The Echinoderm Fauna
of Ascension Island,
South Atlantic Ocean

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SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES e NUMBER 2

The Echinoderm Fauna
of Ascension Island,
South Atlantic Ocean

David L. Pawson

SMITHSONIAN INSTITUTION PRESS
City of Washington
1978

JN, 1831S) TE IR ANG IE

Pawson, David L. ‘The Echinoderm Fauna of Ascension Island, South Atlantic
Ocean. Smithsonian Contributions to the Marine Sciences, number 2, 31 pages,
Il figures, 1978.—Two recent intertidal collecting expeditions and existing
museum collections have added much to knowledge of the Ascension Island
echinoderm fauna. Twenty-five species are now known from Ascension; eight are
new records. One new species, Holothuria (Halodeima) manningi, and one new
subspecies, Echinometra lucunter polypora, are described. Diadema ascensionis
Mortensen is regarded as a subspecies of D. antillarwm Philippi, and Pseudo-
boletia atlantica H. L. Clark is regarded as a subspecies of P. maculata ‘Troschel.

The echinoderm fauna of Ascension Island includes 8 amphi-Atlantic species,
3 western Atlantic species, 4 eastern Atlantic species, 5 circumtropical species,
4 species shared only with St. Helena, and 1 endemic species. There are in addi-
tion three endemic subspecies. ‘Twelve species are shared with St. Helena, and
both islands are closely similar in terms of numbers and relationships of their
faunal components. Colonization of both islands by planktonic larval stages is
suggested. Dendrochirotid holothurians, which lack such larval stages, are- not
represented at either St. Helena or Ascension. The structure of the Ascension
fauna seems to have been determined by vagaries of ocean surface and sub-
surface currents. In contrast, Bermuda, which sits astride the Gulf Stream, has
a fauna that is entirely typical of the West Indian region to the south.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded
in the Institution’s annual report, Smithsonian Year. SERIES COVER DESIGN: Seascape along the
Atlantic coast.

Library of Congress Cataloging in Publication Data

Pawson, David Leo, 1938—

The Echinoderm fauna of Ascension Island, South Atlantic Ocean.
(Smithsonian contributions to the marine sciences ; 2

Bibliography: p.

1. Echinodermata—Ascension Island (Atlantic Ocean) I. Title. II. Series.
QL383.35.A8P38 593’.9 77-20941

Contents

Page

LTD TOCOCHOM 2.6 oo.cls Sig bolo Sete ohne ais eee eRe Oe ce aera ]
Previous Records of Echinoderms from Ascension ..................- 2
ChecklistofeNscensioni island Echinoderms =... 45.5. 2-2.002:-s5005--: 4
Composition of the Ascension Echinoderm Fauna ..............+.++: At
Nelationshipsyof the Ascension Echimoderm Fauna ...5....-.:.......- 4
HOWvael Solatecmlsw ASCENSION mn era ies. sole ce selelstensie, 2) 2 Se avcvaldin aida ote 6
IMateniall Ta<enriVEGl 0:6 oi 5 bicine ome ce Os Ce Orn DE eens Ee nS an 8
Class QS ROTMDIIN as oie ccs octet Cleo ee ON Ren Ie cece a ee cae ern a 9
SUlGlassmASER OID EA ais arya ee tatsercia/<eisinlicns\arsle Sune esis a 8 Side ietelatunnete le 9
NRG PASI TUL ONTTOVAS yar She soles ec aS EOE Es Se ere TO Perera 9
Ranh? AST RODCIDINIDYND: Aenean cd boat eer 6 Ae nore area one 9

LT ARVOSKEP MARDELEUS (ell) case oe noose noeneb acne a odeE aor y)

Opralar WATINWATEIIYN "ore o-0'o 8 a EE Oe CRO OLOIG Chee SIR TE ET ce ea ae 9
Harmilyal CHAE TASTERIDAEM pease ee lieiais sieisetey« ls sole sls «cies said wera o8's 9
Ghactasicrmoncepesp (INCIZUS)) meetin eames te els 9
anni yal OPE IDUASTER ID AR. © qe icra) eusle ois 4 0 sare) eie ciate cudversieisie abs bias a ursle 10
Opiiidrastcrm cus demons Graver rereatse oA tr atria toes soo 10
ILAQELHO: BONNIE BORON CHEF oe Sage COO AS eet OTRO 8 OO 10
SuDelissm OPMIOROIDEAS cess 1 siete celle = els eieraisieis oe ce wdes oe on wenesslo sib 11
Oi dere ORTON Meer ee ey eee ce aes a nal wis mesos 1]
arly OREWA CIID AK Serie toe oy ih ren ek Ls eas Bate 1]
Opiniacisesauvenyi«(Meullersxand dsroschel)) 55... 2.225 +1 11
Ophiachissiymanimbjungman) ae eee). seen econ ese: 1]
Arai Va AMPHTURIDAE NS one esas can ste ya an bene nes cee eene es 11

Alii DOCOURD: CONGO IYPWEIN co cconanaoooaodagceuesnnee (peenoor 1]
Opliosinma Coinoimme (UymeI) | oa. deosoesaosaseneoguoe so05c 1]

Hani ilyeOPHIOMARICIDAE) one yc se ao sas ale cee sob ye oe eee 12
Ophiothrix (Ophiothrix) roseocaerulans Grube ...........--++ 12

Cllags TECHTINGRDBA, 3 co Getee anee ee en so ee See Ee arene 12
Order CIDARO I AM ey ena nas ie. vats i as SAR a ea ea nee s IZ
Rannlhy CHOAIVNG 3 Manse cane Ode tees oeis aa a i are 12
Enicidanismclavata@uNlontensenes 44.580 sess eee eee 12
NineLocid anisms pivosamMOrtensem) sn. hes 2 1 oe oo 16

Oreo res LADENUAT OUD AWM SNE ete cet, 0) eich si4 Sah, Siete bus Suersiesie @ ads) xe 17
arnt hye VAD FEN UANTID AE peter Stet RES oe.) icc, 5)-eleyeisieventiau me cus ies BR See 17
Diadema antillarum ascensionis Mortensen ............++-+++- 17

Oriel CEP ARB A CLOUD AM meee gona iicre seo ais 105) ay tueue uals: se) opauaetre or eiaie, sureieua.8 ¢ 17
Family ARBACIIDAE ..... n'a) betibeeeaty kid OC ROTC CID eee eee aeeene 17
AGOCMOPICUnUSs TOMGANUS AGASSIZ) enya eee esas 5 sos: 17
OrdemelEMNOPEEUROWU Age eer etic Aims at cele ochre tie esas Siete 17
anal yaahOXORNEWSIMDAENE = 5 sacrum + /iclet Mera a exe ee hie chs ele Slee eam 17
Rscudouolentaumacwlatavatiantica @lark.... =): a6. sss ceo 17

ili

iv SMITHSONIAN CONTRIBUTIONS TO MARINE SCIENCES

Page

Dripneustes, vewtinicosvise amare) rarer ee een eeee 20
Order “EGHINOIDA: if): )2aiusie situs soe Suse tes ces ace ee. alee ee 20
Family /ECHINOMETRIDAE) sec cose cae ane as tee eae 20
Echinometra lucunter polypora, new subspecies ............... 20
Echinometra Specimens yes ase ee ee ae eee 22
Summary of Echinometra Morphometric Data ................ 22
Order. ‘HOLECTYPOIDA,)..Jncyecaiash inet na. De aren eee 23
Family -ECHINONEIDAR :).iss).-5 scaures som ee a siete ie cies eee 23
Echinoneus cyclostomusmbeskeyo aie eee ee ee eee 23
Order (GLY PEASTEROMDA 4). /50)c/-0 ees ceae tic laiels ieee) Sep ee eee 23
Family JROTULIDAE - 3). 4c soreness ape eke eee Cee ee eee 23
Helhiophora orbiculus (Minnaeus)) 35 c eeeree 23
Order SPATANGOIDA 0.2, s)S.n0 since eo Suna meee ee ee ee De
Family BRISSIDAB 5: s2050. vee Wee, Seer eee eee 24
Brissus unicolor (Leske)) .:.75) oe en dsen. heer eee 24
Glass: FIOLOTHUROIDEA (4/60/05, .93.0/5 ocisi's one seiner os bree ae Oe eee 24
Order ASPIDOCHIROTIDAY (21/5 <5 cio c-estoce canis ere St Ree 24
Family: HOLOTHURIIDAE, (2.000.595 oe se cree see eee 7a:
Holothuria (falodeima) grisea Selenka. a: ae 4410 eee jus
Holothuria (Halodeima) manningi, new species .............. 26
Holothuria (Thymiosycia) arenicola Semper .................- 27
Holothuria (Platyperona) sanctori Delle Chiaje ............... 27
Family “STICHOPODIDAE #32. 3.0)0 oi): etc. sce ae eae ee Or eee 27
Tsostichopus badionotus,(Selenka), 00s) eee 2
Order \APODID Aw: 2.0s/.92%s a5 sa) s)sie ele svensts be oteleh tas OGRE a On ee ee ae 28
Family) SYNAPTIDAB) (000 2.0.e0Sas cote. oleesne ices See a Ree roe eee © 4%
Euaptalappas (Mullen 05... 3.15-:ca aoe eo eee 28

Literature -\Gited\ ose eee ee ie ee 29

The Echinoderm Fauna
of Ascension Island,
South Atlantic Ocean

David L. Pawson

Introduction

Ascension Island lies in the South Atlantic at
07°57’S, 14°22’W, approximately 150 kilometers
west of the centerline of the mid-Atlantic Ridge
(Wilson, 1963). It is an entirely volcanic island,
comprising the uppermost part of a cone which
rises from a depth of about 3000 meters below sea
level (Atkins et al., 1964). Daly (1925) has noted that
the only nonvolcanic material on the island are
“some small patches of beach material thrown up
by storm waves.”’ Much of this material is calcareous,
derived from calcareous algae and mollusk shells.
The island is approximately circular, with an area
of about 97 square kilometers (Figure 1). In the
absence of fossils and other reliable indicators, it is
difficult to determine the length of time that Ascen-
sion has stood above sea level. Estimates range from
approximately 1.5 million years to considerably less
than 1 million years (Sullivan, 1974; Daly, 1922;
J. D. Bell (in litt.) in Chace and Manning, 1972).

The nearest large land masses are a considerable
distance away; the coast of Brazil lies 2200 km to
the west, West Africa is 1300 km to the northeast
(Figure 2). To the south, 1100 km away, lies St.
Helena, another volcanic island, but considerably
older (20 million years; Wilson, 1963) than Ascen-
sion,

David L. Pawson, Department of Invertebrate Zoology, Na-
tional Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560.

Until very recently, no extensive collections of
invertebrates have been made at Ascension, and
our knowledge of the echinoderm fauna stems from
one or two small collections made during the late
1800s, and visits by Antarctic research vessels during
the first half of this century. Dr. R. B. Manning,
National Museum of Natural History, visited Ascen-
sion in 1971 and made extensive collections of in-
vertebrates. The interesting material obtained led
to the organization of a second Smithsonian expedi-
tion in 1976. This paper is based upon material
collected during those two expeditions: Asc. = col-
lection stations for 1976 expedition; RBM = col-
lection stations for 1971 expedition; USNM = cata-
log numbers using abbreviation for former United
States National Museum, collections of which are
in the National Museum of Natural History, Smith-
sonian Institution.

ACKNOWLEDGMENTS.—I am grateful to Dr. R. B.
Manning for making a fine collection of echino-
derms at Ascension in 1971. A second expedition to
the island in 1976 by Dr. Manning, Dr. M. L.
Jones, Dr. J. Rosewater and myself, of the National
Museum of Natural History, and Dr. A. J. Pro-
venzano, Jr., of the Old Dominion University, Nor-
folk, resulted in the amassing of an extensive series
of shallow water marine invertebrates, among them
numerous echinoderms. Mr. Ross Simons of the
Office of the Assistant Secretary for Science, Smith-
sonian Institution, was especially helpful in making
necessary arrangements with the U.S. Air Force and

no

ENGLISH BAY

FORT HAYES

COLLYER POINT.
7°s7's —

MCARTHUR POINT y
TURTLE SHELL BEACH ~ ZS
y

SOUTH WEST Bay

SHELLY BEACH

SMITHSONIAN CONTRIBUTIONS TO MARINE SCIENCES

NORTH EAST BAY

10° 22'w

Ficure 1.—Outline map of Ascension Island, showing collecting localities.

with the Administrator of Ascension Island. The
Administrator, Mr. Jeffrey C. Guy, the U.S. base
commanders Major Henry Spangler and Lieutenant
Colonel Thomas Morris, and the head of the NASA
tracking station, Mr. Jefferson Speck, were most
helpful to us during our stay at Ascension. Our
expenses were met by a grant from the Smithsonian’s
Fluid Research Fund; we are grateful to Mr. S.
Dillon Ripley, Secretary of the Smithsonian Institu-
tion, for his support.

For the loan of additional study material, I wish
to thank Miss Ailsa M. Clark of the British Museum
(Natural History), Dr. F. Jensenius Madsen, Uni-
versitetets Zoologiske Museum, Copenhagen, and
Dr. Lowell P. Thomas, University of Miami. The
photographs were made by Mr. V. Krantz, the maps
and graphs by Irene Jewett. Miss Tamara A. Vance
assisted with measurements of echinoids and _ col-
lation of data. I thank Drs. M. L. Jones and R. B.
Manning for reviewing the manuscript. Partial sup-
port for preparation of this paper was derived from
a grant made by the Smithsonian Research Founda-
tion (SRF 71500525).

PREVIOUS RECORDS OF ECHINODERMS FROM ASCENSION

The first record of an echinoderm from the island

was that of Cuninghame (1699:298), who made the
following observations:

One small warted Barbadoes sea egg. Echinus ovarious Bar-
bad. verrucis plurimis minoribus Mus. -Petiver 123.

The spines of these are purplish, especially the tips, the
largest I have yet seen, exceed not a crow-quill in thickness,
and are scarce an inch long; they end pointed, and are finely
striated if strictly viewed. The naked shell of this was some-
what more than six inches in circumference, and about 5
broadways and 514 lengthways.

Bell (1881:437) suggested that this “small warted
Barbadoes sea egg’’ was undoubtedly a specimen of
Echinometra subangularis Leske (= E. lucunter
(Linnaeus)); Cunninghame’s description fits that
species admirably. In the same paper Bell listed the
following species from Ascension Island:

Bell’s name Currently accepted name

Eucidaris clavata Mortensen

Diadema antillarum
ascensionis Mortensen

Tripneustes ventricosus
(Lamarck)

Echinometra lucunter
polypora Pawson

Cidaris metularia Lamarck
Diadema setosum Gray

Tripneustes angulosus Leske

Echinometra subangularis
Leske

Echinoneus cyclostomus Leske Same
Rotula dentata Leske Heliophora orbiculus
(Linnaeus)

Linckia diplax Miller and
Troschel

Linckia guildingi Gray

NUMBER 2 : 3

20 N

See
°

oie ae fe
CAPE VERDE 's.

GULF OF |GUINEA —

SAO TOME
\

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o
-
ATOL DAS ROCAS

wean ANNOBON nN

VaeeenaNee DE NORONHA

ASCENSION

10 Ss

ST. HELENA__

NE eae

T r 7 Saal 1

60 w 50 w aow 30 w 20 w 10 Ww o 10 =

FicurE 2.—Central Atlantic Ocean, showing principal islands.

dema ascensionis Mortensen (now D. antillarum
ascensionis) at Ascension, according to Mortensen
(1909).

As a result of a visit by the Discovery, Fisher
(1940) noted the presence of Ophidiaster guildingii
at Ascension, and Mortensen (1936) listed the fol-

During the early part of this century, several
ships paused briefly at Ascension, when they were
returning from expeditions in the southern oceans,
and made collections, usually by dredging. Koehler
(1908) reported the following species from Ascen-

sion, collected by the Scotia:

Koehler’s name

Moiraster magnificus (Bell)

Chaetaster longipes (Retzius)

Amphiura capensis Lyman

Tretocidaris spinosa
Mortensen

Cidaris minor, new species

?Coelopleurus floridanus
Agassiz

Echinometra subangularis
(Leske)

Pseudoboletia maculata
Troschel

The Deutsche Sudpolar-Expedition collected Dia-

Currently accepted name

Tethyaster magnificus (Bell)
Same
Same
Same

Eucidaris clavata Mortensen
Same

Echinometra lucunter
polypora Pawson .
Pseudoboletia maculata

atlantica Clark

lowing species:

Mortensen’s name

Ophiothrix roseocaerulans
Grube

Ophiactis savignyi (Miiller
and Troschel)

Ophiostigma abnorme
(Lyman)

Eucidaris tribuloides
(Lamarck)

Diadema antillarum var.
ascensionis Mortensen

Echinometra lucunter
(Linnaeus)

Currently accepted name

Same

Same

Same

Eucidaris clavata Mortensen

Diadema antillarum
ascensionis Mortensen

Echinometra lucunter
polypora Pawson

Thus, at the time of writing of this paper, 17
species of echinoderms were known from Ascension.
The following checklist lists all 25 known species
in systematic order; new records are marked with an
asterisk (*).

CHECKLIST OF ASCENSION IsLAND ECHINODERMS

CRINOIDEA
None
ASTEROIDEA
Family OPHIOTHRICIDAE
Tethyaster magnificus (Bell)
Family CHAETASTERIDAE
Chaetaster longipes (Retzius)
Family OPHIDIASTERIDAE
Ophidiaster guildingi Gray
Linckia guildingi Gray
OPHIUROIDEA
Family OPHIACTIDAE
Ophiactis savignyi (Miller and Troschel)
*Ophiactis lymani Ljungman
Family AMPHIURIDAE
?Amphiura capensis Lyman
Ophiostigma abnorme (Lyman)
Family OPHIOTHRICIDAE
Ophiothrix (Ophiothrix) roseocaerulans Grube
ECHINOIDEA
Family CIDARIDAE
Eucidaris clavata Mortensen
Tretocidaris spinosa Mortensen
Family DIADEMATIDAE
Diadema antillarum ascensionis Mortensen
Family ARBACIDAE
2Coelopleurus floridanus Agassiz
Family ‘TOXOPNEUSTIDAE
Pseudoboletia maculata atlantica H. L. Clark
Tripneustes ventricosus (Lamarck)
Family ECHINOMETRIDAE
Echinometra lucunter polypora Pawson
Family ECHINONEIDAE
Echinoneus cyclostomus Leske
Family ROTULIDAE
Heliophora orbiculus (Linnaeus)
Family BrissipAE
*Brissus unicolor (Leske)
HOLOTHUROIDEA
Family HOLOTHURIIDAE
*Holothuria (Halodeima) grisea Selenka
*Holothuria (Halodeima) manningi, new species
*Holothuria (Thymiosycia) arenicola Semper
*Holothuria (Platyperona) sanctori Delle Chiaje
Family STICHOPODIDAE
*Isostichopus badionotus (Selenka)
Family SYNAPTIDAE
*Euapta lappa (Miller)

The record of Amphiura capensis Lyman is doubt-

SMITHSONIAN CONTRIBUTIONS TO MARINE SCIENCES

ful (see p. 11). The single juvenile specimen of
Coelopeurus recorded from Ascension was identi-
fied by Koehler (1908) as C. floridanus Agassiz; this
identification must remain in doubt until adult
specimens are found.

COMPOSITION OF THE ASCENSION ECHINODERM FAUNA

The fauna as it is known today comprises 25
species (Table 1), of which one, Amphiura capensis,
is a doubtful record. The identity of Coelopleurus
floridanus has yet to be confirmed. Eight species are
reported from Ascension for the first time, but only
one of these, Holothuria manningi, is new.

There are some notable absentees from the fauna.
No members of the holothurian Order Dendro-
chirotida have yet been found, despite the fact that
these “cucumaria-type’”’ holothurians are common
on both sides of the Atlantic Ocean (see below for
discussion). No crinoids are known from Ascension.
It is also surprising that many species that are
amphi-Atlantic in distribution and can tolerate a
variety of habitats are apparently absent from
Ascension. These include such common shallow-
water forms as Linckia bouvieri Perrier, Axiognathus
squamatus (Delle Chiaje), Ophiocoma pumila
Litken, Ophioderma appressum (Say), and Ophio-
lepis paucispina (Say).

RELATIONSHIPS OF THE ASCENSION ECHINODERM
FAUNA

As can be seen from Table 1, the relationships of
the echinoderm fauna of Ascension are complex:
8 species (32%) are amphi-Atlantic; 1 species (4%)
is restricted to Ascension; 3 species (12%) are also
known from the western Atlantic only; 4 species
(16%) are also known from the eastern Atlantic
only; 5 species (20%) are circumtropical in distribu-
tion; 4 species (16%) are also known from St.
Helena only; 12 species (48%) are shared by Ascen-
sion and St. Helena.

These figures are similar to those for the mol-
lusks given by Rosewater (1975). The only notable
difference lies in the fact that 5 of the 89 Ascension
mollusk species (6%) occur at Ascension and St.
Helena only, while 4 of the 25 echinoderms (16%)
have this distribution pattern. There is some indi-
cation here that Ascension-St. Helena are more

NUMBER 2

TABLE 1.—Relationships of Ascension echinoderms (+ = present; O = present as species, but
not as subspecies listed; ? = doubtful record)

Ascension echinoderms

Tethyaster Magnificusssccccersscverevcscerece
Chaetaster longpipesss.ercccerceneecesoecece
Ophidiaster guildingi..r.crecccscesseesseoces
iinlaielcte), eabelilc binaeal GGG uo OdGOUCGDOOOUOOGOOOOG000
Ophiactis SAVIGNVl eecvscecccvecescnscceseece
Ophiactis Lymanisesercccccesscccccscccvescens
Amphiura Capensise.ceccrccrecccccccscssccces
Ophiostigma AbNOTME.cevcererccccccercsssoces
Ophiothrix (0.) roseocaerulans...sssseessees
Bucidaris Clavatas.cscccccccscccacscccssccccs
PRELOCLMALIS SPINOSA.e.reccsscccccvcarecsens
Diadema antillarum ascensionises.ccsscecccece
Coelopleurus floridanusesceccescsccsescecves
Pseudoboletia maculata atlantica.s..c.ccccosoee
TYLPNEUSTES VENTLICOSUSssceccecreccccreccece
Echinometra lucunter polyporasrcrersesescccae
Behinoneus cyclostomus.cecccccrecccerseccens
Heliophora orbiculus....cesecscccnsecscevcce
BIUSSUS (UME COMO. «cic. e.c sie cle ese 6/sc sicise vie e sles
Holothuria (Halodeima) griseasssssceseeesees
Holothuria (Halodeima) manningi....ceccseoons
Holothuria (Thymiosycia) arenicola.ssesseees
Holothuria (Platyperona) sanctoris.s.ssssceees
Esostichopus badilonotus.<.cssccccsessveceses

HUVEC AMPLUD DeNelelevelevelicielslelslelelelsisleiols(ellejeieleielsielsieleiere

WOalL spIslesocacdandccono0GGDUOGGDOG

“isolated” for the echinoderms than for the mol-
lusks, but little weight can be attached to such
slender evidence.

The relationships expressed in the above per-
centages imply that propagules have reached Ascen-
sion from the east and from the west. It seems
likely that colonization of Ascension was mediated
by surface and subsurface transport of planktonic
larval stages.

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Briggs (1974) has pointed out that Ascension lies
on the “northern limb of the South Atlantic gyre
where the water movement is relatively rapid and
the direction of flow almost due west.” He notes
that if this pattern of surface water flow were con-
stant, then one would expect that for the fishes
there would be a strong relationship with West
Africa. But this is not so, and Briggs suspects that
Ascension comes under the influence of the east-

ward-flowing Equatorial Countercurrent for at least
part of the year.

The equatorial undercurrent, flowing eastward at
velocities of around 0.8 to 2.9 knots (Metcalf et al.,
1962; Sturm and Voigt, 1966; Voigt, 1975), with a
core which may range from the surface to (usually)
around 75 meters, has been suggested by Chesher
(1966) as a likely route for transportation of larvae
of echinoids across the Atlantic from west to east,
thereby maintaining genetic continuity between
populations of amphi-Atlantic species. Chesher cal-
culated that larvae might make the trans-Atlantic
trip in a minimum of 43 days and a maximum of
70 days. Scheltema (1968, 1971) suggested that the
undercurrent would be a potent dispersal agent for
his so-called teleplanic molluscan larvae. ‘The true
extent of the equatorial undercurrent is not known.
Most authors estimate that it extends southwards to
a latitude of around 2°S. Possibly then the current
might not directly influence the fauna of Ascension,
which lies at latitude 7°57’S, some 600 km away.

Some alternative mechanisms for trans-Atlantic
transport of pelagic larval stages have been dis-
cussed by Scheltema (1968, 1971) for the North
Atlantic; similar patterns of current flow exist for
the South Atlantic. For successful transport of larvae
in the North or South Atlantic gyres, the larvae
must be eurythermal and long-lived. Six of the ten
mollusk species discussed by Scheltema in 1971 have
larval lives of at least three months; two others can
reach the settling state in less than two months;
these two, plus the remaining two species, can
probably delay metamorphosis until conditions are
suitable for settlement.

Regrettably, very little is known about the actual
larval life span of echinoderms. The excellent in-
vestigations of Mortensen (1921, 1931, 1938) and
others are mostly descriptive.

For Ascension Island species, the following in-
formation is available. Linckia guildingi: In the
related L. multifora the late brachiolaria stage is
reached after 27 days (Mortensen, 1938); in L.
laevigata metamorphosis can take place after 22
days (Yamaguchi, 1973). Ophiactis savignyi: Late
ophiopluteus is reached after 21 days (Mortensen,
1931). Eucidaris clavata: In the closely related E.
tribuloides metamorphosis takes place after about
25 days (McPherson, 1968). Tripneustes ventricosus:
Fully developed larva after 22 days (Mortensen,
1921). Echinometra lucunter: Metamorphosis takes

SMITHSONIAN CONTRIBUTIONS TO MARINE SCIENCES

places after 19 days (Mortensen, 1921). Holothuria
species: In H. impatiens metamorphosis can take
place after 21 days, and in H. difficilis after 14 days
(Mortensen, 1938).

Devaney (1973) extracted estimates of average
lengths of larval life for tropical echinoderms from
Thorson (1946) as follows:

Number of days Percentage of species
0-9
10-20 33
21-30 26
31-40 17
41-50 13

While the figures given above provide some use-
ful guidelines, it must always be borne in mind that
metamorphosis can be delayed for considerable
periods of time if proper food and/or proper sub-
strate are not available, or if physical factors such
as temperature and salinity are not optimal (Thor-
son, 1950).

The fact that Ascension appears to have received
colonists from the east and from the west leads to
the suggestion that the important surface and sub-
surface currents mentioned above are not exerting
a continuous influence on the island’s fauna. Lati-
tudinal shifts in currents accompanied by extensive
midocean eddies (Robinson, 1976) over the past one
million years have no doubt had important effects
upon composition of the Ascension fauna.

How “IsoLaTepD” Is ASCENSION?

Oceanic islands are useful natural laboratories for
investigation of effects of isolation upon their resi-
dent populations of animals and plants. MacArthur
and Wilson (1967) noted that new populations
arriving on an island might pass through three
overlapping evolutionary phases. During the first
phase, genetic divergence from the “mother” popu-
lation might take place (founder effect: see Mayr,
1963) because the colonizing population is small,
and contains fewer genes than the “mother popula-
tion” (the founder principle.) There is some experi-
mental evidence in support of the founder prin-
ciple (see MacArthur and Wilson, 1967) but some
critics have suggested that many colonizations are
made by successive waves of propagules, and that
large fractions of the genetic variation of the mother
population can be inserted into the founder popula-

NUMBER 2

tion, thus reducing genetic drift to a low level.
During the second phase, adjustments are made to
the new environment. The third phase comprises
speciation, secondary emigration, and radiation.

It is assumed that Ascension has been available
for colonization only within the past one million
years or so. The Ascension fauna offers some excel-
lent possibilities for exploration of the effects of
isolation. Many of the Ascension echinoderm species
are consistently different in a variety of ways from
their conspecifics elsewhere. The reader is referred
to the systematic section of this paper for details,
but some species are worth mentioning here in this
context.

Ophiactis lymani: Ventral arm plates with convex
distal edges in St. Helena and Ascension forms.
Specimens of this species from the eastern and
western Atlantic do not have convex distal edges
on ventral arm plates.

Diadema_antillarum ‘Tridentate
pedicellariae in Ascension and St. Helena popula-
tion (also Fernando de Noronha ?) with curved
valves. Valves essentially straight in eastern and
western Atlantic populations of this species (Dia-
dema antillarum antillarum).

Pseudoboletia maculata’ atlantica: ‘There are
usually five pore-pairs to the arc in Ascension and
St. Helena populations of this species. In Pseudo-
boletia maculata maculata there are typically four
pore-pairs.

Echinometra lucunter polypora: Ascension and
St. Helena populations have seven pore-pairs to the
arc, and the test is purplish adapically. In Echino-
metra lucunter lucunter there are typically six pore-
pairs, and the test is greenish or white adapically.

Holothuria (Halodeima) grisea: In comparison
with West Indian specimens of this species, Ascen-
sion specimens have smaller buttons with more
numerous larger perforations, and the ossicles in
the tube feet are a different shape.

Holothuria (Platyperona) sanctori: In Ascension
and St. Helena specimens of this species, the but-
tons have numerous small knobs. In contrast, speci-
mens from the eastern Atlantic and Mediterranean
have buttons with very few knobs or with none.

These striking and consistent differences from
presumed ‘“‘mother” populations prompt enquiry
as to whether they might be environmentally or
genetically induced. In the absence of experimental
evidence (rearing of specimens from different At-

ascensionis:

ih

lantic localities in uniform environments would be
most informative), it is almost impossible to answer
the question. If the differences are genetic, one
must presume that Ascension is genetically isolated
from the eastern and western Atlantic, but not from
St. Helena, and that recruitment of newly settling
juveniles is entirely internal.

There is some evidence to indicate that Ascension
is genetically isolated to a considerable degree.
Amphi-Atlantic populations of many species are
virtually identical morphologically, and a genetic
continuity across the Atlantic has been postulated
for the echinoids by Chesher (1966). But in the case
of Ascension, it is notable that (1) several amphi-
Atlantic species appear not to have been able to
colonize this island, and that (2) many taxa, in-
cluding Eucidaris clavata, Diadema_ antillarum
ascensionis, and Echinometra lucunter polypora
appear to have originated from populations else-
where in the Atlantic, but have not been “swamped”
by continuing recolonization, nor have populations
of these taxa apparently become established in other
parts of the Atlantic (Fernando de Noronha may be
the exception; this area of the Atlantic requires
further investigation).

It also seems evident that colonization by plank-
tonic larval stages is the only method by which the
Ascension echinoderms have reached the island. ‘The
absence of dendrochirotid holothurians is attribu-
table to the fact that these animals generally lack
planktotrophic larvae. Some dendrochirotids dis-
perse by rafting on floating weed. In the tropics, and
certainly in the case of Ascension, this method of
colonization is not common.

The Ascension-St. Helena fauna might be inter-
preted as being in the third evolutionary phase in
the sense of MacArthur and Wilson (1967). This is
the phase of “speciation, secondary emigration, and
radiation.” The process of speciation is obviously
slow, but establishment of subspecies, especially in
the echinoids, is proceeding rather rapidly.

One might contrast the fauna of Ascension with
that of Bermuda, another isolated island, which
lies 1075 km away from the nearest land-mass. ‘The
known echinoderm fauna of Bermuda now com-
prises approximately 50 species (Pawson, in Ms);
none of these are endemic to Bermuda, and all are
common in the West Indian region. Only one
species, Lytechinus variegatus (Lamarck), shows
slight but consistent differences from its West

Indian conspecifics. Bermuda is a relatively much
older island than Ascension (36 million years), and
thus might be expected to have developed some
endemic species. The single overwhelming factor
influencing the Bermuda fauna, however, is the
Gulf Stream; this gigantic river must be rapidly
transporting endless numbers of larval and early
juvenile stages of echinoderms from the West Indies
to Bermuda. No equivalent physical factor exists
for Ascension.

MATERIAL EXAMINED

This report is based upon material from several
sources. In 1970, Storrs Olson, National Museum
of Natural History, took time from his search for
fossil birds to make some shore collections; these
included species of echinoderms. Stimulated by Dr.
Olson’s discoveries, R. B. Manning visited the
island in May 1971, and made general collections
of shallow-water marine invertebrates; these in-
cluded an excellent series of echinoderms. Further
material was obtained during a second visit in July
1976, to Ascension by a party comprising R. B.
Manning, M. L. Jones, J. Rosewater, A. J. Pro-
venzano, Jr., and me. A specimen of Ophidiaster
guildingi collected at Ascension by Mr. A. Love-
ridge, together with ophiuroids and asteroids col-
lected by the U.S, Eclipse Expedition to West Africa
in 1890 are described in this report. Some echino-
derms collected a few years ago by Mrs. K. M.
Hutchfield, a former resident of Ascension, and
deposited in the British Museum (Natural History)
were kindly sent to me for study by Miss Ailsa M.
Clark. The small collection in the Ascension His-
torical Society Museum at Fort Hayes was examined
during July 1976.

In the station list below only the 1971 Manning
expedition and the 1976 expedition stations are
listed. Miscellaneous records are given under the
relevant species accounts. The number of specimens
is indicated after the author of the species.

R. B. MANNING’s 1971 EXPEDITION TO ASCENSION

RBM 3: 18 May 1971, Turtle Shell Beach, beach and rocky
point (MacArthur Point) at northern edge of South West
Bay, coarse sand beach and rocky intertidal, 1900-2030, RBM
and D. Rogers

Holothuria grisea Selenka, 1
RBM 5: 19 May 1971, North East Bay Beach and tidepools

SMITHSONIAN CONTRIBUTIONS TO MARINE SCIENCES

on rocky point west of main beach, temperature 27°C, salinity
38%o0, 0900-1100, RBM, D. Rogers, P. Kashulines
Ophidiaster guildingi Gray, 1
Ophiothrix (Ophiothrix) roseocaerulans Grube, 1
Eucidaris clavata Mortensen, 1 (found dry on beach,
not represented in collections)
Echinoneus cyclostomus Leske, 2
RBM 9: 20 May 1971, Turtle Shell Beach (MacArthur Point),
northern edge of South West Bay, tidepool with sand bottom
in lava flow area, 0800-0930, RBM and P. Kashulines
Isostichopus badionotus (Selenka), 1
RBM 10: 20 May 1971, rocky point at northern edge of
English Bay, intertidal pools and subtidal rocky shore, some
coarse sand bottom, temperature 27°C, salinity 39%o0, 1430-
1900, RBM and P. Kashulines
Eucidaris clavata Mortensen, 1
Holothuria (Halodeima) grisea Selenka, 1
RBM 12: 21 May 1971, Turtle Shell Beach, northern edge
of South West Bay (MacArthur Point), tidepool with sand
bottom in lava flow area, 1200-1630
Ophiothrix (Ophiothrix) roseocaerulans Grube, 7
Echinometra lucunter polypora Pawson, 1
RBM 13: 22 May 1971, rocky point off Fort Hayes, George-
town, algal mats in intertidal, 0800-1100, RBM and D.
Rogers
Holothuria (Thymiosycia) arenicola Semper, 1
RBM 14: 22 May 1971, rocky flat near Collyer Point, rock
surface at low tide with dense algal mat, area locally known
as Cable and Wireless Beach, 1100-1130, RBM and D. Rogers
Echinometra lucunter polypora Pawson, 1
RBM 15: 22 May 1971, same locality as sta 10, but second
tidepool from shore as well as more isolated tidepool, 1330-
1600, RBM, D. Rogers, K. Double
Ophiothrix (Ophiothrix) roseocaerulans Grube, 1
Eucidaris clavata Mortensen, 1
Echinoneus cyclostomus Leske, 1
Holothuria (Platyperona) sanctori Delle Chiaje, 2
RBM 16: 22 May 1971, Shelly Beach, flat exposed at low tide
on open beach proper, 1830-2330, RBM and D. Rogers
Eucidaris clavaia Mortensen, 1
Diadema antillarum ascensionis Mortensen, 2
RBM 18: 23 May 1971, Shelly Beach, tidepools in flat exposed
at low tide on open beach, temperature 27°C, salinity 40%o,
1000-1200, RBM, D. Rogers, K. Double
Ophiothrix (Ophiothrix) roseocaerulans Grube, 8
Schinometra lucunter polypora Pawson, 2
RBM 20: 23 May 1971, same as sta 14, 1900-2030, RBM
Eucidaris clavata Mortensen, 1
Echinometra lucunter polypora Pawson, 2
RBM 21: 24 May 1971, rocky point at northern edge of Eng-
lish Bay, tidepools on rocky flat, 0800-1200
Ophiothrix (Ophiothrix) roseocaerulans Grube, 4
Ophiactis savignyi Miiller and Troschel, 2
Isostichopus badionotus (Selenka), 1
RBM 22: 24 May 1971, Turtle Shell Beach, northern edge of
South West Bay (MacArthur Point), tidepool about 20 feet
in diameter with clear, soft sand bottom, 1300-1700
Ophiothrix (Ophiothrix) roseocaerulans Grube, 1
Eucidaris clavata Mortensen, 1

NUMBER 2

Diadema antillarum ascensionis Mortensen, 1

Echinometra lucunter polypora Pawson, 3

Holothuria (Thymiosycia) arenicola Semper, 2
RBM 23: 25 May 1971, same as sta 22, 0800-1030, RBM, K.
Double

Eucidaris clavata Mortensen, 1

Diadema antillarum ascensionis Mortensen, 5
RBM 24: 25 May 1971, same as sta 14, 1030-1230, RBM and
K. Double

Diadema antillarum ascensionis Mortensen, 7

Echinometra lucunter polypora Pawson, 17
RBM 24: 25 May 1971, same as sta 14, temperature high pool
42°C, shore pool 38°C, salinity high pool 32%o0, shore pool
28%0, 1300-1530

Ophiothrix (Ophiothrix) roseocaerulans Grube, 1

TueE 1976 EXPEDITION TO ASCENSION

Asc. 1-76: 11 Jul 1976, MacArthur Point, temperature 25°C,
salinity 340
1B, associated with rocks
Ophiothrix (Ophiothrix) roseocaerulans Grube, 12
Ophiactis savignyi (Miller and Troschel), 2
Eucidaris clavata Mortensen, 7
Echinometra lucunter polypora Pawson, 27
Holothuria (Halodeima) grisea Selenka, 8
1C, in clumps of coralline algae
Ophiothrix (Ophiothrix) roseocaerulans Grube, 12
Ophiactis savignyi (Miller and Troschel), 1
Echinometra lucunter polypora Pawson, 3
Asc. 3-76: 12 Jul 1976, English Bay, temperature 28°C (inner
pool), 25°C (outer pool), salinity 34%o
3A, sand, inner tidepool
Isostichopus badionotus (Selenka), 1
3G, associated with rocks beyond outer pool
Ophiothrix (Ophiothrix) roseocaerulans Grube, 10
Ophiactis savignyi (Miller and Troschel), 2
Eucidaris clavata Mortensen, 1
8D, associated with rocks, outer pool
Eucidaris clavata Mortensen, 13
Holothuria (Halodeima) grisea Selenka, 6
Holothuria (Platyperona) sanctori Delle Chiaje, 3
3G, collected by snorkeling in 5-10 m beyond outer tide
pool, calcareous sand and rock bottom
Holothuria (Halodeima) manningi Pawson, 3
Asc. 5-76: 13 Jul 1976, Shelly Beach, temperature 31°C, salin-
ity 34%o0
5A, isolated tide pools in back of open shore
Ophiothrix (Ophiothrix) roseocaerulans Grube, 2
Ophiactis savignyi (Miller and Troschel), 6
Holothuria (Halodeima) grisea Selenka, 1
Holothuria (Thymiosycia) arenicola Semper, 2
Asc. 6-76: 14 Jul 1976, Collyer Point, temperature 26°C,
salinity 34%
6A, isolated tide pools :
Ophiothrix (Ophiothrix) roseocaerulans Grube, 48
Ophiactis lymani Ljungman, 1
Echinometra lucunter polypora Pawson, 2
Echinoneus cyclostomus Leske, 1

6B, associated with exposed rocks
Ophiothrix (Ophiothrix) roseocaerulans Grube, 3
Eucidaris clavata Mortensen, 4
Diadema antillarum ascensionis Mortensen, 4
Asc. 7-76: 14 July 1976, off Collyer Point, diving, collected by
Mr. K. Jourdan
Ophidiaster guildingi Gray, 1
Ophiothrix (Ophiothrix) roseocaerulans Grube, 2
Echinometra lucunter polypora Pawson, 1
Asc. 8-76: 15 Jul 1976. MacArthur Point, isolated tide pool
Ophiothrix (Ophiothrix) roseocaerulans Grube, 3
Asc. 9-76: 16 Jul 1976, Northeast Bay, temperature 25°C,
salinity 34960
9G, isolated tide pool
Ophiothrix (Ophiothrix) roseocaerulans Grube, 7
Ophiactis savignyi (Miller and Troschel), 1
9D, associated with rocks in the intertidal
Holothuria (Platyperona) sanctori Delle Chiaje, 1

Class STELLEROIDEA
Subclass ASTEROIDEA
Order PAXILLOSIDA
Family ASTROPECTINIDAE

Tethyaster magnificus (Bell)

Archaster magnificus Bell, 1882:440.

Moiraster magnificus—Koehler, 1908:630, pl. 12: figs. 107—
110.—Mortensen, 1933a:422, pl. 21: figs. 1-2, pl. 22: fig. 1.
Tethyaster magnificus—A. M. Clark and A. H. Clark, 1954:

16, pls. 9, 10: figs. 2e-g.

MATERIAL EXAMINED.—None.

DisTRIBUTION.—This species is known only from
St. Helena (40 meters) and Ascension Island (72
meters).

Order VALVATIDA
Family CHAETASTERIDAE

Chaetaster longipes (Retzius)

Chaetaster longipes——Koehler, 1908:632.—Mortensen, 1933a:
431.

MATERIAL EXAMINED.—USNM_ 17473, Ascension Island, 77
m, collected by William Harvey Brown, 28 Mar 1890, U.S.
Eclipse Expedition to West Africa, 4 specimens.

10

RemMarks.—The four specimens are typical of the
species in all respects. Largest specimen has R =
52 mm.

DistRIBUTION.—Madsen (1950) gives the distri-
bution of this species as western Mediterranean,
European Atlantic coast as far north as Bretagne,
off West Africa, Morocco and Cape Palmas, St.
Helena, the Cape Verdes, the Canaries, and the
Azores, in 30-1140 meters.

HaBiTAT AT ASCENSION.—Not known.

Family OPHIDIASTERIDAE

Ophidiaster guildingi Gray
FIGuRE 3
Ophidiaster guildingii—Clark, 1921a:79.—Fisher, 1940:269.

MATERIAL EXAMINED.—RBM _ 5, 1 specimen; Asc. 1B-76, 6
specimens; Asc. 3D-76, 1 specimen; Asc. 7-76, 1 specimen.
Ascension Island, collected A. Loveridge, 31 Aug 1963, 1
specimen.

RemMarks.—The smallest specimen has R = 10
mm (station RBM 5); the largest has R = 52 mm.
The ratio R/r = approximately 1:8. When alive,
all specimens were mottled light and dark orange-
brown, and none were unicolor red. The adambu-

Ficure 3.—Left, Ophidiaster guildingi Gray from Ascension.
Right, O. ophidianus (Lamarck) from the Azores. Note con-
spicuous difference in shape of adambulacral spines.

SMITHSONIAN CONTRIBUTIONS TO MARINE SCIENCES

lacral spines (see Figure 3) conform exactly to the
description of H. L. Clark (1921a), and, thus, in all
respects the Ascension specimens concur with Clark’s
concept of this species. Fisher (1940) was of the
same opinion concerning a specimen collected at
Ascension by the Discovery. Mortensen (1933a) re-
corded O. ophidianus (Lamarck) from St. Helena,
noting that his specimens were either uniformly red
or had mottled coloration; he doubted that the two
species could be separated on the basis of color
alone. Chapman (1955) records O. ophidianus from
the Azores without comment, and this species also
occurs at the Cape Verde Islands (see A. M. Clark,
1955). The question of the validity of the two
species and their limits of variation can be settled
only by a thorough study of several scattered popu-
lations.

DisTRIBUTION.—Common throughout the West
Indies (Clark, 1921a), and recorded from Ascension
by Fisher (1940). O. ophidianus is known from the
Mediterranean, West Africa, and the islands men-
tioned above.

Hasitat at AsceNs1on.—On underside of rocks
or exposed in intertidal zone. Also present sub-
tidally (Asc. 7-76).

Linckia guildingi Gray

Linckia diplax—Bell, 1881:437.
Linckia guildingii—Clark, 1921a:67.—Madsen, 1950:216.

MATERIAL EXAMINED.—British Museum (Natural History)
Catalogue No. 1972.1.21.3, collected Mrs. K. Hutchfield, 1
specimen; Catalogue No. 81.10.27.15, collected T. Conry, 1
specimen.

Remarks.—Both specimens undoubtedly repre-
sent this widespread species. One specimen (Cata-
logue No. 81.10.27.15) is that which was identified
by Bell (1881) as Linckia diplax.

DistripuTIoN.—As Clark (1921) points out, this
seems to be “a truly tropicopolitan species of sea-
star, absent only from the western coast of America.”
It has not been reported from St. Helena; according
to Mortensen (1933a) another species which he
named Linckia formosa occurs there. L. formosa is
apparently most closely related to the amphi-
Atlantic species, L. bouviert Perrier.

Hasitat at Ascension.—Not known. Probably
same as for preceding species.

NUMBER 2

Subclass OPHIUROIDEA
Order OPHIURIDA
Family OPHIACTIDAE

Ophiactis savignyit (Miller and Troschel)

Ophiactis savignyi—Mortensen, 1933a:442; 1936:264.—Clark,
1955:36.—Madsen, 1970:207, fig. 33.

MATERIAL EXAMINED.—RBM 21, 4 specimens; Asc. 1B-76,
2 specimens; Asc. 1C-76, 1 specimen; Asc. 3C-76, 2 specimens;
Asc. 5A-76, 6 specimens; Asc. 9C-76, 1 specimen.

REMARKS.—AI] specimens have six arms and all
possess the very large radial shields that render this
species readily recognizable.

DistripuT1on.—According to Madsen 1970) this
species has a circumtropical distribution in littoral-
sublittoral depths. It has been recorded from Ascen-
sion by Mortensen (1936) and also from St. Helena
(Mortensen, 1933a).

Hapirat aT AsceNston.—Under rocks in tide
pools. Not uncommon in clumps of coralline algae.

Ophiactis lymani Ljungman

Ophiactis lymani—Mortensen, 1933a:442, fig.
1955:35, fig. 12.—Madsen, 1970:208, fig. 34.

15 —Clark,

MATERIAL EXAMINED.—Asc. 6A-76, 1 specimen.

Remarks.—This species has been well illustrated
by all of the authors cited in the synonymy above.
The single specimen from Ascension is typical in
most respects. Disc spines are virtually absent, there
being only one or two at the base of each arm.
Mortensen (1933a) pointed out that in St. Helena
specimens the ventral arm plates have convex distal
margins, whereas in populations from elsewhere,
the distal margins tend to be concave. In the pres-
ent specimens the distal margins are convex on all
ventral arm plates until approximately the 12th
arm joint, where the margins become concave, and
then they remain so throughout the length of the
arm. In alcohol the disc is light and dark gray
mottled, and the arms are light brown and dark
gray mottled.

DistRiBuTION.—Madsen (1970) described _ this

11

species as an amphi-Atlantic sublittoral form, known
from several localities off tropical West Africa, from
the Cape Verdes to northern Angola in 0-90 meters,
from the Virgin Islands in the West Indies, and
from St. Helena (10-110 meters). The present
record from Ascension is therefore not unexpected.

HapitTat AT ASCENSION.—Under rock in isolated
tide pool.

Family AMPHIURIDAE

Amphiura capensis Lyman

Amphiura capensis ——Koehler, 1908:634.
MATERIAL EXAMINED.—None.

Remarks.—Koehler recorded this species from
Ascension Island, and it has not been collected there
since. Mortensen (1933a) seriously doubted the valid-
ity of this record, and suggested that there had
been a mistake with the locality labels. Further
doubt is cast upon the validity of Koehler’s identi-
fication by Mortensen’s (1933b) restudy of Koehler’s
(1914) presumed Amphiura capensis from Angola
and Senegal; Mortensen found that these specimens
comprised two genera, neither of which was an
Amphiura.

Ophiostigma abnorme (Lyman)

Ophiostigma abnorme.—Mortensen, 1936:293.—Clark, 1955:
38.—Madsen, 1970:200, fig. 29.

MATERIAL EXAMINED.—USNM_ 17488 Ascension Island, 36—
54 m, collected by William Harvey Brown, 25 Mar 1890, U.S.
Eclipse Expedition to West Africa, 1 specimen; USNM 17489,
as above, 72 m, 5 specimens.

ReMARKS.—The six specimens are in fair condi-
tion; they are typical examples of this species. None
have numerous spines on the upper surface of the
disc; at best they are sparsely scattered.

DisTRIBUTION.—Madsen (1970) notes that this is
an amphi-Atlantic sublittoral species. It is also
known from the Cape Verdes. Mortensen (1936)
first recorded O. abnorme from Ascension.

Hasirat At AscENston.—Not known; has been
taken at 16-27 m (Mortensen 1936) and at 36-72 m.

12
Family OPHIOTHRICIDAE

Ophiothrix (Ophiothrix) roseocaerulans Grube

Ophiothrix roseocaerulans—Koehler, 1904:97.—Mortensen,
1933a:440, pl. 22: figs. 5-7; 1936:2262.
Ophiothrix (Ophiothrix) roseocaerulans—A. M. Clark, 1967:

647.

MATERIAL EXAMINED.—RBM_ 5, 1 specimen; RBM 12, 7
specimens; RBM 15, 1 specimen; RBM 18, 9 specimens;
RBM 21, 5 specimens; RBM 22, 1 specimen; RBM 25, 1
specimen. Asc. 1B-76, 12 specimens; Asc. 1C-76, 12 specimens;
Asc. 3C-76, 10 specimens; Asc. 5A-76, 2 specimens; Asc. 6A-76,
48 specimens; Asc. 6B-76, 3 specimens; Asc. 7-76, 2 specimens;
Asc. 8-76, 3 specimens; Asc. 9C-76, 7 specimens. South West
Bay (MacArthur Point), sandy bottom tide pool, collected by
Storrs Olson, 12 Jul 1970, 2 specimens; Ascension Island,
from dead Cymatium shell, collected K. Jourdan, Jul 1976, 1
specimen.

REMARKS.—These are typical specimens of this
beautiful species. The color is highly variable, as
Mortensen (1933a) had noted for his St. Helena
material. Most commonly shades of blue to purple
are represented, but light blue and even pink speci-
mens were collected at Ascension. A. M. Clark
(1967) in her revision of the family Ophiotrichidae
referred this species to the typical subgenus Ophi-
othrix (Ophiothrix).

DistRIBUTION.—The species is known to occur
only at St. Helena (see Mortensen, 1933a) and at
Ascension.

Hapitat aT AscEeNsion.—Almost ubiquitous in
the intertidal, encountered in clumps of coralline
algae, under rocks, and in crevices. It was common
at all stations sampled.

Class ECHINOIDEA
Order CIDAROIDA
Family CIDARIDAE

Eucidaris clavata Mortensen
Ficures 4-7

Cidaris metularia—Bell, 1881:436.

Cidaris tribuloides juv.—A. Agassiz, 1881:36, pl. 1: figs. 3, 5, 6.

Cidaris minor Koehler, 1908:638, pl. 16: figs. 136-138. [AI-
though C. minor Koehler, 1908, seems to be the oldest
available name for E. clavata, it is not available, as Mor-
tensen (1928) has pointed out, because it is preoccupied
by at least two earlier homonyms.]

SMITHSONIAN CONTRIBUTIONS TO MARINE SCIENCES

Eucidaris tribuloides——Clark, 1925:21 [in part] —Mortensen,
1936:213, pl. 1: figs. 13-15 [in part].

Eucidaris clavata Mortensen, 1928:408, pl. 42: figs. 1-4; pl.
67: figs. 10-11, pl. 73: fig. 2, pl. 86: fig. 15; 1932:169, pl. 4:
figs. 3-5; 1933a:464. =

Not Cidaris minor Koehler, 1914:217 [=Eucidaris tribuloides
africana].

ReMARKS.—The status of the species E. clavata
has been in doubt for some time. On the basis of
St. Helena material, Mortensen described his new
species EF. clavata in 1928, and later (1932, 1933a)
reinforced his opinion of the validity of the spe-
cies. Clark (1925) called attention to the clavate
spines of his St. Helena specimens, but also noted
that not all specimens had such spines. Later,
Mortensen (1936) described some small specimens
from Ascension as E. tribuloides and noted their
similarity to specimens from Annobon, West Africa
(now Pagalu, Equatorial Guinea), which he re-
garded as representing a “var. africana.” Compara-
tive measurements made upon small numbers of
specimens (2 from Annobon, 6 from Ascension, 4
from West Indies) of small size (largest specimen
18 mm h.d.) tended to support his contention. Ex-
amination of larger series of mature specimens for
the present paper show that Mortensen’s conclu-
sions were erroneous, that the Ascension Eucidaris
is identical to the St. Helena form, and quite dis-
tinct from the West African—West Indian species.

A complete description of £. clavata is not in-
cluded here; details can be found in Mortensen
(1928, 1932). However, it is necessary to discuss
those characters that are of apparent systematic
importance.

Relationship between Areole Width and Width
of Median Area of Interamb: Mortensen (1936)
measured the width of a primary areole at the
ambitus, and compared this with the width of the
median area of the interamb. He found that in
two West African species measured the areole width
exceeded slightly the width of the median area;
by contrast, in three West Indian specimens of
E. tribuloides the areole width was less than the
width of the median area. This supposed differ-
ence, together with some others of a minor nature,
led Mortensen to erect a new variety, E. tribuloides
africana, for the West African forms. I have made
additional measurements of some “‘typical’” West
Indian E. tribuloides, and these are presented
graphically herein (Figure 4). It is apparent that

NUMBER 2

o0.3—

RATIO OF WIDTH OF MEDIAN INTERAMB AREA
TO WIDTH OF AMBITAL AREOLE

30

13

a0 50 60

HORIZONTAL DIAMETER OF TEST (mm)

Ficure 4.—Comparison of Eucidaris tribuloides (Lamarck) with £. clavata Mortensen
(for details, see text).

tribuloides @

the relationship between dimensions of median
area and areole approaches unity (median area
width as percentage of areole width: mean 97%)
and that, in fact, Mortensen’s use of this character
to distinguish a new variety africana was not justi-
fiable. Chesher (1972) has plotted the same parame-
ters to distinguish E. tribuloides from E. thouarsit;
his graph shows the same results for the relation-
ship between areole and median interamb’ area in
E. tribuloides sensu stricto as does Figure 4 herein.
In the case of Ascension and St. Helena Eucidaris,
the situation is somewhat different (see Figure 4).

clavata (Ascension) @

clavata (St. Helena) ©

Here, the areole is almost twice as wide as the
median interamb area (interamb width as_per-
centage of areole width: mean 51.5%). It is be-
lieved that this character is important in distin-
guishing E. clavata from E. tribuloides.
Relationship between Diameter of Peristome and
Diameter of Apical System: As shown in Figure
5, for E. tribuloides from West Africa and from
the West Indies area, the peristome and apical sys-
tem have approximately the same diameter (apical
system diameter as percentage of peristome diame-
ter: mean 102%). The West African and West

14

RATIO OF APICAL SYSTEM DIAMETER TO PERISTOME DIAMETER

10 20

SMITHSONIAN CONTRIBUTIONS TO MARINE SCIENCES

ol

|
| |

a

BO

a # =
ae &
a
@ Beeees SEB B B

30 ao 50 60

HORIZONTAL DIAMETER OF TEST (wm)

FicurE 5.—Comparison of Eucidaris tribuloides (Lamarck) with E. clavata Mortensen
(for details, see text).

tribuloides (West Indies)
clavata (Ascension) @

Indian specimens are indistinguishable in respect
to this relationship. In contrast, in Ascension and
St. Helena Eucidaris, the peristome is considerably
larger than the apical system (apical system diame-
ter as percentage of peristome diameter: mean
77%). This character also is regarded as important
in distinguishing Ascension-St. Helena Eucidaris

(West Africa) []
(St. Helena) ©

from its congeners on each side of the Atlantic.
Shape of the Spines: Mortensen (1928, 1932)
regarded the shape of the spines as a feature of
paramount importance in distinguishing E. clavata
from E. tribuloides. He noted that while many of
his St. Helena specimens had the club-shaped spines
(such as those shown in Figure 6a—0 herein) others,

NUMBER 2

a

FicurE 6.—Eucidaris clavata Mortensen: a, St. Helena, 20-30 meters; b, St. Helena, 20-30
meters; c, St. Helena, shore; d, Ascension, shore; e, Ascension, shore; f, St. Helena, 20-30 meters
(Specimens a-c, f are in the Zoological Museum, Copenhagen.)

iB

16

particularly from the intertidal area, had more
slender, almost cylindrical, spines (Figures 6c-e, 7).
In Ascension specimens the spines are almost in-
variably cylindrical (Figures 6e, 7); on occasional
specimens a tendency towards clavate form is ex-
hibited (Figure 6d), but in no case were specimens
with truly club-shaped spines found. No deeper
water specimens have yet been recovered at Ascen-
sion (apart from Koehler’s (1908) Cidaris minor),
so the true range of spine shape for the Ascension
Eucidaris is not yet known. Examination of a great
variety of specimens of western Atlantic FE. tribu-
loides from a variety of habitats in the collections
of the National Museum of National History re-
vealed not one specimen with a tendency to form
club-shaped spines similar to those from St. Helena.

It is believed that, while presence of club-shaped
spines is a useful indicator of the identity of this
species, this character is highly variable in its
expression, and, consequently, is not completely
reliable.

Affinities of E. clavata: E. clavata is readily dis-
tinguished from E. tribuloides on the basis of rela-
tionships between dimensions of peristome and
periproct, relationships between width of ambital
areoles and width of median interamb areas, and,
to a lesser extent on the shape of the spines.
McPherson (1968) measured some specimens of
Eucidaris from Ascension to complement his study
of shape and growth of E. tribuloides, and he re-
garded the Ascension forms as conspecific with
E. tribuloides sensu stricto. He did not analyze his

46

ae

FicuRE 7.—Spines from specimens of Eucidaris clavata
Mortensen, Ascension Island.

SMITHSONIAN CONTRIBUTIONS TO MARINE SCIENCES

data, however, in a way which would have indicated
the obvious differences that exist, for such an analy-
sis was not relevant to his study.

E. clavata may be derived from E. tribuloides,
as has been suggested by Mortensen (1933a). The
eastern Pacific species E. thouarsti closely resembles
E. clavata in terms of the relationship between
diameter of peristome and apical system, and in
relationship between areole width and width of
median interamb area. The two species are dis-
tinctly separated on the basis of color alone. A form
from the Galapagos Islands, often referred to as
E. galapagensis (Doderlein) resembles E. clavata in
having club-shaped spines. The interesting parallel
was noted by Mortensen (1928), who commented in
some detail upon the status of E. galapagensis.

The complex of taxa, tr7buloides-clavata-thouarsii-
galapagensis, requires a thorough analysis, based
upon large series of specimens. The present arrange-
ment of the species is even now not completely
satisfactory. The intriguing question about forma-
tion of club-shaped spines at isolated oceanic
islands has yet to be investigated; a genetic basis is
suggested by the slender evidence available.

DisTRIBUTION.—E. clavata is known only from
Ascension and St. Helena, in depths of 0-60 meters.

HapitatT AT AscENsion.—All specimens were
found either lodged in rock crevices or under rocks,
always on hard substrates. Mostly smaller specimens
of up to 38 mm h.d. were found intertidally. Sub-
tidally, within snorkel diving range, considerably
larger specimens were found (50+ mm h.d.), usually _
wedged inextricably in deep crevices. Off St. Helena,
Mortensen (1932) found numerous specimens lodged
in holes and cavities of the sponge Chondrosia
plebeia Schmidt at depths of 20-30 meters.

Tretocidaris spinosa Mortensen

Tretocidaris spinosa Mortensen, 1903:17, 28, pl. 10: figs. 10,
11, 16.—Koehler, 1908:636, pl. 16: figs. 163, 164.—Morten-
sen, 1928:317, pls. 33: figs. 1-3, pl. 34: figs. 1-2, pl. 67: fig.
9, pl. 71: fig. 15, pl. 82: figs. 32-36; 1932:154, pl. 4: figs.
6-12, 15, pl. 13: fig. 3; 1933a:465.

. MATERIAL EXAMINED.—None.

DistRIBUTION.—Known from Ascension (72 me-
ters) and St. Helena (50-60) meters.

NUMBER 2

Order DIADEMATOIDA
Family DIADEMATIDAE

Diadema antillarum ascensionis Mortensen

Diadema setosum—Bell, 1881:436.

Diadema ascensionis Mortensen, 1909:55, pl. 7: fig. 10, pl. 16:
figs. 1, 4, 8, 16-17, 21-23; 1940:279, pl .48: fig. 2, pl. 54:
fig. 4, pl. 61: figs. 6-11, pl. 73: figs. 14-16—Tommasi, 1966:
11, figs, 3-5——Lima-Verde, 1969:10.—Brito, 1962:5; 1971:
264, fig. 1.

Diadema antillarum.—Clark, 1925:42 [in part].

Diadema antillarum var. ascensionis—Mortensen, 1933a:465;
1936:216.

MATERIAL EXAMINED.—RBM 16, 2 specimens; RBM 22, 1
specimen; RBM 23, 5 specimens; RBM 24, 7 specimens; Asc.
6B-76, 4 specimens. Turtle Shell Beach (MacArthur Point),
rocky tide pools, collected by Storrs Olson 18 Jun 1970, 1
specimen.

Remarks.—The status of the Ascension-St.
Helena populations of Diadema has been the sub-
ject of considerable discussion, mainly on the part
of Mortensen (1909, 1933a, 1936, 1940). His final
conclusion was that these populations should be re-
garded as a separate species from D. antillarum
Philippi. The main character upon which the
separation was based is the shape of the tridentate
pedicellariae, the blades of which are distinctly
curved in D. ascensionis, while they are more or
less straight in typical D. antillarwm. This char-
acter appears to be reasonably consistent, although
a survey of USNM material of D. antillarum revealed
that slightly curved tridentate pedicellariae are com-
mon. Mortensen (1940) also stated that in D. ascen-
sionis the median series of primary tubercles in the
interambs are arranged almost in a straight line,
whereas in D. antillensis the series remain distinct;
thus, in D. ascensionis there appear to be five series
of tubercles in each interamb. Again, this character
seems to be consistent for Ascension populations,
but numerous specimens of D. antillarum in USNM
collections show exactly the same type of tubercle
arrangement. Mortensen (1940) also notes that the
spines of St. Helena specimens are ‘“‘somewhat
coarser’ than those of D. antillarum; this is not a
reliable character, for the size of the spines can vary
considerably within populations.

In view of the broad distribution of D. antillarum
(see below), and the apparent restriction of D. ascen-
sionis to Atlantic islands or island groups, it would
seem wisest to regard D. ascensionis merely as a
subspecies of D. antillarum.

17

DisTRIBUTION.—D. a. antillarum occurs on both
sides of the Atlantic, in the west from Bermuda
south to Brazil, perhaps to 23°S (Tommasi, 1966),
and in the east from Cape Verde to the Gulf of
Guinea (Angola) on the African coast, also the
Azores, Canaries, and Madeira Islands. D. a. ascen-
sionis is known from Ascension (Mortensen, 1909)
and St. Helena (Mortensen, 1933a), Fernando de
Noronha (Clark, 1925; Mortensen, 1940), also Isla
Trindade (Brito, 1971) and Atol das Rocas (Lima-
Verde, 1969). Atol das Rocas is only 200 km from
the coast of Brazil, and possibly the Diadema there
is antillarum rather than ascensionis. Bathymetric
range is 0-360 meters.

HasitTaT AT ASCENSION.—Common in tide pools,
in rock crevices, and partially concealed under
rocks.

Order ARBACIOIDA
Family ARBACIIDAE

?Coelopleurus floridanus Agassiz

Coelopleurus floridanus.—Koehler, _1908:640.—Mortensen,
1935:612, pl. 68: figs. 5-7, p. 88: figs. 17-18, 31-34.

MATERIAL EXAMINED.—None.

REMARKS.—The juvenile specimen, 9 mm in
diameter, upon which Koehler (1908) based his
identification is the only specimen of this genus
so far known from Ascension. As Mortensen (1935)
points out, only additional adult material will settle
the question of the identity of the species. He also
notes that the Ascension form may be C. floridanus,
or the African species C. interruptus Déderlein, or
perhaps a new species.

DistRiBuTION.—Ascension Island, 72 meters
(Koehler, 1908); western Atlantic from West Indies
to Cape Hatteras, U.S.A., in 90-2380 meters (Mor-
tensen, 1935).

Order TEMNOPLEUROIDA
Family TOXOPNEUSTIDAE

Pseudoboletia maculata atlantica Clark

Pseudoboletia maculata—Koehler, 1908:641, pl. 15: figs. 139-
142, pl. 16: fig. 165.

18

Pseudoboletia atlantica—Clark, 1912:344; 1925:131.—Morten-
sen, 1933a:467; 1943:534, pl. 32: figs. 1-5, pl. 39: fig. 6, pl.
40: fig. 6, pl. 42: figs. 1-3, pl. 43: figs. 1-2, pl. 44: fig. 1,
pl. 55: figs. 1, 3, 19, 20.

MATERIAL EXAMINED.—No specimens in collections being
described here.

Remarks.—Koehler (1908) was the first to record
this species (as P. maculata) from Ascension, the
two specimens being collected at a depth of 40
fathoms off Point Pyramid. Clark (1912) regarded
the Ascension specimens as a new species, P. atlan-
tica, and Mortensen (1933a) reported the species
from St. Helena.

Study of the literature of this species and of an-
other Atlantic Ocean species, P. occidentalis Clark,
together with examination of specimens of both
(Table 1), has led to the conclusion that P. atlantica
should be regarded as subspecies of P. maculata
Troschel and that P. occidentalis is a junior syno-
nym of P. maculata. A brief analysis is given here
of the features used in the past to distinguish these
taxa.

Pseudoboletia MATERIAL EXAMINED.—P. occiden-
talis Clark: holotype (USNM E4531); 4 specimens
(USNM E12353) from off Venezuela, 83-186 meters,
diameters 63, 69, 78, 79 mm; 3 specimens (USNM
E16203, E16204) from off Florida north of Miami,
19-21 meters, diameters 60, 76, 85 mm. P. atlantica
Clark: 3 specimens (USNM E16096, diameter 71
mm; USNM E11732, diameter 82 mm; USNM
E5953, diameter 89 mm) from St. Helena.

Spicules in Gills: In the gills of both P. atlantica
and P. occidentalis spicules of the bihamate and C-
shaped type are numerous. The C-shaped spicules
appear to be. more numerous than the bihamate
type in P. atlantica, and the reverse might appear
to be true in P. occidentalis, but it was found that
the relative proportions of the types of spicules can
depend to a great extent upon the area of gill that
is being studied. The same is probably true for
P. maculata. It is concluded that this character is
unreliable. Mortensen (1943:537) referred to the
“remarkable differences from [the Indo-Pacific spe-
cies] indiana and maculata...” in the matter of the
spicules of P. atlantica, but apparently did not
regard a supposed difference as being important sys-
tematically.

Number of Plates on the Periproct: Clark (1921b:
116-117) noted that “In maculata the periproct is

SMITHSONIAN CONTRIBUTIONS TO MARINE SCIENCES

covered by about 30 plates,” and that in P. occi-
dentalis there are “about 20 plates.’’ In the four
additional specimens of P. occidentalis examined
there are more than 30 plates in the periproct, and
the same situation applies in specimens of P. at-
lantica, Clearly, no reliance can be placed upon
this character, as has already been pointed out by
Mortensen (1943).

Size and Shape of Primary Spines: Clark (1921b)
noted that the ambital primary spines of P. occi-
dentalis are “about 12 mm long, terete basally, and
becoming flattened only slightly near tip”; this sit-
uation he contrasted with that in P. maculata,
where the spines are “markedly flattened, with
bluntly chisel-shaped tips (though with a terminal
concavity), and are 14-16 mm long.” The. primary
spines of the additional specimens of P. occidentalis
examined are typically 15 mm or more in length,
often approaching 20 mm. They are terete basally,
and the degree of flattening near the tip is highly
variable. Ambital spines of P. atlantica are similiar,
but tend to be shorter, averaging approximately 14
mm in length.

First Appearance of Inner Primary Tubercle:
Clark (1921:117) notes that in P. maculata the
inner primary tubercle in the ambs appears first
on the seventh, eighth, or ninth plate from the
ocular plate in specimens 52-55 mm in diameter,
whereas in his specimen of P. occidentalis it occurs
first on the tenth-twelfth plate. In the additional
specimens of P. occidentalis, the tubercles first ap-
pear on the eighth to the 14th plate from the
ocular; the number of plates are correlated to some
extent with size of the specimen. In a specimen of
63 mm h.d., the first tubercle appears on plates
8-9 from the ocular, while in a specimen of 78 mm
h.d. it appears on plates 11-14. In P. atlantica the
tubercles appear on plates 8-10 from the ocular.
These figures confirm Clark’s (1921b) own sugges-
tion that this character is “inconstant and unreli-
able.”

Color of Test: The dark brownish or greenish
blotches which appear to characterize P. maculata
are also present on the holotype of P. occidentalis.
They are present and conspicuous in most specimens
of the additional material of P. occidentalis. Mor-
tensen (1943) notes that in P. atlantica the blotches
may be absent, or that there may be some darker
spots or bands in the median spaces aborally. It is

NUMBER 2

evident that in the Atlantic Pseudoboletia the test
coloration is somewhat variable.

Color of Spines: In all specimens of P. occidentalis
examined, the color of the spines has faded some-
what, but in most cases the spines are greenish
basally and pale fawn elsewhere, or almost totally
white. In the holotype and most non-type specimens,
spines located on the greenish blotches on the test
are mostly dark green, light only at the tips, a
feature already noted by Clark (1921b) for the
holotype. In P. atlantica, according to Mortensen
(1943) the spines may be whitish, with green or
brown basal areas, or they may be banded with dull
green or brown. Clark (1921b) noted that in P.
maculata the spines are “green at base and red-
purple or reddish at tip . . . quite different from
the pale colors of occidentalis.”

Pedicellariae: In his original description of P.
occidentalis, Clark (1921b) did not describe the
pedicellariae in detail, and Mortensen (1943) ex-
pressed regret that Clark had not illustrated the
tridentate pedicellariae he had described as “slender,
with valves 1.25 mm long.” I could find no such
pedicellariae on the holotype or on the additional
specimens of P. occidentalis. All of the tridentate
pedicellariae found were of the broad-bladed type,
none of them exceeding 1.1 mm in total length. In
all features these pedicellariae closely resembled
those of P. maculata, P. atlantica, and P. indiana.
It is impossible to determine from Mortensen’s
(1943) figures of tridentate pedicellariae from these
three species exactly how they differ from each
other, although he mentions (p. 533, 537) that they
are essentially the same in P. atlantica and P. indt-
ana, and that those of P. indiana and P. maculata
differ. I do not believe that these differences, what-
ever they might be, are significant.

The globiferous pedicellariae also present a con-
fusing picture. Mortensen (1943:531) notes that the
globiferous pedicellariae in P. indiana ‘‘are as in
P. maculata,” but two pages later (p. 533) he states
that the smaller globiferous pedicellariae “differ
rather conspicuously in the length of the blade, it
being distinctly longer than the basal part in
maculata, scarcely as long as the basal part in in-
diana.” His figures show a conspicuous difference in
the pedicellariae of these two species. The same
type of pedicellariae in P. atlantica are virtually
identical to those of P. maculata, and in P. occiden-

19

talis a condition resembling that in P. maculata was
found, although the blade appears to be relatively
shorter.

An extensive study of large series of pedicellariae
from many parts of the world might settle the
several problems that exist with regard to the
pedicellariae in this genus, but on the basis of the
admittedly scanty evidence now available, it would
seem that they do not afford the best characters
for distinction of species in Pseudoboletia.

Number of Pore-Pairs to the Pore-Arc: All species
in the genus Pseudoboletia have four pore-pairs in
each pore-arc, except P. atlantica, which has five.
In this respect, P. atlantica forms an interesting
parallel to Echinometra from Ascension (see p. 22).
The presence of five pore-pairs has been noted by
Mortensen (1943) in a large specimen of P. indiana
75 mm in diameter, and, conversely, P. atlantica
frequently has arcs of four pore-pairs. Mortensen
(1943) states:

The pore-pairs are in general arranged in arcs of 5.... But
this is by no means constantly so; some specimens have in
the main only 4 pore-pairs, or there may be here and there
a plate with only 4 pairs, most of the plates having 5 pore-
pairs. This has nothing to do with age; thus the specimen of
94 mm h.d.... has in the main 4-geminate plates, whereas
the specimens of 62 and 68 mm have regularly 5-geminate
plates.

In the present collection, all specimens of P. occt-
dentalis have four pore-pairs to the arc, and all
specimens of P. atlantica have five.

It is concluded that the only reasonably consistent
difference between P. atlantica and other species in
the genus is the presence of five pore-pairs in each
pore-arc. There appears to be no evidence for main-
taining P. occidentalis as a species distinct from
P. maculata; all of the distinguishing features enu-
merated by Clark fall within the range of variation
of P. maculata. Further, in view of the fact that
some other echinoid species at Ascension Island
have undergone some phenetic changes as a result
of isolation, environment, or a combination of the
influences of these factors, the presence of five pore-
pairs would not appear to be sufficient to warrant
maintenance of P. atlantica as a separate species,
distinct from P. maculata, and it is here recognized
as a subspecies of P. maculata. The status of
P. maculata in relation to P. indiana cannot be
determined here.

20

DIsTRIBUTION.—P. maculata atlantica is so far
known only from Ascension and St. Helena in
20-40 meters. P. maculata maculata is known from
Ceylon, the Philippines and the Macclesfield Bank,
and the Banda and Timor Sea in 20-70 meters
(Mortensen, 1943), also southern Japan (Utinomi,
1954); in the western Atlantic, P. m. maculata is
known from off Barbados (holotype of P. occiden-
talis), from off Venezuela in 83-186 meters, and
from off southeastern Florida in 19-21 meters.

Tripneustes ventricosus (Lamarck)

Tripneustes angulosus —Bell, 1881:437.
Tripneustes esculentus—Clark, 1925:124.
Tripneustes ventricosus—Mortensen, 1943:490.

MATERIAL EXAMINED.—None.

Remarks.—Bell (1881) first reported this species
(as T. angulosus) from Ascension and Clark (1925)
referred again to Bell’s specimen, confirming its
identity as T. esculentus (=T. ventricosus). No fur-
ther specimens of the species have been taken at
Ascension, in spite of assiduous intertidal and sub-
tidal collecting in what would seem to be suitable
habitats for the species.

Mortensen (1933a) was inclined to believe that
the record of this species from St. Helena (Cunning-
ham, 1910) was erroneous, for he found no further
material in his collecting around that island. In
fact, he suggested that the material of this species
which was described by Cunningham may have
been collected at Ascension.

DisTRIBUTION.—Common in the West Indies and
the west coast of Africa (Mortensen, 1943). The rec-
ords from Ascension and St. Helena must be re-
garded as questionable.

Order ECHINOIDA
Family ECHINOMETRIDAE

Echinometra lucunter polypora, new subspecies
Ficures 8-10

Echinometra acufera Mellis, 1875:219.

Echinometra subangularis—Bell, 1881:437W—Koehler, 1908:
640.—Cunningham, 1910:125.

Echinometra lucunter—Clark, 1925:143 [in part].—Morten-

SMITHSONIAN CONTRIBUTIONS TO MARINE SCIENCES

sen, 1933a:468; 1936:224 [in part]; 1943:357, pl. 41: figs. 1-5,
pl. 42: figs. 12-14, pl. 43: figs. 1-13, pl. 44: fig. 9, pl. 64:
figs. 17, 20-24 [in part].

MATERIAL EXAMINED.—Holotype: USNM E16206, RBM 24,
25 May 1971, Ascension Island, rocky flat near Collyer Point,
rock surface at low tide with dense algal mat, collected
by R. B. Manning and K. Double. Paratypes: USNM E16190,
RBM 24, 16 specimens (same data as holotype); USNM
E16186, RBM 12, 1 specimen; USNM EI16179 (4), USNM
E16191 (1), RBM 14, 5 specimens; USNM E16184, RBM 18,
2 specimens; USNM Ei16185, RBM 20, 2 specimens; USNM
E16187, RMB 22, 3 specimens; USNM E16180, Asc. 1B-76,
27 specimens; USNM E16181, Asc. 1C-76, 3 specimens; USNM
E16189, Asc. 6A-76, 2 specimens; USNM E16182, Asc. 7,
1 specimen; USNM E16183, Turtle Shell Beach (MacArthur
Point), rocky tide pools, collected Storrs Olson 18 Jun 1970,
3 specimens.

Dracnosis.—Like E. 1. lucunter, except that there
are predominantly 7 pore-pairs to the arc, when
greatest test diameter of approximately 32 mm is
reached. Upper surface of test with conspicuous
tinge of purple; no trace of green color adapically.

Remarks.—While I am reluctant to erect sub-
species in a group that is poorly known in so many
ways, there seem to be excellent grounds for regard-
ing the Ascension and St. Helena populations of
E. lucunter (Linnaeus) as distinct from the “‘typi-
cal” amphi-Atlantic populations. Mortensen (1933a,
1943) has already called attention to differences that
he noted in St. Helena specimens of this species,
and my examination of larger samples of speci-
mens from Ascension and elsewhere tend to confirm
Mortensen’s observation (for the St. Helena speci-
mens) that they are ‘about to develop into a sepa-
rate variety’ (1943:366). Mortensen’s observation
(1943:366) that the Ascension Echinometra are dif-
ferent from St. Helena specimens in some respects
are contradicted by my present findings, and I could
find no morphological differences between Ascen-
sion and St. Helena.

DIsTRIBUTION.—Ascension and St. Helena, inter-
tidal and shallow subtidal.

HasiTaT AT ASCENSION.—This is by far the most
common intertidal species at Ascension. Specimens
were found in great numbers in high splash pools,
in pools between tidemarks, and in rock faces at
and slightly below low tide level. While most were
occupying shallow burrows in the volcanic rock,
many were lodged in crevices or were exposed on
the rock surface (Figure 8). Commonly, specimens
were found in association with Diadema antillarum

NUMBER 2

FIGURE 8.—Two contrasting habitats for Echinometra lucunter polypora at Collyer Point,

Ascension Island: upper, specimens on surface of rock; lower, specimens occupying burrows.
(Photos: R. B. Manning.)

21

22

ascensionis. J. Rosewater (pers. com.) found small
clusters of the gastropod mollusk Hipponyx anti-
quatus (Linnaeus) in the bottoms of several of the
burrows after removal of Echinometra. This may
be merely a fortuitous association, for numerous
specimens of the gastropod were also found in other
habitats, not associated with Echinometra. Approxi-
mately 20 specimens of Echinometra were washed
in formaldehyde in search of commensals; none
were found.

Echinometra Specimens

Cotor oF TrEst.—In all specimens from Ascen-
sion, the upper part of the cleaned test has a dis-
tinct purple tinge; the test is white elsewhere. By
contrast, all specimens from the western Atlantic
examined have the test either more or less white,
or more often with a greenish tinge to the upper
part of the test. Color of specimens from Annobon,
off West Africa (now Pagalu, Equatorial Guinea) is
variable; some specimens have greenish tests, some
have a very pale purplish tinge, while others are
virtually white. According to Koehler (1914), the
West African specimens he examined are indistin-
guishable from the West Atlantic forms. Mortensen
(1943) noted that West African specimens are
“darker,” but apparently he was referring to color
of the spines.

Specimens from St. Helena are all purplish adapi-

Ficure 9.—Ambulacrum of Echinometra lucunter polypora,
new subspecies, showing arcs of 7 pore-pairs.

SMITHSONIAN CONTRIBUTIONS TO MARINE SCIENCES

cally (Mortensen, 1933a) as they are in USNM col-
lections.

Cotor oF SpiInEs.—Mortensen (1943) noted that
the spines “show all intermediates between wholly
black and wholly white. . . . It is very noteworthy
however that all the numerous specimens from
St. Helena, Ascension, and West Africa that I have
seen are black.’ The living specimens at Ascension
appeared quite black upon superficial examination.
Preserved and dried specimens have dark green
spines with purplish tips. The same coloration can
be found in specimens from Annobon, off West
Africa (USNM E12263).

NuMBER OF PoreE-Pairs.—Mortensen (1943) noted
that in West Indian specimens of E. lucunter the
number of pore-pairs is “more generally 6 (though
often 7) to each arc,” while in St. Helena specimens
there are “almost constantly 7 (or often 8... .” The
number of pore-pairs in Ascension specimens were
counted and results are given below, together with
data gathered from series of E. lucunter from
the eastern Atlantic (Annobon) and the western
Atlantic (St. Croix, Virgin Islands; Key West, Flor-
ida; and Curacao). The pore-pairs counted were
those immediately above the ambitus. See Figure 9.

Summary of Echinometra Morphometric Data

ASCENSION IsLAND.—Number of specimens exam-
ined, 56. Greatest length 9-64 mm, width 8-64 mm.
Average width as percentage of length 90%. In 12
specimens the product of length X width was less
than 900, and 10 (84%) of these specimens have
6 pore-pairs to the arc. In the 44 specimens where
length X width exceeded 900, 24 (approximately
55%) have 7 pore-pairs to the arc, 3 (7%) have pre-
dominantly 7 pore-pairs and 10 (23%) have 6 and
7 pore-pairs present in approximately equal propor-
tions. None of the latter specimens have 6 pore-pairs
to the arc throughout.

EASTERN ATLANTIC.—Number of specimens exam-
ined, 15. Greatest length 10-60 mm, width 9-50
mm. Average width as percentage of length 91%.
In I1 specimens the product of length < width ex-
ceeded 900, and in all of these (100%) the number
of pore-pairs to the arc is 6. No specimens have
arcs with 7 pore-pairs.

_ WESTERN ATLANTIC.—St. Croix Population: Num-
ber of specimens examined, 110. Greatest length
25-62 mm, width 23-58 mm. Average width as per-

NUMBER 2

centage of length 88%. In 74 specimens the product
of length X width exceeded 900, and in 66 (89%)
of these there are 6 pore-pairs to the arc. One speci-
men had predominantly but not exclusively 7 pore-
pairs, and in only 7 specimens (9%) arcs with 7
pore-pairs were present in equal numbers with arcs
with 6. No specimens have 7 pore-pairs to the arc
throughout.

Key West and Curagao Specimens: Number of
specimens examined, 30. Greatest length 19-62 mm,
width 17-54 mm. In 23 specimens the product of
length X width exceeded 900, and in 15 specimens
(65%) there are 6 pore-pairs to the arc. In 4 speci-
mens (17%) there are 7 pore-pairs, and in 4 speci-
mens (17%) 6 and 7 pore-pairs were present in
approximately equal proportions.

SumMMARY OF Data.—The data summarized above

eee MOSTLY 6 PORE-Pa~IRS

6=7 PORE-PAIRS

\\\ MOSTLY 7 PORE -PAIRS

100

TKN]
SN
<XX
RX

8

oS
TX
me

KD
So
=
x

ZS
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SS

oO

o,

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ZX
4M

%

"0%
en
XX
OO

60

\7
gh
XD
LRKS

KS
Kes he

Slo OF SAMPLE
<x
ORK
Oe
REX

CC
ese
0%

410

AA
05
LR
XX
$55

XRD
SRLS
LXXKXD
SS

OQ
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SS
5S

20

\7
x
xx
S&

RK

QO
Q

55

x

Rey

¥

Xx

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y

Ficure 10.—Numbers of pore-pairs to the arc in three popu-
lations of Echinometra lucunter (Linnaeus): a, Western At-
lantic (St. Croix); 6, Ascension; c, Eastern Atlantic (Anno-

bon). (For details, see text.)

23

and presented graphically in Figure 10 show that the
populations of E. lucunter at Ascension are con-
sistently different from those examined from the
eastern and western Atlantic in terms of number of
pore-pairs to the arc. After a length of approxi-
mately 32 mm is reached (at which point length X
width will approximate 900), the typical configura-
tion for Ascension specimens is to have 7 pore-pairs,
while eastern and western Atlantic forms typically
have 6.

The combination of characters here regarded as
taxonomically important (number of pore-pairs and
color of test) warrant the erection of a new sub-
species to accommodate the central Atlantic popula-
tions of the species.

Order HOLECTYPOIDA
Family ECHINONEIDAE
Echinoneus cyclostomus Leske

Echinoneus cyclostomus.—Bell, 1881:437——Mortensen, 1948:
75, fig. 60, pl. 1: figs. 14, 26; pl. 12: figs. 21-23.

MATERIAL EXAMINED.—RBM 5, 2 specimens; RBM 15, 1
specimen; Asc. 6A-76, 1 specimen.

Remarks.—The four specimens (total lengths 20,
26, 31, 41 mm) are typical of this well-known
species, and need no further comment here.

DisTRIBUTION.—A truly “tropicopolitan” species,
which ranges the Indo-Pacific and the western At-
lantic; it has not yet been recorded from West
Africa (Mortensen, 1948:79).

Hapsitat at Ascension.—Under rocks on hard or
calcareous sand substrate.

Order CLYPEASTEROIDA
Family ROTULIDAE

Heliophora orbiculus (Linnaeus)

Rotula dentata.—Bell, 1881:436.

Rotula orbiculus—Clark, 1925:175——Mortensen, 1948:459,
pl. 57: fig. 1, pl. 72: fig. 4; 1951b:298—Cherbonnier,
1959: 49, pl. 7: fig. K; 1963:187.

a:

Heliophora orbiculus—Durham, 1955:185; 1966:U491, fig.
377, 4a—c.

MATERIAL EXAMINED.—None.

Remarks.—The inclusion of this species in the
faunal list for Ascension rests upon the record of
Bell (1881) of two bare tests. No further specimens
have been collected at Ascension. The species is not
known from St. Helena.

DIsTRIBUTION.—From Senegal to Angola (Mor-
tensen, 1940), also Ascension Island (Bell, 1881) and
the Cape Verdes (Mortensen, 1951b).

Order SPATANGOIDA
Family BRISSIDAE

Brissus unicolor (Leske)
Brissus unicolor.—Mortensen, 1951a:509, pl. 38: fig. 10.

MATERIAL EXAMINED.—Ascension Island, collected by divers,
Ascension Historical Society, approximately 1972, specimen
on display in Ascension Historical Society Museum, Fort
Hayes, Ascension Island.

RemMark.—The single specimen of Brissus uni-
color was apparently collected by diving at Ascen-
sion Island, and was initially identified by Ailsa M.
Clark of the British Museum (Natural History). No
other specimens of the species have been recorded
from Ascension. The bare test was photographed,
and approximate dimensions (based upon the photo-
graph) are as follows: length 115 mm, greatest width
87 mm, greatest height 52 mm, height at level of
apical system 42 mm, length of left posterolateral
petal 44 mm, length of left anterolateral petal
29 mm.

DistripuTion.—Mortensen (195la) records the
species from the “warmer regions of both the
Western and Eastern Atlantic.” The record from
St. Helena (Clark, 1952) is exceedingly doubtful
although, in the light of the known occurrence of
the species at Ascension, it seems quite likely that
it could also occur at St. Helena. Chesher (1972)
notes that some specimens of Brissus from the Gulf
of California cannot be distinguished from B. uni-
color, and he considers that this species ranges into
the eastern Pacific, where its relationships with
B. obesus Verrill have yet to be determined.

SMITHSONIAN CONTRIBUTIONS TO MARINE SCIENCES

Class HOLOTHUROIDEA
Order ASPIDOCHIROTIDA
Family HOLOTHURIIDAE

Holothuria (Halodeima) grisea Selenka
FicurEe lla-e

Holothuria grisea—Deichmann, 1930:76, pl. 5: figs. 1-4.
Holothuria (Holothuria) grisea—Panning, 1934a:31, fig. 23.
Ludwigothuria grisea—Deichmann, 1958:311.

Holothuria (Halodeima) grisea.—Rowe, 1969:138.

MATERIAL EXAMINED—RBM 3, 1 specimen; RBM 10, 1
specimen. Asc. 1B-76, 8 specimens; Asc. 5A-76, 1 specimen;
Asc. 6A-76, 6 specimens.

DisTRIBUTION.—In external features, specimens
conform with previous descriptions. Color in life
generally reddish ground color, with yellowish and
brownish patches marking presence of dorsal tube
feet. Color in alcohol mottled greyish. Ventral feet
form conspicuous sole, light greenish-yellow in life,
brownish in alcohol.

Ossicles in body wall numerous tables and buttons
(rosettes). Tables (Figure 1la) with small disc carry-
ing few perforations; generally four large central
perforations surrounded by four to twelve smaller
peripheral holes. Edge of disc with approximately
12 blunt projections. Spire short, with one crossbar,
and terminating in 12 spines, of which four project
more or less vertically. Dimensions: average diam-
eter of disc 58 um (SD 6.75; SE 2.04); average height -
of spire 51 um (SD 4.28, SE 1.29). Buttons (rosettes)
apparently derived from basic form with 4-6 larger
holes and variable number of smaller holes (Figure
11d); edges of buttons indented. Length ranges
from about 36 to about 45 wm; average length 39
pm (SD5372) SE, 1°24):

Ventral tube feet contain well developed end
plates, together with rods (Figure 11b) with per-
forated ends; some rods develop four large lateral
perforations as well. Rods average approximately
115 zm in length.

Tentacles contain rods (Figure Ile) with perfo-
rated ends and with short spines near perforations.
Tentacle rods greatly variable in size, up to maxi
mum length of approximately 250 pm.

Remarks.—The ossicles of the Ascension repre-
sentatives of this species show some consistent dif

NUMBER 2 25

100 um

50 um

Ficure 11.—Holothuria (Halodeima) grisea Selenka: a, tables from body wall; b, rods from
ventral tube feet of Ascension specimens; c, rods from ventral tube feet of specimen from the
Bahamas (USNM 32489); d, buttons from body wall; e, rods from tentacles. Holothuria (Halo-
deima) manningi, new species: f, tables from body wall; g, buttons from body wall; h, rods
from tentacles; i, perforated plates from tube feet. Holothuria (Platyperona) sanctori Delle
Chiaje: j, buttons from body wall of specimen from Naples (USNM 17025); k, tables from body
wall; J, buttons from body wall. Isostichopus badionotus (Selenka): m, tables from body wall
of juvenile specimens.

26

ferences from those of West Indian specimens. The
tables are virtually identical, although those from
Ascension may be slightly higher. The buttons or
rosettes are smaller in the Ascension form (Deich-
mann, 1930:77, notes that they are about 50 pm
in diameter in West Indian forms), and appear to
have more numerous larger holes. While very few
of the ossicles illustrated by Deichmann (1930, pl. 5:
fig. 1) can be found in the ventral tube feet, most
are the shape shown in Figure 11b. By contrast,
tube foot ossicles from USNM 32489, a specimen
from Andros Island, Bahamas (Figure llc), are
quite different, with more irregular shape, and few
complete perforations in the ends. It can be seen,
however, that both of these ossicles follow the same
basic pattern. The tentacle rods are quite variable
in both populations, although the Ascension speci-
mens tend to have rods with more numerous per-
forations.

DistTRIBUTION.—Common in the tropical western
Atlantic (Deichmann, 1930), and also known from
West Africa (Greef, 1882).

HasitaT at AsceNnsion.—Fairly common under
rocks or on sand under rockly ledges. Despite bright
coloration, difficult to see because of covering of
particles of sand.

Holothuria (Halodeima) manningi, new species
Ficure 11f-i

MATERIAL EXAMINED.—Holotype: USNM_ E16167, total
length 90 mm. Asc. 3G-76, 12 Jul 1976, English Bay, Ascen-
sion Island, collected by snorkeling from depths of approxi-
mately 5 m in broad surge channel; substrate rock or coarse
coralline “sand.” Water temperature 25°C, salinity 34%o0.
Paratypes: USNM E16168, total length 95 and 80 mm. Same
locality data as holotype.

EtymMoLocy.—The species is named for Dr. Ray-
mond B. Manning, Department of Invertebrate
Zoology, National Museum of Natural History,
who has contributed so much to our knowledge of
Ascension Island invertebrates by his assiduous col-
lecting and who was the first to discover the speci-
mens described herein.

DtAGnosis.—Deposits include tables of average
height 54 wm with completely reduced disk and
with spire surmounted by 12 projections, and
minute circular buttons of average diameter 22 «m,
typically with 2 large and 2 small perforations, but

SMITHSONIAN CONTRIBUTIONS TO MARINE SCIENCES

nature and disposition of perforations highly vari-
able.

DescripTION.—Body cylindrical, lacking con-
spicuous tubercles dorsally. In life, dark brown
dorsally, darker along mid-dorsal interradius, fading
to lighter brown on flanks; ventral surface light
brown. In alcohol, color similar. Tentacles olive
green in life, yellowish green in alcohol. Skin
slightly rough to touch due to presence of numer-
ous ossicles. Dorsal and ventral tube feet identical.
Dorsally, feet sparsely scattered; ventrally, feet much
more numerous, but not forming conspicuous sole.
Largest specimen with conspicuous naked band
along mid-ventral radius, but naked area not obvi-
ous in smaller specimens.

Ossicles in dorsal and ventral body wall identical,
consisting of tables and buttons. Tables with com-
pletely reduced disc; basal portion rounded (Figure
I1f). op of spire with “‘maltese cross” comprising
12 projections, none of which appear to point ver-
tically. Spire with one crossbar. Average height of
tables 54 um (SD 3.63). Buttons minute, tending to
be circular. Typical form (Figure 1lg) has four per-
forations, two larger and two smaller, but nature
and disposition of perforations highly variable.
Most buttons with only two perforations. Average
diameter 22 ym (SD 2.28).

Tube feet contain end plates, tables, buttons, and
flat perforated plates (Figure 117), usually with two
larger perforations and numerous small perforations
at ends. Developmental stages of these plates com-
mon.

Tentacles contain scattered minute straight or
curved spinous rods (Figure 11h), averaging approxi-
mately 50 um in length.

Remarks.—This species falls within the subgenus
Halodeima of the genus Holothuria as defined by
Rowe (1969). It is immediately distinguished from
all species listed by Rowe, except Holothuria
(Halodeima) edulis Lesson, in having tables with a
completely reduced disc. This Indo-Pacific species
has tables that closely resemble those of Holothuria
(Halodeima) manningi, but the buttons in both
species are very different, for in H (H.) edulis the
buttons tend to be elongate, have generally larger
holes, and are frequently reduced to form X-shaped
granules. Further, this latter species has distinctive
coloration, being brown dorsally and rose red ven-
trally when alive (Clark, 1946); this color is quite
consistent and distinctive.

NUMBER 2

While relationship with H. (H.) edulis might be
postulated, H. (H.) manningi appears to bear no
close relationship to any species of Holothuria in
the Atlantic Ocean.

The species was found only at English Bay, and
only in minimum depths of 3-4 meters. Further
searching in the English Bay area revealed that the
species is relatively common and conspicuous, lying
exposed on rocks and calcareous sand.

Holothuria (Thymiosycia) arenicola Semper

Holothuria arenicola——Deichmann, 1930:66, pl. 4: figs. 1-9.
Holothuria (Holothuria) arenicola—Panning, 1935:88 fig. 73.
Bradtothuria arenicola—Deichmann, 1958:290.

Holothuria (Thymiosycia) arenicola—Rowe, 1969:145.

MATERIAL EXAMINED.—RBM 13, 1 specimen; RBM 22, 2
specimens; Asc. 5A-76, 2 specimens.

DEscrIPTION.—Specimens typical of species. Speci-
men from RBM 13 is juvenile, with tables only in
body wall; buttons absent. In adult specimens
tables average 61.5 um in diameter (SD 2.70, SE
0.85) and buttons average 48.8 »m in length (SD
1.69, SE 0.56).

Remarks.—In all characters examined, specimens
from Ascension conform well with descriptions of
the species. Dimensions of ossicles fall within the
range of variation for H. arenicola. Deichmann
(1930) noted that there was surprisingly little vari-
ation in ossicle size throughout the geographic range
of the species.

DIsTRIBUTION.—This species is cosmopolitan in
the tropics, generally living under rocks or burrow-
ing into sandy substrates,

HapiTaT aT AsceNsion.—R. B. Manning noted
that the specimens collected at RBM 22 were found
“burrowing deep in bottom of pool.” The speci-
mens from Asc. 5A-76 were found under rocks,
burrowing into coarse calcareous rubble.

Holothuria (Platyperona) sanctori Delle Chiaje
FicurE 11j-1

Holothuria farcimen.—Mortensen, 1933a:471, fig. 29.

Holothuria sanctori—Panning, 1934b:74, fig. 55.—Tortonese,
1965:61, figs. 21A, 22.

Microthele sanctori—Deichmann, 1958:287.

Holothuria (Platyperona) sanctori—Rowe, 1969:145. -

MATERIAL EXAMINED.—RBM 15, 2 specimens; Asc. 30-76,
3 specimens; Asc. 90-76, 1 specimen.

21

DEscRIPTION.—Specimens typical of species in
most respects, although ossicles show some differ-
ences. Tables apparently normal (Figure 11k), but
buttons consistently possess several small knobs
(Figure 111).

RemaRKs.—Most typical specimens of H. sanctori
have smooth buttons, with no trace of knobs on
their surface. A single specimen from Naples
(USNM 17025) possesses buttons with few knobs
(Figure 117); approximately six knobs per button
was the largest density observed. The presence of
numerous knobs on the buttons was also observed
by Mortensen (1933a) in the St. Helena specimens
he described as Holothuria farcimen Selenka.

DisTRIBUTION.—Mediterranean Sea, Azores (Se-
lenka, 1867; Chapman, 1955), St. Helena (Morten-
sen, 1933a). According to Tortonese (1965), the
bathymetric range is 5-30 meters. At St. Helena the
species was collected intertidally, and at a depth of
40 meters (Mortensen, 1933a). The Ascension rec-
ords are all intertidal.

HasiraT AT ASCENSION.—Under rocks on rocky
substrate.

Family STICHOPODIDAE

Isostichopus badionotus (Selenka)
Ficure 11m

Stichopus badionotus—Deichmann, 1930:80, pl. 5: figs. 30-36.
—Cherbonnier, 1976:631, pl. 1a-c: fig. 14-, fig. 2H—-o.
Isostochopus badionotus.—Deichmann, 1958:280.

MATERIAL EXAMINED.—RBM 9, 1 specimen; RBM 21, 1
specimen; Asc. 3A-76, 1 specimen (juvenile).

DESCRIPTION.—Specimens from station RBM 9
and RBM 21 are typical of the species. Juvenile
specimen from Asc. 3A-76, white alive and in alco-
hol, total length 10 mm. Body wall soft, with scat-
tered tube feet. Ossicles exclusively tables (Figure
11m) with four central perforations, surrounded by
eight or more marginal perforations. Spire tall, with
four or more crossbars. Diameter of tables variable,
usually 50-60 wm; spire with same dimensions.

REMARKS.—Clark (1922) described the ossicles of
a juvenile 20 mm in length, and in that specimen
the tables were 75 wm across and the spire 50 «m
high, with two crossbars and with teeth on the top.

28

Differences from the ossicles of the Ascension speci-
men are obvious. However, Clark’s specimen was
approximately twice as long as the present one, and
he noted (Clark, 1922:57) that growth changes in
the ossicles can be quite dramatic. It does not seem
likely that two species of Isostichopus occur at
Ascension Island; thus the juvenile specimen is
assigned to J. badionotus.

DisTRIBUTION.—Deichmann (1930) notes that this
species is distributed over “most of the West Indian
Seas, from Bermuda to Panama.” Greef (1882) re-
ported the species (as Stichopus maculata) from
West Africa, as did Bell (1883) (as S. assimilis), and
Cherbonnier (1975) confirmed that J. badionotus
occurs in West African waters. No stichopodids are
known from St. Helena.

HasiTtat AT ASCENSION.—Exposed on sand or
rock. Not common intertidally.

Order APODIDA
Family SYNAPTIDAE

Euapta lappa (Miller)

Euapta lappa—Clark, 1924:464, pl. 1: figs. 5-7—Heding,
1928:136, figs 8.2, 9.4, 10.5, 10.6—Mortensen, 1933a:471.

SMITHSONIAN CONTRIBUTIONS TO MARINE SCIENCES

MATERIAL EXAMINED.—British Museum, unregistered holo-
thurian, English Bay, Ascension Island; Ascension Island
Historical Society, Fort Hayes Museum, Dec 1972, one frag-
ment, anterior end of body.

DEscRIPTION.—Specimen comprises anterior end
of body approximately 220 mm in length. Tentacles
14, Color mottled brown. Ossicles comprise anchors
and anchor plates, all of approximately the same
size. Anchors average 368 ym in length and 210 pm
in width, and plates average 233 ym in length and
166 wm in width.

ReMARKS.—This is a typical specimen of E. lappa.
The dimensions of the anchors and plates conform
well with those given by Clark (1924) and Heding
(1928).

The label with the specimen notes: “white sea-
slug apparently is never observed during daylight
hours. This specimen was recovered during dark-
ness.” In spite of searching in a variety of habitats
during the 1976 expedition, no further specimens
were found. Apparently the species is not common
in shallow water.

DistRIBUTION.—E. lappa is common in the West
Indies and the Caribbean area generally. Theel
(1886) records the species or a close relative, from
off ‘Teneriffe in the Canary Islands, and Mortensen
(1933a) described what was probably this species
from St. Helena.

Literature Cited

Agassiz, A.
1881. Report on the Echinoidea Dredged by H.MSS.
Challenger during the Years 1873-76. Report on
the Scientific Results of the Voyage of H.M.S.
Challenger, Zoology, 3(9): 321 pages, 45 plates.
Atkins, F. B., P. E. Baker, J. D. Bell, and D.G.W. Smith
1964. Oxford Expedition to Ascension Island. Nature,
204:722-724.
Bell, F. J.
1881. Report of a Collection Made by Mr. T. Conry in
Ascension Island: Echinodermata. Annals and Maga-
zine of Natural History, series 5, 44:436-438.

1882. Description of a New Species of the Genus Archaster
from St. Helena. Annals and Magazine of Natural
History, series 5, 48:440-441.

1883. Studies on the Holothuroidea, II: Descriptions of
New Species. Proceedings of the Zoological Society
of London 1883:58-62, plate XV.

Briggs, J. C.

1974. Marine Zoogeography. x + 475 pages. New York:

McGraw-Hill Book Company.
Brito, I. M.

1962. Ensaio de catalogo dos Equinodermas do Brasil.
Universidade do Brasil Faculdade Nacional de
Filosofia, Centro de Estudos Zoologicos, 13:1-10, 3
plates.

1971. Contribuicao ao conhecimento dos Equinodermas

de Ilha la Trinidade, Brasil. Archivos do Museu
Nacional, 54:261-265.
Chace, F. A., Jr., and R. B. Manning
1972. Two New Caridean Shrimps, One Representing a
New Family, from Marine Pools on Ascension
Island (Crustacea: Decapoda: Natantia). Smithso-
nian Contributions to Zoology, 131:1-18, 11 figures.
Chapman, G.
1955. Aspects of the Fauna and Flora of the Azores,
IV: Echinodermata. Annals and Magazine of Natu-
ral History, series 12, 8:398—400.
Cherbonnier, G.
1959. Echinides. Expedition Oceanographique Belge dans
les Eaux Coutieres Africaines de VAtlantique Sud
(1948-1949), 3(6):35-59, plates I-X.

1963. Echinodermes des cotes du Cameroun récoltés par
A. Crosnier en Decembre 1962-Janvier 1963. Bul-
letin du Muséum National d‘Histoire Naturelle,
35(2):179-193, 3 figures.

1975. Note sur la presence dans la Golfe de Guinee, de

VHolothurie Aspidochirote Stichopus badionotus
Selenka (—St. maculatus Greef). Bulletin du Mu-
seum national d’Histoire naturelle Zoologie, 210:
631-638, I plate, 2 figures.

29

Chesher, R. H.

1966. The R/V Pillsbury Deep-Sea Biological Expedition
to the Gulf of Guinea, 1964-65, 10: Report on the
Echinoidea Collected by the R/V Pillsbury in the
Gulf of Guinea. Studies in Tropical Oceanography,
Institute of Marine Sciences (University of Miami),
4(1):209-223.

1972. The Status of Knowledge of Panamanian Echinoids,
1971, with Comments on Other Echinoderms. Bul-
letin of the Biological Society of Washington, 2:
139-158, figures 1-2.
Clark, A. M.
1955. Echinodermata of the Gold Coast. Journal of the

West African Science Association, 1(2):16—-56, figures
1-23.

1967. Notes on the Family Ophiothricidae (Ophiuroidea).
Annals and Magazine of Natural History, series 13,
9:637-655.

Clark, A. M., and A. H. Clark

1954. A Revision of the Sea-Star of the Genus Tethyaster.
Smithsonian Miscellaneous Collections, 122(11):1-27,
12 plates, 2 figures.

Clark, H. L.

1912. Hawaiian and Other Pacific Echini: The Pedinidae,
Phymosomatidae, Stomopneustidae, Echinidae, Tem-
nopleuridae, Strognylocentrotidae, and Echinometri-
dae. Memoirs of the Museum of Comparative Zool-
ogy, 34(4):209-383, 32 plates.

The Echinoderm Fauna of Torres Strait: Its Com-
position and Origin. Carnegie Institution of Wash-
ington Publication, 214(10): viii + 224 pages, 38
plates.

Report on the Echinoidea of the Barbados-Antigua
Expedition of 1918. Bulletin from the Laboratory
of Natural History of the University of Iowa 9(5):
103-121, 2 plates.

The Holothurians of the Genus Stichopus. Bul-
letin of the Museum of Comparative Zoology, 65(3):
39-74, 2 plates.

The Holothurians of the Museum of Comparative
Zoology: The Synaptinae. Bulletin of the Museum
of Comparative Zoology, 65(13):459-501, 12 plates.
A Catalogue of the Recent Sea-urchins (Echinoidea)
in the Collection of the British Museum (Natural
History). xxviii + 250 pages, 12 plates. London:
British Museum (Natural History).

1921a.

1921b.

1922.

1924.

1925.

1946. The Echinoderm Fauna of Australia: Its Composi-
tion and Origin. Carnegie Institution of Washington

Publication, 566: iv + 567 pages.

30

Cuninghame, J.

1699. A Catalogue of Shells, etc., Gathered at the Island
of Ascension by Mr. James Cuninghame Chirurgeon,
with What Plants He There Observed; Communi-
cated to Mr. James Petiver, Apothecary, and Fel-
low of the Royal Society. Philosophical Transactions
of the Royal Society of London, 21:295-300.

Cunningham, J. T.

1910. On the Marine Fishes and Invertebrates of St.
Helena. Proceedings of the Zoological Society of
London, 1910:86-131, plates IV-VII, figures 3-6.

Daly, R. A.

1922. The Geology of Ascension and St. Helena Islands.
Geological Magazine, 59:146-156.

1925. The Geology of Ascension Island. Proceedings of
the American Academy of Arts and Sciences, 60(1):
1-80.

Deichmann, E.

1930. The Holothurians of the Western Part of the At-
lantic Ocean. Bulletin of the Museum of Compara-
tive Zoology, 71(3):43-226, 24 plates.

1958. The Holothurioidea Collected by the Velero II
and IV during the Years 1932 to 1954, Part II:
Aspidochirota. Allan Hancock Pacific Expeditions,
11:249-349, 9 plates.

Devaney, D. L.

1973. Zoogeography and Faunal Composition of South-
eastern Polynesian Asterozoan Echinoderms. In R.
Fraser, editor, Oceanography of the South Pacific
1972, pages 357-366. Wellington: New Zealand
National Commission for UNESCO.

Durham, J. W.

1955. Classification of Clypeasteroid Echinoids. University
of California Publications in Geological Sciences,
31(4):73-198, plates 3-4.

1966. Clypeasteroids. In R. C. Moore, editor, Treatise on
Invertebrate Paleontology, Part U (Echinodermata
3):U450-491. Lawrence, Kansas: University of Kansas
Press.

Fisher, W. K.

1940. Asteroidea. “Discovery” Reports, 20:69-306, figures

A-M, .23 plates.
Greeff, R.

1882. Echinodermen, beobachtet auf einer reise nach der
Guinea-Insel Sao Thome. Zoologischer Anzeiger,
5:114-120, 135-139, 156-159.

Heding, S. G.

1928. Synaptidae. Videnskabelige Meddelelser fra Dansk
naturhistorisk Forening Kjgbenhavn, 85:105-323,
69 figures, plates IT, III.

Koehler, R.

1904. Ophiures nouvelles ou peu connues. Memoires
Societe Zoologique Francaise, 17:54-119.

1908. Asteries, Ophiures et Echinides de l’expedition ant-
arctique ecossaise. Transactions of the Royal Society
of Edinburgh, 46:529-649, 18 plates.

1914. Echinoderma, I: Asteroidea, Ophiuroidea et Echi-
noidea. Beitrige zur Kenntnis der Meeresfauna
Westafrikas (Hamburg), 1(2):129-303, plates iv—xv.

SMITHSONIAN CONTRIBUTIONS TO MARINE SCIENCES

Lima-Verde, J. S.

1969. Primeira contribuicao ao inventario dos Equinoder-
mas do Nordeste Brasileiro. Archivos de Ciencias
do Mar, 9(1):9-13.

MacArthur, R. H., and E. O. Wilson

1967. The Theory of Island Biogeography. xi + 203
pages, 60 figures. Princeton, New Jersey: Princeton
University Press.

McPherson, B. F.

1968. Contributions to the Biology of the Sea Urchin
Eucidaris tribuloides (Lamarck). Bulletin of Marine
Science, 18(2):400-443, 26 figures.

Madsen, F. J.

1950. The Echinoderms Collected by the Atlantide-Expe-
dition 1945-46, I: Asteroidea. “Atlantide” Report,
1:167-222, plates XIV-XVI, 11 figures.

1970. West African Ophiuroids. “Atlantide” Report, 11:
151-243, 49 figures.

Mayr, E.

1963. Animal Species and Evolution. xiv + 797 pages.

Cambridge: Belknap Press of Harvard University.
Melliss, J. C.

1875. St. Helena: A Physical, Historical, and Topographi-
cal Description of the Island; Including Its Geol-
ogy, Fauna, Flora, and Meteorology. xiv + 426
pages, 62 plates, 1 map. London.

Metcalf, W. G., A. D. Voorhis and M. C. Statcup

1962. The Atlantic Equatorial Undercurrent. Journal of

Geophysical Research, 67(6):2499-2508.
Mortensen, T.

1903. Echinoidea. The Danish Ingolf-Expedition, 4(1):
1-194, 21 plates, 12 figures. Copenhagen: Bianco
Luno.

1909. Die Echinoiden. Deutsche Sudpolar-Expedition
1901-1903 XI (Zoologie, 1): 113 pages, 19 plates.

1921. Studies of the Development and Larval Forms of
Echinoderms. 261 pages, 33 plates, 102 figures.
Copenhagen: G.E.C. Gad.

1928. A Monograph of the Echinoidea, Cidaroidea. Vol- _
ume 1, 551 pages, 88 plates, 173 figures. Copen-
hagen: Bianco Luno.

1931. Contributions to the Study of the Development
and Larval Forms of Echinoderms I-II. Danske
Videnskabernes Selskabs Skrifter, Naturvidenskabelig
og mathematisk Afdeling, series 9, 4(1):1—-39, plates
I-VII, 12 figures.

1932. New Contributions to the Knowledge of the Cida-
rids I-II. Danske Videnskabernes Selskabs Skrifter,
Naturvidenskabelig og mathematisk Afdeling, series
9, 4(4):145-182, 18 figures, 10 plates.

1933a. The Echinoderms of St. Helena. Videnskabelige
Meddelelser fra Dansk naturhistorisk Forening 1
Kjébenhavn, 93:401-472, plates XX-XXII, 29 figures.

1933b. Echinoderms of South Africa (Asteroidea and Ophi-
uroidea). Videnskabelige Meddelelser fra Dansk
naturhistorisk Forening i Kjgbenhavn, 93:215-400.

1935. A Monograph of the Echinoidea, Bothriocidaroida,

Melonechinoida, Lepidocentroida, and Strirodonta.
Volume 2, 647 pages, 377 figures, 89 plates. Copen-
hagen: Bianco Luno.

NUMBER 2

1936. Echinoidea and Ophiuroidea. “Discovery” Reports,
12:199-348, plates I-IX.

1938. Contributions to the Study of the Development
and Larval Forms of Echinoderms, IV. Danske
Videnskabernes Selskabs Skrifter, Naturvidenskabelig
og mathematisk Afdeling, series 9, 7(3):1-59, plates
I-XII, 30 figures.

1940. A Monograph of the Echinoidea: Aulodonta. Vol-
ume 3, number 1, 370 pages, 197 figures, 77 plates.
Copenhagen: Bianco Luno.

1943. A Monograph of the Echinoidea: Camarodonta, 1.
Volume 3, number 2, 553 pages, 321 figures, 56
plates. Copenhagen: Bianco Luno.

1948. A Monograph of the Echinoidea: Clypeastroida.
Volume 4, number 2, 471 pages, 258 figures, 72
plates. Copenhagen: Bianco Luno. i

195la. A Monograph of the Echinoidea: Spatangoida, 2.
Volume 5, number 2, 593 pages, 286 figures, 64
plates. Copenhagen: Bianco Luno.

1951b. Report on the Echinoidea Collected by the “At-
lantide” Expedition. Atlantide Report 2:293-303,
plates I-II.

Panning, A.

1934a. Die Gattung Holothuria, 2. Mitteilungen aus dem
Zoologischen Staatsinstitut und Zoologischen Mu-
seum in Hamburg, 45:24-50, figures 22-44.

1934b. Die Gattung Holothuria, 3. Mitteilungen aus dem
Zoologischen Staatsinstitut und Zoologischen Mu-
seum in Hamburg, 45:65-84, figures 45-71.

1935. Die Gattung Holothuria, 4. Mitteilungen aus dem
Zoologischen Staatsinstitut und Zoologischen Mu-
seum in Hamburg, 45:85-107, figures 72-102.

Robinson, A. R.
1976. Eddies and Ocean Circulation. Oceanus, 19(3):2-17.
Rosewater, J.

1975. An Annotated List of the Marine Mollusks of
Ascension Island, South Atlantic Ocean. Smithso-
nian Contributions to Zoology, 189:1-41, 24 figures.

Rowe, F.W.E.

1969. A Review of the Family Holothuriidae (Holothuri-
oidea: Aspidochirotida). Bulletin of the British
Museum (Natural History), Zoology, 18(4):119-170,
21 figures.

Scheltema, R. S.

1968. Dispersal of Larvae by Equatorial Ocean Currents
and Its Importance to the Zoogeography of Shoal-
water Tropical Species. Nature, 217:1159-1162.

1971. Larval Dispersal as a Means of Genetic Exchange
between Geographically Separated Populations of
Shallow-water Benthic Marine Gastropods. The
Biological Bulletin, 140(2):284-322.

31

Selenka, E.

1867. Beitrége zur Anatomie und Systematik der Holo-

thurien. Zeitschrift fur Wissenschaftliche Zoologie,
_ 17:291-374, plates 17-20.
Sturm, M., and K. Voigt

1966. Observations on the Structure of the Equatorial
Undercurrent in the Gulf of Guinea in 1964.
Journal of Geophysical Research, 71(12):3105-3108.

Sullivan, W.

1974. Continents in Motion. 399 pages. New York: Mc-

Graw-Hill Book Company.
Theel, H.

1886. Report on the Holothurioidea Dredged by H.M.S.
Challenger during the Years 1873-1876, Part 2.
Report on the Scientific Results of the Voyage of
H.M.S. Challenger, Zoology, 14(39):290 pages, 16
plates.

Thorson, G.

1946. Reproduction and Larval Development of Danish
Marine Bottom Invertebrates with Special Reference
to the Planktonic Larvae in the Sound (Oresund).
Meddelelser Kommissionen Havundersggelser Kjob
benhavn, Plankton, 4(1):343-367.

1950. Reproductive and Larval Ecology of Marine Bottom
Invertebrates. Biological Reviews of the Cambridge
Philosophical Society, 25(1):1-45, 6 figures.

Tommasi, L. R.

1966. Lista dos Equinoides recentes do Brasil. Contribui-
coes avulsas do Instituto Oceanografico Sao Paulo,
11:1-50, 9 plates, 72 figures.

Tortonese, E.

1965. Echinodermata. Fauna d'Italia. Volume 6, xv +

422 pages, 186 figures. Bologna: Edizioni Calderini.
Utinomi, H.

1954. A Checklist of Echinoids Found in the Kii Region.
Publications of the Seto Marine Biological Labora-
tory, 3(3):339-358.

Voight, K.

1975. The Atlantic Equatorial Undercurrent. Technical
Series Intergovernmental Oceanographic Commis-
sion, 11:12-19.

Wilson, J. T.

1963. Evidence from Islands on the Spreading of Ocean

Floors. Nature, 197:536-538.
Yamaguchi, M.

1973. Early Life Histories of Coral Reef Asteroids, with
Special Reference to Acanthaster planci (L.). In
Jones and Endean, editors, Biology and Geology
of Coral Reefs, I1(Biology, 1):369-387. New York:
Academic Press.

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