MÉMOIRES DU MUSÉUM NATIONAL D’HISTOIRE NATURELLE

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Source : MNHN, Paris

■'■■■ -

Source : MNHN, Paris

Révision of the northeast Atlantic

and Mediterranean Stylasteridae

( Cnidaria: Hydroyoa)

Source : MNHN, Paris

ISBN : 2-85655-192-X

ISSN : 0078-9747

Source : MNHN, Paris

MÉMOIRES DU MUSÉUM NATIONAL D’HISTOIRE NATURELLE

l 260 t

SÉRIE A

ZOOLOGIE

TOME 153

Helmut ZlBROmus

Station marine d’Endoume

rue Batterie des Lions

15007 Marseille, France

Stephen D. Cairns

Department of Invertebrate Zoology

Smithsonian Institution

Washington D.C. 20560, USA

Révision of the northeast Atlantic

and Mediterranean Stylasteridae

(Cnidaria: Hydro^oa)

ÉDITIONS

DU MUSÉUM

PARIS

1992

Source : MNHN, Paris

Contents

Pages

Abstract. 9

Résumé. 11

Introduction. 13

Abbreviations. 14

Material used for the révision. 15

Acknowledgements. 16

Historical review. 17

Biogeography. 19

Symbiotic associations. 21

Records of fossil stylasterids from Europe. 23

Taxonomical révision (with key to the généra of Stylasteridae from the study area). 26

Lepidopora eburnea (Calvet, 1903) (Fig. 1-2). 27

Lepidopora sp. A (Fig. 1,3-4). 31

Lepidopora sp. B (Fig. 3-4). 36

Pliobolhrus symmetricus Pourtalès, 1868 (Fig. 5-6) including Pliobothrus sp. (Fig. 7). 38

Pliobothrus gracilis n. sp. (Fig. 5, 8). 44

Errina aspera (Linnaeus, 1767) (Fig. 9-10). 46

Errina dabneyi (Pourtalès, 1871) (Fig. 11-13). 53

Errina atlantica Hickson, 1912 (Fig. 14-15). 58

Stylaster norvegicus (Gunnerus, 1768) (Fig. 16-17). 62

Stylaster rosaceus (Greeff, 1886) (Fig. 18). 69

Stylaster blatteus (Boschma, 1961) (Fig. 19-20). 72

Stylaster maroccanus n. sp. (Fig. 21-22). 76

Stylaster gemmascens (Esper, 1794) (Fig. 23-24). 79

Stylaster ibericus n. sp. (Fig. 25-26). 84

Stylaster erubescens Pourtalès, 1868. 87

Stylaster erubescens groenlandicus n. ssp. (Fig. 27-28). 89

Stylaster erubescens britannicus n. ssp. (Fig. 29-30). 92

Stylaster erubescens meteorensis n. ssp. (Fig. 31-32). 96

Stenohelia maderensis (Johnson, 1862) (Fig. 33-34). 99

Stenohelia sp. A (Fig. 35). 104

Crypthelia affinis Moseley, 1879 (Fig. 36-37). 106

Crypthelia medioatlantica n. sp. (Fig. 36, 38). 112

Crypthelia vascomarquesi n. sp. (Fig. 39-40). 114

Crypthelia tenuiseptata Caims, 1986 (Fig. 41-42). 117

Note added in press. 122

List of deep-water stations from océanographie cruises. 123

Référencés. 128

Index. 135

Source : MNHN, Paris

ABSTRACT

ZIBROWIUS H. & S. D. CAIRNS, 1992. REVISION

OF THE NORTHEAST ATLANTIC AND MEDI-

TERRANEAN STYLASTERIDAE (CNIDARIA:

HYDROZOA). Mém. Mus. natn. Hist. nat., (A), 153:

1-136. Paris isbn: 2-85653-192-X.

Published March 20th, 1992.

In the northeastern Atlantic, from the Cape Verde

Islands to the Mid-Atlantic Ridge at 23°31'N, the

Azores, Iceland and northern Scandinavia, 19 species

of stylasterids hâve been recognized, one of which is

represented by 3 subspecies. Complementary records

extend the study area to Greenland. In addition, 2

species are included from the islands of the Gulf of

Guinea, équatorial eastern Atlantic. Of ail these

species and subspecies, 20 are fully described whereas

3 that are incompletely known, are not yet named. The

new taxa are: Pliobothrus gracilis n. sp., Stylaster

maroccanus n. sp., S. ibericus n. sp., S. erubescens

groenlandicus n. ssp., S. erubescens britannicus n. ssp.,

S. erubescens meteorensis n. ssp., Crypthelia medioat-

lantica n. sp., and C. vascomarquesi n. sp.

Compared with the scleractinian corals présent in

the same areas, most of the studied stylasterid species

appear to hâve a rather narrow geographical range.

Only 3 of the 21 species (15 %) recorded from the

northeastern and équatorial eastern Atlantic are

known in the western Atlantic.

Although covering a much wider area, the northeas¬

tern Atlantic stylasterid fauna is considerably less

diversified than its West Indian counterpart, which

comprises about twice as many species in 8 généra.

Among the northeastern Atlantic régional faunas, that

of the Azores is the richest, with 9 deep-water species.

Only one species ( Errina asperà) occurs in the south-

western Mediterranean.

Although présent in the islands of the Gulf of

Guinea, shallow-water stylasterids are missing in the

northeastern Atlantic and Mediterranean where the

shallowest records are from about 100 m. A few

records exceed depths of 2000 m.

In the study area various symbionts leave characte-

ristic traces on the stylasterid skeleton or cause

modifications: the gastropod genus Pedicularia (on 8

species) and polynoid and eunicid polychaetes (each

on one species).

The reliable record of fossil stylasterids is scarce in

Europe and the Mediterranean basin (as it is elsewhe-

re). It ranges from the Lower Paleocene to the

Plio-Pleistocene whereas Pedicularia , an obligate sym-

biont of stylasterids, is known from the Messinian

(Upper Miocene) and from the Lower Pleistocene.

Source : MNHN, Paris

RÉSUMÉ

ZIBROWIUS H. & S. D. CAIRNS, 1992. REVISION

OF THE NORTHEAST ATLANTIC AND MEDI-

TERRANEAN STYLASTERIDAE (CNIDARIA:

HYDROZOA). Mém. Mus. nain. Hist. nat.. (A), 153:

1-136. Paris isbn: 2-85653-192-X.

Publié le 20 Mars 1992.

Révision des Stylasteridae de l'Atlantique nord-

oriental et de la Méditerranée. Dans l'Atlantique

nord-oriental, des îles du Cap-Vert à la dorsale

Médio-Atlantique par 23°31'N, aux Açores, à l’Islande

et au nord de la Scandinavie, 19 espèces de Stylaste¬

ridae ont été reconnues, dont une représentée par 3

sous-espèces. Des signalisations complémentaires éten¬

dent la zone étudiée jusqu'au Groenland. En plus, sont

inclues ici 2 espèces des îles du golfe de Guinée,

Atlantique équatorial oriental. De toutes ces espèces et

sous-espèces, 20 sont décrites en détail tandis que 3

autres, incomplètement connues, n’ont pas encore reçu

de nom. Les nouveaux taxa sont: Pliobolhrus gracilis

n. sp., Stylaster maroccanus n. sp., S. ibericus n. sp., S.

erubescens groenlandicus n. ssp., S. erubescens britan-

nicus n. ssp., S. erubescens meteorensis n. ssp., Crypt-

helia medioatlantica n. sp., C. vascomarquesi n. sp.

Les auteurs du 18e siècle connaissaient trois espèces:

deux en provenance des fjords de Norvège, et une

obtenue en Méditerranée par les pêcheurs du corail

rouge. Avant la fin du 19e siècle, du matériel de la

plupart des espèces avait été récolté, y compris par les

expéditions du " Challenger ”, du “ Talisman ” et du

Prince de Monaco. Mais au-delà de notes dispersées

(dont notamment celles de H. Boschma) à propos de

quelques espèces, l'ensemble de cette faune n’avait

jamais été l’objet d’une synthèse. Trois espèces avaient

même été décrites à titre de bryozoaires (“ Hornera ”).

La synthèse présentée ici s’appuie aussi sur de

nombreuses campagnes océanographiques plus récen¬

tes (y compris des prélèvements par submersibles),

telles que les campagnes biaçores, balgim, seamount

1, hydrosnake et celles de la “ Thalassa".

Pour la plupart, les espèces étudiées semblent avoir

une répartition géographique assez étroite, comparé à

celle des scléractiniaires présents dans les mêmes

zones. Seulement 3 espèces sur 21 (15 %) inventoriées

dans l’Atlantique nord-oriental et équatorial oriental

sont connues aussi dans l'Atlantique occidental. Au¬

cune de ces espèces atlantiques ne semble exister dans

les autres océans. Tout comme les scléractiniaires, les

Stylasteridae sont absents au-delà de la périphérie du

bassin arctique, ce qui contraste avec la diversité des

deux groupes dans l’océan Antarctique aux conditions

plus stables depuis des millions d'années.

La faune de Stylasteridae de l’Atlantique nord-

oriental est considérablement moins diversifiée que

celle de l'Atlantique tropical américain qui comprend

environ deux fois plus d’espèces dans 8 genres. Parmi

les faunes régionales, celle des Açores est la plus riche,

avec 9 espèces de profondeur. Une seule espèce ( Errina

aspera) vit dans la Méditerranée et y est limitée à la

partie sud-occidentale. Elle est typique de fonds ro¬

cheux exposés à de forts courants (détroit de Messine,

détroit de Gibraltar).

Des Stylasteridae littoraux ou de faible profondeur

existent aux îles du golfe de Guinée, mais sont absents

de l’Atlantique nord-oriental et de la Méditerranée où

les signalisations les moins profondes correspondent à

environ 100 m de profondeur. De rares récoltes

correspondent à des profondeurs dépassant 2000 m,

dont une par le submersible “ Nautile ” sur la dorsale

Médio-Atlantique.

Dans la zone étudiée divers symbiontes laissent des

traces caractéristiques sur le squelette des Stylasteridae

ou causent des modifications: gastropodes prosobran-

ches du genre Pedicularia (sur 8 espèces) et polychètes

Polynoidae et Eunicidae (chacun sur une espèce). Le

contour de la coquille de Pedicularia correspond

étroitement à la configuration et aux irrégularités de la

branche à l’endroit occupé. Après la disparition du

symbionte cet emplacement précis reste évident par un

dépôt de calcaire. Les galles induites par des Polynoi¬

dae, formant des galeries le long des branches et à

orifices latérales, sont exceptionnellement rares dans

l'Atlantique nord-oriental. Eunice norvegica, sym¬

bionte d’une espèce de Stylasteridae aux Açores, est le

même qu’on trouve dans des colonies de scléractiniai¬

res ( Lophelia , Madrepora, Solenosmilid). Son tube

souple d’une matière organique est recouvert de la

même façon par le sclérenchyme de l’hydrocoralliaire.

Les signalisations fiables de Stylasteridae à l’état

fossile pour l'Europe et le bassin méditerranéen sont

rares (comme elles le sont aussi ailleurs). Elles vont du

Paléocène inférieur au Plio-Pléistocène tandis que

Pedicularia, symbionte obligatoire de Stylasteridae, est

connu dans le Messinien (Miocène supérieur) et dans

le Pléistocène inférieur. Les faunes tertiaires compren¬

nent, en plus de genres encore présents dans l'Atlan¬

tique nord-oriental, des genres qui font partie de la

faune actuelle de l'Atlantique occidental et de l’Indo-

Pacifique. Errina aspera (accompagnée de Pedicularia)

vient d’être découvert dans le Plio-Pléiostocène de

Sicile et de Calabre, dans des conditions sédimentaires

analogues à celles de son milieu actuel (détroit de

Messina). Cette découverte tardive dans un secteur en

principe bien prospecté depuis plus d’un siècle,

confirme le peu d’attention porté aux Stylasteridae par

les paléontologistes.

Source : MNHN, Paris

INTRODUCTION

The stylasterids, along with the milleporids and scleractinians, are classed as “ hard ” or

“ stony ” corals, an obviously polyphyletic assemblage including various taxa in two of the three

classes of cnidarians. Stylasterids were originally considered as scleractinians and it was only by 1873

that G.O. Sars began to question this relationship. Finally, during the “ Challenger " expédition

(1873-1876) one of the naturalists aboard, H.N. Moseley, discovered that stylasterids were highly

modified hydroids and thus only distantly related to the anthozoan scleractinians. His results, based

on observation of live material carried out during the circumnavigation, were published in a sériés

of preliminary and very elaborate papers (Moseley, 1876b, 1877, 1879, 1881).

A check list of ail stylasterid taxa then known (Cairns, 1983b) included 23 généra, 1 subgenus,

184 valid species (20 of them fossil), 11 formae or faciès, 42 junior synonyms, 4 nomina nuda, and

4 unnamed “ species Subsequently the number of described species has increased by 35 (Cairns,

1985, 1986a, 1986b, 1987, 1988). Furthermore, we are aware of several new généra and many new

species yet to be described in rich collections from South Africa, the southwestern Indian Océan, New

Caledonia, and New Zealand. Notwithstanding this expected further increase in number of described

taxa, species of stylasterid corals will always remain far less numerous than species of scleractinian

corals.

Correspondingly, the literature is less abundant on stylasterids than on scleractinians, and the

list of major authors is considerably shorter for stylasterids. Extensive bibliographies compiled by

Boschma (1957a) and by Vervoort & Zibrowius (1981) document the previous literature.

This révision treats mainly the stylasterid fauna of the northeastem Atlantic and the

Mediterranean. In the northeastem Atlantic (comprising to the east the coasts of Europe and Africa,

and to the west Iceland, the Azores, seamounts Southwest and south of the Azores, the Mid-Atlantic

Ridge south to 23°31'N, and the Cape Verde Islands) we recognized 19 species, one of which is

represented by 3 subspecies. Of ail these taxa, 18 are fully described herein, whereas 3 that are known

only from poor material are not named. Only one of these Atlantic species also occurs in the

southwestern Mediterranean. For comparison, from about the same area, Zibrowius (1980) studied

85 species of scleractinians (which did not represent the entire fauna known at that date); and out

of these the Mediterranean scleractinian fauna comprises about 30 species.

In this paper we also include 2 species from the islands of the Gulf of Guinea, équatorial

eastern Atlantic (Principe, Sào Tomé). These species are separated by approximately 15° of latitude

from the nearest recorded stylasterids in the Cape Verde Islands, and by approximately 34° of latitude

from the more diversified South African stylasterid fauna. Both species from the Gulf of Guinea

occur in shallow water whereas only deep-water species exist in the Mediterranean and the

northeastem Atlantic (shallowest records from about 100 m depth).

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

ABBREVIATIONS

The following abbreviations are used in the text

— for institutions:

amnh American Muséum of Natural History, New York

bmnh The Natural History Muséum, London; formerly: British Muséum (Natural History)

imfb Institut für Meeresforschung, Nordseemuseum, Bremerhaven

irsnb Institut Royal des Sciences Naturelles de Belgique, Bruxelles

mcm Museu Carlos Machado, Ponta Delgada, Azores

mcz Muséum of Comparative Zoology, Cambridge, Mass.

mhng Muséum d’Histoire Naturelle, Genève

mnhn Muséum national d’Histoire naturelle, Paris

mzuc Museo Zoologico, Université di Catania

mzus Musée Zoologique, Université de Strasbourg

nhmw Naturhistorisches Muséum, Wien

rmnh Nationaal Natuurhistorisch Muséum, Leiden; formerly: Rijksmuseum van Natuurlijke

Historié

rsm Royal Scottish Muséum, Edinburgh

smf Natur-Museum und Forschungs-Institut Senckenberg, Frankfurt a.M.

smnh Naturhistoriska Riksmuseet, Stockholm

umzc University Muséum of Zoology, Cambridge, U.K.

usnm National Muséum of Natural History, Smithsonian Institution, Washington, D.C.

vsm Det Kgl. Norske Videnskabers Selskab Museet, Trondheim

ypm Yale Peabody Muséum, New Haven, Conn.

zma Zoologisch Muséum, Amsterdam

zmb Muséum für Naturkunde, Berlin

zmuk Zoologisk Muséum, Kobenhavns Universitet

zmuo Zoologisk Muséum, Universitet i Oslo

zsm Zoologische Staatssammlung, München

— for stylasterid morphology:

h:w height to maximum width ratio of gastrostyles

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

MATERIAL USED FOR THE REVISION

Many samples of stylasterids from various localities throughout the study area were available

for this révision. These included types and other samples referred to in the literature, previously

unpublished samples in muséum collections, abundant new material primarily from French

océanographie cruises since 1958, and a few samples received from individual collectors. For

complementary information see also Acknowledgements and List of deep-water stations from

océanographie cruises.

We thus studied stylasterids obtained by various océanographie expéditions corresponding to

more than a century of exploration in the northeastern Atlantic, starting with the “ Challenger "

circumnavigation in 1873. Material collected by submersibles is also included (“ Pisces III" on

Rockall Bank; “ Nautile " cruise hydrosnake on the Mid-Atlantic Ridge). The most diversified

collections, in term of number of species, are those from the “ Talisman " expédition in 1883, from

the Prince of Monaco expéditions (1888 to 1905), and from the “ Jean Charcot " cruise biaçores in

1971. While the greater part of the “ Talisman ” stylasterids had been published already (but was in

great need of a révision), those of the Prince of Monaco expéditions had remained unpublished,

except for three mistakenly identified as bryozoans.

Stylasterids from the northeastern Atlantic and the Mediterranean are best represented in

muséums of Europe, but we also found several samples in American muséums. A list of the

institutions (with abbreviations used in the text) housing material studied herein is provided above.

One of us (H.Z.) had access (at the rmnh, in 1978 and 1980) to the abundant collections

amassed by the late H. Boschma before they were returned to the respective institution of origin.

Among these collections originally loaned to Boschma was abundant material from Norway

belonging to the zmuo and vsm, and various samples from the Faroes to Greenland (mainly from the

“ Ingolf" and “ Dana " expéditions) belonging to the zmuk.

Other institutions that provided many important samples for our révision were the bmnh,

possessing many old samples including types; the mom, possessing the samples from the Prince of

Monaco expéditions; and the mnhn, possessing the greater part of the “ Talisman " stylasterids and

various other old samples including types. Surprisingly, there is only one lot of stylasterid from the

“ Travailleur " expéditions. In addition, the mnhm now houses the abundant material from the French

océanographie cruises in the Atlantic between 1958 and 1988 (some duplicates were given to other

institutions) and various samples originally given to H.Z.

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

ACKNOWLEDGEMENTS

Assistance from many people in the institutions listed above (see Abbreviations) is gratefully

acknowledged, their help including guidance in the collections, loan of specimens, and checking

various data. Spécial thanks are extended to W. Vervoort (retired director, rmnh), G. Behrmann

(imfb), C. Carpine (mom), P.F.S. Cornélius (bmnh), D. Kühlmann (zmb), and M.E. Christiansen

(ZMUO).

Part of the “ Thalassa ”, “ Cryos ” (cruise balgim), and “ Noroit ” (cruise seamount 1)

stylasterids, and the “ Nautile ” (cruise hydrosnake) stylasterids were kindly provided by M.

Segonzac (Centre de Tri d’Océanographie Biologique, Brest). We further want to thank S. Gofas

and J. Laborel for specimens collected by diving in the Gulf of Guinea; L. Saldanha for

transmitting samples from the Gulf of Guinea and from off the Azores; J. Stirn for material from

Morocco; A. Freiwald for material from Denmark Strait; J.B. Wilson for material from Rockall

and Anton Dohm Seamount; T.P. Scoffin for material from Porcupine Bank; C. Alvarez-Claudio

for material from the south of the Bay of Biscay; G. Bellan for material from Hyères Seamount; P.

Colantoni, I. di Geronimo and G. Fredj for specimens from the Straits of Messina and

corresponding information; P. Barrier for the rich fossil stylasterid fauna from Carboneras,

southeastem Spain; F. Kézirian for fossil stylasterids and Pedicularia from Sicily; G. Hartmann-

Schrôder for identification of symbiotic polychaetes; and P.M. Arnaud for information on the

symbiotic gastropod genus Pedicularia.

H.Z. also acknowledges the opportunity of taking part in cruises that permitted the collection

of stylasterids: “ Jean Charcot ” cruise biaçores in 1971 (chief scientist J. Forest), “ Thalassa ” in

1972 and 1973 (chief scientist L. Cabioch), “ Cryos ” cruise balgim in 1984 and “ Noroit ” cruise

seamount 1 in 1987 (chief scientist P. Bouchet).

The scanning électron micrographs were taken by S.D.C. in the s.e.m. Laboratory of the

National Muséum of Natural History, Smithsonian Institution.

Source : MNHN , Paris

HISTORICAL REVIEW

The first stylasterid species to be described from the study area was Errina aspera (Linnaeus,

1767), from the Mediterranean (also reported, by error, from Norway). Next reported were Stylaster

norvegicus (Gunnerus, 1768) from Norway, and Stylaster gemmascens (Esper, 1794), most likely also

from Norway although its origin was given as the Indian Océan.

Allopora oculina Ehrenberg, 1834, was reported from an unknown origin, but most likely was

Stylaster norvegicus from Norway; the name A. oculina was later used again by Norwegian authors

for S. norvegicus.

In their sériés of “ monographies ” and “ histoire naturelle des coralliaires ”, in which Milne

Edwards & H aime (1848-1860) gave a rather complété inventory of the scleractinians then known

from the Mediterranean and the northeastem Atlantic, they were less successful with the stylasterids,

which at that time were still included in the scleractinian family Oculinidae. From their own

expérience the authors knew Stylaster gemmascens and Allopora oculina (probably identical with

Stylaster norvegicus) and described these in some detail, but they mentioned Stylaster norvegicus

(under Allopora norvegica) as a doubtful species and overlooked Errina aspera (Milne Edwards &

Haime, 1850, 1857).

Stenohelia maderensis (Johnson, 1862), originally described from Madeira, was later reported

also from the Cape Verde Islands (Sa ville Kent, 1872; Greeff, 1884).

Next to be reported, from the Faroes — Hébrides area (Duncan, 1870, 1873), was Pliobothrus

symmetricus Pourtalès, 1868, a species originally described from the western Atlantic.

Errina dabneyi (Pourtalès, 1871) was the first species to be described from the Azores, an area

from which the stylasterid fauna subsequently was found to be more diversified than in the other

areas investigated.

Crypthelia pudica Milne Edwards & Haime, 1849, originally described from the Philippines,

was mistakenly reported by Moseley (1879, 1881) from south of the Canary Islands (“ Challenger ”

expédition), and by Filhol (1885) without indication of the locality from the “ Talisman ” expédition.

In fact, the “ Challenger ” stylasterid was Crypthelia afjinis Moseley, 1879, which had been figured

and confusedly named (but not described) under that name, whereas the “ Talisman ” stylasterid was

most likely Crypthelia tenuiseptata Cairns, 1986.

Of the two species reported by Greeff (1884) from the islands of the Gulf of Guinea, one was

described as new, Stylaster rosaceus (Greeff, 1884), whereas the other was misidentified as the South

African Stylaster subviolaceus (Saville Kent, 1871) and was much later recognized as a distinct species,

Stylaster blatteus (Boschma, 1961).

Another exotic species mistakenly reported by Thornely (1897) was the southeastem Pacific

Stylaster sanguineus Milne Edwards & Haime, 1850, which is, in fact, Stylaster gemmascens (Esper,

1794) from Rockall Bank.

Lepidopora eburnea (Calvet, 1903) from the Azores was originally described as a bryozoan of

the genus Hornera (independently described as Lepidopora hicksoni Boschma, 1963). In addition,

Calvet (1903, 1911) redescribed Errina dabneyi and Pliobothrus symmetricus (see above) as

bryozoans: Hornera verrucosa and Hornera gravieri, respectively. Two more stylasterid species

{Crypthelia) hâve now been discovered in the Prince of Monaco collection, which heretofore had not

been the subject of a synthesis.

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Errina atlantica Hickson, 1912, was described long after the material had been collected by the

“ Talisman ” expédition in 1883. In the same paper (Hickson, 1912b) the Philippine Stenohelia tiliata

Hickson & England, 1905, was mistakenly reported from the Cape Verde Islands (a confusion with

S. maderensis) and the western Atlantic Pliobothrus tubulatus (Pourtalès, 1867) mistakenly reported

from the Azores (a confusion with Lepidopora eburnea). Altogether Hickson’s (1912b) report on (part

of) the stylasterids of the “ Talisman ” expédition included 6 species (2 of them misidentified) from

the Cape Verde Islands and the Azores.

Broch’s (1914a) report on the stylasterids from the “ Ingolf" expédition was more exemplary

and included 4 species (one misidentified) from high latitudes. He mistakenly reported the West

Indian Sty/aster roseus (Pallas, 1766) from the Greenland - Iceland area; this identification was

corrected by Boschma (1955b, 1965c) as Stylaster erubescens Pourtalès, 1868, the latter species being

previously known only from the western Atlantic. In this paper the North Atlantic S. erubescens

sensu Boschma is considered as a distinct subspecies, S. erubescens groenlandicus n. ssp., differing

from the nominotypical West Atlantic 5. erubescens erubescens (the latter redescribed by Cairns,

1986a).

Starting in 1951, Boschma published original data on a greater number of stylasterid species

worldwide than any other author before (see Vervoort & Zibrowius, 1981), including on species

from the Mediterranean, the northeastern Atlantic, and the Gulf of Guinea. The analysis of a register

and the collections (comprising material sent to him on loan from various muséums) found at the

rmnh after Boschma's death shows that he had access to 12 out of the 23 species and subspecies

revised here. Unfortunately Boschma’s observations had not been the subject of a synthesis and his

published results on this fauna are dispersed throughout many smaller notes.

In the original description of Crypthelia tenuiseptata Cairns, 1986, which was based on western

Atlantic material, the distribution was given as amphiatlantic, including the Azores, Hyères

Seamount, and Madeira. This indication is here corrected: in the eastern Atlantic C. tenuiseptata is

known only from the Azores, whereas the Crypthelia from Hyères Seamount and Madeira is C.

vascomarquesi n. sp. (it also occurs in the Azores). The other new species and subspecies described

here are Pliobothrus gracilis n. sp. from Hyères Seamount, Stylaster maroccanus n. sp. from off the

Atlantic coast of Morocco, Stylaster ibericus n. sp. from off northwest Spain, Stylaster erubescens

groenlandicus n. ssp. from south and east of Greenland to northwest and southeast of Iceland, 5.

erubescens britannicus n. ssp. from southeast of Iceland to the Celtic Sea, S. erubescens meteorensis

n. ssp. from the Great Meteor Seamount and Southwest of the Azores (imprécise locality), and

Crypthelia medioatlantica n. sp. from the Azores and the Mid-Atlantic Ridge south of the Azores.

In addition, a flattened form of Lepidopora from the Canary Islands collected by the

“ Challenger ” in 1873 (previously unpublished and considered as Lepidopora sp. B herein) definitely

is a distinct species, whereas a clavate form of Lepidopora from off Mauritania collected by the

“ Talisman ” in 1883 (previously identified as L. eburnea and considered as Lepidopora sp. A herein),

remains problematical, but has at least a look distinct of closely related typical L. eburnea. There is

also a Stenohelia from the Azores, unfortunately represented only by a small colony, which differs

from S. maderensis (considered as Stenohelia sp. A herein).

Although most of the 23 species and subspecies studied here had already been collected before

the end of the 19th century (3 of which had even been named as early as in the 18th century), the

présent révision is the first attempt of a représentative faunistic study for this area.

Source : MNHN, Paris

BIOGEOGRAPHY

In our analysis of the distribution patterns and affinities of the northeastern Atlantic and

Mediterranean stylasterid fauna, the Scleractinia are used as the référencé group. These other

“ hard ” or “ stony ” corals are well known in the investigated area (Zibrowius, 1980) where they are

represented by considerably more species than the stylasterids.

The northern limit in the Atlantic of stylasterids ( Stylaster rtorvegicus, S. gemmascens, S.

erubescens) roughly coincides with that of the scleractinians. Both groups attain northern Norway,

the Faroes, Iceland, and Southern Greenland (Denmark Strait), but are absent from the Arctic basin.

This is in contrast to the presence of numerous species of both groups in Antarctic waters (Cairns,

1982, 1983a) where conditions favorable to benthic life were not largely disrupted during the

Pleistocene.

The Mediterranean has only one stylasterid species ( Errina aspera), which equates to a much

lower proportion of the Atlantic species than the approximately 30 species of scleractinians in

common to the Mediterranean and the northeastern Atlantic. However, the greater part of the

scleractinian species living in the Mediterranean occur in shallow water, whereas the stylasterids from

corresponding latitudes in the Atlantic are deep-water species. In terms of number, the shallow-water

scleractinians are more successful in the présent Mediterranean than the deep-water species, but the

latter were more diversified during the Pleistocene when the Mediterranean deep hydrology was more

similar to that of the présent northeastern Atlantic (Zibrowius, 1980; Barrier et al., 1989). The

deep-water stylasterids may also hâve been more successful in the Pleistocene Mediterranean, but thus

far the fossil records are missing.

Errina aspera is the only stylasterid species living in the Mediterranean, where it appears to

be limited to the southwestern part (including the Straits of Messina); it thereby shows a common

distribution pattern with some scleractinians.

Concerning rarely collected stylasterids, it may still be prématuré to compare distribution

patterns. Nevertheless, some trends appear sufficiently évident from the available data.

Few of the 23 stylasterid species and subspecies recorded from the northeastern and équatorial

eastern Atlantic hâve a wide latitudinal range, the exceptions (group 1) being: Stenohelia maderensis

from the Cape Verde Islands through Madeira, Galicia Seamount, and the Bay of Biscay to the

Faroes Channel (range 45° of latitude; the northernmost occurrence needing confirmation by new

records); Stylaster erubescens with three subspecies from Great Meteor Seamount through the Celtic

Sea and the Faroes to Iceland and Greenland, Denmark Strait (range 36° of latitude); Pliobothrus

symmetricus from Madeira and the Azores through the Celtic Sea and the Faroes to Southwest of

Iceland (range 31° of latitude); Errina aspera from the Cape Verde Islands through the Straits of

Gibraltar to the Straits of Messina (range 23° of latitude); and Lepidopora sp. A from Mauritania

through Morocco to Galicia Seamount (range 23° of latitude).

Although they are proportionally rather widespread (compared with group 3 below), some

other species (group 2) appear to hâve a considerably narrower range in latitude. Stylaster norvegicus

and S. gemmascens are known to occur only in high latitudes, from Norway to Iceland and

Greenland, respectively, but apparently do not extend south beyond Rockall Bank (at about 58°N).

Crypthelia vascomarquesi is known from three distant areas (Hyères Seamount, Azores, Madeira

Archipelago) that are not widely separated in latitude (range only 7°); this species is perhaps more

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

widely destributed north and south of Madeira (poorly known représentatives of Crypthelia from

Selvagens Archipelago and Joséphine Seamount).

Available data suggest that the remaining species (group 3) occur in a more limited area.

Several species are thus far known from the Azores and in part from the Mid-Atlantic ridge to the

Southwest: Lepidopora eburnea, Errina dabneyi, E. atlantica, Stenohelia sp. A, Crypthelia affinis, C.

medioatlantica, and C. tenuiseptata. Pending additional records and further investigations, forms

similar to Lepidopora eburnea and Errina atlantica from seamounts between Portugal and Madeira

are not formally referred to the Azorean species. Crypthelia affinis is provisionally included here; in

fact its type locality (given as far Southwest of the Canary Islands with the unusual depth of 2790 m)

may be incorrect and thus the species is positively known only from the Azores. Pliobothrus gracilis

was obtained only from Hyères Seamount, Lepidopora sp. B from one station off the Canary Islands,

Stylaster maroccanus only once off Morocco, and S. ibericus exclusively northwest of Spain.

The shallow-water species Stylaster rosaceus and S. blatteus from the islands of the Gulf of

Guinea hâve not been found elsewhere.

With 9 species ( Lepidopora eburnea, Pliobothrus symmetricus, Errina dabneyi, E. atlantica,

Stenohelia maderensis, Crypthelia affinis, C. medioatlantica, C. vascomarquesi, C. tenuiseptata), the

fauna of the Azores is the most diversified. For comparison, the stylasterid faunas of the Cape Verde

Islands, the Canary Islands, and the Madeira Archipelago appear poor with one to three species thus

far recorded. The situation is similar concerning the deep-water scleractinian fauna of these

archipelagos (Zibrowius, 1980). In part this impression may be due to a more intense investigation

of the Azores, notably by the “ Talisman ”, the Prince of Monaco expéditions, and “ Jean Charcot ”

cruise biaçores.

Three of the 21 stylasterid species (15%) recorded from the northeastern and équatorial

eastern Atlantic also occur in the western Atlantic (Cairns, 1986a). Among these, Pliobothrus

symmetricus and Stylaster erubescens (with distinct subspecies) range from the West Indies to the

eastern margin of the Atlantic and attain high latitudes, whereas Crypthelia tenuiseptata is known

only from the West Indies and the Azores. None of the specifically northeastern Atlantic stylasterids

is known from any other area. Furthermore, there are no species in common with the South African

fauna or that of the Indo-Pacific. Proportionally more deep-water scleractinians hâve amphiatlantic

distributions (Cairns, 1979; Zibrowius, 1980), and some species hâve even wider géographie ranges

extending into the Indo-Pacific.

With 21 species in 6 généra, the stylasterid fauna of the northeastern Atlantic (to which are

herein annexed the 2 species of the Gulf of Guinea) is considerably less diversified than the

geographically nearest stylasterid fauna of the West Indies, which comprises 42 species in 8 généra

(Cairns, 1986a). The généra Lepidotheca and Distichopora are not represented in the northeastern

Atlantic. In the northwestern Atlantic no stylasterids are thus far recorded from between North

Carolina and Greenland, whereas in the northeastern Atlantic no similar gap of recorded distribution

exists.

The West Indian stylasterid fauna has only one shallow-water species ( Stylaster roseus );

likewise, shallow-water species occur in the eastern Atlantic in the Gulf of Guinea ( Stylaster rosaceus,

S. blatteus). However, shallow-water stylasterids are not confined to tropical areas; some do exist on

the coast of South Africa and in the fjords of New Zealand. A preliminary study (H.Z.) of the South

African stylasterid fauna, largely based on the collections of the South African Muséum, Cape Town,

shows that, in a smaller geographical area this fauna is about as diversified in term of number of

species as the northeastern Atlantic stylasterid fauna.

Source : MNHN, Paris

S YM BIOTIC ASSOCIATIONS

Attention has been focused by Zibrowius (1981) on the diversity of symbiotic associations

involving stylasterid corals and other organisms. That preliminary inventory, which was worldwide

and based on records from the literature and on new observations, listed the following symbionts

adapted to life on a stylasterid host (simple epibionts on dead parts of skeleton excluded):

— a nemertean coiled around tops of branches and causing an unusual structure of the

irritated zone;

— a species of Polydora (spionid polychaete) perforating living branches;

— a species of Autolytus (syllid polychaete) living in blister-like galls;

— various species of polynoid polychaetes causing gallery-like gall-tubes with latéral openings

along stems and branches (a particular new case reported by Cairns, 1987);

— gastropods of the prosobranch genus Pedicularia having their shell contour adapted to the

précisé place where they settled on the coral branch;

— tiny pycnogonid larvae found inside gastrozooids (larger pycnogonid larvae inside the

gastropore tube hâve been found subsequently by H.Z. on material from New Zealand);

— siphonostomatoid copepods which cause galls each of which envelop a cyclosystem;

— a thoracic cirriped largely overgrown by its host coral;

— an acrothoracic cirriped perforating live branches; and

— aplacophoran molluscs coiled around stylasterid branches or inside polynoid gall-tubes

hâve recently been reported by Cairns (1986b, 1987).

When the northeastern Atlantic and Mediterranean stylasterid fauna was investigated in

detail, symbiotic associations were especially noted and three types were found, one of them here

reported for the first time.

1) Eunice norvegica (Linnaeus, 1767) (identified by G. Hartmann-Schrôder), an eunicid

polychaete, lives in colonies of Errina atlantica from the Azores. The rather large worm (several cm

long) builds its own tube of a soft organic material which, providing an additional substrate for the

coral, is subsequently covered by the coenosteum. The worm tube thus influences the shape of the

colony by inducing growth along a preferential axis. Covered by the coral skeleton, the tube becomes

équivalent to a strong trunk of the colony (Fig. 14 A).

Previously, Eunice norvegica was known as a symbiont of three colonial deep-water

scleractinian corals (Zibrowius, 1980), ail of which cover the worm tube as described above for the

stylasterid: Madrepora oculata Linnaeus, 1758; Lophelia pertusa (Linnaeus, 1758); and Solenosmilia

variabilis Duncan, 1873.

2) Although polynoid polychaetes are probably the most widely distributed type of stylasterid

symbiont, they are poorly represented in the northeastern Atlantic stylasterids. Only one colony of

Stenohelia sp. A from an unknown station in the Azores shows an irregular growth caused by the

presence of a gall-tube inhabited by Harmothoe sp. (identified by G. Hartmann-Schrôder; Fig. 35

G). The infested colony is not flabellate as normal Stenohelia colonies. For comparison, in the

western Atlantic several species of stylasterids hâve a polynoid symbiont (Cairns, 1986a). In the case

of the polynoid — stylasterid association, the polychaete does not produce its own tube of organic

material which is subsequently covered by the coral. Instead, the gall-tube is entirely an induced

production of the coral.

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

3) The prosobranch gastropod Pedicularia causes a characteristic, very localized modification

of the stylasterid branch surface. It deposits a layer of lime under which the sealed off skeleton cannot

grow in thickness, unlike the surrounding skeletal areas. This may resuit in a slight dépréssion, the

contour of which fits that of the shell. Generally the crust of lime deposited by the snail (apparently

by its foot) comprises conspicuous prominent crests which considerably enlarge the crust surface and

on which the foot of the snail adhères more effïciently. These elongate subelliptical or more irregular

Pedicularia traces (Fig. 1 C, 2 C, 5 C, 9 G, 11 H, 14 B, 25 A, D, E, 33 C) are easily recognizable

and provide information on the occurrence of the snail, even when the latter had been lost, or when

the stylasterid was already dead when collected.

Pedicularia gastropods are the most common symbionts of the northeastern Atlantic and

Mediterranean stylasterids. Their characteristic traces, if not the molluscs themselves, hâve been

found on 8 species and subspecies: Lepidopora eburnea, Pliobothrus symmetricus, Errina aspera, E.

dabneyi, E. allantica, Sty las ter ibericus, S. erubescens britannicus, and Stenohelia maderensis. In the

investigated area the Pedicularia — stylasterid association has thus been recognized in the Azores,

Madeira Archipelago, Canary Islands (only isolated disassociated Pedicularia collected here). Cape

Verde Islands, Celtic Sea, off northwestern Spain, Atlantic coast of Morocco, Straits of Gibraltar,

and Straits of Messina. Indirect records of Pedicularia (traces on Pliobothrus symmetricus and

Stylaster erubescens britannicus) from the Celtic Sea at 48°37.0'N to 48°38.2'N are the northernmost

ones known worldwide.

Pedicularia from the northeastern Atlantic area hâve been reported in the malacological

literature (Bellon-Humbert & Gofas, 1977; Dautzenberg, 1889, 1927; Locard, 1897; etc.) under

various spécifie names: P. sicula Swainson, 1840; P. decussata Gould, 1855; P. decurvata Locard,

1897; and P. sicula var. sublaevigata Locard, 1897. Bouchet & Waren (1992) synonymize ail these

under P. sicula.

The incidence of Pedicularia on the northeastern Atlantic and Mediterranean stylasterid fauna

is considerably higher than on that of the western Atlantic. In the western Atlantic, Pedicularia traces

are rare (not mentioned by Cairns, 1986a) and are known only from 4 species: Stylaster complanatus

Pourtalès, 1867, from Cuba near Havana; Stylaster erubescens Pourtalès, 1867, from the Blake

Plateau; Conopora sp. from northwestern Brazil; and Crypthelia peircei Pourtalès, 1867, from

Guadeloupe and St. Vincent. Apparently no specimen of western Atlantic Pedicularia has yet been

seen on its host coral.

Source : MNHN, Paris

RECORDS OF FOSSIL STYLASTERIDS FROM EUROPE

Records of fossil stylasterids are worldwide but comparatively rare. In emerged areas

neighbouring the Mediterranean and the northeastern Atlantic, fossil stylasterids are thus far known

only from Europe.

The geologically oldest record, from the Maastrichtian (uppermost Cretaceous) of Denmark,

is only briefly mentioned by Floris (1979) in a paper dealing mainly with scleractinians. Alluding to

previous records from the Danian (lowermost Paleocene, see below), Floris indicated that “ rare

finds of stylasterine hâve now also been made in the Maastrichtian however, these stylasterids from

the Maastrichtian are not even tentatively referred to généra.

Nielsen (1919) distinguished 8 new species from the Danian of Fakse (= Faxe), Denmark,

originally considered as uppermost Cretaceous but now known to be Lower Paleocene: Sporadopora

faxensis, Pliobothrus dispergens, P. laevis, Spinipora irregularis, Labiopora lobata, Congregopora

nasiformis, Astylus crassus, and Conopora arborescens. Boschma (1951a) reproduced the description

and figures of Congregopora nasiformis and discussed the affinities of this form. Nielsen’s 8 species,

with subséquent citations by other authors (no original new data) are listed by Boschma (1957a),

who also transferred 3 of them into different généra: Spinipora irregularis and Labiopora lobata into

the genus Errina, Astylus crassus into the genus Astya. Cairns (1983b) followed Boschma, except for

transferring Labiopora lobata into the genus Errinopora (not Errina) and considered most species

(except Errina irregularis , formerly under Spinipora) as of uncertain generic placement, or incertae

sedis. Some of Nielsen’s species hâve been reported again from Fakse by Bernecker & Weidlich

(1990).

Distichopora antiqua Defrance, 1826, from the Eocene of the Paris basin (list of references in

Boschma, 1957a) is a typical représentative of the genus Distichopora (specimen at mnhn labelled

“ calcaire grossier de Chaumont ”).

Allopora compressa (Rômer, 1863), as revised by Boschma (1951b) (detailed synonymy also in

Boschma, 1957a), has been described from the Oligocène of Lattorf near Kassel, Germany, under

various names: Dendracis compressa Rômer, 1863; D. multipora Rômer, 1863; D. pygmaea Rômer,

1863; D. tuberculata Rômer, 1863; and Cryptaxis alloporoides Reuss, 1865. Cairns (1983b) included

this species in Stylaster (Group A).

Stylaster priscus Reuss, 1872, has been described and figured in some detail from material from

the Miocene of Porzteich near Nikolsburg (now Mikulov), Moravia, Central Czecho-Slovakia.

Boschma (1951b, 1957a) listed it as Allopora prisca. Cairns (1983b) included this species in Stylaster

(Group B).

Stylaster antiquus Michelotti in Sismonda, 1871, was summarily described (not figured) from

material from Sassello near Savona, Liguria, northwestern Italy Originally referred to the Lower

Miocene, it has later been considered as from the Oligocène. This species was also mentioned by De

Angelis (1895) and Boschma (1951a, 1957a). Cairns (1983b) included this species in Stylaster

(Group C).

Gignoux (1913: 650) mentioned Distichopora sp. from what he considered as upper Pliocène

(now identified as Lower Pleistocene) at Musalà near Villa San Giovanni, Calabria, Southern Italy.

Mirigliano (1949) also mentioned Distichopora sp. in the species list of a Plio-Pleistocene fauna from

the province of Salerno, Southern Italy. No further information was provided, and the corresponding

Source : MNHN, Paris

HELMUT ZIBROW1US & STEPHEN D. CAIRNS

material could not be located at Naples (university and muséum) where Mirigliano carried out his

study (I. di Geronimo, in litt. 1979).

Heretofore unreported fossil stylasterids include the following new géographie records from

the Mediterranean basin.

A remarkably rich fauna (in fact the most diversified assemblage of fossil stylasterids presently

known) was discovered in 1989 by P. Barrier in the “ red brecchia ” of Messinian âge (Upper

Miocene) at Carboneras, southeastern Spain (Barrier et al., 1992). It comprises at least 14 species

in at least 8 généra: Calyptopora, Conopora, Crypthelia, Distichopora, Lepidopora, Pliobothrus,

Stenohelia, Stylaster (Groups A, C, sensu Cairns, 1983b). In addition, one shell of the gastropod

Pedicularia, an obligate symbiont of stylasterids, was found with that fauna; this is the oldest record

of Pedicularia known.

Ironically, Errina aspera, the only stylasterid species now living in the Mediterranean and

occurring abundantly in the Straits of Messina, had previously not been reported as a fossil from this

area rich in Plio-Pleistocene fossiliferous deposits. According to Montenat & Barrier (1985) and

Barrier, di Geronimo & Montenat (1988), a narrow passageway similar to the présent Straits of

Messina in canalizing strong tidal currents, already existed in the Plio-Pleistocene. Most likely, idéal

conditions for E. aspera occurred there in the past as they do at présent.

Although E. aspera was heretofore missing from the faunal lists of the Plio-Pleistocene, the

obligate symbiont of stylasterids, Pedicularia, was already known as a fossil from Sicily: Seguenza

(1865) described P. deshayesiana from what he considered as Miocene (in fact, Plio-Pleistocene) of

Rometta near Messina. A few years ago, a second fossil specimen of Pedicularia was obtained by I.

di Geronimo (pers. comm.) from the Lower Pleistocene near Capo deU’Armi, Calabria (locality

Canale).

Specially searched for, E. aspera has finally been obtained (in 1990; most specimens by F.

Kézirian) on both sides of the Straits of Messina: in Calabria from the Lower Pleistocene near Capo

dell’Armi (same locality Canale as for Pedicularia-, see above); in Sicily (province of Messina) from

the Upper Pliocène at Salice, and from the Lower Pleistocene at Giampilieri and at Casino di Falcone

(together with Pedicularia). In addition, when studying fossil scleractinians (H.Z.) from Sicily

described by Seguenza (1864) and donated by him to the Wien muséum, one small colony of E.

aspera was recently discovered attached to a fragment of Lophelia defrancei sensu Seguenza (nhmw

1864/XXI/242). This sample, with Seguenza's original label indicating “ dintorni di Messina”,

should be considered as of imprécise origin (Messina area) and of Plio-Pleistocene âge although a

later muséum label refers it to Seguenza’s “ Miocene ” of St. Filippo near Messina.

The Lower Pleistocene deposit at Casino di Falcone (Sicily; see above) also contains a second

species of stylasterid of which only a tiny fragment is presently known: it is a typical Stenohelia,

probably referable to S. maderensis, a species still living in the north-eastern Atlantic.

The records of fossil stylasterids from Europe are too scarce to permit an analysis of how the

stylasterid fauna of European seas evolved since the late Cretaceous. The generic attribution of most

of the Lower Paleocene stylasterids from Denmark remains questionable due to incomplète

conservation of essential structures. Two of the généra possibly represented in the Fakse fauna do still

exist in European seas: Pliobothrus and Errina. A few représentatives of the large genus Stylaster

(Groups A, B, C, sensu Cairns, 1983b) are well documented in the Oligocène and Miocene of central

Germany, Central Czecho-Slovakia (Moravia), northwestern Italy, and southeastern Spain. Related

forms of Stylaster (Groups A, B) still live in European seas.

In addition to généra still represented in Europeans seas ( Crypthelia, Lepidopora, Pliobothrus,

Stenohelia, Stylaster), the rich Messinian fauna of Carboneras comprises généra that no longer occur

there. Calyptopora is known only in the Indo-Pacific and Conopora is widespread throughout the

Indo-Pacific and in Antarctic and Sub-Antarctic waters, the nearest location to Europe being off

Brazil.

The genus Distichopora deserves spécial attention. In the Indo-Pacific it is now represented by

several shallow- and deep-water species (Boschma, 1959), and in the western Atlantic (Caribbean)

by several deep-water species (Cairns, 1986a). It is absent from the northeastem Atlantic and

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

adjacent waters. The Eocene D. antiqua from the Paris basin and the unnamed Distichopora from the

Messinian of Carboneras are authentic Distichopora, whereas we may doubt the correct generic

identification of Gignoux’s (1913) and Mirigliano’s (1949) Distichopora sp. from the Plio-

Pleistocene of Southern Italy (specimens unavailable). Although Distichopora is easily recognizable

among ail other stylasterids, Gignoux and Mirigliano were surely not experts on this group. A

confusion with morphologically convergent bryozoan colonies is most probable, especially since

“ transphyletic ” confusions hâve happened to trained bryozoan and coral workers (see Zibrowius,

1982, and Cairns, 1983b: 441, for examples).

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

TAXONOMICAL REVISION

The 21 species included in this révision belong to 6 généra. Because Cairns (1983b) has already

published a generic révision, general information on these généra is kept to minimum.

The présentation of each species (and subspecies) entails the following arrangement: valid name

followed by author and date, synonymy and chresonymy (see Smith & Smith, 1972), types, material

studied, description, comparisons, remarks, distribution and ecology, and symbionts. Under

“ types ”, information is provided on ail nominal species subsequently synonymized. Under

“ material studied ” only cursory data are given for deep-water stations from océanographie

expéditions; detailed station data for ail expéditions are compiled in a general station list. The passage

“ remarks ” includes historical comments (see also under types) and rectification of misidentifications

in the previous literature.

The morphological terminology used in the descriptions is that reviewed, augmented and illustrated

by Cairns (1983b, 1985, 1986a, 1986b); however an additional new term is introduced here. A binary

ampulla is a large female ampulla, usually elongate in shape, with two efferent pores on opposite

vertices and, presumably, two planulae within (see Stenohelia maderensis).

Key to the généra of Stylasteridae from the study area

la Gastro- and dactylopores not arranged in cyclosystems. 2

lb Gastro- and dactylopores arranged in cyclosystems. 4

2a Gastropore without gastrostyle. Pliobothrus

2b Gastropore with gastrostyle. 3

3a Dactylopores are (low) apically perforate cônes. Lepidopora

3b Dactylopores are U-shaped spines with slit. Errina

4a Cyclosystem (partially) covered by lid;

gastropore tube double-chambered;

gastro- and dactylostyles absent. Crypthelia

4b Cyclosystem without lid;

gastropore tube cylindrical;

gastro- and dactylostyles présent. 5

5a Cyclosystems unifacially arranged;

gastropore tube long and curved;

ampullae usually clustered near cyclosystems. Stenohelia

5b Cyclosystems randomly or sympodially arranged;

gastropore tube (usually) short and (nearly) straight;

ampullae scattered randomly over coenosteum. Stylaster

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Genus LEPIDOPORA Pourtalès, 1871

Diagnosis. — Coordination of gastro- and dactylopores usually random; however, in some

species dactylopores serially arranged on branch edges, and gastropores serially arranged on anterior

or antérolatéral branch faces. Coenosteal texture quite variable. Gastropores often bordered by

proximal lip; gastro- and dactylopore tubes long. Gastrostyles usually not ridged; H:W ratio high.

Dactylopores apically perforate mounds; no dactylostyles.

Type species: Errina glabra Pourtalès, 1867, from the western Atlantic.

Lepidopora was formerly considered as a subgenus of Errina Gray, 1835. Lepidopora is

represented in the study area by one well known species from the Azores, L. eburnea (Calvet, 1903),

and two unnamed forms. Under Lepidopora sp. A, we provisionally group problematical material (in

part poorly preserved) close to L. eburnea, from Mauritania, Morocco, and Seine, Gorringe and

Galicia Seamounts. In contrast, Lepidopora sp. B, known only from one well preserved but

incomplète specimen from the Canary Islands, definitely is a distinct species.

Lepidopora eburnea (Calvet, 1903)

Fig. 1 A-F, 2 A-J

Synonymy:

Hornera eburnea Calvet, 1903: 162, pl. 18, fig. 5a-c.

Errina ( Lepidopora ) hicksoni Boschma, 1963a: 339-342, text-fig. 1, pl. 1, fig. 1-3.

Chresonymy:

Hornera eburnea — Calvet, 1906: 479 (part, not “ Talisman ” drag. 96, Mauritania); 1931: 45. — Borg,

1944: 203. — Belloc, 1960: 12.

Errina ( Lepidopora ) eburnea — Zibrowius, 1981: 982 (part, NOT “ Talisman ” drag. 96, Mauritania).

Errina ( Lepidopora ) hicksoni — Boschma, 1963b: 395-396; 1964a: 61; 1967: 335-336; 1968c: 207. —

Vervoort & Zibrowius, 1981: 27.

Lepidopora hicksoni — Cairns, 1983b: 428.

Pliobothrus tubulatus — Hickson, 1912b: 465, pl. 8, upper fig.

Types

Hornera eburnea : In the original description, Calvet (1903) mentioned “ a beautiful sample ”

from the Azores (Prince of Monaco stn 229), apparently the figured colony (40 mm high, 35 mm

wide). Being the only specimen mentioned, it must be considered the holotype. Its depository is

unknown. The mom possesses 3 smaller topotypic specimens (stn 229) not specially mentioned by

Calvet (lower part of 2 colonies and 1 slender distal branch; part of them figured herein, Fig. 1 D-E).

Type locality: Prince of Monaco stn 229, 16.8.1888, 38°22'N, 28°14'24"W, 736 m. Azores.

Errina ( Lepidopora ) hicksoni : In the original description, Boschma (1963a) designated the

holotype and a paratype: as holotype the colony figured by Hickson (1912b, pl. 8, upper figure with

the explanation “ Pliobothrus tubulatus, Azores, 56 meters ”), and as paratype a colony figured in his

own paper (pl. 1, fig. 3), also part of the material previously referred to P. tubulatus. The holotype

was known to Boschma only from Hickson’s paper and was presumed to be deposited at the mnhn;

in fact, this specimen had been retained by Hickson and later transferred to the bmnh (1964.9.17.11).

The specimen designated paratype had also been retained by Hickson and deposited at the

Manchester muséum (from where Boschma had it on loan and to where he returned it in 1969,

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Fig. 1. — Lepidopora eburnea (A-C, from “ Talisman ” drag. 123, mnhn; D-E, topotypes of H ornera eburnea , mom; F, from

“ Jean Charcot ” 1971. stn 161, mnhn): A, colony ( * 3.0); B, detail of A showing gastro- and dactylopore arrangement,

gastropore lips ( x 18); C, detail of A, opposite face showing Pedicularia trace ( x 5.9); D, colony ( x 3.1); E, slender

colony with bulging female ampullae ( x 4.6); F, colony ( x 3.3).

Lepidopora sp. A, branched specimens close to L. eburnea (G-H, from " N or oit " dw-21, mnhn; I-K, from

“ Noroit ” dw-78, mnhn): G, branched distal fragment showing gastro- and dactylopore arrangement ( x 3.8); H, nearly

complété bifurcate colony showing bulging ampullae (x 4.1); I, colony bifurcate in upper part (x 4.1); J, repeatedly

branched specimen showing gastro- and dactylopore arrangement and bulging ampullae (x 4.1); K, branched distal

fragment, like J (x 4.1).

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTER1DAE

according to his annotations preserved at the rmnh). Additional topotypic specimens are at the bmnh

(1977.8.2.1, 3 colonies) and at the mnhn (10 colonies, branches and minor fragments).

Type locality: Given by Boschma (1963a) as “ Talisman ” expédition dredging no. 120, Azores,

560 m. The data available to Boschma were incomplète and partly incorrect. The correct data are:

“Talisman” drag. 123, 13.8.1883, 38°23'N, 28°49'45"W, 560 m. Azores.

Material studied

Azores: Prince of Monaco stn 229, 2 small colonies + branch, topotypes of Hornera eburnea

(mom); stn 597 ( ?), 3 small branches, labeled [by Calvet ?] Hornera eburnea (mom). — “ Talisman ”

drag. 123, about 15 colonies, branches and fragments including holotype of E. (L.) hicksoni (bmnh

1964.9.17.11, 1977.8.2.1; mnhn); drag. 128, 3 small branches (mnhn). — “ Jean Charcot" 1971, stn

159, 3 colonies (mnhn); stn 161, 2 colonies (mnhn); stn 213, 4 colonies (mnhn); stn 229, 3 small

colonies + branch (mnhn); unknown station, about 60 colonies, branches and fragments (most

mnhn; usnm 75600). — “ Bartlett" 1975, stn 2, branch (zmuk); stn 4, branch (zmuk).

Description

Colonies uniplanar, with regular dichotomous

branching forming V-shaped to U-shaped axils

(Fig. 1 A-E, 2 A). Examined specimens up to 25

mm high and 35 mm wide; holotype of Hornera

eburnea slightly taller (40 mm). Branches circular

in cross section, tapering to slender tips about

0.45 mm in diameter. Coenosteum white and

linear-imbricate (Fig. 2 F-G), composed of well-

ordered parallel strips each 0.11-0.12 mm wide.

Strips not well defined, bordered by an alignment

of narrow coenosteal pores 20-30 pm long. Pla-

telets irregular in width but rarely extend across

entire strip.

Gastropores occur primarily on anterior face,

each about 0.25 mm in diameter and often

bordered by a small proximal lip. Gastropore lip

does not actually cover any of gastropore (Fig. 1

B) but, rather, gives anterior directionality to

gastrozooid. Illustrated gastrostyle (Fig. 2 J) 0.50

mm tall and 0.16 mm in basal diameter (H:W =

3.13); gastrostyles measured by Boschma (1963a)

slightly more slender (H:W = 3.75). Gastrostyle

unridged and acutely conical with a pointed tip.

Extremely large spines cover gastrostyle in a

random arrangement; spines up to 68 pm long

and 10 pm in diameter. Gastropore tube smooth;

no ring palisade. According to Boschma (1963a),

gastrostyles occupy only lower one-sixth of gas¬

tropore tube.

Mound shaped dactylopores (Fig. 1 B, 2 D)

occur mainly on anterior branch faces and latéral

branch edges. Dactylopores often linearly arran-

ged on branch edges in sériés of 5-9 but ran-

domly arranged on branch faces. Dactylopores

50-70 pm in diameter, quite short (rarely more

than 60 pm tall), and projecting perpendicularly

to slightly anteriorly from branch.

Female ampullae massive, superficial, elliptical

mounds (Fig. 1 E, 2 H) about 0.9-1.1 mm in

diameter, the greater axis aligned with branch

axis. When mature, female efferent pore located

at distal vertex of ellipse, a concavity about 0.16

mm in diameter. Female ampullae occur on both

branch faces and often in sériés of two or three.

Male ampullae smaller, about 0.7 x 0.5 mm in

diameter, superficial on branch tip but internai in

larger diameter branches.

Comparions

Eleven valid species of Lepidopora (13, counting the 2 unnamed species included herein) hâve

been described: 6 from the Atlantic, 2 from the Subantarctic off South America, 1 from South Africa,

and 2 from New Zealand (Cairns, 1983b, 1985, 1986a). L. eburnea can be distinguished from ail

congeners by its dactylopore arrangement: short rows on latéral branch edges in addition to

dactylopores randomly scattered on anterior face. It is also characterized by having very short

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Fig. 2. — Lepidopora eburnea (A-B, D. F, G, from “ Talisman ” drag. 123, bmnh 1977.8.2.1; C, E, H-J, from “ Jean Charcot ”

1971, unknown station. Azores, lemale. usnm 75600): A, distal branch showing arrangement of gastro- and dactylopores

(x 18); B. branch edgc illustrating a row of dactylopores (x 29); C, Pedicularia trace (x 17); D, conical dactylopore

( x 252); E. detail of branch surface showing gastropore, several aligned dactylopores, and coenosteal texture ( x 33,

stereopair); F-G, coenosteal texture (x 121, x 300, respectively); H, female ampulla (x 41); I, gastrostyle spines

( x 289); J, gastrostyle ( x 97 stéréo pair).

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

dactylopore spines and a relatively low gastrostyle H:W ratio. Only one other species of Lepidopora

is known to hâve linear-imbricate coenosteal texture, the western Atlantic L. carinata (Pourtalès,

1867), which can be distinguished by its distinctively shaped, ridged dactylopore spines; ring palisade;

and complété absence of gastropore lips. Comparisons of L. eburnea to the 2 unnamed species

{Lepidopora sp. A and sp. B) are made in the accounts of the latter.

Remarks

L. eburnea was considered at first as a bryozoan and described as Hornera eburnea by Calvet

(1903). Calvet (1906) also recognized H. eburnea in the “ Talisman " collection, still considering it

as bryozoan. We agréé that his material from “ Talisman ” drag. 128 (Azores) is typical L. eburnea,

but we consider the stylasterid from “ Talisman ” drag. 96 (Mauritania) to be problematical, possibly

a distinct species (see Lepidopora sp. A). Only much later Calvet became aware that his Hornera

eburnea , as well as two other supposed bryozoans described by him ( H. verrucosa and H. gravieri —

see Errina dabneyi and Pliobothrus symmetricus), were stylasterid hydrocorals (Calvet, 1931: 45-46).

This rectification had been reiterated in the bryozoan literature by Borg (1944: 203) and Cook (1968:

238).

Hickson (1912b) erroneously referred specimens from the Azores (“ Talisman") to Pliobo¬

thrus tubulatus (Pourtalès, 1867), a species known only from the western Atlantic (Cairns, 1986a).

Intending to correct this confusion, and being unaware of Calvet’s Hornera eburnea, Boschma

(1963a) erected a new species, Errina ( Lepidopora ) hicksoni, which he mentioned again in later papers.

It lists as Lepidopora hicksoni in Cairns’ (1983b) check list, in which the former subgenus Lepidopora

had been elevated to generic rank. The “ transphyletic ” synonymy of Boschma’s stylasterid with

Calvet’s bryozoan was noted by Zibrowius (1981).

Distribution and ecology

The typical branched form of L. eburnea is known from about 10 stations in the Azores at

depths ranging from 480 m to 983 m.

Some branched colonies of Lepidopora from Seine Seamount (235 m) and Gorringe Seamount

(460-545 m) between Madeira and Portugal resemble the Azorean L. eburnea (see Lepidopora sp. A).

Symbionts

In the Azores, L. eburnea is frequently inhabited by Pedicularia. Two small specimens of this

gastropod hâve been obtained from branches from an unknown station (“ Jean Charcot ” 1971, cruise

biaçores) and the characteristic traces (Fig. 1 C, 2 C) hâve also been found on material from several

other stations (“ Talisman" drag. 123; “ Jean Charcot ” 1971, cruise biaçores, stn 161, 213, 229;

“ Bartlett ” 1975, stn 2, 4). Material with Pedicularia traces includes the holotype, paratype, and

topotypic specimens of Errina ( Lepidopora ) hicksoni ; the trace on the paratype is obvious on two

illustrations published by Boschma (1963a, pl. 1, fig. 2-3).

Lepidopora sp. A

Fig. 1G-K, 3 A-U, 4 A-E

Chresonymy:

Hornera eburnea — Calvet, 1906: 479 (part, “ Talisman " drag. 96, Mauritania).

Errina ( Lepidopora ) eburnea — Zibrowius, 1981: 982 (part: “ Talisman" drag. 96, Mauritania).

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Fig. 3. — Lepidopora sp. A (A-E, from “ Talisman " drag. 96, mnhn; F, from “ Cryos ” CP-95, mnhn; G-J, from “ Calypso ”

1958, sme-1277, mnhn; K-M, from “ Noroît " de-10, mnhn; N-0, from “ Noroît ” cp-12, mnhn; P, from “ Noroît ” cp-1 1,

mnhn; Q-S, from “ Noroît ” de-13, mnhn; T-U, from “ Noroît ” dw-108. mnhn): A, clavate form, same specimen as Fig.

4 A, opposite face illustrating gastro- and dactylopore arrangement ( x 11); B, clavate form, more eroded specimen with

less regular gastro- and dactylopore arrangement (x 11); C-D, slender form, distal branch fragments showing gastro-

and dactylopores ( x 11); E, clavate form, larger eroded specimen ( x 8.3); F, lower part of colony with bulging ampulla

and eroded surface ( x 14); G-H, lower part of colonies ( x 9.1, x 3.8, respectively); I-J, two faces of distal end of H

showing gastro- and dactylopore arrangement, I also showing internai ampullae ( x 9.8); K, eroded strongly clavate

colony showing gastropores and large efferent pores of internai ampullae ( x 3.5); L, nearly complété slightly clavate

colony showing gastro- and dactylopore arrangement (x 4.1); M. nearly cylindrical curved colony showing bulging

ampullae and distal dactylopores arranged in several ridges (x 4.1); N, cylindrical colony dichotomous in distal part

( x 2.8); O, distal part of N, opposite face ( x 4.6); P. eroded clavate colony showing gastropores and large efferent

pores of internai ampullae ( x 4.1 ); Q, incomplète clavate, like P ( x 4.1 ); R, eroded clavate colony, incomplète, showing

many large efferent pores of internai ampullae (x 4.1); S-T, slender cylindrical colonies (x 4.1); U, eroded clavate

colony showing gastro- and dactylopores and large efferent pores of internai ampullae (x 4.1).

Lepidopora sp. B (V, from “ Challenger ", stn 85, bmnh 1890.4.11.24): V, unique specimen, branch face

illustrating gastro- and dactylopore arrangement ( x 5.7).

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Source : MNHN, Paris

,f »

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Material STUDIED

Galicia, Gorringe and Seine Seamounts: “ Calypso ” 1958, stn sme-1277, 5 incomplète colonies,

dead (mnhn). — “ Noroit ” 1987, cruise seamount 1, stn dw-8, 6 specimens, basis or lower part of

colonies (mnhn); stn de- 10, 9 specimens, mostly fragments, lower or upper parts (mnhn); stn cp-11,

18 specimens, including bases, incomplète and complété unbranched colonies, dead (mnhn); stn

cp- 12, 17 specimens, including lower and distal parts, unbranched colonies (mnhn); stn de- 13, ca. 70

specimens including bases and nearly complété unbranched colonies (mnhn); stn dw-21, 45

specimens, including bases, various fragments and incomplète colonies, many pièces branched

(mnhn); stn cp-30, 3 specimens, including complété unbranched colony (mnhn); stn dw-78, 45

specimens including bases, lower parts and various fragments, many pièces branched (mnhn); stn

dw- 108, 13 specimens including various fragments, from base to nearly complété unbranched colony

(mnhn); stn dw-111, 2 incomplète unbranched colonies (mnhn); stn dw-116, lower part of colony

(mnhn).

Morocco: “ Cryos ” 1984, cruise balgim, stn cp-95, incomplète colony, dead (mnhn).

Mauritania: “ Talisman ” drag. 96, 12 colonies, some of them incomplète or in two pièces

(mnhn).

Description

The first samples of Lepidopora obtained from

Gorringe Seamount (“ Calypso ” 1958, stn sme-

1277) comprise the lower part of 5 colonies (part

of them considerably corroded). They are un¬

branched, up to 12 mm high, cylindrical, and

decrease in diameter to the distal fracture (Fig. 3

G-J). Gastropores and dactylopores occur on ail

sides. Ampullae are internai.

Further material obtained from Gorringe Sea¬

mount (“ Noroit ” 1987, cruise seamount 1, stn

dw-8, de-10, cp-11, cp-12, de-13, dw-21, cp-30)

is morphologically variable (Fig. 2 G-H, 3 K-S),

but includes many tiny fragments, often in poor

condition. Many pièces, including complété co¬

lonies, are similar to the earlier “ Calypso ”

material: cylindrical and unbranched, up to 15

mm high and 1.5-3.3 mm in diameter. Ampullae

internai to slightly bulging near distal end. Other

unbranched specimens of similar size (up to 18

mm high) tend to a clavate shape, with interme-

diates from almost cylindrical to considerably

widened in the upper part. Ampullae typically

bulging, even vaguely packed into four straight

Unes in some specimens. A large transverse

orifice up to twice as wide as a gastropore can be

found in the distal part of bulging ampullae, but

may be obstructed by calcareous deposit (Fig. 3

K, P, Q, R, U).

In addition to the unbranched, either slender

cylindrical or clavate specimens, some lots from

Gorringe Seamount (stn dw-10, cp-12, dw-21)

comprise branched fragments or incomplète co¬

lonies, with either internai or bulging ampullae.

The better preserved bifurcating specimen from

stn de-10 is 7.5 mm high and its terminal

branches are slightly clavate with distinct bulging

ampullae (Fig. 3 N-O). The unique bifurcating

colony from from stn cp-12 is 22.5 mm high and

cylindrical ail along, with an average diameter of

2.4 mm. It bifurcates only in the upper third.

Ampullae are not évident extemally. Most pièces

of the large lot from stn dw-21, up to 18 mm

high and 3 mm in diameter, are branched

(occasionally more than once) and show distinct

ampullae.

Material from Seine Seamount (stn dw-78,

Fig. 1 I-K) consists of one larger lot of mostly

branched (and corroded) specimens similar to

those from Gorringe Seamount (stn dw-21, Fig.

1 G-H). A few slender cylindrical unbranched

specimens are also présent, but no clavate ones.

Material from Galicia Seamount (stn dw-108,

dw-111, dw-116) includes subcylindrical to cla¬

vate unbranched specimens (maximum size 15.5

mm, Fig. 3 T-U). A poorly preserved specimen

looks branched, unless it results from specimens

being attached one on the other.

Ail well preserved Lepidopora from the three

seamounts resemble the Azorean L. eburnea by

having white, linear-imbricate coenosteum

composed of parallel strips. Gastropores may

hâve distinct proximal lip. Gastrostyles are simi¬

lar to those of L. eburnea, as are the dactylopo¬

res.

Source : MNHN, Paris

NE ATLANTIC AND M EDITERRANEAN STYLASTERIDAE

The unique specimen from Morocco (“ Cry-

os" 1984, cruise balgim, stn cp-95), in very bad

condition, is the lower part (6.5 mm high) of a

small mature colony (ampullae présent) (Fig. 3

F). Its diameter increases from the narrowest

zone just above the encrusting base to the distal

fracture and thereby is similar to clavate spéci¬

mens from Gorringe Seamount and Mauritania.

Material from Mauritania (“ Talisman ” drag.

96) first mentioned by Calvet (1906) comprises

15 small unbranched pièces in poor condition.

Some of these, belonging together, can be re-

combined into more complété colonies. Two

forms can be distinguished on the basis of their

general aspect.

The slender form (Fig. 3 C-D, 4 C-D) is slightly

sinuous, thin, subcircular in cross section, and of

rather uniform diameter. It is represented by 3 spé¬

cimens, the largest of which (incomplète, base mis-

sing) is 7 mm high and up to 1.8 mm wide. The cla¬

vate form (Fig. 3 A-B, D, 4 A-B) is rather massive,

straight, subcircular to slightly flattened in cross

section, and notably wider in the upper part (Fig. 4

A). Itis represented by 12 pièces, corresponding to

9 colonies, the largest of which (incomplète, base

missing), is 10 mm high and up to 2.5 mm wide in the

upper part. Distal branch tip of this form about 1.6

mm in diameter.

Both forms hâve white, linear-imbricate coen-

osteum composed of parallel strips about 0.15

mm wide. Their platelet structure (Fig. 4 C),

although worn in ail specimens, is very similar to

that of L. eburnea.

Gastropores 0.32-0.44 mm in diameter, each

bordered by a proximal lip, which is slightly

more prominent in several of the clavate speci¬

mens (Fig. 4 A-B, 4 D-E). Gastrostyles could not

be examined.

Conical to tubular dactylopore spines occur in

lines on latéral branch edges as well as sparsely

scattered on branch faces (Fig. 4 A-B, 4 D-E).

Dactylopores 67-80 pm in diameter and up to

0.25 mm ta.ll, directed perpendicular to branch

surface. Dactylopore centers about 0.37 mm

apart.

Ampullae (female ?) superficial, about 0.55

mm in diameter. No efferent pores noted.

Comparisons

The northeastern Atlantic stylasterids here provisionally grouped under Lepidopora sp. A

appear to reflect a disconcerting range of colony shape. Although we can see différences especially

between the end points, i.e. the typical Azorean L. eburnea and the distinctive clavate form from

Gorringe Seamount and Mauritania, we hesitate to draw a line. Ail these forms are clearly similar

in coenosteal texture; dactylopore arrangement ; gastropore lips; female ampullar size, location, and

efferent pore location; and gastrostyle shape. Structural uniformity thus constrasts with variation in

colony shape. Typical Azorean L. eburnea is repeatedly branching, whereas the clavate form is

typically unbranched (as is the slender cylindrical form, with intermediates tending to the clavate

one). But there are transitional specimens (occasionally with a clavate tip) that hâve long sections

without bifurcation resulting in consequently smaller colonies. We are not sure that slight size

différence, such as larger gastropores, higher and larger dactylopore spines, and wider coenosteal

strips are consistent and should be considered as essential arguments for species distinction.

Our comparison may also be biased by the lack of early ontogenetic stages and young colonies

of typical L. eburnea from the Azores, sorting of dredge contents on “ Jean Charcot ” cruise biaçores

having been less detailed than on “ Noroit ” cruise seamount 1. Obviously more well preserved

specimens, including from intermediate areas, must be examined to better understand this group.

Remarks

Stylasterids from 2 stations of the “ Talisman ” were referred by Calvet (1906) to the species

he had previously described from the Azores and mistaken for a bryozoan. In fact, Calvet’s

specimens from drag. 128 (Azores) are typical Lepidopora eburnea, whereas those from drag. 96

(Mauritania) are here included under Lepidopora sp. A (distinction not yet made by Zibrowius,

1982).

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Distribution and ecology

Stylasterids here grouped under Lepidopora sp. A are known from Galicia Seamount

(675-1125 m), Gorringe Seamount (470-2075 m), Seine Seamount (235 m), off Morocco (1378 m), and

off Mauritania (2320-2330 m), i.e. from a wider depth range than typical L. eburnea in the Azores.

Unbranched specimens, slender cylindrical and clavate, occur down to the greatest depths, whereas

specimens more similar to L. eburnea were obtained on the seamounts only at shallower depths

(235-545 m).

No symbionts are known.

Lepidopora sp. B

Fig. 3 V, 4 F-H

Material studied

Canary Islands: “ Challenger ” stn 85, dorsoventrally flattened fragment (bmnh 1890.4.11.24);

3 minor cylindrical fragments from same lot are specifically different and not considered here

(accidentai mixture in muséum collection ? indication of origin reliable ?).

Description

The study material consists of one dorsalven-

trally flattened, slightly sinuous unbranched up-

per branch fragment (Fig. 3 V) without the distal

end, 10.5 mm high and 1.5 mm wide at the widest

point. Coenosteal strips 0.12 mm wide, covered

by small granules. Five gastropores arranged

along midline of anterior face, each bordered by

a very broad crescent shaped proximal lip (Fig. 4

G-H) about 0.7 mm wide and 0.3 mm deep.

Elongate gastrostyles présent but not available

for detailed study. Dactylopore mounds linearly

arranged along each latéral branch edge, their

centers about 0.43 mm apart. Dactylopores el-

liptical (e.g. 78 x 56 pm in diameter), the greater

axis aligned with the branch axis (Fig. 4 G).

Dactylopore mounds about 0.15 mm tall and

0.18 mm in diameter. Superficial swellings (?

female ampullae) occur proximal to each gastro¬

pore on anterior face and in a staggered arran¬

gement on posterior face, each about 0.7 mm in

diameter. No efferent pores observed.

Comparisons

Lepidopora sp. B is similar to Lepidopora sp. A (clavate form), particularly regarding its

flattened branches and latéral rows of dactylopore mounds. Lepidopora sp. B differs in coenosteal

texture (granular, not linear-imbricate), having extremely prominent gastropore lips, lacking

dactylopores on branch faces, and having elliptical dactylopores (not circular).

Compared to typical L. eburnea from the Azores, Lepidopora sp. B has much broader

gastropore lips, larger dactylopores, and only two latéral rows of dactylopores. It is similar to the

western Atlantic L. glabra (Pourtalès, 1867) in dactylopore - gastropore coordination, but the

coenosteal texture is quite different.

Remarks

At the bmnh the unique sample of Lepidopora sp. B was found labeled “ Hydrocorallinae

Stylasteridae ”. Moseley (1881) probably had not seen it when he studied the “ Challenger ”

stylasterids.

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Fig. 4. — Lepidopora sp. A (A-E, from “ Talisman ” drag. 96, mnhn): A, clavate form. distal branch face illustrating gastro-

and dactylopores ( x 23); B, clavate form, distal branch edge showing aligned dactylopores and proximal gastropore

lip (x 26); C, slender form, coenosteal texture (x 116); D-E, slender form. branch faces of two female (?) specimens

(x 28, x 29, respectively).

Lepidopora sp. B (F-H, from “ Challenger ” stn 85, bmnh 1890.4.11.24): F-H, latéral, oblique, and top views of

branch fragment illustrating dactylopores and proximal gastropore lip ( x 23, x 40, x 40, repectively).

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Distribution and ecology

Lepidopora sp. B is known from only one station in the Canary Islands, at a much greater

depth (2100 m) than the typical Azorean L. eburnea.

No symbionts are known.

Genus PLIOBOTHRUS Pourtalès, 1871

Diagnosis. — Gastro- and dactylopores randomly arranged. Coenosteal texture linear-

imbricate; coenosteal pores large. Gastropore tube double-chambered (see Cairns, 1983b: 439); no

gastrostyles. Dactylopore spines conical or tubular; dactylopore tubes quite long; no dactylostyles.

Ampullae usually internai.

Type species: Pliobothrus symmetricus Pourtalès, 1868, from the western Atlantic.

Pliobothrus is represented in the study area by 2 species, including the type species.

Pliobothrus symmetricus Pourtalès, 1868

Fig. 5 A-G, 6 A-G

Synonymy:

Pliobothrus symmetricus Pourtalès, 1868: 141.

H ornera gravieri Calvet, 1911: 7, fig. 5.

Chresonymy:

Pliobothrus symmetricus — Duncan, 1870: 290, 297; 1873: 336, pl. 49, fig. 7. — Pourtalès, 1871: 57,

pl. 4, fig. 7-8; 1878: 211. — Moseley, 1879: 469, 479, 440-442 (Florida), pl. 41, fig. 2 (part, not “ Challenger ”

stn 23, Sombrero Island); 1881: 47-50, 78, 84, pl. 8, fig. 2 (part, not “ Challenger " stn 23, Sombrero Island).

— Agassiz, 1888: 138-139, fig. 444. — Nutting, 1895: 84. — Hickson, 1912b: 465-466. — Broch, 1914a: 3-7,

pl. 1, fig. 1-3, pl. 3, fig. 19-20, 28-29, pl. 4, fig. 34, 40-42, text-fig. A-B. — Dons, 1939: 196-197, fig. 1. —

Boschma, 1956b: F104, fig. 85.1a-b; 1957a: 59-60; 1967: 333-335, pl. 1, fig. 5-6. — Squires, 1965: 24, pl. 2,

fig. 7-8. — Zibrowius, 1981: 269; 1982: 982 (part, not Hyères Seamount). — Zibrowius & Cairns, 1982: 211,

212 (part, not Hyères Seamount). — Cairns, 1983b: 428, 439-441, fig. 3A-H, 24G, 25G, 27B (part, not stn j-s

43); 1986a: 16-17, fig. 6A-H (part, not Hyères Seamount).

Hornera gravieri — Calvet, 1931: 46. — Borg, 1944: 203.

Types

Pliobothrus symmetricus : In the original description Pourtalès (1868) simply reported P.

symmetricus as being “ not rare between 100 and 200 fathoms ofif the Florida reef ”. In a later more

detailed and illustrated description (Pourtalès, 1871) he reported the species from 7 dredging

stations in the same area. Cairns (1983b: 441) selected one of these as type locality and designated

a lectotype (colony illustrated by Pourtalès, 1871, pl. 4, fig. 7; Cairns, 1983b, fig. 3 A; Cairns,

1986a, fig. 6 A) and a paralectotype, among material deposited at the mcz (5529, 5530, respectively).

The ypm possesses another paralectotype (Cairns, 1986a).

Type locality: Restricted by Cairns (1983b) to “ Bibb ” stn 64, 11.5.1868, 24°17'N, 81°43'W,

262 m. Off Sand Key/Key West, Florida.

Hornera gravieri: Calvet (1911) reported H. gravieri from 11 stations in the Azores (Prince

of Monaco stn 568, 584, 597, 616, 618, 683, 712, 719, 838, 866, 869) and mentioned “ several beautiful

colonies and many fragments ”. Calvet did not designate the type locality and select types. None of

the specimens seen by Calvet could be found and ail may be lost (see Zibrowius, 1982).

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Fig. 5. — Pliobothrus symmetricus (A, from “ Talisman " drag. 123, mnhn; B, from “ Jean Charcot ” 1971, unknown station,

Azores; C-D, from “ Thalassa ” z-435, mnhn; E-F, from “ Thalassa " z-407, mnhn; G, from “ Challenger II" 1977, stn

134, bmnh 1986.11.5.2): A, large colony ( x 1.3); B, colony ( x 1.1 ); C, distal branch with Pedicularia trace ( x 3.3); D,

branch illustrating gastro- and dactylopore arrangement ( x 3.3); E, distal part of colony ( x 1.8); F, detail of E

illustrating gastro- and dactylopores (x 4.1); G, dichotomous branch (x 2.4).

Pliobothrus gracilis (H-L, from “ Calypso ” 1959, drag. 6, mnhn; M-N. from “ Calypso " 1959, drag. 4. mnhn) :

H-I, holotype, opposite faces (both x 1.7); J-K, details from holotype with bulging female ampullae (both x 3.0); L-N,

paratypes (ail x 1.3).

Source : MNHN, Paris

HELMUT ZIBROW1US & STEPHEN D. CAIRNS

Material STUDIED

W Atlantic: Lectotype and paralectotype of P. symmetricus and additional specimens

identified by Pourtalès (mcz; bmnh 1869.1.25.16, 1891.2.4.42, 1891.12.18.1).

Faroes: “ Michael Sors" 25.2.1904, colony + fragments (vsm).

Faroes - Hébrides area: “ Porcupine” 1869, station data uncertain, colony + branch

(BMNH 1883.12.10.131, 1898.5.7.17).

W Ireland: “ Challenger II” 1977, stn 134, 4 branches (bmnh 1986.11.5.2). “ Challenger II”

1981, stn 30, colony (bmnh 1989.6.16.1).

Celtic Sea: “ Thalassa ” stn z-407, branch (mnhn); stn z-415, distal branch fragment (mnhn);

stn z-435, 31 colonies + 60 branches/fragments (most mnhn; usnm 77121).

S Bay of Biscay: “ Travailleur ” drag. 70, branch (mnhn).

Galicia and Joséphine Seamounts: “ Noroit ” 1987, cruise seamount 1, stn dw-56, 6 fragments

(mnhn); stn dw-58, 3 fragments (mnhn); stn dw-111, 2 poorly preserved fragments, probably of P.

symmetricus (mnhn).

Madeira Archipelago: “ Jean Charcot ” 1966, stn 49, 4 poorly preserved fragments probably

of P. symmetricus (mnhn).

Azores: “ Talisman ” drag. 123, colony (mnhn). — “ Jean Charcot ”1971, cruise biaçores, stn 34,

branch (mnhn); stn 161 , colony + 2 branches (mnhn); stn 197, colony (mnhn); stn 213, colony (mnhn); stn

218,4 colonies + fragments (most mnhn; usnm 75601); stn 229,5 colonies + 2 branches (mnhn); stn 231,

12 colonies + 9 branches (most mnhn; usnm 77120); stn 232,4 colonies + 2 branches (mnhn); stn 240,15

small colonies on pebbles, provisionally included here, identity uncertain—see comparions (most mnhn;

usnm 77119); unknown station, 2 colonies + fragments (mnhn).

Imprécise locality Southwest of the Azores: said to corne from seamount 260 miles ( ?) [475 km]

Southwest of Faial, ca. 500 m, fishing boat “ Tomiguel ”, coll. J. G. Pereira, Sept. 1976, 2 colonies

+ 8 branches/fragments (mnhn).

Description

Eastern Atlantic specimens: colonies unipla-

nar, up to 60 mm high and 68 mm wide (Fig. 5

A-B). Branches robust, cylindrical to slightly

flattened along branching plane, 2.5-3.0 mm in

diameter. Branches gradually taper to blunt tips;

branch axils U-shaped. Coenosteal strips 80-

95 pm wide, separated by large, round to elon-

gate pores up to 65 pm wide. Platelets variable in

ornamentation, ranging from broad fiat platelets

(Fig. 6 F), to corrugated narrow platelets (Fig. 6

D-E), to an irregular arrangement of imbricating

spines (Fig. 6 G). Sometimes two different tex¬

tures occur on the same colony.

Gastro- and dactylopores occur on ail branch

surfaces but are more common on anterior

branch face (Fig. 5 D-G). Gastropores round to

slightly elliptical and 0.30-0.45 mm in

Comparisons

Of the 3 other species in the genus (ail occurring in the Atlantic), P. symmetricus is

distinguished from P. tubulatus (Pourtalès, 1867), by its shorter dactylopore spines and more robust

diameter, although most colonies hâve some

smaller diameter gastropores (e.g. 0.20 mm),

which may reflect developing gastropores or a

different size class. Dactylopore spines tubular,

up to 0.20 mm tall and 0.15 mm in diameter,

with a wall thickness of about 38 pm.

Female ampullae internai and 0.8-0.9 mm in

diameter, communicating to branch surface by a

tube that terminâtes in a spongy area on coenos-

teum 0.15-0.25 mm in diameter. Male ampullae

also internai and round to elliptical, with greater

axis perpendicular to branch surface. Male am¬

pullae 0.3-0.4 mm long, terminating in a slit like

efferent pore 0.05-0.11 mm long aligned with a

coenosteal slit. Ampullae from western Atlantic

material figured by Cairns (1986a) were female.

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Fig. 6. — Pliobothrus symmelricus (A-C, from “ Jean Charcot" 1971, stn 218, male, usnm 75601; D-E, from “ Thalassa"

z-435, male, usnm 77121; F, from imprécise “ Tomiguel" locality, male, mnhn; G, from “ Talisman " drag. 123, mnhn):

A, branch face illustrating gastro- and dactylopores (x 11); B, gastropore and several dactylopores (x 33); C,

dactylopore and coenosteal texture ( x 96); D, branch face illustrating large gastropores, smaller dactylopores, and even

smaller irregularly shaped coenosteal pores ( x 19); E, detail of large gastropore and one dactylopore, coenosteal texture

of narrow imbricate platelets (x 55, stéréo pair); F, coenosteal texture of broad fiat platelets (x 121); G, dactylopore

and rough coenosteal texturè of imbricate spines (x 119).

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

branches; from P. echinatus Cairns, 1986, by its nonspinose coenosteum and smaller gastropore

diameter; and from P. gracilis n. sp., by its internai female ampullae and larger gastropores.

One lot of 15 specimens of Pliobothrus from the Azores (“ Jean Charcot ” 1971, cruise

biaçores stn 240) is specially discussed here (Fig. 7 A-I). These specimens are unlike any of the other

known species of Pliobothrus in a number of characters. Their gastropores are circular and only

0.16-0.18 mm in diameter. Coenosteal pores are very narrow (e.g. 9 pm) producing a very dense

coenosteal texture, not porous as in the other species. The dactylopore spines are conical mounds

instead of thin walled tubes and ail of the colonies are short and sparsely branched (or unbranched)

with a firmly attached base. Several colonies bear low superficial female ampullae about 1 mm in

diameter, invariably with an efferent pore on the upper (distal) edge of the ampulla.

Remarks

P. symmetricus has occasionally been confused with each of the other species of Pliobothrus

occurring in the western and eastern Atlantic (rectification in Cairns, 1986a, and herein). Moseley

(1879, 1881) erroneously attributed to P. symmetricus a specimen of P. tubulatus (Pourtalès, 1867)

from “ Challenger ” stn 23 (Sombrero Island, West Indies). Nevertheless, he based his detailed study

of the skeleton and of the soft parts on authentic P. symmetricus (specimens from Florida, sent to

him by Pourtalès, cf. Moseley, 1879: 440; 1881: 47). Cairns (1983b) mistakenly included under P.

symmetricus a record which he subsequently (1986a) tranferred to a new species, P. echinatus Cairns,

1986. In the eastern Atlantic Pliobothrus gracilis n. sp. from Hyères Seamount, described in this

paper, had also erroneously been included under P. symmetricus (Zibrowius, 1982 ; Zibrowius &

Cairns, 1982; Cairns, 1986a).

In the eastern Atlantic, P. symmetricus has been mistaken for a bryozoan, described as

Hornera gravieri Calvet, 1911. Only much later Calvet (1931: 45-46) became aware that his H.

gravieri was a stylasterid hydrocoral. This rectification was mentioned again by Borg (1944: 203) and

Cook (1968: 238). The “ transphyletic ” synonymy of Calvet’s (1911) bryozoan with Pourtalès’

(1868) stylasterid was noted by Zibrowius (1982).

Distribution and ecology

In the western Atlantic, which includes the type locality of P. symmetricus, distribution and

depth range hâve been studied by Cairns (1986a). The species occurs mostly between 150 m and 400

m from the Blake Plateau off South Carolina through the Lesser Antilles, including the Pourtalès

Terrace off Florida.

In the eastern Atlantic P. symmetricus is known from many stations between 63°35'N and

31°26'N: southeast of Iceland, east of the Faroes, between the Faroes and the Hébrides, Norway

(62°3TN and 62°15'N), west of Ireland, Celtic Sea, Bay of Biscay, Galicia and Joséphine Seamounts,

Madeira Archipelago (to be confirmed, presently available material in poor condition), Azores, and

an unidentified seamount Southwest of the Azores. We hâve seen material from most of these areas

except from southeast of Iceland and from Norway; however, the specimens reported from these two

areas hâve been adequately described and figured (Broch, 1914a; Dons, 1939) and are included here

without hésitation.

Its depth range in the eastern Atlantic is from 80 m and 250-300 m for the shallowest stations

off Norway (Dons, 1939) to 1550 m and 1600 m for the deepest stations in the Azores (Calvet, 1911,

as Hornera gravieri). It is unknown whether live material had been obtained at these deepest stations;

in the Azores, with locally very irregular and steep slopes, dead specimens are occasionally collected

at uncommonly great depths. The deepest confirmed record of live material is 1050 m in the Celtic

Sea (“ Thalassa ” stn z-435).

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Fig. 7. — Piiobothrus sp. (A-I, frora “ Jean Charcot" 1971, stn 240, mnhn): A, tip of small unbranched colony illustrating

several gastropores and more numerous smaller dactylopores ( x 17); B, coenosteum bearing one gastropore and four

dactyloporcs ( x 49); C, detail of branch showing a dactylopore and coenosteal texture ( x 76); D-F, small unbranched

colonies (x 4.1, x 4.1, x 3.9, respectively); G-H, larger branched colonies(x 2.7, x 2.6, respectively); I, distal branch

tip illustrating gastro- and dactylopore arrangement ( x 3.9).

Symbionts

In the northeastem Atlantic, P. symmetricus is the host of Pedicularia, but, curiously, among

the abundant material studied here, only 3 branches from 2 stations in the Celtic Sea hâve been found

with the characteristic traces of the gastropod (“ Thalassa ” stn z-415, z-435; Fig. 5 C).

Moseley (1879: 469; 1881: 78) reported parasitic organisms thought to be pyenogonid larvae

from the gastric cavities of'gastrozooids of P. symmetricus dredged off Florida by Pourtalès. No

similar association is yet known from the northeastem Atlantic.

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Pliobothrus gracilis new species

Fig. 5 H-N, 8 A-G

Chresonymy:

Pliobothrus symmetricus — Zibrowius, 1982: 982 (part: Hyères Seamount). — Zibrowius & Cairns,

1982: 212 (part: Hyères Seamount). — Cairns, 1986: 17 (part: Hyères Seamount).

Types

Holotype and paratypes of Pliobothrus gracilis from 2 stations on slope of Hyères Seamount,

“ Calypso ” 1959, drag. 4 and drag. 6. Holotype a female colony collected alive from drag. 6, without

base (Fig. 5 H-I). Paratypes 3 colonies (2 with base) from drag. 4 and 3 colonies or branches (ail

without base) and 2 minor fragments from drag. 6; paratypes up to 30 mm high and 35 mm wide.

Holotype and most paratypes at mnhn, 1 paratype from drag. 6 at usnm (77118).

Type locality: “ Calypso ” 1959, drag. 6, 13.8.1959, 31°27.7'N, 28°55.6'W, 620-700 m, Hyères

Seamount.

Material studied

Hyères Seamount: holotype and paratypes.

Description

Colonies uniplanar: holotype 38 mm high and

36 mm wide, paratypes smaller. Branches cylin-

drical, gradually tapering through regular dicho-

tomous branching to blunt, rounded branch tips.

Branches slender, subterminal branches 1.25-1.75

mm in diameter; branch axils U-shaped. Coenos-

teum linear-imbricate (Fig. 8 E), the strips 65-

80 pm wide and of variable length, producing a

very porous coenosteum. Platelets fiat and irre-

gularly shaped, about 25 pm wide, occurring

three or four across a strip. Granular coenos¬

teum sometimes also présent on same specimen

bearing linear-imbricate texture, the granules

about 18 pm in diameter (Fig. 8 F).

Gastropores occur in low density on ail branch

surfaces but are most common on anterior face.

COMPARISONS

Gastropores small and usually circular (e.g.

0.18-0.25 mm in diameter) but may also be

elliptical, the greater axis parallel to branch axis

(e.g. 0.29 x 0.19 mm). Dactylopore spines also

scarce but most common on anterior branch

face. Dactylopore spines tubular, up to 0.10 mm

tall and 0.15 mm in diameter, with thin walls

about 38 pm thick (Fig. 8 B).

Female ampullae superficial mounds (Fig. 5

J-K, 8 G) 1.0-1.2 mm in diameter occurring on

both anterior and posterior branch faces.

Efferent pores 0.17-0.19 mm in diameter and

usually slightly off center from apex of bulge.

Male ampullae not discerned in limited material

available.

P. gracilis is distinguished from the other 3 species of Pliobothrus by its superficial female

ampullae and very small gastropores. It is further distinguished from P. symmetricus by its very

slender branches and low density of gastro- and dactylopores. P. gracilis is similar to the Azorean

Pliobothrus from " Jean Charcot" 1971, cruise biaçores stn 240 (see P. symmetricus, Comparisons)

in having superficial female ampullae and small circular gastropores, but differs in other characters,

such as: placement of efferent pore, coenosteal texture, colony size, and polyp density.

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Fig. 8. — Pliobothrus gracilis (A-G, from “ Calypso ” 1959, drag. 6, female paratype, usnm 77118): A. branch face showing

eflerent pores of three female ampullae, no gastropores évident ( * 18, stéréo pair); B, oblique view of coenosteum

illustrating elliptical gastropore and conical dactylopores ( x 66); C, top view of coenosteum illustrating three

dactylopores and coenosteal texture (x 61, stéréo pair); D-E, coenosteal texture (x 143, x 235, rcspectively); F,

granular coenosteum and two dactylopores ( x 87); G, female ampulla with large apical eflerent pore ( x 40).

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

E. aspera

E. dabneyi

E. atlantica

colony shape

uniplanar to slightly

bushy

uniplanar

bushy, sparsely

branched

branches: taper; distal

graduai taper;

abrupt taper;

graduai taper;

branch diameter

0.7 mm

0.6 mm

0.9 mm

coenosteal texture

reticulate-granular but

imbricate dactylopore

spines

reticulate-granular

gastropore lip

absent

présent

absent

gastrostyle shape

ridged

H:W = 1.5-3.3

ridged

H:W = 3.5

ridged

H:W = 2.3-3.3

dactylopore spines:

some clustering; up to

individual; 0.13-0.15

individual; 0.14-0.16

individuality; height,

width

0.65 mm, 0.27 mm

mm, 0.15 mm

mm, 0.25-0.27 mm

male ampullae:

internai; one

superficial; 1-3 apical

primarily internai; one

position; efferent pore

location

irregularly shaped

efferent pore 30-40 pm

in diameter

efferent pores

round efferent pore

50-60 pm in diameter

other diagnostic

many small dactylopo-

coenosteal papillae

larger colonies with

characters

res without associated

spine

sometimes présent,

branch anastomosis

occasionally

eunicid polychaete

Remarks

The spécifie name given to the new species refers to the slender form of the branches.

Previously, this form from Hyères Seamount had not been distinguished from the presumed

widespread P. symmetricus.

Distribution and ecology

P. gracilis is known only from 2 stations on the slope of Hyères Seamount, depth 600-700 m.

No symbionts are known.

Genus ERRINA Gray, 1835

Diagnosis. Gastro- and dactylopores usually randomly arranged. Coenosteal texture

reticulate-granular or linear-imbricate. Proximal gastropore lips common; gastrostyles présent,

having a moderate H:W ratio. Dactylopore spines U-shaped, with groove directed proximally; walls

of dactylopore spines thick; no dactylostyles. Ampullae superficial.

Type species: Millepora aspera Linnaeus, 1767, from the Mediterranean.

Errina is represented in the study area by 3 species, including the type species.

Errina aspera (Linnaeus, 1767)

Fig. 9 A-I, 10 A-H

Synonymy:

Millepora aspera Linnaeus, 1767: 1283.

Errina aspera mascarina Boschma, 1965a: 3-6, text-fig. 1, pl. 1, fig. 1 - 4 .

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

E. cochleata

E. altispina

uniplanar

abrupt taper;

graduai taper;

0.3-0.7 mm

0.4 mm

linear-imbricate,

linear-imbricate, broad

multiloculate platelets

platelets

présent, prominent

not ridged

H:W = 3.6

H:W = 4.9

individual; 0.10 mm,

individual, two types

0.10-0.13 mm

of spines

* superficial; several

1) 0.10 mm, 0.11 mm

2) 0.50 mm, 0.12 mm

* similar to that of E.

apical efferent pores

cochleata

giving ampullae spiny

aspect

gastropores restricted

coenosteal papillae

to center of anterior

sometimes présent

face and branching

axils

Table 1. — Distinguishing characters of the five North

Atlantic species of Errina (with new information on

two species front the West Indies *)

Chresonymy:

Millepora aspera — Esper, 1790: pl. 18, fig. 1-4; 1795: 106. — Lamarck, 1816: 202. — Risso, 1826:

347-348. — Darwin, 1854: 477, pl. 19, fig. 5a. — Aradas & Benoit, 1876: 301.

Errina aspera — Gray, 1835: 85. — Dana, 1848: 570-571. — Saville Kent, 1871: 282-283. — Moseley,

1879: 479; 1881: 84. — Carus, 1885: addendum. — Fol, 1885: 668-669. — Boschma, 1953a: 32-33; 1953b: 301,

etc.; 1954: 143, etc., pl. 1-3; 1956a: 283, 284, 286, 288; 1956b: F100; 1957a: 50-51; 1963a : 337, etc.; 1964f: 284;

1964g: 288; 1965b: 19; 1967: 329, 330, 331, 333. — Arnaud & Zibrowius, 1979: 123-124. — Fredj & Giermann.

1982: 284-285, photo 1-8. — Cairns, 1983b: 428, 459-462, fig. 11A-G. — di Natale & Mangano, 1985: 344.

— Giacobbe & Leonardi, 1985. — di Geronimo & Fredj, 1988: 243, pl. 1, fig. 1-4.

Errina aspera aspera — Boschma, 1965a: 1-3, text-fig. 2, pl. 1, fig. 5-7.

Errina ( Labiopora ) aspera — Hickson, 1912a: 888-889, pl. 95, fig. 6; 1912b: 462.

Errina aspera mascarina — Vervoort & Zibrowius, 1981: 29.

Hydrocoralliaire — Pérès, 1964: 21.

not Errina aspera — Verrill, 1864: 46 (Azores). — Pourtalès, 1867: 116 (Azores).

not Erina aspera — Boschma & Lowe, 1969: 15, pl. 5, map 2.

not Errina (Eu-Errina) aspera — Broch, 1942: 40, fig. 10.

Types

Millepora aspera : In the original description, Linnaeus (1767) characterized a structure which

we now recognize as typical dactylopore spines (prominent outgrowths with slit-like pore on the lower

side). This is a detail that he could not hâve found in the previous literature (Marsili, 1725;

Gualteri, 1742) and proves that he had before him at least one authentic specimen. The type could

not be found and is probably lost. Hickson (1912a: 888) said that he examined “ the type-specimen

of this species [aspera] in the British Muséum ” on which Gray (1835) founded the genus Errina. This

simply means that he has seen the specimen previously studied by Gray who made Linnaeus’ species

the type of the genus Errina.

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Type locality: Given by Linnaeus (1767) as Mediterranean and Norwegian Sea. This should be

restricted to Mediterranean. As pointed out by Boschma (1953a, 1953b), the indication Norwegian

Sea was due to a misunderstanding arising from a correspondence Linnaeus had with Gunnerus:

referring to Linnaeus, Gunnerus (1768) added to his description of Millepora norvegica (as the name

says, a species from Norway) that he considered this as being the same as Linnaeus’ M. aspera. In

fact, these are distinct species (see account of Stylaster norvegicus).

Errina aspera mascarina : This subspecies is said to corne from the Indian Océan (Mauritius)

and to differ from typical Mediterranean E. aspera aspera ; it was described by Boschma (1965a) from

2 slender colonies up to 37 mm high and 33 mm wide, one designated by him holotype, the other

paratype, both at mnhn. Originally these colonies were attached to a telegraph cable (substrate not

mentioned by Boschma).

Type locality: Given by Boschma (1965a) as Mauritius (Indian Océan), from unrecorded depth.

The origin, as indicated on the label found with the specimens (“ Spinipora, île Maurice, 1883, échange Vi-

mont ’*), is certainly misleading. The 2 types look like slender Mediterranean material and closely resem¬

ble another slender colony of E. aspera at the mnhn that also had grown on a telegraph cable. The latter spé¬

cimen (Fig. 9 B), found unidentified in the bryozoan collection, was labeled “ Toulon - sur le cable

transatl. ”, meaning probably that it had been detached from a telegraph cable in the Mediterranean (or

Straits of Gibraltar ?) and eventually landed at Toulon. Possibly the types of E. aspera mascarina and the

“ Toulon ” specimen were collected together from the same cable.

Material studied

Specimens of various sizes (colonies, branches, fragments) of E. aspera without exact collecting

data are présent in many muséum collections (e.g. in coral collections, or together with Pedicularia

sicula in mollusc collections): many colonies (mzuc). — 5 colonies already mentioned by Boschma

(1965a), Mediterranean or without indication (mnhn); types of E. aspera mascarina (mnhn); colony

“ Toulon - sur le cable transatl. ” (mnhn); branch with P. sicula, Palermo (mnhn). — colony (mzus).

— Messina, H. Fol, colony (mhng). — 2 colonies (nhmw 15714). — 2 branches with P. sicula (irsnb

IG 10591). — colony studied by Boschma (1954) (zma Coel 7401). — branch with P. sicula (bmnh

1846.12.8.1) and several colonies, either Sicily or without locality (bmnh 1842.3.29.47/49,

1843.3.6.147/148/150).— 2 colonies (uzmc). — colony (zmuo B 879). — photo (rmnh; by Boschma)

of colony in Manchester Muséum mentioned by Hickson (1912 a, as sent by J. Morgan). — colony

usnm 75602 (ex bmnh 1842.3.29.47). — Messina, branches with P. sicula (usnm 189433, 189435). —

Sicily, branches with P. sicula (amnh 14084, 14085). — fragment (ypm 5368).

Straits of Messina, more recently collected samples: G. Fredj, 1964, colony (mnhn). —

"Bannock " 1972, stn 23, off Punta Pezzo, 95 m, photo received from P. Colantoni showing colony

bearing P. sicula. — I. di Geronimo, 1986, several colonies from rocky bottom + many small

colonies on fouled fishing line, ca. 100 m (mnhn; usnm 75603).

Straits of Gibraltar and approaches: Geological cruise, W.N. Nesteroff (no details), small

colony (mnhn). — “ Calypso ” 1958, stn sme-1282, several colonies + branches (most mnhn; usnm

59931). — “ Al Mounir " stn b6-d6, b8-d2, dead colonies + branches (mnhn; usnm 48889). —

Cryos ’ 1984, cruise balgim stn dr-40, dr-49, dr-1 15, dr-1 16, dead branches + fragments (mnhn).

Atlantic coast of Morocco: “ Al Mounir" stn b10-d3, several colonies + branches (most

mnhn; usnm 48889; bmnh 1980.1.4.1).

Cape Verde Islands: “ Talisman ” drag. 114, 4 branches (mnhn) + fragment (bmnh

1950.1.11.94).

Description

Colonies primarily uniplanar but sometimes

bushy (Fig. 9 A-B). Examined specimens up to

20 cm high and wide. Branches cylindrical,

gradually tapering to pointed tips about 0.7 mm

in diameter. Coenosteum white and porous.

Coenosteal texture predominantly reticulate-

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Fig. 9. — Errina aspera (A, from Messina, coll. H. Fol, mhng; B, froin unknown locality, “ Toulon ", mnhn; C-D, from

“ Calypso " 1958, sme-1282, mnhn; E, from “ Al Mounir " b10-d3, usnm 48889; F-I, from “ Talisman ” drag. 114, F-H:

mnhn, I: bmnh 1950.1.11.94): A, large colony (x 0.7); B. small slender colony very similar to types of E. aspera

mascarina ( x 1.8); C-E, details of branches showing dactylopore spines and gastropores with gastrostyles (ail x 10);

F, four branches and fragments ( x 1.1); G, distal part, opposite face of larger branch under F showing two Pedicularia

traces ( x 1.1); H, tip of slender branchlet from larger branch under F ( x 12); I, distal part of another branch ( x 14).

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

granular (Fig. 10 E), but sides of dactylopore

spines and inner gastropore tube surface usually

imbricate (Fig. 10 F, H), a rare combination of

coenosteal textures in one species. Coenosteal

strips 45-75 pm wide; granules irregular in shape.

Gastropores circular, 0.18-0.25 mm in diame-

ter, without proximal lip. Gastropore tube cylin-

drical, lacking a ring palisade. Gastrostyle oc¬

cupas lower half of gastropore tube. Gastrostyle

lanceolate: up to 0.27 mm tall and 0.09 mm in

diameter, H:W ratios ranging from 1.5-3.3. Gas¬

trostyles bear short, vertical ridges, which are

covered by large pointed spines up to 26 pm

long. Individualized dactylopore spines project

perpendicularly from branch surface, predomi-

nantly with groove directed proximally (Fig. 9

C-E); however, some dactylopore spines directed

obliquely, particularly when 2 or 3 are clustered.

On distal branches dactylopore spines occur on

ail sides of branch but are rare on posterior face

of larger basal branches. Dactylopore spines up

to 0.65 mm tall; width, about 0.27 mm; width of

dactylopore groove, 0.06-0.08 mm, or about

one-quarter width of spine. Smaller slit-shaped

dactylopores, unaccompanied by spines, are

common, measuring 0.13-0.16 x 0.06 mm.

Female ampullae hemispherical, 0.5-0.7 mm in

diameter, and often spinose as a resuit of short

dactylopore spines. Female efferent pores about

0.11 mm in diameter, but were rarely observed.

Male ampullae internai (Fig. 6 G), elliptical in

shape, and about 0.4 mm in greater internai

diameter. Each male ampulla communicates to

surface by a narrow irregularly shaped efferent

pore 30-40 pm in diameter (Fig. 10 E).

Comparisons

There are 17 valid Recent species of Errina, a genus known from the Atlantic, off South

Africa, the Galapagos, and particularly from the New Zealand région, Subantarctic, and Antarctic

(Cairns, 1983b, 1986a, 1986 b). The 5 Atlantic species are compared in Table 1. E. aspera is most

similar to E. atlantica but can be distinguished by its taller dactylopore spines (which are imbricate

in texture); the presence of numerous unspined dactylopores; and a tendency to be uniplanar in

colony shape.

Remarks

E. aspera is recognizable in the brief original description of Millepora aspera by Linnaeus

(1767), which certainly was based on material from the Mediterranean. Linnaeus also referred to

Marsili (1725) and to Gualteri (1742), believing that his species had already been figured by these

authors. However, as pointed out by Boschma (1953b), who analyzed and reproduced the pertinent

passages and figures from ail these older publications, Marsili’s and Gualteri’s illustrations are

difficult to interpret, and bryozoans may be involved as well, at least in part.

The oldest known stylasterid from European seas, E. aspera has been the subject of

considérable confusion. Confusion with the morphologically very different Stylaster norvegicus is

explained above (see Types).

E. dabneyi (Pourtalès, 1871), from the Azores, had first been mentioned as E. aspera by

Verrill (1864) and Pourtalès (1867) before it was described as a distinct species.

Boschma’s (1965a) subspecies mascarina is discussed above (see Types).

E. aspera sensu Boschma & Lowe (1969) from Antarctica proved to be E. gracilis Marenzeller,

1903, as indicated by Cairns (1983a: 98, 100; 1983b: 459).

E. aspera sensu Broch (1942), said to corne from the West Indies, was also misidentified. The

small branch of slightly pink colour (smnh 45) does not belong to either the Mediterranean — eastern

Atlantic E. aspera , or to any species known from the West Indies (see Cairns, 1986a), but represents

some other species, possibly of Antarctic or Subantarctic origin.

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Fig. 10. — Errina aspera (A, C, H, from Straits of Messina, female, usnm 75603; B, D, F, from “ Calypso ” 1958, sme-1282,

usnm 59931; E, G, from “ A! Mourir ” b10-d 3, female, usnm 48889): A, branch segment illustrating a gastropore and

numerous dactylopore spines ( x 17, stéréo pair); B, C, D. F, aspects of dactylopore spines and coenosteal texture

(x 45, x 47, x 120, x 120, respectively, C being a stéréo pair); E, coenosteal texture (x 120); G, transverse branch

fracture revealing two gastrostyles and internai male ampullae ( x 36); H, gastrostyle ( x 140).

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Distribution and ecology

As demonstrated above (see Types, Remarks), E. aspera has been reported by error from

Norway, the Azores, the West Indies, Mauritius, and Antarctica.

The occurrence of typical E. aspera is confirmed only for the Straits of Messina, the Straits

of Gibraltar with its Atlantic approaches, and a locality further south on the Atlantic coast of

Morocco (33°43.5'N). A few branches from the Cape Verde Islands differ slightly from the typical

Mediterranean form (see below).

In the Mediterranean the occurrence of E. aspera is sufficiently confirmed only for the Straits

of Messina. Aradas & Benoit (1876) reported the associated gastropod Pedicularia sicula (on

Millepora aspera) as frequently obtained there by the red coral fishermen. Abundant material of

Errina, probably that examined by those authors, is présent in the collection of the mzuc. Fol (1885),

who observed the red coral fisheries aboard a boat from Messina, collected Errina and Pedicularia

(mhng). Thanks to the red coral fisheries once prosperous in and near the Straits of Messina, the

fortuitously caught E. aspera and Pedicularia sicula are represented in many muséum collections,

generally with poor (if any) indications on the origin, such as Mediterranean, Sicily, or, at best,

Messina.

Referring to observations from submersible dives, Fredj & Giermann (1982) reported E.

aspera to be very abundant on rocky bottom exposed to strong currents at depths of about 110-130

m in the straits between Ganzirri (Sicily) and Punta Pezzo (Calabria). In this part of the straits (off

Punta Pezzo, 95 m) the hydrocoral with its gastropod symbiont has also been dredged by the

“ Bannock ” in 1972 (photo communicated by P. Colantoni; Arnaud & Zibrowius, 1979; Selli et

al., 1980), and more recently by I. di Geronimo (material studied). The depth range of the species

was extended by di Natale & Mangano (1985) on the basis of sampling down to 181 m, and of a

video-record from a submersible dive down to 236 m ofT Capo Annunciata, south of Messina (Sicily).

The characteristic E. aspera hard bottom of the straits was again mentioned by Giacobbe &

Leonardi (1985).

On the basis of dredgings and the previously mentioned submersible observations, di

Geronimo & Fredj (1988) summarized the depth distribution of E. aspera as being roughly 80-90 m

to 200 m. They further pointed out that the stylasterid occurs together with the large barnacle

Pachylasma giganteum (Philippi, 1836), which also abounds and coexists with E. aspera in the Straits

of Gibraltar on deep hard bottoms. Previously Darwin (1854) had mentioned and illustrated the

co-occurrence of both species (** Sicily, deep water ”).

E. aspera has been recorded as a Plio-Pleistocene fossil from several localities in Sicily and

Calabria (see Records of fossil stylasterids from Europe).

From various old collections and mainly old malacological literature (presence in the

Mediterranean of Pedicularia implies that of its host E. aspera) the following list of hypothetical

Errina localities has been compiled: coast of North Africa, Alger, Maltese Islands, Palermo, Lipari

or Eolian Islands, Capri, Nice area, Hyères, Toulon (the latter 3 localities on the coast of France).

These indications do not prove the extended range of E. aspera ; specimens may hâve been obtained

there by naturalists from red coral fishermen or dealers, or the origin may be confused for other

reasons. It is particularly unlikely that the stylasterid (included by Risso, 1826, in the fauna of the

Nice area) occurs in the northern Mediterranean. It is more likely to be found along the coast of

North Africa since it also occurs in the Straits of Gibraltar.

Like the Straits of Messina, the Straits of Gibraltar is known for its hard bottom exposed to

strong currents. Although few samples of Errina hâve yet been collected in the straits and its

approaches (see Material studied) — the irregular and steep bottom is difficult to dredge — we can

conclude that the stylasterid must be rather common there. In fact, shells of Pedicularia hâve been

found abundantly (together with rare fragments of Errina) in coarse sédiments under the

Mediterranean outflow slightly west of the sill (“ Cryos ” 1984, cruise balgim). Apparently, the coral

branches, when dead and detached, remain close to where they lived, whereas the lighter shells are

sorted by the current and transported further away.

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

In the nearby Atlantic, E. aspera has been obtained from Spartel Bank, in the western

approaches of the Straits of Gibralter (first mentioned by Pérès, 1964, as “ Hydrocoralliaire ”), and

from farther south on the coast of Morocco (33°43.5'N). Pedicularia sicula from that station had been

reported independently by Bellon-Humbert & Gofas (1977), who did not mention that it was an

obligate symbiont of the stylasterid.

The occurrence of E. aspera in the Cape Verde Islands requires further confirmation. The few

branches obtained there by the “ Talisman ” at the unusual depth of 600 m differ from typical

Mediterranean E. aspera by their more spinous aspect, which we hâve interpreted as intraspecific

variation (Fig. 9 F-I).

Symbionts

Throughout its geographical range E. aspera is found with traces of Pedicularia. The symbiotic

P. sicula has been obtained, either on live branches or as dead shells, from the sédiment in the Straits

of Messina, the Straits of Gibraltar and its western approaches (Spartel Bank), and further south on

the Atlantic coast of Morocco (33°43.5'N). Only the traces of the symbiont are known from the Cape

Verde Island material (Fig. 9 G).

In the Straits of Messina P. sicula is a common symbiont. Dead shells swept away by the

currents hâve been obtained from the sédiments at greater depths and at greater distances north and

south of the sill (Selli et al., 1980): towards the Tyrrhenian Sea at 329 m off Scilla, and at 220 m

towards the Ionian Sea off Villa San Giovanni.

Pedicularia is known as a fossil from the Lower Pleistocene of Sicily and Calabria (see Records

of fossil stylasterids from Europe).

Errina dabneyi (Pourtalès, 1871)

Fig. 11 A-I, 12 A-F, 13 A-B

Synonymy:

Lepidopora dabneyi Pourtalès, 1871: 41, pl. 7, fig. 10-11.

Hornera verrucosa Calvet, 1903: 161-162, pl. 18, fig. 6a-c.

Errina amoena Boschma, 1956a: 281, text-fig. 1-3, pl. 1-2, pl. 3, fig. 1-4.

Chresonymy:

Errina aspera — Verrill, 1864: 46. — Pourtalès, 1867: 116.

Errina dabneyi— Moseley, 1879: 479; 1881: 84. — Hickson, 1912a: 893 ; 1912b: 463-464, pl. 8, lower

fig. — Boschma, 1953b: 313, 314; 1954: 148; 1956a: 286-288; 1957a: 53; 1963a: 337; 1963c: 398, text-fig. 1-4, pl.

1, fig. 1-8; 1964f: 284-285; 1965a: 2; 1967: 331; 1968b: 113. — Cairns, 1983b: 428.

Errina dabneyae — Boschma, 1953b: 313.

Errina ( Errina ) dabneyi — Zibrowius, 1982: 981-982.

Hornera verrucosa — Calvet, 1931: 45. — Borg, 1944: 203. — Belloc, 1960: 12.

Errina amoena — Boschma, 1957a: 50; 1963a: 335.

Errina ( Errina ) amoena — Vervoort & Zibrowius, 1981: 26.

Errina spec. 2 — Boschma, 1967: 330-331.

Types

Lepidopora dabneyi: In the original description, Pourtalès (1871) mentioned several large

colonies sent to the mcz by Miss Dabney (of the family of the American consul at Faial) that were

obviously collected by local Azorean fishermen. No précisé type status was given to the specimens by

Pourtalès. The type lot (here considered as syntypes) preserved at the mcz comprises 3 large

fan-shaped colonies 20-30 cm high and wide, and various smaller branches and minor fragments, at

least in part broken off the larger colonies. Two fan-shaped branches from the type lot (the larger

58 mm high, 27 mm wide)'were tranferred to the bmnh (1891.2.4.29). Some branches and fragments

from the type lot are also at rmnh (Coel 13907) (Fig. 11 A) and usnm (75608).

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Type locality: Faial, Azores. Upper bathyal depths (no depth indication in Pourtalès, 1871;

but remarks on Coryophyllia cyathus as epifauna permit this bathymétrie extrapolation).

Hornera verrucosa: Calvet (1903) mentioned 4 colonies, ail from one station, and figured one

(29 mm high, 22 mm wide). No précisé type status was given to these specimens, here considered as

syntypes. Only part of the material mentioned was found at the mom: 8 small fragments (possibly

from a single branch or colony, but apparently not from the figured one), the largest about 20 mm

high (Fig. 11 E).

Type locality: Prince of Monaco stn 247, 30.8.1888, 38°24'N, 28°0T25"W, 318 m. Azores.

Errina amoena : Boschma’s (1956a) original description was based on a single, large,

fan-shaped colony (29.5 cm high, 31.5 cm wide) of which larger parts and smaller sectors hâve been

figured. Being the only specimen mentioned, it must be considered as the holotype, which is preserved

at mnhn, together with some detached branches. Some smaller branches from the holotype at rmnh

(Coel 15855) (Fig. 11 B) and usnm (75605).

Type locality: Given by Boschma (1956a) as “ China Sea, depth not recorded ” based on old

label found with type colony (“ Stylaster flabelliformis, Mer de Chine, Mr. Montigny ”). In view of

the otherwise confirmed distribution ( E. amoena being a synonym of E. dabneyî), it is most likely that

the indicated origin was wrong, as was the identification on the old label (S. flabelliformis being a

large fan-shaped, morphologically convergent, species from the Mascarene Islands; see Boschma,

1957b), and that the holotype of E. amoena came from the Atlantic.

Material studied

Azores: Types of Lepidopora dabneyi, Hornera verrucosa, and Errina amoena. — Azores,

Sa vigny, 1843, 4 pièces of large fan-shaped colonies (zmb 1066). — Origin not given but undoubtedly

from local Azorean fisheries, large fan-shaped colony on piece of volcanic rock (mcm 63). —

“ Talisman" drag. 123, ca. 15 colonies, branches, fragments (mnhn); branch (figured by Hickson,

1912b: pl. 8, lower fig.) + small branch (bmnh 1950.1.11.93; 1964.9.17.10). — “ Jean Charcot ” 1971,

cruise biaçores stn 4y, 1 5 small branches + trag

Mid-Atlantic Ridge: “ Bartlett" 1975, stn

Description

Colonies uniplanar, up to 30 cm high and

wide; branches cylindrical and occasionally anas¬

tomose. Small diameter branches project perpen-

dicularly from the larger diameter main bran¬

ches. Distal branch tips about 0.60 mm in

diameter. Coenosteum white and reticulate-

granular in texture. Strips 55-100 pm wide, co-

vered by rounded granules about 8 pm in diame¬

ter. Coenosteal papillae 63-100 pm in diameter

and up to 0.15 mm tall sometimes occur on and

adjacent to male ampullae (Fig. 11 G, 12 E).

Papillae are assumed to contain additional ne-

matocysts or a different type of nematocyst and

therefore to serve a défensive function.

Gastropores circular, 0.15-0.20 mm in diameter,

and bordered by a broad proximal lip. Gastropore

tube cylindrical, lacking a ring palisade; gastrostyle

occupies lower half of chamber. Gastrostyle lan-

ceolate: illustrated style (Fig. 13 B) 0.31 mm high

(most mnhn; usnm 75606).

1, 4 branches (zmuk).

and 0.091 mm wide (H:W = 3.4). Gastrostyle

strongly ridged, the ridges bearing pointed spines

up to 27 pm long. Dactylopore spines individuali-

zed, almost exclusively with groove directed proxi-

mally, and occurring uniformly on ail sides of distal

branches but infrequently on larger diameter bran¬

ches. Dactylopore spines also very common on un-

dersides of gastropore lips (Fig. 12 C). Dactylopore

spines short, projecting perpendicularly from

branch surface up to 0.13-0.15 mm. Width of spine

about 0.15 mm; width of groove 44-50 pm, or about

one-third spine width. There are no unspined

dactylopores.

Female ampullae (Fig. 12 F) hemispherical,

0.6-0.7 mm in diameter, with an efferent pore

diameter of about 0.17 mm. Male ampullae (Fig.

12 E) smaller, slightly irregular protubérances,

0.4-0.5 mm in diameter, each with 1-3 small

apical efferent pores.

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Fig. 11. — Errina dabneyi (A, syntype of Lepidopora dabneyi, rmnh Coel 13907; B, holotype of Errina amoena, rmnh Coel

15855 ; C-D, from “ Jean Charcot ” 1971, stn 49, mnhn; E, syntype of Hornera verrucosa , mom; F-G, from “ Talisman

drag. 123. mnhn; H, from “ Talisman" drag. 123. bmnh 1950.1.11.93; I, from “ Barilett 1975, stn 14, zmuk): A.

colony fragment ( x 2.0); B, colony fragment (3.0); C-D, branches with anastomoses (both * 1.9); E, distal part of

slender branchlet ( x 15); F, colony having overgrown colony of Pliobothrus ( x 1.5); G, detail of F showing abundance

of coenosteal papillae ( x 4.0); H, Pedicularia trace (x 13); I, branch with abundant coenosteal papillae (x 3.0).

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

COMPARISONS

Among the Atlantic Errina, E. dabneyi is clearly most similar to the western Atlantic E.

cochleata Pourtalès, 1867 (Table 1) but can be distinguished by its ridged gastrostyles, reticulate-

granular coenosteal texture, and occurrence of numerous dactylopore spines on the underside of each

gastropore lip.

Remarks

Previous to its description as a distinct species, the type sériés of E. dabneyi had been referred

to E. aspera (Linnaeus, 1767) by Verrill (1864) and Pourtalès (1867).

Pourtalès’ (1871) original description of E. dabneyi (as Lepidopora) is very brief. A more

detailed description, based on the types, was given by Boschma (1963c), who also mentioned this

species in several other papers.

E. amoena Boschma, 1956, based on an old muséum specimen said to corne from the China

Sea, proved to be identical with E. dabneyi (Boschma, 1963c, 1964f, 1968b). As for the unlikely

disjunct distribution, we présumé that the type of E. amoena was in fact of Atlantic origin and had

been mislabeled.

Yet another synonym of E. dabneyi is Hornera verrucosa Calvet, 1903, described as a bryozoan

from the Azores. Only much later Calvet (1931: 45-46) became aware that his H. verrucosa was a

stylasterid hydrocoral. This rectification has been reiterated in the bryozoan literature by Borg (1944:

203) and Cook (1968: 238). The “ transphyletic ” synonymy of Calvet’s (1903) bryozoan with

Pourtalès’ (1871) stylasterid was noted by Zibrowius (1982).

Hickson (1912b) tentatively referred specimens from the Azores (“ Talisman ”) to E. dabneyi,

a species which at that time had not yet been illustrated, and of which he had not seen the types.

Having seen these, Boschma (1963c, 1967) concluded that Hickson’s identification of the

“ Talisman ” material was incorrect, and that the form in question ( Errina sp. 2 in Boschma, 1967)

was more similar to E. aspera (Linnaeus, 1767). We hâve reexamined the 2 branches seen by

Boschma, as well as additional material from the same station; we cannot follow his interprétation

and refer the whole lot to E. dabneyi.

Distribution and ecology

E. dabneyi is known from the Azores (recorded depths of 3 stations ranging from 215-225 m

to 500 m) and from the Mid-Atlantic Ridge Southwest of the Azores (recorded depths of one station

140-2200 m).

For other samples the depth had not been recorded. According to Pourtalès (1871),

Caryophyllia cyathus (Ellis & Solander, 1786) was growing on the thicker branches (type lot of

Lepidopora dabneyi ) received from Faial. In the Azores this scleractinian is common in depths of a

few hundred meters (Zibrowius, 1980). Jourdan (1895: 10) also mentioned a specimen of C. cyathus,

and Dautzenberg (1889: 39) specimens of Pedicularia taken from the same coral substrate

(unidentified “ polypier ”) from off Faial, 400-500 m), and offered to the Prince of Monaco by W.S.

Dabney in 1887. The coral substrate in question (no longer preserved either with C. cyathus or

Pedicularia at the mom) might well hâve been another colony of E. dabneyi.

To summarize, in the Azores E. dabneyi appears to be a species mainly of upper bathyal depths.

Symbionts

The characteristic traces of Pedicularia (Fig. 11 H) hâve been found on branches of most lots

of E. dabneyi from the Azores (or presumably from the archipelago), including the types of

Lepidopora dabneyi, Hornera verrucosa, and Errina amoena, and branches from “ Talisman ” drag.

123 and “ Jean Charcot ”1971, cruise biaçores stn 49. When examined in 1979, a large colony at the

mcm (no. 63) still had several small Pedicularia attached.

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Fig. 12. — Errina dabneyi (A-B, holotype fragment of Errina amoena, USNM 75605; C, from “ Jean Charcot" 1971, stn 49,

female, usnm 75606; D, F, syntype of Lepidopora dabneyi, female, usnm 75608; E, from “ Talisman " drag. 123, male):

A, branch face illustrating gastro- and dactylopores ( x 27, stéréo pair); B, branch tips ( x 26); C, branch face showing

dactylopore spines and underside of proximal gastropore lip ( x 38, stéréo pair); D, dactylopore spines ( x 158); E, male

ampulla ( x 38, stéréo pair); F, female ampullae ( x 32).

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Fig. 13. — Errina dabneyi (A-B, from “ Jean Charcot " 1971, stn 49, female, usnm 75606): A, gastrostyle and gastropore tube

(x 91); B, gastrostyle ( x 175, stéréo pair).

Errina atlantica Hickson, 1912

Fig. 14 A-F, G-I ?, 15 A-G

Synonymy:

Errina atlantica Hickson, 1912b: 464-465.

Chresonymy:

Errina atlantica — Boschma, 1957a: 52; 1967: 331-333, text-fig. 3a-b, pl. 1, fig. 7-10; 1968c: 206. —

Cairns, 1983b: 428.

Errina ( Lepidopora) atlantica — Boschma, 1963a: 338-339; 1963b: 395; 1964a: 60-61.

Types

Errina atlantica : In the original description (Hickson, 1912b), which was based on several

branches from the Azores (“ Talisman ”), no specimens had been figured or otherwise individually

characterised by size, or given a précisé type status. Material identified by Hickson as E. atlantica

is présent at the mnhn (various branches and fragments, including those figured by Boschma, 1967:

pl. 1, fig. 7-8), at the bmnh (1950.1.11.87, 1950.1.11.99, small branch and tiny fragment), and at the

usnm (75609, fragment from bmnh 1950.1.11.87). When rediscovered in 1977, the largest pièces at the

mnhn were branches 90 mm and 60 mm high, but hâve subsequently been accidentally broken into

smaller pièces. The various old labels with these samples indicating dredge number, date, depth, and

locality, are confusing and at variance with the official station list of the expédition. The greater part

of the material certainly cornes from “ Talisman ” drag. 128 (syntypes; Fig. 14 E), the remaining part

possibly from drag. 127.

Type locality: Herewith designated as “ Talisman ” drag. 128, 16.8.1883, 38°07'N,

27°11'45"W, 983 m. Azores.

Material studied

Azores: Origin not given but undoubtedly from the local Azorean fisheries, large colony (mcm

62). — “ Talisman ” drag. 127 (?), branch originally 90 mm high, now broken (mnhn); drag. 128,

syntypes of Errina atlantica (see above). — “ Jean Charcot ” 1971, cruise biaçores stn 197, several

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Fig. 14. — Errina atlantica (A-D, from “ Jean Charcot ” 1971, stat 240, mnhn; E-F, syntypes of E. atlantica, “ Talisman " drag.

128, mnhn): A, bushy colony with incorporated tube of Eunice norvegica (x 0.8); B, Pedicularia trace (x 14); C-D,

branch tip and branch segment illustrating gastro- and dactylopore arrangement (both x 18); E, branch with bulging

ampullae ( x 2.8); F, branch tip illustrating gastro- and dactylopore arrangement ( x 8.6).

Errina atlantica ? (G, from “ Noroit" de-13, mnhn; H-I. from “ N or oit ” dw-21, mnhn): G, eroded branch

(x 9.3); H, small incomplète colony (x 4.6); 1, branch (x 11).

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

large pièces and many branches from at least one big colony (mnhn); stn 212, branch, dead (mnhn);

stn 230, central piece of a colony, dead (mnhn); stn 240, several large pièces and many branches of

several colonies (most mnhn; usnm 75610).

Gorringe Seamount: Material provisionally included here, identification uncertain (see

Comparisons). “ Noroit ” 1987, cruise seamount 1, stn de-13, 11 fragments, dead (mnhn); stn dw-21,

5 fragments, including 1 live specimen (mnhn).

Description

Colonies bushy and sparsely branched (Fig. 14

A); largest pièces of colonies examined up to 12

cm high and wide, complété colonies apparently

considerably larger. Branches cylindrical, gra-

dually tapering to rather thick distal branches

0.8-1.0 mm in diameter. Coenosteum white,

reticulate-granular in texture (Fig. 15 C-E).

Strips 65-95 pm wide; granules 10-14 pm in dia¬

meter.

Gastropores circular, 0.22-0.30 mm in diame¬

ter, without a proximal lip. Gastropores occur

predominantly on anterior face and latéral

branch edges but not always in linear sequence as

suggested by Boschma (1967). Gastropore tube

lacking ring palisade (Fig. 15 G). A slender,

sharply pointed gastrostyle occupies lower 50-60

% of gastropore tube. Illustrated style (Fig. 15

G) 0.31 mm tall and 0.093 mm wide (exclusive of

projecting spines) for a H:W ratio of 3.3. Two

gastrostyles measured by Boschma (1967: 332)

had lower ratios of 2.3-2.6. Gastrostyle vertically

Comparisons

As previously discussed, E. atlantica is most similar to E. aspera (see Table 1 and E. aspera,

Comparisons).

Tiny pièces (18 mm maximum length and 1.4 mm maximum diameter) of Errina from

Gorringe Seamount (“ Noroit ” 1987, cruise seamount 1 stn de-13, de-21) are provisionally

considered as a délicate form (young stages only ?) of E. atlantica. Available material consists of

basal parts, unbranched colonies, branched fragments, and elongate straight terminal fragments (Fig.

14 G-I). Compared with Azorean material, the Gorringe specimens hâve slightly smaller dactylopore

dimensions and gastropore diameter, and show no coenosteal luster (presumed to be eroded away).

But some typical Azorean E. atlantica also hâve elongate, slender, nonbifurcating branches very

similar to the Gorringe specimens.

Remarks

The original description of E. atlantica by Hickson (1912b) was brief and not illustrated. This

description was reproduced by Boschma (1967), who also reexamined part of Hickson’s material and

described some additional characters. Having seen original material, Boschma (1967, 1968c)

concluded that E. atlantica was a typical représentative of Errina sensu stricto. In previous papers he

had referred it, on the basis of Hickson’s description, to Lepidopora , then a subgenus of Errina but

later elevated to generic rank by Cairns (1983b).

ridged and spinose, as in the two previously

described species. Dactylopore spines individua-

lized, almost exclusively with groove directed

proximally, and occurring primarily on the an¬

terior face and latéral branch edges of distal

branches and much less frequently on larger

diameter branches. Dactylopore spines quite

short, only about 0.14-0.16 mm tall on distal

branches and virtually flush with coenosteal

surface on larger diameter branches. Width of

dactylopore spines 0.25-0.27 mm; width of

groove 0.09-0.12 mm, or almost one half width

of spine.

Female ampullae hemispherical (Fig. 15 B, F),

about 0.70 mm in diameter with a latéral efferent

pore about 0.20 mm in diameter. Male ampullae

superficial on branch tips, becoming internai on

larger diameter branches; mature ampullae about

0.5 mm in diameter, each with 1-3 apical efferent

pores 50-60 pm in diameter.

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Fig 15 - Errina atlantica (A-G, from “ Jean Charcot ” 1971, stn 240, tonale, usnm 75610): A, branch tip with gastropore

' and several dactylopore spines ( * 37); B. rrerJ'aiivpïv^W^wn^atfvIrnnire'stdnes 79*^1^

pair) ; C, E, reticulate-granular branch coenosteum ( x 35, x 165, re

tranverse section of female ampulla (x 51); G, gastrostyle (x 158, s

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Distribution and ecology

E. atlantica is known from a few stations in the Azores. The depths of the 4 stations of “ Jean

Charcot" 1971, cruise biaçores range between 610 m and 825 m; the depths of the type-locality is

983 m, if correctly recorded. Additional live material obtained by the “ Talisman ” possibly cornes

from a deeper station (1237 m).

Errina from two stations on Gorringe Seamount, off Portugal (460-480 m, 1110-1180 m), could

be the same species.

Symbionts

Larger colonies of E. atlantica from the Azores had Eunice norvegica (Linnaeus, 1767) as a

symbiont (“ Jean Charcot ” 1971, cruise biaçores stn 197, 230, 240; colony mcm 62). Covered by the

coenosteum, the polychaete tube typically extends as an irregular hollow trunk through the greater

part of the colony (Fig. 14 A).

Traces of Pedicularia (Fig. 14 B) hâve been found on branches of E. atlantica from the Azores,

from several stations (colony mcm 62; “ Talisman " drag. 128; “ Jean Charcot " 1971, cruise biaçores

stn 197, 240). Rather large specimens of Pedicularia collected at stn 197 and 240 are likely to hâve

been symbionts of E. atlantica , on which traces are correspondingly large; at stn 240 one large

Pedicularia was found still on the coral in the collection.

Genus STYLASTER Gray, 1831

Diagnosis. — Gastro- and dactylopores arranged in cyclosystems. Cyclosystems variable in

location, ranging from a uniform coverage of ail branch surfaces (Group A) to a strictly sympodial

arrangement (Group C), with many intermediate arrangements (Group B). Coenosteal colour and

texture variable: most common textures reticulate-granular and linear-imbricate. Gastro- and

dactylostyles présent, the latter robust in Group A and moderate to rudimentary in Groups B and

C. Gastrostyles usually ridged and bear long, pointed spines. Ring palisade often présent; gastropore

inner shelf sometimes présent in Group C. Ampullae usually superficial, usually with distinct efferent

pores.

Type species: Madrepora rosea (Pallas, 1766), from the western Atlantic (a member of Group

B).

The 3 groups suggested by Cairns (1983b) are terms of convenience, not meant to imply

taxonomie categories. Cairns (1983b) included Allopora Ehrenberg, 1834 (corresponding essentially

to his Group A), as a synonym of Stylaster.

In the eastem Atlantic, Stylaster is represented by 3 species of Group A (S. norvegicus, S. rosaceus,

and S. blatteus) and 3 species of Group B (S. gemmascens, S. ibericus, and S. erubescens, the latter

with subspecies groenlandicus n. ssp., britannicus n. ssp., and meteorensis n. ssp.). S. maroccanus n.

sp. is not assigned to a group because so little material was available.

Stylaster norvegicus (Gunnerus, 1768)

Fig. 16 A-G, 17 A-I

Synonymy:

Millepora norvegica Gunnerus, 1768: 64-67, pl. 2, fig. 20-22.

Stylaster ( Allopora ) norvegicus forma atlantica Broch, 1936: 49-52, text-fig 14 ol 7 fia 20-21

[?] Allopora oculina Ehrenberg, 1834: 147. 6 '

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Chresonymy:

Allopora norvegica — Sars, 1873: 115-118. — Storm, 1882: 26. — Hickson, 1890: 594. — Norman, 1893:

349 — Boschma, 1956b: F100, fig. 80.5; 1957a: 24-26; 1962: 196-203, text-fig. ld-e, 3a-i, pl. 2, fig. 1-4; 1964e:

109, 115.

Stylaster norvegicus — Broch, 1914a:. 15-19, text-fig. D, pl. 2, fig. 12-15, 18, pl. 3, fig. 23, 27, 31, pl. 4,

fig. 35, 37-38, pl. 5, fig. 44-45. — Nordgaard, 1915: 5. — Cairns, 1983b: 429; 1986a: 57.

Stylaster ( Allopora) norvegicus — Broch, 1928: 55, fig. 46c. — dons, 1939: 197.

Allopora oculina — [ ?] Milne Edwards & Haime, 1850: 99, pl. 4, fig. 4; [ ?] 1857: 132. — Duncan, 1870:

290, 295 (part). — Moseley, 1879: 480; 1881: 85 (part: Norway).

Stylaster gemmascens — Duncan, 1873: 332 (part), pl. 49, fig. 1.

not Stylaster (Allopora) norvegicus — Hickson, 1915: 544-545, pl. 1, fig. 3 (British Columbia). — Broch,

1935: 59, fig. 2 (Okhotsk Sea).

NOT Stylaster (Allopora) norvegicus forma pacifica Broch, 1936: 52-54, text-fig. 15, pl. 6, fig. 18-19

(Okhotsk Sea, British Columbia)

not Allopora norvegica pacifica — Fisher, 1938: 522-524, pl. 53, fig. 2-2b, pl. 54, fig. 1, pl. 76, fig. 3-4

(Okhotsk Sea). — Eguchi, 1941: 1181-1182 (Japan). — Boschma. 1953a: 166, 170-171; 1957a: 26. — Naumov,

1960: 533, fig. 417, pl. 25, fig. 3 (Kurile Islands).

Types

Millepora norvegica : Gunnerus’ (1768) description was based on specimens from Norway.

According to Broch (1914a: 18), syntypes (number of specimens not given) are présent at the vsm.

One of these is figured by Broch (1914a: pl. 2, fig. 12).

Type locality: Gunnerus’ specimens came from Nordmor, west coast of Norway.

Allopora oculina : Ehrenberg’s (1834) description was based on a colony (herein considered to

be the holotype) of unknown origin at the zmb. This has been redescribed in more detail and figured

by Milne Edwards & Haime (1850). The type could not be found in 1984, and may be lost.

Type locality: Unknown (probably Norway, see Remarks).

Stylaster (Allopora) norvegicus forma atlantica : the original rank of Broch’s (1936) forma

atlantica (opposed to forma pacifica) was clearly meant to be that of a geographical subspecies.

However, the North Atlantic subspecies being the nominotypical one, its name must be S. norvegicus

norvegicus; accordingly, Broch’s naming of forma atlantica was an unnecessary taxonomie action.

Material studied

Denmark Strait: “ Ingolf ” stn 52, colony (zmuk). — “ Poséidon ” stn 12/1, ca. 450 branches

+ fragments, only one alive (smf 6470; usnm 88824); stn 14/1, ca. 120 branches + fragments, dead

(smf 6473; usnm 88827).

Iceland or Faroes?: Müller, 28.11.1900, colony (zmuk).

Faroes: “ Dana ” stn 6009, small colony (zmuk). — “Anton Dohrn" 18.11.1973, colony

(imfb). — “ Anton Dohrn ” 19.11.1973, branch (imfb).

Between Faroes and Hébrides: “ Porcupine ” stn 54, 4 branches (bmnh 1880.11.13.4-5/8-9). —

“ Triton" stn 3, several big colonies + fragments (bmnh 1891.5.7.1, 1889.12.14.1, 1980.1.6.1).

Hébrides: off Butt of Lewis, June 1913, 2 colonies + fragments (rsm 1913.113.2).

Rockall: ca. 165 m, J. Cordeaux (bmnh 1896.8.3.3-5; usnm 75620).

Norway: abundant samples in zmuo and vsm, from many localities along the Norwegian coast,

Senja, Steinavaer, Lofoten, Rost, Sklinna, Trondheimsfjord (including Agdenes, Brettingnes),

Fjellvaeroy, Leksa, Kvalholmen/Hemna, Skredeness/ Sunndalsfjord, Bud, Aukra, Giske/Sunnmore,

Âlesund, Storegga (= Havbroen), Hardangerfjord, etc. — Normans Nasa, NW Storegga, 128-183 m,

von Yhlen, colony (smnh 28). — Storegga (= Havbroen), Sars, small colony (rsm 1884.37.158). —

Rodberg, TrondheimsÇord, M. Norman, 1893, 1 fragment (bmnh 1922.2.2.6a). — Norwegian Sea, 2

colonies (rmnh Coel 15387^ — “ ? N Atlantic ”, Frank, 1897, 4 colonies + fragments (rmnh Coel

15388). — Agdenes, Trondheimsfjord, 60-80m, C. Dons, 23.6.1935, branches (rmnh Coel 15381). —

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CA1RNS

Table 2.

— Comparisons of eastern

cyclosystem orientation

(group)

dactylostyle

coenosteal texture;

colour

S. maroccana

(group ambiguous) i

S. norvégiens

random

(group A)

robust

reticulate-granular

but smooth;

white to off white

S. rosaceus

random

(group A)

rudimentary

reticulate-irregular

granular; pink

random

(group A)

rudimentary

reticulate-granular

(granules irregular);

purple

rudimentary

reticulate-granular

white

dactylopores per cyclosystem: 3-10, 6.33, 6 (500)

range, mean, mode (N) 5-11, 6.92 7 (738)

cyclosystem shape and size circular;

0.9-1.1 mm

5-12, 8.57, 9 (70)

circular to slightly

elliptical; 0.6-0.7 mm

6-12, 8.88, 9 (500)

circular to slightly

elliptical; 0.8-0.9 mm

9-14, 11.77, 11 (27)1

circular to elliptical:

1.0-1.4 mm

gastrostyle shape; ridges;

H:W ratio

ring palisade

ampullae: female, male

giobular to conical; not

ridged;

0.9-2.6

absent

9 unknown

c? internai (0.5-0.6 mm)

elongate-conical;

ridged;

3.5

well-developed, éléments

cylindrical

9 superficial (0.7 mm)

unknown

lanceolate; deeply

ridged;

2.0-3.9

well-developed, éléments

vertical carinae

9 superficial

(0.5-0.65 mm)

c? superficial with apical

efferent pores

(0.3-0.45 mm)

elongate cylindrical:

ridged;

5.5

well-developed, elem

vertical carinae

9 superficial (0.45 m:

0.65 mm)

c? unknown

distribution

other diagnostic characters

Denmark Strait to

Rockall and northem

Norway, 75-1400 m

branch tips robust

Gulf of Guinea: Sâo

Tomé,

37 m

isolated dactylopores

présent;

deep slits between

coenosteal strips

Gulf of Guinea: Sâo

Tomé and Principe,

0-10 m

isolated dactylopores

very common; deep slits

between coenosteal

strips

off Morocco (Atlai'j

1378 m

colonies small and

bushy; early sexual

maturity

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

sier and S. erubescens erubescens

r^gënmascens

S. ibericus

S. erubescens

erubescens

S. erubescens

groenlandicus

S. erubescens

britannicus

S. erubescens

meteorensis

primarily on latéral

branch edges but

some on anterior and

(sterior branch

posterior branch

faces

posterior branch

faces

posterior branch

faces

posterior branch

faces

posterior branch

faces

(group B)

jdimentary

robust

reticulate granular

reticulate-granular

coarse granules

reticulate-smooth

but smooth;

(granules irregular in

(granules rounded);

(40-60 pm wide);

(strips with numerous

hite to light yellow

glistening white

shape);

white

symmetrical latéral

protubérances),

unique;

white

[•17,11.76, 12(71)

4-11, 7.71, 8 (685)

7-15, 11.45, 11 (350)

5-14, 10.04, 10 (175)

7-12, 9.34, 9 (270)

9-15, 11.98, 11 (206)

icular to quite

circular to slightly

circular to irregular;

circular to slightly

circular to elliptical;

circular to irregular;

ïgular; e.g.

3mm, 1.6mm *

elliptical;

0.8-1.1 mm

1.3-1.7 mm

elliptical;

1.2-1.4 mm

0.9-1.4 mm

0.9-1.2 mm

lanceolate, broadest

squat; not ridged;

lanceolate; not ridged

lanceolate; broadest

ovate; unridged, very

Iged;

M.7

at mid-height; not

ridged; 1.3-1.9

0.8-1.4

but latéral tiers of

spines; about 2

at mid-height; not

ridged; 1.7-2.0

long spines; 1.1-1.3

S'use, éléments

unded

absent

superficial

$ very low in relief

$ superficial, often

$ superficial, large.

Ç superficial (1.1 mm)

$ superficial, often

7-0.8 mm)

(0.75-1.0 mm)

with efferent tube

sometimes with

cJ low. irregular

with efferent tube

superficial, carinate

65-0.75 mm)

cJ internai (0.5 mm)

(1.0 mm)

c? superficial, with

apical efferent pores

(0.6 mm)

efferent tube, occasio-

nally binary (1.0-

1.3 mm)

J superficial latéral

efferent pore (0.7-

0.8 mm)

mounds, apical

efferent pore

(0.6-0.7 mm)

(0.8 mm)

cJ superficial, apical

efferent pores (0.6-

0.7 mm)

:nmark Strait to

western Atlantic: off off Southern and

off SE Iceland to

seamounts south/

ickall and northern

irway, 40-665 m

450-545 m

SE U.S.A,

146-530 m

eastern Greenland to

Iceland, 326-713 m

Celtic Sea,

350-1080 m

Southwest of Azores,

293-500 m

:nosteal papillae

t ucommon, espe-

lly on or near am-

llae; cyclosystems

red;

fferent pore large

host of Pedicularia

colonies uniplanar,

branches often

thickened; host of

Pedicularia and

polynoid polychaete

colonies uniplanar,

branches often

thickened

colonies uniplanar,

branches moderate in

thickness; deep coen-

osteal slits on pseu-

dosepta (Fig. 30 E);

host of Pedicularia

colonies bushy,

branches slender

«M

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

l ^mZ e œüC 8i Z^ mï Trondhe.rnsljordnofurther details, vsm; C, from Trondheimsfjord, Brettingnes,

BaiZ n f r ™^ p ; 9 9 ' D ’J rom He bndes, rsm 1913.113.2; E-F. from Lofoten, Rest! Kunstmaler

cvlindricàl branches (x 1 5V B ^hhint tram-h . aMNH r A - slightly bushy colony with rather slender

^cZ7np naUencd near thehit I n - n " P ° f A , Wlth c >; clos y st . e ™ < x l7 )i C, larger colony with branches

teseTxl OV E colonv withhlnn, gef ? nlplanar colon y Wlth branches nattened and coalescent near the

s iyf F ' br ™ h ° r *■

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Norway, colony received from Hickson (mnhn). — Norway, colony (nhmw 7970). — Norway,

fragment received from G.O. Sars (usnm 6854, erroneously labelled as cotype). — “ Anton Dohrn "

10.3.1972, large colony at imfb known only from photograph.

Description

Colonies mostly uniplanar, up to 10 cm high

and 15 cm wide (Fig. 16 A, C-D, F). Branches

cylindrical and robust, 2-3 mm in diameter at

blunt distal tips. Coenosteum white to off-white

and smooth, a resuit of a reticulate-granular

texture with very low relief granules. Figure 17 F

shows a transition from an early stage of coe-

nosteal development characterized by irregularly

shaped granules to the more advanced, smooth

stage in which the granules are united into strips

and covered with a smooth coenosteum. Coenos-

teal strips 45-75 pm wide. Cyclosystems circular,

0.9-1.1 mm in diameter, occurring with equal

frequency on ail branch faces. Cyclosystems

flush with coeosteum or raised only slightly

above coenosteum. According to Boschma’s

(1962) analysis of 500 cyclosystems, there is a

range of 3-10 dactylopores per cyclosystem,

mean 6.33, and mode 6. Our analysis of 738

cyclosystems revealed a slightly higher number of

dactylopores per cyclosystems: range 5-11, mean

6.92, and mode 7. There are no diastemas.

Gastropore tubes 0.7-2.2 mm long, slightly

constricted at level of gastrostyle tip; no ring

palisade (Fig. 17 G). Gastrostyle (Fig. 16 F)

occupies a variable percentage of gastropore

tube, depending on height of style and length of

tube (see Boschma, 1962: fig. 3). Gastrostyle

variable in shape, ranging from rotund (wider

than high) to conical, with H:W ratios ranging

from 0.9-2.6. Tallest known gastrostyle 0.75 mm;

widest, 0.52 mm (Boschma, 1962). Gastrostyles

covered by extremely long, cylindrical, blunt

spines up to 110 pm long and 15 pm in diameter,

which are sometimes laterally fused into tiers

(Fig. 17 H). Ridging of gastrostyle, if présent,

obscured by tall and abundant gastrostyle spines.

Dactylotomes 0.10-0.15 mm wide. Dactylostyles

(Fig. 17 C) robust, composed of blunt cylindrical

éléments much like gastrostyle spines, each élé¬

ment up to 75 pm tall and about 11 pm in

diameter. Dactylostyles up to 0.6 mm long and

65 pm wide (4 or 5 éléments across width),

terminating about 0.2 mm from the tip of

dactylopore.

Ampullae (male ?) internai to slightly

superficial (Fig. 17 E), 0.5-0.6 mm in internai

diameter. Efferent pores round, about 0.13 mm

in diameter. Female ampullae unknown.

Comparisons

5. norvegicus is easily distinguished from the other two eastern Atlantic species of Stylaster

(Group A), by a large number of characters (Table 2). Within the Atlantic, it is most similar to

S. miniatus (Pourtalès, 1868), known from off the southeast coast of the United States from 146-

530 m (Cairns, 1986a). Points of similarity include: cyclosystem diameter, gastrostyle ornamentation

and H:W ratio, dactylostyle shape, and characteristics of the male ampullae. Nonetheless, S.

norvegicus can be distinguished by its coenosteal texture and higher number of dactylopores per

cyclosystem.

Remarks

S. norvegicus is recognizable in Gunnerus’ (1768) illustrated description from Norway (as

Millepora). Broch (1918) commented upon this description and reproduced some of the figures.

Referring to a correspondence he had with Linnaeus, Gunnerus (1768: 67) added the erroneous

remark that his Millepora norvegica was the same as Millepora aspera Linnaeus, 1767 (for which the

distribution had been given as Mediterranean and Norwegian Sea). This was a misunderstanding

(Boschma, 1953a, 1953b, 1965a); in fact, Linnaeus’ coral (now known as Errina aspera) is a

Mediterranean species and does not occur along the coast of Norway.

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

fragmeit musttati5'1ïo <A c°ios y sMs?m°™Fà” 0 Sj | ^^ m f R “ ka J'’ «««« 1896.8.3.3-5): A, branc

SS2^t^gS5>a^|SsS S3PSïSSS&?r? , Sf

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Broch (1936) erroneously included Allopora miniata Pourtalès, 1868, in the synonymy of S.

norvegicus. As Boschma (1962) had already argued, Broch’s reference specimens were not authentic

western Atlantic A. miniata, but mislabeled northeastern Atlantic S. norvegicus ; in fact, Broch had

not seen the single authentic small branch of A. miniata in the zmuo. Typical A. miniata (redescribed

by Cairns, 1986a, as Stylaster miniatus) differs from S. norvegicus in various aspects, such as surface

structure, depth of gastropores, higher number of dactylopores (see Comparisons).

Allopora oculina Ehrenberg, 1834, of unknown origin, is most likely a synonym of S.

norvegicus. The original description and a redescription by Milne Edwards & Haime (1850) may

apply to the Norwegian stylasterid, which was easily available to the older authors. A. oculina, as

repeatedly reported from Norway in the 19th century, should be referred to Stylaster norvegicus. A.

oculina sensu Studer (1878), from South Africa (zmb 1654), is very different, close to Stylaster

bithalamus Broch, 1936.

The first adéquate description of S. norvegicus, according to modem standards, was given by

Broch (1914a) from Norwegian and other northern Atlantic material (including Gunnerus’ types).

Norwegian material was again analyzed in detail by Broch (1936) and Boschma (1962).

S. norvegicus, often attributed to the genus Allopora, is mentioned in various publications on

the fauna of Norway, where 5. norvegicus and S. gemmascens frequently occur together but hâve not

always been distinguished; many lots in muséum collections under one or the other name hâve proved

to be a mixture of both species.

S. norvegicus is one of four species reported by Duncan from between the Faroes and the

Hébrides, first (1870) as Allopora oculina, then (1873) as Stylaster gemmascens (the other species are

S. gemmascens, S. erubescens britannicus, and Stenohelia maderensis).

Stylasterids from the northern Pacific (Japan to British Columbia) hâve occasionally been

referred to Stylaster (or Allopora) norvegicus, mostly as a distinct forma or subspecies pacifica as

opposed to atlantica (Hickson, 1915; Broch, 1935, 1936; Fisher, 1938; Eguchi, 1941; Naumov,

1960). One of these records (British Columbia) was subsequently reidentified as Stylaster verrillii Dali,

1884 (Fisher, 1938: 524; Broch, 1942: 6 ). Likewise, Cairns (1983b: 429) tentatively included Allopora

norvegica pacifica Broch, 1936, in the synonymy of S. verrillii.

Distribution and ecology

S. norvegicus is known from many stations in a wide area of the North Atlantic: from

Denmark Strait, the northwest, Southwest, and southeast of Iceland, Faroes and Hébrides, Rockall,

and along the coast of Norway. Its occurrence in Icelandic waters, already reported by Broch

(1914a), is here confirmed. The northernmost records are 66°18'N in Denmark Strait, and 69°14'N

off Norway.

In Norway the species is common in depths of 80-300 m, frequently found together with the

bank-forming scleractinian Lophelia pertusa. Elsewhere, the shallowest record is from 75 m in the

Faroes. In the Iceland — Faroes area the species has been obtained as deep as 1040 m and 1400 m.

No symbionts are known.

Stylaster rosaceus (Greeff, 1886)

Fig. 18 A-H

Synonymy:

Allopora rosacea Greeff, 1886: 19-20.

Chresonymy:

Allopora rosacea — Broch, 1914b: 38-40, text-fig. 12, pl.l, fig.4-5. -

Stylaster rosaceus — Cairns, 1983b: 429.

- Boschma, 1957a: 27; 1961: 219-220.

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

* %&B2£SSZ

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Types

Allopora rosacea : In the original unillustrated description, Greeff (1886) mentioned several

small colonies up to 35 mm high. Having not been given a précisé type status, they are considered

as syntypes. After Greeff’s death at least part of his collections were transferred from the Zoological

Institute, Marburg, to the Zoological Muséum, Hamburg, where Broch (1914b) was able to study

one of the syntypes. This specimen was lost when the muséum burnt in 1943. Another smaller female

syntype (12 mm tall, 13 mm broad) exists at the zmb (4725), probably deposited there by Greeff

himself. This is the only specimen known to be preserved (Fig. 18 F-H).

Type locality: Passage between Sào Tomé and Ilha das Rolas, Gulf of Guinea, depth ca.

37 m (R. Greeff, 1880, dredge).

Material studied

Sâo Tomé: 1 syntype (ZMB 4725).

Description

The only specimen available (Fig. 18 F-H) is a

slender incomplète colony (most branch tips

broken), 12 mm tall, uniplanar, with a thin

encrusting concave base about 8 mm wide, the

latter originally encrusting a cylindrical subs¬

trate. On the same base, next to the main stem,

two smaller branches are broken away. Branches

cylindrical, gradually tapering to slender branch

tips the diameter of a cyclosystem. Coenosteum

pink, mature (female) ampullae less pigmented to

whitish; branch tips may also be of lighter

colour. Coenosteum reticulate, strips 55-80 pm

wide, but not granular, rather covered with

irregularly shaped elongate carinae about 30 pm

long and 7 pm wide (Fig. 18 D-E).

Cyclosystems occur on ail branch surfaces;

they are circular to slightly elliptical, 0.6-0.7 mm

in diameter, almost flush with coenosteum. No

diastemas observed. Based on 70 cyclosystems

from encrusting base and branches, the range is

5-12 dactylopores per cyclosystem, mean 8.57,

and mode 9.

COMPARISONS

Based on the single preserved syntype, S. rosaceus differs from the better known sympatric S.

blatteus in the following characters: colour pink versus purple; slender branches; smaller cyclosystem

diameter; cyclosystems flush and widely spaced versus slightly exsert and denser; dactylotomes open

apically versus opening into gastropore tube; diastema absent versus occasionally présent; ring

palisade consisting of cylindrical éléments versus vertical carinae; gastrostyle spines tall and slender

(e.g. 41 x 7 pm) versus short and blunt (e.g. 25 x 12 pm).

Coenosteal texture, gastrostyle, and range, mean and mode of dactylopores per cyclosystem

are not significantly differefit in the two species. However, the impression of a similar coenosteal

texture is based on only a very small piece of the basal encrustation of S. rosaceus used for sem study.

Gastropore about 0.2 mm in diameter, with a

robust ring palisade (Fig. 18 C) composed of

cylindrical éléments about 25 pm in height and

diameter, not vertical carinae. Gastrostyles

elongate-conical (Fig. 18 C), about 0.35 mm tall

and 0.10 mm in diameter (H:W = 3.6), occup-

ying basal half of gastropore tube. Gastrostyles

vertically ridged, each ridge bearing slender

elongate spines up to 41 pm long and

8 pm in diameter. Dactylotomes about 70 pm

wide, separated by low pseudosepta 1-1.5 times

width of dactylotomes; no diastemas. Dactyloto¬

mes open apically, rarely directly into gastropore

chamber. Isolated dactylopores présent but

sparse. Dactylostyles not examined.

Female ampullae superficial hemispheres

about 0.70 mm in diameter, clustered on both

anterior and posterior faces. Efferent pore about

0.15 mm in diameter. Male ampullae unknown.

Source : MNHN, Paris

HELMUT Z1BR0WIUS & STEPHEN D. CAIRNS

Greeff’s syntypes of S. rosaceus (only one of which is preserved) are the only pink stylasterids

from the Gulf of Guinea, thereby strangely contrasting with the many specimens of S. blatteus (types

included) available for the présent study, ail of which hâve the purple colour described by Boschma

(1961). If pink S. rosaceus and purple S. blatteus are just colour variations of one species, they would

be expected to occur together, but this apparently is not the case. Some other stylasterid species do

show colour variations, generally even among material from one station, e.g. the South African

Errina diffusa Boschma, 1963b, which may be either white or pink.

The occurrence of S. rosaceus at a depth of about 37 m may also point to a spécifie différence

from S. blatteus, which is always collected from infralittoral rocks at depths not exceeding 10 m.

Remarks

Specimens corresponding to the pink S. rosaceus as described by Greeff (1886, under

Allopora) hâve never been subsequently collected. Broch (1914b) reproduced Greeff’s description and

added a few details on the surface structure and on the shape of the cyclosystems. In order to preserve the

single available syntype undamaged, he forsook any detailed study that required dissection.

According to Greeff (1886), the main characters distinguishing S. rosaceus from S. blatteus

are the colour (pink in rosaceus, deep violet or wine-red in blatteus) and the absence of isolated

dactylopores between the cyclosystems. Greeff was wrong on the second point: we found isolated

dactylopores on the only preserved syntype. Broch (1914b) mentioned a différence of the surface

structure: in S. rosaceus roughly reticulate, the small “ fields ” being separated by “ stripes ” of lighter

colour; in 5. blatteus with a fine punctuation (but most specimens of the latter were beach worn).

Indications on the cyclosystems by both authors (width, number of dactylopores) are apparently

based on a rough comparison and only a few countings. Cyclosystems of S. rosaceus are said to be

less regularly circular, occasionally also deformed by adjacent ampullae. There are in fact some less

regular cyclosystems on the remaining syntype, but this should not be considered significant;

irrégularités are also observed in S. blatteus.

Distribution and ecology

S. rosaceus is known only from one record at Sào Tomé (type locality), depth ca. 37 m.

No symbionts are known.

Stylaster blatteus (Boschma, 1961)

Fig. 19 A-J, 20 A-F

Synonymy:

Allopora blattea Boschma, 1961: 210-221, pl. 3-6.

Chresonymy:

Allopora subviolacea — Greeff, 1886: (11-13)16-19. — Broch, 1914b: 40-41, pl 1 fie 6-7 —

Boschma, 1957a: 28-89 (part).

Allopora blattea — Vervoort & Zibrowius, 1981: 27.

Stylaster blatteus — Cairns, 1983b: 429.

Types

Allopora blattea: The original description by Boschma (1961) was based on many specimens

from several localities at Principe and Sâo Tomé, islands in the Gulf of Guinea. One of these colonies

(“ Cal yP so " 1956, stn 70, Sào Tomé) was designated by Boschma (pl. 3, fig. 1-2) as holotype, ail

other colonies from the same and other localities (“ Calypso ” 1956, stn t-25, stn 67, stn 74, stn 111;

Sao Tome, I Marche-Marchad, 1956) as paratypes (pl. 4-6, and other unfigured specimens)!

Holotype and most paratypes at mnhn. Additional paratypes at rmnh (“ Calypso ” 1956, stn t-25,

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Fig. 19. - Stylaster blatteus (A-C. from Praia das Conchas, Sào Tomé, male, usnm 75614; F-H, same locality, mnhn; D-E,

from Sào Tomé. coll. cpas, mnhn; I-J, from Principe, Ilheu dos Mosteiros, coll. J. Laborei.. mnhn): A, gastrostyle and

ring palisade ( x 169); B, dactylostyle ( x 253); C, male ampulla with apical efferent pore ( x 63); D, robust colony with

short stout branches ( x 1.2); E, detail of D illustrating cyclosystems and ampullae ( x 4.3); F-G, anterior and posterior

face of colony with rather slcnder branches (both x 0.6); H, slender branch of another colony ( x 4.4); I-J, young

colonies ( x 2.9, x 2.0. respectively).

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

dry colony Coel 13750A, alcohol preserved material Coel 13750B; Sào Tomé, I. Marche-Marchad,

1956, 2 colonies Coel 13902) and usnm (“ Calypso ” 1956, stn t-25, 2 fragments 75613). In addition,

the rmnh has fragments from “ Calypso ” 1956, stn t-25 and stn 70 that hâve been used by Boschma

to make préparations, but were not given type status (Vervoort & Zibrowius, 1981: 27).

Type locality: Holotype (and several, but not ail paratypes) from “ Calypso ” 1956, stn 70,

19.6.1956. Praia Santa Catarina on the west coast of Sâo Tomé, 3-10 m.

Material studied

Ail specimens listed by Boschma (1961) from the “ Calypso ” cruise in 1956 to the islands in

the Gulf of Guinea (see Types), and additional more recently collected specimens.

Principe: “ Calypso ” 1956, stn 111, 7.7.1956, Uheu dos Mosteiros, 3-10 m, 8 colonies

(paratypes). — Ilheu dos Mosteiros, overhang 4-6 m, 18 small colonies, J. Laborel, Jan. 1971 (most

mnhn; usnm 77124).

Sào Tomé: “ Calypso ” 1956, stn t- 25, 15.6.1956, Punta Furada, 3-8 m, 4 colonies + several

fragments (paratypes); stn 67, 18.6.1956, Punta Diogo Vaz, 6-10 m, 3 colonies (paratypes); stn 70,

19.6.1956, Praia Santa Catarina, 3-10 m, 8 colonies + several fragments (holotype + paratypes); stn

74, 21.6.1956, Sào Miguel bay, 6-10 m, 2 colonies (paratypes); same cruise, précisé locality not given,

I. Marche-Marchad, several small branches + fragments (paratypes); same cruise, northwestern

coast of Sâo Tomé, branch. — Sâo Tomé, without further details, colony collected by divers of

Centro Portugues de Actividades Subaquaticas [ =

Conchas, Guadalupe, 10 m, S. Gofas, Nov. 1983,

Description

Colonies robust and primarily uniplanar (Fig.

19 D, F-G): holotype 10.7 cm high and 10.5 cm

wide, with a massive basal branch and encrusting

base. Branches cylindrical, their distal part fol-

lowing last ramification still rather stout before

tapering to tip the diameter of a cyclosystem.

Coenosteum purple, reticulate in texture, the

strips 35-55 pm wide and separated by deep slits

up to 11 pm wide. Strips covered with irregularly

shaped pointed granules and short carinae; the

granules as small as lOpm in diameter, the

carinae up to 25 pm long and about 7 pm wide

(Fig. 20 B).

Cyclosystems occur on ail branch surfaces.

They are round to slightly elliptical, 0.8-0.9 mm

in diameter, rather exsert on their proximal

margin and occasionally develop a narrow dias-

tema (Fig. 20 A). Based on 500 cyclosystems,

Boschma (1961) found a range of 6-12 dactylo¬

pores per cyclosystem, mean 8.88, and mode 9.

Gastropore about 0.3 mm in diameter and

0.65-1.55 mm deep, with a well-defined ring

palisade at level of gastrostyle tip. Ring palisade

composed of 12-15 vertical carinae (Fig. 20 E-F)

up to 80 pm long and 15 pm wide arranged

parallel to one another. Gastrostyles 0.30-0.65

cpas], transmitted by L. Saldanha. — Praia das

7 colonies (most mnhn; usnm 75614; bmnh).

mm tall and 0.13-0.24 mm wide (H:W = 2.0-

3.9), occupying basal 40-60 % of gastropore.

Gastrostyle deeply ridged, bearing short, blunt

spines about 25 pm long and 12 pm in diameter.

Well developed high dactylotomes about 73 pm

wide, opening straight into steep-walled dactylo-

pore tube devoid of shelf; pseudosepta one to

four times width of dactylotomes. Narrow distal

diastemas occasionally présent, especially on

large diameter branches. Circular to irregularly

shaped isolated dactylopores very common (Fig.

20 C), 60-90 pm in diameter. Dactylostyles

rudimentary (Fig. 20 B), composed of a single

line of well-spaced cylindrical to clavate élé¬

ments, each up to 35 pm tall and about 12 pm in

diameter. Isolated dactylopores do not hâve

dactylostyles.

Female ampullae (Fig. 20 C-D) smooth,

superficial hemispheres 0.50-0.65 mm in diame¬

ter, each with an efferent pore about 0.11 mm in

diameter. Male ampullae (Fig. 19 C) also

superficial, 0.30-0.45 mm in diameter, with 1-3

apical efferent pores, each about 22 pm in

diameter. Ampullae of both sexes often clustered

(Fig. 19 E, H, 20 D) on both anterior and

posterior faces.

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Fig. 20. — Stylaster blatteus (A, B-F, from Praia das Conchas. Sào Tomé, male and female respectively usnm 75614): A

branch segment showing cyclosystems and isolated dactylopores ( x 14); B, coenosteal texture ( x J28, .nf-'n

C-D, coenosteum illustrating cyclosystems, female ampullae. and isolated dactylopores ( x 32, x 24. respectively, D

being a stéréo pair); E-F, gastrostyle and ring palisade (x 164, x 64, respectively, F being a stéréo pair).

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

COMPARISONS

Boschma (1961), who knew S. rosaceus only from the literature, questioned whether Greeff’s

(1886) pink S. rosaceus and the sympatric purple S. blatteus were distinct species.

We conclude that these forms are different species: S. blatteus has purple coenosteum; stouter

distal branches; and more densely crowded and exsert larger cyclosystems with higher dactylotomes.

Other structures of both species are similar, such as coenosteal texture and gastrostyle, as well as

range, mean and mode of dactylopores per cyclosystem.

Young colonies of S. blatteus (Fig. 19 I-J) similar in size to the only preserved syntype of S.

rosaceus are different in aspect, being stouter with more crowded cyclosystems.

Remarks

Greeff (1886), the first collector of the purple stylasterid from the Gulf of Guinea, mistakenly

referred it to Allopora subviolacea Saville Kent, 1871, at that time known only from the type from

unknown origin. Broch (1914b), who reexamined part of Greeff’s specimens (beach worn pièces

only), had little to add to the previous description.

Boschma (1961) finally had access to abundant new material from the Gulf of Guinea and

described it as Allopora blattea (the name meaning purple), while his student Goedbloed (1962a, b)

studied the dactylozooids and gonophores of this species. Boschma (1966) also examined the type of

A. subviolacea and redescribed this species from new material from South Africa. Having seen ail

samples studied by Boschma, as well as additional specimens from the Gulf of Guinea and South

Africa, we confirm Boschma’s conclusion on the obvious différence of the two species (now absorbed

into the genus Stylaster).

Distribution and ecology

S. blatteus is known only from the two central islands in the Gulf of Guinea, Principe and Sâo

Tomé. The presence of the stylasterid is best documented for Sâo Tomé, which has been more

intensively investigated. Ail records are from shallow water (depth not exceding 10 m), especially

from steep cliffs, under overhangs, and in vertical fissures of zones exposed to strong currents

(Forest, 1959: 10).

No symbionts are known.

Stylaster maroccanus new species

Fig. 21 A-D, 22 A-G

Types

Type lot comprising 8 specimens collected dead, 3 of them representing lower parts of colonies

with the preserved base, the other 5 being colonies without base or detached branches. Holotype

largest specimen, colony about 8 mm high, without base, with 8 cyclosystems (Fig. 21 A-B). The other

7 specimens are paratypes. Holotype and most paratypes at mnhn; 1 paratype at usnm (77125).

Type locality: “ Cryos ” cruise balgim, stn cp-95, 8.6.1984, 34°24.7'N, 7°39.3'W, 1378 m. Off

Atlantic coast of Morocco.

Material studied

Morocco: holotype and paratypes (see above).

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Fig. 21. — Stylasier maroccanus (A-D. from “ Cryos ” cp-95, mnhn): A-B, opposite views of female holotype illustrating

ciustering of ampullae on posterior face (both x 9.0); C-D, paratypes ( x 9.0, x 13 , respectively).

Description

The few fragmentary specimens available sug-

gest a délicate, bushy corallum. Largest available

specimen has only 8 cyclosystems. Branches

cylindrical; 0.5-0.7 mm in diameter. Ail speci¬

mens were dead when collected, with poorly

preserved coenosteum, some specimens appa-

rently suffering from bioerosion. Coenosteum

white and reticulate-granular in texture (Fig. 22

F), the strips about 70 pm wide. No nematopores

or isolated dactylopores apparent.

Cyclosystems sympodially arranged on some

branches or parts of branches and restricted to

anterior face of other branches. Cyclosystems

circular to slightly elliptical in shape, 1.0-1.4 mm

in diameter. Based on 27 cyclosystems (ail avai¬

lable), there is a range of 9-14 dactylopores per

cyclosystem, mean 11.77, and mode 11. Some

cyclosystems hâve a narrow diastema.

Only one poorly preserved gastrostyle was

examined (Fig. 22 C): it is 0.40 mm high and

0.073 mm wide (H:W = 5.47) and not ridged.

Little more detail is known of the gastrostyle;

however, the ring palisade is well developed.

Elongate éléments up to 0.1 mm long, 30 pm

wide, and 30 pm high encircle upper third of

gastrostyle (Fig. 22 G). Gastrostyle tip extends

slightly beyond the ring palisade zone into the

slightly expanded upper gastropore chamber.

Dactylotomes about 65 pm wide; pseudosepta

1-3 times width of dactylotomes and often

slightly concave, especially on their outer edges.

Dactylostyle rudimentary, composed of a single

line of cylindrical éléments.

Female ampullae (Fig. 22 A, D) elliptical in

outline, about 0.65 x 0.45 mm, the circular latéral

efferent pore at one of the vertices of ellipse.

Female efferent pore 0.12 mm in diameter. Male

ampullae unknown. Mature female ampullae

présent on very small branches, indicative of

early maturation and/or small colony size.

Comparisons

S. maroccanus is a very distinctive species among the eastem Atlantic Stylaster (Table 2) based

on its cyclosystem orientation', elongate gastrostyles, and very small corallum size. It is perhaps most

similar to S. blatteus, both having similar dactylostyle and ring palisade morphology; however, it is

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

F,G ' ^^sever^^amp^illa^^'^^stereo palrV^B 6 to^view^nfr ” fe ” a,e > ” SNM 11 cyclosystem surrounded by

ampullae and eflerent pore above^yclosystem ( x 46V C , calclfi . ed lo . wer gastroporc tube, female

pair); D. female ampdla and efferent ST>< 87?È bnSinaT fmcînrf of7!'f a " d nng pa ! sade ( X 146 i s,e f reo

(x 50); F. poorly preserved coenosteum sLwing LeVosion t Vw i 23) r, won Jt?^ H rCVeal,n f- gaStr0S,yle of F

removed ( x 113). ë osion ( ' ( ui >’ well-developed nng palisade. gastrostyle

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STVLASTERIDAE

easily distinguished by its coenosteal color, larger cyclosystems, and greater average number of

dactylopores per cyclosystem, as well as by a very different depth and géographie range.

Remarks

The spécifie name given to the new species refers to its géographie distribution.

Distribution and ecology

Species known from one station only (type locality), off the Atlantic coast of Morocco, depth

1378 m.

No symbionts are known.

Stylaster gemmascens (Esper, 1794)

Fig. 23 A-H, 24 A-F

Synonymy:

Madrepora gemmascens Esper, 1790: pl. 55, fig. 1-2; corresponding text 1794: 60.

Chresonymy:

Madrepora virginea — Gunnerus, 1768: 56, pl. 8, fig. 2-4.

Oculina gemmascens — Ehrenberg, 1834: 303.

Allopora gemmascens — Dana, 1848: 696.

Stylaster gemmascens: Milne Edwards & Haime, 1850: 98; 1857: 130-131. — Saville Kent, 1871: 281.

— SARS, 1873: 115. — Duncan, 1873: 332 (part), pl. 49, fig. 4-7. — Storm, 1879: 24; 1882: 25-26. — Moseley,

1879: 480; 1881: 86. — Norman, 1893: 349. — Thomson, 1910: 61. — Ritchie, 1912: 281. — Nordgaard, 1912:

7. — Arndt, 1913: 122. — Broch, 1914a: 8-12, text-fig. C, pl. 1, fig. 4-7, pl. 2, fig. 16, pl. 3, fig. 21, 24-26, 30-31,

pl. 4, fig. 32-33, pl. 5, fig. 46, 49-50; 1918: 9, fig. C. — Dons, 1932: 15. — Boschma, 1955a: 22-31, text-fig. 1-3,

pl. 1-2; 1956b: F98; 1957a: 10-11; 1958: 71-72. — Bourdon-Jones & Tambs-Lyche, 1960: 7.— Cairns, 1983b:

430.

Stylaster ( Eustylaster ) gemmascens — Dons, 1939: 197.

Allopora oculina — Duncan, 1870: 290, 295 (part).

Stylaster sanguineus — Thornely, 1897: 81.

not Stylaster gemmascens — Hickson & England, 1905: 12 (Sulu Islands).

not Stylaster cf. gemmascens — Eguchi, 1941: 1176 (Japan).

not Stylaster gemmascens alaskanus Fisher, 1938: 500-501, pl. 47, pl. 48, pl. 54, fig. 2 (Aleutian Islands).

— Boschma, 1953a: 166; 1957a: 11. — Naumov, 1960: 534, fig. 430-431 (Kurile Islands).

Types

Madrepora gemmascens : As shown by Grasshoff & Scheer (1991) Esper’s description (1794)

was published after the illustration (1790), but opinion No. 574(3b) of the International Commission

on Zoological Nomenclature (1959) had already rejected ail uses of the binomen Madrepora

gemmascens prior to Esper (1794). It was based on presumably only one specimen. Boschma (1955a)

described and figured in detail the specimen found in Esper’s collection and regarded it as the

“ type ”; this colony is massive, 10 cm high, 8.5 cm wide, 1.7 cm basal branch diameter (Fig. 23 C-D)

and is a male, not a female as suggested by Boschma. In a later publication (Boschma, 1958) it was

referred to as the “ holotype ” and also “ selected as the lectotype ”. It is now deposited at the SMF,

where it had been transferred from Erlangen (Vervoort & Zibrowius, 1981: 10).

Type locality: According to Esper (1794), his material came from the East Indian Sea,

information which was reiterated by, among others, Milne Edwards & Haime (1850, 1857). The

origin given by Esper was certainly incorrect: the species is known only from the North Atlantic.

Most likely, Esper’s type came from Norway, where the species occurs in relatively shallow water,

and where it was already known to Gunnerus (1768). Accordingly, Norway should be considered as

the type locality.

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

F '°' Ü'bT W,»*" Harfangeriiord. „ale, „ S »„ ,5618;

1909.191.13): A, lemale àmpuÙa S'ŒS.-JTÏ màfc'.™,,. S T" C °" '- 5 ?S «• F >™>- «w

( x 0.6): D. detail from C iHustrating cyclosvstem shane anHmflp fm P n 25, jt re0 pair); C ’ massive male colony

and posterior faces of 5n, ÏÏÏÏÏ î? P a P illae < « 3.0) ; E-F. anterio,

branch sepnen, il.n.babng cyclosys.en,, and <*’ it

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Material studied

Denmark Strait: “ Ingolf ” stn 94, many small branches + fragments, dead (zmtjk). —

“ Poséidon ” stn 12/1, ca. 430 branches + fragments, dead (smf 6471; usnm 88825); stn 14/1, ca. 70

branches + fragments, dead (smf 6474; usnm 88828).

Faroes: off Faroes, taken by fishing boat, colony + fragment (RSM 1909.191.3). — “ Michael

Sars ” 1902, stn 43, long line fishing, branch (zmuk). — “ Dana ” stn 5835, 3 colonies on pebbles

(zmuk); stn 6005, branch (zmuk). — “ Ingolf ” stn 1,2 branches (zmuk); stn 2, 2 branches (zmuk).

Between Faroes and Hébrides: “ Porcupine ” 1869, stn 54, 4 branches (bmnh 1880.11.13.1-3/7).

Rockall: “ Granuaille ” 1896, branch + fragment (= Stylaster sanguineus sensu Thornely,

1897; bmnh 1898.6.6.1). — J. Cordeaux, 2 colonies (bmnh 1898.8.25.1). — “ Pisces III" dive 73-5,

fragment of a big photographed colony.

Norway: Various old muséum specimens of confused origin are likely to corne from Norway.

Holotype of Madrepora gemmascens (smf). — Colony labeled “ Mer des Indes ” (mnhn). — Colony

labeled “ Océan Indien ” (rmnh Coel 15806). — Abundant samples in zmuo and vsm, from many

localities along the Norwegian coast, Hjelmsoystauren, Malangen, Steinavaer, Brettesnes/Lofoten,

Sklinna, Rodoy, Trondheimsfjord (including Brettingnes, Hysnes, Roberg, Stornesset, Tômmerda-

len), Mefjordsgrunnen/Beian, Sunde/Trondelag, Kristiansund, Skredeness/Sunndalsfjord, Bud, Gis-

ke/Sunnmore, etc. — Trondheimsfjord, Kroyer, colony (nhrm 33). — Rodberg, Trondheimsfjord,

ca. 300 m, T. Mortensen, 27.7.1911, colonies (zmuk). — Trondheimsfjord, 40-170 m, 31.8.1961, 10

colonies + branches (rmnh Coel 17414). — Rodberg, Trondheimsfjord, 150-300 m, “ Gunnerus ”,

16.3.1926 (rmnh Coel 15335). — Bud, 200 m, “ Gunnerus ” 6.7.1931, 3 branches + fragments (rmnh

Coel 15338). — Norway, deep water, colony (bmnh 1896.7.31.1). — Trondheimsfjord, 457 m, C.

Bovallius, several colonies + branches (bmnh 1891.5.11.1-3). — Rodberg, Trondheimsfjord, A.M.

Norman, 1893, several colonies (bmnh 1898.5.7.15). — Rodberg, Trondheimsfjord, A.M. Norman,

1893 (bmnh 1922.2.2.6b). — Hardangerfjord, A.M. Norman, 1879, ca. 20 colonies on pebbles +

branches (most bmnh 1910.10.1.71, 1912.11.30.4-6; usnm 75618). — Norway, colony + 2 branches

from Univ. Copenhagen (ypm 1358). — Norway, 2 colonies (usnm 15275). — No locality, female

colony (usnm 52229).

Description

Colonies uniplanar to slightly bushy (Fig. 23

C, E-F, H), up to at least 11 cm high and wide.

Branches cylindrical; branch tips are diameter of

terminal cyclosystems. Coenosteum white to

light yellow, reticulate-granular in texture (Fig.

24 C-E). Coenosteal strips uniformly 75-85 pm

wide and covered with granules irregular in size

and shape, producing a rough texture. Coenos¬

teal papillae (specialized nematocyst structures

?) common on some specimens, particularly

around ampullae (Fig. 23 B, G). Papillae cylin¬

drical or carinate: up to 0.8 mm long, 0.2-0.4 mm

high, and about 0.10 mm wide.

Cyclosystems occur primarily on latéral edges

of distal branches in a sympodial manner, but on

larger diameter branches they occur with greater

frequency on anterior and posterior branch faces

(Fig. 23 C-F). Cyclosystems cirular to irregular

in shape, some quite compressedi' circular cy¬

closystems about 1.3 mm in diameter; a compres-

sed cyclosystem might measure 1.6 x 0.7 mm.

Proximal dactylopores of cyclosystem project up

to 0.8 mm, whereas distal dactylopores usually

flush with coenosteum, tending to give the

cyclosystem a proximal orientation. Based on

100 cyclosystems examined from the holotype,

Boschma (1955) found a range of 6-15 dactylo¬

pores per cyclosystem, mean 10.19, and mode 10.

Based on 71 cyclosystems examined from five

other localities, we found a range of 7-17 dacty¬

lopores per cyclosystem, mean 11.76, and mode

12. Boschma's slightly lower average and range

may be attributable to his counting cyclosystems

in which the distal dactylopores had become

obsolète (diastema).

Lower gastropore chamber narrow and cylin¬

drical, about 0.30 mm in diameter, which gives

the cyclosystem a flared aspect. Diffuse ring

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

fâTwSâZnm(i ÎFx'zf i £~i&’ÿîl£?ï 7 t 5618: D '- \'r°? “ Dknown locaIit ï.

« , 2 , « d. f. .«u &*. a.’SB&îdra ^js&^srssssr,^

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

palisade présent at mid-gastrostyle level, compo-

sed of éléments about 30 pm in height and

diameter. Gastrostyle lanceolate; figured style

(Fig. 24 D-E) 0.56 mm high and 0.15 mm in

diameter (H:W = 3.7). Boschma (1955) illustra-

ted 15 gastrostyles with H:W ratios ranging from

1.9 to 4.7. Gastrostyles not ridged but bear

sharply pointed spines up to 42 pm long. Dacty¬

lotomes about 84 pm wide; pseudosepta one to

three times width of dactylotome. Dactylostyle

rudimentary, composed of a single line of blunt

cylindrical éléments up to 57 pm high and 11 pm

in diameter.

Female ampullae superficial hémisphères (Fig.

24 A) 0.7-0.9 mm in diameter, each with a large

efferent pore about 0.24 mm in diameter. Male

ampullae also superficial, 0.65-0.75 mm in dia¬

meter, often covered by 1-5 papillae or carinae

(Fig. 24 B). Male efferent pore not observed.

Comparisons

S. gemmascens is one of the most easily distinguished eastem Atlantic Stylaster, characterized

by a rough coenosteal texture, flared cyclosystems, narrow gastropore tubes, and carinate male

ampullae (Table 2).

Remarks

As pointed out by Broch (1918), S. gemmascens had been well characterized by Gunnerus

(1768) from material from Norway but had been referred to the scleractinian Madrepora virginea

(Linnaeus, 1758). Figures from Gunnerus were reproduced by Broch (1918). The same stylasterid

species was again recognizably figured and described by Esper (1790/1794) as Madrepora gemmascens

from material said to corne from the East Indian Sea (description and figures reproduced by

Boschma, 1955a). S. gemmascens was then reported from the North Atlantic, starting with Sa ville

Kent (1871), Duncan (1873), and Sars (1873). It was often mentioned in papers on the fauna of

Norway. Although Esper’s name gemmascens was thus in common use for the North Atlantic

stylasterid, it became available only by suppression of the senior homonym, Madrepora gemmascens

Wilkens, 1787, for a scleractinian (Boschma, 1955a, 1958; International commission on zoological

nomenclature, 1959: opinion No. 574).

In the northern Atlantic, S. gemmascens has occasionally been confused with other species.

Material from between the Faroes and the Hébrides reported by Duncan, first (1870) as Allopora

oculina, and later (1873) as Stylaster gemmascens, proved to be a mixture of 4 species: S. gemmascens,

S. norvegicus, S. erubescens britannicus, and Stenohelia maderensis. Stylaster sanguineus sensu

Thornely (1897), from Rockall Bank, proved to be S. gemmascens ; the true S. sanguineus Milne

Edwards & Haime, 1850, with a pink skeleton, is confined to the southwestern Pacific (Zibrowius,

1981).

S. gemmascens and S. norvegicus frequently occur together and hâve not always been properly

distinguished; many lots in muséum collections under one or the other name hâve proved to be a

mixture of both species.

The fact that S. gemmascens had been reported, first (mistakenly) from the East Indian Sea

and then recognized in the northern Atlantic, encouraged other authors to accept its wide géographie

range, and to uncritically refer somewhat similar forms from other areas to the same species.

The North Pacific S. gemmascens alaskanus Fisher, 1938, first reported from the Aleutian and

subsequently from the Kurile Islands (Naumov, 1960), should not be considered as S. gemmascens.

Cairns (1983b: 430) listed it as a distinct species, S. alaskanus, in Group C (and not Group B) within

the genus Stylaster.

S. gemmascens sensu Hickson & England (1905), from the Sulu Islands (southem

Philippines), is also different (zma Coel 7386). The insufficiently known S. cf. gemmascens sensu

Eguchi (1941) from Japan is also undoubtedly misidentified.

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CA1RNS

Distribution and ecology

S. gemmascens is known from a wide area in the North Atlantic, from Denmark Strait east

of Greenland (off Angmagssalik) and northwest of Iceland through the Faroes - Hébrides area to

Norway and Rockall. The occurrence in Icelandic waters is here reported based on Broch (1914a);

from ail other areas material was available for the présent study. The species is new for eastern

Greenland. The northernmost records are 66°18'N in Denmark Strait, and 71°07'N in Finmark,

Norway.

Along the Norwegian coast the species is common in depths of about 40 m to 400 m,

frequently found together with the bankforming scleractinian Lophelia pertusa. Elsewhere, the species

has been collected down to 621 m northwest of Iceland, and at 665 m between the Faroes and

Hébrides. On Rockall Bank large colonies hâve been photographed and collected by the submersible

“ Pisces III" in depths of 160-190 m.

No symbionts are known.

Types

Stylaster ibericus new species

Fig. 25 A-O, 26 A-G

Ail available specimens from seven closely adjacent stations of northwestem Spain are given

type status: “ Thalassa " stn u-807, 2 colonies fused at the base; stn y-428, 4 encrusting bases of

colonies; stn y-430, 21 colonies + 10 branches + fragments; stn y-431, branch; stn y-432, colony +

3 branches; stn y-438, dead branch; stn y-440, 28 colonies, branches, fragments, mostly dead. A

rather complété fan-shaped colony 30 mm high and 42 mm wide from stn y-430 is designated

holotype (Fig. 25 H), ail other specimens from stn y-430 and the other stations paratypes. Holotype

and most paratypes at mnhn, 4 paratypes from stn y-430 at usnm (75619).

Type locality: “ Thalassa " stn y-430, 4.9.1972, 44°1.6'N, 8°40.6'W, 500 m. Off northwestem

Spain.

Material studied

NW Spain: Holotype and paratypes (see above).

Description

Colonies uniplanar, up to 40 mm high and

wide (Fig. 25 F-H, L-O). Branches cylindrical,

about 1.5 mm in distal branch diameter. Coe-

nosteum glistening white. Coenosteal texture

smooth, resulting from a reticulate-granular coe-

nosteum with very few low granules, each about

11 pm in diameter. As in 5. norvegicus, some

colonies show a transition of coenosteal textures

from coarse, disjointed granules (Fig. 26 D) to a

smooth, almost porcelaneous coenosteum (Fig.

26 C). Coenosteal strips 45-95 pm wide.

Cyclosystems circular to slightly elliptical, 0.8-

1.1 mm in diameter, occurring primarily on

latéral to antérolatéral branch edges. Occasio-

nally some cyclosystems occur on anterior face

and, more rarely, posterior face. Cyclosystems ail

slightly exsert, extending about 0.4 mm above

coenosteum. Based on 685 cyclosystems, there is

a range of 4-11 dactylopores per cyclosystem,

mean 7.71, and mode 8. These data include the

single branch from stn y- 438 which has many

cyclosystems with a low number of dactylopo¬

res : based on 70 cyclosystems, its range is 4-8,

mean 6.00, and mode 6. There are no diastemas.

Gastrostyle occupies lower half of gastropore

chamber, which is constricted at level of gas¬

trostyle tip (Fig. 26 E). Gastrostyle ovate and

sharply pointed, its widest section being at

mid-height, caused by the perpendicular projec¬

tion of very long spines. Illustrated style (Fig. 26

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Fig. 25. Stylaster ibericus (A-B, paratype from “ Thalassa " y- 430, male, usnm 75619; C, paratype from “ Thalassa " y- 440,

female' mnhn; D-K. from “ Thalassa ” y-430, mnhn; L, from “ Thalassa ” u-807. mnhn; M-N. from “ Thalassa ” y-440.

mnhn; O, from “ Thalassa ” y-438, mnhn): A. Pedicularia trace ( x 13); B, dactylostyle ( * 125); C, female ampulla and

efferent pore ( x 39); D, Pedicularia trace ( x 6.8); E, detail of paratype with two Pedicularia traces ( x 2.5); F-G,

opposite views of paratype, Pedicularia trace near thc base (both x 1.1 ); H, holotype ( x 1.1); I-K, branches of holotype

showing sympodial arrangement of cyclosystems (x 3.0, x 3.1, x 4.9 respectively); L, two paratypes on cirripcd shell,

next to base of right colony some very young stages consisting of a single cyclosystem (x 1.9); M-N, stouter dead

paratype ( x 1 . 3 , x l . 5 , respectively); O, stouter dead paratype characterized by cyclosystems with low dactylopore

number (x 2.1).

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

g-* «*«. «m. »»

rrsie reveali “ E

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

F-G) 0.46 mm high and 0.24 mm in diameter

(H:W = 1.9); another style with H:W = 1.3.

Gastrostyle spines cylindrical and blunt, up to 60

pm long and 11 pm in diameter. Spines often

laterally fused into transverse or oblique tiers.

Gastrostyle not ridged. Dactylotomes about 0.12

mm wide. Dactylostyles robust, composed of

blunt, cylindrical éléments up to 50 pm tall and

15 pm in diameter. Dactylostyle éléments basally

fused and occur 3 or 4 across the width of

dactylostyle (Fig. 26 B).

Male ampullae internai, about 0.5 mm in

internai diameter. Male efferent pores small

concavities about 0.15 mm in diameter. Female

ampullae elongate and very low in relief (0.75-1.0

mm in diameter and about 0.25 mm tall); efferent

pore about 0.18 mm in diameter, latéral in

position but inclined upward (Fig. 25 B).

Comparisons

S. ibericus is similar to S. erubescens britannicus, particularly regarding cyclosystem diameter,

gastro- and dactylostyle morphology, and internai male ampullae. These are the only two eastern

Atlantic taxa of Stylaster known to hâve internai male ampullae. S. ibericus is distinguished by its

smooth, porcellaneous coenosteum; higher average number of dactylopores per cyclosystem (9.34

versus 7.71), and normally shaped pseudosepta (Table 2).

Remarks

The spécifie name given to the new species refers to its géographie distribution.

Distribution and ecology

5. ibericus is known from 7 closely adjacent dredging stations in the La Coruna area,

northwestem Spain. The species lives solidly attached to big boulders apparently exposed to bottom

currents (no sédiment in dredge).

Symbionts

Specimens of Pedicularia hâve been found on live colonies of 5. ibericus from 2 stations

(“ Thalassa ” stn y-430, y-432). Dead branches from a third station (y-440) also show typical

Pedicularia traces (Fig. 25 D, 26 D).

Stylaster erubescens Pourtalès, 1868

Material from the study area identified as S. erubescens differs in several aspects from

specimens of the nominotypical western Atlantic form (north to North Carolina) previously described

by Cairns (1986a). Largely a matter of degree, these différences appear insufflcient to justify the

description of distinct species; accordingly, these geographically nonoverlapping forms are presented

here as subspecies of S. erubescens: groenlandicus n. ssp., britannicus n. ssp., and meteorensis n. ssp.,

the western Atlantic nominotypical subspecies being 5. erubescens erubescens.

Stylaster erubescens erubescens Pourtalès, 1868

Synonymy:

Stylaster erubescens Pourtalès, 1868: 135-136.

Chresonymy:

Stylaster erubescens: Pourtalès, 1871: 34, pl. 4. fig. 10-11; 1878: 210. — Broch, 1914a: 12 (only in

synonymy of Stylaster roseus). — Boschma, 1955b: 135-138 (part: NOT North Atlantic); 1957a: 8 (part: NOT

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

North Atlantic); 1965c: 235-236, 245-247, pl. 4, fig. 1-4 (part: not North Atlantic). — Zibrowius & Cairns,

1982: 210, 212 (part: not East and North Atlantic). — Cairns, 1983b: 430; 1986a: 58-61, fig. 26A-H (part: not

N ortheast Atlantic).

Types

Stylaster erubescens: In the original description (Pourtalès, 1868) the species is qualified as

“ rather common between 120 and 324 fathoms [220-592 m] off the Florida reef ” (area known as

Pourtalès Terrace); this information was reiterated in a later publication (Pourtalès, 1871). Types

were not specially designated. According to Cairns (1986a), there is only one larger syntype

(fan-shaped colony about 80 mm high, 117 mm wide) at the mcz, with data corresponding to the 1868

description (Florida, 120-324 fathoms). He also recognized additional syntype branches at usnm

(71822), ypm, and rmnh. Five branches at bmnh (1869.10.25.11, 1891.2.4.15, 1894.12.22.6) also

deserve syntype status.

Type locality: Pourtalès Terrace (Florida), 220-592 m.

Material studied

For western Atlantic material north to North Carolina see Cairns (1986a).

Description

See Cairns (1986a), description based on the western Atlantic nominotypical subspecies.

Comparions

See Table 2 and Comparisons of the 3 new subspecies.

Remarks

Broch (1914a) incorrectly included S. erubescens as a synonym of S. roseus (Pallas, 1766). As

already demonstrated by Boschma (1955b, 1957a, 1965c), S. erubescens and S. roseus are distinct

species. The latter (redescribed by Cairns, 1986a), is primarily a shallow-water species widespread

throughout the Caribbean and ranging south to Brazil.

S. erubescens has been mistakenly reported by early authors from beyond the confirmed range

of its four subspecies presently known in the northwestem and northeastem Atlantic: by Moseley

(1876b: 94) from “ Challenger ” stn 320 in the southwestem Atlantic (subsequently described as a

distinct species, S. densicaulis Moseley, 1879; redescribed by Cairns, 1983a; types bmnh

1880.11.25.175/199); by Thomson (1877: vol. 2, 267) from “ Challenger ” stn 344 off Ascension Island

(mentioned by Moseley, 1879: 450; 1881: 58) as S. granulata (nomen nudunr, material not at bmnh);

and by Moseley (1881: 81) from “ Challenger " stn 170 off the Kermadec Islands (in reality Conopora

verrucosa (Studer, 1878), bmnh 1880.11.25.178).

Distribution and ecology

S. erubescens (nominotypical subspecies erubescens) was known to Pourtalès (1868, 1871,

1878) exclusively from off southeastern Florida. Cairns (1986a) provided a station list for this form

that he qualified as the most commonly collected stylasterid of the western Atlantic. The distribution

was given as comprising the continental shelf and slope of the southeastern United States, from the

Blake Plateau off South Carolina to off southwestem Florida, and extending to Little Bahama Bank,

Cay Sal Bank, and Arrowsmith Bank off Yucatan Peninsula, depth 146-965 m, but most common at

650-850 m.

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Symbionts

In the western Atlantic S. erubescens erubescens is one of the rare stylasterids known as the

host of Pedicularia; traces hâve been found on material from one of the northern localities

(“ Albatross" stn 2416, off Georgia, 505 m; usnm 10542).

In the western Atlantic, Cairns (1986a) found about half of the colonies with gall-tubes

induced by a polynoid polychaete, an association first noticed by Pourtalès (1869).

Stylaster erubescens groenlandicus new subspecies

Fig. 27 A-H, 28 A-H

Chresonymy:

Stylaster roseus — Broch, 1914a: 12-15, pl. 1, fig. 8-9, pl. 2, fig. 10, 11, 17, pl. 3, fig. 22, pl. 4, fig. 36,

39, pl. 5, fig. 43, 47-48 (part); 1936: 15.

Stylaster erubescens — Boschma, 1955b: 135-138 (part: North Atlantic); 1957a: 8 (part: North Atlantic);

1965c: 236, 246 (part: North Atlantic). — Zibrowius & Cairns, 1982: 212 (part: East and North Atlantic). —

Cairns, 1986a: 61 (part: Northeast Atlantic).

Types

Most available material from Greenland through Denmark Strait (“ Poséidon " excepted) to

southeast of Iceland is given type status: a male colony 61 mm high and 40 mm wide from “ Ingolf ”

stn 15 is designated holotype (Fig. 27 C), ail other specimens from “ Ingolf ” stn 15 and various other

stations paratypes (see Material studied). Holotype deposited at zmuk, together with most paratypes

from type locality.

Type locality: “ Ingolf ” stn 15, 4.6.1895, 66°18'N, 25°59'W, 621m. Denmark Strait, northwest

of Iceland.

Material studied

Denmark Strait and Greenland: “ Ingolf ” stn 15, 18 colonies + 26 branches + numerous

fragments (holotype + paratypes, zmuk, usnm 77123); stn 16, 2 branches (paratypes zmuk). — East

Greenland Expédition 1900, off Angmagssalik, 263 m, 5 colonies + 2 branches (paratypes, zmuk,

usnm 77122). — “ Walther Herwig" stn 538-24, 2 male branches (paratypes, imfb). “ Venta" stn

v17/rd-29, 7 colonies + 5 branches + fragments, male + female (paratypes, usnm 60004). —

“ Poséidon ” stn 12/1, ca. 260 branches + fragments, most dead, only one alive (smf 6472; usnm

88826); stn 14/1, ca. 50 branches + fragments, dead (smf 6475; usnm 88829).

Iceland: “ Ingolf ” stn 7, 2 colonies + 7 branches (paratypes, zmuk); stn 52, 16 branches

(paratypes, zmuk). — “ Bartlett ” 1975, stn 52c-5, male branch (paratype, usnm 60005).

Description

Colonies uniplanar (Fig. 27 A, C), up to 80

mm high with a basal diameter of 7 mm (East

Greenland exped. specimens); branches someti-

mes thickened (Fig. 27 C) as in nominotypical

subspecies but evidence of polynoid symbiont

not observed. Coenosteum white and reticulate-

granular in texture (Fig. 28 D). Strips 70-90 pm

wide and fairly continuous, covered with small

rounded granules 8-10 pm in diameter.

Cyclosystems primarily on latéral and antéro¬

latéral branch edges but some occasionally on

anterior and posterior branch faces. Cyclosys¬

tems circular to slightly elliptical in shape, 1.2-1.4

mm in diameter (Fig. 28 A-B). Based on 175

cyclosystems, there is a range of 5-15 dactylopo¬

res per cyclosystem, mean 10.15, and mode 10.

Gastrostyle conical (Fig. 28 E-F), about 0.50

mm tall and 0.25 mm wide in greatest width,

with H:W ratios around 2. Style covered with

blunt, cylindrical spines about 40 pm long and 15

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Fig. 27. — Stylasler erubescens groenlandicus (A-D. from “ Ingolf" stn 15, zmuk; E, from “ Bartletl" 1975, 52c-5, USNM

60005; F-H. from “ Vema ” v17/rd-29, usnm 60004): A, fcmale paratype with abundant clustercd ampullae ( x 1.0); B,

detail of A illustrating large female ampullae ( x 3.2); C, male holotype ( x 1.0); D, detail of holotype illustrating small

male ampullae (x 3.2); E. male paratype with particularly thickened branches (x 2.1); F, male paratype (x 2.9); G,

detail of female paratype illustrating large ampullae and gastrostyles in depth of cyclosystems ( x 4.3); H. female

paratype with large ampullae ( x 3.0).

(im in diameter, the bases of which are fused into

transverse or oblique tiers as in S. norvegicus and

5. ibericus. Dactylotomes about 0.12 mm wide;

dactylostyles robust. Pseudosepta usually equal

to dactylotomes width (Fig. 28 G). Diastemas

présent on cyclosystems positioned at branch

axils (Fig. 28 C) and on larger diameter bran¬

ches.

Female ampullae large superficial hémisphères

(Fig. 27 B, 28 C) 1.0-1.3 mm in diameter,

sometimes with a short latéral efferent tube.

Female efferent pore about 0.20-0.25 mm in

diameter; binary ampullae sometimes présent.

Male ampullae (Fig. 27 D, 28 H) superficial

mounds 0.7-0.8 mm in diameter, becoming inter¬

nai on large diameter branches. Male efferent

pore circular, 80-90 pm in diameter, and located

laterally (Fig. 27 D, 28 H), not apically. Both

genders of ampullae often densely clustered,

especially the males, on anterior and posterior

branch faces (Fig. 27 A-H).

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

D, coenosteal texture (x 92); E. g’astrostyle ( x ( 116, stéréo pair), F gastrostyle surrounded by damaged da^lostyles

( x 69); G. outer edges of pseudosepta. coenosteal texture ( x 56); H, male ampulla with efferent pore ( 56).

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

COMPARISONS

S. erubescens groenlandicus is clearly most similar to the nominotypical subspecies, both

having the same coenosteal texture, thickened branches, and relatively large cyclosystems (Table 2).

Their différences are largely a matter of degree: S. erubescens groenlandicus has a slightly lower

average number of dactylopores per cyclosystem, a higher gastrostyle H:W ratio, and larger ampullae

of both sexes. One qualitative différence concerns the male efferent pores, which are latéral in S.

erubescens groenlandicus and apical in S. erubescens erubescens.

S. erubescens groenlandicus can be distinguished from the other two subspecies on the basis of

coenosteal texture, cyclosystem diameter, gastrostyle shape, and ampullar position and size (Table 2).

Remarks

The name given to the new subspecies refers to its géographie distribution.

Broch (1914a) was the first to report corals referable to S. erubescens groenlandicus from the

northem Atlantic (specimens from 3 stations figured), but he incorrectly included S. erubescens as a

synonym of S. roseus (Pallas, 1766). Boschma (1955b, 1957a, 1965c) corrected this identification to

S. erubescens, but did not examine Broch’s material.

Material mentioned by Broch (1914a) from 3 stations between Greenland and Iceland

(“ Ingolf" stn 17, 94; “ Thor " 1904) was not available for the présent study; most likely this was ssp.

groenlandicus, also obtained from other stations in the same area.

Distribution and ecology

5. erubescens groenlandicus is separated from the nominotypical subspecies by a broad

géographie hiatus extending from South Carolina (32°24'N) to Greenland (60°27'N). It is

geographically nearest to S. erubescens britannicus which extends north to southeast of Iceland.

Broch (1914a) reported S. erubescens groenlandicus (misidentified as S. roseus) from 7 stations

(mainly “ Ingolf") in the North Atlantic, from east of Greenland to northwest and southeast of

Iceland, ail north of 60°N. We hâve seen additional material of the same form from 6 stations.

Broch's data combined with ours cover a depth range from 263 m to 1440 m. The northernmost

record is 66°18'N in Denmark Strait, northwest of Iceland.

No symbionts are known.

Stylaster erubescens britannicus new subspecies

Fig. 29 A-G, 30 A-G

Chresonymy:

Allopora oculina — Duncan, 1870: 90, 95 (part).

Stylaster gemmascens — Duncan, 1873: 322 (part), pl. 49, fig. 8-10.

Stylaster erubescens — Wilson, 1979: 157. — Zibrowius & Cairns, 1982: 212 (part: East and North

Atlantic). — Cairns, 1986a: 61 (part: Northeast Atlantic).

Types

Ail available material (except “ Dana ” stn 6001) from southeast of Iceland through Rockall

Trough to the Celtic Sea is given type status: a female specimen (17 mm high, 24 mm wide) collected

by C. Wandel, SE Iceland, is designated holotype (Fig. 29 A), ail other specimens from the same and

various other stations paratypes (see material studied). Holotype deposited at zmuk, together with

most paratypes from type locality.

Type locality: 64°16'N, 11°15'W, 350 m, C. Wandel, 19.9.1891. Southeast of Iceland.

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Fig. 29. — Stylaster erubescens britannicus (A-C, from southeast of Iceland, coll. C. Wandel, zmuk; D-E, from “ Porcupine ”

stn 54. bmnh 1883.12.10.92. 1880.11.13.6; F. from '■•John Murray " site 4/dredge, bmnh 1986.11.5.1; G, from

- Thalassa ” Z-435, usnm 75614): A, female holotype ( x 2.1); B, detail of A ( x 3.2); C. male paratype ( x 2.3); D-E,

male paratypes (x 1.6, x 2.0. respectively); F, male paratype (x 1.8); G, paratype (x 3.6).

Material studied

SE Iceland: 6 branches from Type locality (holotype + paratypes, zmuk, usnm 75617). —

“ Dana ” stn 6001, tiny poorly preserved colony, tentatively assigned to this subspecies (zmuk).

N Faroes: “ Ingolf" stn 144, fragment (paratype, zmuk).

Between Faroes and Hébrides: “ Porcupine ” stn 54, 3 branches (paratypes, bmnh 1880.11.13.6/6a,

1883.12.10.92).

Rockall Trough: “ John Murray ", site 4/dredge, male branch (paratype, bmnh 1986.11.5.1).

Celtic Sea: “ Thalassa " stn z-430, dead male branch (paratype, mnhn); stn z-435, 2 branches

(paratypes, mnhn, usnm 75614). '

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Description

Colonies uniplanar (Fig. 29 A-F), up to 43 mm

high and 28 mm wide. Branches délicate, not

thickened. Coenosteum white, composed of

coarse granules (Fig. 30 B) 40-60 pm in diameter,

some of which are fused into short strips up to

0.25 mm long. Smaller granules about 8 pm in

diameter occur on lower edges of coarse granu¬

les.

Cyclosystems primarily on latéral and antéro¬

latéral branch edges (Fig. 29 A, C), although

some occasionally présent on anterior and pos-

terior branch faces. Cyclosystems circular to

elliptical, 0.9-1.4 mm in diameter. Based on 270

cyclosystems, there is a range of 7-12 dactylopo¬

res per cyclosystem, mean 9.34, and mode 9.

Gastrostyles lanceolate, up to 0.6 mm high

and 0.32 mm wide ; H:W = 1.6-1.8. Lower half

of style lacks spines or has only very short spines;

midstyle greatly flared caused by long perpendi-

cularly projecting spines; upper third of gas¬

trostyle with large cylindrical spines (up to 66 pm

long and 20 pm in diameter), which are directed

upward (Fig. 30 F-G). Dactylotomes about 0.12

mm wide; dactylostyles robust. Pseudosepta one

to two times width of dactylotome and compo¬

sed of slender (20-30 pm), elongate, labyrinthine

strips that are separated by very wide coenosteal

slits (Fig. 30 E).

Female ampullae (Fig. 29 B) hemispherical,

about 1.1 mm in diameter with an efferent pore

diameter of 25-28 pm. Male ampullae (Fig. 29 C,

30 D) smaller, more irregularly shaped hémi¬

sphères, rapidly becoming internai with a slight

increase in branch diameter; apical efferent pore.

COMPARISONS

S. erubescens britannicus is most easily distinguished from the other subspecies by its coarse

coenosteum. Other distinctive characters are its low number of dactylopores per cyclosystem,

distinctively shaped gastrostyle, unusual pseudoseptal architecture, and primarily internai male

ampullae.

Remarks

The name given to the new subspecies refers to its géographie distribution around the British

Isles.

The earliest record of S. erubescens britannicus in the northeastern Atlantic (Faroes -

Hébrides area) was contemporaneous with the description of the nominotypical subspecies in the

western Atlantic, but this record was not duly recognized until now. In fact, S. erubescens britannicus

is one of the 4 species from one station identified by Duncan, first (1870) as Allopora oculina, then

(1873) as Stylaster gemmascens (the other species of the mixture are S. norvegicus, S. gemmascens, and

Stenohelia maderensis).

From the literature Broch (1914a) already presumed that S. erubescens (incorrectly

synonymized with S. roseus) was included under S. gemmascens sensu Duncan (1873), but he

incorrectly quoted the corresponding illustrations from Duncan (pl. 49, fig. 13-15, instead of fig.

8-10). Broch (1914a) had not seen specimens of S. erubescens britannicus, ail his material (reported

as S. roseus) being the other northern subspecies, S. erubescens groenlandicus.

Distribution and ecology

Geographically S. erubescens britannicus is nearest to 5. erubescens groenlandicus. S. erubescens

britannicus has been confidently identified from 6 stations ranging from southeast of Iceland

(64°16'N) through the Faroes - Hébrides area and the Rockall Trough to the Celtic Sea (48°37'N),

depth range 350-1080 m.

Unidentified Stylaster from Porcupine Bank (W of Ireland) may belong here. In a

sedimentological study, Scoffin & Bowes (1988: 130) mentioned Stylaster sp., dredged and observed

from the submersible “ Cyana ”. The dredged material C Challenger II” 1981, stn 30, bmnh

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Fig. 30. — Stylasler erubescens britannicus (A. D. G, paratype from “ Thalassa " z-430, mnhn; B. male paratype from “ John

Murray'' site 4/dredge, bmnh 1986.11.5.1; C, E, F. paratype from " Porcupine" stn 54. bmnh 1880.11.13.6): A,

cyclosystem and coenosteal texture (x 17); B. coenosteal texture (x 71. stéréo pair); C, cyclosystem (x 30); D,

longitudinal fracture through cyclosystem revealing gastrostyle and internai male ampulla ( x 38, stéréo pair); E,

pseudosepta (x 70); F-G, gastrostÿles (x 71, x 116, respectively).

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

1989.6.16.2-3), which consists of two highly eroded pièces of larger colonies with poor traces of

cyclosystems on partly anastomosing branches, cannot be identified to the species level. The photos

from the “ Cyana" dive referred to (cruise cyaporc, dive 8, 22.7.1986, 50°42'N, 11°07'W, 700 m)

unfortunateiy do not show the observed stylasterid colonies.

Symbionts

A branch of S. erubescens britannicus from the Celtic Sea (“ Thalassa ” stn z-430) shows a

typical trace of Pedicularia.

Stylaster erubescens meteorensis new subspecies

Fig. 31 A-H, 32 A-H

Types

Ail available material from Great Meteor Seamount (“ Meteor ”, “ Chain ”) is given type

status: a small male branch (11 mm high, 11 mm wide) with 19 cyclosystems, laterally overgrowing

a pteropod shell, from “ Meteor ” stn 129/dd-94 is designated holotype (Fig. 31 E), ail other

specimens paratypes (see Material studied). Holotype and most paratypes deposited at zsim.

Type locality: “ Meteor ” cruise m 19, stn 129/dd- 95, 17.2.1970, 29°59'N, 28°33'W, 290 m.

Great Meteor Seamount.

F, °' (A-Ç. fom imprécise " Tomtguer station, mnhn; E-H, from “ Meleor" stn

0.8); B. detail from A (x 1.7); C-D. other branches (a 1.4. a 1.5, respective!,);

E, holotype ( x 3.1), F-H, paratype colonies (x 2.3, x 2.4, x 2.2, respectively). 3

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Material STUDIED

Great Meteor Seamount: “ Meteor" stn 129/dd-94, 129/dd-95, 131/dd-98, 20 small colonies

and branches + minor fragments, male + female (holotype + paratypes, zsim). — “ Chain ” stn

ch7/pd-24, small colony (paratype, usnm 75612).

Imprécise locality Southwest of the Azores: said to corne from seamount 260 miles ( ?) [475 km]

Southwest of Faial, ca. 500 m, fishing boat “ Tomiguel", coll. J.G. Pereira, Sept. 1976, several

colonies + larger fragments, male + female (most mnhn; usnm 75611).

Description

Colonies from Great Meteor Seamount small

and bushy (Fig. 31 E-H), up to 25 mm high;

colonies from “ Tomiguel ” bushy (Fig. 31 A), up

to 15.5 cm high and 10 cm wide. Branches

slender (Fig. 31 C-D). Coenosteum white,

composed of oddly shaped convex strips 65-75

gm wide. Strips highly anastomotic, forming a

maze of interconnections, and characterized by

symmetrical protubérances on either side of the

strip (Fig. 32 B). Strips smooth (granules not

présent), producing a porcelaneous aspect.

Cyclosystems primarily on latéral branch ed-

ges but some occasionally présent on anterior

and posterior faces. Cyclosystems circular to

irregular in shape, 0.9-1.2 mm in diameter (Fig.

32 A, C). Based on 206 cyclosystems, there is a

range of 9-15 dactylopores per cyclosystem, a

mean of 11.98, and a mode of 11.

COMPARISONS

S. erubescens meteorensis is most easily distinguished from the other subspecies by its unique

coenosteal texture and bushy colony shape. Other distinctive characters are its very long gastropore

spines, high number of dactylopores per cyclosystem, and slender pseudosepta (Table 2).

Gastrostyles ovate, illustrated style (Fig. 32

G-H) 0.39 mm tall and 0.28 mm wide (H:W =

1.39). Style covered with extraordinarily long

and often bifid spines, up to 120 gm long and 15

gm in diameter. The long closely spaced spines

make the gastrostyle a very délicate structure.

Dactylotomes about 0.11 mm wide; pseudoseptal

width equal to or less than that of dactylotomes.

Dactylostyles robust, composed of cylindrical

éléments up to 60 gm tall and 15 gm in diameter,

arranged 3 or 4 across width of dactylostyle (Fig.

32 E-F).

Female ampullae superficial (Fig. 32 A), 0.6-

0.8 mm in diameter, often with a short efferent

tube leading to efferent pore, which is about 0.2

mm in diameter. Male ampullae superficial

mounds 0.6-0.7 mm in diameter, each having 1

or 2 apical efferent pores 25 gm in diameter.

Remarks

The name given to the new subspecies refers to its géographie distribution (Great Meteor

Seamount).

Distribution and ecology

There is a wide géographie hiatus between S. erubescens meteorensis, the western Atlantic

nominotypical subspecies, and the geographically nearest eastem Atlantic subspecies, S. erubescens

britannicus.

The only précisé locality from which S. erubescens meteorensis is known is the Great Meteor

Seamount (300 m). The seamount (ca. 500 m) visited by the fishing boat “ Tomiguel ” could not

precisely be identified with one of those situated between the Azores, the Mid-Atlantic Ridge, and

Great Meteor Seamount.

No symbionts are known.

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Fig. 32. — Stylaster erubescens meieorensis (A, D, female paratype from imprécise “ Tomiguel" station, usnm 75611; B-C,

E-H, male paratype from “ Meteor ” stn 129/DD-95, zsm): A, cyclosystcm and female ampulla (x 32, stéréo pair); B,

D, coenosteal texture ( x 88, x 92, respectively); C, cyclosystem ( x 36); E-F, dactylostyle viewed end-on, from above

cyclosystem (x 745, x 418, respectively); G, gastrostyle and dactylostyle (x 62); H, gastrostyle (x 124).

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

Genus STENOHEL1A Saville Kent, 1870

Diagnosis. — Gastro- and dactylopores arranged in cyclosystems, which occur exclusively on

anterior branch face. Cyclosystems without lips or lids. Coenosteum white or light brown, either

linear-imbricate or reticulate-granular in texture. Gastropores long and usually curved; gastrostyles

présent, usually encircled by a robust ring palisade. Dactylostyles rudimentary. Ampullae superficial,

often clustered around base of cyclosystem. Ampullar efferent pores of both sexes usually well

distinguished.

Type species: Allopora maderensis Johnson, 1862, from Madeira.

Genus represented in the study area by the type species and a poorly known form (unnamed)

that appears to be a distinct species.

Stenohelia maderensis (Johnson, 1862)

Fig. 33 A-L, 34 A-J

Synonymy:

Allopora maderensis Johnson, 1862: 196, fig. 1-3.

Chresonymy:

Stenohelia maderensis — Saville Kent, 1870: 120; 1871: 277, pl. 24, fig. 3, 3a-c. — Boschma, 1957a:

31-32; 1964b: 64-65, 67, 68, 71, 72 (part: only eastem Atlantic), text-fig. la-g; 1964d: 80-84 (part: only eastern

Atlantic); 1967: 325-329 (part: only eastern Atlantic), text-fig. 2a-b, pl. 1, fig. 3-4, pl. 2, fig. 3-4, pl. 2, fig. 3-4;

1968d: 437, 438 (part: only eastern Atlantic). — Cairns, 1983b: 431, 487-489, fig. 20 A-B, D-G.

Allopora madeirensis — Studer, 1878: 633; 1879: 676.

Stylaster madeirensis — Moseley, 1879: 481.

Stenohelia madeirensis — Moseley, 1879: 503; 1881: 88. — Greeff, 1886: 20. — Studer, 1889: 6-7.

Allopora oculina — Duncan, 1870: 290, 295 (part).

Stylaster gemmascens — Duncan, 1873: 332 (part), pl. 49, fig. 13-15.

Stylaster tiliatus — Hickson, 1912b: 461.

Types

Allopora maderensis : According to Johnson (1862) the unique figured colony (considered the

holotype) on which the description was based was 89 mm high and 57 mm wide. Although the bmnh

was indicated as depository, the specimen could not be found there and may be lost. A smaller colony

(51 mm high, 29 mm wide) at the bmnh (1872.6.26.7, Madeira, J.Y. Johnson) registered as Johnson’s

type does not correspond to the larger holotype as previously figured, and may be a later acquisition

from Johnson; it is here designated the neotype (Fig. 33 A). Fragment of the neotype usnm 75625.

Type locality: Madeira. No additional indications in Johnson’s text.

Material studied

Between Faroes and Hébrides: “ Porcupine" stn 54, 2 small branches (bmnh 1880.11.13.10-

11 ).

S Bay of Biscay — NW Spain: “ Thalassa ” stn t-503, small branch; stn x-340, fragment; stn x-341,

3 fragments; stn x-342, fragment; stn x-347, 3 fragments; x-352, 3 fragments; stn x-353, 5 fragments;

stn x-362, 14 fragments; stn x-363, 7 fragments; stn y-434, fragment (ail mnhn). — Off Cabo Penas

43°58.06'N, 5°43.95'W, 769 m (stn h-5), C. Alvarez-Claudio, 10 small colonies and branches (usnm

85078).

Galicia Seamount: “ Nor oit” 1987, cruise seamount 1 stn dw-108, 85 pièces, mainly branch

Source : MNHN, Paris

100

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Fig. 33. — Sienohelia maderensis (A-C, from Madeira, bmnii 1872.6.26.7; D-E, from Madeira. BMNH 1873.7.9.6; F, from “ Jean

Charcot” 1966, stn 29, mnhn; G-H, from “ Thalassa" t-503, mnhn; I, from “ Porcupine” stn 54, bmnh

1880.11.13.10-11; J, from “ Talisman ” drag. 103, mnhn; K-L, from “ Gazelle” stn 2, zmb 1772): A, neotype ( x 1.3);

B, branch segment of neotype with three cyclosystems (x 18); C, Pedicularia trace on neotype (x 20); D, small

incomplète colony (x 2.3); E, branch segment of D with four cyclosystems (x 15); F, branchlet illustrating

cyclosystems (x 15); G, small colony (x 3.8); H, detail of G with gastrostylc visible in gastropore (x 14); I, colony

with clustering female ampullae ( x 3.5); J, colony with clustering female ampullae ( x 3.2); K, branch ( x 2.9); L, detail

of K with ampullae near cyclosystems ( x 14).

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

101

fragments with 1 to 6 cyclosystems, and a few tiny colonies with up to 3 cyclosystems (mnhn; usnm

88326); stn dw- 116, tiny colony with 3 cyclosystems (mnhn).

Madeira Archipelago: Neotype. — Madeira, R.T. Lowe, colony (bmnh 1873.7.9.6). —

“ Gazelle ” stn 2, 2 branches (zmb 1772). — “ Jean Charcot ” 1966, stn 21, 7 fragments (mnhn); stn

29, 12 fragments (mnhn); stn 49, dead fragment (mnhn).

Cape Verde Islands: “ Talisman ” drag. 103, ca. 60 colonies, branches, fragments (most mnhn;

branch usnm 75627; branch bmnh 1950.1.11.81). — “ Calypso ” 1959, stn 16, 4 branches (mnhn); stn

91, ca. 20 branches + fragments (most mnhn; 3 branches usnm 75626).

Description

Colonies uniplanar (Fig. 33 A, D, I-K), largest

specimen known (Johnson’s holotype of Allo-

pora maderensis) 89 mm high and 57 mm wide;

another incomplète colony 66 mm wide. Bran¬

ches with inversed unifaciality of cyclosystem

arrangement occasionally occur. Branches cylin-

drical and delicately branched, tapering to

branch tips equalling diameter of cyclosystem.

Coenosteum white, linear-imbricate in texture

(Fig. 34 G). Strips 75-100 pm wide, bordered by

deep continuous slits about 10 pm wide (Fig. 34

F). Platelets broad and corrugated, extending

across width of strip. Coenosteal papillae occur

on posterior face of some specimens.

Cyclosystems elliptical to irregular in shape

(Fig. 33 B, E-F, H, L, 34 E), with greater axis

perpendicular to branch axis. A typical gastro-

pore measures 1.1 x 0.85 mm in diameter. Based

on 727 cyclosystems, there is a range of 8-20

dactylopores per cyclosystem, mean 14.61, and

mode 14. The component data for each of the 5

areas included are: Madeira Archipelago, 179

cyclosystems (from 5 stations), range 12-20,

mean 14.80, and mode 15; Cape Verde Islands,

326 cyclosystems (from 3 stations), range 11-20,

mean 14.75, and mode 14; Galicia Seamount, 89

cyclosystems (“ Nor oit ” 1987, cruise seamount 1

stn dw-108), range 8-20, mean 14.29, and mode

15; south of Bay of Biscay, 87 cyclosystems (off

Cabo Penas), range 11-18, mean 14.46, and mode

15; Faroes - Hébrides area, 46 cyclosystems

(“ Porcupine” stn 54), range 12-16, mean 13.70,

and mode 14. As observed on material from

Galicia Seamount, tiny colonies may hâve regu-

lar cyclosystems with a particularly low number

of dactylopores.

Gastropores deep and curved, but gastrostyle

tip always visible in undamaged cyclosystem

(Fig. 33 H). Diffuse ring palisade at level of

gastrostyle tip composed of irregularly shaped

éléments up to 43 pm in diameter and 60 pm tall,

but éléments more commonly only about 15 pm

in diameter. Gastrostyle conical (Fig. 34 B-C), up

to 0.40 mm tall and 0.16 mm in diameter, with

H:W ratios ranging from 2.5-3.6. Style slightly

ridged and very spinose, the long slender spines

up to 55 pm long. Dactylotomes about 80 pm

wide; pseudosepta one to two times dactylotome

width; no diastemas. Dactylostyle composed of a

row of cylindrical to clavate éléments up to 46

pm tall and 14 pm in diameter (Fig. 34 D).

Female ampullae hemispherical (Fig. 33 I, 34

J), 0.70-0.85 mm in greater diameter, with 1 or 2

efferent pores, each 0.16-0.18 mm in diameter.

Binary ampullae are elongate (elliptical in cross

section), with one efferent pore occurring at each

of the vertices. Male ampullae (Fig. 34 H-I)

slightly less prominent superficial mounds 0.45-

0.60 mm in diameter, with 1-3 apical efferent

pores, each about 40 pm in diameter. Male

ampullae often occur on posterior face opposite

cyclosystems, whereas female ampullae are

usually clustered on anterior face near a cyclosys¬

tem, often with their efferent pores directly

adjacent to cyclosystems.

COMPARISONS

Of the 11 valid species of Stenohelia (see Cairns, 1983b, 1986a), 3 occur in the Atlantic. The

western Atlantic S. profunda Moseley, 1881, and S. pauciseptata Cairns, 1986, are discussed and

compared to S. maderensis by Cairns (1986a). To reiterate, S. maderensis differs from S. profunda by

having a lower average number t>f dactylopores per cyclosystem, a shorter gastropore tube (allowing

Source : MNHN, Paris

102

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

“ n f 8T a E G°mil TaB ”? an ” drag - 103 ' BMNH <950.1.11.81; B-C, H-I, male froi

female ampullae ( x i 9 ); BCgastroi.ilefi gf'xl^ “f tyP f- 75625): A, brknch.jp beS foi

stéréo pair); F-G, coenosteal texture (x 65 x 183 resnecHwiv'rh' dactylostyle < x 358 )i E, cyclosystcm (x 2i

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

103

a view of the gastrostyle tip), smaller cyclosystems, and rugose ampullae. It differs from S.

pauciseptata by having a higher average number of dactylopores per cyclosystem, a smaller H:W

ratio, much larger gastrostyle spines, and larger ampullae.

Remarks

On the basis of Johnson’s (1862) holotype from Madeira, Saville Kent (1870) emended the

species description and erected the genus Stenohelia. Studer (1878, 1879, 1889) reported a new record

from Madeira, and Saville Kent (1871) and Greeff (1886) new records from the Cape Verde

Islands. Boschma (1964b) analyzed the descriptions by Johnson (1862) and Saville Kent (1870,

1871) and reproduced the figures contained in these papers.

By error, Hickson (1912b) referred material from the Cape Verde Islands (“ Talisman") to

Stenohelia tiliata (Hickson & England, 1905), a species based on material from the Sulu Islands

(Southern Philippines). That misidentification was corrected by Boschma (1967), who examined

Hickson’s material and concluded that it was typical S. maderensis.

Although he had access to authentic S. maderensis from the eastern Atlantic, Boschma (1964b,

1964c, 1964d, 1967, 1968d) incorrectly included under S. maderensis a different species from the West

Indies (well figured by him 1964b: pl. 1, fig. 13-14; 1964d, text-fig. la-c, pl. 2, fig. 1-4). As pointed

out by Cairns (1986a), the western Atlantic species in question is S. profunda Moseley, 1879.

Boschma frequently used average numbers of dactylopores per cyclosystem and proportions

of gastropore tube and gastrostyle in species diagnosis. It is therefore surprising that he did not take

into account as significant the remarkably different values characerizing the eastern and western

Atlantic Stenohelia. In one of Boschma’s papers (1968d), S. maderensis (with an average of 13.50

dactylopores per cyclosystem) is said to corne from the Azores. This was a lapsus; in fact, the material

in question came from the Cape Verde Islands (“ Talisman ”; previously mentioned by Hickson,

1912b, under 5. tiliatus).

Unfortunately, the holotype of S. maderensis, as characterized and figured by Johnson (1862),

was not available for the présent study, and additional material from Madeira is scarce: 2 small

colonies at the bmnh (including designated neotype (Fig. 33 A-B); 2 small branches from the

“ Gazelle ” expédition (Fig. 33 K-L); and a few small fragments collected by the “ Jean Charcot ” in

1966 (Fig. 33 F). These specimens appear conspecific with Johnson’s missing holotype, and are herein

considered as topotypic.

Material is much more abundant from the Cape Verde Islands and includes pièces from larger

colonies (originally exceding 10 cm ?). Only small fragments (ail dead, the largest comprising 11

cyclosystems) are available from the “ Thalassa ” stations off northwestern Spain and in the south of

the Bay of Biscay (ail sorted out from sédiment samples) but live colonies up to 25 mm high and 30

mm wide are known from 2 stations off Cabo Penas (cruises cocace of the University of Oviedo in

1987; part of material studied here, the other station being 43°56.50'N, 5°48.90'W, 893 m; C.

Alvarez-Claudio, in litt. 1990).

5. maderensis is one of the 4 species from between the Faroes and the Hébrides confused by

Duncan, first (1870) as Allopora oculina, then (1873) as Stylaster gemmascens (the other species are

S. gemmascens, S. norvegicus, and S. erubescens britannicus).

Distribution and ecology

S. maderensis is known from between the Faroes and the Hébrides (665 m; one old record;

presence in the area to be confirmed), northwestern Spain and the south of the Bay of Biscay (490-910

m), Galicia Seamount (985-1125 m), Madeira Archipelago (depth recorded from 4 stations, 110-500

m; dead fragments from the deeper stations), and the Cape Verde Islands (150-400 m). This is a wide

geographical range (from about ISSN to 60°N) and a wide depth and température range (ca. 7-15°C).

The shallowest records are 110-128 m at Madeira, and 150 m in the Cape Verde Islands; the deepest

Source : MNHN, Paris

104

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

occurrences in the Bay of Biscay and on Galicia Seamount, comprising live specimens, appear

correctly recorded. Greater depths are linked with higher latitudes, which would seem to be an

anomaly.

In the Cape Verde Islands S. maderensis occurs together with the precious scleraxonian gorgonian

Corallium rubrum. In 1883 the “ Talisman ” obtained abundant material of the stylasterid from

several dredge hauls (150-275 m) in the area off Sào Tiago where the precious coral was commercially

fished at that time (cf. Zibrowius, Monteiro Marques & Grasshoff, 1984). Greeff (1886), who in

other papers reported on the red coral from the Cape Verde Islands, received a colony of S.

maderensis at Sào Tiago, most likely from red coral fishermen. S. maderensis and Corallium rubrum

hâve again been dredged together in the Cape Verde Islands by the “ Calypso ” in 1959 between Maio

and Boavista (185 m).

A tiny fragment of Stenohelia, probably S. maderensis, is known from the Lower Pleistocene

of Sicily (see Records of fossil stylasterids from Europe).

Symbionts

In the Madeira Archipelago, 5. maderensis is the host of Pedicularia. Johnson (1862) already

noticed this association (2 specimens of the symbiont reported from the holotype colony). The

neotype (bmnh 1872.6.26.7), another old colony from Madeira (bmnh 1873.7.9.6), and a small branch

(“ Jean Charcot ” 1966, stn 29) each hâve one distinct trace of Pedicularia (Fig. 33 C).

Stenohelia sp. A

Fig. 35 A-H

Material studied

Azores: “ Jean Charcot ” 1971, cruise biaçores unknown station, small male colony now

broken into 6 branches and fragments (most mnhn; 1 fragment usnm 77126).

Description

The only small colony available was irregular

in shape, not uniplanar as typical Stenohelia

colonies known so far, the deformation being

due to the presence of a gall-tube induced by a

polynoid polychaete. The colony has been bro¬

ken in order to extract the polychaete; resulting

fragments (Fig. 35 F-H) up to 25 mm tall, highly

modified by cavernous polychaete tube. Indivi-

dual branches with cyclosystems on anterior face

only (Fig. 35 F), as typical for Stenohelia.

The coenosteum is linear-imbricate (Fig. 35

C-D) covered by broad fiat corrugated platelets,

but along the midline of every third or fourth

strip is a prominent ridge or row of blunt spines,

producing a distinctive carinate branch texture

(Fig. 35 A). Cyclosystems are about 1.1 mm in

diameter. Based on 62 cyclosystems, there is a

range of 11-18 dactylopores per cyclosystem,

mean 14.18, and mode 15. Male ampullae (Fig.

35 A-B) are large, about 0.8 mm in diameter, and

covered with tall blunt spines; 1 -3 apical efferent

pores occur on each ampulla, each about 60 pm

in diameter.

Comparisons

Stenohelia sp. A differs from S. maderensis primarily in its unusual carinate coenosteum and

large spiny male ampullae. Commensal relationship with a polynoid polychaete may also be a

distinctive character.

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

105

Fig. 35. — Stenohelia sp. A (A-H, male from “ Jean Charcot" 1971, unknown station. Azores, mnhn): A. branch fragment

illustrating coenosteal ridges and male ampulla ( * 33, stéréo pair); B, two male ampullae with eflerent pores ( x 37);

C-D, coenosteal texture ( x 80, x. 150, respectively); E, cyclosystem ( x 35); F-H, three fragments of original colony;

G, illustrating gall tube induccd Bÿpolynoid polychaete (x 3.7, x 3.7, x 2.9, respectively).

Source : MNHN, Paris

106

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Remarks

The form documentée! here is not identified as 5. maderensis, but not enough material is

présent to justify the description as a new species.

Distribution and ecology

A single colony from the Azores, unknown locality and depth (bathyal).

Symbionts

The single colony is deformed by a gall-tube (Fig. 35 G) induced by a polynoid polychaete

(Harmothoe sp.).

Genus CRYPTHELIA Milne Edwards & Haime, 1849

Diagnosis. — Gastro- and dactylopores arranged in cyclosystems, which usually occur

exclusively on anterior branch face. Cyclosystems partially or entirely covered by one or more fixed

lids. Coenosteum white or light brown, linear-imbricate in texture, and often spinose as well.

Nematopores usually présent, especially on cyclosystem lids, pseudosepta, and ampullae. Gastropore

double-chambered; no gastro- or dactylostyles. Ampullae usually superficial and large, occurring in

various positions and with a variety of efferent pore locations. Cairns (1986b) described 3 female

ampullar types and 8 male ampullar types, for 24 possible permutations; however only 9

combinations (not 12 as reported by Cairns, 1986b), hâve been discovered thus far: A-Al, A-A2,

A-C4, B-B, B-Cl, B-C2, B-C3, B-C4, C-D (as shorthand notations these are called ampullar

formulae).

Type species: Crypthelia pudica Milne Edwards & Haime, 1849, from the western Pacific.

Genus represented in the study area by 4 species.

Crypthelia affinis Moseley, 1879

Fig. 36 A-I, 37 A-I

Synonymy:

Cryptohelia affinis Moseley, 1879: legend on pl. 42.

Cryptohelia moseleyi Hickson & England, 1905: 21.

Chresonymy:

Cryptohelia pudica - Moseley, 1876a: 548, 557 (part: 2798 m); 1879: 462-467, 482 (part: 2798 m), pl. 35,

fig. 7, pl. 42 (with legend on plate Cryptohelia affinis), pl. 44, fig. 1, 3-8; 1881: 71-76, 82-83, 88 (part:

“ Challenger ” stn 3, 2790/2798 m), pl. 2, fig. 7, pl. 9 (with legend on plate Cryptohelia affinis); pl. 11, fig. 1, 3-8,

pl. 12, fig. 7. — Thomson, 1877: vol.l, 271 (part: “ Challenger ” stn 3, 2790 m), fig. 65.

Cryptohelia affinis — Moseley, 1881: legend on pl. 9.

Crypthelia affinis - Boschma, 1951b: 455-456; 1953c: 171-172; 1956b: F100, fig. 81.2; 1957a: 34; 1968a: 106

(part: not “talisman ”). — Cairns, 1983b: 431; 1986b: 24-25.

Crypthelia moseleyi — Fisher, 1938: 534. — Broch, 1936: 94.

Types

Cryptohelia affinis : There is no original description, strictly speaking. The name C. affinis

appears only on a plate published twice (Moseley, 1879, 1881) and is not used in the corresponding

explanation or in the text, where the figured stylasterid (from “ Challenger ” stn 3) is incorrectly

ascribed to Cryptohelia pudica Milne Edwards & Haime 1849. Since Moseley’s species is different.

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

107

it has to be designated by the first available name. C. affinis Moseley, 1879, fullfills this condition,

since this name is clearly associated with an illustration in which the species is recognizable.

Moseley (1879, 1881) did not indicate the number of specimens from “ Challenger " stn 3, but

6 branches (5 collected alive, 1 dead) and some smaller fragments, here considered as syntypes (part

of them figured herein, Fig. 36 F-G), are preserved at the bmnh (1880.11.25.188). The largest of the

live pièces (45 mm high, 16 mm wide, 40 cyclosystems) was part of the larger piece figured by

Moseley (1881, pl. 12, fig. 7) and together with another branch (20 cyclosystems) was part of the

colony figured by Thomson (1877: vol. 1, fig. 65; erroneously referred to stn 23).

Type locality: “ Challenger " stn 3, 18.2.1873, 25°45'N, 20°12'W, 2790 m. Far Southwest of the

Canary Islands. Locality and depth doubtful (see below Distribution and ecology).

Crypthelia moseleyi : Hickson & England (1905) were right in considering Crypthelia pudica

sensu Moseley (as described and figured from “ Challenger ” stn 3) as a distinct species. Being not

aware of the availibility of affinis, they proposed the name moseleyi. The latter thus is an objective

synonym of the former, each being based on the same name-bearing type material.

Material studied

SW Canary Islands: syntypes of Cryptohelia affinis (see above).

Azores: Prince of Monaco stn 203, dead branch (mom); stn 233, 4 dead branches + fragments

(mom). — “ Jean Charcot ” 1971, cruise biaçores stn 25, dead branch (mnhn); stn 74, colony +

fragment (mnhn); stn 102, dead colony (mnhn); stn 112, 5 colonies + branches (most mnhn; usnm

75624); stn 135, dead branch (mnhn); stn 180, 7 colonies + branches + fragments (mnhn); unknown

station, dead female colony (mnhn).

Description

Colonies uniplanar, up to 53 mm high and 79

mm wide (Fig. 36 A-F). Branch anastomosis

common. Branches with inversed unifaciality of

cyclosystem arrangement occasionally occur

(Fig. 36 B). Coenosteum primarily linear-

imbricate (Fig. 37 F-G), becoming reticulate-

imbricate near cyclosystems and on ampullae

(Fig. 37 D). Strips 75-90 pm wide, covered by

coarse, irregularly shaped platelets, producing a

rough microtexture. Nematopores randomly

scattered over branch coenosteum, pseudosepta,

ampullae, and lids; nematopores about 65 pm in

diameter.

Cyclosystems circular to slightly elliptical in

shape, 1.4-1.8 mm in diameter (Fig. 36 G-H).

Based on 88 cyclosystems from the syntypes

(“ Challenger ” stn 3) there is a range of 12-18

dactylopores per cyclosystem, mean 14.51, and

mode 15; based on 341 cyclosystems from speci¬

mens from the Azores (9 stations) the range is

12-22, mean 16.26, and mode 17.

Maximum width of upper gastropore chamber

about 0.65 mm, which narrows to a gastropore

ring constriction of about 0.50 mm in diameter.

Lower chamber (Fig. 37 H) about 0.9 mm in

greatest diameter and about 0.1 mm deep. Cy¬

closystem lid tongue-shaped and horizontal, co-

vering 50-100 % of cyclosystem when viewed

from above. Intact lid approximately 1 mm wide

and quite thin in male colonies (Fig. 37 B) but

invariably inflated in mature female cyclosystems

(Fig. 37 C). Dactylotomes about 75 pm wide;

pseudosepta of equal width and concave.

Each female cyclosystem has an ampulla

consisting of a large swelling in proximal cy¬

closystem wall (Fig. 37 C), which often extends

into lid and partially around the cyclosystem.

Efferent pore circular and quite large (0.35 mm

in diameter), occurring on lower wall of upper

gastropore chamber in proximal cyclosystem

région. Male ampullae consist of 1-5 less conspi-

cuous swellings that encircle cyclosystem. Male

ampullae most common on proximal cyclosys¬

tem wall adjacent to lid (Fig. 37 D). Each male

ampulla bears a large apical steep sided conca-

vity 0.20-0.25 mm in diameter, the center of

which is an efferent pore about 45 pm in

diameter. According to the classification of

Cairns (1986b), the ampullar formula is B-Cl,

the most common of the nine known combina¬

tions, shared by five other species.

Source : MNHN, Paris

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Fig. 36. — Crypthelia affinis (A, from “ Jean Charcot " 1971, stn 74, mnhn; B-D, from “ Jean Charcot ” 1971, stn 112, mnhn;

E-H, syntypes of C. affinis, bmnh 1880.11.25.188; I, from Prince of Monaco stn 233, mom): A, colony ( * 0.7); B, branch

iliustrating alternation of umfaciality of cyclosystem arrangement ( x 2.4); C, branch ( x 3.6); D, colony ( x 1.8); E-F,

colony fragments (x 2.0, x 1.6, respectively); G, syntype cyclosystem having lost the lid (x 15); H, syntype

cyclosystem with lid intact ( x 15); 1, part of massive dead colony ( x 3.6).

Crypthelia medioatlantica (J-L, holotype, zmuk; M, paratype, mom): J, colony still with soft tissues ( x 2.8); K-L,

branch segments of J with cyclosystems having lost their lids (both x 13); M, branch (x 4.8).

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

109

Fig. 37. Cryplhelia affinis (A-B. D, F-H. male from “ Jean Charcot ” 1971, unknown station, Azores, mnhn; C, E, I, female

from "Jean Charcot" 1971, stn 112, usnm 75624): A-B, male cyclosystem viewed from above and from side (x 21,

x 33, respectively); C, side view of female cyclosystem ( x 35); D, male cyclosystem with broken lid, two efferent pore

dépréssions in upper right (x 20, stéréo pair); E, pseudosepta (x 75); F-G, coenosteal texture (x 148, x 97,

respectively, F being a stéréo pair); H, longitudinal section of male cyclosystem showing gastropore chambers ( x 26,

stéréo pair); I, longitudinal section of fema^cyclosystem showing gaslropore chambers and large efferent pore (x 19).

Source : MNHN, Paris

110

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

C. affinis

C. medioatlantica

coenosteum: width + relief of

strips; width of platelets

75-90 pm, sligthtly convex;

platelets irregularly

shaped, coarse

70-85 pm, sligthtly convex;

narrow platelets (4-14 pm)

nematopores: size; location

65 pm, random on coenos¬

teum, ampullae, lid,

pseudosepta

absent

cyclosystem: average size and shape

1.4-1.8 mm, circular to

slightly elliptical

1.2-1.4 mm, circular

dactylopores per cyclocystem:

range mean, mode (N)

12-22, 16.54, 16 (420)

14-19, 15.86, 15 (22)

cyclosystem lid: shape; % cover of

cyclosystem; inclination

tongue-shaped; 50-100 %;

horizontal

digitiform; 0-20 %;

inclined

pseudosepta: width relative to

dactylostomes; concavity

esqual; concave

esqual; very concave

ampullar formula (see Caims,

1986b and text)

B-Cl

?-Cl (female ampullae

unknown)

other diagnostic characters

female efferent pore opens

quite low in gastropore

chamber

male ampullae carinate

COMPARISONS

Within the Atlantic, C. qffinis is most similar to C. medioatlantica and is compared to that

species in the account of the latter and in Table 3.

Remarks

Samples of several species of Crypthelia collected by the “ Challenger ” at 4 stations (stn 3,

Canary Islands, 2790 m; stn 24, West Indies, 713 m; stn 171, Kermadec Islands, 1097 m; stn 236,

Japan, 1417 m) were ail referred by Moseley (1876a, 1879, 1881) to C. pudica. Moseley’s (1879, 1881)

detailed descriptions and illustrations of what he believed to be C. pudica were based on the more

abundant material from stn 3, part of which had been collected alive and provided the soft parts for

an exemplary anatomical study.

A lapsus occurred in the sentence indicating the origin of the material (Moseley, 1879: 462;

1881: 71): “ The specimens, the anatomy of which is here described, were dredged off the mouth of

the La Plata This should read “ off the Canary Islands In fact, Moseley did study the anatomy

of other stylasterid species dredged at “ Challenger ” stn 320, in the Southwest Atlantic off the mouth

of the Rio de la Plata, but no Crypthelia had been obtained there. In addition to the anatomical

structures of the soft parts, Moseley figured a branch of the coral, the origin of which is clearly

indicated in the explanation as being the Canary Islands station (Moseley, 1881: 226, pl. 12, fig. 7).

Throughout his text, Moseley (1879, 1881) uses the name C. pudica, but on a plate (pl. 42 of

1879, reprinted as pl. 9 of 1881) showing the anatomy of the soft parts, the name C. qffinis occurs.

In ail probability, Moseley at first was convinced that the Crypthelia from the Canary Islands station

belonged to a species different from C. pudica (the latter described from the West Pacific and type

of the genus Crypthelia), and accordingly intended to name it C. affinis, whereas later he concluded

that the Canary Islands material was conspecific with the type species. After his change of opinion

the name C. affinis was superfluous, but already engraved on the plate and was not corrected.

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

111

C. tenuiseplata

C. vascomarquesi

120 pm, slightly convex;

broad (up to 68 pm) + fiat

85-120 pm; random on coen-

osteum, ampullae, lid +

upper outer pseudosepta

2.1-2.6 mm, circular to ellipti-

cal

14-23, 19.38, 21 (47)

tongue-shaped; 30-40 %;

horizontal + concave

narrow (1/4 dactylotome

width); sligthly concave

B(?)-C4 + A2

250-350 pm, convex to

carinate; broad + fiat or

irregular around

cyclosystem

220 pm; random on coen-

osteum and lid edge

3.8 mm, circular to irregu¬

lar

15-27, 19.75, 19 (529)

tongue-shaped; 60-90 %;

horizontal + concave

narrow (1/2 dactylotome

width); not concave

A-C4 + Al

Table 3. — Comparions

of eastern Atlantic Crypihelia

female efferent pore

enormous

Hickson & England (1905) were convinced that the species described in detail by Moseley

was not C. pudica, and therefore renamed it C. moseleyi. In fact, it must bear the name C. affinis,

which had already been ressurected by Boschma (1951b) and subsequently used by him in other

papers.

Boschma (1956b) reproduced (partly) a figure from Moseley (1881: pl. 12, fig. 7) but

erroneously indicated the distribution of C. affinis as the West Indies. La ter (Boschma, 1968a), when

mentioning C. affinis as a distinct species of the eastern Atlantic, he mistakenly included material

from “ Talisman ” drag. 128 (mnhn), which, in fact, is referable to C. tenuiseptata and C.

vascomarquesi.

Distribution and ecology

C. affinis has been obtained from several stations in the Azores, depth 712-1557 m. Specimens

were alive as deep as 1300 m.

According to Moseley (1879, 1881) the types of C. affinis (including live material) came from

“ Challenger ” stn 3, far Southwest of the Canary Islands, at a depth of 2790 m. This is the greatest

depth ever recorded in the literature for a stylasterid coral, and about twice the depth at which C.

affinis has been obtained in the Azores. The “ Challenger ” locality and depth should therefore be

viewed with circumspection; a confusion of the collecting locality is not unlikely. Regardless, the types

of C. affinis apparently came from the eastern Atlantic; the species is not known from the western

Atlantic where several other représentatives of the genus Crypthelia occur.

No symbionts are known.

Source : MNHN, Paris

112

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Crypthelia medioatlantica new species

Fig. 36 J-M, 38 A-H

Types

Holotype (Fig. 36 J) a délicate subterminal male branch from “ Bartlett ” 1975, stn 14 (zmuk).

Originally 23 mm high and 6 mm wide, comprising 17 cyclosystems, it was subsequently fractured for

sem studies. Paratype (Fig. 36 M) a small branch with 5 cyclosystems from Prince of Monaco stn 242

(mom).

Type locality: “ Bartlett ” 1975, stn 14, 16.12.1975, 36°50.9'N, 32°57.9'W, 1400-2200 m.

Mid-Atlantic Ridge.

Material studied

Azores: Paratype.

Mid-Atlantic Ridge: At 36°50.9'N, Holotype. — “ Nautile ” 1988, cruise Hydrosnake dive

hs-16, 3 small colonies, dead and manganèse coated, largest 34 mm high, with 4, 5, 9 cyclosystems,

respectively + 3 isolated cyclosystems (mnhn; usnm 88325).

Description

Only small pièces comprising a total of 42 well

preserved cyclosystems (+ one incomplète),

from 3 stations in 3 areas are available for the

description of this species.

Colonies uniplanar, probably small and déli¬

cate. No complété colony available; largest spé¬

cimen (holotype) with 17 cyclosystems. Subter¬

minal branches only 0.45 mm in diameter.

Coenosteum exclusively linear-imbricate in tex¬

ture (Fig. 38 C-E). Strips 70-85 pm wide and

slightly convex, covered with numerous quite

narrow platelets 4-14 pm wide. Nematopores not

observed.

Cyclosystems circular, 1.2-1.4 mm in diameter

(Fig. 36 K-L). Based on 42 cyclosystems (from 3

stations), there is a range of 14-19 dactylopores

per cyclosystem, mean 16.12, and mode 15.

Maximum width of upper gastropore chamber

about 0.60 mm, which leads to a gastropore ring

constriction of about 0.35 mm width (Fig. 38 G).

Lower chamber about 0.65 mm wide at greatest

diameter and about 0.10 mm deep. Cyclosystem

lid digitiform and inclined upwards, covering

0-20 % of the cyclosystem (the lid is sometimes

absent). Lid about 0.30 mm wide. Dactylotomes

about 0.10 mm wide; pseudosepta of equal width

and quite concave (Fig. 36 K-L, 38 A-B, H).

Male ampullae discrète, conspicuous hemi-

spheres (Fig. 38 A-B, F) about 0.6 mm in

diameter, 1-4 of which occur around each cy¬

closystem wall starting in the proximal cyclosys¬

tem area adjacent to lid (arrangement Cl of

Cairns, 1986b). Ampullae covered by a reti-

culate System of carinae, about 60 pm high,

which also extends to upper lid. At apex of each

male ampulla is a shallow dépression about 0.18

mm in diameter, in the center of which is a small

raised papilla about 0.10 mm in diameter. The

papilla has an apical pore 40 pm in diameter: the

male efferent pore (Fig. 38 A). Female ampullae

unknown; however, the 6 other species of Crypt¬

helia with Cl-type male ampullae ail hâve B-type

female ampullae, i.e. ampullae confined to proxi¬

mal cyclosystem wall, with an efferent pore

opening into upper gastropore chamber beneath

lid.

COMPARISONS

Of the 9 Atlantic species of Crypthelia (see Cairns, 1986a), only one other has Cl-type male

ampullae: C. affinis. Both species are also similar in their average number of dactylopores per

cyclosystem, coenosteal strip width, and morphology of pseudosepta and gastropore chamber.

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

113

Fig. 38. - Crypthelia mejloatlmllca (A-H, pan of holotype, zmuk):_A-B, cyctaystem surroundcd by male ampiÿtoe, efcanl

nores visible lid broken in A ( x 34, x 38, respectively, A being a stéréo pair); C-E, coenosteal texture ( x 73, 140,

x 389, respectively); F, oblique view of cyclosystem and male ampulla (x 49); G, longitudinally fractured cyclosyste

revealing gastropore chambers (x 37); H, pseudosepta (x 125).

Source : MNHN, Paris

114

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

C. medioatlantica is distinguished most readily by its smaller cyclosystems and branch diameter; its

smaller, inclined lids; and its differently shaped platelets and male ampullae.

Remarks

The spécifie name given to the new species refers to its géographie distribution.

Distribution and ecology

C. medioatlantica is known only from the Azores (861 m) and from the Mid-Atlantic Ridge

at 36°50.9'N (1400-2200 m) and at 23°31'N (2644 m).

No symbionts are known.

Crypthelia vascomarquesi new species

Fig. 39 A-J, 40 A-J

Chresonymy:

Crypthelia affinis — Boschma, 1968a: 106 (part: “ Talisman”).

Crypthelia — Zibrowius & Cairns, 1982: 212 (“ unnamed ”, part: Hyères Seamount, Azores/part,

Madeira).

Crypthelia tenuiseptata — Cairns, 1986a: 117 (part, not Hyères Seamount and Madeira).

Types

Ail specimens available from the Azores are given type status: “ Talisman ” drag. 128, 3 small

subterminal branch fragments comprising 6, 4, and 4 cyclosystems, respectively, the largest one

designated holotype (Fig. 39 C-D), the smaller ones paratypes (mnhn). — Prince of Monaco, stn 242,

small branch with 7 cyclosystems (paratype, mom). — “ Jean Charcot ” 1971, cruise biaçores 1971,

stn 232, 3 small dead fragments, with a total of 5 cyclosystems (paratypes, mnhn).

Type locality: “ Talisman ” drag. 128, 16.8.1883, 38°07'N, 27°11'45"W, 983 m. Azores,

between Sào Miguel and Faial.

Material studied

Azores: Holotype and paratypes (see above).

Hyères Seamount: “ Calypso ” 1959, drag. 4, small branch with originally 7 cyclosystems (Fig.

39 A-B) subsequently fractured (5 cyclosystems mnhn; 2 cyclosystems usnm 75622).

Madeira Archipelago: “ Jean Charcot ” 1966, stn 12, small fragment with 1 cyclosystem

(mnhn); stn 17, small branch with 4 cyclosystems (mnhn); stn 19, 6 small fragments with a total of

10 cyclosystems (mnhn).

Description

Only 16 small pièces comprising a total of 48

cyclosystems from 7 stations in 3 widely separa-

ted areas are available for the description of this

species.

Colonies uniplanar, probably small and déli¬

cate; known only from subterminal branch frag¬

ments with one to a few cyclosystems (Fig. 39

A-J); tallest fragment about 15 mm, bifurcate,

and comprising 7 cyclosystems. Subterminal

branches very thin (0.5-0.7 mm in diameter),

particularly relative to the large cyclosystems

they support. Coenosteal strips broad and

slightly convex (Fig. 40 B), some up to 0.12 mm

wide. Platelets well defined and fiat (Fig. 40 C),

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

115

Fig. 39. — Cryplhelia vascomarquesi (A-B, from “ Calypso ” 1959. drag. 4. mnhn; C-F, from “ Talisman " drag. 128. mnhn;

G-H, from Prince of Monaco stn 242, mom; I. from “ Jean Charcot ” 1966, stn 19, mnhn; J, from “ Jean Charcot " 1966,

stn 17, mnhn): A-B, anterior and posterior branch faces with bulging female ampullae (both x 3.6); C, holotype

(x 4.5); D, part of C. oblique side view (x 4.2); E-F, two views of paratype branch with four cyclosystems (x 4.5,

x 4.2, respectively); G-H, two views of paratype branch ( x 3.9); I, branch ( x 4.2); J, branch ( x 4.2).

10-68 pm wide. Large, shallow nematopores,

85-120 pm in diameter occur randomly over

coenosteum, ampullae, and lids, and in a more

orderly manner on almost every upper outer

pseudoseptum (Fig. 40 A, H-I).

Cyclosystems circular to elliptical: circular

cyclosystems 2.1-2.6 mm in diameter, elliptical

ones about 2.2-2.5 x 1.6-2.0 mm. Cyclosystems

very exsert on branch, being elevated by the

underlying ring like ampullae (Fig. 39 B, D, F,

40 D, G-H). Based on 47 cyclosystems (out of

48), there is a range of 14-23 dactylopores per

cyclosystem, mean 19.38, and mode 21.

Maximum width of upper gastropore chamber

about 0.83 mm; gastropore ring constriction

about 0.65 mm in diameter. Lower chamber

about 1.0 mm wide, with very fiat floor. Cy¬

closystem lid tongue-shaped and horizontal, co-

vering 30-40 % of cyclosystem (Fig. 39 A, C).

Lid up to 1.1 mm wide and highly concave.

Dactylotomes wide (0.16 mm); pseudosepta cor-

respondingly narrow, about 0.045 mm wide, and

only slightly concave. Pseudoseptal edges slightly

flared, overhanging underlying ampullar bulge

(Fig. 40 H-I).

Male ampullae produce a continuous band

encircling cyclosystem and also extend into

proximal lid (Fig. 40 A, D-E). Each ampulla

appears to be 0.55 mm in diameter, discrète in

young cyclosystems but merging into a conti¬

nuous band in mature cyclosystems. Each am¬

pulla has an efferent pore about 70 pm in

diameter, which opens into a contiguous dacty¬

lotome within the cyclosystem. Mature cyclosys¬

tems hâve efferent pores opening into almost

every dactylotome (Fig. 40 F). Presumed female

Source : MNHN, Paris

116

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Fig. 40. — Crypthelia vascomarquesi (A, D-F, H-I, male paratype from “ Talisman ” drag. 128, mnhn; B-C, G, J, female (?)

paratype from Prince of Monaco stn 242, mom): A, distal branch with fractured cyclosystem encircled by male ampullac,

pseudoseptal and coenosteal nematopores présent (x 11); B-C, coenosteal texture (x 61, x 140, respectively); D.

oblique view of male cyclosystem (x 20, stéréo pair); E, longitudinally fractured cyclosystem revealing gastropore

chambers and male ampullae (x 19); F, pseudoseptal area within cyclosystem showing male efferent pore in middle

dactylotome (x 53); G, J, female ( ?) cyclosystems and ampullae ( x 24, x 26, respectively); H, side view of male

cyclosystem illustrated in D (x 19); 1, side posterior view of cyclosystem illustrated in fig. D and H (x 19).

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

117

ampullae (Fig. 39 B, 40 G, J) more discrète,

larger, hemispherical structures up to 1 mm in

diameter. One to six female ampullae occur per

cyclosystem, concentrating near proximal cy-

closystem wall and proximal lid région. Female

efferent pores also appear to open to a dactylo¬

tome within the cyclosystem; however, more

specimens are needed to fully understand the

ampullar arrangement of this species.

COMPARISONS

C. vascomarquesi can be distinguished from the other Atlantic congeners by its distinctive

ampullar formula (B-C4 + A2) and various other characters (see Table 3). Within the Atlantic, it is

most similar to the western Atlantic C. glossopoma Cairns, 1986, both species sharing the same

ampullar formula (the only 2 of 28 species in the genus) and having similar cyclosystem lids, large

nematopores, and very slender, flared pseudosepta. C. vascomarquesi is distinguished by its larger

cyclosystems, thinner terminal branches, better defined platelet structure, and more prominent

ampullae with male efferent pores occurring around the entire circumference of the cyclosystem.

Remarks

This new species is named in memory of Vasco Monteiro Marques, Portuguese marine

biologist (14.9.1953-19.12.1985).

Having previously resurrected the name Crypthelia affinis Moseley, 1879, for another eastern

Atlantic stylasterid, Boschma (1968a) incorrectly attributed to that species material from the

“ Talisman ” expédition (drag. 128) preserved at the mnhn; in reality, this lot consisted of C.

vascomarquesi and C. tenuiseptata.

Records of C. vascomarquesi from Hyères Seamount and Madeira Archipelago had previously

been included under C. tenuiseptata by Cairns (1986a).

Distribution and ecology

C. vascomarquesi is known from the Azores, the Hyères Seamount, and the Madeira

Archipelago, depths respectively 390-983 m, 600 m, and 990-1520 m.

The species is probably more widely distributed in the northeastern Atlantic. A small branch

of Crypthelia, badly preserved, from near Selvagem Grande (“ Tydeman ” 1983, cruise cancap 3 stn

3.099, 585 m; rmnh) possibly belongs to C. vascomarquesi. Likewise, the unnamed Stylasteridae from

Joséphine Seamount (622 m) mentioned and schematically figured by Lindstrôm (1877: 15, pl. 2, fig.

25) could also be this species; unfortunately the tiny branch at the smnh was found entirely

decomposed into powder (R. Olerôd in litt., 1977).

No symbionts are known.

Crypthelia tenuiseptata Cairns, 1986

Fig. 41 A-J, 42 A-I

Synonymy:

Crypthelia tenuiseptata Cairns, 1986a: 115-117 (part, NOT Hyères Seamount, Madeira), fig. 52 A-G, 53

Chresonymy:

[?] Crypthelia pudica — Filhol, 1885: 268, pl. 1.

Crypthelia affinis — Boschma, 1968a: 106 (part: “ Talisman").

Crypthelia — Zibrowius & Cairns, 1982: 212 (“ unnamed ”, part: Azores/part. not Hyères Seamount,

Madeira).

Crypthelia tenuiseptata — Cairns, 1986b: 24-25.

Source : MNHN, Paris

118

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

F '°- “U'& «wr* D - " r '"™” *» 127 ». «™» ; E-i

penpheral branch ( x 4.3); C male branchTx 2 SV n Monaco stn 233, mom): A, female branch ( x 1.3 ); b,

ofE(x 3.1); G, massive colony(x 0.8); H deUH S’g (x 3 l ‘\ k°' 9)î Aw°i° ny ( * °-8); f » detail

colony ( x 0.6). ' A ’ aetal1 01 0 ( x 3.1), I, penpheral branch ( x 4.0); J, detail of massive dead

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

119

Types

Crypthelia tenuiseptata : The original description (Cairns, 1986a) is based on 3 small branches

(up to 21 mm high and 25 mm wide) from 3 stations in the western Atlantic (“ Blake ” 1878/79, stn

131, Santa Cruz; stn 230, St. Vincent; stn 264, Grenada). These are the designated holotype (stn 264)

and paratypes, ail deposited at mcz and usnm (71812 from stn 230).

Type locality: “ Blake ” stn 264, 1.3.1879, 12°03'15"N, 61°48'30" E, 761 m. Grenada.

Material studied

W Atlantic: holotype and paratypes of Crypthelia tenuiseptata (see above).

Azores: “ Talisman ” drag. 127 (?), 3 branches (mnhn); drag. 128, ca. 20 branches + minor

fragments (mnhn). — Prince of Monaco, stn 203, 18 dead branches + fragments (mom); stn 233, big

colony (mom); stn 616, 3 colonies + branches (mom). — “ Jean Charcot ” 1971, cruise biaçores stn

180, 10 colonies, branches + fragments (most mnhn; usnm 75623); stn 196, ca. 10 dead fragments

(mnhn); unknown station, colony (mnhn).

Description

Colonies uniplanar (Fig. 41 A, D, E, G);

well-preserved specimens up to 92 mm high and

69 mm wide; a poorly preserved incomplète

colony (Prince of Monaco stn 233) was conside-

rably larger: it is still 80 mm high and 72 mm

wide, and measures 20 mm across the encrusting

base and 12 by 15 mm in the lower part of the

massive trunk (Fig. 41 J). Branches with inversed

unifaciality of cyclosytem arrangement occasio-

nally occur. Branches often anastomose; subter¬

minal branch diameter about 0.9 mm. Coenos-

teal strips extremely wide (250-350 pm), and

convex to carinate along strip midline (Fig. 41

H). Platelets broad and fiat but sometimes

irregular in size and shape near cyclosystems

(Fig. 42 I). Nematopores extremely large and

shallow (Fig. 42 C), about 220 pm in diameter,

occurring randomly on branch coenosteum and

along lid edge. Invariably there is a tiny coenos-

teal pit in the center of the nematopore.

Cyclosystems circular to irregular in shape, up

to 3.8 mm in diameter. Old cyclosystems flush

with coenosteum (Fig. 41 F, H); younger cy¬

closystems only slightly exsert (Fig. 41 B-C, I).

Based on 502 eastern Atlantic (Azores) cyclosys¬

tems, there is a range of 15-27 dactylopores per

cyclosystem, mean 19.75, and mode 19.

Maximum width of upper gastropore chamber

about 1.1 mm; gastropore ring constriction

about 0.8 mm in diameter. Lower, compressed

chamber about 1.8 mm wide. Cyclosystem lid

tongue-shaped and horizontal, covering 60-90 %

of cyclosystem. Lid up to 2.3 mm wide and

ordinarily quite thin and concave unless inflated

with an ampulla. Dactylotomes quite wide (0.20

mm); pseudosepta narrow, about 0.1 mm wide,

and not concave.

Female ampullae massive, restricted primarily

to cyclosystem lid (Fig. 41 A, F, 42 B, E). Female

efferent pores circular and extremely large (0.5

mm in diameter), opening on underside of verti¬

cal segment of lid. Male ampullae form a

continuous, rather smooth bulge, encircling cy¬

closystem and occasionally extending into proxi¬

mal lid. Male efferent pores circular, about 0.13

mm in diameter, opening into adjacent dactylo¬

tome within cyclosystem (Fig. 42 C). Some

cyclosystems hâve one male efferent pore per

dactylotome (Fig. 42 C). Also shown in Fig. 21 C

is the apical efferent pore of the male ampulla in

the lid.

Comparions

C. tenuiseptata is distinguished from the other 8 Atlantic Crypthelia by its ampullar formula

(A-C4 + Al); other characters are listed in Table 3 and by Cairns (1986a). Only one other species

has an A-C4 ampullar formula: C. gigantea Fisher, 1938, from the Galapagos. They are also similar

Source : MNHN, Paris

120

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Fie. 42. — Crypthelia tenuiseptata (A-B. D-E, female from “ Jean Charcot ” 1971, stn 180, usnm 75623; C, F-I, male from

" Talisman ” drag. 127 ?, mnhn): A, branch segment illustrating cyclosystems, cyclosystem lid, gastropore chambers,

and gross coenosteal texture ( x 8.4, stéréo pair); B, oblique view of female cyclosystems with ampullar bulge in lid

( x 13); C, intact male cyclosystems showing numerous efferent pores opening into upper dactylotomes (x 11); D,

pseudosepta ( x 47); E, longitudinal fracture of female cyclosystem illustrating gastropore chambers and female ampulla

(x 10, stéréo pair); F, side posterior view of male cyclosystem of C (x 16); G, branch coenosteum with numerous

coenosteal pores and three shallow nematopores (x 44); H-l, coenosteal texture (x 124, x 397, respectively).

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

121

in having massive coralla with very large cyclosystems, wide coenosteal strips, and relatively thin

pseudosep ta. C. tenuiseptata is distinguished from C. gigantea by its very large nematopores (C.

gigantea has none), even broader coenosteal strips, and a lower average number of dactylopores per

cyclosystem (see Cairns, 1986b).

Remarks

C. tenuiseptata was described by Cairns (1986a) from 3 small branches from the western

Atlantic, where larger colonies are still unknown. Previously, it had been mentioned by Zibrowius

& Cairns (1982) as an unnamed amphiatlantic species of Crypthelia from 3 areas in the eastem

Atlantic (Azores, Hyères Seamount, Madeira). Although the species is amphiatlantic, the indicated

distribution was partly incorrect because it included a second form now distinguished as C.

vascomarquesi. In fact, in the eastem Atlantic the occurrence of C. tenuiseptata is confirmed only for

the Azores, whereas C. vascomarquesi also occurs in the other areas (Hyères Seamount, Madeira

Archipelago).

C. tenuiseptata, which had much earlier been collected in the Azores, had not been recognized

as a distinct species by previous authors. A colony of unknown origin (“ Talisman ”, depth 800 m)

figured by Filhol (1885) and misidentified as C. pudica, may well belong to C. tenuiseptata, which

indeed is represented in the “ Talisman ” collection by several large specimens.

Having previously resurrected the name Crypthelia ajfinis for another eastem Atlantic

stylasterid, Boschma (1968a) incorrectly attributed to that species material from the “ Talisman"

expédition (drag. 128) preserved at the mnhn; in reality, this lot consisted of C. tenuiseptata and C.

vascomarquesi.

Distribution and ecology

C. tenuiseptata is noteworthy as one of the rare amphiatlantic stylasterids. It is known from

the Virgin Islands, Lesser Antilles (761-1080 m), and from the Azores (983-1557 m). In the Azores

live specimens were obtained as deep as 1257 m.

No symbionts are known.

Source : MNHN, Paris

122

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

Note added in press

We refer to a large collection of stylasterids (currently being studied by H.Z.) obtained around

the Faroe Islands by the biofar project (1987-1990), in fact the richest collection ever obtained in high

latitudes of the North Atlantic, biofar found ail four species previously known from high latitudes

but not ail material is yet sorted: Pliobothrus symmetricus (8 stations), Stylaster norvegicus (45

stations), Stylaster gemmascens (21 stations), Stylaster erubescens (25 stations).

In addition, some samples of stylasterids from poorly collected areas (West Africa and Canary

Islands) were discovered in the collections of the zmuk and the Institute of Océanographie Sciences

(the latter samples now transferred to the bmnh).

Stenohelia maderensis

Sahara: “ Discovery ” stn 7975, 19.7.1972, 26°23.64'N, 14°51.10'W, 785-834 m, 5 dead

fragments, the largest 1 1 mm high (bmnh); stn 7984, 20.7.1972, 25°26.00' N, 16° 10.25' W, 811-890 m,

24 specimens (bmnh), ranging from a small incomplète colony with base (distal cyclosystems broken

ofl), 13 mm high and 10 mm wide through branchlets to fragments with only very few or even one

cyclosystem; few specimens alive, but some of the dead ones fresh-looking.

These records from the upper slope off West Africa fill in a wide gap of latitudinal distribution

between the Cape Verde Islands and Madeira.

Crypthelia vascomarquesi

Canary Islands: “ Dana ” stn 4011, 25.3.1930 (coll. Th. Mortensen), 9 nautical miles SE of

Las Palmas, Gran Canaria, 670-1100 m; small dead colony (zmuk) with base, 11 mm high, with 5

cyclosystems, the basalmost being filled in by sclerenchyme; distal cyclosystems broken off (missing);

two cyclosystems hâve 17 dactylopores, the other damaged ones also more than, respectively, 14 and

This record from the “ Dana ” circumnavigation is the first confirmed record of a stylasterid

from the Canary Islands. It extends the range of the species farther south (ca. 3 degrees of latitude).

Source : MNHN, Paris

LIST OF DEEP-WATER STATIONS FROM OCEANOGRAPHIC CRUISES

This is an inventory of stations from océanographie cruises in the northeastem Atlantic Océan,

in the straits of Gibraltar, and in the Mediterranean from which stylasterid corals and/or their

Pedicularia symbionts (or occasionally only the traces of Pedicularia), were available for the présent

study, or are quoted here from the previous literature.

Examples: “ Lepidopora eburnea + trace of Pedicularia ” means that only the trace, not the

symbiont itself, was found on the stylasterid; “ Errina atlantica + Pedicularia ” means that the

symbiont was found on its host; “ Errina dabneyi + trace of Pedicularia, Pedicularia ” means that

only the trace was found on the stylasterid, but that Pedicularia from this station was no longer

attached to a stylasterid host.

The vessels are listed in alphabetical order; under each vessel the cruises and stations are

chronologically arranged, following the current station numbers, either continuous for the vessel, or

distinct for each cruise. Prince of Monaco stands for ail the cruises carried out by Albert I, Prince

of Monaco, on several vessels. Whenever possible, the (main) depository of material is indicated for

each vessel or cruise.

The présent list summarizes the greater and more diversified part of the stylasterid records

from the investigated area. Additional records not from these cruises are found under “ material

studied ” of most species sections. These additional records comprise samples of imprécise origin in

old muséum collections, collections by fishermen, etc.

“ Al Mounir ” 1969 (collector J. Stirn; material in mnhn, usnm)

b6-d6: 2.7.1969, 35°55'00"N, 5°34'55"W, 350m, Straits of Gibraltar: Errina aspera

b8-d2: 5.7.1969, 35°54'15"N, 5°46'00"W, 365-390m, Straits of Gibraltar: Errina aspera

b10-d3: 21.7.1969, 33°43'30"N, 6 o 21'00"W, 200m, NW Morocco: Errina aspera + Pedicularia

“ Anton Dohrn ” (collector G. Behrmann; material in imfb)

stat. ?: 10.3.1972, 68°30'N, 12°19'E, 300m, Norway: Stylaster norvégiens

stat. ?: 18.11.1973, 63°00'N, 6°30'W, 1040m, Faroes: Stylaster norvegicus

stat. ?: 19.11.1973, 62°07'N, 6°27'W, 75m, Faroes: Stylaster norvegicus

“ Bannock ” 1972, cruise joti 72: Straits of Messina (collector P. Colantoni)

23: 7.1972, 38°14.56'N, 15°37.7'E, 95m: Errina aspera + Pedicularia

34: 7.1972, 38°12.7'N, 15°36.4'E, 220m: Pedicularia (fide Selli et al., 1980; stylasterid host not recorded,

but undoubtedly Errina aspera)

47: 7.1972, 38°17.2'N, 15°44.0'E, 329m: Pedicularia (same remark as for stn. 34)

“ Bartlett ” 1975 (collector J. Knudsen; material in usnm)

2: 30.1.1975, 37°13.8'N, 28°44.5'W, 480m, Azores: Lepidopora eburnea + trace of Pedicularia

4: 3.2.1975, 38°09.8'N, 28°53.4'W, 750-800m, Azores: Lepidopora eburnea + trace of Pedicularia

14: 16.2.1975, 36°50.9'N, 32°57.9'W, 1400-2200m, Mid-Atlantic Ridge: Errina dabneyi, Crypthelia

medioatlantica

“ Bartlett ” 1975 (collector Byerly; material in usnm)

52c-5: 23.9.1975, 61°59.7'N, 26°35’-36'W, 585m, SW Iceland, Reykjanes Ridge: Stylaster erubescens

groenlandicus

“ Calypso ” 1958 (material in mnhn)

sme-1277: 28.8.1958, approx. 36°30'N/11°30'W, 510m, Gorringe Seamount: Lepidopora sp. A

sme-1282: 30.8.1958, 35°54'N, 6°00'W, 110m, NW Morocco, Spartel Bank: Errina aspera + Pedicularia

Source : MNHN, Paris

124

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

“ Calypso ” 1959 (collector G. Bellan; material in mnhn, usnm)

drag. 4: 13.8.1959, 31°26.6'N, 28°55.4'W, 600m, Hyères Seamount: Pliobothrus gracilis, Crypthelia

vascomarquesi

drag. 6: 13.8.1959, 31°27.7'N, 28°55.6'W, 620-700m, Hyeres Seamount: Pliobothrus gracilis

“Calypso" 1959: Cape Verde Islands (material in mnhn)

16: 17.11.1959, W Sào Tiago, NW Ponta Geneanes, 235-400m: Stenohelia maderensis

91: 27.11.1959, 15°34.5'N, 23°U.5'W, 185m, between Maio and Boavista: Stenohelia maderensis

“ Chain " 1959, cruise 7 (material in usnm)

pd-24: 31.7.1959, 30°00'N, 28°25'N, 295m, Great Meteor Seamount: Stylaster erubescens meteorensis

“Challenger " 1873 (material in bmnh)

3: 8.2.1873, 25°45'N, 20°12'W, 2795m, far SW Canary Islands: Crypthelia affinis

85: 19.7.1873, 28°42'N, 18°06'W, 2100m, Canary Islands, W Palma: Lepidopora sp. B

“ Challenger II" 1977 (collector J.D. Gage; material in bmnh)

6a- 134: 9.4.1977, 54°05'N, 12°06'W, 800m, W Ireland: Pliobothrus symmetricus

“ Challenger II" 1981 (collector T.P. Scoffin; material in bmnh)

30: 7.1981, 53°16.76'N, 14°35.47'W, 520m, Porcupine Bank: Pliobothrus symmetricus, Stylaster sp.

“ Cryos" 1984, cruise balgim (collector H. Zibrowius; material in mnhn)

dr-37: 1.6.1984, 36°17.8'N, 7°15.4'W, 860-868m, western approaches Straits of Gibraltar: Pedicularia

(stylasterid host unknown, most likely Errina asperà)

dr-40: 2.6.1984, 35°49.9'N, 6°08.6'W, 362m, Straits of Gibraltar, W sill: Errina aspera, Pedicularia

dr-49: 3.6.1984, 35°53.0'N, 6°32.8'W, 524-578m, western approaches Straits of Gibraltar: Errina aspera

dw- 50: 3.6.1984, 35°52.7'N, 6°31.9'W, 518-524m, western approches Straits of Gilbraltar: Pedicularia

(stylasterid host unknown, most likely Errina aspera)

cp-95: 8.6.1984, 34°24.0'N, 7°39.3'W, 1378m, NW Morocco: Lepidopora sp. A, Stylaster maroccanus

dr-115: 11.6.1984, 35°47.5'N, 6°04.2'W, 332m, Straits of Gibraltar, W sill: Errina aspera

dr-116: 11.6.1984, 35°48.6'N, 6°04.2'W, 322-365m, Straits of Gibraltar, W sill: Errina aspera

dr-153: 17.6.1984, 35°55.8’S, 5°35.3'W, 568-604m, Straits of Gibraltar, E sill: Pedicularia (stylasterid host

unknown, most likely Errina aspera)

“ Dana ” 1938 (material in zmuk)

5835: 14.5.1938, 62°35'N, 7°52'W, 400m, W Faroes: Stylaster gemmascens

6001: 24.7.1938, 63°33'N, 11°25'W, 322m, SE Iceland: Stylaster erubescens (britannicus 7)

6005: 25.7.1938, 62°19'N, 8°51'W, 475-504m, W Faroes: Stylaster gemmascens

6009: 26.7.1938, 61°14'N, 7°04'W, 220m, S Faroes: Stylaster gemmascens

“Gazelle ” 1874: Madeira Archipelago (material in zmb)

2: 16.7.1874, S Madeira, 2 miles off shore, 110-128m: Stenohelia maderensis

“ Ingolf " 1895-1896 (material in zmuk)

1: 11.5.1895, 62°30'N, 8°21'W, 249m, W Faroes: Stylaster gemmascens

2: 12.5.1895, 63°04'N, 9°22'W, 493m, NW Faroes: Stylaster gemmascens

7: 17.5.1895, 63°13'N, 15°41'W, 1130m, SE Iceand: Stylaster erubescens groenlandicus

15: 4.6.1895, 66°18'N, 25°59'W, 621m, NW Iceand: Stylaster norvégiens, S. gemmascens (both fide Broch,

1914a), S. erubescens groenlandicus

16: 5.6.1895, 65°43'N, 26°58'W, 471m, NW Iceland: Stylaster gemmascens, S. erubescens groenlandicus

17: 6.6.1895, 62°49'N, 26°55'W, 1403m, SW Iceland: Stylaster norvegicus, S. erubescens (groenlandicus 2)

(both fide Broch, 1914a)

52: 15.5.1896, 63°57'N, 13°32'W, 791m, SE Iceland: Stylaster norvegicus, S. erubescens groenlandicus

55: 19.5.1896, 63°33'N, 15°02'W, 595m, SE Iceland: Pliobothrus symmetricus, Stylaster norvegicus (both

fide Broch, 1914a)

57: 20.5.1896, 63°37'N, 13°02'W, 659m, SE Iceland: Pliobothrus symmetricus (fide Broch, 1914a)

94: 26.6.1896, 64°56'N, 36°19'W, 384m, E Greenland, off Angmagssalik: Stylaster gemmascens, S.

erubescens (groenlandicus ?) (fide Broch, 1914)

144: 11.8.1896, 62°49'N, 7°12'W, 520m, N Faroes: Stylaster erubescens britannicus

“ Jean Charcot” 1966, cruise zarco: Madeira Archipelago (material in mnhn)

12: 13.7.1966, 32°36.2'N, 17°07.7'W, 1520m, S Madeira: Crypthelia vascomarquesi

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

125

17: 15.7.1966, 32°58.5'N, 16°26.5'W, 1630-1690m, SW Porto Santo: Crypthelia vascomarquesi

19: 15.7.1966, 33°00.2'N, 16°20.0'W, 990m, SW Porto Santo: Crypthelia vascomarquesi

21: 15.7.1966, 33°01.2'N, 16°24.9'W, 220-290m, SW Porto Santo: Stenohelia maderensis

29: 16.7.1966, 33°01.4'N, 16°15.5'W, 300-340m, SW Porto Santo: Stenohelia maderensis + trace of

Pedicularia

49: 18.7.1966, 32°27.3'N, 16°32.0'W, 450-490m, SW Deserta: Pliobothrus symmetricus ?, Stenohelia

maderensis

“ Jean Charcot ” 1971, cruise biaçores: Azores (collector H. Zibrowius; material in mnhn, usnm)

25: 9.10.1971, 38 0 21'N, 28°49.5'W, 800-1020m: Crypthelia affinis

34: 10.10.1971, 38°09.5'N, 29°15.0'W, 650-670m: Pliobothrus symmetricus

49- 12.10.1971, 37°56'N, 29°12'W, 215-225m: Errina dabneyi + trace of Pedicularia, Pedicularia

74: 15.10.1971, 38°28.5'N, 27°54.5'W, 1235-1310m: Crypthelia affinis

102: 19.10.1971, 39°30'N, 31°04.5'W, 712-750m: Crypthelia affinis

112: 20.10.1971, 39°34'N, 31°19.5'W. 806-825m: Crypthelia affinis

135: 25.10.1971, 39°24.5'N, 31°05.5'W, 760-860m: Crypthelia affinis

159: 31.10.1971, 37°26'N, 25°51'W, 525-600m: Lepidopora eburnea

161: 31.10.1971, 37°39.5'N, 25°50.5'W, 590m: Lepidopora eburnea + trace of Pedicularia, Pliobothrus

symmetricus

180: 3.11.1971, 37°57.5'N, 25°33'W, 1069-1235m: Crypthelia affinis, C. tenuiseptata

196: 5.11.1971, 37°50'N, 24°55.5'W, 1146-1191m: Crypthelia tenuiseptata

197: 5.11.1971, 37°49.5'N, 25°01.5'W, 815m: Pliobothrus symmetricus, Errina atlantica + trace of

Pedicularia, Pedicularia

212: 7.11.1971, 37°18'N, 24°45.5'W, 610m: Errina atlantica

213: 7.11.1971, 37°21.5'N, 24°32.5'W, 895m: Lepidopora eburnea + trace of Pedicularia, Pliobothrus

symmetricus

218: 8.11.1971, 36°54'N, 25°08'W, 772-800m: Pliobothrus symmetricus

229: 10.11.1971, 37°01.5'N, 25°14'W, 600m: Lepidopora eburnea + trace of Pedicularia, Pliobothrus

symmetricus

230: 10.11.1971, 36°54'N, 25°09.5'W, 665-712m: Errina atlantica

231: 10.11.1971, 36°55'N, 25°10'W, 380-440m: Pliobothrus symmetricus

232: 10.11.1971, 36°55'N, 25°H'W, 390-620m: Pliobothrus symmetricus, Crypthelia vascomarquesi

238: 11.11.1971, 37°25'N, 25°45'W, 506m: Pliobothrus symmetricus

240: 12.11.1971, 37°35'N, 25°32.5'W, 810-825m: Pliobothrus sp., Errina atlantica + Pedicularia

stat. ?: 1971, Azores: Lepidopora eburnea + trace of Pedicularia, Stenohelia sp. A

“ John Murray ” 1972 (collector E.J.W. Jones; material in bmnh)

site 4/dredge: 3./4.1972, 57°24'N, 10°45'W, 1500m, Anton Dohrn Seamount: Stylaster erubescens

britannicus

“ Joséphine ” 1869: Joséphine Seamount (material in smnh)

stat. ?: 1869, approx. 36°46'N/14°07'W, 622m: Crypthelia sp. (fide Lindstrôm, 1877)

“ Meteor ” 1970, cruise Ml 9: Great Meteor Seamount (material in zsm)

129/dd- 94: 17.2.1970, 29°59'N, 28°33'W, 293-296m: Stylaster erubescens meteorensis

129/DD-95: 17.2.1970, 30°00'N, 28°31.5'W, 293m: Stylaster erubescens meteorensis

131/DD-98: 17.2.1970, 30°06'N, 28°33'W, 303m: Stylaster erubescens meteorensis

“ Michael Sars ” 1902: Faroes (material in vsm)

43: 4.7.1902, 62°31'N, 5°14'W, 320m, NE Faroes: Stylaster gemmascens

stn ?: 25.2.1904, 16 miles E Faroes, 376m: Pliobothrus symmetricus

submersible “ Nautile ” 1988, cruise hydrosnake (material in mnhn, usnm)

hs- 16: 5.7.1988, 23°31'N, 45°10'W, 2644m, Mid-Atlantic Ridge: Crypthelia medioatlantica

“ N or oit ” 1987, cruise seamount (collector H. Zibrowius; material in mnhn)

dw-8: 22.9.1987, 36°28.5'N, 11°37.1'W, 470-485m, Gorringe Seamount: Lepidopora sp. A

de- 10: 23.9.1987, 36°27.4'N, 11°35.0'W, 500-545m, Gorringe Seamount: Lepidopora sp. A, Pedicularia

cp- 11: 23.9.1987, 36°26.4'N, 11°40.2'W, 805-830m, Gorringe Seamount: Lepidopora sp. A

CP-12: 23.9.1987, 36°24.2'N, 11°43.2'W, 1005-1040m, Gorringe Seamount: Lepidopora sp. A

de-13: 23.9.1987, 36°23.0'N, 11°42.5'W, 1110-1180m. Gorringe Seamount: Lepidopora sp. A, Errina

atlantica ? . _ .

dw-21: 24.9.1987, 36°34.9'N, 11°28.4'W, 460-480m, Gorringe Seamount: Lepidopora sp. A, Errina

atlantica ?

CP-30: 26.9.1987, 36°44.3'N, 11°23.0'W, 1940-2075m, Gorringe Seamount: Lepidopora sp. A,

Source : MNHN, Paris

126

HELMUT ZIBROWIUS & STEPHEN D. CAIRNS

dw-56: 7.10.1987, 36°42.3'N, 14°21.6'W, 360-425m, Joséphine Seamount: Pliobolhrus symmetricus

dw-58: 7.10.1987" 36°45.9'N, 14°20.4'W, 340-380m, Joséphine Seamount: Pliobothrus symmetricus

dw-78: 10.10.1987, 33°48.7'N, 14°22.6'W, 235m, Seine Seamount: Lepidopora sp. A

de-98: 12.10.1987, 35°03.2'N, 12°55.4'W, 300-325m, Ampère Seamount: Pedicularia (stylasterid host

unknown)

dw-108: 19.10.1987, 42°50.9'N, 11°53.1'W, 1100-1125m, Galicia Seamount: Lepidopora sp. A, Stenohelia

maderensis

dw-111: 19.10.1987, 42°39.9'N, 11°35.8'W, 675-685m, Galicia Seamount: Lepidopora sp. A, Pliobothrus

symmetricus ?, Pedicularia

dw-116: 20.10.1987, 42°52.44'N, 11°50.6'W, 985-1000m, Galicia Seamount: Lepidopora sp. A, Stenohelia

maderensis, Pedicularia

submersible “ Pisces III" 1973 (collector J.B. Wilson; material in bmnh)

P73-5: 24.6.1973, 57°54.9'N, 13°52.3'W, 160-190m, Rockall Bank: Stylaster gemmascens

“ Porcupine" 1869: between Faroes and Hébrides (material in bmnh)

54: 1869, 59°56'N, 6°27'W, 665m: Stylaster norvegicus, S. gemmascens, S. erubescens britannicus,

Stenohelia maderensis

“ Poséidon ” 1990, cruise 175/1: Denmark Strait (collector A. Freiwald; material in smf, usnm)

12/1: 18.10.1990, 65°26.76'N, 30°50.31'W, 477m: Stylaster norvegicus, S. gemmascens, S. erubescens

groenlandicus

14/1: 18.10.1990, 65°29.70'N, 30°01.92'W, 433m: Stylaster norvegicus, S. gemmascens, S. erubescens

groenlandicus

Prince of Monaco 1888-1905 (material in mom)

203: 30.7.1888, 39°27'05"N, 30°55'05"W, 1557m, Azores: Crypthelia affinis, C. tenuiseptata

229: 16.8.1888, 38°22'N, 28°14'24"W, 736m, Azores: Lepidopora eburnea, Pedicularia

233: 18.8.1888, 38°33'21"N, 28°08'39"W, 1300m, Azores: Crypthelia affinis, C. tenuiseptata, Pedicularia

242: 22.8.1888, 38°48'30"N, 27°58'45"W, 861m, Azores: Crypthelia medioatlantica, C. vascomarquesi

247: 30.8.1888, 38°24'N, 28°01'25''W, 318m, Azores: Errina dabneyi + trace of Pedicularia, Pedicularia

553: 3.7.1895, 37°42'40"N, 25°05'15"W, 1385m, Azores: Pedicularia (stylasterid host unknown)

568: 11.7.1895, 37°54'N, 25°35'25"W, 550m, Azores: Pliobolhrus symmetricus (fide Calvet, 1911)

584: 16.7.1895, 38°31'N, 26°49'15"W, 845m, Azores: Pliobothrus symmetricus (fide Calvet, 1911),

Pedicularia

597: 23.7.1895, 38°27'N, 28°03'25"W, 523m, Azores: Lepidopora eburnea, Pliobothrus symmetricus (fide

Calvet, 1911), Pedicularia

616: 1.8.1895, 38°47'40"N, 28°17'05"W, 1022m, Azores: Pliobothrus symmetricus (fide Calvet, 1911),

Crypthelia tenuiseptata

618: 1.8.1895, 38°52'45"N, 28°06'00"W, 1142m, Azores: Pliobothrus symmetricus (fide Calvet, 1911)

683: 7.7.1896, 38°20'N, 28°04'45"W, 1550m, Azores: Pliobothrus symmetricus (fide Calvet, 1911)

712: 24.7.1896, 39°39'40"N, 31°00'55"W, 1424m, Azores: Pliobothrus symmetricus (fide Calvet, 1911)

719: 27.7.1896, 39°11'N, 30°24'15"W, 1600m, Azores: Pliobothrus symmetricus (fide Calvet, 1911)

838: 22.7.1897, 37°55'N, 25°22'45"W, 880m, Azores: Pliobothrus symmetricus (fide Calvet, 1911)

866: 2.8.1897, 38°52'50"N, 27°23'05"W, 599m, Azores: Pliobothrus symmetricus (fide Calvet, 1911)

869: 3.8.1897, 39°03'N, 27°42'45"W, 1240m, Azores: Pliobothrus symmetricus (fide Calvet, 1911)

1349: 19.8.1902, 38°35'30"N, 28°05'45"W, 1250m, Azores: Pedicularia (stylasterid host unknown)

1713: 1.8.1904, 28°04'N, 16°49'30"W, 1320-1530m, Canary Islands, between Tenerife and Gomera:

Pedicularia (stylasterid host unknown)

2210: 1.9.1905, 39°25'N, 31°22'30"W, 1229m, Azores: Pedicularia (stylasterid host unknown)

“ Talisman" 1883 (material in mnhn, bmnh)

drag. 96: 15.7.1883, 19°19'N, 18°01'45"W, 2320-2330m, Mauritania: Lepidopora sp. A

drag. 103: 23.7.1883, Cape Verde Islands, S Sâo Tiago, off Praia, 150-275m: Stenohelia maderensis

drag. 114: 30.7.1883, 16°51'N, 25°09'45"W, 598-633m, Cape Verde Islands: Errina aspera + trace of

Pedicularia

drag. 123: 13.8.1883, 38°23'N, 28°49'45"W, 560m, Azores: Lepidopora eburnea + trace of Pedicularia,

Pliobothrus symmetricus, Errina dabneyi

drag. 127: 15.8.1883, 38°38'N, 28°20'45"W, 1257m, Azores: Crypthelia tenuiseptata

drag. 128: 16.8.1883, 38°07'N, 27°11'45"W, 983m, Azores: Lepidopora eburnea, Errina atlantica + trace

of Pedicularia, Crypthelia vascomarquesi, C. tenuiseptata, Pedicularia

drag.129: 16.8.1883, 38°00'N, 27°02'45"W, 2155-2220m, Azores: Pedicularia (confused, from drag. 128?;

stylasterid host unknown)

Source : MNHN, Paris

NE ATLANTIC AND MEDITERRANEAN STYLASTERIDAE

127

“ Thalassa" 1967-1973 (1972-1973 collector H. Zibrowius; material in mnhn, usnm)

t-503: 10.8.1967, 44°00.7'N, 7°06.9'W, 490m, SW Bay of Biscay: Stenohelia maderensis

u-807: 18.8.1968, 44°11'N, 8°40,2'W, 450-500m, NW Spain: Stylaster ibericus

x-340: 16.10.1971, 44°07'N, 4°29.8'W, 860-910m, S Bay of Biscay, Le Danois Bank: Stenohelia maderensis

x-341: 16.10.1971, 44°07.3'N, 4°30.7'W, 800-840m, S Bay of Biscay, Le Danois Bank: Stenohelia

maderensis

x-342: 16.10.1971, 44°07.5'N, 4°36.2'W, 700m, S Bay of Biscay, Le Danois Bank: Stenohelia maderensis

x-347: 16.10.1971, 44°07.3'N, 4°44'W, 640-910m, S Bay of Biscay, Le Danois Bank: Stenohelia maderensis

x-352: 17.10.1971. 44°06.5'N, 4°45.2'W, 545-580m, S Bay of Biscay, Le Danois Bank: Stenohelia

maderensis

x-353: 17.10.1971, 44°06.8'N, 4°45.1'W, 635-655m, S Bay of Biscay, Le Danois Bank: Stenohelia

maderensis, Pedicularia

x-362: 17.10.1971, 44°06.5'N, 4°50.9'W, 585-600m, S Bay of Biscay, Le Danois Bank: Stenohelia

maderensis

x-363: 17.10.1971, 44°06'N, 4°53.2'W, 545-630m. S Bay of Biscay, Le Danois Bank: Stenohelia maderensis

y-428: 4.9.1972, 44°11.8'N, 8°40.6'W, 500m, NW Spain: Stylaster ibericus

y-430: 4.9.1972, 44°11.6'N, 8°40.6'W, 500m, NW Spain: Stylaster ibericus + Pedicularia

y-431: 4.9.1972, 44°12.6'N, 8°40.7'W, 540m, NW Spain: Stylaster ibericus

y-432: 4.9.1972, 44°12'N, 8°40.6'W, 515m, NW Spain: Stylaster ibericus + Pedicularia

y-434: 4.9.1972, 44°12'N, 8°40.8'W, 620m, NW Spain: Stylaster ibericus

y-436: 4.9.1972, 44°12.6'N, 8°40.8'W, 620m, NW Spain: base of tiny stylasterid, unidentified

y-438: 4.9.1972, 44°12'N, 8°40.3'W, 545m, NW Spain: Stylaster ibericus

y-440: 4.9.1972, 44°12.2'N, 8°40.2'W, 490m, NW Spain: Stylaster ibericus + trace of Pedicularia

z-407: 23.10.1973, 47°43.6'N, 8°07.5'W, 1085-1115m, Celtic Sea: Pliobothrus symmetricus

z-415: 24.10.1973, 48°07.2'N, 8°26.2'W, 380m, Celtic Sea: Pliobothrus symmetricus + trace of Pedicularia

z-430: 25.10.1973, 48°37.0'N, 9°52.2'W, 1080m, Celtic Sea: Stylaster erubescens britannicus + trace of

Pedicularia

z-431: 25.10.1973, 48°38.2'N, 9°47.3'W, 800m, Celtic Sea: Pliobothrus symmetricus

z-435: 26.10.1973, 48°39.7'N, 9°53.2'W, 1050m, Celtic Sea: Pliobothrus symmetricus + trace of

Pedicularia, Stylaster erubescens britannicus

“ Thor ” 1904: W Iceland (material not located)

stn ?: 1904, 65°50'N, 26°53'W, 392m: Stylaster norvegicus, Stylaster erubescens (groenlandicus ?) (both

fide Broch, 1914a)

“ Travailleur" 1882: S Bay of Biscay (material in mnhn)

drag.70: 28.8.1882, 43°59'N, 5°34'15"W, 1000m: Pliobothrus symmetricus

“ Triton" 1882: between Faroes and Hébrides (material in bmnh, usnm)

3: 8.8.1882, 60°39'30''N, 9°06'W, 159m, SW Faroes Bank: Stylaster norvegicus

“ Tydeman" 1977-1978, cruises cancap 2 and 3 (material in rmnh)

2.160: 10.9.1977, 27°36'N, 17°59'W, 550m, Canary Islands, S. Hierro: Pedicularia (stylasterid host

unknown)

3.099: 23.10.1978, 30°07'N, 15°52'W, 585m, S Selvagem Grande: Crypthelia sp.

“ Ve ma " 1961, cruise vl7 (material in usnm)

rd-29: 9.4.1961, 60°27'N, 48°31'W, 326-366m, SW Greenland: Stylaster erubescens groenlandicus

“ Walther Herwig" 1982, cruise 54 (coll. G. Behrmann; material in imfb)

538-24: 24.9.1982, 65°25.2'N, 30°13.7'W, 700-713m, Denmark Strait: Stylaster erubescens groenlandicus

Source : MNHN, Paris

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INDEX

135

INDEX

The index covers taxonomie categories. Bold characters indicate the présent generic placement of spccies, bold numerals

major référencés. An asterisk (*) indicates taxa other than Stylasteridae.

•acrothoracic cirriped 21.

affinis (Crypthelia) 17, 20, 106-111, 112, 114, 117, 121.

alaskanus (Stylaster) 79, 83.

Allopora 17, 23, 62, 69, 71, 72, 76, 79, 83, 92, 94, 99, 101, 103.

alloporoides <*Cryptaxis) 23.

allispina (Errina) 47.

amoena (Errina) 53, 54, 55, 56, 57.

antiqua (Dislichopora) 23, 25.

antiquus (Stylaster) 23.

*aplocophoran mollusc 21.

aspera (Errina, *Millepora) 17, 19, 22, 24, 46-53, 56, 60, 67.

Astya 23.

Astylus 23.

ailantica (Allopora, Stylaster) 62, 63.

allantica (Errina, Lepidopora) 18, 20, 21, 22, 46, 50, 58-62.

*Autolytus 21.

bithalamus (Stylaster) 69.

blatteus (Allopora, Stylaster) 17, 20, 62, 64, 71, 72-76, 77.

britannicus (Stylaster) 18, 22, 62, 65, 69, 83, 87, 92-%, 97,

103.

carinata (Lepidopora) 31.

Calyptopora 24.

*Caryophyllia 54, 56.

cochleata (Errina) 47, 56.

compressa (Allopora, *Dendracis, Stylaster) 23.

Congregopora 23.

Conopora 22, 23, 24, 88.

*Corallium 104.

crassus ( Astya, Astylus) 23.

*Cryptaxis 23.

Crypthelia 17, 18, 19, 20, 22, 24, 26, 106-121, 122.

*cyathus (*Caryophyllia) 54, 56.

dabneyi (Errina) 17, 20, 22, 31, 46, 50, 53-58.

*decurvata (*Pedicularia) 22.

*decussata (*Pedicularia) 22.

*defrancei (*Lophelia) 24.

*Dendracis 23.

densicaulis (Stylaster) 88.

*deshayesiana (*Pedicularia) 24.

diffusa ( Errina) 72.

dispergens (Pliobothrus) 23.

Distichopora 20, 23, 24, 25.

ebrunea (Errina, *Hornera, Lepidopora) 17, 18, 20, 22, 27-31,

34, 35, 36, 38.

echinatus (Pliobothrus) 42.

Errina 17, 18, 19, 21, 23, 24, 26, 27, 31. 46-62, 67, 72.

Errinopora 23.

erubescens (Stylaster) 18, 19, 20, 22, 62, 65, 69, 83, 87-98,

103, 122.

Eu-Errina 47.

*Eunice 21, 59, 62.

Eustylaster 79.

faxensis ( Sporadopora) 23.

flabelliformis (Stylaster) 54.

gemmascens (Eustylaster, *Madrepora, *Oculina, Stylaster)

17, 19, 62, 63, 65, 69, 79-84, 92, 94, 99, 103, 122.

gigantea (Crypthelia) 119, 121.

*giganteum (*Pachylasma) 52.

glabra (Errina, Lepidopora) 27, 36.

glossopoma (Crypthelia) 117.

gracilis (Errina) 50.

gracilis (Pliobothrus) 18, 20, 39, 42, 44-46.

granulata (Stylaster) 88.

gravieri (* H ornera, Pliobothrus) 17, 31, 38, 42.

groenlandicus (Stylaster) 18, 62, 65, 87, 89-92, 94.

*Harmothoe 21, 106.

hicksoni (Errina. Lepidopora) 17, 27, 29, 31.

* H orner a 17, 27, 28, 29, 31, 38, 42, 53, 54, 56.

ibericus (Stylaster) 18, 20, 22, 62, 65, 84-87, 91.

irregularis (Errina, Spinipora) 23.

Labiopora 23, 47.

laevis (Pliobothrus) 23.

Lepidopora 17, 18, 19, 20, 22, 24, 26, 27-38, 53, 54, 56, 58, 60.

Lepidotheca 20.

lobata (Labiopora) 23.

*Lophelia 21, 24, 69, 84.

maderensis (Allopora, Stenohelia, Stylaster) 17, 18, 19, 20, 22,

24, 26, 69, 83, 94, 99-104, 106, 122.

*Madrepora 21, 62, 79, 81, 83.

maroccanus (Stylaster) 18, 20, 62, 64, 76-79.

mascarina (Errina) 47, 48, 49, 50.

medioatlantica (Crypthelia) 18, 20, 108, 110, 112-114.

meteorensis (Stylaster) 18, 62, 65, 87. 96-98.

*Millepora 46, 48, 50, 52, 61, 67.

miniatus (Stylaster) 67, 69.

moseleyi (Crypthelia) 106, 107, 111.

multipora (*Dendracis) 23.

nasiformis (Congregopora) 23.

•nemertean 21.

*norvegica (*Eunice) 21, 59, 62.

norvegicus (Allopora, *Millepora, Stylaster) 17, 19, 48, 50,

62-69, 83, 84, 91, 94, 103, 122.

*oculata (* Madrepora) 21.

oculina (Allopora, Stylaster) 17, 63, 64, 69, 79, 83, 92, 94, 99,

103.

*Oculina 79.

•Oculinidae 17.

* Pachylasma 52.

pacifica (Allopora, Stylaster) 63, 69.

pauciseptata (Stenohelia) 101, 103.

•Pedicularia 21, 22, 24, 28, 30, 31, 39, 43, 48, 49, 52, 53, 55,

56, 59, 62, 64, 85, 87, 89, 96, 100, 104.

peircei (Crypthelia) 22.

*pertusa (* Lophelia) 21, 69, 84.

Pliobothrus 17, 18, 19, 20, 22, 23, 24, 26, 27, 31, 38-46, 55,

122.

*Polydora 21.

*polynoid polychaet 21.

priscus (Allopora, Stylaster) 23.

profunda (Stenohelia) 101, 103.

pudica (Crypthelia) 17, 106, 107, 110, 111, 117, 121.

*pycnogonid 21.

Source : MNHN, Paris

136

INDEX

pygmaea (*Dendracis) 23.

rosaceus (Allopora, Stylaster) 17, 20, 64, 69-72. 76.

roseus (“ Madrepora. Stylaster) 18, 20, 62, 87, 88, 89, 92, 94.

*rubrum (*Corallium) 104.

sanguineus (Stylaster) 17, 79, 81, 83.

*Scleractinia 19.

*sicula (*Pedicularia) 22, 48, 52, 53.

*siphonostomatoid copepod 21.

*Solenosmilia 21.

sp. A (Lepidopora) 18, 19, 27, 28, 31-36, 37.

sp. A ( Stenohelia ) 18, 20, 21. 104-106.

sp. B (Lepidopora) 18, 20, 27, 31, 32, 33, 36-38.

sp. (unidentified, Stylaster) 94.

sp. (unnamed. Pliobothrus) 42, 43.

*Spinipora 23, 48.

Sporadopora 23.

Stenohelia 17, 18, 19, 20, 21, 22, 24, 26, 69, 83, 94, 99-106,

Stylaster 17, 18, 19, 20, 22, 23, 24, 26, 48, 50, 54, 62-98, 99.

103, 122.

*sublaevigata (*Pedicularia) 22.

subviolaceus (Allopora, Stylaster) 17, 72, 76.

symmetricus (Pliobothrus) 17, 19, 20, 22, 31, 38-43, 44, 46,

122.

tenuiseptata (Crypthelia) 17, 18, 20, 111, 114, 117-121.

*thoracic cirriped 21.

tiliata (Stenohelia, Stylaster) 18, 99, 103.

tuberculata (*Dendracis) 23.

lubulatus (Pliobothrus) 18, 27, 31, 40, 42.

‘variabilis (*Solenosmilia) 21.

vascomarquesi (Crypthelia) 18, 19,20,111, 114-117, 121, 122.

verrillii (Stylaster) 69.

verrucosa (*Hornera, Lepidopora) 17, 31, 53, 54, 56.

verrucosa (Conopora) 88.

*virginea (*Madrepora, Stylaster) 79, 83.

Source : MNHN, Paris

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Stylasteridae are calcified and highly modified hydroids, occurring worldwide over a wide range of

depth. Some stylasterids resemble bryozoans and others colonial scleractinians, convergences that hâve

caused confusion in recent and ancient faunas, and may also hâve limited our knowledge of their geo-

logical record. The présent monograph covers the 20 species in 6 généra known from the northeastern

Atlantic, only one of which also occurs in the Mediterranean. Ail species are described in detail, with

abundant illustrations of the colony shape and of the minute structures essential for identification.

Helmut Zibrowius is a marine biologist of the Centre d’Océanologie de Marseille (CNRS) and is

specially interested in deep-water faunas and in stylasterid and scleractinian corals together with their

symbionts. Stephen D. Cairns is the curator of stony corals at the National Muséum of Natural History,

Smithsonian Institution, Washington D.C., his research activity also centred on stylasterids and deep-

water scleractinians. The authors hâve cooperated since 1975 and published many papers on corals,

worldwide.

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