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1969 Chesher - Contributions To The Biology of Meoma Ventricosa
1969 Chesher - Contributions To The Biology of Meoma Ventricosa
RICHARD H. CHESHER2
Museum of Comparative Zoology, Harvard University
ABSTRACT
The biology of Meoma ventricosa (Echinoidea; Spatangoida) is de-
scribed on the basis of a two-year study conducted in Florida, the Bahama
Islands, Panama, and Colombia. The habitat, behavior, food and feeding,
growth, reproduction, predators, parasites, commensals, abnormalities,
internal anatomy, and relation to the substrate of this large, abundant
echinoid are discussed.
INTRODUCTION
Meoma ventricosa was described by Lamarck in 1816 but until Kier &
Grant (] 965) published data on its distribution and behavior, nothing was
known about the ecology of this large and abundant animal. M. ventricosa
is a member of the largest and most successful order of sea-urchins, the
Spatangoida. Spatangoids live in the mud and sand bottoms of all oceans
and range in depth from the intertidal zone to about 5000 meters. The
literature and systematics of the group are in excellent order, having been
summarized by Mortensen (1950, 1951). The fossil history of spatangoids
is perhaps superior to that of any other echinoderm group. One of the
few known examples of long-term, continuous evolution is that of a spat-
angoid from the Chalk of England (Kermack, 1954; Nichols, 1959).
In spite of such impressive attributes, very little is known about spatan-
goid biology. What data is available shows this to be a rewarding area of
study. Moore (1936), Nichols (1959, 1962), Buchanan (1966, 1967),
Brattstrom (1946), and Vasseur & Carlsen (1949) have studied ecological
aspects of European spatangoids, and Moore & Lopez (1966), Chesher
( 1963 ), and Kier & Grant (1965) have studied ecological aspects of some
tropical spatangoids.
One of the major difficulties in studying spatangoids has been their habit
of burrowing, which makes them difficult to find and capture. This is
particularly true in shallow water, where maneuvering is dangerous for
ships with adequate dredging facilities. The development and popularization
of free diving apparatus has opened the way for intensive ecological studies
in these and other areas which are relatively inaccessible from the surface
(Riedl, 1967).
M. ventricosa (Fig. 2) is a burrowing sea-urchin common to the coral
1 Contribution No. 986 from the Institute of Marine Sciences, University of Miami.
" Present address: Department of Biology, College of Guam, Agana, Guam.
1969J Chesher: Biology of Meoma ventricosa 73
reef areas of the West Indies; it is one of the largest and most accessible
of the sand-dwelling, tropical spatangoids. Large populations found in
Florida, the Bahamas, and Panama provide an excellent opportunity to
examine the urchins in their natural habitat. Using various diving tech-
niques, which ranged from snorkeling to night diving with small submarines,
populations of M. ventricosa were examined from Fort Lauderdale to Key
West along the Florida coast, and in a variety of locations in the Bahama
Islands, in Panama, and in eastern Colombia. Monthly observations and
collections were made near Molasses and Alligator reefs in the Florida
Keys. Distribution, behavior, reproduction, growth rates, and ecological
parameters were examined in the field. Morphology, internal anatomy,
and details of behavior were studied in the laboratories of the Institute of
Marine Sciences, Miami, Florida. The study began in 1964 and continued
through the summer of 1966.
I
FIGURE I.-A, portable air-lift dredge for capturing small specimens of burrow-
ing animals. (n, net bag to retain specimens; h, clamp; p, plastic tube, 3 meters
long, 6.5 cm diameter; i, intake end; c, collar to provide an even flow of air
into the intake end of the dredge; v, valve controlling the air flow; t, high pres-
sure rubber hose; y, yoke to fasten hose to scuba tank.)-B, measuring board for
tagging urchins underwater. Construction is of %-inch clear Plexiglas. Ca,
movable arm; I, pencil; 95, the tag number on the urchin and marked on the
board; s, pencil sharpener.)
1969] Chesher: Biology of Meoma ventricosa 75
the end with the net was directly above the diver and the end which re-
ceived the flow of air was held next to the sand. The dredge sucked up
large amounts of sand and could operate continuously for about 15 to 20
minutes. The scuba tank was held on the diver's back with a double tank
harness, which also held another tank for the diver's breathing air. The
air-lift was quite useful for digging small specimens out of sand pockets
and grassy areas where other methods of digging were difficult. Alan
Emory, of the Institute of Marine Sciences, subsequently used the same
apparatus to sample planktonic organisms from the small niches within the
coral reef structure.
Finding small urchins in the broad, open sand areas presented still an-
other problem. The air-lift was too slow, and digging by hand was un-
economical. Surface trawls did not cover enough distance along the bottom
(they became clogged within a few meters). A portable 3.5-h.p. water
pump was placed in an outboard motorboat, and a 60-meter length of fire-
hose with a brass nozzle was used to direct a stream of water at the sand.
The jet of water cleared away the sand rapidly, exposing the burrowing
animals without harming them. The jet was directed in long, slow sweeps
from side to side as the diver moved ahead. By working with the direction
of the bottom currents in the area, the silt that was stirred up was carried
away from the diver.
Tagging, feeding, and behavior experiments were performed in situ, and
the methods are described below in the sections on growth, feeding and
behavior.
Ciliary currents were discerned by using a dissecting microscope and
biologically inert fluorescent particles (Tracer-Glow pigments available
from Wildlife Supply Co., Saginaw, Michigan) and food-coloring dyes.
Small dots of the food dye were placed on the test with a hypodermic
syringe; the progress of the ciliary currents was easily traced by the move-
ment of the dye. Movement of sand particles was determined by using
black grains of sand. After the animal had begun to burrow into the zone
containing the black grains, the burrow was opened up and the progress
of these grains was then visible. Movement of particles and feeding methods
were observed through a glass-bottomed aquarium.
Analyses of the sediment were based on the techniques described by
Krumbein & Pettijohn (1938). Core samples were taken by pushing a
plastic core tube, 20 cm in length, into the substrate. Both ends of the
core tube were then stoppered with rubber plugs. The cores were stored
in an ice-cooler and frozen within two hours. H~S was measured with a
kit produced by Hatch Chemical Co. Particle size was measured by wet
screening through a nest of sieves with mesh sizes of 2.0, 1.0, 0.5, 0.25,
0.125, and 0.062 mm. Silt and clay fractions were separated via liquid
suspension (Krumbein & Pettijohn, 1938). Porosity of the substrate was
76 Bulletin of Marine Science [19(1)
determined by measuring the loss in weight of the sand upon drying. Per-
meability was measured in a permeameter as described by Stearn (1927).
Total organic carbon present was measured by ignition of a preweighed
sample to 500°C, at which temperature the organic carbon is driven off as
CO2. CaCOx does not burn until a much higher temperature is reached.
Crucibles were pre-ignited to 1000°C, cooled in a desiccator, and weighed.
Samples were cut from the central portion of a frozen core, using a sterile,
stainless steel knife, and then were dried in a vacuum desiccator at 28
Ibsjin2 and 50°C before being weighed prior to ignition.
HABITAT
Meoma ventricosa has the greatest ecological diversity of all the West
Indian Spatangoida. It has been collected from the intertidal zone to depths
of 200 meters, in sediments ranging from fine, heavily silted sand to coral
rubble, and in bionomes ranging from beds of Thalassia to coral reef tops
and the deeper sea floor. Geographically, this species is known from Santa
Marta, Colombia, along the American coastline to Fort Lauderdale, Flor-
ida, and from most of the West Indian islands. A few specimens have
been taken from Bermuda.
Four principal habitats can be delimited for Meoma ventricosa: Thalassia
grass with pockets of sand, shallow-water sandy areas with grass and
patches of coral, coral reef areas, and deep-water sandy areas.
1969] Chesher: Biology of Meoma ventricosa 77
Thalassia Grass with Pockets of Sand (0-14 Meters Depth).-Specimens
of M. ventricosa were found buried or partially buried in patches of cal-
careous sand within the beds of Thalassia. They tended to congregate to
one side of the sand patches, close to the grass. Urchins were found buried
between the Thalassia plants where these were not too dense, and occa-
sionally were found covered only with bits of shell and grass in dense
grass beds. Where siltation was pronounced (i.e., in Hawk Channel off
Key Largo, Florida) M. ventricosa was often found completely uncovered.
Under such conditions, abnormal individuals and stunted adults occurred
frequently (see section on abnormalities, below).
In the pockets of sand, the urchins had a profound effect on the particle-
size distribution. The burrowing action of M. ventricosa tended to shift
the larger particles to the surface of the substrate and, where the urchins
were congregated, the surface of the sand was composed of coarse particles,
whereas a few meters away the sand was fine and poorly sorted.
Wave action in this habitat is mild; the waves seldom reach more than
a meter in height. Tidal currents of up to one-half knot are common. The
sandy, grassy habitat where M. ventricosa was found was either part of a
reef tract (Florida Keys, Andros, New Providence, the Berry Islands,
Abaco, Panama, Grand Bahama) or in protected bay areas (Bimini,
Andros, Colombia). Few particles of silica were found in the sand of
these areas, and little or no H2S.
Other echinoids found in these areas were Clypeaster rosaceus (found
mainly on the grass beds, but also buried in the sand), Clypeaster sub-
depressus (common only in the deeper areas and covered with a thin layer
of sand), Lytechinus variegatus, Tripneustes ventricosus, and Arbacia
punctuJata (on the grass beds), Eucidaris tribuloides and Diadema antil-
larum (usually associated with sponges and small coral heads), and Leodia
sexiesperforata (sandy areas).
Shallow-Water Sandy Areas with Grass and Patches of Coral (5-15 Meters
Depth).-This area, bordered on one side by Thalassia beds and on the
other by shallow-water coral reefs (or deep water) was found to support
large populations of M. ventricosa distributed primarily along its borders.
Occasionally, a population was found in the center of an extremely large
sandy area, but such populations were moving and in time probably con-
gregated along a grass or coral boundary. Along the Florida reef-tract,
such sandy areas are often several miles long and average about 400 meters
wide. The sand, itself, is clean, white, calcareous, and firmly packed. It
usually has deep ripples caused by occasional heavy wave action. The
waves reach a maximum of about 1.5 to 2 meters in height. The pattern
of ripples is irregularly broken by the traces of the macroinvertebrates liv-
78 Bulletin of Marine Science [J 9(1)
Of the four environments, the boundary between the sand and coral reef
or Thalassia bed supported the largest populations. Heavily silted sand and
sand with large amounts of silica were the poorest habitats for M. ven-
tricosa. It is evident that silt and silica sand, which cause an increase in
H2S and decrease in pH and permeability, limit the dispersal of M. ven-
tricosa.
BEHAVIOR
rowing in the sand between the grass blades or near coral outcroppings,
and urchins of greater than 50 or 60 mm in test length were burrowing in
the open sandy areas near the grass.
Urchins 60 mm in test length were occasionally found emerging from
the sand at night. Changes in morphology, growth rates, and pigmentation
accompany this increase in mobility. Young M. ventricosa were not com-
monly found emerging at night until they were about 100 mm long.
Depth of Burrowing.-During the daylight hours, the urchins are buried
in the sand. The depth to which they are covered depends on size,
gonadal condition, rate of movement, the nature of the sand, wave action,
and possibly water temperature. As the urchins must move water for
respiration through the interstitial spaces of the substrate, they cannot
burrow deeper than the length of their apical spines in fine sand or in silty
conditions (Fig. 3,B,C). In areas of clean, unsilted sand, these urchins
were found as much as 10 cm below the surface of the sand (Fig. 3,D).
Wave action, and the subsequent disturbance of the surface of the sand,
caused M. ventricosa to burrow deeper into the substrate. M. ventricosa
tended to burrow deeper during the colder months of the year, while dur-
ing the summer months almost all of the urchins were just below the sur-
face of the sand. Urchins which had high respiratory demands (large, ripe,
or actively moving urchins) were not buried as deeply as urchins with
lower respiratory demands (small, spent, or motionless urchins). The
specimens of M. ventricosa which were buried the deepest (their apical
spines covered to a depth of 6 to 10 em) were almost motionless during
the periods of observation.
TABLE 1
FEEDING RATES AND MOVEMENTS OF Meoma
(7/16/66, 1100 to 1600 hrs, 6 meters depth, Molasses Reef, Station RHCK 60)
average depth of the lower part of the animals at Sta. 1 was 6 em, and the
daytime rate of movement was 5 em/hr. The amount of sand turned over
by each echinoid was, therefore, 11 x 6 x 5, or 330 cc per hour. One
cubic meter of sand 6 em deep contains 6 x 104 cc of sand. At a density
of three specimens of M. ventricosa per square meter, 990 cc of sand are
overturned each hour within one meter. Thus, it requires only 60.6 hours
for the M. ventricosa to disturb their entire habitat within that particular
patch of sand.
1969] Chesher: Biology of Meoma ventricosa 87
The average rate of feeding was 25 cc per hour per urchin. Therefore,
it would require 800 hours before all of the sand would be passed through
their intestines. Since the continual overturn of the substrate helps to in-
crease the water content and provide fresh surfaces for algal and bacterial
growth, the activities of the urchins probably stimulate microbenthic ac-
tivity and so help to maintain the amount of food available. Since M. ven-
tricosa is the largest and most abundant macroorganism in the sand of
many areas, its influence on the distribution of sedimentary particles and
on the productivity of the sandy areas must be pronounced.
Each M. ventricosa disturbs about 48 square meters of substrate to a
depth of 6 cm each year. Assuming that the individuals of M. ventricosa
do not cover the same areas again and again (as they do in enclosed pockets
of sand), an area with a density of more than one M. ventricosa per 48
square meters would be completely disturbed in one year.
GROWTH
14
12
-
"co"
-
'=:8
...•..~
6
o
o 2 3 4
Years
FIGURE 6. The growth curve of M. ventricosa derived from tagging experi-
ments.
dicated that this method of tagging did not alter the growth rate of the
urchins significantly.
During March and April of 1965, 150 specimens of M. ventricosa were
tagged. Forty-eight additional small urchins were tagged as they were
found and were added to the population. Of the 198 specimens of M.
ventricosa tagged, 117 urchins were recaptured (some of those recaptured
were removed from the population while others were retagged and left in
the population). Of the 117 individuals examined for a period of two
months or longer, 48.6 per cent showed some growth and 51.4 per cent
showed no growth. Growth was followed for 37 individuals for one year
or more, and, of these, only 37.9 per cent showed no growth. Urchins
which had apparently stopped growing ranged in size from 109 to 144 mm
test length.
Since an estimate of the maximum growth rate of M. ventricosa was
sought, urchins which had already stopped growing were not included in
the calculation of the growth curve. The growth curve (Fig. 6) was cal-
culated from a Walford Line (Ricker, 1958) using the 57 specimens which
showed some growth during a two-month (or greater) period. If the in-
terval of time was greater than two months, the growth vs. time was
plotted and interpolated to the standard two-month period. The least-
square line for the Walford plot (Fig. 7) is L = 0.84Lt + 21.82. Loo,the
maximum size to which M. ventricosa is likely to grow, is 136.4 mm. Thus,
1969] Chesher: Biology of Meoma ventricosa 89
15
14
13
~ on
=12 0
E
N
,:!:.11
-
:;10
..•
••••
c
...
-'9
7 8 9 10 11 12 13 14 15
Test Length (em) at T
FIGURE 7. Walford plot showing the length at tagging (T) versus the length
at recapture 2 months later (T + 2 months).
.6
.5
>1 ,,
9> .4 ,,
0•....
...• ,
.3 0-_ •• __ ""'0..
1965 -66
.2
.1
o
MAR APR MAY JUN JUL AUG
100
o
1964 1965
FIGURE 8. Changes in gonadal indices for female specimens of M. ventricosa
during 1964 to 1966, and the changes in gonadal condition from 1964 to 1965.
by ignition) was 2.2 per cent, by weight, of the sand in the silty habitat
and 1.8 per cent, by weight, of the sand in the open sandy areas. Popula-
tion densities in the open sandy areas were as great as, or greater than, in
the silty areas. The total available organic material per urchin was, there-
fore, found to be higher in the inshore station where the urchins were
growing more slowly, and to a smaller size, than the urchins from the off-
shore, open sandy areas. Bader (1952) found a peak concentration of the
sand dollar, Echinarachnius parma, in areas with an organic content of
0.917 to 0.977 per cent of the total substrate. The numbers of echinoids
decreased as the organic content increased.
The inshore urchins spend more time on the surface of the sand than
do the offshore urchins, probably because of the increased silt content of
the substrate and the pronounced changes in oxygen concentration asso-
ciated with the grass beds. Some populations in deeper areas, particularly
those in depths greater than 30 meters, were found completely buried at
1969] Chesher: Biology of Meoma ventricosa 91
night. Apparently, these were not subjected to the same environmental
pressures as populations in shallower areas, and, although the feeding rates
of these deeper water populations were not measured, it is believed they
were feeding normally while buried. If this is true, one explanation for the
size differential could be that the large urchins from the deep-water stations
feed continuously, whereas those in shallower water spend considerable
time feeding slowly, if at all, on the surface of the sand.
The low concentration of oxygen found at night in the areas containing
Thalassia may, in addition to slowing the growth by decreasing the amount
of time spent feeding, act directly upon the growth rates. If the urchins
are sufficiently affected by the low concentration of oxygen to stop their
feeding, it is likely that other physiological activities might be slowed or
stopped during the period of stress.
REPRODUCTION
Gonadal Volume.-Changes in gonadal volume were followed from Sep-
tember, 1964, to August, 1966. The gonadal indices (10 X gonad volume/
test volume) for female individuals of M. ventricosa are plotted in Figure
8. Male gonadal development closely paralleled the female development
and averaged about 8 per cent less in volume than female gonads. The
degree of ripeness was determined by gonadal smears. Some ripe males
(indicated by viable sperm in the gonadal smear) were available all year.
Development of female gonads was recorded as spent (no ova), small ova
(developing, nucleate ova less than 0.19 mm in diameter), large ova (0.19
to 0.29 mm in diameter but still nucleate), and some ripe ova. Only one
individual was found with more than a few ripe ova.
Spawning.-In March of both years, individuals of M. ventricosa were
completely spent, and had a gonadal index of 0.18. Gonads developed
rapidly, reaching a peak index of 0.55 in July of 1965, and a peak of 0.46
in August of 1966. Some urchins with ripe ova were present in August,
as were a few spent individuals. Spawning began, therefore, sometime in
August and continued until February. The peak spawning period was
from November to January. Almost all males were ripe from June to
February.
On January 3, 1964, at 4 P.M., spontaneous spawning of a male M.
ventricosa was observed. The urchin was buried to the peripetalous fas-
ciole. No external stimulus was noted. The animal spurted a milky column
of sperm about 20 em into the water. This was repeated twice at intervals
of about two minutes. The sperm dispersed and drifted away with the
slow bottom current. Other urchins nearby (including those down-current
from the spawning individual) were not spawning and showed no reaction.
The male remained motionless for one hour, at which time observation was
discontinued.
92 Bulletin of Marine Science [19(1)
Moore & Lopez (1966) indicated that the lack of fully ripe females in
Moira atropos may be an indication of repeated, limited spawning periods
which are possibly correlated with lunar phases. Thus, a proportion of
the ova ripen and are released monthly. If this were so, the ripe ova should
be concentrated near the oviducts and the unripe ova confined to the areas
adjacent to the gonadal walls. Sections of female gonads with some ripe
ova show that the ripe ova are randomly distributed among the nucleate
ova. The ripe ova can be found in any portion of the gonad, and it is
doubtful that these could be sorted out just prior to a monthly spawning.
When spent females are found, they are thoroughly emptied of ova
(with the exception of a few large ova which are in the process of breaking
down). On August 28, 1963, a specimen of Moira atropos was found
which contained about 85 per cent ripe ova 0.12 mm in diameter. When
the ovaries were removed, the ripe ova were ejected from the torn oviduct
in a steady stream. The ova were fertilized and larvae developed to the
five-armed pluteus stage. On November 21, 1964, one M. ventricosa was
found with about 80 per cent ripe ova. As with the specimen of Moira
atropos, the ova were ejected in a steady stream from the oviduct. This
would indicate that when Meoma ventricosa or Moira atropos spawn, they
release the genital contents completely and not in small portions.
Since fully ripe individuals have not been found and since the gonadal
volume of the entire population decreases at a steady rate during the
spawning period (Fig. 8), it is evident that spawning must occur on an
individual basis and not as a single mass spawning. The evidence further
suggests that the ova are kept in an immature state and, when the proper
stimulus occurs, ripen in a few hours (or minutes) before being shed.
Chaet (1966) has shown that the nucleate ova of asteroids can ripen in a
few minutes after the addition of an extract from the radial nerves. Spawn-
ing may well occur at night, explaining the lack of ripe females in the
samples which have been taken during the day.
Holland (1967), for the cidarid Stylocidaris affinis, noted a phenomenon
similar to that now found for M. ventricosa, and suggested a rapid matura-
tion of ova just prior to spawning. Holland's contention that this type of
gametogenesis is evidence supporting separation of the cidarids into a sub-
class Perischoechinoidea is poorly based. Meoma ventricosa and Moira
atropos have a gametogenic process that is very similar to that of Stylo-
cidaris affinis, whereas gametogenesis in Echinocardium corda tum is similar
to that of other "Euechinoidea" (Moore, 1936).
Maturation.-Although genital pores appear in M. ventricosa at a test
length of between 39 and 52 mm, gonads do not mature until the test length
is about 90 mm. As in Moira atropos (Moore & Lopez, 1966), larger first-
year individuals of M. ventricosa may spawn to a limited degree, but major
1969] Chesher: Biology of Meoma ventricosa 93
spawning probably begins during the second year of life. Larger specimens
of M. ventricosa produce more gonadal material relative to the test volume
than do smaller individuals. Thus, the mean maximum gonadal index for
those specimens of M. ventricosa that were 120 to 125 mm long was 0.49
in 1965, while those that were 135 to 140 mm long had a maximum
gonadal index of 0.68 during the same period. One specimen, 165 mm
in test length, had a gonadal index of 0.77.
Comparison of Gonadal Production to That of other Spatangoids.-AI-
though there is a considerable difference in adult size between Meoma
ventricosa (130-140 mm test length) and Moira atropos (40-50 mm test
length), their relative gonadal production is similar. The maximum gonadal
index for Moira atropos was found to be 0.62 in 1964, and 0.47 in 1965
(Moore & Lopez, 1966). The maximum gonadal index for Meoma ven-
tricosa was 0.55 in 1965, and 0.46 in 1966. Echinocardium cordatum in
England reaches a gonadal index of 1.1 (Moore, 1936). Moore & Lopez
( 1966) attributed the larger gonadal index of the northern urchin to the
fact that it has four gonads, whereas Moira atropos has two. Meoma ven-
tricosa, however, also has four gonads. It is interesting that the two tropical
spatangoids, despite differences in size, habits, and habitat, have similar
gonadal indices, although the similarity may be coincidental.
PREDATORS
The adult Meoma ventricosa has few predators. Its large size, sturdy
test, burrowing habit, and chemical makeup are effective protection against
many echinoid predators. Animals which were observed feeding on Meoma
ventricosa include a loggerhead turtle (Caretta caretta caretta), a large
stingray (Dasyatus americana), helmet shells (Cassis madagascariensis),
a crab (Calappa flammea), a hogfish (Lachnolaimus maximus), and a
parrot fish (Sparisoma aurofrenatum).
The turtle, which was observed in a broad sandy area off Lower Mate-
cumbe Key, Florida, bit a large M. ventricosa, crushing the test into sev-
eral large pieces, and then swam off, leaving the broken test scattered on
the bottom. Only a few fragments of the test and portions of the intestine
and gonads were missing from the remains, indicating that the turtle was
either frightened off by my proximity or did not find the urchin acceptable
food.
The stingray, observed on the sand near a turtle-grass bed, was first
seen lying motionless on the surface of the sand. When prodded, it swam
off, disclosing the remains of a partially eaten adult M. ventricosa. About
half the test was broken away, and a portion of the internal organs was
missing from the urchin.
The crab was observed eating a specimen about 85 mm long. One claw
94 Bulletin of M'arine Science [19(1)
was used to hold the urchin against the anterior portion of the crab while
the other claw was used, much like a can opener, to break away the dorsal
portion of the test. Only the gonads and a portion of the intestine were
eaten by the crab during about one hour of feeding. Numerous specimens
which had been similarly "opened" were found during the course of this·
study, and it is thought that this crab may be the major predator of small
individuals of M. ventricosa in some areas.
The helmet shell, Cassis, feeds by drilling a hole through the test and
digesting all of the urchin, leaving only a bare test behind (Moore, 1956).
Cassis madagascariensis, which normally feeds on Plagiobrissus grandis,.
was found eating a M. ventricosa (112 mm test length). Several other tests
which had been drilled by Cassis were found during the study, but the much
greater incidence of drilled tests of Plagiobrissus grandis indicated that M.
ventricosa is not the preferred food of Cassis. Some of the drilled speci-
mens of M. ventricosa had been partially eaten, and some had not been
eaten at all. The conclusion is that Meoma ventricosa is not commonly
preyed upon by Cassis.
The hagfish, Lachnolaimus maximus, was seen biting an adult M. ven-
tricosa. Teeth marks could be clearly seen in the injured specimen but the
test was not broken. Kier (Kier & Grant, 1965) observed a small parrot
fish (Sparisoma aurofrenatum) feeding on the spines of M. ventricosa.
Kier also observed a starfish (Oreaster reticulatus) eating M. ventricosa.
This starfish is a general omnivore which eats any available injured or
decaying material. The injuries described by Kier & Grant (1965) as
occurring on many specimens of M. ventricosa can be caused by an infec-
tion which occurs when, for prolonged periods of time, the animals are
unable to burrow. Such injuries also developed in a number of specimens
kept in aquaria. The spines of the infected area dissolved or fell off, and
the integument turned purple-brown; often a depression was dissolved in
the test and the limits of infection were clearly shown by an abrupt change
from healthy to infected tissue. Boolootian et al. (1959) reported a similar
"peculiar disorder" in specimens of the regular echinoid Allocentrotus
fragilis kept in aquaria. While the starfish could have been preying on a
healthy M. ventricosa, it may have been feeding upon an infected animal.
In any case, no other instances of starfish feeding on M. ventricosa were
found.
Even the other animals listed above do not seem to be active predators
of M. ventricosa, and none are known to depend on it as a source of food.
In contrast to what occurs for other echinoids, when the test of M. ven-
tricosa is broken and scattered about underwater, fish do not rush in to
eat the remains.
When disturbed, M. ventricosa secreted large amounts of a yellowish
pigment which was repellent to small fish and killed both small fish and
1969] Chesher: Biology of Meoma ventricosa 95
crustaceans in confined conditions. The pigment is water soluble and tastes
strongly of iodine. The degree of pigmentation increases with age. No
other spatangoid is known to produce such a large amount of pigment and
it is probable that the rapid secretion of pigment when the animal is phys-
ically injured or disturbed acts as a deterrent to predators.
PEDICELLARIAE
The gut, between the origin of the primary siphon (j) and the junction
of the secondary siphon (S2), is highly vascularized. At the end of this
vascularized area, the secondary siphon removes additional water from the
sand, returning it again a short distance farther along the intestine. Just
opposite the origin of the secondary siphon, the caecum (c) attaches to
the anterodorsal portion of the intestine. The caecum is a highly vas-
cularized, thin, convoluted sac which occupies a major portion of the coe-
lom between the intestine and the gonads. Sand does not enter the caecum.
Sand progresses through the lower intestine to the upper intestine, and is
100 Bulletin of Marine Science [19(1)
1969] Chesher: Biology of Meoma ventricosa 101
finally pushed through the rectum to the anus, which dilates periodically
to release the sand.
The haemal system is similar to that described by Koehler (1883) for
Brissus unicolor. An inner vessel (v) adjoins the primary siphon. It divides
in the vicinity of the oesophagus, and one branch connects to the caecum
while the other connects to the haemal ring encircling the peristome. The
outer vessel (ov) originates at the peristome in interambulacrum 4. It
attaches to the mesenteric system of the oesophagus and lower intestine.
The heavy, sand-filled gut is supported by mesenteries located along the
entire course of the intestine. The inner portions of the intestinal loops
are attached to one another by mesenteries, dividing the coelom into upper
and lower sinuses.
The coelomic fluid is circulated by ciliary action of the peritoneum
(Fig. 13,D). A strong current was detected moving anteriorly along the
oesophagus. This current originates from fluids flowing centrally from the
median, ventral portion of the first loop of the intestine. With the excep-
tion of this current, which seems to be a localized circulation in the vicinity
of the mouth, the currents radiate outward over the intestine and mesen-
teries to the wall of the test. In general, the currents flow in an opposing
direction along the adjacent parts of the wall. Currents are most pro-
nounced on the oesophagus, the highly vascularized portion of the intes-
tine, the caecum, the ampullae of the tube-feet, and the gonads.
RESPIRA TORY CURRENTS
The respiratory currents of the petaloid area are shown in Figure 12.
The solid arrows indicate the direction of flow of currents on the external
and internal portions of the test, as determined from observing the flow
of small concentrations of food dye and small particles of fluorescent plas-
tic. The dotted arrows indicate the flow of currents within the tube-feet
and ampullae as determined by the observation of the movements of small
pigmented particles which occur naturally within the water-vascular sys-
tem. A counter-current system was found similar to that recorded from
the respiratory organs of many aquatic animals including oysters and fish
(Prosser & Brown, 1962). The effect of this system is to increase the
efficiency of oxygen exchange through the respiratory membranes. The
respiratory tube-feet (g) and the ampullae (a) are subdivided internally
into numerous small sinuses and the surface of the tissue is convoluted,
particularly on the ampullae. The currents flowing through and around this
system are vigorous.
Current flow through the feeding tube-feet is somewhat different. There
is only one pore through the test, and the currents flow into the bulb-
shaped ampulla from the tube-foot along the walls of the pore and am-
pulla, and leave through the center of the ampulla and pore.
102 Bulletin of Marine Science [19(1)
Figure 13,A-C shows the movement of sand and water within the bur-
row of M. ventricosa. The dotted lines represent movement of sand and
the solid lines represent movement of water. Circles with a dot in the cen-
ter represent areas where the current leaves the surface of the test.
Since the burrow of M. ventricosa does not communicate with the sur-
face of the sand, the circulation of water for respiration is a problem.
Water is obtained by drawing the surface water through the interstices of
the sand (or, in silty areas, through the opening kept open by the apical
spines). The water flows outward from the apical system, as shown in
Figure 13,A, past the respiratory tube-feet, and through the peripetalous
fasciole. The water must then be forced out into the interstices of the sur-
rounding sand. Since the permeability of the substrate normally decreases
1969] Chesher: Biology of Meoma ventricosa 103
FIGURE 13. A-C, movements of water (solid arrows) and particles (dotted
arrows) within the burrow of M. ventricosa. Circles with a dot in the center
indicate areas where the water currents leave the burrow. D, the flow of
coelomic fluids within the test of M. ventricosa, as viewed from the ventral side.
104 Bulletin of Marine Science [19(1)
PPF
FIGURE 14. The function of the peripetalous fasciole. The diagram shows the
fasciole (PPF) with the respiratory section (R) of the burrow on the left, and
the drainage area (D) on the right. Water currents are represented by dotted
arrows. Mucus (MS), produced from the ends of the clavulae, is shown ad-
hering to the sand.
with increasing depth (Krumbein & Pettijohn, 1938), it follows that dis-
posing of the water presents a greater problem than obtaining it. This is
particularly true when the apical spines maintain an opening to the surface.
The mechanics of the circulatory system of the burrow of M. ventricosa
are, therefore, easier to understand if they are thought of as comprising a
system for eliminating water from the burrow, rather than obtaining it.
The fascioles play an important role in the circulation of respiratory
currents within the burrow, and a description of their function is essential
to an understanding of the burrowing habits of echinoids. Fascioles are
comprised of numerous, small, modified spines calIed clavulae. These
tightly packed spines form narrow bands. In M. ventricosa, one band en-
circles the petaloid area on the dorsal surface (the peripetalous fasciole),
and another band encircles the epiplastronal area under the anal system
(the subanal fasciole, which is incomplete in adult M. ventricosa). Each
clavula has a thin shaft with two bands of cilia along each side and a bulb-
1969] Chesher: Biology of Meoma ventricosa 105
shaped distal portion which secretes a considerable quantity of mucus. The
mucus adheres to sand grains and to the tips of the clavulae; thus are formed
one continuous mucous "wall" that surrounds the petaloid area, and an-
other that surrounds the epiplastronal area. These walls of mucus extend
to the walls of the burrow and divide it into three main sections; a res-
piratory section encircled by the mucus of the peripetalous fasciole, an
anal section isolated by the mucus of the subanal fasciole, and the remain-
ing section, the locomotory section, where the spines used for movement
and the tube-feet used for feeding are located. Water, propelled by the cilia
of the clavulae, is moved from the petaloid area into the locomotory area
where most of it drains out into the surrounding sand. Residual water is
removed in the subanal drainage area. This process is shown diagram-
matically in Figure 14.
When the water passes through the peripetalous fasciole, some of it
stays close to the test and flows over the ambitus to the ventral side. A
large proportion, however, is diverted upwards, away from the test. The
upward flow seems to be controlled by currents approaching the fasciole
from the locomotory area, propelled by ciliary action along the lower parts
of the spines. It is possible, however, that the fasciole may also contribute
to the upward direction of these currents. The resultant upwardly directed
current causes an area of drainage between the fasciole and the ambitus.
Two other major drainage areas in the locomotory section of the burrow
occur along the anterior ambitus and the median section of the plastron
where opposing currents flow together. Burrowing movements in these
areas probably increase the drainage facility of the burrow.
The subanal fasciole provides the mechanism for ridding the burrow of
surplus water, a problem that is apparently more serious for the juveniles,
which burrow deeply and continuously in the sand. Water flows into this
fasciole from the locomotive section of the burrow. Water approaching
the fasciole from inside the fasciolar area directs the flow of water away
from the test into the disturbed sand through which the animal has just
passed.
The adult urchins, which migrate to the surface of the sand at night and
live near the surface of the sand during the day, have an incomplete sub-
anal fasciole, and drainage from this area is less than in juveniles, which
remain buried at all times (Fig. 3,A). Juveniles have a well-defined sub-
anal fasciole and construct two small drainage tunnels behind them as they
move through the sand. These tunnels are constructed by the joint effort
of the spines within the subanal fasciole and the subanal tube-feet and aid
drainage of the burrow by increasing the surface area of the burrow. The
adult urchins do not construct drainage tunnels, and the subanal fasciole
is degenerate as are the subanal tube-feet.
106 Bulletin of Marine Science [19(1)
A large portion of the staff and student body of the Institute of Marine
Sciences contributed in various ways to the completion of this study, and
I offer thanks to each of them. In particular, I thank Dr. F. M. Bayer for
his supervision and direction of this study and Dr. H. B. Moore for his
advice and contributions. Dr. and Mrs. W. A. Starck deserve special thanks
for their continued help with the field operations. The urchins of Colombia
were studied through the generosity of the Corporaci6n Aut6noma Regional
de los Valles del Magdalena y del Sinu, Bogota, Colombia. The M. ven-
tricosa of Panama were studied during ecological comparisons of the trans-
isthmian echinoids sponsored by the Smithsonian Institution.
The major portion of this study was conducted with the support of
National Science Foundation Grant GB 2037. Part of the manuscript was
revised during an N.S.F. Postdoctoral Fellowship at the Museum of Com-
parative Zoology, Harvard University. The statistical analysis of the growth
data was supported by a Milton Fund grant to Dr. H. B. Fell.
1969] Chesher: Biology of Meoma ventricosa 107
SUMARIO