The Echinoids of
Carrie Bow Cay, Belize
PORTER M. KIER
m
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ? NUMBER 206
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S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y ? N U M B E R 2 0 6
The Echinoids of
Carrie Bow Cay, Belize
Porter M. Kier
SMITHSONIAN INSTITUTION PRESS
City of Washington
1975
ABSTRACT
Kier, Porter M. The Echinoids of Carrie Bow Cay, Belize. Smithsonian Con-
tributions to Zoology, number 206, 45 pages, 8 figures, 12 plates, 1975.?Twenty
species were found living off Carrie Bow Cay. Echinometra lucunter (Linnaeus),
E. viridis Agassiz, and Diadema antillarum Philippi are the most common species
on the reef. In the lagoon, the most common in the Thalassia beds are Lytechinus
variegatus (Lamarck), Eucidaris tribuloides (Lamarck), and Clypeaster rosaceus
(Linnaeus). The deeper mud fields have large numbers of the spatangoids Paraster
doederleini Chesher, fewer Moira atropos (Lamarck), and Bnssopsis elongata
Mortensen. Meoma ventricosa (Lamarck) occurs in sand fields both in the lagoon
and on the reef. Cassidulus cariboearum Lamarck lives in the lagoon's coarse sand
in water one meter deep. This species has never been seen alive in situ before.
Immature specimens were found of Plagiobrissus grandis (Gmelin) and Meoma
ventricosa (Lamarck). These are the smallest specimens ever described of these
species. Their growth changes are described. Rod-like structures are reported in
P. grandis, which probably serve to strengthen its very thin test.
OFFICIAL ITBI.ICATION DATE is handstampcd in a limited number of initial copies and is recorded
in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 5261. SERIES COVER
DESIGN: The coral Montastrea cavernosa (Linnaeus).
Library of Congress Cataloging in Publication Data
Kier. Porter M.
The echinoids of Carrie Bow Cay, Belize.
(Smithsonian contributions to zoology, no. 206)
Supt. of Docs, no.: SI 1.27:206
1. Sea-urchins?British Honduras?Carrie Bow Cay. I. Title. II. Series: Smithsonian Institution,
Smithsonian contributions to zoology, no. 206.
QL1.554 no. 206 [QL384.E2] 591'.08s [563'.95] 74-23845
For sale by the Superintendent of Documents, U.S. Government Printing Office
Washington, D.C. 20402?Price $0.00 (paper cover)
Contents
Page
Introduction 1
Plagiobrissus grandis (Gmelin) 3
Meoma ventricosa (Lamarck) 7
Paraster doederleini Chesher 9
Paraster cf. Paraster floridiensis (Kier and Grant) 15
Moira atropos (Lamarck) 15
Brissopsis elongata Mortensen 16
Agassizia excentrica Agassiz 16
Cassidulus cariboearum Lamarck 16
Eucidaris tribuloides (Lamarck) 16
Diadema antillarum Philippi 17
Arbacia punctulata (Lamarck) 17
Echinometra viridis Agassiz 17
Echinometra lucunter (Linnaeus) 17
Lytechinus variegatus (Lamarck) 17
Lytechinus williamsi Chesher 17
Tripnenstes ventricosus (Lamarck) 18
Clypeaster rosaceus (Linnaeus) 18
Clypeaster subdepressus (Gray) 18
Leodia sexiesperforata (Leske) 18
Echinoneus cyclostomus Leske 18
Literature Cited 18
Plates 21

The Echinoids of
Carrie Bow Cay, Belize
Porter M. Kier
Introduction
Scientists from the Smithsonian Institution are
making an ecosystem study of Carrie Bow Cay, a
small island located on the barrier reef that ex-
tends along the coast of Belize (formerly British
Honduras). This study will be an integrated pro-
gram emphasizing the biological and geological
interactions of the system. Initially the diversity
and distribution of the benthic organisms will be
studied, as well as their role in the construction
and destruction of the reef. Specialists in each
animal and plant group will first describe the fauna
and flora. They will then try to formulate the
interaction between the individual groups and in
turn their involvement in the development of the
reef itself.
This report on the echinoids follows one month
of diving and collecting in 1973 and 1974. Most
of the dives were made from the reef directly east
of Carrie Bow Cay and, in particular, from a tran-
sect that will be studied by all participants in the
program (Figure 1). The transect extends 600
meters from the Cay to a point over the reef drop-
off. Many dives were also made in the lagoon west
of Carrie Bow. Twenty species of echinoids were
found. None of them are new, but several of them
have never been seen alive before in their natural
habitat. Of particular interest was the discovery
of extremely small specimens of two spatangoids,
Plagiobrissus grandis (Gmelin) and Meoma ventri-
Porter M. Kier, Department of Paleobiology, National
Museum of Natural History, Smithsonian Institution, Wash-
ington, D.C. 20560.
cosa (Lamarck). They reveal new information
about the development of these echinoids, such as
the presence of rod-like structures in P. grandis,
which probably serve to reinforce the very thin
test.
The distribution of the echinoids is similar to
that described for other sites in the Caribbean.
Echinometra lucunter (Linnaeus), E. viridis Agassiz,
and Diadema antillarum Philippi are the most
common species occurring on the reef, although D.
antillarum is not as numerous as elsewhere in the
Caribbean. In the lagoon's Thalassia beds, Lytech-
inus variegatus (Lamarck), Eucidaris tribuloides
(Lamarck), and Clypeaster rosaceus (Linnaeus) are
the most common species occurring together with
herds of Diadema antillarum. Spatangoids occur in
great numbers in the mud fields of the lagoon,
including many Paraster doederleini Chesher,
fewer Moira atropos (Lamarck), and Brissopsis
elongata Mortensen.
An interesting feature of the echinoid fauna is
the presence of only a few individuals of a large
number of species. Only one dead specimen was
found of Clypeaster subdepressus (Gray), one live
specimen of Leodia sexiesperforata (Leske), no
adults of Plagiobrissus grandis, no living specimens
of Agassizia excentrica Agassiz, two specimens of
Paraster cf. P. floridiensis (Kier and Grant), a few
Cassidulus cariboearum Lamarck, and two speci-
mens of Arbacia punctulata (Lamarck). Although
only one specimen of Lytechinus williamsi Chesher
was found, it may be common. I simply was not
looking for it.
ACKNOWLEDGMENTS.?I thank William M. Kier,
who served as my diving partner and did all the
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Location of
research area
KILOMETERS
Water Cay Range
HONDURAS
South Water Cay
LA GOON
TRANSECT
V Carrie Bow Cay
(I6?48'N 88?O5'W)
FIGURE 1.?Map showing location of Carrie Bow Cay and location of transect.
NUMBER 206
underwater photography. Many of the specimens
described in this paper were collected by him.
Thomas R. Waller, Frederick Hotchkiss, and Doug-
las Putman also collected some of the echinoids.
Mary H. Lawson did the laboratory preparations
and photography. The SEM photography was done
by Walter R. Brown, and the map and graphs
were drawn by Lawrence B. Isham. David L. Pawson
and Richard H. Chesher critically read the manu-
script and made many useful suggestions for its
improvement.
This paper is contribution number 15 of the
Smithsonian Institution Investigations of Marine
Shallow-Water Ecosystems Project.
Plagiobrissus grandis (Gmelin)
FIGURES 2-5; PLATES 1-3; PLATE 4: FIGURES 1, 2
Echinus grandis Gmelin, 1791:3200. [For synonymy, see
Mortensen, 1951:496.]
Immature specimens of Plagiobrissus grandis
(Gmelin) and Meoma ventricosa (Lamarck) were
found living buried in the sand at a depth of 50
to 100 mm, in sand fields in the grooves of the
Spur and Groove Zone on the transect off Carrie
Bow Cay at a depth of 10 m.
The immature of the two species resemble each
other in general appearance (compare Plate 1:
figure 1 with Plate 6: figure 1); but the young of
Plagiobrissus grandis can be distinguished by their
whiter spines and lack of large, black pedicellariae
on their dorsal surface. If all specimens found had
been less than 6 mm, it would have been difficult
to identify these species; but specimens nearly 20
mm long are enough like the adults to make identi-
fication possible. These identifications were con-
firmed by the comparison of specific features in the
tests of both immature and adult that are not
affected by growth. Chesher (1968a) has shown that
the fascioles remain on the same plates throughout
the growth of the echinoid, that the number of
plates beyond the petals are constant, and that the
periproct remains enclosed by the same plates.
These features were compared between the imma-
ture specimens and specimens of adults believed
to be the same species and no differences were
found.
No adult specimens of either species were living
with these immature. Considering the great differ-
ences between the immature and the adult, it is
not surprising that they live in different environ-
ments.
Three immature specimens were found of P.
grandis varying in length from 5.8 to 20 mm. The
smallest specimen is far smaller than any of this
species previously described (Kier and Grant, 1965:
36, described a specimen 35 mm long) and shows
features never seen before. These features and
changes that occur in the larger specimens are
described in detail below.
SHAPE.?The smaller specimens have a higher,
B
FIGURE 2-?Plagiobrissus grandis (Gmelin). Side views showing
the decrease in relative height of the test in the larger
specimens; A, specimen USNM E137SO, 6.9 mm long, X 10;
B, specimen USNM E10304, 35 mm long, X 2; c, specimen
USNM El3737, 150 mm long, X \/t. The two larger speci-
mens (B and c) are from the Florida Keys.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
NUMBER 206
more smoothly rounded test with a far larger
periproct and peristome relative to the test's total
length. The specimen 5.8 mm long has a very high
test?a height equal to 67 percent of the length.
In a specimen 35 mm long, the height is 45 percent
of the length; and in an adult 150 mm long, only
33 percent (Figure 2). The test is smoothly rounded
in the smallest specimen; but in an adult the margin
is sharp, the adoral surface is more flattened, and
the dorsal interambulacra are slightly ridged.
The periproct in the smallest specimen is very
large, with its height equal to 38 percent of the
length of the test and its width equal to 24 percent.
In contrast, the periproct of a specimen 35 mm
long has a height of only 18 percent and a width
of only 10 percent. Finally, in an adult 150 mm
long, the height of the periproct is only 10 percent;
and the width is only 7 percent of the length of the
test.
The periproct remains surrounded by the same
interambulacral plates throughout its growth. The
surrounding plates just become fewer in the adult.
The smallest specimen's periproct occurs within
interambulacral plates 5 through 10. An adult's
falls within plates 5 through 8. The anus is situated
so far dorsally in the smaller specimens (Figure 4A;
Plate 1: figure 1) that it is almost entirely visible
when viewed from the top. In an adult (Plate 1:
figure 2) it has shifted down to a posterior trunca-
tion where most of it is now visible from a ventral
view.
FIGURE 3.?Plagiobrissus grandis (Gmelin): A, petaloid area of
specimen 5.8 mm long showing single pores in the paired
petals, and medially constricted pores in ambulacrum in,
USNM E13731, X 22; B, ambulacrum HI of same specimen un-
der greater magnification showing nature of constricted
pores suggesting that petaloid pores are originally single
and then are divided in two, X 100; c, petaloid area of
specimen 6.9 mm long, USNM E1373O, showing presence of
pore-pairs in all petals, X 22; D, apical system in smallest
specimen 5.8 mm long, USNM El3731, has no genital pores
and only a single madreporic pore, X 40; E, specimen 20 mm
long, USNM E13732, has small genital pores, and 16 madre-
poric pores, x 25; F, specimen 150 mm long, USNM E13737
(from the Florida Keys) has greatly enlarged genital pores,
hundreds of madreporic pores, and genital 2 extends pos-
teriorly, X 6; G-I, the peristome in the smaller specimens is
much larger, and the plastron wider (G, specimen 6.9 mm
long, USNM E13730, X 10; H, specimen 20 mm long, USNM
E13732, X 3.5; i, specimen 150 mm long (from Florida
Keys), USNM E13737, X i/2.
B
FIGURE 4.?Plagiobrissus grandis (Gmelin). A specimen 5.8 mm
long (A) differs from one 20 mm long (B) in lacking pore-pairs
in the petals and in having its periproct on an adapical
surface: A, USNM E13731, X 14; B, USNM E13732, X 5.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
The peristome, too, is much larger (Figure 3c-i;
Plate 1: figures 3, 4) relative to the length of the
test in the very young. The width of the peristome
in the smallest specimen is 34 percent of the length
of the test; in a specimen 20 mm long, only 13
percent. A decrease also occurs in the height of the
peristome. The smallest specimen, 5.8 mm long,
has a height equal to 22 percent of the length of
the test; whereas, the peristome in a specimen 20
mm long is only 11 percent of the total length, and
is only 4 percent in an adult 150 mm long. Part
of this reduction in the height of the peristome
is due to the development of a protruding labrum
in the adult.
PLASTRON.?The plastron on the smaller speci-
mens is far wider than that in the adult. A specimen
6.9 mm long has a plastron with a width equal to
43 percent of the length of the test (Figure 3c);
whereas, a specimen 20 mm long has a plastron
width of only 30 percent (Figure 3H). In an adult
the width is only 20 percent (Figure 3i).
The plastron in the immature specimens bears
many large tubercles, but in the adult the tubercles
are smaller than those on the broad interambulacral
areas. Presumably, the spines attached to the plas-
tron in the immature (Plate 3: figure 2) are the
primary means of locomotion and burrowing. Dur-
ing growth a change takes place, and these locomo-
tive spines in the adult are attached to the in-
ter ambulacra.
APICAL SYSTEM.?The apical system of an im-
mature specimen differs from an adult in being
much larger relative to the size of the test, lacking
genital pores, having only one madreporic pore,
and lacking an extension of the madreporite (geni-
tal 2) into the posterior interambulacrum.
The apical system in the smallest specimen (5.8
mm long) is 0.99 mm wide (measured at its greatest
width across oculars n and iv). This is 17 percent
of the length of the test. The next largest speci-
men (6.9 mm long) has an apical system 0.98 mm
wide or 14 percent of its length. A specimen 20
mm long has a system 1.2 mm wide or 6 percent
of its length. An adult 150 mm long has a system
4.7 mm wide, which is only 3.1 percent of its length.
Although the apical system does increase in size
during the growth of the animal, its rate of growth
is much less than the growth of the test.
No genital pores are present in the two smallest
specimens, 5.8 and 6.9 mm long (Figure 3cp); but
small pores occur in a specimen 20 mm long (Figure
3E). They are quite large in a specimen 35 mm
long; in an adult 150 mm long, they are extremely
large, occupying most of the genital plates exclud-
ing the madreporite (Figure 3F).
Only one madreporic pore is present in each of
the two smaller specimens (Figure 3C,D), and this
occurs in the right anterior portion of genital 2.
Sixteen pores are present in a specimen 20 mm long
(Figure 3E); 35 in a specimen 35 mm long, and
hundreds in an adult 150 mm long (Figure 3F).
Although the single pore in the two smaller speci-
mens is large relative to the size of the test, the
madreporic pores in all the specimens are approxi-
mately the same size, 0.03 to 0.04 mm in diameter.
The madreporite (genital 2) extends very far
into the posterior interambulacrum in the adult
specimen (Figure 3F), but in the smaller specimens
(Figure 3D,E) this posterior extension does not
occur.
PETALS.?The ambulacral plates of the petals
are just beginning to be developed in the smallest
specimen 5.8 mm long. These petaloid plates can
be distinguished from the ambulacral plates be-
yond the petals by their smaller size and by their
slightly larger pores (Plate 1: figure 1). The pores
in the ambulacral plates beyond the petals are so
small that they cannot be distinguished from the
holes in the meshwork of the calcite plates. Each
of the petaloid ambulacra have six to eight plates.
The pore in each plate is single, not double as in
an adult. Each of these pores was studied under
high magnification (Plate 4: figures 1, 2) with a
scanning electron microscope, and in none of them
is there any indication of a partition developing
to divide the pore in two. However, in the non-
petaloid ambulacrum m, each pore is constricted
medially (Figure 3B; Plate 4: figure 1), suggesting
that the ambulacral pores are originally single and
then are divided in two by this constriction.
In the next larger specimen, 6.9 mm long, most
of the petaloid pores are double (Figure 3c). The
pores, though, are still very close together; and
the ambulacra are not petal-like in appearance.
Only eight plates are present in each petaloid
ambulacra. A specimen 20 mm long has well-
developed petals with the pores elongated trans-
versely.
The petals are slightly depressed, not flush to the
general surface of the test as in the smaller sped-
NUMBER 206
mens. Each anterior petal has 26 plates, and each
posterior has 32. The petals in an adult 150 mm
long (Plate 1: figure 2) are flexuous, not straight as
in the smaller specimens. There are 70 plates in
each anterior paired petal and 90 in each posterior.
FASCIOLES.?The peripetalous fasciole is only
slightly developed in the immature specimens, but
the subanal fasciole is distinct and clearly func-
tional. The fascioles increase in size, not by an
increase in the size of the fasciole tubercles but by
an increase in the number of these tubercles.
The peripetalous fasciole on a specimen 5.8 mm
long is composed of a single row of tubercles. A
second row of tubercles is present for part of the
length of this fasciole in a specimen 6.9 mm long
(Plate 1: figure 1). Most of this fasciole has 2 rows
of tubercles in a specimen 20 mm long; 5 rows in a
specimen 35 mm long; and 17 rows in an adult 150
mm long.
The subanal fasciole is very distinct in the small-
est specimen (5.8 mm long) and consists of 2 to 3
rows of tubercles. There are 3 to 5 rows in a speci-
men 20 mm long; 9 to 13 in a specimen 35 mm
long; and 30 to 35 in an adult 150 mm long.
The tubercles in both fascioles do not increase
in size during the growth of the echinoid. They
are between 0.05 and 0.06 mm in diameter in all
the specimens regardless of the size of the echinoid.
FIGURE 5.?Plagiobrissus grandis (Gmclin). Adoral view of
specimen USNM E13732, 20 mm long, showing the rod-like
structures that are herein considered to be for reinforce-
ment, x 3.5.
DORSAL TUBERCLES.?The large dorsal tubercles
that typically occur within the peripetalous fasciole
of an adult are absent or just forming in the small-
est specimen. The plates that bear them in the
adult are just beginning to be developed in the
smaller specimens. A specimen 20 mm long has
approximately 20 of these larger tubercles. A speci-
men 35 mm long has less than 50, and an adult
has nearly 200.
VENTRAL RODS.?Rod-like structures (Figure 5)
are very apparent on the ventral side of the smaller
specimens, particularly when the specimens are
wettened with xylene (Plate 2: figure 2). Under high
magnification it can be seen that the rods are
formed of the calcite matrix of the test and are
produced by thickening of the calcite with a cor-
responding reduction in the size of the holes in the
calcite meshwork (Plate 2: figures 3, 6). Where a
rod meets the suture of an adjacent plate, it is
overlapped by a rod extending from that plate
(Plate 2: figures 4, 5). The rods are very conspicuous
on the smaller specimens because of their thin tests,
but much less apparent on an adult.
I suspect that these rods serve to strengthen the
test, which is very thin in this species, especially in
the immature. The rods are most developed where
large tubercles occur, suggesting that they are espe-
cially needed here because of the strain exerted on
the test by the strenuous movement of the ventral
spines. The rods are not present in the immature
of Meoma ventricosa (described below), but the
test in that species is considerably thicker and
perhaps needs no such reinforcement.
ADULTS.?Although I found no adults in the
region, Richard Chesher (pers. comm., 1974) points
out that, in his experience, they are one of the most
difficult echinoids to find because they bury very
deep, have no surface burrow, and do not come
up at night.
Meoma ventricosa (Lamarck)
FIGURE 6; PLATE 4: FIGURES 3, 4; PLATE 5; PLATE 6:
FIGURES 1-3
Spatangus ventricosus Lamarck, 1816:323. [For synonymy, see
Mortenscn, 1951:529, and Chesher, 1970:737. The biology
and living habits have been described by Kier and Grant,
1965:38, and Chesher, 1969, 1970.]
This species occurs off Carrie Bow Cay in sand
grooves in the reef and in the lagoon west of the
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
B
NUMBER 206
island. Its distribution is sparse in the reef where
only three specimens were found alive. It is quite
common in the deeper water (over 7 m) in the sand
fields of the lagoon where specimens reach their
greatest size. Here the specimens live completely
buried in the sediment. Although it occurs in
Thalassia beds where the water is over 5 m deep,
specimens are less common and smaller. Individuals
are not completely buried in the sediment, pre-
sumably because the network of Thalassia roots
prevents burial. Here the echinoid lives much like
Clypeaster rosaceus.
Immature specimens were found along the tran-
sect in the Spur and Groove Zone in sand fields
at a depth of 10 meters. The echinoids were buried
in the sand, and were only 5.8 to 17.5 mm long.
They occur with the young of Plagiobrissus grandis
and can be distinguished from them by their black
pedicellariae, darker color due to yellow tips on
their spines, and red tubefeet in the anterior
ambulacrum. No adults were found in this zone.
The changes from the growth of a specimen 7
mm long to adult dimensions have been described
by Kier and Grant (1965) and Chesher (1969, 1970).
Only the changes from the smallest specimen 5.8
mm long to one 17.5 mm long are described here.
APICAL SYSTEM.?The apical system is very large
relative to the size of the test in the immature. The
specimen 5.8 mm long has an apical system (Figure
6A; Plate 6: figure 1) with a width equal to 16
percent of the length of the test. In the specimen
17.5 mm long, the width of the apical system is
only 9 percent of the length of the test. No genital
pores are present in either of these specimens. Kier
and Grant (1965:44) report small genital pores in a
specimen of this species 44 mm long. Only a single
madreporic pore is present in the specimen 5.8
mm long, but 27 occur in the specimen 17.5 mm
long.
PETALS.?The petals are just beginning to be
developed in the specimen 5.8 mm long (Figure
FIGURE 6.?Meoma ventricosa (Lamarck): A, B, adapical and
adoral views of the smallest specimen (5.8 mm long) showing
features of the immature test such as the lack of pore-pairs
in the petal, the absence of genital pores, the presence of a
single madreporic pore, adapical position of the periproct
and relatively large peristome, USNM E13733, X 13; c, D,
adapical and adoral views of larger specimen 17.5 mm long,
USNM E13734, x 6.
6A; Plate 6: figure 1). Each petal has only six to
eight pores. All appear to be single, although some
of them are constricted medially. This indicates
that their bifurcation into pore-pairs would have
occurred soon. The specimen 17.5 mm long (Figure
6c; Plate 6: figure 2) has well-developed petals
with 35 to 36 pore-pairs in the anterior petals n
or iv. Thirty-one to 33 pore-pairs compose posterior
petals v or i.
PLASTRON.?Although the plastron is relatively
wider in the immature than in the adult, this dif-
ference is not nearly as great as in Plagiobrissus
grandis. The smallest specimen, 5.8 mm long, has
a plastron (Figure 6B) whose width is equal to 45
percent of the length of the test. The width of the
plastron in the specimen 17.5 mm long (Figure 6D)
is 40 percent of the length, and in an adult it is
35 percent.
FASCIOLES.?The peripetalous fasciole is very
slightly developed in the smallest specimen (5.8
mm long) and is composed of a discontinuous single
row of small tubercles (Plate 6: figure 1). This
fasciole is well developed in the specimen 17.5 mm
long and is composed of one to three rows of
tubercles (Plate 6: figure 2).
The subanal fasciole is distinct in the smallest
specimen and is composed of two rows of tubercles.
The specimen 17.5 mm long has up to nine rows
in this fasciole.
PEDICELLARIAE.?Approximately 10 globiferous
pedicellariae (Plate 5) were present on the dorsal
side of the specimens 5.8 mm and 17.5 mm long.
They are very conspicuous on the smallest speci-
men because of their black color and easily dis-
tinguish the young of this species from the young
of Plagiobrissus grandis. Although Mortensen
(1951:531) reports no globiferous pedicellariae in
an adult of Meoma ventricosa, Chesher (1969:99)
has found them in an adult specimen from Ber-
muda.
Paraster doederleini Chesher
FICURES 7, 8; PLATE 7; PLATE 8: FIGURES 1-5
Paraster doederleini Chesher, 1972a: 10-25, figs. 1-9.
The Carrie Bow specimens are indistinguishable
from the Floridian holotype of this species that
previously was reported only from Florida and
Colombia. Chesher (1972a:21) noted that the Co-
U- UJ
GROUP I
GROUP 31
X British Hon
O Holotyp*
? Porotyp??
. ? . . . ! ! i i I I ? ? I I ? I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I
it n M m M ii it n si HI
LENGTH OF TEST (MM)
UJ
I
UJ
o
UJ
Q.
in
<
OL
<r
GROUP I
x ._
GROUP 31
X British Honduras sp*clmsna
O Holotyp*
? Porotypm
CO ' ? i i i . i i i i i i i i i . i . i i i i . . i i i i
41 SI II
LENGTH OFTEST(MM)
91 H I
FIGURE 7.?The type-specimens of Paraster doederleini Chesher can be divided into two groups:
one having curved posterior petals (group n) and the other having straight posterior petals
(group i). As shown in the above scattergrams the group i specimens, including the specimens
of Carrie Bow Cay, have longer anterior petals than the specimens of group II.
NUMBER 206 11
TABLE 1.?A comparison of the dimensions of the two "groups" of specimens
of Paraster doederleini Chesher
CHARACTER
AH (height periproct)
AP (length anterior petal)
APE (span anterior petal)
APW (width anterior petal)
FF (apical to fasciole along
ambulacra III)
AW (width periproct)
LF (apical to fasciole along
interambulacra 4)
LP (labrum to posterior test)
NW (width anterior notch)
PA (peristome to anterior test)
PP (length posterior petal)
PPE (span posterior petal)
PPFL (length periproct fasciole)
PPW (width posterior petal)
TW (width test)
Will (width ambulacra III)
Group
14.9
35.2
58.2
11.8
52.7
9.7
16.7
76.3
4.6
20.32
19.9
16.5
26.1
203.4
9.0
96.3
15.6
MEAN PERCENT OF
I (Min-Max)
(11.3-18.6)
(33.6-37.8)
(54.8-62.2)
(09.5-14.0)
(48.0-55.9)
(07.5-11.8)
(14.1-18.5)
(71.4-82.0)
(03.4-15.6)
(15.2-25.1)
(15.2-25.1)
(13.7-19.5)
(22.8-28.1)
(186.0-215.1)
(06.9-10.6)
(92.5-99.0)
(11.2-22.3)
LENGTH
Group
12.3
24.8
39.8
9.1
44.2
8.2
11.5
81.3
2.7
18.2
17.5
13.6
17.9
159.5
6.95
94.1
9.4
II (Min-Max)
(10.5-13.8)
(21.2-28.5)
(35.7-44.1)
(07.8-11.1)
(41.6-46.0)
(07.1-09.2)
(10.7-13.1)
(79.1-83.3)
(01.8-03.7)
(17.0-18.1)
(16.8-18.1)
(12.1-15.0)
(16.1-21.1)
(140.9-177.4)
(06.0-07.7)
(92.2-95.8)
(07.4-12.8)
T-TEST OF SIGNIFICANCE
OF DIFFERENCE
03.5-.010
09.7-.001*
10.5-.001*
04.3-.001
08.1-.001
03.2-.010
12.0-.001
03.2-.010
02.8-.010*
03.9-.001*
05.1-.001*
05.3-.001
09.6-.001
07.0-.001*
05.5-.001
02.3-.050
06.1-.001
Group I ? Carrie Bow Cay specimens (25) plus Chesher's E11376, 8397C, and 8397D. All these specimens have straight
posterior petals.
Group II -- Chesher's E11378, E11377, 8398, 8396B+, 8396B, and 8396A. All these specimens have curved posterior petals.
?Because of few specimens in Group II, variance so high as to indicate in f-test that
t-test may not be valid. However, there is no overlap between means in Group I and II
(min-max) and differences appear to be very significant ? see scattergrams for AP and APE.
lombian specimens were slightly different from the
Floridian specimens and postulated that the "dif-
ferences between the two populations are not
significant and will probably prove to be part of
a morphological cline when more specimens are
found." However, two of Chesher's Floridian para-
types are similar to the Colombian specimens. This
suggests that a morphological cline based on geo-
graphic separation probably does not occur between
the Floridian and Colombian populations. The
Carrie Bow Cay specimens are indistinguishable
from the holotype and two of the paratypes but
readily differentiated both statistically and by visual
inspection from the Colombian specimens and two
of the Floridian paratypes. The fact that no in-
termediates between these two groups are present
either among Chesher's types or among the Carrie
Bow Cay specimens and that the two have never
been found living together suggests that these
groups should probably be differentiated taxonom-
ically. However, until we know more of the geo-
graphic distribution of the two groups and the
variation within the "Colombian population" it
seems best to postpone any taxonomic decision.
The two groups are easily distinguished visually
by the straight posterior petals and longer anterior
petals found in the Carrie Bow Cay specimens, the
holotype, and paratypes USNM El 1376, MCZ
8397c, and MCZ 8397d (these specimens henceforth
are referred to as group i). The specimens in group
TABLE 2.?The dimensions of specimens of Paraster floridiensis (Kiev and Grant),
P. cf. P. floridiensis, and P. doederleini Chesher (all statistics except the total
length are the percent of the total length)
CHARACTER
TL (length)
TW (width)
H (height)
Will (width amb. Ill)
AP (length ant. petal)
APE (span ant. petals)
APW (width ant. petal)
PP (length post, petal)
PPE (span post, petals)
PPW (width post, petal)
AX (apical syst. to
ant. end)
PG (post genital pores)
ND (notch depth)
NW (notch width)
PPFL (length fasciole)
FF (apical syst. to fasc
along amb. Ill)
LF (apical syst. to fasc
along inter. 4)
PF (apical syst. to fasc
along inter. 5)
LAF (latero-anal fasc.)
AH (anus height)
AW (anus width)
PL (plastron length)
B (plastron width)
C (plastron width)
D (plastron width)
L (labrum length)
LP (ant. labrum to
post, test)
PW (peristome width)
PH (peristome height)
PA (ant. peristome to
ant. test)
APW/AP
PPW/PP
PP/AP
AW/ AH
D/PL
Paraster cf. P.
USNM E13741
X6 ? 8 nun
86.90
75.00
14.88
39.88
62.50
13.09
20.83
29.16
10.11
58.33
-
1.78
14.88
250.00
64.88
20.23
22.02
136.90
20.83
16.66
57.14
30.95
36.90
39.88
9.52
74.44
-
-
21.42
32.82
48.57
52.23
79.98
69.79
floridiensis
USNM E13742
12.8 mm
88.52
71.88
16.64
34.61
57.27
11.41
17.42
27.81
8.83
55.94
6.25
-
21.33
228.91
54.14
16.41
18.98
145.31
19.22
15.31
57.81
34.06
42.66
42.19
7.81
72.89
25.00
9.61
18.44
32.90
50.60
50.33
79.66
72.98
P. floridiensis
Chesher (1972)
11.60 mean
92.66
75.24
12.35
38.52
66.19
-
17-24
30.19
-
53.61
-
-
-
253.72
55.41
19.21
19.35
152.46
20-28
-
56.61
25.00
35.00
41.27
6.48
64-75
-
-
21.00
35.82
48.75
50.59
79.07
72.88
Paraster floridiensis
MCZ 8494
15.87 mean
95.59
77.95
14.68
40.14
65.34
12.60
19.09
28.36
9.45
56.27
2.52(?)
-
-
266.54
59.42
17.64
19.72
165.72
21.87
15.75
56.52
-
-
-
-
labrum missing
24.13
-
24.58
31.39
49.50
47.56
72.02
-
MCZ 8495
16.03 mean
91.08
73.61
12.66
39.74
67.19
13.10
19.78
31.19
10.42
57.39
1.56
-
-
238.93
59.45
18.72
21.21
128.95
22.46
18.72
59.45
-
-
-
6.05
72.55
26.64
9.36
23.52
32.96
52.68
49.77
83.35
-
P. doederleini
(holotype)
63.43
96.58
70.73
11.19
33.84
55.75
9.83
17.08
26.54
7.88
49.92
3.63
4.05
20.34
198.63
48.24
17.34
18.29
179.19
13.24
7.51
50.18
25.85
35.47
38.41
9.25
82.03
17.75
4.26
15.24
29.05
46.14
50.47
56.72
76.54
NUMBER 206 13
II (the Colombian paratypes and the Floridian para-
types, USNM El 1378 and MCZ 8398) have flexuous
posterior petals and much shorter anterior petals.
These differences are readily apparent when one
compares the photographs of specimens equivalent
in size in both groups on Plate 7: figures 3 (group
I) and 6 (group II).
Statistically, specimens in group i differ from
group II in having longer and wider anterior and
posterior paired petals with a greater span and
more pore-pairs. They have a wider ambulacrum m,
larger periproct, and a greater distance between
the apical system and the peripetalous fasciole as
measured along the center of ambulacrum in and
interambulacrum 4. The peristome is more pos-
terior; the anterior notch, deeper and wider; the
peripetalous fasciole, longer; and the test, slightly
wider. The statistics of these characters for the
two groups together with the results of t-tests of
the significance of the differences are presented on
Table 1. The difference between the two groups
in the span and length of the anterior petals is
shown graphically in Figure 7.
Although the groups are easily differentiated,
the taxonomic separation is probably not as great
as might seem. Specimens of both groups have
their fascioles crossing the same plates in each
ambulacrum and interambulacrum, a feature which
Chesher has proved to be of great significance.
Because Chesher's description and statistics are
based on specimens of both groups, I am including
a description and statistics of 25 specimens from a
single population from Carrie Bow Cay and the
measurements (Table 2) of the holotype.
MATERIAL.?Statistics from 25 specimens col-
lected at same locality.
SIZE AND SHAPE.?Specimens 14.7-51.9 mm long,
mean 35.49 mm, width 96.47 percent TL (SD 1.72,
CV 1.8), height 73.78 percent TL (SD 3.10, CV
4.2); posterior truncated obliquely, dorsally over-
hanging so that periproct visible adorally; greatest
width of test anterior at anterior series of inter-
ambulacra 1 and 4; greatest height posterior at
interambulacrum 5 between posterior petals.
APICAL SYSTEM.?Posterior of center, distance
from anterior margin 52.86 percent of TL (SD
2.39, CV 4.5); ethmolytic, tetrabasal (Figure 8B)
with four genital pores, anterior pair about one-
third diameter of posterior pair; posterior pair
present on all specimens including smallest 14.7
mm long, anterior pair absent on smaller speci-
mens 14.7 and 17.7 mm long, present on next larger
specimen 20.2 mm long and all other specimens;
14 specimens with plate sutures of apical system
visible, 13 tetrabasal, one specimen tribasal with
genital 1 missing (Figure 8B), 10 specimens with
genital 2 extending posteriorly joining ocular v
and separating genital 4 from genital 2 (Figure 8B),
one specimen with genital 3 extending posteriorly
into interambulacrum 5 instead of genital 2.
AMBULACRA.?Anterior ambulacrum in in sunken
furrow extending from apical system to peristome
FIGURE 8.?Paraster doederleini Chesher: A, adoral view
showing plate arrangement of USNM E13735, from Carrie
Bow Cay, x 5; B, apical system of same specimen, X 14.
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
forming pronounced notch in front of test with
depth 4.61 percent TL (SD 2.36, CV 51.1), width
20.23 TL (SD 2.15, CV 10.6), slightly sunken at
edge of peristome; pore-pairs in single series, not
conjugate, pronounced node separating pores of
pair; pores situated parallel to plates but due to
curvature of plates outer pore of pair slightly pos-
terior to inner; greatest width of ambulacrum in
14.92 percent TL (SD 2.22, CV 13.9); peripetalous
fasciole crosses ambulacrum m at distance from
center of apical system equal to 52.96 percent TL
(SD 1.64, CV 3.1); ambulacral plates beyond peri-
petalous fasciole with single pore in each plate.
Anterior petals (II and iv) slightly flexuous, length
35.36 percent TL (SD 1.03, CV 2.9), greatest width
12.01 percent TL (SD 0.98, CV 8.1), ends of an-
terior petals separated apart distance equal to 58.34
percent TL (SD 1.74, CV 3.0). Posterior petals
(v, L) straight, short, length 16.60 percent TL (SD
1.58, CV 9.5), greatest width 9.19 percent TL (SD
0.61, CV 6.6), ends of petals separated by a distance
equal to 26.30 percent TL (SD 1.11, CV 4.2).
INTERAMBULACRA.?In smaller specimen, 17.7 mm
long, 10 plates in interambulacrum 3a, 8 in 4a, 13
in 5a; few plates added during growth with only
12 plates in interambulacrum 3a in specimen 30.5
mm long, 9 in interambulacrum 4a, 13 in 5a. Inter-
ambulacrum 2b and 3a separated from ocular in
in specimen only 19.6 mm long indicating that
production of new plates had ceased in these inter-
ambulacra (Kier, 1956).
PERISTOME.?Anterior, distance from anterior
edge of peristome to anterior margin of test 19.87
percent TL (SD 1.39, CV 7.0); not sunken; width
16.66 percent TL (SD 1.74, CV 10.4), height 4.26
percent TL (SD 0.94, CV 15.2). Labrum well de-
veloped, short, length 11.26 percent TL (SD 1.22,
CV 10.8), extending back to posterior portion of
first adjoining ambulacral plate or anterior por-
tion of second (Figure 8A); distance from anterior
tip of labrum to posterior of test 76.05 percent TL
(SD 2.13, CV 2.8); 33 or 34 oral tubefeet arising
from single pores, on some larger specimen large
ridge rising alongside pore or crossing pore divid-
ing pore in two.
PLASTRON.?Length 47.58 percent TL (SD 1.98,
CV 4.2), width at junction of second and third
adjacent ambulacral plates 25.29 percent TL (SD
1.99, CV 7.9), width at junction of third and fourth
adjacent ambulacral plates 36.09 percent TL (SD
1.70, CV 4.7), width at junction of fourth and fifth
adjacent ambulacral plates 38.23 percent TL (SD
1.60, CV 4.2).
PERIPROCT.?Small, ovoid, height 14.91 percent
TL (SD 1.78, CV 11.9), width 9.81 percent TL (SD
1.23, CV 12.5), situated near middle or slightly
above midpoint on posterior.
FASCIOLES.?Peripetalous fasciole curving slightly
towards apical system in interambulacra 2, 3, more
deeply in interambulacra 4 and 1; straight across
interambulacrum 5; greatest width where passing
around petals n and iv; distance from apical system
to posterior portion 18.28 percent TL (SD 0.91,
CV 5.0), to lateral portion (along center of inter-
ambulacrum 4) 16.87 percent TL (SD 0.86, CV
5.1); length of fasciole 203.42 percent TL (SD
6.17, CV 3.0); Lateroanal fasciole passes below anus,
at distance from lower portion of anus slightly
greater than height of anus; length of fasciole
170.45 percent TL (SD 4.31, CV 2.5).
See Chesher (1972:20) for plates on which fasci-
oles occur.
PEDICELLARIAE.?Tridentate common, largest oc-
curring near peristome with stalks 2 mm long,
valves 1.2 mm long (Plate 7: figures 10, 11); rostrate
(Plate 7: figures 12-14) less common with stalk 1.3
mm long, valves 0.75 mm long, seen on periproctal
plates, in apical region near edge of petals; globif-
erous (Plate 7: figures 7-9) rare, with a circle of
small spines surrounding the terminal opening.
ECOLOGY.?This species occurs in large numbers
buried in mud fields west and south of Water Cay
Range. The water depth is between 12 and 18
meters. These mud fields can be identified from
the surface on a clear day by the azure-blue color
of the water. The mud fields are white, with worm
mounds and the algae Penicillus and Udotea con-
glutinata (Ellis and Solander). The echinoids live
buried 20-100 mm below the surface. I could see
no evidence of their presence at the surface. Their
density varied from locality to locality but was
greatest in the finer sediments, decreasing where
the sediment contained some silt or sand. Maximum
density was approximately one to two specimens
every square meter. The echinoids are light yellow-
ish tan with bright red tubefeet in the anterior
ambulacrum (in) and orange tubefeet in the phyl-
lodes. They bury beneath the surface in approxi-
mately 15 minutes (Plate 8: figures 1-3). Three
other spatangoids are present with them: Brissopsis
NUMBER 206 15
elongata Mortensen, Moira atropos (Lamarck) and
Par aster cf. Paraster floridiensis (Kier and Grant).
Paraster doederleini occurs in far greater numbers
than any of these species. In 20 minutes of collect-
ing, 30 specimens were found of P. doederleini, six
of M. atropos, one of B. elongata, and none of P.
cf. P. floridiensis.
Paraster cf. Paraster floridiensis (Kier and Grant)
PLATE 10: FIGURES 1-5, 8-11
Schizaster (Paraster) floridiensis Kier and Grant, 1965:50-54,
fig. 15, pi. 13: figs. 4-6; pi. 14: figs. 1-9.?Kier, 1966:9.
Paraster floridiensis.?Chesher, 1966:1-19, figs. 1-6, tables 1-2;
1972a, tables 1, 2.
In 1973 we collected in the same mud field with
Paraster doederleini, Moira atropos, and Brissopsis
elongata, one specimen of a small spatangoid very
similar to Paraster floridiensis. When we returned
to the same site in 1974, we were determined to
find more specimens. After 11 dives and approxi-
mately 7 hours of searching, we located only one
more. Because of the scarcity of the specimens, we
thought that perhaps we were on the fringe of a
larger population present in adjacent areas. Ex-
tensive searching within two to three miles of the
site revealed no more specimens.
These specimens may be conspecific with Parasler
floridiensis; but until more specimens are found,
their reference to this species must be tentative.
They may represent a new subspecies. They are
similar in most of their dimensions (Table 2) to
specimens from Florida referred to P. floridiensis
by Kier and Grant, and Chesher. Their peripetal-
ous fasciole and lateroanal fascioles also cross the
same plates as in the Floridian specimens. They
differ in that their tests are narrower, 86.90-88.52
percent of their total length as opposed to 91.08-
95.59 percent of the total length in two Floridian
specimens of approximately the same length
(Chesher, 1972a: 19, reports a mean of 92.66 for all
the specimens he measured). The anterior paired
petals in the Carrie Bow Cay specimens are
straighter than in P. floridiensis. This difference is
readily seen in a comparison of photographs (Plate
10) of the Carrie Bow Cay specimens and a speci-
men referred to P. floridiensis by Chesher (1966)
of approximately the same size. The anterior petals
in the Carrie Bow Cay specimens curve more an-
teriorly; whereas they curve more posteriorly in
Chesher's specimen. Finally, the tridentate pedicel-
lariae in the Carrie Bow Cay specimens (Plate 10:
figures 10, 11) differ in having longer, better defined
serrated heads than those figured by Chesher for
P. floridiensis.
The Carrie Bow Cay specimens strongly resemble
P. rotundatus Doederlein from the Galapagos.
Their anterior petals are similar, not flexuous, as
in P. floridiensis; but their tests are narrower and
their anus larger. The tridentate pedicellariae in
the Carrie Bow Cay specimens are intermediate in
shape between those found in P. floridiensis and
P. rolundalus, having their heads longer with bet-
ter defined serrated heads than in P. floridiensis
but shorter than in P. rotundatus.
Until more specimens have been found from
Carrie Bow Cay, and of P. rotundatus from the
Galapagos, it is impossible to know the significance
of the differences between the Carrie Bow Cay
specimens, P. floridiensis and P. rotundatus. At first
glance I thought that the differences between the
Carrie Bow Cay specimens and the Floridian speci-
mens of P. floridiensis and the similarity shared by
the Carrie Bow specimens and the Galapagos spe-
cies suggested that the Carrie Bow Cay specimens
were more similar to the Pacific species because of
their nearer geographic location. However, a speci-
men of P. floridiensis recently collected by Dr.
David Meyer from the San Bias Islands off the
Republic of Panama (USNM El3753) is indistin-
guishable in the width of its test and flexuous
petals from the Florida specimens, even though it
occurs nearer to the Galapagos Islands than the
Carrie Bow Cay specimens. Perhaps, as suggested
by Chesher (1972b: 149) the Pacific counterparts
of the Atlantic species exist but remain to be found.
LOCALITY.?The two specimens were buried 10-
15 mm beneath and surface at a depth of 15 m in
mud fields west and south of Water Cay Range.
Chesher (pers. comm., 1974) points out that his
Floridian specimens were all found in coarse sand.
Moira atropos (Lamarck)
PLATE 8: FIGURES 6-10
Spatnngus atropos Lamarck, 1816:32. [For a synonymy, see
Mortensen, 1951:329. Chesher, 1963, described the feeding
mechanism; and Moore and Lopez, 1966, the ecology and
productivity.]
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
This species lives in the same mud fields with
Paraster doederleini Chesher, P. cf. P. floridiensis
(Kier and Grant), and Brissopsis elongata Morten-
sen. They live buried 40-100 mm in the mud, and
they can bury themselves (Plate 8: figures 8-10)
beneath the surface in 12 minutes. They may bury
deeper than 100 mm but, because of the mud's
great density, we were not able to dig deeper with
our hands in search of them.
This species has previously been reported from
the coast of North Carolina south to the coast of
Brazil from littoral depths to 145 meters.
Brissopsis elongata Mortensen
PLATE 9: FIGURES 1-6
Brissopsis elongata Mortensen, 1907:163, pi. 3: figs. 4, 14, 15,
19; pi. 4: figs. 1, 4, 13, 18; pi. 18: figs. 2, 15-17, 21, 28;
pi. 19: figs. 12, 17. [See Chesher, 1968a:63, for synonymy,
detailed description, and illustrations.]
This species occurs in mud fields with Paraster
doederleini Chesher, Paraster cf. P. floridiensis, and
Moira atropos (Lamarck). It is not common. We
collected nine specimens, and it took approximately
20 minutes to find each one. They live buried 40-
100 mm in the mud. Burrowing beneath the surface
takes 10 minutes, faster than any of the other
spatangoid species occurring with them. Observa-
tions of live B. elongata, in situ have never been
published before, and photographs of it burying
itself in its natural environment are on Plate 9:
figures 4-6.
Brissopsis elongata has been previously reported
from the South American coasts of Venezuela to
Panama and from Puerto Rico at depths of 13-72
m (Chesher, 1968a:76).
Agassizia excentrica Agassiz
PLATE 9: FIGURES 8, 9
Agassizia excentrica Agassiz, 1869:276. [See Mortensen, 1951:
345, for synonymy.]
One dead and denuded specimen was collected
by Miss Annie Bowman in February 1974, on a
beach on Carrie Bow Cay. This species is previ-
ously known from the West Indies (Cuba and the
Florida Reef to Barbados) from depths of 45-900
meters (Mortensen, 1951:346).
Cassidulus cariboearum Lamarck
PLATE 6: FIGURES 6-9
Cassidulus cariboearum Lamarck, 1801:549. [For a synonymy
see Mortensen, 1948a:205.]
Frederick Hotchkiss found six live specimens of
this species while screening the coarse sand that
lies in one meter of water 5-10 meters from the
shore on the northeast side of Carrie Bow Cay. This
discovery is particularly important because this
species has never been seen before alive in situ.
Nothing was known for certain of its living habits.
The species was known to range from a depth of
46 cm (Kier, 1962:21) to 197 m (Agassiz, 1872:343).
Although Mortensen (1948a: 193, 209) reports that
observations are lacking on living specimens, he
predicted that it would be found living buried in
coarse sand. As is so often the case with Mortensen's
predictions, he was correct. After Hotchkiss dis-
covered the specimens, he placed them in a glass
vial with some of the coarse sand; and within 3-4
minutes they had buried out of sight. The speci-
mens reached a maximum depth of 3.4 cm suggest-
ing that this may be their normal depth of burial.
Hotchkiss found only six specimens after hours of
digging through square meters of sand; therefore,
the species does not occur in large numbers. Two
dead tests that were washed up on the beach on
Carrie Bow Cay were found in April 1974 by Miss
Annie Bowman; but I found no specimens in four
weeks of diving in 1973 and 1974.
Hotchkiss reports that the specimens were white
when alive but turned green when placed in
formaldehyde.
Eucidaris tribuloides (Lamarck)
PLATE 11: FIGURES 4, 5
Cidarites tribuloides Lamarck, 1816:56. [For a synonymy, see
Mortensen, 1928:400. The biology of this species has been
described by McPherson, 1968.]
This echinoid
lagoon. It lives
following zones:
reef, rubble and
groove. It is not
ments. It occurs
particularly east
beds where the
lives both on the reef and in the
in crevices in the coral in the
reef crest, sand and rubble, patch
pavement, buttress, and spur and
common in any of these environ-
in great numbers in the lagoon,
of Water Cay Range in Thalassia
grass is especially luxurious and
NUMBER 206 17
the water depth is between 5 and 7 meters. Here
the echinoid feeds on sponges and Thalassia and
has a density of approximately one specimen per
square meter. This distribution is not regular?
commonly several specimens occur close together
eating the same sponge.
Diadema antillarum Philippi
Cidaris antillarum Philippi, 1845:355. [For a synonymy, see
Mortensen, 1940:269. For the biology of this species see
Lewis, 1964, and Randall, Schroeder, and Starck, 1964.
For illustrations, see Kier and Grant, 1965, pi. 2: figs. 4-7.]
This urchin occurs both on the reef and in the
lagoon. It lives throughout the length of the tran-
sect in cavities in the coral or rock. It is most
common in the Patch Reef Zone with a density of
6-7 per square meter. Rare at greater depths, no
specimens were observed deeper than 40 meters.
However, it has been reported elsewhere at far
greater depths. Large herds of 20-40 specimens occur
in profusion in the lagoon's Thalassia beds.
Arbacia punctulata (Lamarck)
PLATE 11: FIGURE 1
Echinus punctulatus Lamarck, 1816:47. [For synonymy, see
Mortensen, 1935:573. For the biology of this species, see
Harvey, 1956; Sharp and Gray, 1962; Kier and Grant,
1965:17; and Boolootian and Cantor, 1965.]
Only two specimens were found of this species.
They both were living in the Thalassia beds east
of Water Cay Range. Because this species does not
cover itself, it is readily visible. In spite of this, a
careful search for many days revealed no more
specimens.
Echinometra viridis Agassiz
PLATE 11: FIGURE 3
Echinometra viridis Agassiz, 1863:22. [For synonymy, see
Mortensen, 1943b:368. Kier and Grant, 1965, and McPher-
son, 1969, describe aspects of the biology of this species.]
This species is readily distinguished from E.
lucunter by its blue-tipped spines. It is much more
common, living in great numbers in niches in the
coral, particularly in beds of Agaricia (leaf coral)
in the Buttress Zone where approximately 10 speci-
mens are present per square meter. No specimens
were found below a depth of 9 meters.
Echinometra lucunter (Linnaeus)
Echinus lucunter Linnaeus, 1758:665. [For a synonymy and
illustrations, see Mortensen, 1943b:357. Kier and Grant,
1965, and McPherson, 1969, describe aspects of the biology
of this species.]
This species is restricted to shallow water less
than 3 meters deep, where it lives in crevices in
the coral. It is more prevalent in a high energy
environment and is absent from the lagoon.
Lytechinus variegatus (Lamarck)
PLATE 11: FIGURES 1, 2
Echinus variegatus Lamarck, 1816:48. [For a synonymy, see
Mortensen, 1943a:437. Subsequent papers describing the
biology and distribution are Moore, Jutare, Bauer, and
Jones, 1963; Moore, 1965; Kier and Grant, 1965; Chesher,
1968b; and Serafy, 1973.]
This species is common to the lagoon in Thalas-
sia beds where the water is more than 4 meters
deep. It covers its test with blades of Thalassia. It
is most common off Water Cay Range, where
it occurs with Clypeaster rosaceus (Linnaeus),
Eucidaris tribuloides (Lamarck), herds of Diadema
antillarum Philippi, few Tripneustes ventricosus
(Lamarck) and rare Arbacia punctulata (Lamarck).
No specimens were found on the reef.
The vast majority of the specimens belong to the
subspecies L. variegatus variegatus, but one speci-
men was collected of L. variegatus carolinus Agas-
siz. It is distinguished by its pink test and more
numerous ambulacral and interambulacral plates.
The subspecies has never been found this far south
(see Serafy, 1973, fig. 1, for map of distribution of
subspecies).
Lytechinus williamsi Chesher
Lytechinus williamsi Chesher, 1968b, figs. 1-4.
Frederick Hotchkiss collected one specimen of
this species (subsequently identified by David L.
Pawson) living in a crevice in coral on the outer
fore-reef on the edge of the outer trough on the
transect approximately 510 meters from its western
end.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Tripneustes ventricosus (Lamarck)
PLATE 12: FIGURE 3
Echinus ventricosus Lamarck, 1816:44. [For synonymy, see
Mortensen, 1943a:490. Lewis, 1958; Moore, Jutare, Jones,
McPherson, and Roper, 1963; Kier and Grant, 1965; and
McPherson, 1965, describe aspects of the biology of this
species.]
This echinoid was found both on the reef and
in the lagoon, but it is rare in both environments.
One large specimen was seen eating algae on the
boulders in the Patch Reef Zone on the transect.
One specimen was under the pier on the lagoon
side of Carrie Bow Cay. Several other specimens
were in the Thalassia beds east of Water Cay Range,
where they were eating the Thalassia.
Clypeaster rosaceus (Linnaeus)
PLATE 12: FIGURES 1, 2
Echinus rosaceus Linnaeus, 1758:665. [For a synonymy, see
Mortensen, 1948b:40. See Kier and Grant, 1965:26, for
description of living habits.]
This species was never found on the reef, but it
is very common in the lagoon in Thalassia beds in
water over 5 meters deep. It occurs with Lytechinus
variegatus, Meoma ventricosa, Eucidaris tribuloides,
Tripneustes ventricosus, and Diadema antillarum.
It lives on top of the sediment or very slightly
buried, but never with most of the thickness of its
test under the surface of the seabottom. It holds
blades of turtle grass or large shell fragments over
its test. The echinoid grazes on Thalassia (many
specimens were seen with blades of Thalassia be-
tween their teeth).
Clypeaster subdepressus (Gray)
PLATE 6: FIGURE 4
Echinanthus subdepressus Gray, 1825:427. [For a synonymy,
see Mortensen, 1948b: 112, and for a description of living
habits, see Kier and Grant, 1965:28.]
Only one specimen (dead) was found of this
species. It was located in water 30 meters deep in
a sand trough on the reef just west of the dropoff
on the transect. A thorough search was made in
similar sand fields but no more specimens were
found.
Leodia sexiesperforata (Leske)
PLATE 9: FIGURE 7
Echinodiscus sexiesperforatus Leske, 1778:199. [For a syn-
onymy, see Mortensen, 1948b:429. Goodbody, 1960, de-
scribes the feeding mechanism in this species and Kier and
Grant, 1965:31, the living habits.]
This is the only sand dollar found off Carrie
Bow Cay (many specimens of Encope emarginata
(Leske) were found living oft the mainland of
Belize along the pier of the Pelican Beach Motel at
Stann Creek). Only one specimen and many dead
were found in sand fields at the western end of
the transect in 2 to 3 meters of water. A careful
search was made in sand fields throughout the
Carrie Bow Cay region, but no more specimens
were found. Local people report that a sand dollar
used to be very common in the sand off the south
end of South Water Cay, but none occur there
now.
Echinoneus cyclostomus Leske
PLATE 6: FIGURE 5
Echinoneus cyclostomus Leske, 1778:173. [For a synonymy,
see Mortensen, 1948a:75.]
This species was found only on the reef, where
it lives in coarse sand under coral rock.
Literature Cited
Agassiz, A.
1863. List of the Echinoderms Sent to Different Institu-
tions in Exchange for Other Specimens, with An-
notations. Museum of Comparative Zoology, 1(2):
17-28.
1869. Preliminary Report on the Echini and Starfishes
Dredged in Deep Water between Cuba and the
Florida Reef, by L. F. de Pourtales, Assist. U. S.
Coast Survey. Museum of Comparative Zoology,
Bulletin, l(9):253-308.
1872. Revision of the Echini. Memoir of the Museum of
Comparative Zoology, 3(1,2): 1-378, plates 1-49.
Boolootian, R. A., and M. H. Cantor
1965. A Preliminary Report on Respiration, Nutrition,
and Behavior of Arbacia punctulata. Life Sciences,
4(16):1567-1571, table 1.
NUMBER 206 19
Chesher, R. H.
1963. The Morphology and Function of the Frontal Am-
bulacrum of Moira atropos (Echinoidea: Spatan-
goida). Bulletin of Marine Science of the Gulf and
Caribbean, 13(4):549-573.
1966. Redescription of the Echinoid Species Paraster
floridiensis (Spatangoida: Schizasteridae). Bulletin
of Marine Science, 16(1): 1-19, figures 1-6, tables 1-2.
1968a. The Systematics of Sympatric Species in West Indian
Spatangoids: A Revision of the Genera Brissopsis,
Plethotaenia, Paleopneustes, and Saviniaster. Stud-
ies in Tropical Oceanography (Miami), 7:1-168,
figures 1-25, plates 1-35, tables 1-13.
1968b. Lytechinus williamsi, a New Sea Urchin from
Panama. Breviora (Museum of Comparative Zool-
ogy), 305:1-13, figures 1-5, tables 1-3.
1969. Contributions to the Biology of Meoma ventricosa
(Echinoidea: Spatangoida). Bulletin of Marine Sci-
ence, 19(l):72-110, figures 1-14.
1970. Evolution in the Genus Meoma (Echinoidea:
Spatangoida) and a Description of a New Species
from Panama. Bulletin of Marine Science, 20(3):
731-761, figures 1-9, tables 1-5.
1972a. A New Paraster (Echinoidea: Spatangoida) from the
Caribbean. (Biological Results of the University of
Miami Deep-Sea Expeditions, 86). Bulletin of Ma-
rine Science, 22(1): 10-25, figures 1-9, tables 1-3.
1972b. The Status of Knowledge of Panamanian Echinoids,
1971, with Comments on Other Echinoderms (in
the Panama Biota: Some Observations Prior to a
Sea-level Canal). Bulletin of the Biological Society
of Washington, 2:139-158, figures 1, 2, tables 1-4.
Gmelin, J. F.
1791. Vermes. Pages 3021-3910 in part 6 of volume 1 in
Caroli a Linne Systema Naturae per Regna tria
Naturae. Lipsiae.
Goodbody, Ivan
1960. The Feeding Mechanisms in the Sand Dollar Mellita
sexiesperforata (Leske). Biological Bulletin, 119(1):
80-86, figures 1-3.
Gray, J. E.
1825. An Attempt to Divide the Echinida, or Sea Eggs,
into Natural Families. Annals of Philosophy, new
series, 10 [or 26]: 423-431.
Harvey, E. B.
1956. The American Arbacia and Other Sea Urchins.
298 pages, 12 figures, 16 plates, 18 tables. Princeton:
Princeton University Press.
Kier, P. M.
1962. A Revision of the Cassiduloid Echinoids. Smith-
sonian Miscellaneous Collections, 144(3): 1-262, fig-
ures 1-184, plates 1-44.
1956. Separation of Interambulacral Columns from the
Apical System in the Echinoidea. Journal of
Paleontology, 30(4):971-974, figures 1-3.
1966. Bredin-Archbold-Smithsonian Biological Survey of
Dominica, 1: The Echinoids of Dominica. Proceed-
ings of the United States National Museum, 121
(3577): 1-10, figures 1-3, plates 1, 2.
Kier, P. M., and R. E. Grant
1965. Echinoid Distribution and Habits, Key Largo Coral
Reef Preserve, Florida. Smithsonian Miscellaneous
Collections, 149(6): 1-68, 15 figures, 16 plates.
Lamarck, J.B.P.A. de M. de
1801. Systeme des animaux sans vertebres, ou tableau
general des classes, des ordres et des genres de ces
animaux. 432 pages. Paris.
1816. Histoire naturelle des animaux sans vertebres,
presentant les caracteres, gene'ralaux et particuliers
de ces animaux, leur distribution, leur classes, leurs
families, leur genres et la citation synonymique des
principales especes qui s'y rapportent. 1st edition,
volume 3, 586 pages.
Leske, N. G.
1778. Jacobi Theodori Klein naturalis dispositio echino-
dermatum . . . , edita et descriptionibus novisque
inventis et synonomis auctorem aucta. 278 pages,
54 plates. Lipsiae.
Lewis, J. B.
1958. The Biology of the Tropical Sea Urchin Tripneustes
esculentus Leske in Barbados, British West Indies.
Canadian Journal of Zoology, 36:607-621, figures
1-9, table 1.
1964. Feeding and Digestion in the Tropical Sea Urchin
Diadema antillarum Philippi. Canadian Journal of
Zoology, 42:549-557, figure 1, tables 1-6.
Linnaeus, C.
1758. Systema Naturae per Regna tria Naturae, secundum
Classes, Ordines, Genera, Species, cum characteribus
differentis, synonymis, locis. 10th edition, revised,
824 pages. Holmiae.
McPherson, B. F.
1965. Contributions to the Biology of the Sea Urchin
Tripneustes ventricosus. Bulletin of Marine Science,
15(l):228-244, figures 1-8, table 1.
1968. Contributions to the Biology of the Sea Urchin
Eucidaris tribuloides (Lamarck). Bulletin of Marine
Science, 18(2):400-443, figures 1-26, tables 1-8.
1969. Studies on the Biology of the Tropical Sea Urchins,
Echinometra lucunter and Echinometra viridis. Bul-
letin of Marine Science, 19(1): 194-213, figures 1-13,
table 1.
Moore, H. B.
1965. The Correlation of Symmetry, Color and Spination
in an Urchin. Bulletin of Marine Science, 15(1):
246-254, figures 1-8.
Moore, H. B., Thelma Jutare, J. C. Bauer, and J. A. Jones
1963. The Biology of Lytechinus variegatus. Bulletin of
Marine Science of the Gulf and Caribbean, 13(1):
23-53, figures 1-22, tables 1-4.
Moore, H. B., Thelma Jutare, J. A. Jones, B. F. McPherson,
and C. F. E. Roper
1963. A Contribution to the Biology of Tripneustes
esculentus. Bulletin of Marine Science of the Gulf
and Caribbean, 13(2):267-281, figures 1-10.
Moore, H. B., and N. N. Lopez
1966. The Ecology and Productivity of Moira atropos
(Lamarck). Bulletin of Marine Science (Miami), 16
(4):648-667, figures 1-14.
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Mortensen, T.
1907. Echinoidea. Part 2 of number 2 in volume 4 of The
Danish Ingot)-Expedition, 1895-1S96. 200 pages,
27 figures, 19 plates. Copenhagen.
1928. Cidaroidea. Volume 1 of A Monograph of the
Echinoidea. 551 pages, 173 figures, 88 plates [under
separate cover]. Copenhagen: Bianco Luno.
193"). Bothriocidaroida, Melonechinoida, Lepidocentroida,
and Stirodonta. Volume 2 of A Monograph of the
Echinoidea. 647 pages, 377 figures, 89 plates [under
separate cover]. Copenhagen: Bianco Luno.
1940. Aulodonta with Additions to Volume 2: (Lepidocen-
troida and Stirodonta). Part 1 in volume 3 of A
Monograph of the Echinoidea. 370 pages, 197
figures, 77 plates [under separate cover]. Copen-
hagen: Bianco Luno.
1943a. Camarodonta. Number 1 of part 2 in volume 3 of A
Monograph of the Echinoidea. 553 pages, 321 fig-
ures, 56 plates [under separate cover]. Copenhagen:
Bianco Luno.
1943b. Camarodonta. Number 2 of part 3 in volume 3 of A
Monograph of the Echinoidea. 446 pages, 215 fig-
ures, 66 plates [under separate cover]. Copenhagen:
Bianco Luno.
1948a. Holectypoida, Cassiduloida. Part 1 in volume 4 of
A Monograph of tlie Echinoidea. 371 pages, 326 fig-
ures, 14 plates [under separate cover]. Copenhagen:
Bianco Luno.
1948b. Clypcastroida. Part 2 in volume 4 of A Monograph
of the Echinoidea. 471 pages, 258 figures, 72 plates
[under separate cover]. Copenhagen: Bianco Luno.
1951. Spatangoida. Number 2 of Part 2 in volume 5 of A
Monograph of the Echinoidea. 593 pages, 286 fig-
ures, 64 plates [under separate cover]. Copenhagen:
Bianco Luno.
Philippi, R. A.
1845. Beschrcibung einiger luucn Echinodermen (Spat-
angus excavata, Sp. cavernosus, Sp. Australia, Sp.
placenta, Sp. canaliferus) nebst kritischen Bcmer-
kungen iiber einigc weniger bekanntc Arten. Archiv
filr Xaturgeschichte, 11:344-359, plate 11.
Randall, J. E., R. E. Schroeder, and \V. A. Starck II
1964. Notes on the Biolog\ of the Echinoid Diadema
antillarum. Caribbean journal of Science, 4(2/3):
421-433, figures 1-3, tables 1-3.
Serafy, I). K.
1973. Variation in the Polytypic Sea Urchin Lytcchinus
variegatus (Lamarck, 1816) in the Western Atlantic
(Echinodermata; Echinoidea). Bulletin of Marine
Science, 23(3):525-534. figure 1, tables 1-3.
Sharp, 1). T., and I. E. Gray
1962. Studies on Factors Affecting the Local Distribution
of Two Sea Urchins, Arbacia punctulata and
Lytechinus variegatus. Ecology, 43(2):3O9-313, figures
1-2.
PLATES 1-12
2 2 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
PLATE 1
Plagiobrissus grandis (Gmelin)
1, 3. Adapical, adoral view of specimen 6.9 mm long. Note immature features such as lack of
petals, large apical system, large tubercles, slight development of peripetalous fasciole,
adapical position of periproct, and large peristome. USNM E1373O, X 13.
2, 4. Adapical, adoral views of adult 150 mm long for comparison with immature specimen.
USNM E13737, from sand field east of Molasses Reef, Florida Keys, depth of 30 meters,
x %?
NUMBER 206 23
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
PLATE 2
Plagiobrissus grandis (Gmelin)
1, 2. Adoral view of specimen 20 mm long (USNM El3732), X 5. Figure 2 shows the specimen
photographed under xylene with the rod-like structures visible. Carrie Bow Cay.
3. More highly enlarged view of same specimen showing nature of rod-like structures, X 16.
4, 5. Views of edge of plate showing extension of rods over suture. USNM E13736, X 20. West
of Molasses Reef, 10 meters depth, Florida Keys.
6. SEM photograph of ventral portion of interambulacrum 1 showing that rods are produced
by thickening of the calcite with a corresponding reduction in size of the holes in calcite
mesh work. USNM El 3730, specimen 6.9 mm long, Carrie Bow Cay, X 100.
NUMBER 206 25
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
PLATE 3
Plagiobrissus grandis (Gmelin)
1, 2, 3. Adapical, adoral, left side of specimen 5.8 mm long, USNM E13731, X 10.
4, 5. Adapical, adoral views of specimen 20 mm long, USNM E13732, X 4.
NUMBER 206 27
2 8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
PLATE 4
Plagiobrissus grandis (Gmelin)
1. Ambulacrum in of immature specimen showing that the pores have begun to divide into
two to form pore-pairs. The edge of the ocular plate is at the upper part of the photo-
graph. USNM El3731, X 200.
2. Petal ii of same specimen showing that the pores have not yet begun to divide, X 200.
Meorna ventricosa (Lamarck)
3, 4. Adapical, adoral views of specimen 17.5 mm long, USNM E13734, X 4.5. A side view of
this specimen is on Plate 6: figure 3.
NUMBER 206 29
1m iiJ
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
PLATE 5
Meoma ventricosa (Lamarck)
1, 2, 3. Adapical, adoral, left side of specimen 5.8 mm long showing the relatively long spines
and large black pedicellariae characteristic of an immature specimen. USNM E13733,
X15.
4. Valve of globiferous pedicellaria from this specimen, X 400.
NUMBER 206 31
go SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
PLATE 6
Meoma ventricosa (Lamarck)
1. Adapical view of specimen 5.8 mm long showing the beginning of the development of
the petals, the large apical system, and dorsal position of the periproct. USNM E13733,
X 17.
2. Adapical view of specimen 17.5 mm long. USNM El3734, x 5.
3. Left side of same specimen before removal of spines, x 4.5. Adapical and adoral views
of this specimen with spines are on Plate 4: figures 3, 4.
Clypeaster subdepressus (Gray)
4. Adapical view of specimen USNM E13752, X i/?.
Echinoneus cyclostomus Leske
5. Adapical view of specimen USNM El3756, X 2.
Cassidulus cariboearum Lamarck
6, 7. Adapical and adoral views of USNM E13755, X 3.
8, 9. Adapical and adoral views of specimen before removal of spines, X 3.
NUMBER 206
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
PLATE 7
Paraster doederleini Chesher
1. Adapical view of the holotype USNM El 1376 from White Shoals, Dry Tortugas, Florida,
X 1.
2. Adapical view of USNM E13743 from Carrie Bow Cay, x 1.
3. Adapical view of USNM El3744 from Carrie Bow Cay, X 2. Compare this specimen with a
specimen from Colombia of similar size in figure 6, below. Note the larger petaloid area,
more divergent posterior petals, and wider ambulacrum HI in the Carrie Bow Cay specimen.
4. Adapical view of paratype USNM El 1378 from Northwest Providence Channel, Bahama
Islands, X 1.
5. Adapical view of paratype USNM El 1377 from off northern Colombia, X 1.
6. Adapical view of paratype MCZ 8396 from off northern Colombia, X 2.
7. Globiferous pedicellaria from Carrie Bow Cay specimen. USNM E13745, X 100.
8. Valve of globiferous pedicellaria from Carrie Bow Cay specimen. USNM El3746, X 75.
9. Enlarged view of teeth of same valve, X 375.
10. Tridentate pedicellaria from Carrie Bow Cay specimen. USNM El3747, X 25.
11. Value of tridentate pedicellaria from Carrie Bow Cay specimen. USNM E13748, x 25.
12. Rostrate pedicellaria from Carrie Bow Cay specimen. USNM E13749, X 50.
13. Valve from rostrate pedicellaria from specimen from Carrie Bow Cay. USNM El3750, X 50.
14. Enlarged view of teeth of same valve, x 375.
NUMBER 206 35
12 13 14
35 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
PLATE 8
Par aster doederleini Chesher
1, 2, 3. Photographs of specimen burrowing into mud at a depth of 15 meters. The pictures
were taken over a period of 15 minutes. USNM El3740, 47 mm long.
4, 5. Adapical and adoral views of USNM E1376O, X 1.
Moira atropos (Lamarck)
6, 7. Adapical and adoral views of USNM El3759, X 1.
i, 9, 10. Burying sequence at same site as for P. doederleini above. The pictures were taken
over a period of 15 minutes. USNM E13738, 42 mm long.
NUMBER 206 37
g a r ; ? ? ; - .?, ? -:?: ^ s 3 $ g*;.
10
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
PLATE 9
Brissopsis elongata Mortensen
1,2, 3. Adapical, adoral and right side of USNM El3757, X 1.
4, 5, 6. Burying sequence of USNM E13739 at a depth of 15 meters. Pictures taken over a
period of 10 minutes. Specimen 58 mm long.
Leodia sexiesperforata (Leske)
7. Adapical view of USNM El3758, X ]/2.
Agassizia excentrica Agassiz
8, 9. Adapical and right side of USNM E13754, X 2.
NUMBER 206 39
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
PLATE 10
Paraster cf. Paraster floridiensis (Kier and Grant)
1, 2, 3. Adapical left side and adoral views of USNM E13741, X 4.
4, 5. Adapical and adoral views of USNM E13742, X 4.
Paraster floridiensis (Kier and Grant)
6. Adapical view of MCZ 8495 from the Florida Keys, X 4.
7. Adapical view of MCZ 8494 from the Florida Keys, X 4.
Paraster cf. Paraster floridiensis (Kier and Grant)
8. Globiferous pedicellaria from USNM E13742, X 175.
9. Valve of globiferous pedicellaria from USNM E13742, X 175.
10. Tridentate pedicellaria from USNM El3742, X 90.
11. Valve of tridentate pedicellaria from USNM E13742, X 100.
NUMBER 206 41
4 2 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
PLATE 11
Lytechinus variegatus variegatus (Lamarck) and Arbacia punctulata Gray
1. The individual on the left (indicated by arrow is L. variegatus variegatus. It is commonly
found almost completely covered with blades of Thalassia. The specimen on the right eating
Thalassia is A. punctulata. Photographed in water 3 meters deep.
2. Lytechinus variegatus variegatus browsing on Thalassia.
Echinometra viridis Agassiz
3. An individual in one meter of water living in a crevice in coral.
Eucidaris tribuloides (Lamarck)
4. Eating Thalassia in water 4 meters deep.
5. Feeding on a sponge.
NUMBER 206 43
4 4 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
PLATE 12
Clypeaster rosaceus (Linneaus)
1. Three individuals (indicated by arrows) covered with blades of Thalassia. Photographed in
water 4 meters deep.
2. Individual holding over its test blades of Thalassia and the dead test of a Lytechinus
variegatus variegatns (Lamarck).
Tripneustes ventricosus (Lamarck)
3. Individual feeding on Thalassia in water 3 meters deep.
NUMBER 206 45



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author when he returns page proof.
Two headings are used: (1) text heads (boldface in print) for major sections and chap-
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headings may be worked out with the editor.
In taxonomic keys, number only the first item of each couplet; if there is only one couplet,
omit the number. For easy reference, number also the taxa and their corresponding headings
throughout the text; do not incorporate page references in the key.
In synonymy, use the short form (taxon, author, date:page) with a full reference at the
end of the paper under "Literature Cited." Begin each taxon at the left margin with subse-
quent lines indented about three spaces. Within an entry, use a period-dash (.?) to separate
each reference. Enclose with square brackets any annotation in, or at the end of, the entry.
For references within the text, use the author-date system: "(Jones, 1910)" and "Jones
(1910)." If the reference is expanded, abbreviate the data: "Jones (1910:122, pi. 20: fig. 1)."
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