Systematics and Ecology of the
Sea-Urchin Genus Centrostephanus
(Echinodermata: Echinoidea)

from the Atlantic and Eastern
Pacific Oceans

DAVID L- PAWSON

P< serum ces ee si maine and i Ste - agian seme .
” JOHN E. “MITEER =" ==

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SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES © NUMBER 20

Systematics and Ecology of the Sea-Urchin
Genus Centrostephanus (Echinodermata:
Echinoidea) from the Atlantic and
Eastern Pacific Oceans

David L. Pawson
and John E. Miller

ISSUED
SEP 26 1983

CMITHSONIAN PUBLICATIONS

SMITHSONIAN INSTITUTION PRESS \\\ | )5U)\
City of Washington |
1983

Ze Jos) JE IR ANGIE

Pawson, David L., and John E. Miller. Systematics and Ecology of the Sea-
Urchin Genus Centrostephanus (Echinodermata: Echinoidea) from the Atlantic
and Eastern Pacific Oceans. Smithsonian Contributions to the Marine Sciences,
number 20, 15 pages, 5 figures, 5 tables, 1983.—Surveys by the JoAnson-Sea-
Link submersibles have revealed the presence of large populations of black
Centrostephanus, superficially resembling Dzadema antillarum Philippi, along the
shelf edge prominences off the east coast of Florida in depths of 48-80 meters.
Typical habitats are aggregations of dead coral rubble, with seasonal growths
of leafy red algae. Some aspects of the biology of these echinoids are described.

We affirm that only a single species of the genus, Centrostephanus longispinus
(Philippi), occurs in the Atlantic Ocean. As Fell (1975) and Serafy (1979) have
shown, western Atlantic populations can be referred to the subspecies Centros-
tephanus longispinus rubicingulus H.L. Clark, which usually differs from the
typical subspecies in possessing uniformly black spines rather than banded
purple and yellowish white spines when fully grown. C. besnard: Bernasconi
from Isla Trindade, Brazil, is herein synonymized with C. longispinus rubicin-
gulus. C. coronatus (Verrill) from California and the Galapagos Islands differs
little from C. longispinus and the two species may yet prove to be subjective
synonyms. There is some evidence to suggest that Gulf of California popula-
tions of C. coronatus differs from California populations at the subspecies level.

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 of eastern North America.

Library of Congress Cataloging in Publication Data

Pawson, David L. (David Leo), 1938-

Sea urchin genus Centrostephanus (Echinodermata: Echinoidea)

(Smithsonian contributions to the marine sciences ; no. 20)

Bibliography: p.

Supt. of Docs. no.: ST 1.41:20

1. Centrostephanus. 2. Echinodermata—Atlantic Ocean. 3. Echinodermata—Pacific
Ocean. I. Miller, John E. II. Title. III. Series.

QL384.E2P38 1983 593.95 83-600054

Contents

JTNTROGINCTHONM | -crh"55 4 nse oA OILS e ocho eG ONS SRE Ne MiSs aera 1
INCKMOW EAST CHItSMMERE ye ne mha Seok: me hdl hot Bota, 2s: 1
COcniiostephanusmeeterssglODD) ee Es gs ee oe oe 1
Cenizosiephanusslonarspinus|(Ehilippi) 9.2)... 4. 62 eee ee 3
Key to Subspecies of C. longispinus (Philippi) ..................... 4
Centrostephanus longispinus longispinus (Philippi) .................... 4
Centrostephanus longispinus rubicingulus H.L. Clark .................. 4
Czatiostephanusycoronatiusn Wenn) pea the gg 2
ILititerranrenns (Caneel: «| a se Se encase ee ees as enter 4

eas

ill

Ts,

Oy

Systematics and Ecology of the Sea-Urchin
Genus Centrostephanus (Echinodermata:
Echinoidea) from the Atlantic and
Eastern Pacific Oceans

David L. Pawson
and John E. Miller

Introduction

During investigations of the structure and
faunas of shelf-edge prominences along the east
coast of Florida (see Avent et al., 1977; Reed,
1980), a long-spined black sea urchin was found
to be a notable and numerous inhabitant of coral
rubble zones. Using Harbor Branch Foundation,
Inc. submersibles Johnson-Sea-Link 1 and II, we
have been able to collect specimens of this urchin,
to observe them in their natural habitat, and
maintain them alive in aquaria. Closer exami-
nation revealed that the urchin is a species of
Centrostephanus, a widespread Atlantic and Indo-
Pacific diadematid genus.

In this paper we describe this newly found
material of Centrostephanus and discuss some as-
pects of its biology and systematic position. In
addition, we compare our Florida specimens with
C. longispinus from the eastern Atlantic and C.
coronatus from the eastern Pacific.

ACKNOWLEDGMENTS.— Determinations of algal
species were kindly made by S.M. Blair, Harbor

David L. Pawson, Department of Invertebrate Zoology, National
Museum of Natural History, Smithsonian Institution, Washington,
D.C. 20560. John E. Miller, Harbor Branch Foundation, Inc., R.R.
1, Box 196, Fort Pierce, Florida, 33450.

Branch Foundation. We thank F.M. Bayer,
Smithsonian Institution, for his helpful insight
with several nomenclatural problems. Loan of
museum specimens was made possible by Robert
M. Woollacott, Museum of Comparative Zool-
ogy, Harvard, and Alberto Larrain, Allan Han-
cock Foundation, University of Southern Califor-
nia. We are indebted to the crews of Johnson-Sea-
Link submersibles and R/V Johnson for their
professional skills during submersible operations.
We are grateful to G. Hendler and R.L. Turner
for critically reviewing the manuscript. Funding
for this study was granted by the Smithsonian
Institution and Harbor Branch Foundation, Inc.
This paper represents Contribution No. 118 from
the Smithsonian Marine Station at Link Port and
Contribution No. 336 from Harbor Branch Foun-
dation, Inc.

Centrostephanus Peters, 1855

Diacnosis.—Like Diadema, but with buccal
plates carrying spines and pedicellariae. Globifer-
ous pedicellariae present.

TypE-SpEciEs.— Centrostephanus longispinus (Phi-
lippi).

ComposITION OF THE GENUS.—Centrostephanus

2 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

APICAL SYSTEM AS % OF HORIZONTAL DIAMETER

10 20 30
HORIZONTAL DIAMETER

40 50 60

Figure 1.—Relationship of apical system to horizontal diameter (in mm) in species of
Centrostephanus (closed circles = C. longispinus longispinus (Philippi) from eastern Atlantic; open
circles = C. longispinus rubicingulus H.L. Clark from east coast of Florida; closed triangles = C.
coronatus (Verrill) from lower California coast; open triangles = C. coronatus from Gulf of

California).

currently comprises ten nominal species, widely
distributed in the Atlantic and Indo-Pacific. In
his monograph of the Echinoidea, Mortensen
(1940) described eight species of Centrostephanus.
Later, Bernasconi (1955a,b) described C. besnardi
from Isla Trindade, Brazil, and Fell (1975) de-
scribed C. syluiae from San Felix and Juan Fer-
nandez Islands, off Chile. Mortensen (1940) noted
that C. longispinus and C. rubicingulus were appar-
ently closely related; he believed that C. rubscin-
gulus could be distinguished from C. longispinus by
the presence in the former species of more nu-
merous and larger secondary interambulacral tu-
bercles. Fell (1975) briefly reviewed the genus and
found that differences in size and number of

secondary interambulacral tubercles were not re-
liable systematic characters. Fell (1975) suggested
that C. rubicingulus should be reduced to a subspe-
cies of C. longispinus, and he noted that the three
nominal Atlantic Ocean species, C. longispinus, C.
rubicingulus, and C. besnardi, along with juvenile C.
coronatus from the eastern Pacific, “cannot be
distinguished morphologically using adult char-
acters. Identifications must be done by locality”
(Fell, 1975:181). Serafy (1979) followed Fell’s sug-
gestion, and reduced C. rubicingulus to a subspecies
of C. longispinus (Phillippi).

CLAVIFORM SPINES IN Centrostephanus.—Several
authors, including Hamann (1887), Mortensen
(1940), and Fell (1975) have called attention to

NUMBER 20

SPINE: WHORLS PER CM LENGTH

20 30

40 50 60

TEST: HORIZONTAL DIAMETER

Ficure 2.—Relationship of whorls of spinelets of spines (verticillations) to horizontal diameter
of test in species of Centrostephanus (circles = C. coronatus from Gulf of California; triangles = C.
coronatus from lower California coast; squares = C. longispinus rubicingulus from east coast of
Florida; open symbols = ambulacral spines; closed symbols = adambulacral spines).

the presence of peculiar club-shaped spines on the
adapical plates in some species of Centrostephanus.
These highly motile spines, tipped with an intense
pink or purple pigment, are found in only four of
the ten nominal species: C. longispinus, C. rubicin-
gulus, C. besnardi, and C. coronatus. These closely
related “species” form a natural (Tethyan?)
group occurring in the eastern Atlantic, the west-
ern Atlantic, and in the eastern Pacific. Fell
(1975) noted that C. asteriscus Agassiz and Clark,
1907, from Hawaii and the Kei Islands, has some
features in common with the species mentioned
above but differs in lacking claviform spines. Fell
(1975) referred the species that lack claviform
spines, C. rodgersiz (Agassiz, 1863), C. tenuzspinus
Clark, 1914, and C. syluiae Fell, 1975, to a single
“superspecies.” If the genus Centrostephanus were
to be split into two genera or subgenera based
upon the presence or absence of claviform spines,
the genus name T7ichodiadema Agassiz, 1863, type-

species rodgerstt Agassiz, is available for species
that lack these spines.

CHANGES PROPOSED FOR THE GENUS Centroste-
phanus.—Results from our studies of live material
and existing museum collections lead us to agree
with the suggestions of Fell (1975). We accept
Serafy’s reassignment of Centrostephanus rubicingulus
to a subspecies of C. longispinus, although the
diagnoses of the subspecies require some modifi-
cation. Further, we refer C. besnardi Bernasconi to
synonymy with C. longispinus rubicingulus H.L.
Clark, thereby restricting Centrostephanus to eight
nominal species.

Centrostephanus longispinus (Philippi)

Diacnosis.—Claviform spines present adapi-
cally. Apical system usually 30%-40% of test di-
ameter. Spines usually with well-developed tra-

+ SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

beculae between wedges. Eastern and western
Atlantic Ocean.

RemMarRKS.—T'wo subspecies of C. longispinus
may now be recognized.

Key to Subspecies of C. longispinus

Specimens of all sizes with banded spines, with purple bands on yellowish
white or light green ground color; from the Mediterranean and eastern
Atlantic: es anes Centrostephanus longispinus longispinus (Philippi)

Small specimens (h.d. <25 mm) with banded spines, reddish brown on whitish
ground color; larger specimens (h.d. >25 mm) with uniformly blackish
spines or with banded spines, the bands brown on lighter brown ground

color; from the western Atlantic ...

secre, eager Centrostephanus longispinus rubicingulus H.L. Clark

Centrostephanus longispinus longispinus

(Philippi)
Ficure 1; TABLE 1

Diadema longispina Philippi, 1845:354.

Centrostephanus longispinus.—Mortensen, 1940:300, pl. 24: figs.
1-11; pl. 25: figs. 11, 12; pl. 75: figs. 5-24 [complete
synonymy up to 1938]; 1951:296.—Tortonese, 1953:214;
1965:311, figs. 143-145; 1976:238.—Chapman, 1955:
399.—Cherbonnier, 1956:17; 1959:44, pl. 4: figs. B-L; pl.
5: figs. A, B; 1963:186.—Millott, 1957, fig. 4.—Repelin,
1962:4, pl. 1: fig. 6; pl. 2: figs. 7, 8.—Chesher, 1966:215.—
Markel, 1970:190, figs. 1, 2, 4, 5, 14a.

?Centrostephanus longispinus.—Alvardo and Alvarez, 1964: 278.

MatTeERIAL EXAMINED.—Universitetets Zoolo-
giske Museum, Copenhagen, Atlantide Expedition,
Sta. 61, 106, 116, Gulf of Guinea, 53-95 meters,
seven specimens. Museum of Comparative Zool-
ogy, Harvard University, Cat. No. 689, 4573,
Northeastern Atlantic, two specimens.

Diacnosis.—Spines banded purple and yellow-
ish white. Mediterranean and eastern Atlantic.

Type-Loca.ity.—Palermo, Sicily.

DeEscrIPTION.—Specimens examined were typ-
ical of the subspecies as described by Mortensen
(1940) and others. Information on numbers of
test plates, diameter of apical system and peris-
tome, and on structure of spines, is presented in
Table 1.

DistripuTion.—Western Mediterranean; east-
ern Atlantic, from Morocco to Gabon; Cape
Verde Islands and the Azores (Mortensen, 1940,
1951; Chesher, 1966). Depth range 42-208 me-

ters.

Remarks.—This subspecies has been _thor-
oughly described by Koehler (1895) and Morten-
sen (1904, 1940), and requires no further com-
ment here. Unlike C. longispinus rubicingulus (see
below), apparently no specimens with spines of a
solid dark color have been found; in even the
largest specimens the spines are banded.

Centrostephanus longispinus rubicingulus

H.L. Clark

Ficures 1-5; TaBves 2, 3

Centrostephanus rubicingulus H.L. Clark, 1921:108, pl. 1: figs. 1,
De

Centrostephanus rubricingulus H.L. Clark, 1925:52.—Mortensen,
1940:308, pl. 36: figs. 3-6, 11, 12, pl. 76: figs. 10-12, 14—
18, 22.—A.H. Clark, 1954:374.—Lewis, 1961:53.—Fell,
1975:180.

Centrostephanus besnardi Bernasconi, 1955a:92; 1955b:56, figs.
1-9, pl. 1: figs. 2, 6; pl. 7: fig. 2a-c.—Brito, 1962:5, pl. 3:
fig. 2; 1968: 20, pl. 9: figs. 2, 3.—Tommasi, 1967:10, figs.
Is 2s

Centrostephanus longispinus rubricingulus.—Serafy, 1979:28, fig.
8.

Species Name.—In his original description of
this species, H.L. Clark (1921:108) spelled the
species name as rubicingulus. In a footnote, Clark
stated that the name was derived from “rubus =
red + cingulus = a zone or band, in references to
the banded spines.” Here Clark was in error, for
rubus translates from Latin as a bramble or black-
berry. In a later paper (Clark, 1925) he spelled

NUMBER 20

TaBLe 1.—Measurements of Centrostephanus longispinus longispinus from the eastern Atlantic (Atl.
= Atlantide Expedition stations; MCZ = Museum of Comparative Zoology, Harvard University;
H.D. = horizontal diameter of test (in mm); Dia. = diameter; % H.D. = peristome or apical
system diameter expressed as percentage of test diameter; Plates A and IA = number of plates
in ambulacral and interambulacral columns, respectively; Wedges A and IA = number of
wedges in ambital ambulacral and interambulacral spines, respectively)

Peristome Apical system Plates Wedges

Specimen H.D

Dia. %H.D. Dias %ZH.D. A IA A IA
Atl. 106 16 7 44 6 38 11 9 14 15
Atl. 116 17 7 4] 6 35 12 10 15 17
Atl. 116 20 9 45 8 40 12 10 15 17
Atl. 116 21 10 48 9 43 12 11 15 18
MCZ 689 24 10 42 9 38 14 11 17 20
Atl. 61 31 14 45 10 32 15 12 19 19
Atl. 61 40 15 38 16 40 16 13 18 21
MCZ 4573 40 16 40 17 43 16 12 18 21
Atl. 61 45 16 36 17 38 17 14 ~ -

the species-name as rubricingulus without com-
ment, and every subsequent author has spelled
the name as rubricingulus. Clark’s original spelling
of the species name does not represent an incor-
rect original spelling in the sense of Article 32(c)
of the International Code of Zoological Nomenclature
(1961 edition). Clark made an incorrect translit-
eration and, according to the provisions of Article
32(a) (ii), this does not qualify as an inadvertent
error; the original spelling must therefore be re-
tained.

MatTeERIAL ExAMINED.—The description given
below is based upon study of numerous live and
preserved specimens collected from the east coast
of Florida (see Table 2). Representative speci-
mens have been deposited at the National Mu-
seum of Natural History, Smithsonian Institution
(USNM E28068, E28069), and the Indian River
Coastal Zone Museum, Harbor Branch Founda-
Hommince, GQRCZM 72:330, 72:331, 72:332,
9223335) 72:334).

Diacnosis.—Smaller specimens with banded
spines, bands reddish brown on a whitish ground
color. Larger specimens, horizontal diameter >
25 mm, with uniformly blackish spines or banded
spines, the bands brown on a lighter brown
ground color. Western Atlantic.

Type-Loca.ity.—Off Barbados, British West
Indies.

DescripTion.—Characters of test and spines
are similar to those described by Mortensen
(1940) for C. rubicingulus, although supposed dif-
ferences between C. longispinus and C. rubicingulus
noted by Mortensen are apparently not signifi-
cant, as pointed out by Fell (1975) and by the
present authors (see “Remarks,” p. 11). Infor-
mation on the numbers of test plates, the diame-
ter of peristome and apical system, and on the
structure of spines, is presented in Table 3.

In natural light, overall color of living speci-
mens appears uniformly black, except for distinc-
tively colored claviform spines and lighter colored
bases of primary spines. Under artificial light, the
apical system is reddish brown, madreporic plate
darker. Anal funnel and spines on the periproct
are dark brown. On the plates surrounding the
apical system, the proximal portion is deep red-
dish brown; the distal portion and all other plates
are lighter reddish brown. Thus a dark ring is
formed around the apical system. Claviform
spines are reddish brown proximally, tipped with
intense pink or purple pigment. Primary and
secondary spines are dark reddish brown to black;
the bases of the primary spines are light reddish

CONTRIBUTIONS TO THE MARINE SCIENCES

SMITHSONIAN

photographed

Florida, 27°49.7’N, 79°58.2’W, at

58 m (note dense aggregations of dead Oculina coral rubble in area).

5)

Upper and lower, Centrostephanus longispinus rubicingulus H.L. Clark

from Johnson-Sea-Link submersible on sea floor off Sebastian,

FicurE 3.—

NUMBER 20

VLE,
“Za
cs
~~

Ficure 4.—Centrostephanus longispinus rubicingulus H.L. Clark, from off Sebastian, Florida: upper,
specimen of 48 mm horizontal diameter; lower, specimen of 34 mm horizontal diameter.

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

Ficure 5.—Upper: Centrostephanus coronatus (Verrill), from Gulf of California, horizon-
tal diameter 33 mm; lower: Centrostephanus longispinus rubicingulus H.L. Clark, light-
colored form from Gulf of Mexico, horizontal diameter 37 mm (note banding of

spines).

NUMBER 20

Tasie 2.—Geographic and bathymetric distribution of Centrostephanus longispinus rubicingulus off

east central Florida, compiled from observations and collections by Johnson-Sea-Link submersibles

Locality Coordinates Depth (m)
Daytona Beach 28°57.7'N, 80°06.5’W 66-84
Cape Canaveral 28°30.0’N, 80° 10.4’W 43

28°29.6’N, 80°01.2’W 70
28°29.6'N, 80°06.5’W 51
28°29.5/N, 80°01.7'W 48
Satellite Beach 28°08.6'N, 79°59.6’W 73
Sebastian 27°50'11.7”N, 79°57'59.8”W 79
27°50'02.4”N, 79°58/08.3”W 67
27°49.7'N, 79°58.2’W 53-58
27°46'07.7”N, 79°58/10.5”W 85
27°46.0’N, 79°58.6’W 54
27°43.8’N, 79°58.4’W 60-70
27°43.8/N, 79°58.0'W 73
St. Lucie Inlet 27°12.5’N, 80°08.2’W 40

brown, uniformly flecked with brown pigment
spots. The podia are dark brown when con-
tracted, flecked with reddish brown pigment spots
when expanded. The peristome is creamy white,
except for triangular areas adjacent to ambs,
which are flecked with light brown spots. The
spines around the peristome are white, with light
orange tips. The cleaned test is green to yellowish
white.

In the USNM collections are several lots of
small (<20 mm h.d.) specimens of this subspecies;
all have reddish brown bands on white ground
color on their primary spines. Two additional
specimens, one of 32 mm h.d. from Sebastian,
Florida, and one of 37 mm h.d. from the Gulf of
Mexico, also have distinctly light-colored, banded
spines, although in these cases the bands are
darker brown on a lighter brown ground color. It
is possible that the black-spined adults are derived
from younger specimens with banded spines, as
in Diadema species (Mortensen, 1940). Alterna-
tively it is possible that populations of specimens
with banded spines occur elsewhere in the Flor-
ida-Caribbean area, and they have yet to be
adequately sampled.

Concerning the black color of the spines in the
east Florida specimens, one of us (J.E.M.) has
noted that when these spines are immersed in

liquid bleach, the black pigment disperses, and it
becomes evident that the spines are banded with
light and dark brown bands. In the field, occa-
sional specimens have a very light-colored aboral
surface and spines. In light of recently published
information, (Weber and Dambach, 1972), on
ameboid movement of pigment cells in C. longis-
pinus longispinus, a study of the pigmentation of
this western Atlantic form would undoubtedly be
rewarding.

DistRIBUTION.— Western Atlantic, from Florida
to Brazil and Gulf of Mexico, 33-310 meters.
Black, long-spined echinoids, usually referred to
as ““Diadema” (those collected or photographed in
depths greater than 40 meters along the U.S. east
coast from North Carolina to Florida), probably
also represent this subspecies.

Hasitrat.—Serafy (1979) found Centrostephanus
longispinus rubicingulus on algal substrata (Litho-
thamnion spp.) in the Gulf of Mexico off Egmont
Key and Sanibel Island, 55-73 m. During this
study we examined specimens collected from
shelf-edge prominences off east central Florida,
from Daytona Beach to St. Lucie Inlet, 40-85 m
(see Table 2). Many of these outcroppings are
covered with a dense layer of rubble, remains of
the reef-building scleractinian coral, Oculina van-
cosa Lesueur. In addition to Centrostephanus, these

10 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

TaBLe 3.—Measurements of Centrostephanus longispinus rubicingulus from western Atlantic (first 11

specimens with uniformly blackish spines, last 4 with banded spines; abbreviations as in Table

1; also Whorls A and IA = number of whorls per 1 cm of spine length in ambulacral and

interambulacral spines, respectively)

Peristome Apical system Plates Wedges Whorls
Locality H.D

Dia %H.D. Dia. %H.D. A IA A IA A IA
Sebastian 24 13 54 8 33 13 11 14 15 18 17
Sebastian 25 13 52 8 32 13 11 14 15 19 18
Sebastian 27 14 52 9 33 13 11 17 19 20 19
Sebastian 32 14 44 10 31 14 12 = = 18 17
Sebastian 37 19 51 14 36 15 12 19 22 18 18
Sebastian 44 22 50 18 40 15 12 18 18 20 20
Sebastian 45 23 51 18 40 16 13 20 21 = =
Daytona Beach 47 23 49 17 36 18 14 20 21 16 14
Sebastian 48 24 50 18 36 18 15 20 21 19 7
Sebastian 48 22 46 17 35 18 15 19 23 19 17
Sebastian 50 24 48 18 35 17 13 20 23 18 18
Puerto Rico 18 8 44 6 33 10 9 13 6 - =
Puerto Rico 22 11 51 9 42 12 11 VW 11 — =
Sebastian 32 = = = = = - 16 16 - =
Gulf of Mexico 37 = = = ~ = = 18 18 — =

rubble zones support a rich and diverse inverte-
brate fauna of mollusks, echinoderms, poly-
chaetes, decapods, and encrusting organisms (see
Avent et al., 1977; Reed, 1980; Reed et al., 1982).
Reed (1980) mapped numerous Oculina reefs, both
living and dead, that occur off east central Florida
in depths of 50-100 m. Our in situ observations
of C. longispinus rubscingulus with the submersibles
confirm that this subspecies is restricted to dead
reefs.

Several possibilities could explain the absence
of Centrostephanus from living reefs. Living Oculina
reefs support large schools of fish that seek shelter
within the confines of the reef matrix. Predation
pressure by these fish, as well as the suspension-
feeding coral polyps, may exclude juvenile and
larval Centrostephanus respectively. Alternatively,
the lack of suitable substratum for the settlement
and metamorphosis of urchin larvae (i.e., living
coral branches) might limit recruitment. In ad-
dition the living coral community provides less
available space for the attachment of leafy algae
and encrusting organisms, thereby limiting po-

tential food resources for the urchins; leafy algae
have not been reported from living Oculina reefs
(J. Reed, pers. comm.).

As noted by Avent et al. (1977), C. longispinus
rubicingulus usually occurs on the summits and
sides of dead Oculina reefs. Though the bases of
these reefs commonly extend to a depth of 100 m,
Centrostephanus restricts its depth to 75-80 m, in
apparent response to a distinct pycnocline and
thermocline occurring at that depth.

Poputation Densit1es.—Throughout the year,
C. longispinus rubscingulus were found in densities
up to 5 individuals per m*. Commonly co-existing
with Centrostephanus are large populations of the
cidaroid urchin, Stylocidaris affinis (Philippi),

- which achieves densities up to 80-100 individuals

per m’*.

BEHAvior.— Laboratory-maintained — speci-
mens are photo-negative. A sudden increase or
decrease of illumination evokes a rapid response
of the primary spines that normally wave or
rotate incessantly. The small claviform spines are
especially sensitive to varying light intensities.

NUMBER 20

Under intense artificial light from a fiber optics
illuminator, the claviform spines initially wave
vigorously and cease all movement after approx-
imately 10 minutes. Reduction of light intensity
induces resumption of normal movement.

Any slight mechanical stimulation also elicits
a rapid reflex action, presumably defensive in
nature. Similar reactions have been noted by von
Uexkull (1896) in C. longispinus longispinus.

FEEpInNGc.—In contrast to its congener, C. coro-
natus, which hides in crevices during the daylight
and emerges only at night to feed (Nelson and
Vance, 1979), C. longispinus rubicingulus actively
forages in daylight. The coral rubble habitat of
adult C. longispinus rubicingulus provides little re-
fuge from predators, although juveniles perhaps
receive some protection in the rubble.

During the months of June through September,
various species of leafy algae have been observed
attached to the coral rubble substratum. An ex-
amination of intestine contents of resident Cen-
trostephanus revealed the remains of several of these
species of red algae:

RHODOPHYTA

DELESSERIACEAE
Searlesia subtropica (Schneider) Schneider and Eiseman
Apoglossum ruscifolum (Turner) J. Agardh
Hypoglossum tenuifolium (Harvey) J. Agardh
Nitophyllum species

KALLYMENIACEAE
Kallymema limminghu Montagne

RHODYMENIACEAE
Leptofauchea rhodymenioides Taylor

Diet during those months in which leafy algae
is absent is unknown. However, the presence year-
round of large, successful populations of urchins
suggests that C. longispinus rubicingulus can also
exist on a varied diet of small invertebrates, en-
crusting organisms, and drift algae. Vance (1979)
reported a similar diet for C. coronatus off Catalina
Island, California.

In the laboratory, we have maintained several
specimens in closed-system aquaria for up to two
years. These specimens fed on fresh shoots of the
seagrass Thalassia testudinum. Starved specimens
have been observed to prey upon the arms of the
sea star Narcissia trigonaria Sladen.

11

SEXUAL Dimorpuism.—In specimens of Florida
C. longispinus rubicingulus, males possess conspicu-
ous tubular genital papillae while females have
short conical papillae. Through a brief survey of
museum specimens, we were able to determine
that the following species of Centrostephanus also
display this type of sexual dimorphism: C. longi-
spinus longispinus (Philippi), C. rodgersa (A. Agas-
siz), and C. coronatus (Verrill). It is probable that
all species in the genus are sexually dimorphic.
This type of sexual dimorphism is probably more
common among echinoids than formerly sup-
posed (Chia, 1977; Pawson and Miller, 1979).

ReMARKS.—In commenting on C. rubicingulus,
Mortensen (1940:310) noted the “general resem-
blance to dongispinus in nearly all characters.” He
listed several supposed differences in the inter-
ambulacral tubercle size and distribution, in the
number of wedges in primary spines, and in the
tridentate and ophicephalous pedicellariae. As
examination of our newly acquired material of C.
longispinus rubicingulus demonstrates, the only con-
sistent difference, at least in adult specimens, lies
in the one character that Mortensen (1940) dis-
missed, namely the coloration of spines.

Bernasconi (1955a,b) based her species C. bes-
nardi on two small specimens, h.d. 12 and 18 mm,
from Isla Trindade, off Brazil. In her detailed
description, Bernasconi noted that C. besnardi dif-
fers from C. longispinus in various characters of the
apical system, the rostrate pedicellariae, and the
spine coloration. In the apical system of C. bes-
nardi, ocular plates are all exsert, and the plates
are essentially naked, except for the presence of a
single secondary tubercle. According to Morten-
sen (1940:303), the position of the ocular plates,
either insert or exsert, as well as the tuberculation
of the apical system plates, can vary considerably
in C. longispinus. The rostrate pedicellariae of C.
longispinus are also variable in shape (Mortensen,
1940:305). Bernasconi’s description (1955b:59)
and figures of rostrate pedicellariae do not pro-
vide a clear indication that the rostrate pedicel-
lariae of C. besnardi are distinctive. Coloration of
the spines in C. besnard: (white and claret colored
bands) seem to differ in no important aspect from
that in juvenile C. longispinus rubicingulus. Al-

Ya SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

TasLe 4.—Measurements of Centrostephanus coronatus, from lower California (see Tables 1 and 3
for abbreviations; also, AHF = Allan Hancock Foundation)

Peristome Apical system Plates Wedges Whorls
Specimen H.D.

Dia. % A.D. Dia. % H.D. A IA A IA A IA
AHF 8.38 28 14 50 — - 14 20 21 18 18
AHF 8.43 41 20 49 10 24 14 2 21 23 15 17
AHF 6.30 46 22 48 11 24 - = 23 24 16 16
AHF 6.30 48 22 46 11 23 17 13 272 25 16 15
AHF 8.34 49 Dl 55 13 27 18 13 22 27 18 16
AHF 8.43 55 25 45 14 25 17 14 27 28 16 17
AHF 8.43 56 25 45 13 23 19 14 25 21 20 19
AHF 8.36 60 25 42 - - - = 26 22 16 16

though type specimens of this species are not
available for study, it seems evident that C. bes-
nardi falls within the range of variation of C.
longispinus rubicingulus, and the species are herein
regarded as synonymous. Fell (1975) has already
suggested that C. besnardi might be at best a
subspecies of C. longispinus.

Centrostephanus coronatus (Verrill)
Ficures 1, 2, 5; TaBies 4, 5

Echinodiadema coronata Verrill, 1867:295.

Centrostephanus coronatus.—Mortensen, 1940:314, pls. 36: figs.
7-10, pl. 76: figs. 1-4 [complete synonymy up to
1938].—H.L. Clark, 1948:237, 244, pl. 39: fig. 10.—Fell,
1975:181.—Morris, Abbott, and Haderlie, 1980:162, pl.
51: fig. 11.2a,b.

MatTeRIAL ExamMiIneD.—Allan Hancock Foun-
dation, University of Southern California: Lower
California (6.30, 8.34, 8.36, 8.38, 8.43), eight
specimens; Gulf of California (8.27, 8.28), six
specimens. National Museum of Natural History,
Smithsonian Institution: Gulf of California
(USNM 17441, USNM 32472), five specimens.

Dracnosis.—Like Centrostephanus longispinus, ex-
cept that diameter of apical system usually 24%-
34% of test diameter. Spines may virtually lack
trabeculae between wedges.

Type-Loca.iry.—Cabo San Lucas, Gulf of
California.

Distrisution.—Mortensen (1940) noted that
C. coronatus was known only from the Gulf of

California to 3 fathoms. The species also occurs
at the Galapagos Islands and in lower California
(including offshore islands) where it is common
in depths of 3-10 meters (H.L. Clark, 1948; see
Pearse, 1972).

Remarks.—H.L. Clark (1921:109) noted that
“in every way” his C. rubicingulus is nearest to C.
coronatus. Mortensen (1940:310) disagreed, and
later (p. 315, et seq.) alluded to differences in
characters of test, spines, pedicellariae, and color
that could distinguish C. coronatus from C. longi-
spinus. Our investigations of additional material
revealed the following information.

In examining the interambulacra, Mortensen
(1940:317) noted that in C. coronatus only the
uppermost interambulacral plates carry clavi-
form spines, whereas in C. longispinus “. . . the two
uppermost plates often carry a claviform spine,
and are without a primary tubercule... .” In our
material of C. longispinus, more often than not,
there is only a single adapical interamb plate
with a claviform spine. Furthermore, in C. coro-
natus, several instances occur in which the two
uppermost interamb plates carry claviform
spines. Thus, this character is not reliable.

Regarding the apical system, Mortensen (1940)
stated that in C. coronatus the apical system is
25%-27% of the horizontal diameter, while in C.
longispinus it is 35%-40%. Our material shows that
in C. coronatus the apical system is 24%-34% h.d.,
while in C. longispinus it is 31%-40% h.d. (see
Tables 1, 3, 4, and 5; Figure 1). There is a slight

NUMBER 20

13

TaBLe 5.—Measurements of Centrostephanus coronatus, from Gulf of California (see Tables 1 and
3 for abbreviations; also, USNM = United States National Museum, collections in National
Museum of Natural History, Smithsonian Institution; Mrtsn. = Mortensen collection; AHF

= Allan Hancock Foundation)

Peristome Apical system Plates Wedges Whorls
Specimen H.D

Dia. %H.D. Dia. %H.D. A IA A IA A IA
USNM 32472 13 6 46 4 31 11 16 18 - =
USNM 17441 26 12 46 8 29 15 12 18 20 27 26
Mrtsn. 1940 28 13 46 7.5 27 14-15 11-12 - - 28 25
AHF 8.27 30 16 53 - - 16 12 20 21 26 30
USNM 17441 32 16 50 11 34 17 12 17 19 25 22
USNM 17441 34 17 50 11 32 17 12 18 20 24 22
AHF 8.27 34 17 50 8 24 15 12 23 26 23 25
AHF 8.27 35 16 46 10 29 16 12 22 27 25 22
AHF 8.27 40 19 48 11 28 15 13 - - ~ =
AHF 8.27 4] 17 4] 10 24 16 13 24 25 22 21
AHF 8.28 43 19 44 10 23 18 13 23 25 23 24
USNM 17441 50 - - - - - - 22 24 - =
Mrtsn. 1940 55 27 49 15 al) 18-19 15-16 - - - =
Mrtsn. 1940 62 - - - - - - - - - -

overlap in these proportions, but apical system
sizes do appear to differ consistently in the two
“species.”

Supposed differences in spine coloration, noted
by Mortensen (1940), are eliminated by our new
Florida material. In large specimens of C. corona-
tus, the spines are nearly black and the same is
true for C. longispinus rubicingulus, at least in East
Florida populations. One usually consistent dif-
ference between the two species lies in the trabe-
cules between the spokes of the spines; in C.
longispinus trabecules are conspicuous, while in C.
coronatus they are virtually non-existent (Morten-
sen, 1940).

The number of verticillations (whorls of
spinelets) on the spines provide some evidence
that there may be two distinct subspecies of C.
coronatus (see Tables 4 and 5; Figure 2). In Gulf of
California specimens, the number of verticiila-
tions in ambulacral and interambulacral spines

usually exceeds 20 per cm of spine length; in
California specimens examined, there are usually
fewer than 20 verticillations per cm of spine
length. A more detailed survey of California spec-
imens from several habitats is required before the
status of the California and Gulf of California
populations is settled. In C. longispinus rubicingulus
spines usually have fewer than 20 whorls of spi-
nelets per cm of spine length, and thus in this
character the western Atlantic Centrostephanus
most closely resembles the California coast pop-
ulations of C. coronatus.

The relatively trivial differences between C.
coronatus and C. longispinus might seem to justify
uniting them under a single species name. We
hesitate to take this step because it would appear
to serve no useful purpose. It is important to note,
however, that these species are indeed closely
related, contrary to Mortensen’s (1940) conten-
tions.

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wh

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contents, (4) foreword and/or preface, (5) text, (6) appendixes,
(7) notes section, (8) glossary, (9) bibliography, (10) legends,
(11) tables. Index copy may be submitted at page proof stage,
but plans for an index should be indicated when manuscript is
submitted.

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