPES.—Two syntypes of Rehberg's D. japonica were 

present at the ZMB in the 1920's (Van der Horst, 1926), but it 

is likely that they were subsequently lost or destroyed. Type 

Locality: "Japan," depth unknown. 

DISTRIBUTION.—Japan: Sea of Japan off northwestern 

Honshu from Tsugaru Strait to Wakasa Bay; Sagami Bay; 

115-250 m. Elsewhere: Banda Sea; North Island, New 

Zealand; 114-549 m. 

Dendrophyllia arbuscula Van der Horst, 1922 

PLATE 38/-/ 

? Dendrophyllia conferta — Rehberg, 1892:28. 

Dendrophyllia arbuscula Van der Horst. 1922:53. pi. 8: fig. 6.—Yabe and 

Eguchi. I932a:388; 1941b: 102.—Eguchi, 1934:367; 1965:294. 2 figs.; 

1968:C55-56. pi. C21: figs. 5, 13.—Not Kikuchi, 1968:9, pi. 5: fig. 6 [= 

Cladopsammia sp.].—Not Utinomi, 1971:220 [=Cladopsammia gracilis].— 

Eguchi and Miyawaki. 1975:54, pi. 6: fig. 4.—? Song, 1991:136, pi. 1: fig. 

4; pi. 3: fig. 1. 

^Dendrophyllia mbtornigera Eguchi. 1934:367 [nomen nudum]; 1968:C64, 

pi. C32: figs. 3. 4. 

^Dendrophyllia xufxornigera cylindrica Eguchi. 1934:367 [nomen nudum]; 

l968:C64-65. pi. C32: figs. 1.2. 

^Dendrophyllia sp—Eguchi. I968:C67, pi. C30: figs. 6. 7. 

Tuhastraea cYK

NUMBER 557 91 

(0.8-0.9 mm), with smooth, vertical inner edges that reach the 

columella only low in fossa. S2 less exsert and slightly less 

wide than an S,, but also have smooth inner edges that almost 

attain the columella. S3 about half width of an S2. Each pair of 

S4 within a half-system bend towards each other, resulting in 

either a loose fusion of their inner edges before their enclosed 

S3 or a closely parallel arrangement leading to the columella. 

Inner edges of S4 slightly porous and laciniate to dentate, the 

S^ being solid septa with entire inner edges. Fossa of 

moderate to shallow depth, containing a well-developed, 

discrete, elongate columella, sometimes constricted into 3 

lobes by inner edges of the 4 lateral Sv Columella composed of 

small, flattened, swirled elements that are tightly fused 

together. 

DISCUSSION.—Dendrophyllia arbuscula belongs to the 

"second group" of Dendrophyllia species described in the 

generic discussion: species having small, bushy colonies, with 

relatively few large corallites. The only other species in this 

group from the Japanese region is D. japonica Rehberg, 1892, 

which differs in having larger corallites, more septa, and a less 

developed columella. 

Dendrophyllia subcornigera Eguchi, 1968 is only tentatively 

synonymized with D. arbuscula, as the type and only 

reported specimen of this species is not available for study. Its 

description and illustrations are consistent with a large, 

well-developed colony of D. arbuscula, much like that of the 

holotype; in his original description, Eguchi (1968) noted the 

resemblance. Likewise, D. subcornigera cylindrica Eguchi, 

1968, which is also known from only one specimen, is also 

tentatively synonymized but this specimen is even more robust 

with long internodes between budded corallites and quite long 

secondary corallites. Its growth form resembles that of the "first 

group" of species of Dendrophyllia. 


USNM 22057, 2, ORI; Alb-3716, 4, USNM 22054; Alb-3718, 

1. USNM 22053; Alb-3720, 2, USNM 22052; Alb-3730, I, 

USNM 22056; Alb-3746, 2, USNM 22055; Alb-4935. 1, 


USNM 92855; TM (KT7811, OT6-2), 1, USNM 92856; 

Okinosc, Sagami Bay, 183 m, Mortensen's Pacific Expedition, 

15 June 1914, 3, ZMC. 

TYPES.—Three syntypes of D. arbuscula are deposited at the 

ZMA (Coel. 1254, 5477). Type Localities: Siboga-260. 277: 

Banda Sea, 45-90 m. 

The holotype of D. subcornigera is deposited at the 


(specimen number 896, reg. no. 40859). Type Locality: 

Enoura Bay, Shizuoka ken. Honshu, depth unknown. 

The holotype of D. subcornigera cylindrica is deposited at 

the Biological Laboratory of the Imperial Household, Tokyo 

(specimen number 897). Type Locality: Off Seto, Kanayamamaru, 

Nishimuragun, Wakayama-ken; depth unknown. 


Kii Strait; off Amakusa Islands; off Cheju Do, East China Sea; 

40-240 m. Elsewhere: Banda Sea; 45-90 m. 

Dendrophyllia boschmai Van der Horst, 1926 

Dfndrophytlia japonica Van der Horst, 1922:51, pi. 7: fig. 6 [junior primary 

homonym of I) japonica Rehberg. 1892]. 

Dendrophyllia htmhmai Van der Horst. 1926:44 [replacement name for D. 

japonica Van der Horst. 1922].—Eguchi, 1934:367—Yahe and Eguchi. 

1936:167; I968:C56-S7, pi. C15: figs. 4. 7; pi. CI6: figs. 3. 4; pi. C17: figs. 

12, 15; pi. C30: fig. 1—Song, 1982:138, pi. 4: figs. 3, 4; 1991:136. 

Dendrophyllia cyathoheloides Eguchi, 1934:367 [nomen nudum) 

Dtndrophyllia boschmai cyathoneloides Eguchi. 1965:294, I fig. 

Dfndrophytlia boschmai cyathohelioides [sic].—Eguchi. 1968:C57. pi. C2: fig. 

I; pi. CI5: figs. 1-3; pi. C15: figs. 1-3, 5. 6. 8. 9. 

Dendrophyllia boschmai cyathoheluxies [sic].—Song, 1991:136, pi. 1: figs. 

7.8. 

DIAGNOSIS.—Colonies robust and uniplanar, up to 9 cm in 

height and width. Corallites primarily sympodially budded 

from branch edges but in a very crowded manner, with 

additional corallites on corallum faces. Cocnosteum costate and 

nonepithecate. Corallites large, up to 9 x II mm in diameter. 

Septa hexamerally arranged in 4 cycles in a typical Pourtales 

Plan, larger corallites having some pairs of S5. Inner edges of 

S1 smooth, those of S2_5 dentate. Columella well developed 

and spongy. Corallum white; polyps red with yellow tentacles. 

DISCUSSION.—No specimens of this species were examined 

and thus the diagnosis above is taken from the literature. D. 

boschmai differs from the other sympodially branched species 

known from Japan, D florulenta, in having larger corallites, a 

more robust corallum, and a much denser budding arrangement. 

MATERIAL EXAMINED.—None. 

TYPES.—The holotype of D. boschmai (= D. japonica of 

Van der Horst, 1922) is deposited at the ZMA (Coel. 5451) 

(Van Soest, 1979). Type Locality: "Japan," depth unknown. 

The holotype of D. boschmai cyatheloides is deposited at the 

TIUS (57443). Type Locality: Off Kowa, Mic-ken, Honshu, 


DISTRIBUTION.—Known only from the Japanese region: 

Sagami and Suruga Bays, Honshu; off Kii Peninsula, Honshu; 

off Shikoku; south of Cheju Do, South Korea; 40-165 m. 

Dendrophyllia florulenta Akock, 1902 

PLATE 39

92 

6.0-6.5 x 4.0-4.5 mm in diameter, Wells' specimen having 

larger corallites up to 9 x 7 mm in diameter. Costae evident, 

separated by narrow intercostal striae, and covered with fine 

granules; epitheca absent. Corallum white. Septa hexamerally 

arranged in 4 complete cycles in a Pourtales Plan. Su2 highly 

exsert and have smooth (entire) inner sdges that attain the 

columella. S3 small; pairs of S4 fuse before their adjacent S3 

and extend to columella. Inner edges of S3-4 dentate. Fossa 

deep, containing a small, spongy columella. 

DISCUSSION.—The specimen reported by Wells (1954) from 

the Marshall Islands differs from typical D. florulenta in 

several characters and thus its identification is queried. It has 

larger corallites; more septa (occasional pairs of S5 in large 

corallites); and a smaller, almost lamellar, columella. Comparisons 

to the only other sympodially branched Dendrophyllia 

from the Japanese region, D. boschmai, are made in the account 

of that species. 

Van der Horst (1922) described D. florulenta as a new 

species apparently unaware that Alcock (1902a) had previously 

described the same species under the same name, making Van 

der Horst's species both a junior homonym and junior 

synonym. Most of Alcock's (1902a,b) brief descriptions of his 

new species from the Siboga collection were republished one 

month later (Alcock, 1902c), but eight of them were not, 

including Dendrophyllia florulenta. I suspect that Alcock's 

type specimens were labelled as D. florulenta in the ZMA 

collections but not recognized as types, such that when Van der 

Horst (1922) later revised the Siboga Dendrophylliidae, he 

used what he thought were Alcock's manuscript types as his 

type specimens and adopted the name as well. 

MATERIAL EXAMINED.—-New Records: TM (KT9015, 

CB1-1), 1, USNM 92875; TM (KT9015, CB1-2), 2, USNM 

92876; TM (KT9015, CB2-2), 2, USNM 92877; 34°20'N, 

\3Q O \QTE (off Okino Shima), 110 m, 18 May 1914, Mortensen's 

1914 Pacific Expedition, 1, ZMC. Previous Records: 

Specimen from Marshall Islands (Wells, 1954), 

USNM 45102. 

TYPES.—The holotype of Alcock's D. florulenta is presumed 

to be deposited at the ZMA, probably one of the two 

syntypes of Van der Horst's D. florulenta, but Alcock (1902a) 

did not document the Siboga station from which his specimens 

came. Type Locality: Unknown, but undoubtedly from the 

Siboga Expedition to Indonesia, probably Siboga-51 or 305. 

Two syntypes of Van der Horst's (1922) D. florulenta are 

deposited at the ZMA (Coel. 1195, 1256). Type Localities: 

Siboga-51, 305: Banda Sea, 69-113 m. 

DISTRIBUTION.—Off Japan: Sagami Bay; Eastern Channel, 

off Mishima and Okinoshima; north and south of Cheju Do, 

South Korea, East China Sea; 70-110 m. Elsewhere: Banda 

Sea, Bonin Islands, ?Marshall Islands; 69-243 m. 

EnaUopsammia Michelotti, 1871 

DIAGNOSIS.—Colonial, arborescent colonies formed by 

extratentacular budding. Corallites often, but not always, 


unifacially arranged. Coenosteum dense, synapticulotheca 

porous only near calice and on distal branches. Septa arranged 

normally. Columella small. 

TYPE SPECIES.—Coenopsammia scillae Seguenza, 1864, by 

monotypy. 

EnaUopsammia rostrata (Pourtales, 1878) 

PLATE 39d-f 

Amphihelia rostrata Pourtalfcs, 1878:204, pi. 1: figs. 4, 5. 

Dendrophyllia amphelioides Alcock, 1902a:l 12-113. 

Amphehelia adminicularis Rehberg, 1892:10, pi. 4: fig. 1. 

Dendrophyllia amphelioides var. cucullata Vaughan, 1907:157, pi. 47: fig. 3; 

pi. 48: figs. 1-4. 

Anisopsammia amphelioides.—Eguchi, 1934:368. 

EnaUopsammia rostrata.—Zibrowius, 1973:44-45, figs. 14, 15.—Cairns, 

1979:186-188, pi. 37: figs. 2, 3, 6; 1982:57-58, pi. 18: figs. 1-4 

[synonymy).—Cairns and Parker, 1992:52-53, fig. 18e-i. 

EnaUopsammia (Anisopsammia) amphelioides.—Eguchi, 1965:296, 1 fig. 

EnaUopsammia amphelioides disticha Eguchi, 1968:C68, pi. C24: figs. 1,6; pi. 

C29: figs. 1, 2.—Zibrowius and Grygier, 1985:134. 

DESCRIPTION.—Colonies usually planar, having close, dichotomous 

branching, which occasionally leads to branch 

anastomosis. Corallites occur on only 1 side of colony and are 

circular to slightly elliptical in shape: up to 5.6 mm in GCD, but 

usually only 4.0-4.5 mm in diameter. Costae better developed 

on acalicular face, being about 0.5 mm wide, slightly convex, 

and finely granular, separated by narrow, porous intercostal 

regions. No septocostal rostra present on Japanese specimens. 


Septa hexamerally arranged in 3 complete cycles (24 septa). 

Sj.g equal in size, nonexsert, and quite slender, having vertical, 

smooth inner edges. S3 only only half width of an S^_2 and thus 

quite slender, usually having a dentate or irregularly shaped 

inner edge. Inner edges of pairs of S3 fuse to their adjacent S2 

near columella. Fossa quite deep, sometimes slightly curved, 

and quite open because of the very narrow septa. Columella 

rudimentary, consisting of several small extensions from 

lowermost inner edges of Su2. 

DISCUSSION.—EnaUopsammia rostrata was first reported 

from Japan by Rehberg (1892) based on one specimen, and 

later by Eguchi (1965,1968) as E. amphelioides disticha, again 

apparently based on one additional specimen. Eguchi's (1968) 

reference to a locality record from Enoshima, Sagami Bay is 

not documented in his papers. The two lots reported herein are 

from the East China Sea southwest of Kyushu, not far from the 

type locality off. a. disticha. 

All Japanese specimens lack the septocostal rostrum that 

characterizes the typical form of this species. But, as earlier 

stated (Cairns, 1982:57), the development of a rostrum seems 

to be a response to an environmental factor, not a species level 

character. The Japanese specimens thus belong to the "amphelioides" 

form of this species, i.e., those specimens lacking a 

rostrum. 

MATERIAL EXAMINED.—New Records: Alb-4891,2 colonies, 

USNM 92849, 1 branch, ORI; Alb-4892, 2 colonies,

NUMBER 557 93 

USNM 92850, 2 branches, CAS 80924. Previous Records: 

Syntypes of A. rostrata and D. a. cucullata, MCZ and 

USNM, respectively. 

TYPES.—The syntypes of A. rostrata are deposited at the 

MCZ (unnumbered). Type Locality: Blake-2: 23°14'N, 

82°25'W (Straits of Florida), 1472 m. 

The syntypes of D. amphelioides are deposited at the ZMA 

and the Indian Museum, Calcutta (Van Soest, 1979). Type 

Localities: Siboga-\56, 177: Pulau Waigeo and Palau Misool, 

Indonesia; 469-1633 m. 

Type material of A. adminicularis is deposited at the ZMB 

(2670, 2691) (H. Zibrowius, pers. comm.) Type Locality: 

"Japan," depth unknown. 

The syntypes of D. amphelioides var. cucullata are deposited 

at the USNM (Cairns, 1991a). Type Locality: Off Hawaiian 

Islands; 426-679 m. 

The holotype of E. amphelioides disticha is presumed to be 

deposited at the Biological Laboratory of the Imperial 

Household, Chiyoda-ku, Tokyo (specimen # 857). Type 

Locality: Off Satsuma Peninusla, Kagoshima-ken, southwestern 

Kyushu, 270 m. 

DISTRIBUTION.—Japan: East China Sea off southwestern 

Kyushu; ?Sagami Bay; 270-331 m. Elsewhere: Cosmopolitan 

except for eastern Pacific; 229-2165 m. 

Tubastraea Lesson, 1829 

DIAGNOSIS.—Colonies dendroid, bushy, or plocoid, all 

achieved by extratentacular budding. Costate, no epitheca. 

Septa arranged normally. Pali absent; columella usually small 

and spongy. 

TYPE SPECIES.—Tubastraea coccinea Lesson, 1829, by 

monotypy. 

Tubastraea coccinea Lesson, 1829 

PLATE 39g-i 

Tubastraea coccinea Lesson, 1829:93.—Wells, 1983:243-244, pi. 18: figs. 1, 

2 [synonymy].—Prahl, 1987:230-231, fig. 8.—Wilson, 1990:137-138, fig. 

1.—Caims, 1991a:26-27, pi. 12: figs, c-c [synonymy].—Cairns and Keller, 

1993:282-284 [synonymy]. 

Lobopsammia aurea Quoy and Gaimard, 1833:195. 

Astropsammia peterseni Verrill, 1869:392. 

Dendrophyllia aurea.—Eguchi, 1934:367. 

Tubastraea tenuilamellosa.—Durham, 1947:38-39, pi. 11: figs. 1, 2, 4. 9; pi. 

12: figs. 6, 7.—Durham and Barnard. 1952:105-106, pi. 12: fig. 50d. 

Tubastraea aurea.—Squires, 1959:427-428.—Utinomi, 1965:257-258; 

1971:220-221.—Eguchi, 1965:295, 1 fig.; 1968:C68-70. pi. C16: figs. 5, 6; 

pi. 17: fig. 17; pi. C26: figs. 2, 3 [synonymy].—Kikuchi, 1968:9.—Eguchi 

and Miyawaki. 1975:54, pi. 7: fig. 3.—?Song, 1982:139-140, pi. 3: figs. 11, 

12; 1991:137-138.—Tribble and Randall, 1986:159. 

Dendrophyllia coccinea.—Eguchi, 1965:296, 1 fig.—Utinomi, 1965:257. 

? Dendrophyllia sibogae.—Kikuchi. 1968:9. 

ITubastraea coccinea.—Eguchi, 1968:C70-71, pi. C2: fig. 3; pi. C14: figs. 4, 

5, 8, 9.—Kikuchi, 1968:9, pi. 5: fig. 4.—Utinomi, 1971:221. pi. 13: fig. 

5a,b— Eguchi and Miyawaki, 1975:54, pi. 7: fig. 2. 

Not Tubastraea coccinea.—Song, 1982:140, pi. 4: figs. 7, 8 [= Dendrophyllia): 

1988:29-30:1991:138. 

DESCRIPTION.—Mature colonies roughly spherical, the 

corallites closely spaced in a plocoid arrangement. Corallites 

bud extratentacularly at colony edge and between older 

corallites, thus maintaining a plocoid structure as colony 

increases in size. Calicular edges often directly adjacent to one 

another, but occasionally corallites project 1-10 mm above 

basal coenosteum. Corallites circular to slightly elliptical in 

shape, the largest examined being about 13 mm in GCD, but 

most are only 7-9 mm in GCD. Costae equal in width 

(0.30-0.35 mm) and coarsely granular, separated by wide 

(0.15-0.20 mm), very deep intercostal furrows that are highly 

porous. Corallum white; coenosarc orange. 

Septa hexamerally arranged in 4 cycles. S^_2 virtually equal 

in size, but S1 slightly thicker and wider, penetrating farther 

into the columella than S2. Upper edges of S^ nonexsert and 

slightly tapered, reaching their greatest width 3-4 mm into 

fossa, where they have vertical, smooth inner edges. S3 much 

smaller than S^, only 0.2-0.3 mm in width, or about 

15%-20% width of an SU2. Inner edges of S3 dentate to highly 

laciniate from top to bottom. S4 rudimentary or of same size as 

the S3, 1 or both lacking from each half-system. Inner edges of 

S4 also laciniate, occasionally loosely fused to adjacent S3. 

Fossa moderate to deep, containing a columella of variable 

size, but usually containing a rather large, spongy columella. 

DISCUSSION.—Tubastraea coccinea is a very common 

shallow-water azooxanthellate with a circumtropical distribution, 

and, probably because of its accessibility, has been 

described under many (approximately 18) names, more 

complete synonymies given by Eguchi (1968) and Wells 

(1983). Its broad distribution may be the result of introductions 

caused by transport on ship hulls. Its apparent spread 

throughout the Caribbean, first noticed in the Netherland 

Antilles and progressively in other parts of the Caribbean, 

would indicate that the western Atlantic was not its original 

range. Coralla of specimens from the Caribbean, Galapagos, 

Japan, and the southwest Indian Ocean are indistinguishable. 

MATERIAL EXAMINED.—New Records: Off Partida Island, 

Espiritu Santo Islands, Gulf of California, 3 colonies, USNM 

91428; off Conception Point, Gulf of California, 4 colonies, 

USNM 91429; off Seto Marine Lab, Shirahama, 2-8 m, 2 

colonies, USNM 83644, 83657; off Tanabe, Kii Strait, 1 

colony, USNM 92851; Misaki, Sagami Bay, 1, ORI; Misaki, 

Sagami Bay, 15 June 1914, Mortensen's 1914 Pacific 

Expedition, 4 colonies, ZMC; TM (KT9202, YT1), 1 colony, 

USNM 92852. Previous Records: Holotype of A. peterseni, 

USNM; specimens reported by Caims (1991a) and Cairns and 

Keller (1993). 

TYPES.—The holotype of T. coccinea is deposited at the 

MNHNP (Wells, pers. comm.). Type Locality: Bora Bora, 

Society Islands, depth unknown. 

The types of L. aurea were not traced. Type Localities: Port 

du Roi George and Port Jackson, Australia, depth unknown. 

The holotype of A. peterseni is deposited at the USNM

94 

(38354). Type Locality: La Paz, Gulf of California, depth 

unknown. 

DISTRIBUTION.—As discussed above, T. coccinea is a 

circumtropical shallow-water species. In the northeastern 

Pacific its northen limit is at the border of the tropical/warm 

temperate region, being found in the Gulf of California, but 

only to Cabo San Lucas on the Pacific side of Baja California 

Sur (Wilson, 1990). In the northwest Pacific, however, records 

extend into the warm temperate region as far north as Sagami 

Bay. It also occurs in the Kii and Bungo Straits; Osumi Shoto, 

northern Ryukyu Islands; off the Amakusa Islands; and East 

China Sea off Cheju Do, South Korea; 0-15 m. Elsewhere: 

0-110 m. 

Schizopsammia, gen. nov. 

DIAGNOSIS.—Dendroid colonies formed exclusively by 

equal, intratentacular budding. Pourtales Plan poorly developed. 

Pali absent; columella spongy. Tabular endothecal 

dissepiments present 

TYPE SPECIES.—Schizopsammia songae, herein designated. 

DISCUSSION.—Only three dendrophylliid genera form colonies 

by intratentacular division: Lobopsammia Milne Edwards 

and Haime, 1848b; Stichopsammia Felix, 1885; and Reussopsammia 

Wells, 1937, all three of which are known only from 

the Eocene to Oligocene of Europe. One Recent species of 

Eguchipsammia, E. gaditana, is also known to bud intratentacularly, 

but only as a rare secondary mode, its primary mode 

being extratentacular. Schizopsammia differs from Stichopsammia 

in having monostomadeal, not polystomadeal, budding; it 

differs from Reussopsammia in having a weak Pourtales Plan, 

not a normal septal insertion. Schizopsammia, however, is quite 

similar to Lobopsammia in coral I urn shape, size, and septal 

arrangement, but differs in having tabular endothecal dissepiments. 

The colony shape of Schizopsammia that results from equal 

intratentacular budding and the presence of tabular endothecal 

dissepiments resembles that of Solenosmilia, a deep-water 

azooxanthellate genus in the suborder Caryophylliina. 

ETYMOLOGY.—The genus name Schizopsammia (Greek 

schizo, meaning "to split" + Greek psammos, meaning "sand," 

a common suffix used in dendrophylliid generic names) refers 

to the equal intratentacular division found within the genus. 

Gender feminine. 

DISTRIBUTION.—Known only from the Western Channel of 

the Korea Strait off South Korea; depth unknown. 

Schizopsammia songae, sp. nov. 

PLATE A0a-e 

DESCRIPTION.—Corallum dendroid, dichotomously branching 

in three dimensions with no branch anastomosis. Holotypic 


colony 46 mm in height and 43 mm broad, with a broken basal 

pedicel 6-7 mm in diameter that supports a colony of 16 

corallites. Branches circular in cross section and fairly uniform 

in diameter (cylindrical), both corallites and branch diameters 

ranging from 5-7 mm. Calices in process of intratentacular 

division quite common. Intratentacular budding initiated by the 

enlargement of 2 opposing lateral S,, which eventually fuse 

across center of calice and form a thin wall between the two 

daughter corallites. Eventually small septa begin to form 

perpendicular to the faces of the bridging Sv Synapticulotheca 

thick (1.2-1.4 mm), and porous only within 4-8 mm of the 

calice; otherwise, theca solid. Costae poorly delineated, each 

costa bearing numerous small, low (40-50 |im) granules 

arranged 4 or 5 across a costa. Intercostal regions porous, the 

pores usually circular and quite deep. Corallum white. 

Septa hexamerally arranged in essentially 4 cycles; however, 

just before intratentacular division a corallite is often elongate 

and may have more than 48 septa, whereas just after division 

corallites usually have fewer than 48 septa. S, nonexsert and 

rather narrow, with straight, vertical, smooth inner edges that 

extend to the columella. S2 about three-quarters width of an S1 

and also have smooth inner edges, but do not extend to the 

columella. S3 rudimentary, each flanked by a pair of S4 not 

much wider than an S3 in upper fossa, but which fuse before the 

S3 lower in fossa and have extremely laciniate inner edges. 

Long (up to 1.5 mm), narrow extensions of lower, inner edges 

of S4 intermingle with columellar elements. Fossa deep, 

containing a small, loose, trabecular columella. Tabular 

endothecal dissepiments present every 1.5-4.0 mm within 

branches, producing a relatively low density corallum. 

DISCUSSION.—See "Generic Discussion." 

ETYMOLOGY.—This species is named in honor of Jun-Im 

Song, the first to review the South Korean coral fauna. 

MATERIAL EXAMINED/TYPES.—Holotype: 1 colony, 

USNM 15847 (Plate 40a-e). Paratypes: 10 colony fragments 

from type locality, USNM 92910, 1 branch, ORI. Type 

Locality: Western Channel of Korea Strait, off Pusan, South 

Korea; depth unknown. Collected by P.L. Joey and received by 

USNM in January 1887. Reference Specimen: Lobopsammia 

cariosa (Goldfuss, 1827) from Eocene of France, USNM 

64630. 


NOTE.—When this paper was in galley stage (May, 1994), I 

received a reprint from J.-I. Song (Song, 1994) published in 

April, 1994, in which she described a new genus and species of 

dendrophylliid from the Korean Strait: Dichopsammia granulosa. 

Her species is the same as my Schizopsammia songae, 

described herein, and therefore must be considered as the senior 

synonym. Her type series of three specimens also came from 

off Pusan, Korea, at a depth of 20-30 m. She considered this 

species to be hermatypic (zooxanthellate).

Station 

2816 

2832 

2862 

2864 

2865 

2876 

2879 

2886 

2888 

2893 

2894 

2895 

2907 

2908 

2913 

2922 

2932 

2935 

2936 

2939 

2940 

2942 

2943 

2944 

2945 

2946 

2948 

2958 

2961 

2962 

2963 

2965 

2968 

2969 

2974 

2975 

2976 

2977 

2978 

2984 

2987 

2999 

3085 

3087 

3088 

3102 

3116 

3124 

3158 

3159 

3160 

3168 

3170 

3174 

Latitude 

l°17'00*S 

24°38'00"N 

50°49'00"N 

48°22'00"N 

48°12'00"N 

48°33WN 

48°53'00*N 

43°59'00"N 

43°58'OO"N 

34°12'30"N 

34°07'00"N 

34°07'00"N 

34°24'30"N 

34°25'25"N 

32°25'30"N 

32°27'15"N 

32°26'15*TM 

32°44'30"N 

32°49'00"N 

33°36'00"N 

33°36'OO"N 

33 O 38'45"N 

34°00'30*N 

34°00 / 00"N 

34°00WN 

33°58'O0"N 

33°55'30"N 

34°04'00"N 

34°22'45*N 

34°23'30"N 

34°23'10"N 

34°2r20*N 

34°2r40"N 

34°20 / 40"N 

34°l9'30"N 

34°01'30"N 

34 o 00WN 

33°59'30"N 

33°59'45'N 

28°57'15"N 

28°54'15"N 

24°54'30"N 

44°29'30"N 

44°28'00*>J 

44°28'00">J 

37 o 40'40 / 'N 

37°05'3(TN 

36°55'10*N 

37°47'30T^ 

37°47'20*N 

37°48'35"N 

38°O1'25'N 

38°17'00"N 

38°15'30">J 

Appendix 

Station List 

USFWS Albatross (Alb) 

Longitude 

90°31 / 30"W 

112°17'30"W 

127°36'30'W 

122°5r00"W 

122°49'00'^V 

124°53'00"W 

125°53'00*W 

124°56'30*W 

124°57'30"W 

120°32'30*W 

12O°33'3O*W 

12O°33'3O'*W 

12O°2O'OO*W 

120°20'00"W 

119°03'30"W 

119 o 05'15"^V 

117°16'15"W 

117°23'OO'W 

117°27'30"W 

118°O9'3O'W 

ns^roc^v 

118°13'45"W 

119°28'30*W 

119°28'30"W 

119°29'30'^V 

119°30'45'^V 

119 o 41'30*W 

120°19'30"W 

119°4O'3O'W 

119°39'30"W 

119°39'40"W 

119°38'30*>V 

U9°38'20"W 

119°37'45*W 

119°44'45'W 

119°29'00'W 

119°26'30^V 

119°25'30*W 

119°22'15'^V 

118°15'45"W 

118°18WW 

110°39'00'^V 

124°17'00~W 

124°26'00"W 

124°25'30*W 

122°59'00*W 

122°19'(XrW 

122°04'00*W 

123°10'4(rW 

123°1O'OO*W 

l2y\2'4O"Vi 

123°26'55"W 

123 o 29'00"W 

123°14'15'^ r 

95 

Depth (m) 

144 

93 

435 

88 

73 

108 

62 

91 

75 

265 

97 

97 

80 

57 

48 

86 

37 

221 

656 

49 

48 

37 

57 

55 

55 

274 

486 

48 

38 

302 

37 

49 

57 

48 

134 

66 

57 

82 

84 

207 

313 

71 

77 

84 

84 

49 

29 

38 

53 

49 

71 

62 

305 

119 

Date 

09 APR 1888 

02 MAY 1888 

01 SEP 1888 

06 SEP 1888 

06 SEP 1888 

25 SEP 1888 

26 SEP 1888 

19OCT1888 

19OCT1888 

05 JAN 1889 

05 JAN 1889 

05 JAN 1889 

08 JAN 1889 

08 JAN 1889 

16 JAN 1889 

17 JAN 1889 

26 JAN 1889 

04FEB 1889 

04FEB 1889 

05FEB 1889 

05FEB 1889 

05 FEB 1889 

06FEB 1889 

06 FEB 1889 

06 FEB 1889 

06 FEB 1889 

07 FEB 1889 

09 FEB 1889 

11 FEB 1889 

11 FEB 1889 

11 FEB 1889 

11 FEB 1889 

11 FEB 1889 

11 FEB 1889 

11 FEB 1889 

12 FEB 1889 

12 FEB 1889 

12 FEB 1889 

12 FEB 1889 

28 FEB 1889 

28 FEB 1889 

16 MAR 1889 

02 SEP 1889 

03 SEP 1889 

03 SEP 1889 

10 MAR 1890 

12 MAR 1890 

13 MAR 1890 

22 MAR 1890 

22 MAR 1890 

22 MAR 1890 

24 MAR 1890 

28 MAR 1890 

28 MAR 1890


Station 

3315 

3317 

3324 

3401 

3443 

3445 

3449 

3451 

3452 

3459 

3465 

3593 

3666 

3707 

3708 

3716 

3718 

3720 

3730 

3738 

3746 

3764 

4225 

4328 

4332 

4359 

4361 

4370 

4373 

4376 

4377 

4397 

4431 

4451 

4463 

4518 

4534 

4543 

4550 

4551 

4552 

4553 

4555 

4768 

4782 

4784 

4788 

4789 

4791 

4792 

4807 

4860 

4891 

4892 

4894 

4903 

4904 

4906 

4908 

Latitude 

54°02'40"N 

53°57'40'"N 

53°33'50"N 

00°59WS 

48°13'30"N 

48°16'00"N 

48°29'40"N 

48°25'10"N 

48°24'40"N 

48°24'20"N 

48°21'00"N 

48°ll'30"N 

36°45'00"N 

35°02'-N 

35°02'-N 

35°02'-N 

35°02'-N 

35°O2'-N 

34°40'-N 

34°58'-N 

35°02'-N 

34°57'-N 

55°--N 

32°--N 

32°--N 

32°--N 

32°--N 

32°--N 

32°--N 

32°--N 

32°--N 

33°1O / 15"N 

34°--N 

36°45'-N 

36°45'-N 

36°45'-N 

36°45'-N 

36°45'-N 

36°45'-N 

36°45'-N 

36°45'-N 

36°45'-N 

36°45'-N 

54°20 / 30"N 

52 o 55'00*N 

52°55'4O"N 

54°50 / 24~N 

54°49'45"N 

54°36'I5*N 

54°36'15"N 

4I°36'I2"N 

36°l8'00"N 

32°27'00~N 

32°27'3O"N 

32°33'OO"N 

32°31'10*N 

32 o 31'20"N 

3I°39'OO1S 

31°4OWN 

Station List.—Continued. 

Longitude 

166°42'00"W 

166°59'00"W 

167°46'50"W 

88°58'30"W 

123°ll'20"W 

123°45'05"W 

124°40'10*W 

124°37'50"W 

124°29'10"W 

124°24'40"W 

123°14'00"W 

122°48'00"W 

12r53'00*W 

138°46'-E 

138°46'-E 

138°46'-E 

138°46'-E 

138°46'-E 

138°20'-E 

138°45'-E 

139°50'-E 

139°53'-E 

130°--W 

117°--W 

117°--W 

117°--W 

117°--W 

117°--W 

117--W 

117--W 

117--W 

121°42'15"W 

120°10'-W 

121°55'-W 

121°55'-W 

121°55'-W 

121°55'-W 

121°55'-W 

121°55'-W 

121°55'-W 

121°55'-W 

121°55'-W 

121°55'-W 

179°O9'3O*E 

173°27'00*E 

173°26'00E 

167°13'00*E 

167°12'30^ 

166°58'15"E 

I66°57'15"E 

140°36'00"E 

129°44'00^ 

128°34'0O*E 

128°33'0O*E 

128°32'linE 

128°33'20"E 

128°32'4O*E 

129°2O'3OTE 

129°29'40*E 

Depth (m) 

507 

302 

199 

722 

177 

183 

247 

193 

229 

225 

88 

68 

124 

115 

110 

119 

119 

115 

62 

205 

90 

80 

272 

104 

113 

179 

166 

181 

174 

170 

232 

4016 

69-82 

? 

88-203 

121 

139 

97 

91 

84 

121 

119 

121 

1397 

104 

247 

102 

102 

132 

132 

80 

223 

331 

331 

174 

196 

196 

675 

796 

Date 

15 AUG 1890 

16AUG 1890 

20 AUG 1890 

28 MAR 1891 

27 AUG 1891 

27 AUG 1891 

28 AUG 1891 

28 AUG 1891 

29 AUG 1891 

02 SEP 1891 

04 SEP 1891 

30 APR 1894 

13 APR 1897 

08 MAY 1900 

08 MAY 1900 

11 MAY 1900 

11 MAY 1900 

11 MAY 1900 

16 MAY 1900 

17 MAY 1900 

19 MAY 1900 

22 MAY 1900 

06 JUL 1903 

08 MAR 1904 

09 MAR 1904 

15 MAR 1904 

15 MAR 1904 

16 MAR 1904 

17 MAR 1904 

17 MAR 1904 

17 MAR 1904 

01 APR 1904 

15 APR 1904 

11 MAY 1904 

13 MAY 1904 

24 MAY 1904 

28 MAY 1904 

01 JUN 1904 

07 JUN 1904 

07 JUN 1904 

09 JUN 1904 

09 JUN 1904 

09 JUN 1904 

03 JUN 1906 

09 JUN 1906 

11 JUN 1906 

14 JUN 1906 

14 JUN 1906 

14 JUN 1906 

14 JUN 1906 

16 JUL 1906 

31 JUL 1906 

09 AUG 1906 

09 AUG 1906 

09 AUG 1906 

10 AUG 1906 

10 AUG 1906 

11 AUG 1906 

11 AUG 1906

NUMBER 557 97 

Station 

4909 

4911 

4912 

4913 

4915 

4916 

4919 

4924 

4925 

4933 

4935 

4936 

4937 

4944 

4956 

4957 

4958 

4959 

4960 

4966 

4967 

4968 

4969 

4970 

4972 

4973 

4975 

4979 

4982 

4994 

5054 

5055 

5056 

5070 

5071 

5083 

5086 

5088 

5091 

5092 

5093 

5094 

5310 

5311 

5312 

5313 

5314 

5315 

5369 

5371 

5381 

5444 

5445 

5582 

5593 

T7414.B2 

r7414,B4 

H8O2.B 

Latitude 

31°38'30"N 

31°38'3O"N 

31°39'40"N 

31°39'10"N 

31°31WN 

30°25'00"N 

30°34WN 

3O°O5'OO"N 

Station List.—Continued 

Longitude 

129°27'30*E 

129°19'00"E 

129°20'(XrE 

129°22'30*t 

129°25'3CTE 

129°06'40"E 

129°19'30T 

130°21'20*E 

Off Yaku Shima, Osumi Shoto 

3O°59'OO"N 

30°57'20"N 

30°54'40"N 

31°13'OO"N 

31°38'15'TM 

32°32'00"N 

32°36'(XrN 

32 o 36 / 20*N 

32°36'30"N 

32°34'00"N 

33°25'20"N 

33°25'10"N 

33°24'5(TN 

33°23'40"N 

33°23'30"N 

33°25'45"N 

33°24'15'TM 

33°21'3(rN 

33°53'OO"N 

43°00WN 

45°27'50"N 

34°52'45"N 

34°53'00"N 

34°57'35"N 

35°O3'25"N 

35°O3'1O"N 

34°04'20"N 

35°O8'15"N 

35°U'25"N 

35°O4'1O*N 

35°O4'5O"N 

35°O3'15"N 

35°04'42*N 

21°33'OO"N 

21°33'OO"N 

21°30WN 

21°3O'OO'N 

21°41'00"N 

2l°40WN 

13°48'00"N 

13°49'40"N 

13°14'15*TM 

12°43'51'TM 

12°44'43'N 

04°19'54"N 

04°02'4O"N 

34°36.6'N 

34°45.8'N 

35°03.9'N 

130°29'50'E 

13O°35'1O^ 

130°37'30"E 

130°43'10"E 

130°46'50TE 

132°25'OOT 

132°23'00"t 

132°24'30"E 

132°23'2(TE 

132°21'45T 

135 O 36'2CTE 

135°37'2O*E 

135°38'40"E 

135°33'OO*E 

135°36'3(TE 

135 o 33'0CTE 

135°30'30*E 

135°38'5O"E 

137°42'00*E 

140°10'30"E 

140°54'0O*E 

138°42'20"E 

138°44'15"E 

138°43'35*E 

m°474(rE 

138°49'50^ 

137°57'3(TE 

139°20'0(rE 

139°28'2(TE 

139°38'12*E 

139°38'18*E 

139°37'42'E 

139°38'20"E 

116°13'00nE 

U6°15'(XrE 

U6°32'00^ 

116°43'00'E 

116°46'00"E 

U6 o 58'00*E 

121°43'00'E 

121°40'15''E 

122°44'45'E 

124°58'50"E 

124°59'5(TE 

118°58'38"^ 

118 o H'20^ 

RA' Tansei Maru (TM) 

138°43.8'E 

138°42.5'E 

138°46.9'E 

Depth (m) 

796 

715 

715 

715 

783 

660 

805 

291 

302 

278 

188 

188 

106 

79 

1317 

799 

741 

741 

1057 

446 

446 

463 

1073 

914 

805 

1097 

997 

1727 

713 

348 

516 

227 

472 

198 

104 

1141 

534 

675 

360 

128 

553 

161 

183 

161 

256 

274 

223 

271 

194 

152 

161 

563 

699 

1628 

70 

150-340 

312-328 

300-370 

Date 

11 AUG 1906 

12 AUG 1906 

12 AUG 1906 

12 AUG 1906 

12 AUG 1906 

13 AUG 1906 

13 AUG 1906 

14 AUG 1906 

14 AUG 1906 

16 AUG 1906 

16 AUG 1906 

16 AUG 1906 

16 AUG 1906 

17 AUG 1906 

23 AUG 1906 

23 AUG 1906 

23 AUG 1906 

23 AUG 1906 

23 AUG 1906 

29 AUG 1906 

29 AUG 1906 

29 AUG 1906 

29 AUG 1906 

30 AUG 1906 

30 AUG 1906 

30 AUG 1906 

31 AUG 1906 

01 SEP 1906 

19 SEP 1906 

22 SEP 1906 

12OCT1906 

12OCT1906 

12OCT1906 

15OCT1906 

15OCT1906 

20OCT1906 

23OCT1906 

25OCT1906 

26OCT1906 

26OCT 1906 

26OCT1906 

26OCT1906 

04NOV 1908 

04 NOV 1908 

04 NOV 1908 

04 NOV 1908 

05 NOV 1908 

05 NOV 1908 

24 FEB 1909 

24FEB 1909 

06 MAR 1909 

03JUN 1909 

03 JUN 1909 

26 SEP 1909 

29 SEP 1909 

19 SEP 1974 

23 SEP 1974 

10 FEB 1978


Station 

KT7802.Z2 

KT7802.Z4 

KT7802,Z61 

KT7811.OT1 

KT7811.OT4 

KT7811.OT6 

KT7811.OT6-2 

KT7818.OT8-1 

KT7818.OT10 

KT7911,OT4 

KT8412.14 

KT8916.T3-2 

KT9015.BS1 

KT9015,BS2 

KT9015.CB0-1 

KT9015.CB1-1 

KT9015.CB1-2 

KT9015.CB2-2 

KT9015.CB6-1 

KT9015.HK2 

KT9015.HK3 

KT9015.HK5 

KT9015.OK1 

KT9015.OK2 

KT9202.AT1 

KT9202.KB3 

KT9202.OS2 

KT9202.OS3 

KT9202.YS1 

KT9202.YS2 

KT9202.YT1 

KT9202.YT2 

KT9202.YT4 

KT9202.YT5 

KT9202.YT6 

1-124 

2-216 

2-224 

3-312 

3-314 

4-426 

5-505 

6-606 

7-721 

8-803 

9-907 

9-910 

10-1007 

10-1017 

2078 

2209 

3353 

5626 

5638 

5640 

Latitude 

34°37.9TM 

34°36.5'N 

34°43.5TM 

34°56.1'N 

34°56.3'N 

35°03.4TM 

35°O3.3'N 

34°55.2'N 

34°53.9'N 

34°46.7'N 

33°29.1'N 

32°19.09TM 

32°50.50TM 

32°43.38'N 

34°52.90'N 

34°54.70'N 

34°54.49'N 

34°58.O8N 

35°45.17N 

34°25.23'N 

34°30.11'N 

34°55.07'N 

35°45.20TM 

35°49.80TS[ 

29°13.67'N 

31°08.41'N 

30°58.42'N 

31°06.17'N 

29°46.59'N 

29°44.02'N 

30°14.80^J 

30°10.10'N 

30°04.64'N 

30°02.43'N 

29°59.28'N 

34°49'N 

34°48'N 

34°41'N 

34°43'N 

34 O 44TM 

34°44'N 

34°43'N 

34°46'N 

34°45'N 

34°44TM 

34°43'N 

34°43'N 

34°45'N 

34"42'N 

44°09TM 

49°46'N 

53°53'N 

45°26'N 

44°36'N 

44°4nM 


Longitude 

138°27.0^ 

138°25.6'E 

138°30.2'E 

138°44.8'E 

138°43.1'E 

138°49.8'E 

138°5O.O^ 

138°44.8'E 

138°43.1'E 

138°41.4'E 

135°28.9'E 

134°00.85'E 

132°05.22'E 

132°06.52'E 

131°11.09'E 

131°07.48'E 

131°07.56'E 

131°04.87'E 

132°52.22'E 

130°46.83'E 

130°52.24'E 

131°18.74'E 

133°09.92'E 

133°26.27'E 

129°45.22'E 

130°39.92'E 

130°31.82'E 

130°34.88TE 

13O°24.11TE 

130°29.12'E 

130°46.10'E 

130°52.50'E 

130°55.73'E 

130°57.37'E 

130°57.88'E 

PULSE Cruises (R/\ New Horizon) 

123°07'W 

123°00'W 

123°08'W 

123°11'W 

123°H'W 

123°08'W 

123°10'W 

U3°06'^ 

123°07'W 

123°07'W 

123°07'W 

123°06'W 

123°04'W 

123°O3'W 

Vityaz (IOM) 

148°09^ 

169°15'E 

154°12'E 

149°07'E 

148°57'E 

Depth (m) 

95-100 

138-145 

70-71 

198-205 

490-500 

105-110 

108-120 

210-245 

382-425 

380-457 

214-235 

1917-1950 

89-90 

193-199 

66-72 

71-78 

70-71 

74-78 

100-101 

86-87 

93-95 

97 

71 

93-96 

1058-1065 

100-101 

237-241 

143-144 

150-151 

238-240 

80-88 

95-98 

402-410 

660-711 

964-1005 

4100 

4100 

4100 

4100 

4100 

4100 

4100 

4100 

4100 

4100 

4100 

4100 

4100 

4100 

1080 

3660 

1680 

5220 

1675-1847 

780 

Date 

08 FEB 1978 

08 FEB 1978 

08 FEB 1978 

13 JUL 1978 

13 JUL 1978 

14 JUL 1978 

14 JUL 1978 

20 NOV 1978 

20 NOV 1978 

19 JUL 1979 

01 SEP 1984 

08 NOV 1989 

03 NOV 1990 

03 NOV 1990 

02 NOV 1990 

02 NOV 1990 

02 NOV 1990 

02 NOV 1990 

01 NOV 1990 

29 OCT 1990 

29 OCT 1990 

02 NOV 1990 

01 NOV 1990 

01 NOV 1990 

18 FEB 1992 

21 FEB 1992 

20 FEB 1992 

20 FEB 1992 

18 FEB 1992 

18 FEB 1992 

16 FEB 1992 

17 FEB 1992 

17 FEB 1992 

17 FEB 1992 

17 FEB 1992 

25 JUN 1989 

26 OCT 1989 

30 OCT 1989 

17 FEB 1990 

18 FEB 1990 

24 JUN 1990 

23 OCT 1990 

20 FEB 1991 

24 JUN 1991 

22 JUL 1991 

01 AUG 1991 

02 AUG 1991 

21 OCT 1991 

25 OCT 1991 

11 MAY 1953 

7 

05 JUN 1955 

20 AUG 1966 

10 SEP 1966 

10 SEP 1966

NUMBER 557 99 

Station 

MV66-II-4 

MV69-VI-9 

MV70-III-22 

MV70-III-1 

MV70-III-6 

20134 

20136 

26499 

26502 

23 

31 

42 

43 

51 

Agassiz MV71-I-1 

Alpha Helix-30 

CR79-1 

CR79-8 

CR79-18 

Horizon MET-123 

//omonMET-133 

Keldish-TiU 

Lets Go-80 

LM49 

MUSORSTOM2-33 

PUIsbury-52\ 

Pillsbury-530 

Pillsbury-E6l 

SEPBOP 18B-764 

Tsuchida-45 

Tcuchida-102 

Tsuchida-197 

Tsuchida-272A 

Tsuchida-851 

Velero 

4-7228-60 

Washington 

MV67-III-22 

PPTU-II 

YO69-3 

YO69-37-39 

YO70-770 

YO70-1009 nF 

67-41 GBR 

69-RD3 

Latitude 

40°30.5'N 

31 O 12TM 

31°19.7'N 

31°47.0'N 

31 "36^1 

60°22'N 

60°21'N 

59° 12.41*4 

59°21.7 

33°21'N 

32°30'N 

32°34'N 

34°25'N 


Longitude 

R/V Melville (SIO) 

119°37'W 

119°39.2W 

120°12'W 

120°07.4'W 

OCSEAP(R/V5eart7i

Abe, N. 

1939. [Ecological Studies on Rhizopsammia minuta var. mutsuensis Yabe 

& Eguchi.] Jubilee Publication for Professor H. Yabe's 60th 

Birthday, 1:175-187. [In Japanese, not seen.] 

Addicott, W.O. 

Literature Cited 

1966. Late Pleistocene Marine Paleoecology and Zoogeography in Central 

California. Contributions to Paleontology, Geological Survey 

Professional Paper, 523G1-21,4 plates. 

Alcock, A. 

1893. On Some Newly-Recorded Corals from the Indian Seas. Journal of 

the Asiatic Society of Bengal, 2(62): 138-149, plate 5. 

1894. Natural History Notes from H. M. Indian Marine Survey Steamer 

"Investigator," series 2, number IS: On Some New and Rare Corals 

from the Deep Waters of India. Journal of the Asiatic Society of 

Bengal, 2(62): 186-188. 

1898. An Account of the Deep-Sea Madreporaria Collected by the Royal 

Indian Marine Survey Ship "Investigator." 29 pages, 3 plates. 

Calcutta: Trustees of the Indian Museum. 

1902a. Diagnoses and Descriptions of New Species of Corals. Tijdschrift 

der Nederlandsche Dierkundige Vereeniging, (2)7:89-115 [published 

in July]. 

1902b. Further Diagnoses and Descriptions of New Species of Corals. 

Tijdscrift der Nederlandsche Dierkundige Vereenining, (2)7:116— 

123 [published in July]. 

1902c. Report on the Deep-Sea Madreporaria of the Siboga- Expedition. 

Siboga-Expeditie, 16a: 52 pages, 5 plates [published in August]. 

Austin, W.C. 

1985. An Annotated Checklist of Marine Invertebrates in the Cold 

Temperate Northeast Pacific, 1: 218 pages. Khoyatari: Khoyatan 

Marine Laboratory, British Columbia. 

Bedot, M. 

1907. Madre"poraires d'Amboine. Revue Suisse de Zoologie, 15:143-292, 

plates 5-50. 

Blainville, H.M.D. de 

1830. Zoophytes. Dictionnaire des Sciences Naturelles, 60: 546 pages. 

Strasbourg: F. G. Levault. 

1834. Manuel d Actinologie ou de Zoophytologie. i-viii + 694 pages, 103 

plates. Paris. 

Boschma, H. 

1923. The Madreporaria of the Siboga Expedition, Part 4: Fungia patella. 

Siboga-Expeditie, 16d: 20 pages, plates 9, 10. 

Bourne, G.C. 

1903. Some New and Rare Corals from Funifuti. Journal of the Unnaean 

Society of London, Zoology, 29:26-37,2 plates. 

1905. Report on the Solitary Corals Collected by Professor Herdman, at 

Ceylon, in 1902. Ceylon Pearl Oyster Fisheries, Supplementary 

Reports, 29:187-242, plates 1-4. 

Briggs, J.C. 

1974. Marine Zoogeography, x + 475 pages. New York: McGraw Hill 

Book Company. 

Broderip, W.J. 

1828. Description of Caryophyllia smithii n. sp. Zoological Journal, 

3:485-486, plate 13. 

Bronn, H.G. 

1837. Lethaea Geognostica. Ire Edition, volume 3. Stuttgart [not seen]. 

Brusca, G, J., and R.C. Brusca. 

1978. A Naturalist's Seashore Guide. 205 pages. Eureka: Mad River Press. 

100 

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Plates 1-42


PLATE \.—Fungiacyathus marenzelleri (aj>. Pulse 2-215, USNM 87616; c4. ViQwz-6143 (paratype of F. 

symmetricus aleuticus Keller, 1976), IOM; e, Vityaz-5605 (paratype of F. symmetricus aleuticus), IOM;/. Pulse 

1-124, USNM 85790): a.b, stereo calicular and basal views of a well-preserved corallum, both x 2.0; c-e, oblique 

and basal views of Vityaz specimens, all x 2.0;/, lateral view of an S, showing trabeculae, synapticulae, basal 

plate, and costa, x 10.

110 


PLATE 2.—Leptopemts discus (a-c, Melville 70-111-1, SIO Co 1271; d, paratype of L. irinae, Viryaz-4158, USNM 

92435): a,b. oblique and stereo lateral views of a corallum, x 12.5, x 13.5, respectively; c, enlargement of 2b 

showing porous network between septa and costae, x 42; d, calicular view, x 3.8. Astrangia haimei (e, lectotype 

of A. haimei, YPM 598a;/, off Monterey wharf, CAS 74819; g, off Coronado, Baja California, SIO Co 1170): e, 

colony, x 2.3;/. small hemispherical colony, x 1.6; g, granular costae, x 45.


PLATE 3.—Astrangia haimei (a-c, off Coronada, Baja California, SIO Co 1170): a. calice, x 12; bx. progressive 

enlargement of columella and paliform lobe region, x 21, x 51, respectively; Oculina profunda (d, holotype, 

USNM 84792; e, paratype, USNM 36372): d, branches, x 1.1; e, stereo view of branch segment showing calices 

and granular costae, x 14. Madrepora oculata (f,h. Rebelling Guyot, USNM 83581; g,off Enoshima, Sagami Bay, 

USNM 92658):/, h, branch fragment and calice, x 0.73, x 15, respectively; g, colony, x0.76.

112 


PLATE 4.—Caryopkyllia arnoldi (ajb. holotype, USNM M1575O9; c. Alb-4463, USNM 77419; d, Alb-4359, 

USNM 92470; e. 49°27'N. 124°18'W, RBCM 976-1030-1): aj>, lateral and calicular views, x 3.0, x 3.1, 

respectively; 14. calicular views of two specimens, both x 3.0 (d is a stereo pair); e, lateral view of costae and 

upper theca, x 3.5. Caryopkyllia ataskensis (f-h. syntype, USNM M547317; j, Alb-4792, USNM 83523):/. h, 

calicular views of a syntype, x 5.3, x 4.5, respectively if is a stereo pair); g, lateral view of same specimen, x 3.2; 

;, calicular view of a specimen, x 3.4. Caryopkyllia japonica: i, V/ryaz-5638, C. ambrosia of Keller (1981 a), IOM, 

x3.0.


PLATE 5.—Caryophyllia sp. A from Alb-4784, USNM 92478: a,b. calicular view and lateral views, x 1.5, x 1.4, 

respectively. Caryophyllia pedroensis, holotype, USNM Ml64736: c.f. lateral and calicular views, x 2.3, x 3.9, 

respectively. Caryophyllia californica. Pleistocene of San Pedro, California, USNM 92479: d,e, lateral and 

calicular views, x 3.0, x 5.2, respectively. Labyrinthocyathus quay lei (g, Alb-4551, paratype, USNM M547417; 

/»,«', holotype, UCMP 30245): g. a cluster of solitary coralla attached to one another, x 3.0; h,i, lateral and calicular 

views of holotype, x 2.6, x 3.6, respectively.

114 


PLATE t.—Labyrinthocyathus quaylei (a, Alb-4551, USNM M547418; b. Monterey Bay, CAS 16405): a, stereo 

view of calice. x 3.7; b. lateral view of corallum, x 3.2. Crispatotrochusfoxi (c, holotype, SBMNH; d,e, Alb-3324, 

USNM 19210): c, calicular view of holotype, x 3.5; d,e, calicular and lateral views of second specimen, x 3.1, x 

2.4. respectively. Paracyathus stearnsii (f. Alb-4431, USNM 92602; g. Bear Rock, IslaSan Martin, SIOCo 1171; 

h, holotype of P. tibwonensis. UCMP 30485; i, off Santa Cruz, SIO CO 1167; j, holotype of P. caltha, YPM 965; 

*, off Pacific Grove, CAS 74821): /. typical calice with five septal cycles, x 3.2; g, large calice with over five 

septa! cycles, x 3.3; h. x 5.6; i. irregularly shaped coralla, x 2.5; j. x 4.7; *. broken corallum exposing paliform 

lobes, x 5.3.


PLATE l.—Paracyathus stearnsii (a-c, off Santa Catalina, CAS 74828; d-f, holotype of P. stearnsii, YPM 

2187): a-c, coral la illustrating early stages of polycyclic development, x 20. x 18, x 42, respectively; d-f, theca, 

calice, and polycyclic base of holotype, x 3.0, x 2.9, x 3.3, respectively. Paracyathus montereyensis (gji. 

Alb-2888, USNM 92604; i, holotype, UCPM 30341): g,h, lateral and calicular views of Oregon specimen, x 3.6, 

x 4.3, respectively; i, calicular view of holotype, x 9.0.

116 


PLATE S.—Coenocyathus bowersi (a. off Santa Cruz, USNM 78631; h, holotype, USNM 21138; c. Alb-2978, 

USNM 19245; d.e. off Bird Rock, SIO Co 1168;/, off La Jolla, USNM 92920): a. cerioid corallum, x 0.95; b, 

holotype, x 1.3; c, branching corallum, x 0.85; d-e. intratentacular division, x 2.8, x 5.9, respectively;/, regular 

calice with four septal cycles, x 5.0. Nomlandia californica: g, stereo pair of holotype, SBMNH 35560, x 6.0. 

Paracyathus pedroensis: h, calice of holotype, USNM Ml64738, x 4.5.


PLATE 9.—Desmophyltum dianthus . neotype. USNM 92475; cjd. Reberling Guyot, USNM 83583): ajb, 

lateral and calicular views, x 0.8, x 1.0, respectively; cA, calice and corallum of Californian specimen having five 

cycles of septa, x 1.0, x 0.9. respectively. Lophelia pertusa (e,f, Alb-2946, USNM 92606; g, TM (KT7414, B2), 

USNM 92611; h,i, holotype of Dendrosmilia nomlandi, SBMNH 35559): e, bushy corallum, x 0.8;/. aspect of 

dense branching, x 2.3; g, specimen from Sagami Bay, x 2.0; h.i, a calice and the entire holotypic branch 

fragment, x 5.6, x 2.1, respectively.

118 


PLATE \O.—Flabellum sp. A: ah. NMCIC 1982-1492, stereo calicular and lateral views of a specimen, x 2.4, x 

2.3, respectively. Javania horealis: c4, stereo calicular and lateral views of holotype, USNM 82019, x 1.3, x 1.2, 

respectively. Javania californica: e,f. stereo calicular and lateral views of holotype, USNM 92613, x 4.0, x 3.7, 

respectively. Javania cailletifg, LM 49. off British Columbia, NMCIC 1900-8362; h,i, Alb-4982, USNM 92747): 

g, a broken calice, x 1.9; h.i. calicular and lateral views of a specimen, x 3.1, x 1.6, respectively.


PLATE 11.—Polymyces montereyensis (a,b, holotype, USNM M547406; c, Alb-2977, USNM 83529; d,e, 

holotype of F. tannerense, SBMNH 35562;/, SEPBOP 18B-764, USNM 83521): a,b, lateral and stereo calicluar 

views of holotype, x 1.9, x 2.0, respectively; c, broken base revealing rootlet development consistent with stage 

diagramed in figure 3a, x 4.7; dje, calicular and lateral views of damaged holotype, x 5.0, x 3.7, respectively,/, 

highly encrusted theca of large Peruvian specimen, x 1.4. Balanophyllia cedrosensis (g, holotype of B. 

tiburonensis, UCMP 14833; h. Velero-1257-4], SBMNH; i.j, holotype of 8. cedrosensis. CAS 29961): g, calice, 

x 5.1; h, calice, x 3.6; i,j, lateral and stereo calicular views of holotype, x 2.7. x 3.9, respectively.

120 


PLATE \2.—Balanophyllia elegam (a. Snipe Bay, Alaska, UA; b,c. Alb-4552, USNM 92627; d, CAS 74809; e.f, 

syntypc. MCZ 5472): a, calice of northernmost record, x 4.5; b,c. calicular and lateral views of largest specimen 

examined, x 3.2, x 2.8. respectively; d, calice, x 4.0; e.f, calicular and lateral views of a syntype, x 4.1, x 3.8, 

respectively. DendrophyUia californica (g.h, holotype, CAS 29960; i. Alb-2832, USNM 38113): g,h, calices of 

holotypic colony, x 1.7, x 3.2 (g is a stereo pair); i, calice, x 3.6. DendrophyUia oldroydae (/. off La Jolla, SIO Co 

1179; *, holotype of Faustino's D. oldroydi. CAS 36397): j. calice with a large columella, x 5.0; *, largest 

fragment of holotypic colony, x 0.6.


PLATE 13.—Dendrophyllia oldroydae (a, holotype of D. cortezi, SBMNH 35564; h, paratype of Faustino's D. 

oldroydi. UCMP 12200): a, colony, x 0.6; b, colony, x 0.7. Madracis sp. A (c-f, Sagami Bay, 110 m, ZMC): c, 

branch fragment, x 2.2; d-f, progressive enlargements of calices, x 15, x 26. x 57, respectively. Fungiacyathus 

stephanus (g,h, Alb-4916. USNM 80124; i, Alb-4919, USNM 80125): gji. lateral and oblique calicular views of 

a concave-based specimen, both x 3.0; i, oblique calicular view of a flat-based specimen, x 1.3.

122 


PLATE 14.—Fungiacyathus paliferus (a. d,e, TM (KT9202, YS1), USNM 92642; b.c. TM (KT9015, OKI). 

USNM 92638; TM (KT9015. BS2), USNM 92635): a, oblique calicular view showing paliform lobes, x 8.3; b,c, 

granular costal base of two Diaseris-fotm specimens, x 6.6, x 3.4, respectively; d, oblique view of a half-system 

showing small S5 and paliform lobes, x 16; e, oblique view of paliform lobes, x 20. Fungiacyathus sp. A: f-h, 

Vifyaz-2209, F. palifera of Keller, 1976, IOM, base and calicular views of larger corallum, all x 1.5.


PLATE 15.—Fungiacyalhus variegatus: a,b, Alb-4911, USNM 82018, calicular and basal views, both x 6.0. 

Lttepsammia formosissima: c,f. TM (CR79-8), USNM 92650. calicular and basal views, both x 2.6. 

Fungiacyathus granulosus: d,e, TM (KT9202, YT4), USNM 92645, calicular and basal views, both x 3.1. 

Leptopenus solidus: g,h, holotype, IOM, basal and stereo calicular views, both x 4.5. Rhombopsammia niphada 

(i,j, TM (KT92O2, YT5), USNM 92647; k, holotype, Alb-4911): i.j, calicular and basal views of a specimen, x 1.3, 

x 1.4, respectively; k, calice, x 1.4.

124 


PLATE Xb.—Stephanophylliafungulws (a-d.f.g.TM (KT9015. BS2), USNM 92652): a,b, oblique calicularand 

basal views, x 8, x 11. respectively; cjd. longitudinal view of fractured coral I urn revealing horizontal thecal wall, 

oblique fulturae. septal faces, and synapticulae, x 9, x 17, respectively;/, granular base at epicenter, x 51; g, 

lamellar columella, x 18. Rhomhopsammia niphada: e, base of holotype, Alb-4911, x 14.


PLATE 17.—Culicia japonka (a-e. off Sesoko, Okinawa, USNM 86826): a. lateral view of epitheca and stolon, 

x 17; b, stereo view of calice, x 17; c, lateral view of epithecate corallum, x 14; d, stereo view of corallites linked 

by stolons, x 2.3; e, paliform lobes on S2.3, x 78. Oulangia stokesiana miltoni: f, TM (KT9015, CB6), USNM 

92646, stereo view of calice, x 9.6.

126 


PLATE \%.—Cyathelia axillaris (a. Monoisa, Sagami Bay, USNM 92665; h,c, Alb-4944, USNM 92660): a, 

colony, x 1.0; bx, calices. x 2.1. x 2.9, respectively. Anthemiphyllia dentata (d,e, TM (KT9202, OS3), ORI;/, 

Alb-5094, USNM 92667): d.e, calicular and basal views, both x 5.0; /, basal view of long-dead corallum, x 2.2. 

Anthemiphyllia frustum (g-i, paratype. TM (KT9202, OS2), USNM 92669): g, oblique calicular view, x 8.5; h. 

oblique view of paliform lobes, x 17; i, edge view of thick septocostae, x 32.


PLATE 19 —Anthemiphyllia frustum: a,b, calicular and basal views of holotype, USNM 92668, both x 6.1. 

Caryophyllia japonica (c,f, Vifyaz-3353, USNM 92678, C. alaskensis of Keller, 1981a; d#, syntype, Enoshima, 

Sagami Bay, NM W 8168; e ,h, Alb-4982, USNM 82158; i, smaller syntype, NM W 8168): c,f, calicular and lateral 

views of a specimen with over 48 septa, x 2.8, x 2.3, respectively; d, g, larger syntype, x 2.3, x 3.4, respectively; 

e,h, lateral and calicular views of a specimen with 48 septa, x 2.2, x 2.9, respectively, i, syntype, x 2.8. 

Caryophyllia sp. cf. C. scobinosa: j-l, TM (KT7911, OT4), USNM 92680, lateral and calicular views of a 

specimen, x 3.7, x 5.1, x 5.3, respectively.

128 


PLATE 20.—Caryophhylia scobinosa: ajb, lateral and calicular views of a syntype from Siboga-45, ZMA Coel. 

574. x 2.4. x 3.0. respectively. Caryophyllia quadragenaria (c.f, TM (KT9202, OS2). USNM 92672; d,g. syntype, 

Siboga-25\, ZMA Coel. 5529; eji, 32° 10TM. \2i°2(fE, ZMC): c, calice. x 7.7; d,g, lateral and calicular views of 

a syntype. x 5.4, x 7.7, respectively; eji. lateral and calicular views of a specimen, x 3.9, x 6.5, respectively;/, 

lateral view of two specimens, x 2.3. Caryophyllia rugosa: i, calice of specimen from 32°21 TV, 128°49Ti, ZMC, 

x 10.5.

PLATE 21.—Caryophyllia rugosa: a, transverse costal rugae of specimen from Plate 20i, ZMC, x 26. 

Caryophyllia jogashimaensis: b.c, lateral and calicular views of specimen from TM (KT92O2. YT2). USNM 

92675, x 3.5, x 4.7. Caryophyllia ambrosia ambrosia {d,g, syntypes, lnvtstigator-\76, USNM 18157; e, Soyo 

Maru-223, TIUS 53625, C. alcocki of Yabe and Eguchi. 1942b; /, TM (KT9202, YT6). USNM 92794; h. 

Alb-4960, USNM 92792): d,g, lateral and calicular views of two syntypes, x 1.7, x 1.9, respectively; e, specimen 

with only 12 pali, x2.2;/, specimen with 18 pali, x 1.5; h, specimen with 20 pali, x 1.6. Caryophyllia grayi: (i-k. 

Kere River, Vanuatu, Pleistocene, USNM 73964): i, calice of a well-preserved specimen, x 2.4; j,k, lateral and 

calicular views of a specimen, x 1.7, x 2.6, respectively. Caryophyllia (A.) spiniger: I, lateral view of syntype from 

"off Japan," BM.x 3.1.

130 


PLATE 22.—Caryophyllia (A.) spiniger (ajb, d, Alb-5371, USNM 92690; c, syntype of Plate 21/. BM): ajb. 

specimen with long, broad edge spines, x 2.0, x 1.9, respectively; c, syntype, x 3.3; d, specimen having additional 

lateral thecal spines, x 2.2. Caryophyllia (P.) compressa: e.f, stereo calicular and lateral views of a corallum from 

TM (KT9015, BS2). USNM 92686, x 6.9, x 3.6. respectively. Crispatotrochus rubescens (g, Soyo Maru-425, 

T1US 53695; h, Soyo Marn-331. TIUS 53694 (Cyathoceras diomedae of Yabe and Eguchi, 1942b)): g, poorly 

preserved corallum with a Stenocyathus vermiformis attached in upper left quadrant, x 2.4; h, well-preserved 

corallum. x 2.2. Paracyathwt pruinosus: i-j, lateral and stereo calicular views of syntype from Siboga-96, ZMA 

Coel. 1307, x 2.1, x 2.7, respectively.


PLATE 23 —Trochocyathus caryophylloides (aji, Alb-4891, CAS 74940; bx, syntype. Siboga-253, ZMA Cod. 

1324): a,h. lateral and calicular views, both x 3.2; b,c. calicular and lateral views of a syntype, x 2.7. x 2.3, 

respectively. Trochocyathus decamera d,e, holotype, USNM 92693, lateral and stereo calicular views, x 5.7, x 

7.0. Trochocyathus cooperi

132 


PLATE 24.—Dtltocyathus vaughani (a-c, Alb-5088, USNM 92721; /, TM (KT7811, OT4), USNM 92723): a-c, 

calicular, basal, and lateral (edge) views of a specimen, x 2.7, x 2.6, x 2.7, respectively; /, specimen with 

ascothoracid gall in right portion of columella, x 3.3. Dellocyathus magnificus (d,e, syntype, Challenger-\92, 

BM; gjt. Tsuchida-102, USNM 92730): djt, only remaining syntype, both x 2.5; gjt, a Japanese specimen, x 1.6, 

x 1.8, respectively. Conotrochus funicolumna: i, Soyo Afarn-268, TIUS 50243, calice, x 4.0. Deltocyathus 

rotulus.jjk. stereo calicular and basal views of specimen from Alb-5079, USNM 92726, both x 1.7.


PLATE 25.—Stephanocyalhus spiniger: a-c, calicular, lateral, and basal views of a specimen from Tosa Bay, 

USNM 92733, all x 1.1. Stephanocyalhus weberianus (d,f, Alb-4958. USNM 92737; e, TM (KT9202, YT6), 

USNM 92743): d,f, specimen having an eroded base, both x 1.4; e, specimen having a strongly tuberculate base, 

x 1.4. Conotrochus funicolumna (g,j, syntype, Siboga-95, ZMA Coel. 709; hX Soyo Maru-323, TIUS 59174, 

syntype of Conotrochus parahispidus; i, Soyo Afaru-268, TIUS 50243: /. off Misake, USNM 92700): g.j. x 2.6. 

x 3.6, respectively; *,*, x 2.9, x 5.5, respectively; i, x 3.8; /, x 4.1.

134 


PLATE 26.—Aulocyathus recidivus: a,b, lateral and calicular views of a specimen from Alb-4958, USNM 92704, 

x 3.7. x 6.4. respectively. Aulocyathus matricidus {c*.f, syntypes. BM 1862.7.16.72; d,g, TM (KT7802. Z4), 

USNM 92703): c.f, both sides of a longitudinally fractured specimen, both x 2.7; e, small corallum budded from 

parent fragment, x 2.6; d.g. lateral and calicular views of a specimen, x 4.1, x 7.7, respectively. Aulocyathus 

juxensecens: h,i, syntypes. ZMB 5064. calice and corallum, x 15. x 5.2. respectively.


PLATE 27.—Coenosmilia sp. cf. C. arbuscula: a,b, calicular views of specimen from TM (KT9202, YS2), USNM 

92709, x 3.3, x 6.2, respectively. Rhizosmilia sagamiensis (c-e, Alb-4944, USNM 92705): c, small phaceloid 

coraJlum, x 1.4; d.e. lateral and calicular views of an individual corallite, x 2.6, x 4.0, respectively. Dasmosmilia 

pacifica: (fj, YO70-1009, ORI; gX Alb-3738, USNM 92710):/, i, regenerated corallum budded from parent 

corallum, x 3.1, x 3.3, respectively; g,h. corallum fragment illustrating costae and fascicular columella, both x 2.9. 

Goniocorella dumosa: j, colony fragment from Sagami Bay, USNM 92708, x 2.3. Notocyathus venustus: k,l, 

lateral and oblique calicular views of a corallum from TM (KT9015, BS2), x 5.0, 7.3, respectively.

136 


PLATE 2%.—Notocyathus conicus: a.b, lateral and calicular views of specimen from TM (KT9O15, CB1-2), 

USNM 92777. x 6.7, x 9.1, respectively. Peponocyathus australiensis (c.f, TM (KT9015, HK5), USNM 92766; 

dx. TM (KT7414, B4). USNM 92917): c.f, calice and costae of typical form, x 9.8, x 66, respectively; d,e. calice 

and basal costae of form japonkus. both x 4.1. Peponocyathus follkulus (g-k, TM (KT9015, CB1-2), USNM 

92774): gji, oblique calicular views 0? is a stereo pair), x 11. x 10, respectively; i.j, enlargements of flat base, x 

11. x 18. respectively; k. costae. x 58.


PLATE 29.—Tropidocyathus lessoni: a,b, lateral and calicular views of a specimen from TM (KT9202, YT2), 

USNM 92779, x 4.6, x 4.9, respectively. Stenocyathus vermiformis (r. 27°24'N, 79°26'W, USNM 46646; /, 

Pillsbury-96\, USNM 46649): c,f. lateral and calicluar views of western Atlantic specimens, x 3.1, x 20, 

respectively. Tropidocyathus pileus: d,e, lateral and caliclular views of a specimen from TM (KT9015, BS2), 

USNM 92781, x 3.5, x 3.9, respectively. Alatotrochus rubescens {g.jjc. TM (KT9015, BS2), USNM 92776; h j.1, 

syntype, Challenger-\92, BM 1880.11.25.163): f>,k. lateral and calicular views of a specimen, x 3.2, x 3.5, 

respectively; h,i,l, oblique basal, calicular, and oblique calicular views of a paratype, all x 2.6; j, lateral view of 

a specimen showing the distinctive costal development, x 3.5.

138 


PLATE 30.—Motrochus kikutii (a-d. Philippine specimen from MUSORSTOM 2-33, USNM 81911): a.b, lateral 

and calicular views, x 20. x 18, respectively; c, calicular edge showing alternation of costae and septa, x 125; d, 

cannae on septal faces, x 60. Truncatoguynia irregularis: e,f. lateral and calicular views of a specimen showing 

characteristic thecal weathering, x 8.2. x 10. respectively. Flabellum pavoninum (g-i, specimens reported as F. 

distinction by Yabe and Eguchi, 1942a): gji, Sagami Bay, coalitum form, TIUS 43409. x 1.3, x 1.5, respectively; 

I. Soyo Maru-429. TIUS 39732, x 1.4.


PLATE 31.—Flabellum pavoninum (a-e, Marenzeller's (1888a) Japanese specimens): a, "off China," NMW 

8201, x 1.1; b.c. "off Japan." "F. distinction." NMW 8197, both x 1.1: d.e. holotype of F. coalitum, "off Japan," 

NMW 8196, x 1.6, x 1.7, respectively. Flabellum angustum.f. syntype, TIUS 43436, Pliocene of Shikoku. x 2.3. 

Flabellum patens: g-i. specimen from YO70-770, USNM 92814, lateral, calicular, and edge views, x 1.2, x 1.2, 

x 1.4. respectively. Flabellum magniftcum (/. F. pavoninum of Yabe and Eguchi, 1942a, Soyo Maru-437, TIUS 

43448; *./. Soyo Maru-416, TIUS 50094):;. x 1.1; *,/, x 0.7, x 0.8, respectively.

140 


PLATE 32.—Flabellum politum: a-c. lateral, calicular. and edge views of specimen from Alb-4903, USNM 

40735. all x 2.7. Flabellum deludens: d.e. lateral and caJicular views of a specimen from Alb-5094, USNM 40761, 

both x 1.2. Flabellum japonkum: g.h. lateral and calicular views of a specimen from YO69-3, USNM 92828, x 

1.05. x 0.95. respectively. FlaMlum apertum horealis (f.i,l, holotype. USNM 40710; jjc. paratype, Alb-5083, 

USNM 40709):/././, calicular. basal, and lateral views, all x 1.5; jjc. lateral and calicular views, both x 1,5.

PLATE 33.—Truncatoflabellum spheniscus: a-d, TM (KT9015, HK2). USNM 92799, lateral, calicular, edge, and 

basal views of a specimen, x 1.8, x 1.9, x 2.3, x 2.6, respectively. Truncatoflabetlum candeanum (e, Soyo 

Maru-465, TIUS 50229, F. ruhrum (in part) of Yabe and Eguchi, 1942a;/. TM (KT9015, HK3), USNM 92802): 

e, three anthocyathi, x 2.7;/, anthocaulus with short anthocyathus still attached, separated by thin white crescent 

one-quater distance from top of corallum, x 3.9. Truncatoflabellum formosum: gji, lateral and calicular views of 

a specimen from TM (CR7918), USNM 92804, x 1.7, x 2.0, respectively. Truncatoflabellum sp. B: /,/. calicular 

and lateral views of a specimen from TM (KT92O2, YT1), USNM 92811, x 10.1, x 3.2, respectively. 

Truncatoflabellum carinatum: j-k, lateral and calicular views of a specimen from Pescadores Islands, Formosa 

Strait, USNM 92806, x 2.1, x 2.2, respectively.

142 


PLATE 34.—Truncatoflabtllum gardineri: ajb, lateral and stereo caiicular views of specimen from TM (KT9202, 

OS3), USNM 92809, x 2.6, x 3.0, respectively. Truncatoflabellum sp. A: c-e, lateral, edge, and caiicular views 

of specimen from TM (KT9202, YT6), USNM 92810, all x 1.3. Placotrnchides scaphula: f-h, lateral, caiicular, 

and basal views of specimen from TM (KT7911, OT4), USNM 92748, x 5.1, x 5.9, x 6.2, respectively. Javania 

insignis: i-k, lateral and caiicular views of a specimen from off Jogashima, Sagami Bay, USNM 92746, x 1.6, x 

2.4, x 2.4. respectively.


PLATE 35—Rhizotrochus typits (a,b, Sagami Bay, USNM 92751; c, Okinose, Sagami Bay, ZMC): a,b, calicular 

and lateral views of a specimen, x 3.2, x 2.7, respectively; c. basal view showing pedicel and ring of six rootlets, 

x 3.5. Balanophyllia cumingii: d,e, lateral and calicluar views of TM (KT9015, HK2), USNM 92887), x4.7, x 5.5, 

respectively. Balanophyllia cornu (f,i, syntypes, Challenger-206, BM 1880.11.25.143; g.h, Formosa Strait, CAS 

74991):/, i, calices of syntypes, x 2.5, x 2.4, respectively; gji. lateral and calicular views of a specimen, x 1.5, x 

2.3, respectively. Balanophyllia gigas (/,/, Alb-5070, USNM 92882; k, holotype, BM 1876.10.11.23): jj. lateral 

and calicular views of a specimen, x 0.7, x 1.9, respectively; *, holotype, x 0.8.

144 


PLATE 36.—Balanophytlia ponderosa: a.b, stereo calicular and lateral views of holotype, BM 1939.7.20.62, x 

2.J. x 1.2, respectively. Balanophyllia teres (c, holotype. USNM 92888; d. Alb-4903, USNM 92889): c4, 

corallum and calice. x 2.4. x 7. respectively. Endopachys grayi: ejt, lateral and calicular views of a specimen from 

Alb-2999 (Gulf of California). USNM 19221, both x 2.0. Eguchipsammia fistula. f,g, lateral and calicular views 

of syntypes from Siboga-\05. ZMA Coel. 564. x 1, x6.4. respectively.


PLATE 37.—Eguchipsammia wellsi (a-c, g, Okinose, Sagami Bay, 110 m, USNM 92897): a, colony, X 1.1; bjc, 

calicular views, x 14, x 13. respectively; g. synapticulotheca, x 22. Eguchipsammia gaditana (d-fM, Okinose, 

Sagami Bay, 110 m, USNM 92894): d. colony, x 1.2; e-f, calicular views, x 18, x 32, respectively; h, 

synapticulotheca covered by epitheca on lower half, x 35. Endopachys grayi: i, TM (KT9015, BS1), USNM 

92866, corallum with five buds on thecal edges, x 2.7.

146 


PLATE 38.—Cladopsammia eguchii: afr, calices from colony off Shirahama, USNM 78646, x 2.1, x 2.8, 

respectively. Dendrophyllia ijimai: c,f, Moroisa, Sagami Bay, USNM 92872, colony and calices, x 0.65, x 1.9, 

respectively. Cladopsammia gracilis: d,e, off Shirahama, USNM 92870. colony and a calice, x 1.1, x 3.0, 

respectively. Dendrophyllia cribrosa: gji, larger syntype, RMNH 9212, colony and one worn corallite, x 0.43, x 

12.2, respectively. Dendrophyllia arbuscula («-/. Alb-3707. USNM 22057): ij, colonies, x 1.2, x 1.3JX calices, 

x 3.4. x 3.3. respectively.


PLATE 39.—Dendrophylliaflorulenta: a-c, TM (KT9O15, CB2-2), USNM 92877, colony and calicular views, x 

1.6, x 7.6, x 3.9, respectively. Enallopsammia rostrata (d.e, Alb-4892, USNM 92850; /. Alb-4891, USNM 

92849): d,e, acalicular and calicular sides of a colony, x 0.48; /. three calices. x 2.7. Tubastraea coccinea (g, off 

Partida Island, Gulf of California, USNM 91428; h,i, off Misake, ZMC): g. colony, xO.72; h.i, calices, both x 1.9.

148 


PLATE 40.—Schizopsammia songae (a-e, holotype, USNM 15847): a, stereo pair of colony, x 1.4; b,c, two other 

views of holotype, x 1.4, x 1.3, respectively; d,e, calices in process of intratentacular division, x 2.3, x 5.4, 

respectively. Rhizopsammia minuta muisuensis if-g, holotypic colony, TIUS 41391):/, stereo view of section of 

rcptoid colony, x 1.3; g. a calice, x 9.3. Rhizotrochus niinoi (h,i, holotype, TIUS 60820): h,i, lateral and calicular 

views, x 5.0, x 5.3. respectively.). Balanophyllia sp. A sensu Yabe and Eguchi (1942b), Soyo Maru-647, TIUS 

59163.x 5.3.


PLATE 41.—Oulangia stokesiana miltoni (ajb, syntypes, TIUS 41051): a, a well-preserved corallite, x 5.9; b, 

stereo view of corallites attached to an oyster shell, x 1.2. Caryophyllia scobinosa decapali (cM. holotype, Soyo 

Marn-210, TIUS 53640): c, corallum, x 2.9; d, stereo view of calice, x 5.3. Flabellum transversale conicum (ejt, 

syntype, TIUS 39727): e. calice, x 4.2; i, corallum, x 2.9. Goniocyathus pacificus (f.g, syntype, TIUS 50086):/, 

calice, x 3.4; g, lateral view of same corallum, x 3.2. i, stereo calicular view of holotype of Deltocyathoides 

japonicus, TIUS 50091, x 3.4.

150 


PLATE 42.—a, Paracyathus pruinosus, Soyo Maru-235, TIUS 53681, x 3.1. Fragilocyathus conotrochoides (b, 

holotype, TIUS 50084; cA. TIUS 58223): b, lateral view showing thecal stripes, x 2.2; c,d, lateral and calicular 

views of a specimen, x 3.2, x 8.0, respectively.

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