FHE NAUTILUS
Volume 107, Number 1
March 24, 1993
ISSN 0028-1344
A quarterly devoted
to malacology.
APR
1 19S3
^
p^^
^^^
iX.:,
^
EDITOR-IN-CHIEF
Dr. M. G. Harasewych
Division of Mollusks
National Museum of
Natural History
Smithsonian Institution
Washington, DC 20560
ASSOCIATE EDITOR
Dr. R. Tucker Abbott
American Malacologists, Inc.
P.O. Box 2255
Melbourne, FL 32902
CONSULTING EDITORS
Dr. Riidiger Bieler
Department of Invertebrates
Field Museum of
Natural History
Chicago, IL 60605
Dr. Robert T. Dillon, Jr.
Department of Biology
College of Charleston
Charleston, SC 29424
Dr. William K. Emerson
Department of Living Invertebrates
The American Museum of Natural
History
New York, NY 10024
Dr. Robert Hershler
Division of Mollusks
National Museum of
Natural History
Smithsonian Institution
Washington. DC 20560
Dr. Richard S. Houbrick
Division of Mollusks
National Museum of
Natural History
Smithsonian Institution
Washington, DC 20560
Mr. Richard I. Johnson
Department of Mollusks
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138
Dr. Aurele La Rocque
Department of Geology
The Ohio State University
Columbus, OH 43210
Dr. James H. McLean
Department of Malacology
Los Angeles County Museum ol
Natural History
900 E.xposition Boulevard
Los Angeles, CA 90007
Dr. Arthur S. Merrill
% Department of Mollusks
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138
Ms. Paula M. Mikkelsen
Harbor Branch Oceanographic
Institution, Inc.
Ft. Pierce, FL 33450
Dr. Donald R. Moore
Division of Marine Geology
and Geophysics
Rosenstiel School of Marine and
Atmospheric Science
University of Miami
4600 Rickenbacker Causeway
Miami, FL 33149
Dr. Gustav Paula\
Marine Laborator\-
University of Guam
Mangilao', (^.uarn 96923
Mr. Richard L. IVtit
P.O. Bo.\ 30
North Myrtle Beach, SC 29582
Dr. Edward J. Petuch
Department of Geology
Florida ,\tlantic University
Boca Raton, FL 33431
Dr. David H. Stansbery
Museum of Zoology
The Ohio State University
Columbus, OH 43210
Dr. Ruth D. Turner
Department of Mollusks
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138
Dr. Geerat J. Vermeij
Department of Geology
University of California at Davis
Davis, CA 95616
SUBSCRIPTION INFORMATION
The subscription rate per volume
is US $25.00 for individuals, US
$40.00 for institutions. Postage
outside the I nited States is an
additional US $5.00 for surface
and US $12.00 for air mail. All
orders should be accompanied by
payment and .sent to: THE
N.4UTILUS, P.O. Box 7279, Silver
Spring, MD 20907-7279, USA.
Change of address: Please inform
the publisher of your new address
at least 6 weeks in advance. All
communications should include
both old and new addresses (with
zip codes) and state the effective
date.
THE NAUTILUS (ISSN 0028-
1344) is published quarterly by
Trophon Corporation, 8911 Alton
Parkway, Silver Spring. MD 20910.
Second Class postage paid at Silver
Spring, MD and additional mailing
offices.
POSTMASTER: Send address
changes to: THE NAUTILUS
P.O. Box 7279
Silver Spring, MD 20907-7279
TH Et^NAUTI LUS
CONTENTS
Volume 107, Number 1
March 24, 1993
ISSN 0028-1344
I IPD/\pv '
Limacosphaera, an I'nusual Mesogastropod (Lamellariidae)
Larva of the Weddell Sea (Antarctica) , 1
)_ Hole, Mass. j
The Reproductive Anatomy of Philomycus sellatits
Hubricht, 1972 and Philomycus virginiciis Hubricht, 1953
(Gastropoda: Philomycidae) 9
Two Confusing Indo-Pacific Cerithiids 14
Rediscovery of Tiirbinella thersites Reeve, 1847, with
Notes on its Taxonomic Position (Gastropoda:
Fasciolariidae) 24
The Rediscovery, Morphology, and Identity of Conus
emersoni Hanna, 1963 29
The Benthic Mollusk Faunas of Two Contrasting Reef
Paleosubenvironments: Falmouth Formation (late
Pleistocene, Last Interglacial), Jamaica 33
Klaus Bandel
Stefan Hain
Frank Riedel
Henry Tieniann
H. Lee Fairbanks
Richard S. Houbrick
R. N. Kilburn
John K. Tucker
James H. McLean
Stephen K. Donovan
D. T. J. Litllewood
THE NAUTILUS 107(l):l-8, 1993
Page 1
Limacosphaera, an Unusual Mesogastropod (Lamellariidae)
Larva of the Weddell Sea (Antarctica)
Klaus Bandel
Gediogiseli-Puliiontiilogisclies
Institut und Museum
Universit) of Hamburg
BundesstraBe 55
2000 Hamburg 13
GERMANY
Slefan Main
A If red- Wegener- Institut fiir
Polar- und Meeresforschung
CoiumbusstraBe
2850 Bremerhaven
GERMANY
Frank Riedel
Geologiseh-PalaoiUulogisches
Institut und Museum
University of Hamburg
BundesstraBe 55
2000 Hamburg 13
GERMANY
Henry Tiemann
Zooiogisches Institut und Museum
University of Hamburg
Martin-Lutlier-King-Piatz 3
2000 Hamburg 13
GERMANY
ABSTRACT
Marsciiiopsis conica Smith. 1915 and M mollis (Smith. 1902)
have pianktotrophic larvae that are unique among gastropods.
They cover the larval shell with a lacunous muscular mantle
that can change its volume by interaction of body fluid and
muscle activity This limacosphaera larva is found in Antarctic
waters and represents the most complex larval strategy within
the "echinospira-group' . Notes on the biology and anatomy,
including histology, of this larva are presented.
Ketj ivords: Lamellariidae, Marseniopsis. larvae, Antarctica.
introduc;tion
The majority of benthic invertebrates of high-Antarctic
seas brood their young or provide postspawning parental
care rather than produce free-Uving larvae (Mileikovsky,
197L Picken, 1980). True pianktotrophic pelagic larvae
are extremely rare in the 200 to 600 m deep high-Ant-
arctic shelf areas.
Plankton samples taken by the R/V polarstern in the
eastern Weddell Sea during several late winter to late
summer cruises yielded only two meropelagic larvae of
benthic gastropods (Piatowski, 1987; Boysen-Ennen,
1987). Both of these larval types were regularly found.
One of these is Capulus subcompressus Pelseneer, 1903,
which had first been observed by Pelseneer (1903) and
is described in detail by Bandel and Hain (in prepara-
tion). The second lueropelagic larvae was first observed
by Simroth (1914) in material from the Davis Sea. Sim-
roth noted anatomical similarity to a planktonic gastro-
pod from the deep sea areas of the Indian Ocean, which
he described earlier as Limacosphaera macdonaldi (Sim-
roth, 1908).
This animal is a spherical, transparent, voluminous
gastropod with two openings: one to allow head and foot
to come out of the shell and the other situated on the
opposite side of the sphere (Simroth, 1908, 1914). Simroth
called the muscular inantle covering the larval shell the
deutoconcha of the limacosphaera. The posterior pore,
which he (1908) called the shell tunnel (Schalengang),
connects the sea water with a cavity surrounding much
of the thin organic shell. Simroth (1914) also suggested
the presence of glandular cells and muscle fibers in the
voluminous deutoconcha that surrounds the shell. He
noticed that the Antarctic forms represented larvae rath-
er than adult gastropods as he had originally assumed
when describing Limacosphaera as a new genus of pe-
lagic gastropods (Pteropoda). Simroth (1914) correctly
placed these larvae into the ontogeny of members of the
"echinospira-group" in the genus Marseniopsis. The term
limacosphaera was retained, now to describe a very pe-
culiar and characteristic larva.
Simroth (1908, 1914) considered that the limaco-
sphaera swam with the aid of their large velum. Another
mode of sw imming, mainly by hydrostatic effects, was
considered the likely mode of propulsion in the water
column by Jevdonin and Minichev (1975).
These authors observed developing spawn of Marsen-
iopsis conica and foimd that the mantle of the embryo
envelops the shell prior to hatching from an egg mass
found in excavations of tunicate surfaces. Jevdonin and
Minichev (1975) also noted that the mantle did not fuse
completely but left an open shell pore. They seem not
Page 2
THE NAUTILUS, Vol. 107, No. 1
Figures 1, 2. Adult animals ol I. Marseniopsis conica Smith, 1915, and 2. M. mollis (Smith, 1902). Dorsal views on left, showing
mantles that are fused ahove the shell and cannot be retracted. Neutral \ lews on right, show foot, parts of the head \\\ illustrations
drawn irom fi.\ed animals. Scale bar = 5 mm for figure I, 10 mm for figure 2.
to have been aware of the earlier cie.scriptions of these
larvae by Simroth (1908, 1914). Jevcloiiin aiul Miiiichev
(1975) suggested that larvae with a size of about 20 mm
in diameter were nearly ready to metamorpho.se. Their
descriptions suggest that their observations had been car-
ried out on preserved material and they most probably
did not .see living larvae or embryos.
We provide a new description of the histology of these
larvae integrated with observations on the li\iiig forms
and their metamorphosis.
MATERIALS AND METHODS
During the cruises polarsirkel 80/81, polarstern ant
I, ANT III/3, ANT V/S and ant \'II/4 larvae of the li-
macosphaera type were caught at about 100 stations in
K. Bandel et al., 1993
Page 3
— grounding linie
Icnntinenlal boundary)
W •" E
Figure 3. Map indicating location of the Wedell Sea and the
area where the larvae of Marseniopsis were caught.
neritic \\ aters above continental shelf areas of the eastern
Weddell Sea (figure 3). Samples were taken with various
plankton gears (mesh sizes of 0.335-4.5 mm) in upper
water layers (300-0 m) and preserved in buffered for-
malin.
Six larvae were kept alive for six weeks during the
expedition PS ant V/3. During this period they were
fed two times with a diatom suspension. Only one of the
larvae survived and metamorphosed four weeks after
capture (November 1986). It was fixed in 70% ethanol.
During the expedition PS ant VII/4 (January to March
1989) 42 living limacosphaera from RMT samples were
photographed for size measurements and transfered in-
dividually to small plastic aquaria (running water system,
temperature adjusted to 0 °C). The larvae were fed once
a week with a suspension of Antarctic diatoms. After 8
and 13 months the photographic measurements of larvae
or juveniles were repeated.
The larval shell, the thick, whitish to semitransparent
tissue covering it (deutoconcha), and the juvenile shell
were studied on 10 critical-point dried specimens using
a SEM.
In May 1989, six living larvae were fixed for histolog-
ical studies in 4% seawater-formalin or 2.5% seawater-
glutaraldehyde. One limacosphaera fixed in formalin was
dehydrated using isopropanol. It was then treated with
benzylbenzoate, embedded in paraplast, serially sec-
tioned (7 ytm) and stained with Helioechtrubin BBL/
Acidgreen 5/Acidorange 10 after Halmi (Adam & Czi-
hak, 1964).
To detect lipids whole larvae without the deutoconcha
as well as parts of the deutoconcha were treated using
the oil-red-niethod (Romeis, 1968). These samples were
embedded in glycerin and examined with a light mi-
croscope.
Another limacosphaera fixed in glutaraldehyde was
osmicated in 1% osmium tetroxide, dehydrated in ace-
Figure 4. Larva of Marseniopsis cf. mollis. Schematic drawing
from several photr)graphs of living specimens. One quarter of
the shell-covering tissue (deutoconcha) is removed, d, deuto-
concha; e, eye; ed, epidermis; f, foot; s, shell; si, siphon; st,
apical tube; t, tentacle; v, velum.
tone and embedded in Spurr's fluid. Sections were cut
at ca. 80 nm to 1 ^m using an ultramicrotome (Reichert).
The 80 nm sections were stained with uranyl acetate and
lead citrate, then examined and photographed using a
TEM (Zeiss). The 1 ^m sections were stained with To-
luidine Blue and Pyronin and examined with a light
microscope.
Voucher material is deposited at the Zoological Insti-
tute and Museum (L'niversity of Hamburg) (cat. no. M
885).
RESULTS
The Living Larva
Description: The shape of the limacosphaera (figures
4-6) is globular with an overall diameter ranging from
1.8 to 20 mm. The actual larval shell is covered by the
deutoconcha. Some of the larvae have granular deposits
in the outer layer of the deutoconcha. On the ventral
side of the larva (figure 6) there is a slit-like opening for
the foot and the four large velar lobes. This opening is
extended anteriorly by a siphon-like, semicircular groove.
A second, small, tunnel-like excavation extends from the
apical region of the shell to the surface of the deutocon-
cha. Observations on living larvae sorted from plankton
samples show that both openings can be closed and that
the deutoconcha produces much hyaline mucus.
During metamorphosis the shell is still barely calcified
(figure 7), becoming solidly calcified right after meta-
morphosis (figure 8). The larval shell reaches 2.3 whorls
with fine spiral threads on the first 1.5 whorls. Earliest
growth lines are visible after 0.8 whorls (figure 9 arrow).
In well fed larvae, the greenish visceral mass can be seen
through the shell and the deutoconcha.
All studied larvae had a well developed foot. The eyes
are at the base of the tentacles. In contrast to Simroth's
original description of Limacosphaera macdonaldi from
Page 4
THE NAUTILUS, Vol. 107, No. 1
Figures 5, 6. Living larvae ul A/(j),Miii(>/;.si.v cj. mollis. 5.
Dorsal view. 6. Ventral view, with foot, velum, tentacles and
eyes visible. Scale bars = 5 mm for liolh pliotograjihs
the tropics, the operculum is absent in the Antarctic
specimens.
Biology
Deposition of egg capsules in the tests of compound as-
cidians is known for temperate lamellarian species (Fret-
ter & Graham, 1962). Antarctic species have the same
spawning habit (Jevdonin & Minichev, 1975; P. Dayton,
personal communication). One compound ascidian with
appro.xiinately 20 egg capsules was obtained in February
1989. Although maintained in an aquarium, the eggs
failed to develop.
Figures 7-9. Shell of Marseniopsis cj. mollis. 7. During meta-
morphosis, the shell is barely calcified and therefore, flexible.
8. .Alter metamorphosis, shell solidly calcified and lias reached
2 4 whorls. 9. Same specimen as in fig, 8. P'irst growth lines
(arrow) are visible at 0 7 whorls of the shell The sjjiral threads
end at 1.4 whorls. Scale bars = 1 mm in all photographs.
Larval size, as well as the time and location of sam-
pling, indicates that hatching of larvae occurs from late
winter to austral summer in the shelf areas of the eastern
Weddell Sea.
The smallest larvae found have a diameter of 1.8 mm.
We suggest that this size is reached shortK after the
larvae have hatched and risen from the l)ottoni. Aquar-
ium observations indicate that the larvae rise by buoy-
ancy. The large velar lobes are not used or have only
secondary function. The buoyancy control mechanism is
uiikuow n.
In the northern shell areas of the Weddell Sea the
amount of food (0.07 ixg chlorophyll -a/liter) is more
K. Bandel et al., 1993
Page 5
Figure 10. ScluMiuitir draw iiig ot the ajMciil lulx' (shell tuiiiu-lj
of Marseniopsis cf. mollis, c, cilia; cc, collagenous cell; cf,
collagen fibers; gc, gland cell; mf, muscular fibers; mv, micro-
villi; n, nuclei; s, shell; st, apical tube
limited even during phytoplankton blooms. In open wa-
ters or polynyas (ice free areas), the phytoplankton con-
centration barely reaches 1.5 ng chi a/liter. It is sus-
pected that growth of larvae in these areas takes a much
longer time.
The total Hpid content of larvae caught in February
(end of austral summer) is low (3.5% dry weight) in
comparison to other planktonic organisms. Seventy per-
cent of these lipids are triacylglycerols (Hagen, 1988).
Although fed only twice, one limacosphaera caught in
October 1986 metamorphosed four weeks later. During
PS Ant VII/4 small larvae with a diameter of 3.9 mm
began to metamorphose in the aquaria after 10 to 24
hours, while specimens of 10 mm diameter remained in
the larval stage for up to 8 weeks. One specimen caught
in the end of February 1989 was still in the larval stage
when it died in August 1990.
Predatory pressure on the larvae seems to be very low.
The shell-covering tissue of juveniles and adults is known
to produce acidic secretions.
HISTOLOGY
Observations with Light Microscope
The deutoconcha of the limacosphaera consists of tissue
that is composed of an outer epidermis, a central, cav-
ernous connective tissue, and an inner epidermis cov-
ering the shell.
The outer epidermis is composed of a single layer of
cells with flattened nuclei and immersed, large, light,
glandular cells with a single nucleus. Below it is a vo-
luminous, spacious connective tissue, consisting of col-
lagenous and muscular fibers with few cells suspended
in it. The inner epidermis is a single layer of cells sup-
ported by connective tissue and muscle fibers. These cells
are stretched in length, and their nuclei are even more
flattened than those of the outer epidermis.
The deutoconcha of the limacosphaera is connected
to the inner mantle of the apertural region of the shell
by an especially strong muscular bridge of tissue. The
outer mantle rests on the shell near the aperture and
forms the connection to the inner mantle covering the
visceral mass and the pallial cavity. Outer and inner
mantle are "divided" from each other by the periostraca!
gland zone characterized by the presence of many nuclei.
On the apical side of the deutoconcha the e.xternal
mantle is pierced by a tube-like canal (figure 10, st)
connecting sea water and shell surface. This canal is lined
with a rugged, ciliated epithelium (figure 10, c) that
continues along the inner mantle for about 0.4 mm, to
form a small cavity between shell and mantle (figure 10).
The tissue below the ciliated epithelium of the canal
contains an agglomeration of large light epithelial glan-
dular cells.
The iimer side of the deutoconcha connects to the
organic shell that is composed of a double-layered peri-
ostracum. This shell in its apical portion is filled with the
visceral mass containing the large digestive gland. Its
diverticula consist of large endodermal cells with basal
nuclei and small cavities. The apical portion of the di-
gestive gland is glandular and contains three different
types of stored substances. The most basal portion con-
tains proteins, somewhat above lie a large number of
spherical bodies. Scale-like storage material that is weak-
ly aeidophilous is suspended between these.
The spherical bodies (6-19 ^m in diameter) are lipids
(analysed by oil-red method). The spheres of lipids were
arranged like strings of pearls of different length
throughout the entire digestive gland of a larva caught
in February 1989 during the phytoplankton bloom.
The outer surface of the deutoconcha was also tested
for the presence of lipids, but was found to lack them.
This test was repeated on another larva that had been
caught prior to the beginning of the phytoplankton bloom
(mid-November 1986). It lacked lipids in either the man-
tle or the digestive gland.
Of the other organs of the larva, the nerve ring proved
to be quite large. The eyes were well-developed, with
lens-like light collectors. The foot is intensely ciliated,
almost to the same degree as the large velum. The radula
is well developed and functional.
Observations with Electron Microscope
The epidermis of the deutoconcha (figure 11) is covered
by a 2 ^m high rim of microvilli. These microvilli are
differentiated into a narrow basal portion with many
filaments and a light, partly bubble-like expanded upper
portion.
These "bubbles" are secretory vesicles, usually linearly
arranged and attaining 2-4 times the width of the mi-
crovilli. They are produced by glandular cells at the base
of the rim of microvilli and are secreted at the outside
of the rim. "Bubbles" open up at the outside of the rim
as could be seen in some sections.
The surface of the rim of microvilli is covered with
fine fibrillar material containing small dark granules.
Page 6
THE NAUTILUS, Vol. 107, No. 1
Between microvilli, larger vesicles with a small internal
membrane are often present. Some larger vesicles are as
high as the microvilli and are connected to the glandular
vesicles of the epidermis.
The epithelium of the deutoconcha consists of a single
layer of flattened cells resting on a distinct basal mem-
brane. The cells are up to 10 ^m in size, and are con-
nected to each other by apical contact zones. Cell bound-
aries are strongly folded and intercalated.
Some nuclei are strongly heterochromatic, others are
large, less densely packed and of irregular shape. All cells
hold large, light vesicles often containing membranous
extensions. These vesicles form a loosely connected sys-
tem that is in contact with the vesicles extruded into the
rim of microvilli. Newly extruded vesicles hold dense
granules.
A voluminous connective tissue is present below the
epidermis and fills the space between inner and outer
layer of the deutoconcha. Within a homogeneous basal
substance, individual, transversally striped fibers of col-
lagen connect the basal membrane of cells within the
tissue to the epidermis.
Large cavities lie within the connective tissue near the
epidermis. These cavities are lined with a thin, conspic-
uously foamy cellular layer resting on a basal membrane.
The nuclei are flattened and elongated.
Nerves, consisting of several axones within a glial cell,
and smooth muscular fibers with connections to the col-
lagen fibers are distributed within the connective tissue.
The intermediate layer between outer and inner ep-
ithelia (gelatinous layer, Simroth, 1908) is shown to con-
tain a network of collagenous fibers connected to muscle
and epithelial cells. This layer functions together with
interconnected blood lacunae as a hydroskeleton that can
change the outline, shape and width of the limacos-
phaera.
The apical tube (figures 4, 10, st) of the deutoconcha
is ciliated, in contrast to the remaining surface of the
mantle. This ciliation continues into the small cavity at
the proximal end of tube above the shell. This cavity is
less extensive than assumed by Simroth (1908) and does
not surround the entire shell surface.
Simroth (1908) noted a folded surface of the shell,
which is an artifact of preservation.
A rim of microvilli on a unilayered epithelium sur-
rounding a structured connective tissue represents a large
surface for resorption as well as for secretion by many
glandular cells that come together in canals opening into
pores. The type of secretion produced by these glands is
unknown. The secretion expelled between the microvilli
probably represent neutral mucus material.
DISCUSSION
The larvae metamorphosed into juveniles that had either
a smooth surface or a tuberculated body surface. Both
juvenile forms were analysed by gel-elect rophoresis (ID-
UISK-SI)S-PA(;E) at the Alfred-Wegener-in.stitut (Dr.
T. Stadler). The results were compared to gel-electro-
Bum
cb
Figure 11. Transmission Electron Micrograph of the deuto-
concha epidernii.s of Marscniopsis cf. mollis, bm, ba.sal mem-
brane; ci), cell boundary; ct, connective tissue; mv, microvilli;
n, nucleus.
phoresis (same method) analyses of adult Marseniopsis
conica and Marseniopsis mollis. Differences in the data
indicate that the predominantly smooth juveniles re-
present M. mollis and the tuberculated ones M. conica.
I'hus it can be concluded that both species have very
similar larvae. It was not possible to section and electro-
phoretically analyse the same individual.
The course of a limacosphaera's development depends
on a variety of factors, including currents in the upper
water layers, melting of sea ice, and patchyness of phy-
toplankton production. Larval survival in captivity for
1.5 years indicates that a limacosphaera could remain in
the plankton for over a year if conditions were not fa-
vorable for metamorphosis.
Analyses of lipid content suggest that larvae either use
most food-energy for growth or they store energy in form
of other biochemical metabolites {e.g., proteins). The
vertical distribution and the stomach contents of larvae
caught during mid-October at 72°S indicate that they
feed under the sea ice where the phytoplankton concen-
tration (0.07 ng chl- a/liter) was seven times larger than
in deeper water (Scharek, personal communication). Due
to the southwcsterK water currents near the surface, the
majority of larvae will be transported to southern parts
of the study area, where very high phytoplankton con-
centrations (50-150 ^g chla/liter) were measured di-
rectly under the sea ice in October 1989. By utilization
of this enormous food resource, larvae living in that area
could grow very rapidly. Due to the fact that the macro-
zoobenthos community of the southern Weddell Sea is
totally different from that of the eastern Weddell Sea
(Voss, 1988) and therefore not suitable for survival of
beiithic stages of Marseniopsis, there nuist be a way of
K. Bandel et al., 1993
Page 7
returning pelagic stages (metamorphosing larvae, juve-
niles or adults) back to the hatching grounds. This may
be accomplished by counter-currents nuining near the
bottom.
Piatkowski (1987) and Boysen-Ennen (1987) reported
the absence of larvae in the oceanic domain between the
tip of tlie Antarctic peninsula and the north-eastern part
of the Weddell Sea. Marseniopsis populations of the
peninsula are likely zoogeographically separated from
populations in the eastern Weddell Sea. Nevertheless
some of the larvae could be transported by eastward wind
drift ot surface water layers near the ice edge from
eastern-Antarctica to the Weddell Sea. Between Feb-
ruary and March 1983, the mean abundance of larvae
in the eastern Weddell Sea was 24 specimens per 1,000
m' (Boysen-Ennen, 1987), with maximum densities of
65 larvae per 1,000 m' in the southern part of the study
area (Piatkowski, 1987). Most larvae were found in water
layers of 200 to 50 m (63%) and 50 to 0 m (35%), only
2% were reported from 300 to 200 m depth (Boysen-
Ennen, 1987). The major function of the deutoconcha is
that of a buoyancy organ. Altering the volume of the
outer mantle may vary the speed of sinking of the li-
macosphaera. According to Stokes formula (Tiemann &
Betz, 1979) this speed depends on the specific weight,
which again is a function of the diameter of the lima-
cosphaera. Histological analyses of the limacosphaera
showed that the deutoconcha contains very little organic
material and is thus very close to the density of sea water.
The deutoconcha can enlarge the diameter and thus the
volume of the larva without decreasing its weight in the
water. A general calculation indicated that the volumi-
nous outer mantle of the limacosphaera decreases the
speed of its sinking to about one half of what it would
be without such a cover around the shell
The ability to float was observed in aquaria for all
developmental stages. Even large adult specimens could
float near the bottom after disturbance (diving obser-
vations at the Antarctic peninsula, W. Wagele, personal
communication).
The function of the deutoconcha's apical tube and
cavity is still unknown. It is evident that water can be
pumped into the cavity and expelled out by ciliary cur-
rents as well as by muscular movements of the deuto-
concha. It is also evident that glands can secrete sub-
stances into the lumen of tube and cavity.
Histological examination of the limacosphaera for the
most part confirms the observations of Simroth (1908,
1914). His assumption that muscular cells are present
within the deutoconcha is substantiated.
The prolific mucus secretion of the entire deutoconcha,
as well as its large size may serve as defensive mechanisms
against carnivorous planktonic groups like copepods or
krill.
The shell size of hatching larvae can only be inter-
preted from the embryonic and the larval shell (figure
8). There are two possibilities. The normal case would
be that the embryo hatches shortly after the Ijeginning
of growth lines (the mantle becomes free from the shell)
(figure 9, arrow). This happens after 0.8 whorls and a
diameter of the shell of 0.65 mm.
The second possibility is that the embryo does not
hatch before reaching 1.4 whorls (with spiral threads).
The shell then has a diameter of 1.5 mm. To reach this
size the embryo v\ould have to feed on extra yolk.
It is proposed that the embryo hatches when the shell
has reached 1.4 whorls. At this time tlie spiral threads
disappear. This is probably the consequence of the man-
tle fusing above the shell. There is no difference in sculp-
ture between the subsequent part of the larval shell and
the teleoconch of the investigated species of Marseniop-
sis.
ACKNOWLEDGEMENTS
Tliis study was carried out with financial support of the
Deutsche Forschungsgemeinschaft (DFG) in the frame
of the Antarctis-research grant and science grant Ba 675/
6-1 We have been greatly aided by our colleagues of
the Alfred-Wegener-Institut (Brenierhaven), Zoological
and Geological-Paleontological departments of the Uni-
versity of Hamburg. We are grateful to Dr. T. Stadler
who carried out the gel-electrophoresis analysis. Dr. M.
G. Harasewych greatly improved st\le and grammar of
the manuscript.
To all persons and institutions involved, we express
our sincere thanks.
LITERATURE CITED
Adam, H. and G, C^lzihak 1964 Arbeilsniethoden der mak-
roskopischen Anatomie. Ein Laboratoriumshandbuch fiir
Biologen, Mediziner und technische Hilfskrafte. Gustav
Fischer Verlag, Stuttgart, 583 p.
Boysen-Ennen, E 1987 Zur Verbreitung des Meso- und Mak-
rozooplankton.s im Oberflachenwasser der Weddell See
(Anlarktis) Berichte zur Polarforschung 35:1-126
Fretter, V. and A, Graham. 1962. British prosobranch mol-
luscs. Ray Society, London. 7.55 p.
Hagen, W. 1988. Zur Bedeutungder Lipide im antarktischen
Zooplanktnn. Berichte zur Polarforschung 49:1-129
Jevdonin, L. A. and J. S. Minichev. 1975. Adaptations of
pelagic gastropods. Malacological Review 11:75
Mileikovsky, S. A. 1971. Types of larval development in ma-
rine bottom invertebrates, their distribution and ecological
significance: a re-evahiation. Marine Biology 10:193-213.
Pelseneer, P 1903 Resultats du vo\age du S Y. "Belgica"
1897-1899. Rapports Sci. Zoologie, Mollusques (Amphi-
neures, Gasteropodes et Lamellibranches) Anvers, 85 p.
I'lalkovvski, L'. 1987. Zoogeographische Untersuchungen und
Gemeinschaftsanalysen an antarktischem Makroplankton.
Berichte zur Polarforschung 34:1-138.
Picken, (; B 1980. Reproductive adaptations of Antarctic
benthic invertebrates. Biological Journal of the Linnean
Society 1467-75.
Ronieis. B. 1968. Mikroskopische Technik. Oldenbourg Ver-
lag, Mijnchen, 16. .\uOage, 7.57 p.
Simroth, H. 1908. Gastropodenlaiche \ind Gastropodenlarven
der Deutschen Tiefsee-Expedition In: C Chun (ed). Wis-
senschaftl Ergebnisse der Deutschen Tiefsee-Expedition
Page 8
THE NAUTILUS, Vol. 107, No. 1
auf dem Dampfer -Valdivia' 1898-1899, Band IX:365- Tiemann, H and K -H Bctz 1979 Elutriation: theoretical
,,Q considerations and methodological improvements Marine
Simroth H 1914. Pelagische Gastropodenlarven der deut- ecology. Progress Series 1277-281.
schen Siidsee-Expedition 1901-1903. /;i: Drygalski, E.v Voss, J. 1988 Zoogeographie and Gemeinschaftsanalyse des
(Hrsg.). Deutsche Sudpolar-Expedition, Zoologie Band VII Makrozoobenthos des Weddellmeeres (Antarktisj. Berich-
15143-160. ''' ^"'' Polarforschung 45:1-14.5.
THE NAUTILUS 107 1 9-13. 1993
Page 9
The Reproclucti\ e Anatom\ of Philomijcus sellatus Hubricht. 1972
and Philomijcus virginicus Hubricht. 1953 (Gastropoda:
Philom\cidae)
H. Lee Fairbanks
Penns\l\ ania State Universits-
Monaca. PA 15061 USA
ABSTRACT
Specimens of Philomycus sellatus and P virginicus were col-
lected at or near their t> pe localities. Their reproducti\ e anat-
om\ is described and compared with that of other species of
Philomycus. The penial anatom\ of P sellatus is unique in
ha\ing two constrictions in the penial wall; P virginicus is
distinguished by a circular pustulose ridge that separates the
distal penis from its atrial opening.
Key Words: SloUusca; Gastrojxxla; landsnail: slug; Philomy-
cldap: Philornvcj'- reproductive anatomy
LVTRODUCTION
.\1] species of the terrestrial slug family Philomycidae
ha\e a mantle that co\ers the entire bod> of the slug.
Three genera of this family. Philomycus. Pallifera. and
Megapallifera. occur in the United States, with most
species being limited to the eastern and south-central
areas of the country . Few original descriptions pro\ide
anatomical data. Neither the description of Philomycus
virginicus Hubricht. 195.3 nor of P. sellatus Hubricht,
1972 discusses or figures an> part of the reproductive
anatomy. Based up>on the mande pattern. Hubricht 19721
compared Philomycus sellatus to P ccirolinianus Bosc.
1S02!. but made no comparisons between P. virginicus
and other species of Philomycus. References to the re-
producti\e anatomy of P. virginicus are limited to two
articles by Branson il96S. 1969; that compared P. vir-
ginicus with P. bisdosus Branson. 196S. Neither contain
figures or measurements. The reproductive anatomy of
P. sellatus has not pre\iousl\- been studied.
The goals of the present study were to describe the
anatomy of the reproductive systems of Philomycus vir-
ginicus and P. sellatus. and to compare them with those
of other sp>ecies of Philomycus.
METHODS AND MATERI.ALS
Sf)ecimens of Philomycus virginicus were collected on
19 May 19S9 near nulepost 47 north of Skyline Drive.
Shenandoah National Park, Madison County , \irginia.
under the loose bark of logs and dead trees, eleiation
appro.ximately 1110 meters. This station was approxi-
mately O.S kilometers east of the tvpe locafity for this
species. Specimens of P. sellatus were collected on 17
May 19S9 from its t>p)e locahty (2.7 kilometers northeast
of Princeton, along Highwa> 65. Jackson County . .Ala-
bama; in beede galleries in rotten logs, elevation ap>-
pro.ximateK 215 meters. For comjjarative purposes, the
following sp>ecie5 were also e.xamined: P. carolinianus
collected in Berkele> Cotmty . South Carolina approxi-
mateK 2-5 kilometers from Charleston, the designated
t>pe iocaht) ,Pilsbry, 1945:754; on 22 June 19SS; P.
bisdosus collected from Breaks Interstate Park. Dick-
enson Coimty. Virginia the type locahty on 22 May
1987: P. flexuolaris collected from Breaks Interstate Park,
Dickenson County. Virginia on 22 Ma> 1987: and P.
togatus collected near Riceville. PittsyKania County,
Virginia on 20 May 1987.
External characteristics of the specimens were com-
pared with the original sp)ecies descriptions to ensure
correct identification. All specimens were drowned in
distdled water and dissected immediately. The repro-
ducti\ e sy stems were removed and the atrium and penis
opened to examine their internal anatom\ . Subsequendy
all material was preserved in 70"^ ethanol. DrowTiings
and dissections were conducted during the month of Jvme
in each > ear to reduce any differences that may be at-
tributable to the stage of the life c>cle. The sjjecimens
were maintained in terraria for approximately two weeks
prior to dissection.
E^ch complete reproductive system was spread out in
a f)etri dish and then projected- \ia an overhead projector,
and traced. The p>enial figures are tracings of photo-
graphic shdes of the opened organs. Voucher sp>ecimens
have been deposited in the Academ\- of Natural Sciences,
Philadelphia ANSP .Al>326 for Philomycus bisdosus,
.ANSP A1S327 for P virginicus. ANSP A13328 for P.
sellatus, and .ANSP Al-3329 for P. carolinianus}.
RESULTS
Four adult specimens each of Philomycus sellatus and
of P. virginicus were collected and used in the studv.
Page 10
THE NAUTILUS, Vol. 107, No 1
/
Figures 1-4. Mantle patterns of living slugs. 1, 2. Philomtjcus
virginicus 3, 4. Philomtjcus sellatiis. Scale bar = 20 mm.
External characteristics of all four specimens of each
species (figures 1-4) agreed with their type descriptions.
The gross reproductive anatomy was similar for all spec-
imens of a species. Measurements of selected reproduc-
tive organs are shown in Table 1.
Genitalia of Phibmycus sellatus Hubricht, 1972
(figures 6, 8)
Atrium glandular on distal half of outer surface, length
approximately 60'/o penial length. Vagina length ap-
pro.ximately 10% of penis. Spermathecal duct approxi-
mately same diameter as free oviduct, length nearly twice
that of free oviduct, slight taper toward spermatheca.
Spermatheca round. Dart sac (with dart) larger than
spermatheca. Penis diameter at proximal end nearly equal
to atrial diameter, gradual taper to diameter of vas de-
ferens at distal end; penial sheath covers approximateK
Figures 5, 6. Genitalia, 5. Philornycus virginicus. 6. Philo-
mtjcus sellatus. Scale bar = 10 mm. A: atrium, AG; albumen
gland, AR: accessory retractor, DS: dart sac, FO: free oviduct,
G: gonad, GP: genital pore, HD; hermaphroditic duct, P: penis,
PR: penial retractor, S: spermatheca, \': vagina, \'D: vas de-
ferens.
90% of penis. InternalK , distal third of penis with several
thin low folds, separated from remainder of penis by
constriction in penial wall; middle third with 3-4 large
pustulose ridges; proximal third with several thin non-
pustulose folds, separated from middle third b\ constric-
tion in penial wall. Vas deferens enters distal end of penis,
length approximately four times that of penis; diameter
of middle third half that of ends. Penial retractor muscle
maximum width 2-3 times diameter of distal end of
penis, length 40% of penis. Accessory retractor muscle
present, located on outer wall of atrium near its junction
with penis.
Genitalia of Philornijcus virginicm Hubricht, 1953
(figures 5, 7)
Atrium length approximately equal to that of penis, ex-
ternal surface of distal half glandular. Base of penis bulg-
es into atrium. N'agina short, approximateK 15% of penial
length. Spermathecal duct diameter greater than that of
free oviduct, slight taper toward spermatheca. Sperma-
theca round. Dart sac (with dart) smaller than sperma-
theca. Penis straight, basal diameter one third of length,
slight taper to junction with vas deferens; penial sheath
reaches to junction of penis with vas deferens. Internally,
H. L. Fairbanks, 1993
Page 11
Figures 7, 8. Internal penial anatomy. 7. Philomycus virgin-
iciis. 8. Philomycus setlatus. Scale bar = 10 mm. Cut surfaces
indicated by oblique lines in all drawings. A; atrium, DS: dart
sac, P: penis, PR: penial retractor, PS; penial sheath, VD: vas
deferens.
penis with 4-6 pustulose ridges, a basal pustulose circular
ridge sets off distal parts of the penis from its opening
into the atrium. Vas deferens encircles distal end of penis,
entering penis terminally. Vas deferens length approxi-
Figures 9, 10. Genitalia 9. Philomycus bisdosus. Scale bar =
10 mm. 10. Philomycus carolinianus. Scale bar = 5 mm. A:
atrium, AG: albumen gland, .\R accessory retractor, DS: dart
sac, FO: free oviduct, G: gonad, GP: genital pore, HD: her-
maphroditic duct, P: penis, PR: penial retractor, S: spermathe-
ca, V: vagina, YD: vas deferens.
mate!) 3.3 times that of penis, approximately same di-
ameter throughout its length. Penial retractor muscle
maximum width one third that of diameter of distal end
of penis but broader at its distal end, length approxi-
mately one third that of penis. Accessory retractor muscle
absent or not distinguishable from typical connective
tissues.
Table I. Measurements (mm) of selected reproductive organs of species of Philomycus. Means with std. deviation and ranges are
P.
sella t us
(4)*
P.
virginicus
(4)*
carolinianus
(5)*
P.
bisdosus
(2)*
flexuolaris
(2)*
P.
togatus
(2)*
Length of
penis
13.1(1.82)
11.0-14.2
5.8 (0.66)
5.4-6.8
6.0(0.61)
5.4-7.0
7,1 (0,00)
7.1
7.3 (0.35)
7.0-7.5
12.9 (2.33)
11.2-14.5
Diameter of
penis (atrial
end)
Length of
vagina
4.4(0.49)
4.1-5.0
3.7(1.07)
2.7-5.0
2.2(0.29)
1.9-2.7
3.0 (0.07)
2.9-3.0
2.8(0.64)
2.3-3.2
3.3 (0.42)
3.0-3.6
1.2(0.15)
1.0-1.3
0,8 (0.29)
0.5-10
1,5(0,41)
1.1-2.2
1.1 (0.49)
0.7-1.4
1.6(0.35)
1.3-1.8
3.1(1.70)
1.9-4.3
Length of
free oviduct
8.8(1.50)
7.8-10.5
10.2(1.87)
8.9-12.9
7.7(1.44)
6.2-10.0
7.6(0.62)
6.9-8.1
7.6 (0.64)
7,1-8.0
7.2(0.28)
7.0-7.4
Length of
\as tleferens
57.5(4.77)
52.0-60.5
21.1 (2.11)
1 8 8-23 0
38.1 (4.96)
33,8-45,1
19.1 (1.02)
17.9-19.8
24.2 (3.04)
22 0-26 3
34.0(0.71)
335-345
* Number ot specimens measured.
Page U
THE NAUTILUS, Vol. 107, No. 1
Figures 1 1, 12. Internal penial anatomy. 1 1. Philomyctis bis-
dosus. Scale bar = 10 mm. 12. Philomycus caroliniantis. Scale
bar = 5 mm. A: atrium, DS: dart sac, P: penis, PR: penial
retractor, PS: penial sheath, VD: vas deferens.
DISCUSSION
When Hubricht (1972) described Philomycus sellatus,
he noted that, based upon the posterior half of the mantle,
it ". . .is most closely related to P. carolinianus (Bosc).",
with which "It sometimes occurs. . .". However, the black
transverse band near the anterior end of the mantle of
F. sellatus (figures 3, 4), which is not present on P.
carolinianus (Pilsbry, 1948 p. 748), easily separates the
two species in the field. Their reproductive systems (fig-
ures 6, 10) are also distinct. The shape of the penes and
their internal anatomies (figures 8, 12) are clearly dif-
ferent, Philomycus carolinianus lacks the two internal
constrictions in the penial wall that are found in P. sel-
latus, and the internal penial pilasters are smaller and
more numerous in P. carolinianus than in P. sellatus.
Additionally, the penis and vas deferens of P. sellatus
are nearly twice as long as those of P. carolinianus (Table
1). Indeed, the black anterior transverse band on the
mantle and the two constrictions in the penial wall are
characteristics unique to P. sellatus.
Hubricht (1953) noted that Philomycus virginicus was
found associated with P. flexuolaris (Rafinesque, 1820)
and P. togatus (Gould, 1841) (as P. carolinianus collinus
in Hubricht's 1953 article). Among these ta.\a, only P.
virginicus has the transverse chevron pattern on the man-
tle (figures 1, 2) and is thus easily separated from the
others in the field. In addition, measurements of the
reproductive systems of P. flexuolaris and P. togatus
demonstrated several differences from that of P. virgin-
icus (Table 1). The penis and vagina of P. virginicus are
shorter than those of P. flexuolaris; the free oviduct of
P. virginicus is much longer than that of P. flexuolaris.
Philomycus togatus has a much longer penis, vagina,
and vas deferens than P. virginicus; the free oviduct of
P. virginicus is longer than that of P. togatus (Table 1).
Branson (1968), in reference to P. virginicus and P.
bisdosus (figures 5, 9), stated that "The genitalia of the
two species also differ.", but no figures nor measurements
were included. Branson (1969) listed some differences
between the reproductive anatomies of these latter two
species, but again without figures or measurements. Fig-
ures 7 and 11 show clearly the differences in the shape
of the penis and the internal penial anatomy of these
two species, and the free oviduct of P. virginicus is nearly
1.5 times the length of the P. bisdosus oviduct (Table
1 ). In addition, P. bisdosus lacks the anterior to posterior
transverse chevrons on the mantle. Comparisons with
previous studies of reproductive anatomv in species of
Philomycus (Fairbanks, 1986, 1989; Pilsbry, 1948) con-
firmed that the internal surface of the penis of P. vir-
ginicus is unique in having a basal circular pustulose
ridge at the atrial end.
ACKNOWLEDGMENTS
Financial support for the field trips associated \\ ith this
study was provided by grants from the Research De-
velopment Grant Fund of The Pennsylvania State Uni-
versity. My thanks go to three anonymous reviewers who
provided valuable suggestions for improving this article.
LITERATURE CITED
Bosc, L. A. G. 1802. Histoire naturelle des coquilles. Conten-
ant leur description, et leurs moeurs. \'ol. I. Paris, 343 p.,
Ipl.
Branson. B. A. 1968. Two new slugs (Pulmonata: Philomy-
cidae: Philomycus) from Kentuck\ and X'irginia. The
Nautilus 81(4):127-133.
Branson, B. A 1969. Genital differences in Philomycus vir-
ginicus Hubricht and P bisdosus Branson. The Nautilus
82:74,
Fairbanks, H. L. 1986. The taxonomic status of Philomycus
togatus (Pulmonata: Philomycidae): a morphological and
electrophoretic comparison with Philomycus carolini-
anus. Malacologia 27(2):271-280,
Fairbanks, H, L. 1989, The reproducti\e anatoms and tax-
onomic status of Philomycus vcniislus Hubricht, 1953 and
Philomycus bisdosus Branson, 1968 (Pulmonata: Philo-
mycidae), The Nautilus 103(I):20-23,
Gould, A, A, 1841. Report on the invertebrates of Massachu-
setts, comprising the MoUusca, Crustacea, Annelida and
Radiata, vol. 3.
H. L. Fairbanks, 1993
Page 13
Hubricht, L. 1953^ Three new species of Philomycidae^ The
Nautilus 66(3):78-80.
Hubricht, L. 1972, Two new North American Puimonata:
Paravitrea seradens and Philomycus sellatus^ The Nau-
tilus 86(1):16-17.
Pilsbry, H. A. 1948. Land MoUusca of North America (North
of Mexico). The Academy of Natural Sciences of Phila-
delphia Monograph Number 3, Vol, II Part 2:759.
Rafinesque, C. S. 1820. .Annals of Nature or annual synopsis
of new genera and species of animals and plants discovered
in North America, p. 10.
THE NAUTILUS 107(l):14-23, 1993
Page 14
Two Confusing Indo-Pacific Cerithiids
Richard S. Houbrick
Department of linertebrate Zoology
National Museum of Natural History
Smithsonian Institution
Washington, DC. 20560
ABSTRACT
Cerithium zebrum Kiener, 1841 and Cerithium boeticum Pease,
1860. although originally described as CerUhiiim species, had
been allocated to Bittium Gray, 1847, in the recent literature
Anatomical investigation has shown that these two species do
not have the characters of Bitlium species, but are are more
like Cerithium taxa, e.xcept for their small size. Cerithium
zebrum is distinguished from Cerithium boeticum. based on
morphological characters derived from the shell, radula and
soft anatomy. Synonymies, descriptions and a discussion of the
differences between the species are presented.
Key Words: Cerithium, Bittium, morphology, anatomy, tax-
onomy.
INTRODUCTION
Small-shelled cerithiids are difficult to identify to the
species-level, particularly poorly-known taxa from the
Indo-Pacific region. Generic and sometimes familial de-
terminations of these small snails are likewise difficult
and controversal, and in most museum collections many
small-sized cerithiids are grouped together and loosely
attributed to the genus Bittium Gray, 1847.
During a recent generic review of the Bittium-gToup
the anatomy of a number of "Bittium" species were
examined. One of the taxa studied is "Bittium" zebrum
(Kiener, 1841), a common species throughout the Indo-
Pacific region that is extremely variably in sculpture and
color pattern. As expected in a common, widely-distrib-
uted species comprising many sculptural phenotypes and
color morphs," Bittium ' zebrum has been zealously ov-
ernamed by authors, as a perusal of its synonymy will
testify.
When living "Bittium" zebrum specimens from Guam,
Enewetak, and Hawaii were examined it was discovered
that none of these snails had an epipodial skirt, charac-
teristic of members of Bittiinae. Moreover, there was no
spermatophore bursa present in the lateral lamina of the
pallial oviduct, a feature distinctive of the Bittium-group.
These animals are thus transferred from Bittitim to Cer-
ithium Briiguiere.
Further comparison of the Hawaiian specimens of
"Bittium" zebrum with those from other Indo-Pacific
regions revealed that there are considerable morpholog-
ical differences between the populations: the shells, rad-
ula and anatomy of the Hawaiian specimens are quite
distinct from those of zebrum specimens from elsewhere.
Subsequent careful examination of many museum lots
of Cerithium zebrum revealed that most lots of Hawaiian
specimens comprised mixtures of two species: the first
having a typical zebrum phenotype, and the second,
more common, species having a different shell physi-
ognomy. It became apparent that the second species is
Cerithium boeticum Pease, 1860, named from the Ha-
waiian Islands and subsequently considered to be con-
specific with "Bittium" zebrum by Kay (1979).
This paper addresses the differences between the two
species, describes and presents a s\ nonymx of each, and
delineates the characters separating and distinguishing
them. Cerithium boeticum is recognized as a valid spe-
cies, different from Cerithium zebrum, and is shown to
be endemic to the Hawaiian Islands.
MATERIALS AND METHODS
Living specimens of Cerithium zebrum were studied at
Enewetak Atoll, Marshall Islands, and at the L^niversity
of Guam Marine Laboratory at Pago Bay, Guam. Cer-
ithium boeticum was collected from shallow, subtidal
coral rubble in Kewalo Basin, Honolulu, placed in aquar-
ia and observed at the Kewalo Laboratorx of the Ha-
waiian Biomedical Research Laboratory, University of
Hawaii, Honolulu, Hawaii. Both species were observed
and dissected under water in wax-filled petri dishes using
a Wild M-5 dissecting microscope. Methylene blue was
used to enhance anatomical features during dissection.
Protoconchs, shells, opercula and radulae were studied
with an Hitachi S-570 scanning electron microscope. The
types of both species and their synonymous nomina were
examined and are discussed below. Man\- museum lots
were examined in order to establish the range of phe-
notypic variation in the shells of each species. Measure-
ments and meristic data of shells were taken using ran-
domly selected specimens from locations throughout the
geographic range of each species.
R. S. Houbrick, 1993
Page 15
Material examined: Cerithiitm hocticum. HAWAIIAN
ISLANDS: (USNM 343522); Oaliu (USNM 12916, types);
Honolulu, Oahu (USNM 335496, 335499, 335497, 335498,
767506); Waikiki Marine Lab, Honolulu, Oahu (USNM
633002); Kewalo Basin, Honolulu, Oahu (USNM 857099);
Honolulu Reef, Oahu (USNM 335289); Quarantine Is-
land, Honolulu, Oahu (USNM 339349, 339342); Waikiki,
Oahu (USNM 343514, 343515, 343519); off Waikiki, Ho-
nolulu, Oahu (USNM 339118); Kewalo Basin, Honolulu,
Oahu, Hawaii (USNM 857099); Diamond Head, Hono-
lulu, Oahu (USNM 339339, 343510); Mokapu Beach,
Oahu (USMN 484570); Maunalua Beach, Oahu (USNM
428173, 428174, 428207, 343520, 343511); Mokoloe Is-
land, Oahu (USNM 339343, 343512); Kaneohe Bay, Oahu
(USNM 472057, 472058, 343513); Coconut Island, Ka-
neohe Bay, Oahu (USNM 771362, 771556); Hospital Point,
Pearl Harbor, Oahu (I'SNM 497938); Pearl Harbor, Oahu
(USNM 484455, 428234, 341308); Pupukea Beach, Oahu
(USNM 484689); Kahana Bav, Oahu (USNM 777967);
Kahala, Oahu (USNM 33951, 339928); Kahala Beach,
Koko Head, Oahu (USNM 343546); Keoke River, Hawaii
(USNM 252339, 252338, 252336); Keokea, Hawaii (USNM
337592); Keokea, Hilo, Hawaii (USNM 339340, 339340);
Hilo. Hawaii (USNM 612281); Kuhio B., Hilo, Hawaii
(USNM 339125); Honaunau, Hawaii (USNM 343517); 5
mi SW Kapoho, Hawaii (USNM 409081, 409095); Lanai
(USNM 612425); Mauele, Lanai (USNM 339341); Han-
alei, Kauai (USNM 340692); Midway Island (USNM
790897, 790898); Frigate Point, Sand Island (USNM
678064).
Cerithiumzebriim. MAURITIUS: (USNM 91247, 91069,
91070); Flicq-en-Flacq, Arsenal Bay (USNM 716523).
SEYCHELLES: Aldabra Atoll, lie Picard (USNM 837490).
COCOS (KEELING) ISLANDS: Alor Pinvu, NW side of
West Island (USNM 656435). AUSTRALIA: Heron Is-
land, Queensland (USNM 8540174); Watson's Bav, Liz-
ard Island, Queensland (USNM 783381). RYUKYU IS-
LANDS: Osima, Osumi (USNM 343873); Odomaria,
Okinawa, Shima (USNM 488199); Yenoshima (USNM
228222); Chichyima, Ogasawara (USNM 175587). BO-
NIN ISLANDS: '(USNM 36950, 343874). PHILIPPINES:
Jamelo Bav, Luzon (USNM 935351). MARIANAS: Saipan
(USNM 486824, 486823, 486795); Guam (USNM 851 104,
851104); Pago Bay, Guam (USNM 774778); Apra Bay,
Guam (USNM 240110); Orote Point, Orote Cliffs, Guam
(USNM 854036). CAROLINES: Moen Island, Truk La-
goon (USNM 842615); Reef at Mutunlik, Kusaie Island
(USNM 609487). LOYALTY ISLANDS: (USNM 253564);
Lifu (USNM 423278, 423260, 423210, 423282, 423281,
423261, 423325, 423283). FIJI: Rat Tail Passage, Suva
Reef (USNM 824808); SAMOA: Pago Pago (USNM
361535, 361536); Ofu, Manu'a Group (USNM 380968,
380965, 380967, 380969, 380962, 380963, 380960,
380966). MARSHALL ISLANDS: Enewetak (USNM
432415, 770725, 542726); Rennit Island, Enewetak
(USNM 542809); Fred, Enewetak (USNM 770687); Aa-
raanbiru Island, Enewetak (USNM 582227, 582265); Tei-
teiripucoki Island, Enewetak (USNM 581543); Girunien
Island, Enewetak (USNM 581534); Engebi Island, Enew-
etak (USNM 743869); Rijoru Island, Enewetak (USNM
581555); E. Rigili Island, Enewetak (USNM 581923);
between Rigili and E. Rigili Islands (USNM 581576,
581575); Ailuk Island, Ailuk Atoll (USNM 615133); Lae
Island, Lae Atoll (USNM 614896); Bock Island, Rongerik
(USNM 594660); Enwertok, Rongerik (USNM 583519);
4 mi. W Bikini Islands (USNM 586086); Bikini (USNM
583949, 585185); Bokororvuru, Bikini (USNM 583883,
583884); Namu Island, Bikini (USNM 580542); Enyu
Island, Bikini (USNM 580945); Eninman Island, Bikini
(USNM 586897); Pigenivarayro, Rongelap (USNM
585588, 585327); Kabelle Island, Rongelap (USNM
582405, 582388, 582125, Burok, Rongelap (USNM
583989); NIUE: Opaahi, S. of Alofi (USNM 854054); Alofi
(USNM 854046); Tuapa (USNM 858164); Tuapa, Avaiki
Cave (USNM 854055). COOK ISLANDS: Mauke Island
(USNM 598174). SOCIETY ISLANDS: Reao Id (USNM
5573, holotype); Makatea, Tuamotu Archipelago, N Ti-
mae Harbor (USNM 819895); Arue, Papeete, Tahiti
(USNM 775927); Mahina, Tahiti (USNM 797267, 791372);
Tahiti (USNM 91068). WALLIS & FUTUNA ISLANDS:
outer reef E of Nukuhifala (USNM 676427); Henderson
Id (G. Pauley coll.). NIUE: off warf, Alofi (USNM 854046);
Avaiki Cave, Tuapa (USNM 854055); Tuapa (USNM
858167); Opaahi, S Alofi (USNM 854054). HAWAII:
French Frigate Shoals (ANSP 195368, 195384); Midway
(USNM 790897); Honaunau, Hawaii (USNM 343509);
Keokea, Hilo, Hawaii (USNM 339353).
Abbreviations: BMNH = British Museum (Natural His-
tory); MNHNP = Museum National d'Histoire Naturelle,
Paris; NMW = National Museum, Wales; SEM = scan-
ning electron microscope; USNM = United States Na-
tional Museum, Smithsonian Institution, Washington,
DC.
RESULTS
Descriptions (Tables 1 & 2, Figs. 1-28), synonymies, and
discussions of both species are presented below, followed
by a table identifying the characters separating the two
species (Table 3).
Cerithium zebrum Kiener, 1841
Cerithium zebrum Kiener, 1841:71, pi. 25, fig. 4 (holo-
tvpe, MNHNP; type locality, Indian Ocean, Mauritius).
Sowerbv, 1855:875-876, pi. 183, figs. 207-209; 1865, pi.
19, fig. 136; Tryon, 1887: 138, pi. 26, figs. 78-82; Kobelt,
1898:210-211, pi. 39, figs. 7-10; Hedley, 1899:434; Vignal,
1903:21-22, pi. 2, fig. 1; Schepman, 1909:162; Dautzen-
berg & Bouge, 1933:313.
Cerithium janlhinum Gould, 1849:121 (holotype, USNM 5573,
8.5 X 3.9 mm; type loclaity. Clermont Tonnere [Reao Id,
Tuamotusj; 1852:152, pi. 10, fig. 173a-b; 1862:63; Johnson,
1964:95.
Cerithium zebrum var. dilectum C.B. Adams in Sowerby, 1855:
896, pi. 33, fig. 207' (15 syntypes, BMNH 1969349; type
locality, Galapagos [in error]); Vignal, 1903:22-23, pi. 2,
fig. 5. ■
Page 16
THE NAUTILUS, Vol. 107, No. 1
R S. Houbrick, 1993
Page 17
Figs. 16-17. Scanning electron micrographs of radula of Cerithium zebrtim Kiener, Pago Bay, Guam (USNM 774778); 16. view
of mid-section of radula, bar = 36 ^m; 17. half row showing details of rachidian, lateral and marginal teeth, bar = 29 ^m.
Cerithium delectum (sic) A. Adams. Sowerbv, 1865, pi 16, fig
112 a-c; Vignal, 1903:23.
Cerithium unilincatum Pease, 1860:432 (lectot>pe, BMNH
19622798, 5.5 x 3 mm, 2 paralectotypes BMNH 1962799;
t\ pe locality. Sandwich Ids [Hawaiian Ids]). Sowerbv, 1866,
pi. 15, fig.99; Trvon, 1887:138, pi. 26, fig. 86; Kobelt, 1898:
224, pl.29, fig. 18.
Cerithium stigmosum Gould, 1861:386 (holotype, ?; type lo-
cahty, Bonin Ids) 1862:141; Johnson, 1964:152.
Cerithium aspersum Deshayes, 1863:97-98, pi. 11, figs. 16-18
(lectotype, here selected, MNHNP, 8.8 mm, 4 paralecto-
types, MNHNP; type locality. Reunion).
Cerithium Crossii Deshayes, 1863:96, pi. 11, figs. 12-14 (lec-
totype, here selected, MNHNP, 8.4 mm, 3 paralectotypes,
MNHNP; type locality. Reunion; not Cerithium cros-
seanum Tiberi, 1863 [= Cerithioipsis sulwylindricum
(Brusina)], nor Cerithium maillardi Crosse, 1863 [unnec-
essary replacement name for Cerithium crosseanum Ti-
ber!].
Cerithium dichroum Melvill is. Standen, 1895: 115, fig. (6 syn-
types NMW 55158200; type locality, Lifu, Lo\aity Idsi
Cerithium zebrum var. pulchra Vignal, 1903:22, pi. 2, fig. 2
(t>pe, MNHNP; type localit> , Isle of Pines, New Cale-
donia); Dautzenberg & Bouge, 1933:315.
Cerithium zebrum var. trijasciata N'ignai, 1903:22, pi. 2, fig.
3 (type, MNHNP; type locality. Isle of Pines, New Cale-
donia).
Cerithium zebrum var. attenuata Vignal, 1903:22, pi. 2, fig.
4 (type, MNHNP; type locality. Isle of Pines, New Cale-
donia); Dautzenberg & Bouge, 1933:313.
Cerithium zebrum var undulata Dautzenberg ic Bouge, 1933:
316 (Type, MNHNP; type locality, Tuamotus).
Cerithium zebrum var. rosea Vignal, 1903:24, pi. 2, fig. 6 (type,
MNHNP; t\pe locality. Isle of Pines, New Caledonia);
Dautzenberg & Bouge, 1933:315.
Cerithium zebrum var. cinerea Vignal, 1903:24, pi. 2, fig. 7
(type, MNHNP; type locality. Isle of Pines, New Cale-
donia); Dautzenberg & Bouge, 1933:314.
Cerithium zebrum var. nivea Vignal, 1903:24, pi. 2, fig. 8 (type,
MNHNP; type locality. Isle of Pines, New Caledonia);
Dautzenberg & Bouge, 1933:315.
Cerithium zebrum var. biUneata Vignal, 1903:25, pi. 2, fig. 10
Figs. 1-15. Cerithium zebrum Kiener, showing shell variation and types of synonymous nominal ta.xa. 1-2. Orote Point, Guam,
7.9 mm length; 3. operculum, 1.3 mm length (ISNM 854036); 4. Orote Point, Guam, 7.5 mm length (USNM 854036); 5. Pago
Bay, Guam, 7.8 mm length (USNM 774778); 6. Enewetak Atoll, Marshall Islands, 7.9 mm length (USNM 770725); 7. lectotype
of Cerithium crossii Deshayes, Reunion (MNHNP), ; 8. Holotype of Cerithium janthinum Gould, Reao, Tuamotus (USNM 5573);
9. SEM of protoconch, bar = 70 ^m; 10. lectotype of Cerithium aspersum Deshayes, Reunion, 8 2 mm (MNHNP); 11-12. Pago
Bay, Guam, apertural and right lateral views under SEM, showing sculptural details, 5 8 tnm length (I'SNM 774778); 13. sculpture
of early whorls, SEM, bar = 0.9 mm; 14—15. Pago Bav, Guam, dorsal and right lateral views under SEM, 5.8 mm length (USNM
774778).
Page 18
THE NAUTILUS, Vol. 107, No. 1
Figs. 18-26. Cerithium boeticum Pease, from Kewalo Basin, Honolulu, Oahu, Hawaii, showing shell variabilit) (USNM 857099).
18-19. Strongly banded morph, 5.8 mm length; 20-21. white morph, 5.7 mm length; 22-23. 5.7 mm length; 24. scanning
electron micrograph showing sculptural details and aperture, 4.9 mm length; 25. operculum. 2 mm length; 26. SEM of protoconch
and early whorl sculpture, bar = 88 fzm
(type, MNHNP; type locality. Isle of Pines, New Cale-
donia); Dautzenberg & Bouge, 1933:314.
Cerithium zelyrum var. triliyieata Vignal, 1903:25, pi. 2, fig.
11 (type, MNHNP; type locality. Isle of Pines, New Cal-
edonia); Dautzenberg & Bouge, 1933315.
Cerithium zebrum var. ornata Vignal, 1903:26, pi. 2, fig. 13
(type, MNHNP; type locality. Isle of Pines, New Cale-
donia).
Cerithium zebrum var. sticta Vignal, 1903:26, pi. 2, fig. 14
(type, MNHNP; type locality, Isle of Pines, New Cale-
donia); Dautzenberg & Bouge, 1933:315.
Cerithium zebrum var mutimaculata Vignal, 1903:26, pi. 2,
fig. 15 (type, MNHNP; type locality. Isle of Pines, New
Caledonia); Dautzenberg & Bouge, 1933:315.
Cerithium zebrum var. maculata Vignal, 1903:26, pi. 2, fig.
16 (type, MNHNP; type locality. Isle of Pines, New Cal-
edonia); Dautzenberg & Bouge, 1933:314.
Cerithium zebrum var. unimacuhita N'ignal, 1903:27, pi. 2 fig.
17 (tspe, MNHNP; type localit\', isle of Pines, New Cal-
edonia).
Cerithium zebrum var. aspersa Deshayes. Dautzenberg &
Bouge, 1933:314.
Bittium zebrum (Kiener) Cernohorsky, 1972:69-70, pi 14. fig.
10; Salvat & Rives, 1975:272, fig. 74.
Bittium zebrum (Kiener) Kay, 1979:120, figs. 45b-c (not Bit-
tium zebrum (Kiener), is Cerithium boeticum Pease 1860).
Description: Shell (Figs. 1-15; Table 1): tiirreted, elon-
gate, attaining 10.4 mm length, 3.2 mm width, and com-
prising about 9 weakly inflated to flat-sided whorls. Pro-
R. S. Houbrick, 1993
Page 19
Figs. 27-28. Scanning electron micrographs of radula of Cerithium boeticum Pease, Kewalo Basin, Honolulu, Oahu, Hawaii
(I'SNM 857099). 27. general view of mid-radular ribbon, bar = 25 ^m; 28. detail of rachidian tooth, bar = 9 nm
toconch (Fig 9) comprising 3 whorls; protoconch 1 nearly
smooth, sculptured with suprasutural spiral cord; pro-
toconch 2 sculptured with series of subsutural plaits fol-
lowed by two strong spiral cords, criss-crossed by minute
lirae up to anterior suture. Early whorls (Fig. 13) sculp-
tured with 3 beaded spiral cords. Adult whorls sculptured
with 4 beaded, major spiral cords each separated from
one another by one fine spiral lira. Penultimate whorl
with 5 beaded spiral cords, having about 26 spiral beads;
beads frequently aligned to form weak a.xial riblets. Body
whorl with 11-12 beaded spiral cords and fine spiral lirae.
Suture weakly defined. Strong white varices randomly
placed on whorls; largest varix on right dorsal side of
body whorl. Aperture circular-ovate, a little over one-
fourth the shell length. Outer lip thick, relatixely smooth,
weakly crenulate at inner edge; columella concave with
weak callus. Anterior canal short, but pronounced, ori-
ented at 45 degree angle to shell a.xis. Weak siphonal
constriction. Anal canal small, defined bv small parietal
plait Shell color white with pink early whorls and adult
w horls w ith brown, tan, rose blotches and spiral stripes,
bands, or dots. Aperture white. Periostracum thin, light
tan.
Animal: Headfoot white, flecked with opaque white,
rose or tan. Snout, mantle edge and siphon \ellow . Mantle
edge with short yellow papillae. Snout long, extensible,
having bilobed tip (lips). Operculum (Fig. 3) tan, cor-
neous, ovate, paucispiral with eccentric nucleus. Lateral
epipodial skirt and operculiferous lobe absent. Osphra-
dium bipectinate, with weakly-defined pectins. Radula
(Figs. 16-17) short, about one-ninth the shell length.
Rachidian tooth (Fig 17) height equals length, having
hourglass-shaped basal plate with spade-shaped basal
projection and two small basal lateral butresses; cutting
edge with large, pointed central cusp flanked by two
small pointed denticles on each side. Lateral tooth with
rectangular basal plate having long lateral projection and
Table 1. Shell statistics for Cerithium zcbrum (measurements
in mm)
Table 2. Shell statistics for Cerithium boeticum (measure-
ments in mm).
Statistic (n = 17)
X
sd
Range
Statistic (n = 17)
X
sd
Range
Length
7.3
1.5
4.2-10.35
Length
6.1
1.8
2.4-10.5
Width
2.4
0.4
1.8-3.2
W^idth
2.3
0.8
1.2-4.6
Aperture length
1.3
0.3
0.8-1.8
Aperture length
1,7
0,7
0.8-2.4
Number of whorls
9.2
1.6
7-13
Number of whorls
8,0
0,9
6-9
Number spiral beads
26,3
6
19-44
Xvunber axial beads
174
1 7
14-20
Page 20
THE NAUTILUS, Vol. 107, No. 1
thick central buttress; cutting edge with one weak inner
denticle, large spoon-shaped cusp, and three outer den-
ticles. Marginal teeth spatulate, curved at tips; inner mar-
ginal tooth with three pointed inner denticles, large cen-
tral cusp and two outer denticles; outer marginal tooth
same but without outer denticles. Esophageal gland pres-
ent; stomach large; style sac and crystalline style long,
about one-third stomach length.
Synonymic remarks: This species has been the recipient
of 23 names, all of which have been based on concho-
logical characters and/or differences in shell pigmenta-
tion. It is obvious that the authors of these nomina did
not employ a modern species concept and failed to rec-
ognize the full range of intraspecific variation when pro-
posing these ta.xa. Examination of the types of the syn-
on\mous nomina re\ealed no significant differences in
shell sculpture from that of Cerithium zebrum (Kiener);
indeed, the nomina proposed by Vignal (1903) as vari-
eties of Cerithium zebrum Kiener were based on mere
intraspecific color variation, as Vignal (1903) clearly stat-
ed. The holotype of Cerithium janthinum Gould (see
Fig. 8) is merely a worn, very wide morph of Cerithium
zebrum. Sowerby (1855) introduced the name dilectum
in the Thesaurus as a variety of Cerithium zebrum Kie-
ner, but the new taxon was indicated only by the itali-
cized name in his "Alphabetical list of species", on p.
896. The name dilectum was initially listed on p. 117 as
a synonym of Cerithium zebrum, and the legend for the
figures of Cerithium zebrum (figs. 107-109) does not
include the name dilectum, although one of the figures
has an asterix (fig. 207-), which presumably is meant to
indicate the varietal taxon, dilectum. Cerithium asper-
sum Deshayes is a highly sculptured morph of C. zebrum,
as illustrated by the lectotype (Fig. 10). Cerithium crossi
Deshayes, 1863, which as shown by the lectotype (Fig.
7), is conspecific with Cerithium zebrum, was incorrectly
considered by Crosse (1863) as a homonym of Cerithium
crosseanum Tiberi, 1863, a Cerithiopsis species, and was
given an unnecessary replacement name, Cerithium
maillardi Crosse, 1863. Cerithium boeticum Pease er-
roneously has been considered conspecific with Ceri-
thium zebrum Kiener by Kay (1979). For a discussion
of this issue, see the discussion under Cerithium boeti-
cum.
Discussion: Cerithium zebrum is a very common spe-
cies, frequently found in the upper intertidal zone among
rubble and algae on benches and platforms associated
with fringing coral reefs of atolls and islands throughout
the Indo-Pacific. In the Hawaiian Islands is also occurs
in deeper waters (Kay, pers. com.). It is characterized
by an elongate, tapering shell, sculptured with many fine
beaded spiral cords and with numerous broad, white,
randomly-placed varices. Cerithium zebrum is a colorful
species, having a rose-colored apex capped with a dark
brown protoconch (see Figs. 1-2,4,6). The adult whorls
are frequently white or light pink with tan blotches or
with brown and tan bands. While the species varies great-
ly in color pattern, shell sculpture is relatively stable
throughout its range. Banded varieties of Cerithium ze-
brum may be confused with banded morphs of Ceri-
thium boeticum, but the latter species has a more coarse-
ly sculptured shell with stronger axial ribs, a more deeply
incised suture, and a thick outer apertural lip (see Figs.
18-26). The animal (headfoot), in contrast to the brightly
pigmented shell, is a uniform white overlain by opaque
white spots, although some shells may also be white.
In contrast to the lecithotrophic development seen in
Cerithium boeticum, Cerithium zebrum undergoes a
lengthy planktotrophic larval phase and has an elabo-
rately sculptured protoconch (Fig. 9) with a deep sinu-
sigeral notch indicative of this de\elopmental mode. The
operculum of Cerithium zebrum is slightK- more ovate,
thicker and opaque, and has fewer spirals than that of
Cerithium boeticum.
Cerithium zebrum occurs from Henderson Id., in the
Pitcairn Group, throughout French Polynesia and the
Hawaiian Islands, and westward throughout the tropical
Pacific from Queensland, Australia, north to the Ryu-
kyus. It is found throughout the archipelagos of southeast
Asia and in suitable localities in the Indian Ocean. In-
deed, specimens from Tamarin Bay, Mauritius (AXSP
273492) look very much like those found in Tahiti (Fig.
6) and Guam (Figs. 1-2, 4-5). While Cerithium zebrum
has occasionally been collected in assemblages from depths
greater than 10 m around the main, volcanic Hawaiian
Islands (Kay, in lit.), it is more common on the shallow
reefs of the Northwestern Hawaiian Islands at French
Frigate Shoals (ANSP 195368, 195384) and Midway
(USNM 790897).
To my knowledge, Cerithium zebrum has not been
recorded as a fossil.
Cerithium boeticum Pease, 1860
Cerithium pusillum Gould, 1851: 120-121 (holotype, USNM
5572, 4 paratypes, MCZ 216606; type locality, Sandwich
Ids [Hawaiian Ids); nol Cerithium pusillum Pfeiffer, 1840,
nor Dunker,1859); Gould, 1852:150, pi 10, fig. 172a-b;
Gould, 1862:62; Dunker, 1882:108.
Cerithium boeticum Pease, I860: 433 (lectotype [Kay, 1965],
BMNH 1962802, 6 x 2.5 mm, 5 paralectotypes, BMNH
1962803; type locality. Sandwich Ids [Hawaiian Ids]); Sow-
erby, 1866 "haeticum", pi. 12, fig. 327; 1866b, pi. 16, fig.
114; Trvon, 1887:143, pi. 27, fig. 26; Kobelt, 1898:236, pi.
41, fig. 14; Kay, 1965:48-49, pi. 10, fig. 8.
Biitium pusillum (Gould) Johnson, 1964: 136; Kay, 1979:120
(not Biitium pusillum [Dunker, 1859]).
Bitlium zebrum (Kiener) Kay. 1979: 120. fig. 45b-c (not ze-
brum Kiener, 1841, is Bittium pusillum [Gould, 1849]).
Bittium boeticum unilineatum Pse. (sic). Mant, 1923: 121.
Bittium boeticum Pils. & Va. (sic). Mant, 1923: 121.
Description: Shell (Figs. 18-24, 26; Table 2): Shell solid,
turreted, reaching 10.5 mm length and 4,6 mm width,
comprising about 8 con\ex whorls. Protoconch (Fig. 26)
one whorl, smooth with straight lip. Early whorls sculp-
tured with two spiral cords and weak axial ribs. Penul-
timate whorl sculptured \\ ith 4 major beaded, spiral cords
R. S. Houbrick, 1993
Page 21
each separated from the other with weak spiral Hrae,
and each bearing about 17 beads aHned to form 17 strong
axial ribs. Bod\ whorl large, sculptured with 9 spiral
beaded cords and weak spiral lirae. Several varices ran-
domly distributed on shell, strongest opposite outer lip
of aperture. Suture deeply impressed. Aperture oval-
elongate, a little over one-third the shell length. Anterior
canal short, well-defined, oriented at 45 degree angle to
shell axis; outer lip thick, strongly denticulate along inner
edge; columella concave with moderate callus. Anal ca-
nal weak, defined b\ parietal plait.
Animal: Headfoot without lateral epipodial skirt; weak-
ly scalloped operculiferous lobe present. Operculum (Fig.
25) corneous, very thin, transparent and paucispiral with
slightly eccentric nucleus. Osphradium very large, about
one-half w idth of ctenidium, strongK bipectinate with
large pectins. H> pobranchial gland well-developed. Me-
dial lamina of pallial oviduct with spermatophore bursa
and large seminal receptacle. Radula (Figs. 27-28) very
short, about one-twelfth the shell length. Rachidian tooth
(Fig. 28) wider than tall, hourglass-shaped, with basal
plate having short basal projection and two small lateral
buttresses; cutting edge with spade-shaped central cusp
flanked on each side with three, sometimes two, small
pointed denticles. Lateral tooth w ith rectangular basal
plate having long lateral projection and short basal but-
tress; cutting edge with small inner denticle, long spoon-
shaped major cusp, and two, sometimes three, outer
pointed denticles. Marginal teeth elongate, spatulate with
curved tips; inner marginal tooth w ith two inner denticles
long, major central cusp and one outer denticle; outer
marginal tooth same but without outer denticle.
Synonymic remarks: Cerithium boeticum has been
confused with several other taxa assigned to Cerithium
or Bittiiim by authors. The earliest name for the species
is Cerithium pusillum Gould, 1851, but as this name is
preoccupied, the next available valid name is Cerithium
boeticum Pease, 1860. Although the figures subsequently
presented by Gould (1852, pi. 10, figs. 172a-b) for Cer-
ithium pusillum are not very good, the holotype (USNM
5572) clearK shows that this taxon is different from Cer-
ithium zebrum Kiener, with which it has been confused
(e.g., Kay, 1979; 120). Cerithium pusillum Dunker, 1859,
and Cerithium pusillum Gould, 1851, are different spe-
cies, but as both taxa have been referred to Bittium by
authors, some confusion has developed about their iden-
tity.
Cerithium boeticum Pease is a name seldom men-
tioned in the literature and has not been a well-under-
stood species. Kay (1965) designated a lectotype and five
paralectotypes from the original type lot of six beach-
worn specimens, but her illustration of the lectotype (1965,
pi. 10, fig. 8) is poor and is not helpful in clarifying the
concept of the species. Kay (1979) subsequently placed
this taxon into the synonymy of Cerithium zebrum, a
species with a wide Indo-Pacific distribution. Further
confusion has developed because another Pease (1860)
name, Cerithium unilineatum, a valid svnonvm of Cer-
ithium zebrum Kiener, was combined with Cerithium
boeticum Pease to form a trinomial In some museum
collections the manuscript name, Cerithium gouldianum
Pilsbry & Vanatta, is found associated with lots of Cer-
ithium boeticum. but the name has no validity.
Discussion: As mentioned above, Cerithium boeticum
had been condsidered to be synonymous with the com-
mon Indo-Pacific species, Cerithium zebrum Kiener, but
the former species has a thicker, less elongate shell with
coarser sculpture, few weak varices, a stronger anterior
canal, and a different kind of protoconch than Cerithium
zebrum (see Figs. 18-26); moreover, Cerithium boeti-
cum appears to be restricted to the Hawaiian Islands,
whereas Cerithium zebrum is distributed throughout the
Indo-Pacific. Shells of the two species frequently share
similar color patterns, especially among the banded,
striped morphs, but when closely compared, they are
seen to be realK' quite different. Radular differences be-
tween the tw o species although slight, are consistent and
serve to distinguish each of them (see Table 3). The very
thin, nearly transparent operculum of Cerithium boe-
ticum (Fig. 25) differs from the thicker one of Cerithium
zebru m .
Gould (1851) mentioned that Cerithium boeticum
(cited as Cerithium pusillum) was "like C. ferrugineum
Sa\-, but much smaller". Cerithium ferrugineum is a
s\nonym of Cerithium lutosum Menke, a small species
from the western Atlantic, and indeed Cerithium boe-
ticum does bear some resemblance to small morphs of
Cerithium lutosum in overall shell morphology (see
Houbrick, 1974, pi. 41).
An unusual anatomical feature of Cerithium boeticum
is the large, well-developed bipectinate osphradium. Al-
though Cerithium zebrum also has a bipectinate os-
phradium, the pectins are poorly defined. A bipectinate
osphradium is common to all members of Cerithium
Bruguiere and Chjpeomorus Jousseaume (see Houbrick,
1974, 1985), but not as common among Bittium species
(pers. obsr). Although Cerithium boeticum has a weakly
scalloped operculiferous lobe, the lateral epipodial skirt,
indicative of Bittium species, is absent. The absence of
a spermatophore bursa in the lateral lamina of the pallial
oviduct, which is the hallmark of Bittium species, is one
of the chief anatomical reasons for reassignment of this
species to Cerithium. However, the specimens studied
herein may not have attained complete reproductive ma-
turity, and this needs reconfirmation.
Cerithium boeticum lives in shallow water on subtidal
rubble where it is frequently found on marine algae.
This species has lecithotrophic development, which is
reflected in its relatively unsculptured protoconch, com-
prising only one whorl with a nearlv straight apertural
Up (Fig. 26).
Cerithium boeticum appears to be endemic to the
Hawaiian Islands. It is geographically sympatric with
Cerithium zebrum, especially in the northern part of the
Hawaiian chain. This species has been recorded from
the Pleistocene of Molokai (Ostergaard, 1939, as Ceri-
thium boeticum).
Page 22
THE NAUTILUS, Vol. 107, No. 1
Table 3. Characters defining Ceritbiurn zclmim and Ceri-
thiitrn hoeticum.
Cerithium zebruni
Cerilhium hoeticum
Sculpture
1. Weak axial riblets Strong axial ribs
2. Many weak spiral beads Fewer strong spiral beads
3. Weakly impressed suture Strongly impressed suture
4. Many broad white varices Few weak varices
5. Protoconch highly sculji- Protoconch unsculptured
tared
6. Thin outer apertural lip Thick outer apertural lip
7. Aperture '74 shell length Aperture 'i shell length
Radula
1. Radula short Radula very short
2. Rachidian height equal to Rachidian wider than tall
width
3. Rachidian cusp flanked Rachidian cusp flanked
with 2 denticles on each with 3 denticles on each
side side
4. Inner marginal with 3 in- Inner marginal with 2 inner
ner and 2 outer denticles and one outer denticle
Osphradium
1. Weakly bipectinate Large, strongly bipectinate
Development
1. Plantotrophic Lecithotrophic
A sumniar) ol the dittereiices between these two spe-
cies is presented below, in Table 3.
CONCLUSIONS
Evidence is presented above to show that the Hawaiian
species, Cerithium hoeticum, is morphologically distinct
from Cerithium zebrum. Although both species had pre-
viously been assigned to Bittium. anatomical evidence
shows that this is incorrect. As both taxa were thought
to be Bittium species, they were omitted from my Indo-
Pacific Cerithium monograph (Houbrick, 1992). Ceri-
thium hoeticum is restricted to the Hawaiian Islands
while Cerithium zebrum has a very wide Indo-Pacific
distribution, including the Hawaiian Islands.
ACKNOWLEDGEMENTS
I thank Michael Hadfield, of the Univeristy of Hawaii,
Honolulu, Hawaii, for the use of laboratory space and
for assistance during field studies in Hawaii. I am grateful
to Alison Kay for valuable discussions about the geo-
graphic distribution of both species throughout the Ha-
waiian Islands. Lou Eldredge, University of Guam, kind-
ly provided me with lab space and logistic support at the
Pago Ray lab of the University of Guam. The work at
the marine laboratory at Enewetak Atoll was made pos-
sible by a grant from the Atomic Energy Commission.
Bernadette Holthuis, University of Guam, kindly ob-
served living Bittium zebrum specimens in the field aiul
passed on her observations to me. I am grateful for the
use the Smithsonian Secretary's Research Opportunity
Fund, w hich covered the costs of transporation and lodg-
ing in both Guam and Hawaii during this study. Susanne
Braden of the Smithsonian Scanning Electron Micro-
scope Lab provided assistance with the SEM micrographs
and V'ictor Krantz, Smithsonian Photographic Services,
helped with photography.
LITERATURE CITED
Cernohorsky, W. O., 1972. Marine shells of the Pacific, Vol.
2, 411 pp., 68 pis. Pacific Publications, Ltd., Sydney.
Crosse, M.H., 1863. Description d'especes nouvelles d'Aus-
tralie. Journal de Conchyliologie 11:84-90, pi. 1.
Dautzenberg, P. & J.L. Bouge, 1933. Les mollusques testaces
marins des etablissements Fran9aise de I'oceanie. Journal
de Conchyliologie 27(2);145-469.
Deshayes, G.P., 1863. Catalogue des mollusques de I'lle de
la Reunion (Bourbon). 144 pp., pis. Paris, Dentu.
Dunker, G. 1859-1860. Neue japanische Mollusken. Mala-
kozoologische Blatter 6:221-240.
Dunker, G. 1882. Index Molluscorum Maris Japonica. Nov-
itates Conchologiae, Supplement to Volume 7. Cassell.
Gould, A. 1851. Descriptions of new species, brought home
by the US, Exploring Expedition Proceedings of the Bos-
ton Society of Natural History 3:118-121.
Gould, A. 1852. Mollusca and shells, X'olume 12, 510 pages,
atlas in US. Exploring Expedition during the vears 1838,
1839, 1840, 1841, 1842 under the command of Charles
Wilkes, U. S.N.Boston: Gould & Lincoln.
Gould, A. A. 1861. Descriptions of shells collected in the North
Pacific Exploring Expedition under Captains Ringgold and
Rodgers. Proceedings of the Boston Societv of Natural
History 7: 385-389.
Gould, A..\. 1862. Otia Conchologica: Descriptions of shells
and mollusks, from 1839 to 1862, 256 pp, Boston, Gould
& Lincoln,
Gray, J, E. 1847. A list of the genera of Recent Mollusca,
their synonyma and types. Proceedings of the Zoological
Society of London (for 1847) 15(I78):129-219 (November
1847).
Hedley, C. 1899. The Mollusca of Funafuti, Part 1, -Gaster-
opoda. Australian Museum, Svdnev, Memoir 3, pp, 397-
488.
Houbrick, R.S, 1974, The genus Cerithium in the western
Atlantic. Johnsonia 5(10):33-84, 48 pis.
Houbrick, R.S. 1985. Genus Clypeomorus Jousseaume (Cer-
ithiidae: Prosobranchia). Smithsonian Contributions to Zo-
ology No,403: iv + 131 pp,, 62 figs,
Houbrick, R.S. 1992. Monograph of the genus Cerithium
Bruguiere in the Indo-Pacific (Cerithiidae: Prosobranchia).
Smithsonian Contributions to Zoology No. 510: 208 pp.,
145 figs.
Johnson, R. 1964, The Recent Mollusca of .Augustus .\ddison
Gould. Bulletin of the United States National Museum 239:
v -I- 182 pp., 45 pis.
Kay, E. A. 1965. Marine mollusks in the Cuming collection,
British Museum (Natural Histor\ ) described by William
Harper Pease. Bulletin of the British Museum (Natural
Histor\ ), ZoologN , Supplement 1, London 96 pp., 14 pis.
Kay, E,A, 1979. Hawaiian Marine Shells Bishop Museum
Press, Special Publication 64(4), Honolulu, Hawaii, 653
PP
Kiener, L.C. 1841. Species genera! et iconograpliie des co-
R. S. Houbrick, 1993
Page 23
quilles vivantes. Genre Cerite Paris vol 5: 10-4 pp , 32
pis.
Kobelt, W. 1888-1898. Die Gattung Ccrithium. in F. H. W.
Martini, & J. H. Chemnitz, Neues systematisches Conchy-
iien-Cabinet ... 1 (pp. 297 pages). Nurnberg: Bauer &
Raspe.
Melvill, J.C. & R. Standen. 1895. Notes on a collection of
shells from Lifu and Uvea, Loyalty Islands, formed b\ the
Rev James and Mrs. Hadfield, with a list of species. Journal
of Conchology 8: 84-132, pis. 2-3.
Mant, C. F. 1923. Mollusca from the dredging operations at
Kewalo Harbor, Honolulu, 1921. The Nautilus 36(4):120-
123.
Ostergaard, J.M. 1939. Reports on fossil Mollusca of Molokai
and Maui. Occasional Papers of Bernice P. Bishop Museum
Honolulu Hawaii 15(6): 67-77.
Pease, W. H. 1861, Descriptions of Forty-seven New Species
of Shells, from the Sandwich Islands, in the Collection of
Hugh Cuming, Esq. Proceedings of the Zoological Society
of London 431-438.
Pfeiffer, L 1840. Ubersichtder im Januar, Februar und Marz,
1839 auf Cuba gesammelten Mollusken. Archiv Natur-
geschite6(l);250-261.
Salvat, B., & C. Rives. 1975. Coquillages de Polynesie. 392
pp, illustrated Papeete, Tahiti.
Schepman, MM. 1909. The Prosobranchia of the Siboga Ex-
pedition. Part 2. Taenioglossa and Ptenoglossa. Leyden,
Brill, 7 pis.
Sowerby, G.B. 1855 Monograph of the genus Cerithium.
.'\danson. In Thesaurus Conch) liorum, or monographs of
genera of shells. London, 2(16);847-899, pis. 176-186.
Sowerby, G.B. 1865-66. Monograph of the genus Cerithium
Adanson in: Reeve, Conchologia Iconica: or illustrations
of molluscous animals. London. 15 (atlas), 20 pis. + index.
Tiberi, N. 1863. Description d'especes nouvelles de la mer
Mediterranee. Journal de Conchyliologie 11: 158-160.
Tryon, G.W. 1887. Manual of Conchology; structural and
systematic with illustrations of the species (Cerithiidae),
First series. Philadelphia, 9: 112-228, 19-39 pis.
Vignal, L. 1903. Sur les varietes du Cerithium zebrum Kiener.
Journal de Conchyliologie 51(l):21-27, pi, 2.
THE NAUTILUS 107(1 ):24-28. 1993
Page 24
Rediscovery of TurhineUa thersites Reeve, 1847, with Notes on
its Taxonomic Position (Gastropoda: Fasciolariidae)
R. N. Kilburn
Natal Museum
P/Bag 9070
Pietermaritzburg 3200, South Africa
ABSTRACT
The holot\pe of Turbinella thersites Reeve, 1847, was believed
to have been collected in China; no further specimens have
subsequently been reported. Comparison with recently trawled
material indicates that it was actually dredged on the Agulhas
Bank. Turbinella thersites is here transferred from the family
Turbinellidae to the genus Fasciolaria, within the family Fas-
ciolariidae.
Key words: Fasciolariidae; Turbinellidae; South African; ne-
ritic; Fasciolaria. Turbinella
INTRODUCTION
Turbinella thersites Reeve, 1847, was based on a single
specimen (Fig. 3) originally in the collection of Captain
Sir Edward Belcher, and supposedK- from China. Noting
that it appeared to be immature, Ree\e obser\ed that
"it rather partakes of the character of Fasciolaria", a
view echoed b\- Melvill (1891), who speculated that it
might belong to either Fasciolaria Lamarck, 1801 (fam-
ily Fasciolariidae) or Tudicla Roding, 1798 i^Turbinel-
lidae). Kobelt (1874) cited it as Turbinella (Plicatella)
thersites, and Tryon (1881) similarly included it within
the genus Latirus Montfort, 1810 (= Plicatella S\\ ainson.
1840). FinalK', Yen (1942), in a paper on C^hinese gas-
tropods in the Natural Histor\ Museum, London, pub-
lished a photograph of the holot\pe, under the name
Tudicla thersites, and quoted the type locality without
comment. No specimens additional to the holot\pe are
on record, and no further reference to the species has
been traced, notwithstanding the considerable bod\' of
literature that has been published in recent decades on
the mollusks of the Sino-japanese region.
,\mong the larger mollusks that are brought up in the
nets of commercial trawlers operating on the .Vgulhas
Bank, the continental shelf south of South Africa, are
several species of Fasciolaria. Most commonly trawled
are F. scholvieni Strebel, 1912, bathymorphs of F. lu-
gubris lugtdnis Ree\e. 1847, and its eastern sulxspecies
F. lugubris heynemanni Dunker, 1871, and the bathyal
F. rutila (Watson, 1882); less commonly seen is F. uat-
tersac Kilburn. 1974. On rare occasions a fifth species.
distinguishable by its sculpture of auriculate ribs, is
trawled. Such ribs similarK' characterize the "lost" Tur-
binella thersites, and comparison of juvenile specimens
of this "fifth species" with the equally juvenile holotype
of T. thersites confirms their identity .
.At the time of description of Turbinella thersites, the
only .\gulhas Bank mollusks known had been dredged
during the cruises of the H.M.S. Sulphur USo6-1842)
and H.M.S. Samarang (1843-1846'. both captained by
Belcher. Confusion of localities in Belcher material from
these voyages is not infrequent, and using the line of
reasoning ad\anced b\ Tomlin (1925: 310). with regard
to the origins of the holotvpe of Turrit ella ferruginea
Ree\e. 1849, one can speculate that the holotv pe of Tur-
binella thersites was originalK amongst the mollusks
dredged by Belcher on the .\gulhas Bank during the
vo\age of H.M.S. Samarang i^this expedition was spe-
cificalK mentioned b\ Ree\e under the description of
Turbinella [= Latirus] belcheri Ree\e. 18471 Indeed
the case of Turrit ella declivis .\dams & Reeve. 1S50. is
directly comparable: although originalK said to have
been dredged b\ the Samarang in the "China Sea", this
species was subsequentK shown i,Tomlin. 1925: 313) to
be characteristic of the Agulhas Bank biota. Although at
present little precise data is a\ ailable for Fasciolaria ther-
sites. indications are that, like Turritclla dcclivis. it is a
continental shelf species endemic to the .\gulhas Bank.
The second known example (Fig. 5) of Fasciolaria
thersites was actually collected in 1962. but its identity
w ent unrecognized, until it w as .sent to me for comment
in 19S(i b\ J. H. McLean of the Los .\ngeles Count)
Mviseum. CuriousK . a.ssociated with it w as a similar error
in localization. Originating from the collection of the late
I'ranz Steiner. the label states: "This shell w as found in
dredged santl. Nacala, Mozambique, on the 19 June,
19ti2, b\ nic . BiogeographicalK , it would be unprece-
dented for such a temperate-w ater .\gulhas Bank species
to live also on the tropical coral coast of northern Mo-
zambique, and its origin is sureK ad\ entitious. It is not
impossible that this specimen w as jettisoned from a ship
docking in Nacala harbor, but although worn, it lacks
the porcellaneous appearance normally seen in shells that
ha\e lain on coral sand for a length) period. It is thus
R. N. Kilburn. 1993
Pa2e25
Figures 1-2. Protoconch of Fasciolaria thersites Reeve. 18471 Scale lines = 1 mm.
more probable that Steiner accidentalK separated it from
a sample of trawled Cape moUusks. and mLxed it with
his Mozambican finds. Indeed, in 1962 or 196-3 I was
personalK told b\ Steiner then a radio-operator in the
merchant navy) that while on the Xacala route he reg-
ularK received trawled shelb from a source in Cape
Town, who accumulated them in barrels of preserv ative
on his behalf.
This species and its temperate-water South .\frican
congeners agree in teleoconch characters with the genus
Fasciolaria Lamarck. 1799. in its broadest sense. Radula
characters, known for all species v Kilbum. 1974. Barnard.
1958). except F. thersites. also agree, .\lthough species
with shoulder nodules are generally referred to subgenus
^sometimes even genus) Pleuroploca P. Fischer. 1SS4.
such a classification does not address the problem of ta.xa
such as Fasciolaria filamentosa \R6ding. 1798) and F.
lugubris Reeve. 1S47. in which nodules are present or
absent according to individual or population. The fi\e
species endemic to South .\frican waters share a close
morphological similarity but differ from all other species
of Fasciolaria in their extremely large and bulbous pro-
toconch (Figs 1-2'. which has a basal diameter of 3.0-
5.0 mm. according to species and individual. Such a
protoconch cannot be dismissed as merely an adaptive
correlation with mode of development, because this is
similarly non-planktotrophic in four of the five other
species of Fasciolaria (= Pleuroploca'' in which devel-
opment is known [see D'.\saro. 1970a. 1970b. Gohar &
Eisawy . 1967. Shepherd & Thomas. 1989). and is there
associated with a much smaller protoconch. .\ second
character of possible significance is egg-capsule shape:
in the sole Cape species in which these are known {F.
lugubris\ capsules differ from those reported for other
species of Fasciolaria in being dorsally rounded without
a demarcated escape hatch ^see Kilburn & Rippey. 1982:
text fig 62 i. Egg capsules have been described for three
species from North .\merica. and one each from the
tropical Indo-Pacific, Mediterranean and southern -\us-
traha ; references given above, also BaccL 1947). In all
of thec-e. the capsule is flat-topped, with a structural es-
cape hatch dorsally. It is possible that recognition of the
South -\f rican species-complex at the subgenxis level may
ultimately prove warranted on grounds of protoconch
size and perhaps capsxile form.
Fasciolaria thersites
Figures 1-7
Reeve. 1S47 , n. comb.
Turbinella thersites Reeve. 1S47: pi. 4. sp 21: MeKiU. 1S91:
409. Type localit> : China [here emended to .\gulfaas Bank].
Turbinella , Flicatella '• thersites: Kobelt in Kster & KobelL 1574:
71. 157. pL 18. fig. 1 ^afte^ Reeve': Kobelt 1S76: 21.
Lc-'":' -rf-rfites: Tryon. 1851: 91. pL 6S. fig. 137 .after Reeve).
7 '-'^tes: Yen. 1942; 237, pL 25. fig. 17S iholot>-pei.
r ^>--- .i^'-.i^ icattersae ynon Kjlbum. 1974 : Richards. 1981: 63,
pi. :35. fig. .300.
Description: Shell fusiform, with bluntly papiUiform
apes, slphonal canal equal in length to aperture, curved
slighdv to right: suture moderately shallow, whorls
strongly convex, except immediately below suture where
they are shaUowly concave. Inner fip with a thin callus
glaze, smooth except for a thin spiral ridge just above
mid-parietal region and three obhque basal pleats; of
these the strongest pleat is situated at entrance to siphonal
canal and the weaker posterior two decrease in strength
and are scarcely visible from without the aperture. Te-
leoconch sculptvtred by strongly prosochne. rather turn id
axial ribs, which project at shoulder in a roundedly au-
riculate manner: in most examples, asials become weak
on later whorls, except sometimes for an occasional strong
rib, but they may remain strong throughout: fine spiral
threads, crossed by microscopic coUabral threads, present
overall. First teleoconch whorl with 8 straight prosocline
asials. the first few low and relatively close, thereafter
in transverse section thick, strongly rounded, slightly
Page 26
THE NAUTILUS, Vol. 107, No. 1
R. N. Kilburn, 1993
Page 27
compressed, more or less equal to intervals, which are
flattened; Second whorl with 6-7 axials, third generally
with 5, fourth with 4. Spiral lirae unequal in width,
groups of fine threads being often separated by groups
of coarser ones, those on siphonal rostrum rather uniform
in strength. Collabral threads ma\ be pliculate in places,
generally rendering intervals between spiral lirae punc-
ticulate and the spirals themselves somewhat scabrous.
Off white, with a thin yellowish-brown periostracum.
Protoconch bulbous, of about IV2 whorls, first whorl
strongl) tilted, smooth, last half whorl with uneven, pro-
socline axial riblets; maximum breadth 3.0-4.8 mm,
height 3.1-4.4 mm (ratio breadth/height 0.87-1.13).
Maximum dimensions 102.0 x 39.6 mm (R. Le Maitre
collection). Holotype (juvenile) 49.0 x 22.1 mm. Oper-
culum oblanceolate with rather obtuse apex, moderately
thick with coarse growth lines; color amber-brown, with
darker outer edge.
Distribution: Agulhas Bank,
Type material: Holotype in the Natural History Muse-
um, London, accession number 1879.2.26.116, from
Lombe-Taylor collection.
Other material examined: Agulhas Bank [Natal Museum
(NM) D3482: R. Le Maitre, also several specimens in
colln Le Maitre]; off Cape St. Blaize area, ex pisce (NM
B1087; R. Le Maitre, juvenile); off Storms River mouth,
69 fathoms [= 126 m], in mud, a juvenile shell (NM
S4953: C. Marais); same data, 64 fathoms [= 117 m], a
broken juvenile (NM S5646; C. Marais); "Nacala harbor,
dredged sand", empty shell (NM K1526: F. Steiner, don.
J. H. McLean).
Remarks: I have seen no adults of this species with an
undamaged lip, and only two specimens (both dead ju-
veniles of about 3 teleoconch whorls) bear a precise lo-
cality. Its exact distribution on the Agulhas Bank, both
geographic and bathymetric, consequently remains to be
established.
Fasciolaria thersites shows much variation in size of
protoconch and extent of axial ribbing on the teleoconch;
too little material is available to judge whether this re-
flects geographic, individual or bathymetric variation.
The smallest protoconchs measured occur in the holotype
and in the specimen purportedK' from Nacala (although
it is more or less worn in both examples); in the latter
individual the axial ribs are unusually strong on later
whorls. Similar variability in protoconch size was re-
ported for F. schotvieni Strebel, 1912, by Kilburn (1974).
In appearance F. thersites is most similar to F. wattersae
Kilburn, 1974, an uncommon species which inhabits the
Mossel Bay-Algoa Bay area at depths of about 100-120
m. In F. wattersae (Fig. 7) the axial ribs are never au-
riculate, the spiral sculpture is markedly coarser, and
pliculate axial striae are totally absent. Although appar-
ently inhabiting similar depths, F. thersites is a much
thinner-shelled species. Among the species of Fasciolaria
endemic to South Africa, the operculum in F. thersites
is unique in its amber-brown color, this being dark brown
in the other species.
ACKNOWLEDGMENTS
I wish to thank Ms. Kathie Way of the Natural History
Museum, London, for the loan of the holotype of Tiir-
binella thersites. Messrs. R. Le Maitre and C. Marais for
providing material from the Agulhas Bank, and Dr. J.
H. McLean for donating the Steiner specimen to the
Natal Museum. Drs. D. G. Herbert and B. R. Stuckenberg
read the manuscript, and Mrs. Linda Davis prepared the
line drawing.
LITERATURE CITED
Bacci, G. 1947. Le capsule ovigere di Cohimbella rustica (L.)
e di Fasciolaria lignaria (L.). Bolletino Zoologico 14; 75-
81.
Barnard, K. H. 1958. Contribution to the knowledge of South
African marine Mollusca. Part I. Gastropoda: Prosobran-
chiata: Toxoglossa. ,'\nnals of the South African Museum
44(4):73-163, figs. 1-30, pi. 1.
D'Asaro, C. N. 1970a. Egg capsules of some prosobranchs
from the Pacific coast of Panama. Veliger 13:37-43,
D'Asaro, C. N, 1970b, Egg capsules of prosobranch mollusks
from South Florida and the Bahamas and notes on spawn-
ing in the laboratory. Bulletin of Marine Science. 20:414-
440.
Gohar, H. A. F. & A. M Eisawy. 1967. The egg masses and
development of five rachiglossan prosobranchs from the
Bed Sea Publications of the Marine Biological Station of
Ghardaqa. 14:215-268.
Kilburn, R. N. 1974, Taxonomic notes on South African ma-
rine Mollusca (3): Gastropoda, Prosobranchia, with de-
scriptions of new taxa of Naticidae, Fasciolariidae, Ma-
gilidae, \'olutomitridae and Turridae, Annals of the Natal
Museum 22(1): 187-220,
Kilburn, R.N. & E. Rippey, 1982. Sea Shells of Southern
Africa. Johannesburg: Macmillan.
Kobelt, W. 1844-1876. Turbinella und Fasciolaria. In Kster,
H. C. & Kobelt, W., Systematisches Conchylien-Cabinet
von Martini und Chemnitz, 2nd Edition, 3(3a):l-164, pis,
1-32,
Melvill, J, C, 1891. An historical account of the genus La^irus
(Montfort) and its dependencies, with descriptions of elev-
en new species, and a catalogue of Latirus and Peristernia.
Memoirs and Proceedings of the Manchester Literar\ and
Philosophic Society [4] 4:365-411.
Figures 3-7. Fasciolaria thersites (Reeve, 1847) and F. wattersae Kilburn, 1974 Fasciolaria thersites: 3. Juvenile holotype of
Turbinella thersites Reeve, 1847, Natural History Museum, London, 1879 2.26.116, 49.0 x 22 1 mm. 4. Immature example from
Agulhas Bank, 78.9 x33,6 mm, in R, Le Maitre colln, 5, Specimen supposedly from Nacala Bay, NM K1526, length 76,2 mm. 6.
Adult example, Agulhas Bank, depth unknown, NM D3482, length 100,2 mm, outer lip damaged. 7. Fasciolaria wattersae Kilburn,
1974, S. E. of Mossel Bay, 119 m, NM E5898, 98.7 x32.3 mm.
Page 28
THE NAUTILUS, Vol. 107, No. 1
Reeve, L. A. 1847. Monograph of the genus Turbinella. Con-
chologia Iconica -1. London: Reeve
Richards, D. 1981. South African Shells. A collector's guide.
Cape Town: Struik,
Shepherd, S. A. & I. M. Thomas. 1989. Marine Invertebrates
of southern Australia. Part II. .Adelaide: S. Australian Gov-
ernment Printing Division.
Tomlin, J. R Le B. 1925. Reports on the marine Mollusca in
the collections of the South African Museum. I. Turritel-
lidae. .Annals of the South .\frican Museum 25:309-316.
Tryon, G. W. 1881. Tritonidae, Fusidae, Buccinidae. In:
Manual of Conchology 3. Philadelphia, privately pub-
lished.
Yen, T. C. 1942. A review of Chinese gastropods in the British
Museum. Proceedings of the Malacological Society of Lon-
don. 24:170-289, pis. 11-28.
THE NAUTILUS 107(1 ):29-32. 1993
Page 29
The Rediscovery, Morphology, and Identity of
Conus emersoni Hanna, 1963
John K. Tucker
llliiKiis Natural Histur\ Survey
LTRMP
P.O. Bo.\ 36S
West Alton, MO 63386, USA
James H. McLean
Natural History of
Los Angeles County
900 E.xposition Boulevard
Los Angeles, CA 90007, USA
ABSTRACT
Conus emersoni is reported from 310 ni off Isla Sanla Maria
(Floreana), Galapagos Islands, the first record subsequent to its
original description. The species is redescribed from eight ad-
ditional specimens; its operculum, raduia, and periostracum
are described for the first time. Affinity of the species to C.
teramachii Kuroda, 1956, is discussed.
Key words: Conidae, Contis emersoni. Galapagos Islands, rad-
uia.
INTRODUCTION
Subsequent to its description 29 years ago (Hanna, 1963)
from two dead, faded and poorly preserved specimens,
Conus emersoni has remained an enigmatic member of
an otherwise extensively studied genus in the Eastern
Pacific region. Even as he proposed the species, Hanna
(1963) suggested that it might be conspecific with the
Indo-West Pacific species C. australis Holten, 1802,
whereas Walls (1979) placed it as a possible synonym of
another Indo-West Pacific species, C. orbigniji Audouin,
1831. Keen (1971) treated the species as valid but made
no comments on its relationships. Finally. Coomans et
al. (1986) considered it to be a tentatively valid species
but noted that it could be based on fossil material.
Here we report upon eight recently collected speci-
mens from moderately deep water at the Galapagos Is-
lands, two of which are in the collections of the Natural
History Museum of Los Angeles County and six in the
American Museum of Natural History. All were provided
by Andre and Jacqueline DeRoy, residents of the Ga-
lapagos Islands. These specimens extend the distribution
from Cabo San Lucas, Baja California Sur, to the Ga-
lapagos Islands, Ecuador, and allow a redescription of
the species with the first report on the morphology of
the radular tooth, the operculum, and the periostracum.
Abbreviations of museums mentioned in the text:
AMNH, American Museum of Natural History, New
York; CAS, California Academy of Sciences, San Fran-
cisco; LACM, Natural History Museum of Los Angeles
County.
SYSTEMATICS
Conus emersoni Hanna, 1963
(figures 1-13)
Contis emersoni Hanna, 1963;25, pi. 1, fig. 2; Walls, 1979:776
[with unnumbered figure of CAS paratypej; Coomans,
Moolenbeek & Wils, 1986:11-4, fig. 718 [holotype, 2 views].
Conus (Asprella) emersoni. Keen, 1971:663. fig. 1497 [holo-
type].
Diagnosis: Spire scalariform throughout its length,
shoulder angle retaining square nodules; sculpture on
final whorl of numerous, closely spaced, shallow sulci;
posterior notch shallow; whorl tops sculptured by one to
two cords that fade in whorl three to be replaced by
numerous fine striae.
Description: Shell elongate-conical, whorl sides flat to
very slightly convex; shoulder angular. Anterior end not
deflected dorsally. Sculpture of numerous (30-50) closely
spaced, shallow sulci, most pronounced at anterior end
and fading in intensity towards shoulder. Color pattern
variable, consisting of three irregularly developed bands,
at shoulder, in area just posterior to midbody area, and
in area just anterior to midbody. Bands variously inter-
rupted and scalloped, producing reticulate or blotched
pattern; bands separated by areas with rows of spiral
dashes or longitudinal reticulations. Dashes and reticu-
lations may be quite pronounced (holotype) or lacking
(Galapagan specimens). Anterior end marked by spirally
elongated blotches or lines. Color markings in fresh spec-
imens medium brown, fading to light brown in dead
specimens.
Spire very slightly convex in profile, scalariform
throughout and moderately elevated, carinate, carina
interrupted by numerous square nodules, 30-40 per whorl;
nodules fading in whorl six although carina may remain
pronounced. Protoconch unknown (not intact in any of
the specimens). One or two spiral cords on first two or
three teleoconch whorls, replaced on later whorls by
numerous, exceedingly fine striae. Whorl tops colored
by variably developed markings, matching those of body
whorl, between nodules or crossing entire width of whorl
Page 30
THE NAUTILUS, Vol, 107, No. 1
Figure§ 1-7. Conus emcrsurii Hanna, 1963 Figures 1-3. Type material dredged off Los Fraiies, Baja California, depth 549
111 L Holotype, AMNH 1052n, length 43 0 mm, faded, dead-collected specimen, lacking periostracum. 2, 3. Paratype, CAS
12405, length 49.0 mm, subfossil specimen (surface gray), showing naticid bore hole. Figures 4-7. Newly reported specimens
dredged off Isla Santa Maria (Floreana), Galapagos Islands, Ecuador, depth 310 m. 4. AMNH 248261, length 46.2 mm, dead-
collected specimen. 5. LACM 146906a, length 33.3 mm, live-collected specimen with periostracum intact. This specimen was used
for opercular and radular illustration. 6. 7. LAC;M 146906b, length 34 0 mm, dead-collected specimen with periostracum removed
to show color pattern.
top. Whorl tops slightly but distinctly concave in cross
section. Posterior notch shallow and C-shaped.
Aperture narrow, white inside e.xcept where exterior
coloration shows through near lip. Interior constrictions
and aperturai flanges absent or at least not developed in
available specimens.
Periostracum thick, dark brown, markedly pilose both
on body and spire. Extremely Bne hairlike extensions of
periostracum not organized into any obvious pattern.
These projections are apparently easily worn off, as they
are pronounced on the dorsal side of one live-collected
specimen (figure 5) but are not obvious on the ventral
side of the same specimen.
Dimensions (.see table I): The known specimens range
in length from 23.0 mm to the 49.0 mm length of the
paratype (figures 2, 3). The ratio of width to length ranges
from 0.43 to 0.50 (table 1 ). The largest Galapagan spec-
imen (figure 4) is 46.2 mm in length. All specimens have
the protoconch eroded, the spire tips filled by secondary
shell deposition.
Operculuni illustrated in figure 8 is 27% of aperturai
length, weakly serrate on outer edge.
Kadular tooth (figures 9-13) small, 285 jum in lengtli;
massive base with pronounced basal spur; waist located
on anterior half of tooth, with small spine on posterior
border in same plane as basal spur. Tooth opening rather
long, extending about one-cjuarter the length. Tip w ith
small barb opposite short blade.
Distribution: The holotype and one paratype were col-
lected oft Los Fraiies, Cape San Lucas, Baja California
in 550 m (given originally as 300 fathoms by Hanna,
1963). The eight newly discovered specimens were
dredged in 310 m (170 fathoms) off Isla Santa Maria
(Floreana), Galapagos Islands, 7 May 1979 by Andre and
Jacqueline DeRoy (table 1).
DISCUSSION
The rediscovery of Conus emersoni in the CJalapagos
Islands should put to rest speculation that this species is
a synonym of either Conus australis (suggested by Han-
na, 1963) or C. orbignyi (suggested by Walls, 1979). Each
of these Indo-Pacific species is unlike C. emersoni in
having persistent sulci on the body whorl along with
persistent cords on the spire \\ horl tops. C'onus emersoni
differs in having the sulci strong only anteriorly and in
having fine spiral striae on the whorl tops. Radular dif-
ferences are that the radular tooth of C. orbignyi has
three anterior barbs (Kilburii, 1973: fig 6), whereas that
J. K. Tucker and J. H. McLean, 1993
Page 31
.-H*
W^
»
m0
10
11
Figures 8-13. Conus emcrsoni Hanna, 1963 8. Operculum, same specimen as figure 5, length 7 3 mm, nucleus lost. Figures
9-13. SEM views of single radular toolli, length 285 m'". from same specimen as in figure .5. SEM photos by H. Chaney 9.
Oblique anterior view showing apical surface, with pronounced basal spur and small spine on posterior border of waist. 10. Enlarged
view of apical surface, with small barb and enrolled blade. 11-13. Three lateral views, showing differing axial rotation of tooth
of C. emersoni has but two. The radular tooth of C.
australis has a serrate shaft along with an enlarged cusp
at the posterior end of the row of serrations (A. J. Kohn,
radular slide collection); neither feature is present on the
radular tooth of C. emersoni.
The scalariform spire and color markings of Conus
emersoni have a superficial resemblance to that of the
Panamic species C. emarginatus Reeve, 1844 (a species
often misidentified as C. recurvus Broderip, 1833). Dif-
ferences are that C. emarginatus has a deep posterior
notch (rather than shallow notch) and a smooth (rather
than pilose) periostracum. The radula of C. emarginatus
(Nybakken, 1970; fig. 5, as Conus recurvus) has three
barbs anteriorly, like that of C. orbignyi, rather than C.
emersoni, in which there are two barbs.
The scarcity of C. emersoni may be due to the great
depth at which it occurs and the paucity of sampling at
depths below 300 m.
It is possible that the specimens from the Galapagos
Islands are specifically or subspecifically distinct. They
differ in color pattern (that of the holotype being more
intricate) and in spire profile (that of the holotype ap-
pearing to be more acute). However, such intraspecific
differences in color pattern are not uncommon in Conus.
In fact, the faintly indicated color pattern of the paratype
that shows on the better preserved dorsal surface (figure
3) is more similar to that of the Galapagan specimens
(figures 5-7) than to that of the holotype (figure 1). The
ground color of the paratype has a gray cast that is
suggestive of fossil or subfossil condition. The holotype
is in fresher condition, although it retains no periostra-
cum; it shows evidence of growth damage to the shoulder
at a stage two whorls above the termination of the lip,
which appears to have altered the profile of the final two
whorls, resulting in a greater downward slope to the spire
and the loss of the nodules in the final two whorls.
The question as to whether the two widely disjunct
records represent the same species can only be answered
after additional specimens from the vicinity of the type
locality become known. It is our opinion, based on the
specimens examined, that there is no clear evidence to
suggest that the Galapagan specimens are not conspecific
with the Mexican specimens.
Conus emersoni is not closely similar to any other
eastern Pacific conid. Shell morphology, including the
Table 1 . Shell dimensions and proportions of the known spec-
imens of Conus emersoni Hanna, 1963.
Width/
Specimen
C:ondition
Length
Width
length
AMNH 92200 (holotvpe
dead
43.0
18.5
0.43
CAS 1240.5 (paratype)
subfossil
49.0
22.5
0.46
LACM l-)6906a
live
33.9
17.0
0.50
LAC:M 146906b
dead
340
15.5
0.46
AMNH 248262
live
35.0
15.7
0.45
AMNH 248263
live
31.5
14.3
0 45
AMNH 248261
dead
46.2
22.3
0.48
AMNH 248169a
live
23.0
10.8
0.47
AMNH 248169b
dead
30.0
149
0.50
AMNH 248169c
live
26 8
12 8
0.48
Page 32
THE NAUTILUS, Vol. 107, No. 1
14
15
Figures 14, 15. Conus teramachii Kuroda, 1956 14. Speci-
men with growth scar producing lowered shoulder angle of
final whorl, periostracum removed. Off NE coast Taiwan,
trawled, depth unknown. LACM 68994, length 79 8 mm. 15.
Operculum with strongly serrate edge, nucleus lost, trawled off
Taiwan. Specimen in H. Chaney collection, length 19.1 mm.
shoulder carina with square nodules, is similar to that of
C. teramachii Kuroda, 1956 (figure 14), a species re-
ported from similar depths off Japan and Taiwan (Ku-
roda, 1956; Walls, 1979). Our illustrations of the radular
tooth of C. emersoni (figures 9-13) are comparable to
those of Azuma (1961: fig. 11) for C. teramachii (as C.
petricosus Azuma, 1961). In addition, C. teramachii also
has a strongly serrate operculum (figure 15), much more
pronounced than that of C. emersoni (figure 8). The
character state of the serrate operculum was stressed in
the original diagnosis of the subgenus Profundiconus
Kuroda, 1956 — type species Chehjconus {Profundico-
nus) profundorum Kuroda, 1956 [= Conus smirna
Bartsch and Rehder, 1943]. The type species of Profun-
diconus also has square nodules on the shoulder carina,
but these are apparent only in young stages.
We refrain, however, from further treatment of the
subgeneric allocation of the species under discussion be-
cause comparison to other available generic level taxa is
beyond the scope of this paper. We recognize that a
generic level classification of Conidae needs to be based
on all the recognized species, including fossils, at a min-
imum treating characters that include adult and juvenile
shell morphology, and, for the living species, the oper-
culum, and radula.
ACKNOWLEDGMENTS
We thank Alan J. Kohn of the L'niversity of Washington
for allowing one of us (JKT) to study his radular slide of
C. australis. William K. Emerson and Walter E. Sage of
the American Museum of Natural History made the ho-
lotype and other specimens of C. emersoni available for
study. Terrence M. Gosliner of the California Academy
of Sciences arranged for loan of the paratype specimen.
We thank Henry W. Chaney of the Santa Barbara Mu-
seum of Natural History for the SEM photos of the rad-
ular tooth of C. emersoni and the photograph of the
operculum of C. teramachii. We are grateful to Henry
W, Chaney, William K. Emerson, Alan J. Kohn and
Walter E. Sage and two anonymous referees for reading
the manuscript and offering helpful suggestions. Finally,
and most importantly, we thank Jacqueline and Andre
DeRoy for providing the specimens of C. emersoni that
form the basis for this paper.
LITERATURE CITED
Azuma, M. 1961. Descriptions of six new species of Japanese
marine Gastropoda. Venus 21(3):296-303.
Coomans, H E., R. G. Moolenbeek, and E. Wils. 1986. Al-
phabetical revision of the (sub)species in recent Conidae.
9. ehraeus to extraordinaritis with the description of Conus
elegans remalhoi. nov. subspecies. Basteria50;93-1.59, figs.
667-760.
Hanna, G. D. 1963. West American mollusks ot the genus
Conus — II. California Academy of Sciences, Occasional
Papers, no. 35, 103 p., 11 pis.
Keen, A. M. 1971. Sea shells of tropical West America, second
edition. Stanford University Press, Stanford, C.-^, 1064 p.
Kilburn, R. N. 1973 Notes on some benthic Mollusca from
Natal and Mozambique with descriptions of new species
and subspecies of Calliostoma, SolarieUa. Latiaxis. Bab-
ylonia, Fusinus. Bathytoma and Conus. Annals of the
Natal Museum 2I(3):557-578, figs. 1-17.
Kuroda, T. 1956. New species of the Conidae (Gastropoda)
from Japan. Venus 19(1): 1-13, 1 pi.
Nybakken, J. 1970. Radular anatomy and systematics of the
West .'\merican Conidae (Mollusca, Gastropoda). Ameri-
can Museum Novitates 2414:1-29, figs. 1-45.
Walls, J. G. 1979. Cone shells, a synopsis of the living Conidae.
T. F. H. Publications, Inc., Neptune City, NJ, 1011 p.
THE NAUTILUS 107(l):33-42, 1993
Page 33
The Benthic Mollusk Faunas of Two Contrasting Reef
Paleosubenvironments: Falmouth Formation (late Pleistocene,
Last Interglacial), Jamaica
Stephen K. Donovan
Department iil Ch'iiIoi;\
University of the West Indies
Mona, Kingston 7 JAMAICA
D.T.J. Littlewood'
Haskin Shellfish Heseareh Laboratory
Institute of Marine and Coastal
Sciences
Cook College/ NJ A ES
P.O. Box B-8
Port Norris, New Jersey 0S349 l!SA
ABSTRACT
The last interglacial (about 125,000 years old) raised reefs of
the Falmouth Formation of Jamaica contain a diverse fauna
of benthic mollusks. Large collections of mollusks ha\e been
made from two contrasting reef localities and paleosubenvi-
ronments. Calcareous muds of lagoonal origin were well-ce-
mented, biassing collecting tow ards larger individuals, whereas
poorlv-cemented santly sediments of a coral framework were
friable and thus amenable to bulk sampling, yielding numerous
micromoUusks. Identification of over 3,000 bivalve, gastropod
and scaphopod specimens shows the molluscan faunas of each
environment to be dominated by different species. The coral
framework fauna is numerically dominated b\ the microgas-
tropod Caecitm piilchcllinu Stimpson, 1851, other epifaunal
gastropods and arks, while Bulla .striata Bruguiere, 1792 and
ceriths dominate the lagoonal sediments. Only Cerithium al-
gicola C.B. ,^dams, 1848, forms more than 2"; of the fauna at
both localities. 17% of the extant species (27% of the common
species) of shallow water benthic mollusks (excluding chitons)
from Jamaica have been identified from these two localities.
Key words Late Pleistocene; benthic reef mollusks; Jamaica.
INTRODUCTION
The present paper is a preliminary report of the mol-
luscan faunas from the late Pleistocene (the last inter-
glacial or Sangamonian) Falmouth Formation (125,000
years B.P.) of northern Jamaica. The mollusks of the
Falmouth Formation are important for two principal
reasons. Firstly, the Neogene and Quaternary fossilif-
erous deposits of Jamaica contain a diverse fauna of
benthic mollusks that have hitherto been largely ignored
' Current address; Department of Palaeontology, British Mu-
seum (Natural Historv), Cromwell Road, London. S\V7 5BD,
ENGLAND
by systematists. Indeed, only the famous Pliocene Bow-
den shell bed in southeastern Jamaica (Woodring, 1925,
1928) and the fauna of the Early Pleistocene Manchi-
oneal Formation (Trechmann, 1930) have so far received
monographic treatment. This lack of published infor-
mation does not reflect the true diversity of the mollusk
faunas through this interval and ma\ thus prove mis-
leading to workers not familiar with the geology of the
island. For example, Petuch (1988:49) considered that
there are few exposures of Miocene age preserved within
the Caribbean and South American region, and did not
include Jamaica in his list of productive sites, yet in a
preliminary survey Jung (1972) listed members of 19
genera from the extensive deposits of the Newport For-
mation (Lower and Middle Miocene) of the island.
Further, molluscan faunas from last interglacial reefs
(and other Pleistocene marine environments; Valentine,
1989) can be a valuable adjunct to studies of extant as-
semblages from the shallow benthos (but see cautionary
comments in Johnson, 1960). The Pleistocene raised reefs
of the Caribbean region are recognised to include large
faunas of fossil mollusks (see, for example, Cerridwen &
Jones, 1991). Unlike those of certain other areas (such
as Aldabra Atoll; Taylor, 1978), Caribbean mollusk fau-
nas have apparently changed little since the last inter-
glacial (Petuch, 1988:119), an observation that may have
important implications for the study of faunal associa-
tions in Recent reefs and associated environments. Mol-
lusk faunas in various biofacies w ithin Pleistocene raised
reefs are generalK easy to sample, outcropping adjacent
to the coast on many Caribbean islands, thus permitting
large collections to be made for systematic and paleo-
ecological analysis.
Geology and geological nomenclalure: The Falmouth
Formation in the Discovery Bay-Rio Bueno Harbour re-
gion of Jamaica's northern coast (figures 1-3) rests un-
conformably on the dolomitized Plio-Pleistocene reef
(Land, 1991) of the Hopegate Fonnation. Lithofacies
Page 34
THE NAUTILUS. Vol. 107, No. 1
2
1^
^
^
0
km
1
' ^-—
^^=^^8=-
^^2
\
^ ^
1^ ^^
%
■^1
DB Jy
^"¥1
i
,54-70
^)
Ftl
W/!?\ TerrigencHJS grainstcxie facies
Mi::::iij Mdluscan algal gralnstone facies
pt'«';1 Mdkjscan biomlcritic wackestone fades
f n Biomicritic packstone facies
I I Coralline boundstone facies
Hopegate Formation
Figures 1-3. Maps ol slud) arta;. 1 Outline map uf JamaiLa, showing the location of the Discovery Bay (DB) region with respect
to Kingston (K), Montego Bay (MB) and Ocho Rios (OR) 2 Topographic map of the Discovery Bay-Rio Bueno Harbour (RBH)
area, showing the positions of localities 1 and 2. Contours in m. 3 Geological map of the Discover) Bay-Rio Bueno Harbour area
(redrawn after Larson. 19So: figure 6). showing the relative distributions of the major sedimentary facies of the Falmouth Formation
in relation to the underK ing Hopegate Formation (key right).
within the Falmouth Formation reef (Larson. 1983) con-
tain associated distinct faunal assemblages which have
been recognised from the evidence of corals (Larson.
1983: Liddell et ai. 1984) and echinoids (Gordon &
Donovan. 1992).
History of research: Hill (1899) named the Falmouth
Formation, and recognized the essential elements of the
mollusk fauna to comprise Lucina, Cardium. Area. So-
len and Bulla. Hill (1899. 155) considered this fauna to
be worthv- of careful stud\ b\ paleontologists, an invi-
tation that has not been taken up in over 90 \ ears. Rob-
inson (1958) noted Area spp. from the elevated reef at
east Rio Bueno Harbour (localitv 1 herein), and Chione
cancellata (Linnaeus. 1767) and Bulla cf. striata Bru-
guiere. 1792. from the lagoonal facies east and west of
Falmouth. Parish of Trelawny (probabh lithologicalK-
similar to locality 2 herein). In 1960. Robinson considered
the Falmouth Formation in the Parish of St. .Ann to
include the moUusks Lueina. Codakia. Chione and Bul-
la, and later commented that this rich fauna is identical
with the living fauna of the Jamaican coastal waters
(Robinson. 19&3:52). Recent studies of the fossil scapho-
pods of Jamaica (Donovan and Littlewood. 1989; Don-
ovan, 1990) ha\e recognised Dentalium sp. cf. D. (An-
atalis) antillarum d'Orbigny, 1842, and D. (Graptacme)
semistriolatum Guilding. 18;34. from the east Rio Bueno
Harbour locality.
The specimens discussed herein are deposited in the
Department of Palaeontology, British Museum (Natural
History). For uncertainties in our taxonomic assignments
(particularly for poorly preserved or fragmentary spec-
imens), we have used question marks thus: Agenus ?spe-
cies signifies that we are uncertain of the specific assig-
nation: ? Agenus aspeeies indicates that some doubt exists
concerning the generic land. thus, also specific! assign-
ment. This differs in detail, but not in spirit, from the
recommendations of Bengston (1988).
MATERIALS AND METHODS
The Falmouth Formation of the Discovery Bay-Rio Bueno
Harbour area has been subdivided into five principal
reef al subenv ironments by Larson (1983; figure 3 herein).
Large collections of benthic mollusks were made from
two contrasting paleosubenvironments ^localities 1 and
2, described below) within this continuum. These par-
ticular subenvironments and localities were chosen be-
cause of their large mollusk faunas and easy access. The
mollusk collections from each locality were made over
a period of 22 months, from .April 19SS imtil February
1990. Both collections are time-averaged, having been
collected from float, the rock face and as bulk samples
from throughout 2-4 m of section in each case, as bedding
planes and other obvious time marker horizons are not
apparent. Howev er. the \ alue of collecting mollusks from
discrete beds (e\ en when preserved) within reefs is de-
batable, as at an\ one time the fauna w ill include shallow
and deep infaunal. sediment surface epifaunal. and (in
the case of the three-dimensional coral framework at
localitv 1) epifaunal taxa living well above the soft sed-
iment surface.
Specimens collected in the field were generally at least
S. K. Donovan and D. T. J. Littlewood, 1993
Page 35
.y .■^.
J'-^
■->^ ^ , x#
^ ^:.'^'
-<; :^<'
^-ii-rv-^is*; '
m
a.
Figures 4-7. Field photographs of molhisks of the Falmouth Formation 4-. Stromhus gigas Linnaeus, 1758, at locahty 1; 0.3 x.
5-7. LocalitN' 2, 5, MainK Bulla striata Bruguiere, 1792; 0 6 x. 6. General view of weathered surface showing abundant (mainly
molluscan) shell hash; 0.6 x 7. MainK Bulla striata Bruguiere, 1792, associated with an articulated shell (arrowed) of Chione
paphia (Linnaeus, 1767); 0.4 x.
5 mm in maximum dimension and even shells of this
size were easily overlooked without repeated scanning
of a limited area at a time. It proved much easier to
detect small specimens during laboratory analysis using
a Wild binocular microscope. Preliminary field identi-
fications were checked in the laboratory using standard
taxonomic references, including Abbott (1954, 1958,
1974), de Jong and Coomans (1988), Fisher (1988), Hum-
frey (1975), Morris (1987), Vaught (1989), and Warmke
and Abbott (1961). Total numbers of whole shells and
(for bivaUes) disarticulated valves are tabulated \\'hole
shell equivalents (discussed in Cerridwen and Jones, 1991 )
were used in calculating relative proportions of moUusks
for graphical analysis.
Locality 1
(Locality 7 of Liddell et a/., 1984; locality 3 of Donovan
6 Gordon, 1989). East side of Rio Bueno Harbour, Parish
of St. Ann, with the Falmouth Formation exposed as a
line of low cliffs (GR 394 572). At this locality the un-
conformable contact with the underlying, dolomitised
Hope Gate Formation (figure 3) formed the hard sub-
strate for Falmouth Formation reef growth. Previous
interpretations have considered this locality to represent
either a patch reef in the back reef lagoon within the
molluscan biomicritic wackestone facies sensii lato (Lar-
son, 1983; figure 3 herein) or perhaps part of the shallow-
fore reef (Robinson, 1958; Liddell et a!., 1984).
Mollusk-rich sediments at this locality occur beneath
an altered caliche cap (Land & Epstein, 1970) in a frame-
work of shallow-water corals, particularly Porites spp.
and Acropora spp. (Liddell et al., 1984: 77). The matrix
between corals is a friable, weakly-cemented carbonate
sand which includes a diverse benthic fauna, including
echinoid (Gordon & Donovan, 1992) and ophiuroid
(Donovan et al., in press) ossicles, crabs (Morris, in press),
disarticulated chiton valves (Donovan et al., research in
progress), scaphopods (Donovan & Littlewood, 1989;
Donovan, 1990), gastropods and bivalves. Due to the
friable nature of the sediment, two collecting methods
were used. Firstly, mollusk specimens that were weath-
ering out of the exposed face (figure 4) were removed
manuallv. The fauna collected by this method was dom-
inated by bivalves and gastropods, with three scaphopod
shells. Secondly , bulk samples were collected by the bag-
ful. In the laboratory these samples were emptied into
shallow trays, dried in an oven overnight, then washed
through a nest of sieves using cold water and redried.
Sieved aliquots were then picked by ey e (coarse fractions)
or under the binocular microscope (fine fractions: frac-
tions finer than coarse sand were not picked). The fauna
collected by this method was dominated by bivalves,
gastropods and (in the finer fractions examined) chiton
valves, but no further scaphopods were found. A total of
1669 benthic mollusks (excluding chiton valves; Donovan
et al., research in progress) have been identified from
this locality (table 1).
Locality 2
(Locality 5 of Liddell et al., 1984). This is an outcrop of
hard, fine-grained, well-lithified, but occasionally chalky,
limestone exposed along the shoreline as a low terrace
east of the Discovery Bay Marine Laboratory, Parish of
St. Ann (GR 405 569). This lithology has been interpreted
as having been formed in a back reef environment (Lid-
dell et al., 1984:76), presumably a lagoon, forming part
of the molluscan biomicritic wackestone facies sensu
stricto of Larson (1983; figure 3 herein).
Page 36 THE NAUTILUS, Vol. 107. No. 1
Table 1. Mollusk fauna collected from locality 1, east of Rio Bueno Harbour. Key: * = includes 11 juveniles, plus 2 shells not
collected; ** = with apparent growth deformity; + = vaKe incomplete; ++ = probably mostly Area iiiibricala Bruguiere; S =
shells or identifiable shell fragments; \' = valves; LV = left valve; RV = right valve; O = operculum; J = juvenile shells.
Gastropods
Diodora listen (d'Orbigny, 1853) 25S
Diodora '^caijencnsis (Lamarck, 1822) 12S
Diodora '■'mintitn (Lamarck, 1822) IS
Fi.ssiirclla harhadcnsis (Gmelin, 1791) IS
Acinuea ?(inlillarinn (Sowvrhy, 1831) IS
Acmaea pustidata (Helbling, 1779) 60S
Cittarium pica (Linnaeus, 1758) ,3S
Tegula fasciata (Born, 1778) ,3S
Astraea ?caelata (Gmelin, 1791) lO
Astraea tccta (Solander, 1786) 4S
Nerita versicolor Gmelin, 1791 IS
Nerita tessellala Gmelin, 1791 2S
Nerita sp. indet, j^S
Neritina virginea (Linnaeus, 1758) IS
Neritina punctulata Lamarck. 1816 2S
Littorina ?nebulosa Lamarck. 1822 2S
Alvania ?aberrans C. B. Adams, 1850 IS
Heliacus injundibulijormis (Gmelin. 1791) IS
Petaloconchus irregularis (d'Orbigny, 1842) 2S
Petaloconchus erecius Dall, 1888 IS
Petaloconchus mcgintyi Olsson and Harbison, 1953 2S
Serpulorbis decussata (CmeYm, 1791) 5S
Serpulorlris ?decussata (Gmelin, 1791) IS
Serpulorbis ?riisei (Morch, 1862) IS
Caecum ?insularum Moore, 1970 IS
Caecum plicatum Carpenter, 1858 IS
Caecum pulchcllum Stimpson, 1851 657S
Modulus modulus (Linnaeus, 1758) 13S
Cerithium ''litteratum (Born, 1778) 7S
Cerithium guinaicum Philippi, 1849 3S
Cerithium ?eburneum Bruguiere, 1792 2S
Cerithium algicola C. B. Adams, 1848 92S
Cerithium sp. 3S
Cerithium sp. indet. 3S
Hipponix antiquatus (lAnna.eus, 1767) ,3,3S
Strombus gigas Linnaeus, 1758 * 36S
Cypraea zebra Linnaeus, 1758 Ij
Cypraea cinerea Gmelin, 1791 2S
Polinices lacteus (Guilding, 1834) 9S
Morum oniscus (Linnaeus, 1767) 3S
Cymatium 'pileare' (Linnaeus, 1767) IJ
?Cymatium nicobariciim (Roding. 1798) IJ
Cymatium ?parthenopeum (von Salis, 1793) 5S
Cymatium sp. sensu lata H
^Cymatium sp. \\
Bursa cubaniarui (d'Orbigny, 1842) IJ
Columbclla mercatoria (Linnaeus, 1758) 81S
F.ngoniophos 'hmicinctus (Say, 1826) IS
?Melongena mclongena (Linnaeus, 1881) 2S
Fasciolaria tulipa (Linnaeus, 1758) 2S
Latirus ?hrevicaudatus Reeve, 1847 IS
Leucozonia nassa (Gmelin, 1791) 17S
Mitra barbadensis (Gmelin, 1791) 6S
Hyalina avena (Kiener, 1834) IS
Conus sp. or spp. 3S
?Mangelia quadrdim-ata (C. B. Adam.s, 1850) 2S
Bulla striata Bruguiere, 1792 2S
Odostomia sp. or Pyramldella sp. IS
Cheilia equestris (Linnaeus, 1758) IS
Total 1,1.3()
S. K. Donovan and D. T, J. Littlewood, 1993
Page 37
Table 1. ('diitiniird
Bivalves
Area ?zebra Swainson. 18S3
Area imhricuta Bruguiere, 1789
Rarlxititi Candida (Helbling, 1779)
Barlmtia ?cancellaria (Lamarck, 1819)
Barhatia teiwra (C B. Adams, 1845)
Anadara sp
.■\rks iiuiet. + +
Brachidonti's cxustus (Linnaeus, 1758)
Smoolli '•'Botida fuaca (Gmelin, 1791)
iJthophaga ''nigra (d'Orbigny, 1842)
hognomvn '•'alalua (CJmelin, 1791)
hognomon radiatus (Anton, 1839)
Isognonwn sp, indet
Lima scabra (Born, 1778)
Ostreola equestris Say, 1834
Codakia orbicularis (Linnaeus, 1758)
Codakia ?orbiculata (Montagu, 1808)
''Codakia sp,
Chama maccrophylla Gmelin. 1791
Chama '''maccrophylla Gmelin, 1791
Pseudochatua radians (Lamarck, 1819)
Americardia media (Linnaeus, 1758)
Americardia guppyi (Thiele, 1910)
Chione cancellata (Linnaeus, 1767)
Pcrighjpta listeri (Gray, 1838)
Tellina listeri Boding, 1798
Tellina sp- cf, Tellina ''tnera Sa\, 1834
Tellins indet.
Tellinacean sp, indet. A
Tellinacean sp, indet B
?Tellinacean sp, indet.**
Areopagia Jausta (Pulteney, 1799)
?Macoma pseudomera Dall and Simpson, 1901
?Macoma sp.
PLucinid or ?pectinacean indet.
?Mactracean indet,
?Periploma papyralium Say, 1822
Bivalve sp indet.
Scaphopods
Dentalium sp cf. Dentalium (Anatalis)
antillanim d'Orbigny, 1842
Dentalium (Graptaeme) semistriolatum CJuilding, 1834
9V +
1V +
1V +
1V +
1V +
Totals
4LV
77LV
25LV
ILV
36LV
ILV
32LV
4LV
ILV
lOLV
4LV
ILV
2LV
ILV
9LV
4LV
ILV
ILV
ILV
2LV
2LV
ILV
3LV
28LV
2LV
61RV
37 RV
4RV
24RV
29RV
2RV
5RV
2RV
IRV
13RV
6RV
IRV
IRV
IRV
IRV
5RV
IRV
IRV
35RV
IRV
IRV
IRV
IRV
IRV
502V
19S
IS
IS
2S
IS
IS
3S
6S
Total
33S
IS
2S
3S
Because of the well-lithified nature of much of this
limestone, it was necessary to use a hammer and chisel
to remove all specimens seen in the exposed faces (figures
5-7). Specimens were not apparent on all surfaces due
to the vagaries of weathering, although breaking open
many of these 'unfossiliferous' exposures showed that
mollusks were nevertheless common. As well as collecting
individual specimens in the field, large fossiliferous slabs
containing dense accumulations of mollusks were taken
back to the laboratory for mechanical breakdown. A total
of 1365 benthic mollusks have been identified from this
locality (table 2), including abundant gastropods and bi-
valves, rare scaphopods, but no chitons.
RESULTS AND DISCUSSION
Taxonomy: Taxa, and number of specimens of each
species, from localities 1 and 2 are tabulated in tables 1
and 2, respectively. It is readily apparent that both fau-
nas, although approximately contemporaneous and sep-
arated by a distance of only about 3 km, show consid-
erable taxonomic divergence (figures 8, 9). Localitv 1
has the more diverse fauna, which is dominated numer-
ically by the micromollusk Caecum pulchellutn Stimp-
son, 1851, arks and epifaunal gastropods (although C.
piilchelltim made only a minute contribution to biomass),
whereas locality 2 is typified by the occurrence of abun-
Page 38
THE NAUTILUS, Vol. 107, No. 1
Table 2. MoUusk fauna collected from locality 2, east of Discovery Bay Marine Laboratory. Key: '
C". B. .-Vdams, 1848; otherwise as for Table I
= encrusting Cerithium algicola
Gastropods
Diodora ?cayenensis (Lamarck, 1822)
Turbo canaliciilahi.s Hermann, 1781
Tricolia sp or spp.
?Smaragdia ciridis ciridenmris Maury, 1917
Rissoina aberans (C. B. .\dams, 1850)
'^Rissoina sp.
Planaxis lineatus (da Costa, 1778)
Modulus carchcdonius (Lamarck, 1822)
Modulus modtdus (Linnaeus, 1758)
Cerithium algicola C. B .\dams. 1848
Cerithium eburneum Bruguiere. 1792
Cerithium litteratum Born, 1778
Cerithium spp. indet. (fragments)
Janthina sp.
?\iso portoricensis Dall and Simpson, 1901
Stromhus gigas Linnaeus, 1758
Polinices lactcus (.Guilding, 1834)
Satica canrena (Linnaeus, 1758)
Naticid sp. indet.
Cymatium "^caribbaeum Clench and Turner. 1957
Ctjmatium muricinum (Boding, 1798)
Cymatium sp. indet. .\
Cymatium sp. indet. B
Cymatium sp indet. C
?Thais ddtoidea (Lamarck, 1822)
Columbella mercatoria (Linnaeus. 1758)
Anachis obesa (C. B. .\dams. 1845)
Anachis ?pretri (Duclos, 1846)
Anachis sp. indet.
S'assarius alhtis (Say, 1826)
Fasciolaria tulipa (Linnaeus, 1758)
Oliiella dealbata (Ree\e, 1850)
Prttnuvi guttatum (Dill\v\n, 1817)
?\'olvarina gracilis (C B. .\dams, 1851)
CoTius '^centurio Born, 1778
Conus verrucosus Hwass, 1792
?Crassispira sp. indet.
Btdia striata Bruguiere, 1792
Retusa candci (d Orbigiu, 1841)
Bivalves
??Suculid sp. indet.
Area imbricata Bruguiere, 1789
Barbatia cancellaria (Lamarck. 1819)
.\rk indet. sp.
Brachidontes citrinus (Boding. 1798)
?lsognomon sp.
Pinna camea Gmelin. 1791
Pecten '^chazaliei Dautzenberg. 1900
?Diplodonta punctata ^Sa>. 1822)
Lucina sp. indet .A
Lucinid sp. indet. .\
Lucinids spp. indet.
?Anodontia alba Link. 1807
Codakia costata (d"Orbigny. 1842)
Codakia orbictdaris (Linnaeus, 175S)
Codakia orbictdata (Montague. 180S)
Dicaricella quadhsulcata (d"Orbign\ , 1842)
Chama macerophulla Gmelin, 1791
Chama sp
Total
ILV
3RV
ILV
IRV
2L\"
IRV
ILV
IRV
ILV
1V +
ILV^
IRV
ILV
2LV
IRV
2L\-
3RV
3L\
3RV
2LV
4LV
3RV
2RV
2LV
2RV
121.V
34RV
IRV
2S
IS
6S
IS
IS
IS
IS
7S
IS
lOlS
3S
33S
58S
IS
35
17S
78S
IS
IJ
IS
IS
IS
IS
IJ
IS
5S
IS
IS
2S
IS
2S
37S
13S
IS
95
IS
IS
6US
IS
LOOS
IS
:i \
S. K. Donovan and D. T. J. Littlewood, 1993
Table 2. ( liiiiliinic
Page 39
fTraihyairdiinu sp
Americardia gupptji (Tliiilc. 1910)
Aniericarilia ntcdici (Linnaeus, 1758)
Lacvictirditini liHiif^iiluin (Linnaeus, 1758)
'^Lacvicardiiun Itui if^dluiu (Liiuiaeus, 1758)
Laevicardium '•'syhdnluuiu Dall, 1886
Chiotie ranci'llata (Liiuiaeus. 17(i7)
Chionc firaniilata ((iinelin. 1791)
Chionc pap}uu (Linnaeus, I7()7)
Transcnnclla '•'cimradimi Dall, 1883
Transennclla ?cubaniana (d'OrliiKny, 18-42)
Pitar allrida (Cmelin, 1791)
Tcllina altcrnala Say, 1822
Tcllina listrri Hoding, 1798
Tcllina similts Sowerby, 1806
Tcllina sp. iiulet
Apohjmelis inlastriata (Say, 1826)
Tagcliis divisus (Spengler, 1794)
Mactra fragilis Gnwhn, 1791
Scaphopods
Cadulus sp
Denialium ''anlillarum dOrliigiu, 1846
Dentalium sp, (smooth-shelled)
1\
Totals
2LV
2RV
ILV
2RV
(>LV
7RV
5S
3LV
IRV'
1V +
lOLV
URV
IRV
IS
2LV
2RV
2S
6LV
2RV
2LV
2RV
13LV
12RV
6S
2LV
2RV
2V-I-
3LV
2RV
2S
18LV
lORV
4V-I-
7LV
3RV
ILV
6S
30LV
29RV
2RV
2S
328V
25S
Total
IS
2S
IS
4S
dant Bulla striata Briiguiere, 1792 and Cerithium spp
with other gastropods and rarer bi\aives.
Paleoecology and preservation: Taxonomic differ-
ences are interpreted as being related to paieoenviron-
mental control. Locality 1 included abundant hard sub-
strates for the attachment of epifaunal bivalves, provided
by the coral framework, as well as a sandy substrate
suitable for a variety of infaunal and epifaunal ta.xa.
Chitons, which generally prefer a hard substrate, are also
common at this locality . In contrast, locality 2 lacks any
evidence of a three-dimensional framework structure;
indeed, corals were small and isolated in this paleoen-
vironment. Consequently , epifaunal species are rare and
the fauna is dominated by infaunal taxa, particularly B
striata, but also including \arious burrowing bi\alves
(habitat preferences of many of the groups tabulated
herein are summarized in Cerridwen & Jones, 1991: table
Table 3. .\ comparison of the taxonomic diversity of the extant
Jamaican moliuscan fauna with nominal species identified here-
in from the Falmouth Formation. Key: -I- = after Humfre\
(1975); * = excluding rare species; 1, 2 = localities.
K\tant-I-
1
,->
1 + 2
Scaphopods
11
•1
3
4
:56
Gastropods
324*
55
37
80
25
558
55
37
80
14
Bivalves
163*
33
34
56
34
246
33
34
56
23
Totals
514*
141)
-">-
831
140
17
3). The only species which forms greater than 2% of both
faunas is Cerithium algicola C B .Adams, 1848 (figures
8, 9; tables 1, 2).
The difference in diversity between the localities may
be real, although it is almost certainly at least part ar-
tifact. This is because of the differing degrees of lithifi-
cation (and, hence, availability of specimens) seen he-
tween the two collecting sites. All specimens from bulk
samples from locality 1 were theoretically collected by
sieving and picking. In contrast, limestones from locality
2 were not amenable to such processing, being well-
lithified, and liberation of specimens was essentially a
destructive process. In addition to those specimens bro-
ken from the rock, there were some mollusks, particularly
those with a fragile shell, that were too easily lost due
to breakage while others were being remo\ed. Further,
many specimens were probably ne\er seen because they
were enclosed within the remaining small chips of rock
that were not broken down further. Micromollusks are
a common component of the fauna from locality 1. main-
Iv recognized during microscopic examination of the
sand-sized sieve fraction. Such a technique is not gen-
erally possible with the sediment from locality 2 and,
even if detected, small shells are difficult to extract and
clean from the hard matrix. The total absence of chiton
vaKes or Caecum shells from locality 2 may be real, but
the limited diversity of micromollusks from this site sug-
gests that preservational and collecting biases were im-
portant factors (in particular, extant Caecum is a com-
mon component of seagrass communities which
presumably occurred in the paleolagoon). Thus, micro-
mollusks were part of the easily collectable fauna only
Page 40
THE NAUTILUS, Vol. 107, No. 1
o
o
a-
o
lOOi
75
50
25
0
a
c3
H.
O
^Myf0(iiyf^'^f!f^^mKy7TmmmmmjTwTw„ym^j
Species
Figure 8. Histogram of relative abundance of mollusk species
of locality 1 (n = 1467; bivalves recalculated as whole shell
equivalents). 89 species are each present as less than 1% of the
total mollusk fauna. • = broken fragments of arks which are
too poorly preserved for even generic identiBcation, but which
are probably dominantly A. inihricala.
at locality 1, although it was still possible to extract some
small shells from the limestones at locality 2 (particularly
robust shells such as Cadtiliis sp. and B. striata juveniles).
Macromollusks from both sections are often exquisitely
preserved. Whilst bivalves are normally (and chitons in-
variably) disarticulated, valves often have a highly glossy
appearance, such as Laevicardium laevigattim (Linnaeus,
1758) at locality 2 and Polinices lacteus (Guilding, 1834)
at both localities, equivalent to that of fresh shells. Fur-
ther, shell coloration and/or color banding is preserved
in, for example, at least some specimens of Leucozonia
nassa (Gmelin, 1791) from locality 1 and B. striata from
locality 2. This indicates that the diagenetic effects lead-
ing to the lithification of the limestones (particularly at
locality 2) have so far had little influence on the organic
molecules that are largely responsible for coloration.
However, many shells are poorly preserved, often being
corroded and/or abraded. Many show the influence of
pre- and post-mortem boring. Pre-mortem borings by
predatory gastropods (?P. lacteus) have been noted in,
for example, Brachidontcs cxustus (Linnaeus, 1758) and
Table 4. ('omparison ol llic extant shallow-vsater molluscan
fauna of Yucatan (PJkdale, 1974) with that of the Pleistocene
Falmouth Knrmation
calitit's
e\rln<liiig chitims.
= lo-
Rocky intertidal zone (14 sp.*)
Lagoonal and mangrove-associated
environments (27 sp.)
Broad, shallow backreef (23 sp.)
High-energy environment (8 sp.)
Open sea assemblage (15 sp.)
(x)smo|)olitan taxa (9 sp.)
Species
in common
vith
1 + 2
1 2
1 + 2
Cc)
3 1
3
21
9 9
11
41
7 10
1 1
10
1
43
13
4 3
4
27
2 4
4
44
100
o
75
e 50
o
o
25:
0
3
I
E.
c ^ 1= _3
■a
^
^
V
-b
cc
o
<3
t-.
r^
VZ^^r77T77xy77777\^
Species
Figure 9. Histogram of relati\ e aliuntlancies of mollusk species
at locality 2 (n = 1226; bivalves recalculated as whole shell
equivalents) 71 species are each present as less than 2''( of the
total mollusk launa. • = broken fragments of dominantly, or
entirely, C. algicola + C. literattim which are too poorly pre-
served for identification below the generic level.
D. sp. cf. D. (A.) antillariim (see Donovan & Littlewood,
1989) from locality 1. Presumably post-mortem borings
produced by Pclionid sponges are common in arks from
the same site, while B. striata and P. lacteus from locality
2 were occasionally bored by polychaetes.
Comparisons with extant faunas: 17''( of the extant,
shallow-water, benthic mollusks known from the island
at the present day have been identified from localities 1
and 2 (table 3). If the rarer taxa listed by Humfrey (1975)
are ignored, this figure rises to 27% (table 3),
Parsons and Brett (1991) have highlighted the fluctu-
ating taxonomic compositions of manv tropical, shallow
shelf mollusk faunas. The assemblages described from
the Falmouth Formation undoubtedly include mollusks
from a number of ephemeral communities, having been
collected from about 2-4 m of section. While the broad
environmental signals of the two facies are obviously
different, it is uncertain how the interplay of changing
community structure at both sites lead to the preservation
of the fossil faunas. Comparisons that we have made
between our Falmouth Formation mollusk assemblages
and published descriptions of Caribbean shallow-water
mollusk communities have been inconclusive, showing
no obvious close parallels. As an example, table 4 com-
pares the Falmouth Formation molluscs with the modern
shallow-water fauna of northeast Yucatan (Ekdale, 1974).
There is no assemblage which shows even a 50% simi-
larity to localities 1-1-2 combined. Faunal lists of the
modern mollusc faunas from particular habitats in Ja-
maica lia\e not been published (Dr. J.D. Woodley, per-
sonal conmnication).
CONCLUSIONS
140 nominal species have been identified from the late
Pleistocene Falmouth Formation of Jamaica. If rarities
S. K. Donovan and D. T. ]. Littlewood, 1993
Page 41
in the extant launa are ignored, this represents over 25%
of the common living species from the island. However,
the Falmouth Formation is an averaged sample from a
number of ephemeral communities and comparisons w ith
extant Caribbean faunas from similar em iromnents are
inconclusive. This preliminary sur\ey highlights the di-
versity of the mollusk fauna across reef environments
and suggests that fossil reefs may be considered as im-
portant sampling points during the reconstruction of an-
cient regional biotas.
ACKNOWLEDGEMENTS
We thank Miss Carla M. Gordon for her considerable
help in collecting samples and providing picked, moilusk-
rich residues surplus to her own research needs. Dr. Rich-
ard Gustafson kindly helped with the identification of
some of the more obscure bivalves from locality 1. We
thank Ms. Dana C. Larson for permission to utilize her
thesis research in Fig. 3 This paper is Discovery Ba\
Marine Laboratory contribution number 530, New Jer-
sey Agricultural Experiment Station publication number
D-32501-1-92 and Institute of Marine and Coastal Sci-
ences contribution number 92-04.
LITERATURE CITED
Abbott, R.T. 1954. American seashelis. Van Nostrand, Prince-
ton. 542 p.
Abbott, R.T. 1958. The marine moliusks of (irand C'ayman
IsiaiKJ, British West Indies. Monograpii ot the Academy
ol Natural Sciences of Philadelphia 11; ISH p.
Abbott, R.T. 1974. American seashelis (2nd ed.). Van Nos-
trand Reinhold, New York. 663 p.
Bengston, P. 1988. Open nomenclature. Palaeuntulogv 31:
223-227.
Cerridwen, S.A. and B. Jones. 1991. Distribution of bivalves
and gastropods in the Pleistocene Ironshore Formation,
Grand Cayman, British West Indies. Caribbean Journal of
Science 27: 97-116.
de Jong, K.M and H.E. Coomans. 1988. Marine gastroputis
from Curasao, Aruba and Bonaire Studies on the Fauna
of Curaq-ao and other Caribbean Islands 69: 261 p
Donovan, S.K. 1990. Fossil Scaphopoda (Mollusca) from the
Cenozoic of Jamaica. Journal of the Geological Society of
Jamaica 27: 1-9.
Donovan, S.K. and CM. Gordon. 1989. Report of a field
meeting to selected localities in St. Andrew and St. Ann,
25 February 1989. Journal of the Geological Society of
Jamaica 26: 51-54.
Donovan, S.K., CM. Gordon, C.J. Veltkamp antl AD Scott
In press. Crinoids, asteroids and ophiuroids in the Ja-
maican fossil record. Geological Society of .Vmerica Mem-
oir 182.
Donovan, S.K. and D.T.J. Littlewood. 1989. A Late Pleisto-
cene scaphopod from Jamaica. Journal of Conchology 33:
185.
Fisher, W.S. (ed.). 1988. Disease processes in marine hi\alve
molluscs. American Fisheries Society Special Publication
18: viii -I- 315 p.
Ekdale, A. A. 1974. Marine molluscs from shallow-water en-
vironments (0 t(i 60 meters) off the northeast Yucatan coast,
Mexico. Bulletin of Marine Science 24: 638-668.
Gordon. CM and S K Donovan. 1992. Disarticulated echi-
iioid ossicles in paleoecology and taphonomy: the last in-
terglacial Falmouth Formation of Jamaica. Palaios7: 157-
166.
Hill, R.T. 1899. The geology and plusical geograph\ of Ja-
maica: a study of a type of .Antillean tievelopment Bulletin
of the Museum ot Coniparati\e Zoolog\ at Harvard 34:
256 p.
Humfrev. M 1975 Sea shells of the West Indies. Collins,
London. .351 p.
Johnson, R.G I960. Enviromnental interpretation of Pleis-
tocene marine species. Journal of Geology 68: 575-576.
Jung, P. 1972. Moliusks from the White Limestone Group of
Jamaica. In: Petzall, C (ed.) Memorias de la VI Confer-
encia Geologica del Caribe, Margarita, Venezuela, 6-14
del Julio de 1971, Impreso por Cromotip, Caracas, p. 465-
468.
Land, L.S. 1991. Some aspects of the late Cenozoic evolution
of north Jamaica as re\ealed b\ strontium isotope stratig-
raph> Journal of the Geological Society of Jamaica 28:
45-48.
Land, L.S. and S. Epstein. 1970. Late Pleistocene diagenesis
and dolomitization, north Jamaica. Sedimentologv 14: 187-
200.
Larson, DC 1983. Depositional facies and diagenetic fabrics
in the Late Pleistocene Falmouth Formation of Jamaica.
M.S. thesis. University of Oklahoma, Norman, Oklahoma.
Liddell, W.D.,S.L. Ohihorstand AG. Coates. 1984. Modern
and ancient carbonate environments of Jamaica. Rosenstiel
School of Marine and Atmospheric Science, University of
Miami, Sedimenta 10: vii -I- 98 p.
Morris, PA. (W.J. Clench, ed.). 1987. A field guide to shells
of the Atlantic and Ckilf Coasts and the West Indies.
Houghton Mifflin, Boston, x.wiii + 330 p.
Morris, S.F. In press. Biostratigraphy of the arthropods of
Jamaica. Geological Society of America Memoir 180.
Parsons, KM. and C.E. Brett. 1991. Taphonomic processes
and biases in modern marine environments: an actualis-
tic perspective on fossil assemblage preservation. In Don-
ovan, S K. (ed ). Fossilization: the processes of taphonomy
Belhaven Press, London p. 22-65.
Petuch, E.J. 1988. Neogene histor\ of tropical .American mol-
iusks. The Coastal Education and Research F"oundation,
C^harlottesville, Virginia. 217 p
Robinson, E. 1958. The younger rocks of St. James and Tre-
lawny. Geonotes 1: 15-17.
Robinson, E. 1960. Observations on the elevated and modern
reef formations of the St. Aim coast. Geonotes 3: 18-22.
Robinson, E 1963. Quaternary deposits. In. Zans, V A., L.J.
Chubb, H R. Versey, J B Williams, E Robinson and D L.
Cooke. Synopsis of the geologv of Jamaica. Bulletin of the
Geological Survey Department of Jamaica 4: 50-54.
Taylor, J.D. 1978. Faunal response to the instability of reef
habitats: Pleistocene molluscan assemblages of Aldabra
Atoll. Palaeontology 21: 1-30.
Trechmann, C T. 1930. The Manchioneal Beds of Jamaica
Geological Magazine 61: 199-218.
N'alentine, J.W 1989 How good was the fossil record? Clues
from the Californian Pleistocene. Paleobiology 15: 83-94.
\aught, KG. (R.T. Abbott and K J Boss, eds) 1989. A clas-
sification of the li\ing Mollusca. American Malacologists
Inc., Melbourne, Florida, xii + 195 p.
Page 42 THE NAUTILUS, Vol. 107, No. 1
VVarmke, G.L. and R.T .Abbott. 1961. Caribbean seashells. Woodring, W.P 1928 Miocene mollusks from Bowden, ja-
Livingston, Narberth, Pennsylvania, x + 346 p. maica, pt. II, gastropods and discussion of results. Carnegie
Woodring, W.P. 1925. Miocene mollusks from Bowden, Ja- Institution of Washington Publication 385: 564 p.
maica. Carnegie Institution of Washington Publication 366:
vii + 222p.
INSTRUCTIONS TO AUTHORS
THE NAUTILUS publishes papers on all aspects of the
biology and systematics of mollusks. Manuscripts de-
scribing original, unpublished research as well as review
articles will be considered. Brief articles, not exceeding
1000 words, will be published as notes and do not re-
quire an abstract. Notices of meetings and other items
of interest to rnalacologists will appear in a news and
notices section.
Manuscripts: Each original manu.script and accompa-
nying illustrations should be submitted in triplicate. Text
must be typed on one side of 8'/2 x 11 inch white paper,
double spaced throughout (including literature cited, ta-
bles and figure captions), with at least 1 inch of margin
on all sides. All pages must be numbered consecutively.
If printed on a word processor, the right margin should
be ragged rather than justified. Authors should follow
the recommendations of the Council oj Biology Editors
Style Manual, which is available from the (Council of
Biology Editors, Inc., 9650 Rockville Pike, Bethesda, MD
20814, U.S.A. The first mention of a scientific name in
the text should be accompanied by the taxonomic au-
thority, including year. Latin names and words to be
printed in italics must be underlined; leave other indi-
cations to the editor. Metric and Celsius units are to be
used.
The sequence of sections should be: title page, abstract
page, introduction, materials and methods, results, dis-
cussion, acknowledgements, literature cited, tables, fig-
ure captions, figures. The title page should include the
title, author's name(s) and address(es). The abstract page
should contain the title and abstract, which should sum-
marize in 250 words or less the scope, main results and
conclusions of the paper. The abstract may be followed
by a maximum of 8 key words. All references cited in
the text must appear in the literature cited section and
vice versa. In the literature cited section, all authors
must be fully identified and listed alphabetically. Follow
a recent issue of THE NAUTILUS for bibliographic style,
noting that journal titles must be unabbreviated. Infor-
mation on plates and figures should be cited only if not
included in the pagination. Tables must be numbered
and each placed on a separate sheet. A brief legend must
accompany each table. Captions for each group of illus-
trations should be typed on a separate sheet and include
a key to all lettered labeling appearing in that group of
illustrations.
All line drawings must be in black, high quality ink,
clearly detailed and completely labeled. Photographs
must be on glo.ssy, high contrast paper. All figures are
to be consecutively numbered (figs. 1, 2, 3, ... , NOT
figs. la. lb, Ic, . . . NOR plate 1, fig. 1 . . .). Illustrations
must be arranged in proportions that will conform with
the width of a page (6% inches or 171 mm) or a column
(3'/4 inches or 82 mm). The maximum size of a printed
figure is 6% by 9 inches or 171 by 228 mm. All illus-
trations must be fully cropped, mounted on a firm, white
backing, numbered, labeled and camera ready. The au-
thor's name, paper title and figure number(s) should ap-
pear on the back. Original illustrations must be between
one and two times the desired final size. It is the author's
responsibility that the line weight and lettering are ap-
propriate for the desired reduction. Original illustrations
will be returned to the author if requested. Color illus-
trations can be included at extra cost to the author.
Voucher Material: Deposition of type material in a
recognized public museum is a requirement for publi-
cation of papers in which new species are described.
Deposition of representative voucher specimens in such
institutions is strongly encouraged for all other types of
research papers.
Processing of Manuscripts: Upon receipt, every manu-
script is acknowledged and sent for critical review by at
least two referees. These reviews serve as the basis for
acceptance or rejection. Accepted manuscripts are re-
turned to the author for consideration of the reviewers'
comments. A finalized version of the manuscript is re-
turned to the editor and sent to press. Two sets of proofs
are sent to the author for correction. Changes other than
typesetting errors will be charged to the author at cost.
One set of corrected proofs should be sent to the editor
as soon as possible. Authors with institutional, grant or
other research support w ill be billed for page charges at
the rate of $60.00 per printed page.
An order form for reprints will accompany the proofs.
Reprints may l)e ordered through the editor.
Manuscripts, corrected proofs and correspondence re-
garding editorial matters should be .sent to: Dr. M.G.
Harasewych, Editor, Division of Mollusks, NHB stop 1 18,
National Museum of Natural History, Smithsonian In-
stitution, 'V\'ashington, DC 20560, USA.
THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.
FHE NAUTILUS
^^.^^p^T
Volume 107, Number 2
July 28, 1993
ISSN 0028-1344
A quarterly devoted
to malacology.
LIBRARY
AUG 61983
Vvoodr} Hole, !V;. 3.
EDITOR-IN-CHIEF
Dr. M. G. Harasewych
Division of Mollusks
National Museum of
Natural History
Smithsonian Institution
Washington, DC 20560
ASSOCIATE EDITOR
Dr. R. Tucker Abbott
American Malacologists, Inc.
P.O. Box 2255
Melbourne, FL 32902
CONSULTING EDITORS
Dr. Riidiger Bieler
Department of Invertebrates
Field Museum of
Natural History
Chicago, IL 60605
Dr. Robert T. Dillon, Jr.
Department of Biology
College of Charleston
Charleston, SC 29424
Dr. William K. Emerson
Department of Living Invertebrates
The American Museum of Natural
History
New York, NY 10024
Dr. Robert Hershler
Division of Mollusks
National Museum of
Natural History
Smithsonian Institution
Washington, DC 20560
Dr. Richard S. Houbrick
Division of Mollusks
National Museum of
Natural History
Smith.sonian Institution
Washington, DC 20560
Mr. Richard I. Johnson
Department of Mollusks
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138
Dr. Aurele La Rocque
Department of Geology
The Ohio State University
Columbus, OH 43210
Dr. James H. McLean
Department of Malacology
Los Angeles County Museum of
Natural History
900 E.xposition Boulevard
Los Angeles, CA 90007
Dr. Arthur S. Merrill
% Department of Mollusks
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138
Ms. Paula M. Mikkelsen
Harbor Branch Oceanographic
Institution, Inc.
Ft. Pierce, FL 33450
Dr. Donald R. Moore
Division of Marine Geology
and Geophysics
Rosenstiel School of Marine and
Atmospheric Science
University of Miami
4600 Rickenbacker Causeway
Miami, FL 33149
Dr. Gustav Paulay
Marine Laboratory
University of Guam
Mangilao, Guam 96923
Mr. Richard E. Petit
P.O. Bo.x 30
North Myrtle Beach, SC 29582
Dr. Edward J. Petuch
Department of Geology
Florida Atlantic University
Boca Raton, FL 33431
Dr. David H. Stansbery
Museum of Zoology
The Ohio State University
Columbus, OH 43210
Dr. Ruth D. Turner
Department of Mollusks
Mu.seum of Comparative Zoology
Harvard University
Cambridge, MA 02138
Dr. Geerat J. Vermeij
Department of Geology
University of California at Davis
Davis, CA 95616
SUBSCRIPTION INFORMATION
The subscription rate per volume
is US $25.00 for individuals, US
$40.00 for institutions. Postage
outside the United States is an
additional US $5.00 for surface
and US $12.00 for air mail. All
orders should be accompanied by
pavment and sent to; THE
NAUTILUS, P.O. Box 7279, Silver
Spring, MD 20907-7279, USA.
Change of address: Please inform
the publisher of your new address
at least 6 weeks in advance. All
communications should include
both old and new addresses (with
zip codes) and state the effective
date.
THE NAUTILUS (ISSN 0028-
1344) is published quarterly by
Trophon Corporation, 8911 Alton
Parkway, Silver Spring, MD 20910.
Second Class postage paid at Silver
Spring, MD and additional mailing
offices.
POSTMASTER; Send address
changes to: THE NAUTILUS
P.O. Box 7279
Silver Spring, MD 20907-7279
THE €9 NAUTILUS
CONTENTS
Volume 107, Number 2
Marine Biological LaboraLw/"'^ ^*' ^^^^
LIBRARY ^% 0028-1344
Silvard P. Kool
The Systematic Position of the Genius W&d&H HolS MasS
(Prosobranchia: Miiricidae; r>r.or.o'kr;n>)p'| ^
43
Jose H. Leal
Donald R. Moore
Thala csperanza. a new Costellariidae (Mollusca:
Gastropoda) from northern Puerto Rico
58
Anionieto S. Tan
Ya-ping Hu
IVIichael Castagna
Richard A. Luiz
Michael J. kennish
Alan S. Pooley
Shell and Pallet Mor|jhology of Early Developmental
Stages of Bankia gouldi (Bartsch, 1908) (Bivalvia:
Teredinidae) ...
63
S. J. kleinschusler
S. L. Swink
A Simple Method tor the In Vitro Oilture of Perkinsus
warinus
76
Harold G. Pierce
On the Identification of Fossil Terrestrial Gastropod
Eggshells
79
THE NAUTILUS 107(2):43-57, 1993
Page 43
The Systematic Position of the Genus NiiceUa
(Prosobranchia: Muricidae: Ocenebrinae)
Silvard P. Kool
Mollusk Department
Museum of Comparative Zoology
Harvard Universitv
Camhriiige. Massachusetts 02138 USA
ABSTRACT
The muricid genus S'ucclla Roding. 1798 has conimonK been
placed in Thaidinae Joiisseaunie, 1888 (Prosobranchia: Muri-
cidae). The Thaidinae (scnsii Kool, 1989) is monoph) letic with
and thus s\ nonymous with Rapaninae Gra\ , 1853 (Kool, 1993,
in press). Comparative anatomical investigations of the type
species of Nucclla Roding, 1798 (Buccinum filosum Gmelin,
1791 [=Nttcella lapillus (Linnaeus, 1758)]) and of Thais Rod-
ing, 1798 (Murex funis Gmelin, 1791 [ = Thais nodosa (Lin-
naeus, 1758)]) as well as other rapanines have revealed that
inclusion of Nucclla in Rapaninae would result in polyphyletic
groups (Kool, 1989; 1993, in press). Studies of the anatomy,
radula, protoconch, shell ultrastructure, and operculum of the
type species of A't/ce/Za, Occnebra Gray, 1847 (Murcx crinaceus
Linnaeus, 1758 [=Ocenebra erinacea]) (Ocenebrinae Coss-
mann, 1903), and Trophon Montfort, 1810 {Murex magellan-
tcMS Gmelin, 1791 [=Trophon geversianus (PaWni, 1774)]) (Tro-
phoninae Cossmann, 1903), indicate that Nucella has close
affinities with Ocenebrinae and Trophoninae Based on cladistic
analyses, it is here proposed that Nucclla be placed in Oce-
nebrinae. Results further reveal that the distinctions between
Ocenebrinae and Trophoninae are less clear than previously
accepted
Key words: Nucclla, Ocenebrinae, phylogeny; systematics;
comparative anatom\
INTRODUCTION
The Thaidinae of authors, also referred to as Thaididae,
Purpurinae/dae Swainsoii, 1840, Driipinae Wenz, 1941,
etc., has been shown to be a conglomerate of disparate
taxa (Kool, 1989; 1993, in press). The ta.xonomic coher-
ence of the Thaidinae and the boundaries of its genera
were based primarily on external shell characters, which
are often convergent, obscuring phylogenetic relation-
ships.
Rigorous cladistic analyses based primarily on char-
acters derived from anatomy, radula, operculum, and
shell ultrastructure, have shown that para- and poly-
phyly were wide-spread in the Thaidinae/dae of authors
(Kool, 1989). Subsequent phylogenetic studies have re-
vealed that the genus Rapana Schumacher, 1817, and
the Thaidinae (in partem) constitute a monophyletic
group, making Thaidinae a junior subjective synonym
of Rapaninae (Kool, 1993, in press). The name Rapaninae
will herein be used for the clade that includes Rapana
and Thais
Several species of the genus Nitcella have been used
extensively in ecological studies (Colton, 1922; Crothers,
1983, 1985; Emlen, 19(56; Etter, 1987; Kincaid, 1957;
Moore, 1936, 1938; Palmer, 1983, 1985; Spight, 1972,
1976). In most of these studies Mucella was regarded as
a subgenus or synonym of Thais. Anatomical studies
(Kool, 1986, 1989) of the type species of Nucella [Buc-
cinum filosum = Nucella lapillus (see Kool & Boss, 1992)]
and Thais [Murex fucus = Thais noc/o.sa] revealed major
differences between these genera. Kool (1988) therefore
excluded Nucella from the Thaidinae and tentatively
placed Nucella in the Ocenebrinae (Kool, 1989) on the
basis of radular (Sabelli & Tommasini, 1987; Bandel,
1977) and protoconch (Bandel, 1975) morphology as well
as anatomical descriptions (Graham, 1941) of Ocenebra
erinacea.
Although the anatomv of Nucella lapillus is well known
(Fretter & Graham, 1962; Kool, 1986, 1989; Oehlmann
et al., 1988), relatively little is known about the soft parts
of Ocenebra erinacea. Aspects of the anatomy of Tro-
phon geversianus were described by Harasewych (1984),
who suggested that similarities (e.g. radular morphology)
between members of Trophon and Nucella may be due
either to convergence resulting from similar environ-
mental conditions or to phylogenetic affinity.
The object of this study is to discern the phylogenetic
affinities among Nucella, Trophon and Ocenebra.
MATERIALS AND METHODS
The following specimens were used for anatomical stud-
ies:
Nucella lapillus- Kittery, Maine, U.S.A. (USNM 857053)
(7 9, 5S).
Trophon geversiatius; Daniel Este, Isia Grande, Tierra
del Fuego, Chile (LACM 86-270.2); Puerto Basil Hall,
Page 44
THE NAUTILUS, Vol. 107, No. 2
Table 1. List of characters and character states
for Murican
•thus.
Thaifi. Troplifm. Nuce
'lla, and Ocenelna.
Character
Mur
Tha
Tro
Nuc
Oce
1. Protoconch whorls
0
0
1
1
2. Calcitic layer
0
1
1
1
3. Number of aragonitic layers
0
1
2
2
0
-4 Position of opercular nucleus
0
3
1
2
5. Opercular shape
0
1
0
1
6. Pigmentation pattern on head-foot re-
gion
0
0
1
1
7. Duct(s) for accessory boring organ and
ventral pedal gland
0
0
0
1
8. Bursa copulatrix
0
2
1
1
0
9. Seminal receptacles at dorsal periphery
of albumen gland
0
1
0
0
0
10 Fenial shape
0
2
1
0
0
11. Penial vas deferens
0
1
0
0
2
12. Prostate
0
1
0
0
0
13. Accessory salivary gland(s)
0
1
1
2
2
14. Straw-like membrane around glancl of
Leiblein
0
0
1
1
1
1.5, Posterior duct of gland of Leiblein
0
0
1
1
1
16. Ontral cusp of rachidian
0
0
0
1
1
17 Margin of rachidian basal plate
0
0
1
1
1
Isla de los Estados, Tierra del Fuego, Argentina (LACM
71-289); Piinta Catalina, Isla Grande, Tierra del Fuego,
Chile (LACM 80-87.2) (4 9, 3 <5).
Ocenebra erinacea; Roscoff, France (MCZ 298425) (2 2.
IS).
Morphological data were compiled from soft tissues,
radulae, shell ultrastructures, protoconchs, and opercula.
Living specimens of Nucella and preserved specimens
of Nucella. Trophon and Ocenebra were dissected.
Radulae (2-4 per species) were cleaned using a potas-
sium hydro.xide solution, rinsed in distilled water, air-
dried, sputter-coated with carbon and gold, and exam-
ined with a Hitachi S-570 scanning electron microscope.
Photomicrographs were taken of the unused, matured
central portion of each radular ribbon.
Shell fragments from at least two individuals of each
species were obtained by crushing the shell. Portions
from the central region of the body whorl about one-
half to three-quarters of a whorl away from the edge of
the apertural lip were mounted, sputter-coated with car-
bon and gold, and their fracture surfaces observed with
a Hitachi S-570 scanning electron microscope. An ap-
parentK amorphous outer layer was interpreted as con-
sisting of calcite, while layers with organization in crystal
lamellar structure were considered aragonitic (data from
x-ray diffraction methods confirm these identifications;
Kool and Harasewych, in preparation).
Cladistic Analysis: Seventeen characters, divided into 41
character states (Table 1 ), were used in a cladistic anaK sis
performed with [Iennig86 (Clopyright J.S. Farris, 1988).
The six multislate characters were entered as unordered.
Most characters were derived from soft tissues (mainly
from the male and female reproductive and alimentarv
systems), the remainder from radulae, opercula, proto-
conchs, and shell ultrastructiire. Two additional species,
Thais nodosa and Muricanthus ftdvescens (Sowerby,
1841), are used in the cladistic analysis, based on data
in Kool (1989; 1993, in press). Muricanthus fulvescens,
a muricine and member of a sister group of Ocenebrinae,
Trophoninae and Rapaninae, is used as outgroup for the
cladistic analysis.
Table 1 lists characters and character states, and re-
flects the sequence in which organs and other morpho-
logical features are described for each of the three spe-
cies.
RESULTS
DESCRIPTIONS OF TAXA
Nucella lapillus.
Shell. Protoconch (Figs. 26, 29) conical, low, of about
l'/4 smooth whorls, with impressed suture; transition to
teleoconch smooth, difficult to discern. Teleoconch high-
ly polymorphic; usually elongate, oval, of 6-7 whorls
(Figs. 1-6, 21, 22). Adult shell to 55 mm in height, 30
mm in u idth. Body whorl rounded, about 80?f of shell
height, smooth or sculptured with pattern of about 15
spiral, occasionally lamellose. ridges. Aperture (to 65%
of shell height) oval; outer lip w ide, smooth, occasionally
with 3-4 denticles on edge of thickened lip. Columella
v\ith moderate callus, flat to concave. Siphonal canal
short, open (Fig. 3) to partly closed (Fig. 1). Siphonal
fascicle poorly developed, adjacent to callus layer. Shell
color variable: white, grey, yellow, brown, orange-red;
often banded; aperture, colmiiella white
Shell Ultraslrncturc. Innermost laverof crossed-lamellar
S. p. Kool, 1993
Page 45
aragoiiite, with crystal planes oriented perpendicular to
growing edge [15-20% of tliiekness; often absent (Fig.
28)]; middle layer of crossed-lamellar aragonite. witfi
crystal planes oriented parallel to grow ing edge [15-25%
of thickness]; outermost laser of calcite [55-85%) of thick-
ness] (Fig, 27).
Operculum D-shaped, with lateral nucleus just below
center right (Figs. 7, 8). Outer surface (Fig. 7) with arch-
shaped growth lines recurved at both ends; inner surface
(Fig. 8) with 3-5 arch-shaped growth lines, with broad
(35-40% of opercular width), callused, glazed outer rim
HeocI and Foot. L'niformly light yellow to white. Ce-
phalic tentacles elongate, thin. Incurrent siphon short.
Mantle edge smooth. Accessory boring organ (Fig. 56,
abo) large, well developed, (in females) anterior to, sep-
arate from equally large ventral pedal gland (Fig. 56,
Pg)-
Mantle Cavity Osphradium slightK more than 'i cten-
idial length, less than l; ctenidial width. Right pectin
usually wider than left. Each lamella (8-10/mm) at-
tached to mantle roof along V2 its base. Anteriormost
portion of ctenidium straight, extending slightly anterior
of osphradium Ctenidial knnellae (9-1 1/mm) wider than
high or equalK wide as high, with strongly convex or
straight lateral edges, translucent. Thick supporting rod
extending beyond lateral edge of each lamella, forming
small papilla.
Female Reproductive System: Vaginal opening round
with slightK swollen edges, located below and posterior
to anus. Bursa copulatrix (Fig. 47, 48, be) small diver-
ticulum, connected to vagina, ventral channel (vc) by
wide ventral passage. Ventral channel formed by two
small interlocking flanges located under ventral lobe of
capsule gland, one arising from left lobe, the other from
ventral epithelium. Single-chambered ingesting gland lo-
cated between capsule gland and albumen gland. Al-
bumen gland (Fig. 49) arch-shaped, elongate, opening
anteriorly into ovi-sperm duct (osd), posteriorly into ovi-
duct (od). Ovary yellow to light golden. Many specimens
with pseudo-penis of variable size (see also Brvan et al.,
1986).
Male Reproductive System: Penis (Fig. 60) simple, elon-
gate, dorso-ventralK flattened, often slightK curved, w ith
abruptly tapering, papilla-like end. Penial vas deferens
(Fig. 62, pvd) minute, simple duct, semi-closed by trans-
verse ridges on overlapping ventral and dorsal sides of
penis. Cephalic vas deferens (Fig. 57, cvd) well devel-
oped, extending from penis (p) to prostate gland (pr).
Prostate gland (Fig. 57, pr) white; prostate duct (prd)
dorso-ventral slit in cross section; duct open to mantle
cavity posteriorly. Posterior vas deferens (along visceral
mass) well developed, white to dirty white, iridescent.
Testis light brow n to golden.
Alimentary System: Paired accessors salivar\ glands ex-
tremely long, usually equal to or slightly longer than one-
half of shell height; left gland intertwined with salivary
glamls, right gland separate from salivary glands, situ-
ated in right anterior corner of buccal cavity. Salivar%
glands in center of dorsal buccal cavity between gland
of Leiblein and short, pear-shaped valve of Leiblein.
Sali\ar> ducts attached to anterior esophagus at some
distance from valve. (Glandular folds in mid-esophageal
region inconspicuous. Connection between mid-esoph-
agus and gland of Leiblein short, thick. Posterior esoph-
agus appressed to left side of gland of Leiblein in loop-
shaped fashion. Gland of Leiblein \ellowish; posterior
blind duct very short ( < Vi length of gland), with small
terminal ampulla. Stomach tubular, with 8-12 large, ra-
dialK oriented folds on wall. Stomach typhlosole ex-
tending dorsally onto left portion of posterior mixing
area. Intestinal t\ phlosole thick, w ide. Tw o digestive di-
verticula present. Rectal gland inconspicuous. Large pa-
pilla overlying equally large anus.
Radula: Ribbon length 30-35% of shell height. Base of
rachidian tooth expanded below fwse of neighboring ra-
chidian tooth; central cusp of rachidian thin, flame-
shaped, leaning more anteriorly (in situ) than lateral
cusps; inner lateral denticle low on base of lateral cusp,
occasionalK bifurcate (w ithin same specimen); outer edge
of lateral cusp w ith se\ eral denticles; large marginal cusp
pointing straight forward and parallel to elongate, lateral
extension at base of rachidian tooth (Figs. 23-25), re-
sulting in bifid appearance of rachidian basal plate. Lat-
eral teeth shorter than rachidian width (Fig. 23).
Egg capsules: Elongate-oval, vase-shaped, up to 9 mm
in height, 3 mm in w iilth. Capsules \ellow, light brow n
or purple (Lebour, 1937), each attached by short, thin
stalk about 1 mm long. Apex tapered with rounded, cap-
shaped top with mucous plug. Capsules interconnected
at base. Number of embryos varying from two (Risbec,
1937) to one thousand (Fretter & Graham, 1962, 1985),
most being nurse eggs (75-95%) (Crothers, 1985; Fretter
& Graham, 1985; Lamv, 1928) (see also Pelseneer, 1911;
Ankel, 1937; Thorson, 1941, 1946; Robertson, 1974).
Ecology: More is known about Nucclla ecology than
about any other muricoidean [for an extensive bibliog-
raphy on the biology (primarily ecology) of Nticella la-
pillus, see Crothers, 1985]. Nucella lapUlus and its west-
ern American congeners have been the topic of man\
comprehensive studies (Crothers, 1985; Emien, 1966; Et-
ter, 1987; Kincaid, 1957; Spight, 1972). Nucella feeds on
barnacles and mussels (Colton, 1922; Connell, 1970;
Crothers, 1973; Graham, 1955; Kool, 1987; Largen, 1967;
Murdoch, 1969; Spight, 1982) in the rocky intertidal zone
and is eaten by crabs and birds (Spight, 1976). Studies
of Agersborg (1929), Colton (1922), and Moore (1936)
show that environmental factors (wave action, food avail-
abilits, etc.) influence shell morphology. Moore (1938)
reported the main spaw ning period to be during winter
and spring; but breeding occurs throughout the year
(Lebour, 1937; Thorson, 1946). Juveniles hatch from the
eggs after 4-7 months (Fretter & Graham, 1985).
Distribution: North Atlantic Ocean from southern Por-
Page 46
THE NAUTILUS, Vol. 107, No. 2
7
^^^
%
^^^
/ ■
/.
11
8
mi'
17
^^m
'^'
18
19 *-
Figures 1-8 .V«C(?//a /api7/us. 1-6. Shells (1,2 MCZ 69192, Freshwater Bay. Isle of Wright. England, height 25 mm; 3,4 MCZ
1 1.509.3. Sullivan, Maine. I S .V . height .35 mm; 5.6 MC.Z 50600. Wales, height 3-1 mm) 7.8 Operculum (MCZ 302404, Braunton,
North Devon, England, height 15 mm); 7. Outer surface; 8. Inner surface. Figures »J-14 Trophon gevcrsianus. Shells (9.10 MCZ
S. p. Kool. 1993
Page 47
tugal to Xovaya Zembl\a [records from western Medi-
terranean, Azores, Morocco, Senegal, and Canary Islands
are suspect (Cooke. 1915)]; Great Britain; Ireland; Ice-
land; Greenland; New Jerse\, L.S.A.. to northern Canada
[for extensi\e list of geographical range and localities,
see Cooke, 1915].
Trophon geversianus:
Harasewych (1984) described aspects of the anatom\ of
Trophon geversianus (Figs. 9. 10, 13, 14. 30. 31 ). Because
my observations were congruent w ith the descriptions in
Harasewych s paper, only the most essential and supple-
mental data are presented to avoid unnecessary dupli-
cation.
Shell ultrastructure: Innermost la\er of crossed-lamellar
aragonite, w ith cr> stal planes oriented perpendicular to
growing edge [10-15!^ of thickness; often absent (Fig.
37)]; middle layer of crossed-lamellar aragonite, with
crystal planes oriented parallel to growing edge [15-20%
of thickness]; outermost layer of calcite [70-80% of thick-
ness] (Fig. 36).
Operculum: 0\ate, with lateral nucleus in lower right
(Figs. 11, 12). Outer surface (Fig. 11) with growth lines
recurved at upper ends, progressiveh upright; inner sur-
face (Fig. 12) with 3-4 narrow horseshoe-shaped growth
lines, broad ( > '3 opercular width ). lightK callused, glazed
outer rim.
Head and foot: UniformK light \ellow Cephalic ten-
tacles elongate, thin. Incurrent siphon short. Mantle edge
smooth, .\ccessory boring organ (Fig. 55, abo) well de-
veloped, sharing common duct with ventral pedal gland
(pg) in females.
Mantle Cavity: Osphradium small ('/3-% ctenidial length.
-/i ctenidial width). usualK partialK overK ing ctenidium.
Right pectin usualK wider than left. Each lamella (9-
10 mm) attached to mantle roof along most of its length.
.•\nteriormost portion of ctenidium straight, extending
slightly anterior of osphradium. Ctenidial lamellae (10-
12 mm) translucent, wider than high anteriorK. equalK
wide as high posteriorly, with straight to convex lateral
edges.
Female Reproductive System: \'aginal opening (Fig. 47,
vo) round, with swollen edges, located below, slightly
anterior to anus. Bursa copulatrix (Figs. 47, 48, be) small
diverticulum, connected to vagina and ventral channel
(vc) by wide duct. Wall of posterior vagina with folds
decreasing in number posteriorly. Capsule gland with
simple, inconspicuous ventral channel posteriorly con-
nected to large, well-de\eloped ingesting gland filled
with w hitish substance. .Albumen gland (Fig. 49) large,
arch-shaped, elongate, opening anteriorly into ovi-sperm
duct (osd), posteriorly into oviduct (od).
Male Reproductive System: Penis (Fig. 59) bulbous, short,
dorso-\entrall\ flattened, w ith large papilla. Penial vas
deferens (Fig. 62. p\d) minute, simple duct, closed by
overlapping ventral and dorsal sides of penis. Cephalic
vas deferens (Fig. 57, cvd) well developed. Prostate gland
(Fig. 57. pr) light yellow; prostate duct (prd) dorso-ven-
tral slit in cross section; duct open to mantle cavit\ pos-
teriorly.
Alimentary System: Paired accessory- salivar\ glands short
(<1 10 shell height); left gland intertwined with left
salivary gland, right gland free, situated in right anterior
corner of buccal cavity. Salivary glands in center of dorsal
buccal ca\it\ between gland of Leiblein and elongate
\al\e of Leiblein. Salivar\ ducts attached to anterior
esophagus immediately anterior to \alve. Glandular folds
in mid-esophageal region well developed. Connection
betw een mid-esophagus and gland of Leiblein short, thick.
Posterior esophagus appressed to left side of gland of
Leiblein in loop-shaped fashion. Gland of Leiblein yel-
low ish; posterior blind duct short, without terminal am-
pulla. Stomach tubular, with 10-15 thin, elevated folds
on w all; posterior ones oriented toward center, anterior
ones merging into elevated section of sorting area. In-
testinal typhlosole thin. Two digestive diverticula pres-
ent. Rectal gland light brown, extending along '2 of pal-
lia] gonoduct.
Radula: Ribbon length 40-45% of shell height. Base of
rachidian tooth expanded below base of neighboring ra-
chidian tooth; central cusp thin, with wide base; inner
lateral denticle small protrusion from base of lateral cusps;
outer edge of lateral cusp straight, with several faint
denticles; large marginal cusp pointing straight forward
and parallel to faint, elongate, lateral extension at base
of rachidian tooth (Figs. 32-34), resulting in bifid ap-
pearance of rachidian basal plate. Lateral teeth shorter
than rachidian width, with wide bases positioned close
together (Fig. 32).
Egg capsules: Discoidal, lateral!) flattened, with wide,
equalK flattened base, up to 20 mm in height (including
base), 12 mm in width (Harasewych, 1984, Fig. 23).
Capsules yellowish in color, and containing 74-112 em-
bryos (MeKill & Standen, 1898). Capsules deposited in
row s w ith flattened edges adjacent to one another (Lamv,
1928; D'Asaro, 1991).
Ecology: This species lives in the rocky intertidal and
subtidal zones where barnacles and mussels are plentiful.
10941.3. Ushaia, Tierra del Fuego, Argentina, height 41 mm; 1:{.I4 MCZ 1-32566. Falkland Islands. Argentina, height 47 mm).
11.12 Operculum iLACM 86-270.2, Daniel Este. Isla Grande. Tierra del Fuego. Chile, height 11 mm); 11. Outer surface; 12.
Inner surface. Figures 15-20 Ocenebra erinacea. Shells (15.16 MCZ 87662. \Ve\mouth. England, height 31 mm; 19,20 MCZ
1724.50. Fos-sur-Mer. Bouches-du-Rhone, France, height 56 mm) Operculum (17.18 MCZ 302405, St. Lunaire. France, height 8 0
mm); 17 Outer surface; 18. Inner surface.
Page 48
THE NAUTILUS, Vol. 107, No. 2
Figures 21-29 \ucclla lapillus. 21,22 Shell (MCZ 09192, Freshwater Bay, Isle ot Wight, England, height 2.5 1 mm). 23-2.S
Kadiila (ISNM 8.570.53, Kittery, Maine, LI.S.A.). 23. Clentral portion of radular ribbon. Scale bar = 15 fim. 24 Rachidian teeth
Scale bar = 10 ^m. 25. Side view of rachidian teeth (right row of lateral teeth removed). Scale bar = 10 fim. 26,29 Protoconch
(MCZ 14184, Isle au Ilaut, Maine, U.S.A.). 26. Apical view. Scale bar = 12 m"' 29. Side view. Scale bar = 12 ^m 27,28 Shell
ultrastructure (view of growing edge; innermost layer on bottom) (MCZ 69192, Freshwater Ba>, Isle of Wight, England) Scale
bars, 4.5 ixn\ and 70 niu, respectively.
S. p. Kool, 1993
Page 49
Figures 30-38. Troplion gcicrsianns. 30.31. Sliell (MC^Z 1.32566. Falkland Islands, Argentina, lu'iglit -JT mm). 32-3 4-. Radula.
(LACM 86-270.2. Daniel Kste. Isia Grande. Tierra del Fuego. Chile) 32. Central portion of radiilar ribbon. Scale bar = 45 nm.
33. Rachidian teeth Scale liar = 25 ^m. 34 Side view of rachidian teeth. Scale bar = 20 ^in 35,38. Protoconch (LACM 86-
270.2, Daniel Este, Isla Grande, Tierra tlel Fuego, Chile). 35. Apical view. Scale bar = 150 ^m. 38 Side view. Scale bar = 150
Mm 36.37 Shell ultrastructure (L.ACM 86-270 2, Daniel Fsle. Isla (Iraiide, Tierra del F'liego, Chile); Scale bars, 60 fim and 150
Mm, respectivel)
Page 50
THE NAUTILUS, Vol. 107, No. 2
4*
^
39 W
i
■?•■,■ ■
40 '
i^BFjlg;
^
atk..:
.«►
f43
IP
^*^.
■^
Fif^ures 39-46. Ocenebra crinai-ciL 39,40, Sliell (MCZ 172450, Fos-sur Mer, Botichfs-dii-Hlioiu", France, height 56 mm) 41-43
Kathila (MCZ 298425, Roseoll, France). 41 Oenlral portion of rihhoii Scale bar = 17 ^ni 42. Rachichan teeth Scale bar = 8 ^m.
43 Side view of rachidian teeth. Scale bar = 9 m"i 44.46 Protoconch (MC:Z 38369, Kent, Fngland) 44. .Apical view. Scale bar
= 9(1 A/ni 46 Side view. Scale bar = 90 niu 45 Shell iillrastnictiire (MCZ 298425, Roscoff, France), Scale bar = 1.30 ^m.
S. p. Kool, 1993
Page 51
Figures 47-56. Anatomical structures of the female repro-
ductive system of Nitcella lapilltis. Trophon geversianus, Oce-
nehra erinacea, Muricanthus fulvescens. and Thais nodosa.
47.50.53. Bursa copulatrix in iV lapillus and T. geversianus
(47), Th. nodosa (50), O erinacea ami M. fulvesccns (53).
48.51.54. Cross sections through bursa copulatrix (location
indicated b>' vertical bar). 49,52. Albumen glands in N. la-
pillus. T gever.'iianus. O erinacea. M Julvescens (49), anil
Th. nodosa (52), 55,56. Sagittal section through foot showing
accessory boring organ and pedal gland of Th. nodosa, T
geversianus, M. julvescens (55), N . lapillus. and O. erinacea
(56).
abo, acces.sory boring organ; be, bursa copulatrix, db, duct to bursa
copulatrix; leg, left lobe of capsule gland, od, oviduct, osd. ovi-sperm
duct; pg, ventral pedal gland; p,sr. posterior seminal receptacles, rcg.
right lobe of capsule gland, tf, transverse fold; vc. ventral channel, \cf,
flange of the ventral chaiuiel, vo. vaginal opening.
Sculpture of the shell may vary v\ itli the t\ pe ot habitat
Distribution: Southern Argentina to Chile.
Ocenebra erinacea:
Shell: Protoconch (Figs. 44, 46) conical, low, of l'/2 smooth
whorls, and with impressed suture; transition to teleo-
conch smooth, difhcult to discern. Teleoconch elongate,
fusiform, biconical (Figs. 15, 16, 19, 20, 39, 40), of 7-8
whorls. Adult shell highly variable in shape, to 55 mm
in height, 25 mm in width (not including spine length;
35 mm including spine length). Body whorl 55-60?c of
shell height, with 3-9 varices, often with frilled edges,
and with (3-8 spiral cords. Aperture (to 30% of shell height)
round to oval; outer lip with crenulated edge Moderately
callused columella. Siphonal canal partly or completely
closed, often nearly equal in length to aperture in larger
Figures 57-64 .■\nal<imical structures ol the male rcjiroductive
system of S'ucclla lapillus. Trophon geversianus. Ocenebra
erinacea. Muricanthus fukescens, and Thais nodosa. Prostate
gland with proximal and distal cross sections in N. lapillus. T.
geversianus. O erinacea, M . julvescens (57) and Th. nodosa
(58). Penial morphologies with cross sections in T. geversianus
(59.62), N. lapillus. M. fulvesccns {f>0,b2). O erinacea (60.63),
and Th nodosa (61,64).
b\ , blood vessel, cvd, cephalic vas deferens; p, penis, pr, prostate, prd,
duct through prostate, pvd, penial vas deferens.
specimens. Siphonal fasciole pointing away from si-
phonal canal. Shell color yellov\ish to cream or dark
brown; aperture, columella white.
Shell ultrastructure: Innermost layer of crossed-lamellar
aragonite, with crystal planes oriented perpendicular to
growing edge [15-20% of thickness]; followed by layer
of crossed-lamellar aragonite, with crystal planes ori-
ented parallel to growing edge [40-45% of thickness];
followed by layer of crossed-lamellar aragonite, with
crystal planes oriented perpendicular to growing edge
[5-8%. of thickness]; outermost layer of calcite [35%. of
thickness] (Fig. 45).
Operculum: D-shaped, with lateral nucleus in lower right
(Figs. 17, 18). Outer surface (Fig. 17) with arch-shaped
growth lines progressively upright, recurved at upper
end; inner surface (Fig. 18) with 4-5 arch-shaped growth
lines, w ith broad (~ Vz opercular w idth), lightly callused,
glazed outer rim.
Head and foot: I'niformly light yellow to white. Ce-
phalic tentacles elongate, thin. Incurrent siphon well de-
veloped. Mantle edge smooth, occasionally with crenu-
lations (possibly an artifact). Sole of foot w ith large lateral
Page 52
THE NAUTILUS, Vol. 107, No. 2
(0
s
(0
(0
c
o
Q.
O
(0
c
O
o
i5
8-1
10-
11 -
4 -
3-
—
8-
16-
-
12 -
11 -
3-
13-
7-
5 -
-
10 -
9-
-
-
8-
17 -
-
5-
15 -
-
4-
14-
-
3 -
6 -
-
4 -
1 -
-
13-
^
2-
—
Fipurp 65 Cladciyram, based on analysis of data in Table 1,
shduiiig high |)ii\l()geiietic affinity between Nucclla and Oce-
nebra. Numbers next to character changes correspond to num-
bers given to characters in Table 1. Alternative, equally par-
simonious character state transformation series are possible for
characters 3, 4, 5, and 8 (see text).
folds. Accessory boring organ (Fig. 56, abo) large, well
developed, (in females) anterior to, separate from equally
large ventral pedal gland (Fig. 56, pg).
Mantle Cavity. Ospliradium (<'/2 ctenidial length, %
ctenidial width) partially extending over ctenidium. Right
pectin usually wider than left. Each lamella ( 10-1 l/mm)
attached to mantle roof along short basal portion. An-
teriormost portion of ctenidium straight, extending an-
teriorly of osphradium. (Ctenidial lamellae (11-12/mm)
basically triangular, longer than high w ith convex lateral
edges posteriorly. Lamellar support rods not well devel-
oped.
Female Reproductive System: Vaginal opening (Fig. 53,
vo) round to elongate, on short, extension of pallial gon-
oduct, situated below, posterior to anus. Bursa copulatrix
(Figs. 53, 54, be) large diverticuhmi (equal in diameter
to capsule gland), connected at its anterior portion to
duct running parallel to ventral channel for some length
prior to connecting with it (Figs. 53, 54, db). Lumen of
bursa copulatrix Oiled w ith loose flocculent material and
iridescent spherules. Capsule gland posterior to, shorter
than bursa copulatrix. Ventral channel (Figures 53, 54,
vc) well developed anteriorly, less distinct posteriorly.
Ingesting gland situated between capsule gland, albumen
gland. Albumen gland (Fig. 49) arch-shaped, elongate,
opening anteriorly into ovi-sperm duct (osd) posteriorly
into oviduct (od). Ovary orange-yellow. Pseudo-penis oc-
casionally present (see also Feral, 1976).
Male Reproductive System. Penis (Fig. 60) simple, dor-
so-ventrally flattened, elongate, slightly curved, with
abruptly tapering, papilla-like end. Penial vas deferens
(Fig. 63, pvd) a wide, straight tube star-shaped in cross
section; tube partialK attached to penial wall Cephalic
vas deferens (Fig. 57, cvd) large, similar to penial vas
deferens in structure, extending from penis (p) to prostate
gland (pr). Prostate gland (Fig. 57, pr) white to yellow;
prostate duct (prd) a dorso-ventral slit in cross section
anteriorK , a triangular large space posteriori)-; duct open
to mantle cavity posteriorly (Fig. 57). Posterior vas de-
ferens white to dirty white, iridescent. Testis yellowish.
Alimentary System: Paired accessory salivary glands very
long (* '/2 shell height); left gland intertwined with left
salivary gland, right gland free, situated in right anterior
corner of buccal cavity. Salivary glands in center of dorsal
buccal cavity between gland of Leibleiii and short, pear-
shaped valve of Leiblein. Salivary ducts attached to an-
terior esophagus at short distance from valve. Glandular
folds in mid-esophageal region swollen, especially well
developed at connecting point between esophagus and
gland of Leiblein. C-onnection between mid-esophagus
and gland of Leiblein short, thick. Posterior esophagus
appressed to left side of gland of Leiblein in loop-shaped
fashion. Gland of Leiblein yellowish; posterior blind duct
very short (<V2 length gland of Leiblein), with promi-
nent terminal am|Milla. Stomach tubular, w ith faint folds
on stomach wall oriented tow ard center of stomach (poorly
preserved in specimens examined). Rectal gland thin,
green, extending along V2 oi pallial gonoduct. Large pa-
pilla overla\ ing anus
Radula: Riblion length 35-40% shell height. Base of ra-
chidian tooth expanded below base of neighboring ra-
chidian tooth; central cusp of rachidian thin, flame-
shaped, leaning more anteriorly (in situ) than lateral
cusps; inner lateral ilenticle low on base of lateral cusp,
occasionally bifurcate (w ithiii same specimen); outer edge
of lateral cusp straight, with several short denticles; large
marginal cus() pointing straight forward and parallel to
elongate, lateral extension at base of rachidian tooth (Figs.
41-43), resulting in bihd appearance of rachidian basal
plate. Lateral teeth with narrow base, wideK spaced,
shorter than rachidian width (Fig. 41).
S. P Kool, 1993
Page 53
Egg capsules: Oval-elongale, vase-shapt-il, trianuulai in
cross section. Capsules \ell()\visli, up to 13 mm in lu-iulit,
6 mm in width, each on narrow stalk, attached b\ Hat
base. Apex with short elevated protuberance with mu-
cous plug Each capsule containing 4-167 embryos, of
which none are nurse eggs (Fretter & Graham, 1985;
Lebour, 1937).
Ecology: Occncbra crinacea li\es in the rocky intertidal
and subtidal zones (Jeffreys, 1867) to 150 meters (Fretter
& Graham, 1985) where it feeds on oysters and other
bivalves (Fretter & Graham, 1985), barnacles ami limpets
(Graham, 1955). Lebour (1937) reported tireediiii; in late
spring ant! summer Juveniles hatch from the eggs after
12-14 weeks (Fretter & Graham, 1985). A comprehensive
ecophysiologicai study was done on Ocenebra erinacea
by Hawkins (1985)
Distribution: Nfirth Atlantic Ocean (Spain to Norway,
Great Britain, Ireland); Mediterranean Sea (southern Eu-
rope, northern Africa).
CLAPISTIC ANALYSIS
The cladistic analysis yielded one tree with a consistency
inde.x of 0.88 (Fig. 65). This cladogram indicates that:
1) placement of \ucclla in Thaidinae or its senior syn-
on\ ni, Rapaninae, creates polyphyletic groups (see also
Kool, 1989); 2) Nucclla is more closely related to Oce-
nebra than to Troplwn and should be placed in Oce-
nebrinae; 3) the subfamilial boundaries between Oce-
nebrinae and Tro[)honinae are much less distinct than
previousK accepted.
Characters and character state distribution among the
taxa Nucclla. Trophon. Ocenebra. Thais, and the out-
group, Muricanthus:
Character 1. Protoconch: 0. multispiral ( > 2 whorls); 1,
paucispiral (< 1': whorls).
Nucella. Trophon and Occncbra (Figs. 26, 29, 35, 38,
44, 46) have a paucispiral, smooth protoconch without a
sinusigeral notch or outwardly-flared lip. This morphol-
ogy reflects direct development. The outgroup and the
rapanines have a multispiral (and generalK sculptured)
protoconch w ith a sinusigeral notch and outwaril-tlaring
lip, t\pical for species v\itli a pianktonic larval stage.
Character 2. Calcitic layer; 0. absent; 1. present.
The outgroup lacks an outer calcitic shell layer that is
present in the other four ta.xa. The presence of calcite
appears to be the derived condition.
Character 3. Number of aragonitic layers: 0. 3; 1. 4; 2. 2
The outgroup and Ocenebra have three layers of ara-
gonite (transverse, collabral, transverse), Thais has an
additional, innermost fourth layer of crystals oriented in
a 45° angle; both Nucella and Trophon have two layers
(transverse, collabral), but may lack the innermost trans-
verse layer (Figs. 28, 37).
Character 4. Nucleus of operculum: 0. terminal nucleus
in lower right; 1. lateral nucleus in lower right; 2. lateral
niiclfus licldw center right; 3 lateral nucleus in center
right.
The outgroup has a terminal nucleus in the lower right.
.All iiigroup taxa have a lateral nucleus, the position of
w Inch varies. The nucleus of the operculum in Nucella
is located below the center right; that of Trophon and
Ocenebra in the lower right (Figs. 7, 11, 17, respectively).
The nucleus is located in the center right in rapanines.
C;haracter 5, Shape of operculum: 0. oval; 1. D-shaped.
The opercula of Nucella, Thais, and Ocenelna are
roughly D-shaped Those of Muricanthus and Trophon
are elongate-oval.
(Character 6, Pigmentation pattern of head-toot region:
0 jjresent; flecked with black and gra\; 1. absent, uni-
formly colored (faint yellow ).
The head-foot regions of Nucella, Trophon, and Oce-
nebra are uniformly faint yellow. Both Thais and the
outgroup are densely flecked with black blotches and
specks (this pattern generalK survives preservation in
alcohol although other colors, such as white and yellow,
fade).
Character 7. Ventral pedal gland and accessory boring
organ: 0. sharing one duct; 1. with separate ducts.
In Nucella and Ocenebra, the accessory boring organ
and ventral pedal gland (Fig. 56, abo, pg) have separate
ducts to the sole of the loot, while in Trophon, rapanines,
and the outgroup these structures share a common duct
(Fig. 55, abo, pg).
CJiaracter 8. Bursa copulatrix: 0. large diverticulum, sep-
arate from capsule gland; 1. small diverticulum, separate
from capsule gland; 2. small chamber with lumen con-
tinuous with capsule gland.
The bursa copulatrix is a small blind sack in Nucella
and Trophon (Fig. 47), a large separate diverticulum in
Ocenebra (Fig. 53) and the outgroup, while in rapanines
the bursa is continuous with the capsule gland (Figs. 50,
51).
Character 9. Seminal receptacles at dorsal periphery of
albumen gland: 0. absent; I. present.
In Thais, a row of posterior seminal receptacles (Fig.
52. psr) at the dorsal periphery of the albumen gland
presumably increases efficiency in the fertilization pro-
cess (Kool, 1988, 1989). These posterior seminal recep-
tacles are absent in Nucella, Trophon. Ocenebra. and in
the outgroup, Muricanthus (Fig. 49). This character is
a synapomorphy for Rapaninae (Kool, 1993, in press).
Character 10. Penial shape: 0. simple, elongate to lightly
curved; 1. bulbous, with papilla; 2 strongK recurved
with pseudo-papilla.
Penial shape in Nucella. Ocenebra. and the outgroup
is elongate (F"ig. 60). The penis in Thais is strongly
recurved, and sinuous (Fig. 61), while that of Trophon
is short, bulbous, with a distinct papilla (Fig. 59).
(Character 11 Penial \as deferens: 0. simple duct; 1.
small, loose duct-w ithin-a-duct; 2. large duct-within-a-
duct, partialK attached to penial iiuier wall
Page 54
THE NAUTILUS. Vol. 107, No. 2
Nucella, Trophon and the outgroiip have a peiiial vas
deferens that is loosely closed, while rapanines have a
■■duct-vvithiii-a-duct" system (Figs. 61, 64) (Kool, 1988,
1989). Ocenehra differs from both types in having a
rather wide inner duct that is partially attached to the
penial inner wall (Fig. 63).
Character 12. Prostate Gland: 0. open to mantle cavity
posteriorly; 1. without opening to mantle cavity.
Males of Nucella. Trophon. Occnelrra. and the out-
group have a prostate gland that is open to the mantle
cavity along its posterior portion (Fig. 57). The prostate
of rapanine males does not open to the mantle cavity
(Fig. 58).
Character 13. Accessory salivary gland length: 0. right
gland small, left gland absent; 1. glands < '/i shell height;
2. glands > V2 shell height.
Character 14. Straw-like membrane around gland of
Leiblein: 0. present; 1. absent.
In most rapanines and the outgroup, the gland of Leib-
lein is covered by a thick membrane of interwoven fibers
of connective tissue, producing a straw-like appearance.
Such a membrane is absent in Nucella. Trophon and
Ocenehra.
Character 15. Posterior duct of gland of Leiblein: 0.
longer than Vz of gland length; 1. shorter than V2 gland
length.
In Nucella, Trophon and Ocenehra. the gland of Leib-
lein tapers posteriorly into a thin, very short posterior
duct that runs adjacent to the posterior esophagus and
is often filled with secretory material from the gland; in
the majority of rapanines and in the outgroup this duct
is much longer, extending into the dorsal branch of the
afferent renal vein.
Character 16. Central cusp of rachidian: 0. oriented in
same plane as lateral cusps; 1. leaning more anteriorly
than lateral cusps.
In Nucella (Fig. 25) and Ocenehra (Fig. 43) the central
cusp on the rachidian leans more anteriorly (in situ ) than
the lateral cusps. In Thais. Trophon (Fig. 34), and the
outgroup, the lateral cusps and central cusp are aligned
in the same plane.
Character 17. Margin of rachidian basal plate: 0. straight;
1. bifid.
The bifid condition of the rachidian basal plate (Figs.
25, 34, most developed in Fig. 43) is found in Nucella,
Trophon and Oceneina, but not in Thais or the outgroup.
Synapornorphies for the Nucella-Ocenehra clade {Fig.
65):
Character 5: The character for opercular shape is ho-
moplastic; a D-shaped operculum occurs in Thais as well.
C]haracter 7: Both Nucella and Ocenehra have separate
openings for the ventral pedal gland and accessory boring
organ (Fig. 56, abo, pg). Female specimens of Trophon
geversianus, Thais nodosa, and the outgroup, have a
single duct and opening for these organs (l"ig. 55, abo.
pg). A shared duct for the accessory boring organ and
ventral pedal gland, as found in Trophon geversianus,
may not be as advantageous as an arrangement as w hen
the ducts originating from the accessory boring organ
and ventral pedal gland are separate. It would appear
that an arrangement where one duct serves both as ven-
tral pedal gland and as a passage for the accessory boring
organ and its secretions during boring activities (Carri-
ker, 1981) prevents the female from boring activity, and
thus perhaps feeding in general, during stages of egg-
laying.
Character 13: Nucella and Ocenehra have a pair of very
long accessory salivary glands (> ','2 shell height). Tro-
p/!o;(andT/ia!shavemuchsmallerglands(< '/^ shell height).
The outgroup has only one extremely small right acces-
sory salivary gland.
Character 16: In Nucella, and to a much greater degree
in Ocenehra, the central cusp leans more anteriorly (Figs.
25, 43, respectively) (in situ) than the lateral cusps,
whereas in Trophon, the central cusp is aligned with the
lateral cusps (Fig. 34).
Synapornorphies for the Trophon-Nucella-Ocenehra
Clade:
Character 1: The paucispiral protoconch is indicative of
having crawl-away larvae, rather than a planktonic larval
stage that is found in Thais and other rapanines (Kool,
1993, in press) and the outgroup.
Character 4: The cladogram suggests that an opercular
nucleus below the center right (Nucella; character state
2) evolved from the ancestral condition for the taxa in
this clade of having a nucleus in the lower right (Oce-
nehra and Trophon; character state 1).
Character 6: All three taxa lack a pigmentation pattern
on their head-foot region that is found in the outgroup,
Thais, and other members of the Rapaninae.
Character 14: The straw-like outer membrane of the
gland of Leiblein is absent in the species of this clade,
but present in the outgroup, and most members of Ra-
paninae (Kool, 1989; 1993, in press).
Character 15: The posterior duct of the gland of Leiblein
is shorter than V2 the length ot the gland itself in this
clade, but much longer in the remaining taxa, reaching
into the dorsal branch of the afferent renal vein.
Character 17: The bifid condition of the basal plate,
especially well developed in Ocenehra (Fig. 43), is absent
in Thais and the outgroup.
Synaponiorphics for the Thais-Trophon -Nucella-Oce-
nehra Clade:
Character 2: An outer layer of calcite is present in all
ingroup taxa, but is absent in the outgroup.
Character 13: The outgroup has only one extremely small
right accessory .sali\ar> gland. A situation of having a
pair of medium-size accessory salivary glands appears to
S. p. Kool, 1993
Page 55
have evolved from the condition described above and to
have given rise to the most deri\ed condition (extremely
long glands).
DISCUSSION
According to the topology of the cladogram (Fig. 65),
two characters have evolved in a parallel manner in
Nucella lapillus and Trophon geversianus. Out of the
context of the cladogram, these similarities would suggest
a closer relationship between these two species than is
suggested by the tree topology:
Character 3: Shell ultrastructure in both Nucella and
Trophon consists of two aragonitic layers and an outer
layer of calcite. Specimens of both may lack the inner-
most (transverse) layer (Figs. 28, 37). More detailed stud-
ies may reveal the cause of this variation. Perhaps en-
vironmental factors may play a role (Etter, personal
communication).
Character 8: The morphology of the bursa copulatrix of
Nucella is very similar to that of Trophon. In both taxa,
a relatively small, muscular blind sack branches off from
the vagina (Figs. 47, 48). In Ocenebra the bursa is thin-
walled and equal in width and height to the capsule
gland, extending for up to V2 the length of the pallial
complex. The ventral channel loops backwards towards
the anterior portion of the bursa in Ocenebra (Fig. 53),
rather than straight up into the bursa as in Trophon and
Nucella (Fig. 47).
The above two characters could be considered synapo-
morphies for an alternative, but less parsimonious, tree
in which Trophon and Nucella would be united in one
clade. However, in the proposed phylogenetic hypothesis
(Fig. 65), four synapomorphies support a clade consisting
of Nucella and Ocenebra. rendering the above two char-
acters as homoplastic.
Fretter and Graham (1962) mention several similari-
ties in egg capsule morphology between Nucella lapillus
and Ocenebra erinacea. Both species lay vase-shaped
capsules, whereas Trophon geversianus produces dis-
coidal egg capsules. However, having discoidal egg cap-
sules is only an autapomorphic trait for Trophon and
does not provide clues about relationship in this case.
Alternative, equally parsimonious transformation series
are possible for characters 3, 4, 5, and 8. For Characters
3, 5, and 8, I chose the scenario involving homoplasy
over one involving a reversal to avoid an "artificial"
increase in synapomorphies. Similarly, for Character 4,
I chose the least linear trans-formation series ("zero state"
evolving into both the "three state" and the "one state").
Zoogeographical data reveal that members of Nucella,
Ocenebra and Trophon occur primarily in colder waters
of the temperate and boreal zones, whereas rapanines
and the outgroup occur primarily in warmer waters of
the (sub)tropics (Kool, 1989).
Nucella lapillus and Ocenebra erinacea overlap for
much of their ranges in primarily temperate western
European waters. In addition, N. lapillus occurs in the
western Atlantic, where O. erinacea does not, and O.
erinacea occurs in the Mediterranean, where N. lapillus
has not occurred since the Pleistocene [Malatesta (1960)
cited records of N. lapillus from the Pleistocene of Sicily].
Cooke (1915), in a comprehensive list of localities for
Nucella lapillus, showed that the southernmost record
for this species is the Algarve coast of Portugal and al-
leged that any records from Northern Africa (see Nord-
sieck, 1982), the western Mediterranean, the Azores, and
the Canary Islands, are highly suspect. Other members
of what can be assumed to be Ocenebra s.s. and Nucella
s.s. are found in the eastern Pacific (Abbott, 1974) and
the northern I'acific. Furthermore, Ocenebra s.s. occurs
in western Africa (Houart, 1989) and South Africa (Kil-
burn & Rippey, 1982). Ranges of these genera may be
revised when more Ocenebra-like taxa (for example from
Japan) and Nucclla-Uke species have been examined with
respect to their anatomy, radula, shell ultrastructure, etc.
Such studies are also necessary to determine if, for ex-
ample, species such as Nucella dubia and Nucella squa-
mosa, both from the South African Province, are indeed
members of Nucella s.s. It appears from preliminary
dissections that the genus Nucella can no longer be con-
sidered restricted to the temperate waters of the northern
hemisphere (Kool, in preparation).
Trophon geversianus, limited to the South American
continent, lives in temperate to boreal waters, as do Nu-
cella lapillus and Ocenebra erinacea.
SYSTEMATIC CONCLUSIONS
The high degree of similarity in anatomy, radula, pro-
toconch, shell ultrastructure and operculum in Ocenebra
erinacea, Trophon geversianus and Nucella lapillus in-
dicates that these three taxa are more closely related to
one another than any one of them is to Thais or other
representatives of Rapaninae. Nucella should therefore
be excluded from Rapaninae to maintain monophyly
(Kool, 1989; 1993, in press). The difficulty of correctly
allocating Nucella to a subfamily is indicative of the
dilemma of our lack of understanding of higher muri-
coidean systematics. The cladistic analysis and the re-
sulting cladogram (Fig. 65) suggest that Nucella is better
placed in Ocenebrinae than in Trophoninae. It is obvious
that the boundaries of groups at the higher taxonomic
categories, traditionally based on shell characters, be-
come less clear after completion of thorough anatomical
studies of members from different genera and subfam-
ilies. Results shown here suggest that Trophon geversi-
anus, the type species of Trophoninae, is closely related
to Nucella and Ocenebra. Perhaps it is more closelv
related to the latter two taxa than to other species hitherto
included in Trophoninae. It is beyond the scope of this
paper to suggest synonymization of Trophoninae with
Ocenebrinae. However, I suspect that future studies will
show that Trophoninae is not a monophyletic group and
that a new name for some of its members may be war-
ranted. The following systematic arrangement is pro-
posed for the taxa treated herein:
Page 56
THE NAUTILUS, Vol. 107, No. 2
MURICOIDEA Rafinesque, 1815
MURICIDAE Rafinesque, 1815
RAPANTNAE Gray, 1853 (sen.su Kool, 1993, in press)
Thais Roding, 1798
OCENEBRINAE Cossmann, 1903
Ocenebra Gray, 1847
Nucella R6ding, 1798
TROPHONINAE Cossmann, 1903)
Trophon Montfort, 1810
ACKNOWLEDGEMENTS
I thank Drs. Richard S. Houbrick, M. G. Harasevvych,
and Kenneth J. Boss for reviewing an early draft of this
manuscript. I thank the staff of the Scanning Electron
Microscope Laboratories at the United States National
Museum of Natural History and the Smithsonian Marine
Station at Link Port, Ft. Pierce, for their assistance. Dis-
cussions with Dr. Diana Lipscomb were of great help in
fine-tuning the section on the cladistic analysis. Dr. An-
ders Waren kindly sent me some well-preserved material
of Ocenebra erinacea; Dr. James H. McLean and Mr.
C. Clifton Coney provided specimens of Trophon gev-
ersianiis. This is Contribution No. 333 of the Smithsonian
Marine Station at Link Port, Ft. Pierce, Florida.
LITERATURE CITED
Abbott, R T, 1974, American Seashells, 2nd edition. \'an
Nostrand Reinhold Company. New York, 663 pp.
Agersborg, H. P. K. 1929. Factors in the evolution of the
prosobranchiate mollusc Thais lapillus. The Nautilus 43(2):
45-49.
Ankel, W. E. 1937. Der feinere Bau ties Kokons der Purpur-
schnecke Nucella lapillus (L I und seine Bedeutung tiir das
Laichleben, V'erhandkingen der Deutsclien Zoologischen
Geselischaft Supplement 1077-86
Bandel, K. 1975. Das Embryonalgehiiuse mariner Prosobran-
chier der Region von Banyuls-sur-Mer I. Teil. Vie et Milieu
25(1 Kser. A):83-118.
Bandel, K. 1977. Neogastropod radulae from Ban\uls-sur-
Mer. Vie et Milieu, 27(2)(ser. A): 191-232
Bryan, G. W., Gibbs, P. E., Hummerstone, L. G,, and Burt, G
R. 1986 The decline of the gastropod Nucella lapillus
around South-West England: evidence for the effect of
tributyltin from antifouling paints Journal of the Marine
Biological Association of the L'nited Kingdom 66:61 1-640.
Carriker, M, R. 1981. Shell penetration and feeding by na-
ticaceaii and muricacean pretlatory gastropods: a synthe-
sis. Malacologia 20(2):403-422,
Colton, H S 1922. Variation in the dogwhelk Thais (Purpura
auct.) lapillus Ecology 3:146-157.
Gonnell, J H 1970. A predator-pre\ system in the marine
intertidai region. 1. lialanus glamlula and several pred-
atory species of Thais. Ecological Monographs 40:49-78
Cooke, A. H. 1915. The geographical distribution of Purjjura
lapillus (L.). Part I: in palaearctic waters. Proceedings of
the Malacological Society of London ll(4):192-209
Cossmann, A. E. M 1903. Essais de Paleoconchologie Coiii-
paree. Vol. 5. Paris, 215 pp.
Oothers, J. H. 1973. On variation in Nucella la])illus (L.):
shell shape in populations from I'cmbrokcshirc, Soulli
Wales. Proceedings of the Malacological Societv of London
40:318-327.
Crothers, J. H. 1983. \ ariation in dog-whelk shells in relation
to wave action and crab predation. Biological Journal of
the Linnean Society 2085- 102
Crothers, J. H. 1985. Dog-whelks: an introduction to the bi-
ology of Nucella lapillus Field Studies 6:291-360.
D'Asaro, C. N. 1991. GunnarThorsons world-wide collection
of prosobranch egg capsules: Muricidae. Ophelia ■351):1-
101.
Emien, J. M. 1966. Time, energy, and risk in two species of
carnivorous gastropod. Ph.D. dissertation. University of
Washington, Seattle, Washington.
Etter, R. J. 1987. The effect of wave action on the biology
of the intertidai snail Nucella lapillus PhD dissertation.
Harvard University, Cambridge, Massachusetts, i-ix + 198
PP
Feral. C 1976 Repartition geographique des femelles a trac-
tus genital male externe chez Ocenebra erinacea (L.), es-
pece gonochorique. Haliotis 7:29-30
Fretter V. and ,\. Graham. 1962. British Prosobranch Mol-
luscs. Ray Society, London, 755 pp
Fretter V. and .\ tJraham. 1985. The prosobranch molluscs
of Britain and Detmiark. Part 8 - Neogastropoda Journal
of Molluscan Studies, Supplement 15:435-556.
Gmelin, J. F. 1791. /n; Linnaeus, Svstema Naturae, Tom 1,
Pars \'l. pp. 3021-3909
Graham, A. 1941. The oesophagus of the stenoglossan pro-
sobranchs. Proceedings of the Ro\ al Societ\ of Edinburgh
61(l)(sec B):l-23.
Graham, A. 1955. Molluscan diets Proceedings of the Mal-
acological Society of London 31(3-4):144-159.
Gray, J. E. 1847. A list of the genera of Recent Mollusca,
their synonyma and t\ pes Proceedings of the Zoological
Society of London 15:129-219.
Gray, J E. 1853. On the division of ctenobranchous gaster-
opodous Mollusca into larger groups and families.The .■\n-
nals and Magazine of Natural History 11 (Ser,2):124-133.
Harasewych, M. G. 1984 C^omparati ve anatom\ of four prim-
itive muricacean gastropods: implications for trophonine
phylogeny. American Malacological Bulletin 3(1): 11-26.
Haw kins, L. E. 1985. Ecoph\siological studies of the Euro-
pean o\ster drill Ocenclna crii^acca (Linnaeus, 1758). Ph.D.
dissertation, L'niversit\ of Southampton, England, 244 pp.
Houart, R 1989 Description of a new species of Ocenebra
(Muricidae: Ocenebrinae) from western .\frica Publica-
9oes Ocasionais da Sociedade Portuguesa de Malacologia
No. 13, pp. 63-64.
Jeffreys, J. G. 1867. British Conchology, X'olume 4. \'an Voorst,
Paternoster Row, London, 486 pp., 8 pis.
Jousseaume, F. 1888. Descriptions des mollusques recueilles
par M. leDr Faurotdansla Mer Rouge etdu Golfed .\den.
Memoires de la Societe Zoologique de F" ranee 112-223.
Kilbnrn, H. and E. Rippe\. 1982. Sea Shells of Southern .Af-
rica South C:hina Printing Co , Hong Kong. 249 pp
Kincaid, T. 1957. Local races and clines in the marine gas-
tropod Thais lamellosa Gmelin. .^ population study. Cal-
liosloma Co., Seattle, Washington, 75 pp , 65 pis.
Kool, S I' 1986. Systematic revision of tliaulid genera based
on anatomy, .\merican Malacological Bulletin 4:110.
Kool, S P 1987 Significance ol raihilar characters in recon-
struction of thaidid ph>logeny (Neogastropoda:Murica-
cea). The Nautilu.s 101 l3l: 117-132.
Kool, S I' 1988. Aspects of the anatonn ol Vluopurpiira
S. p. Kool, 1993
Page 57
patula (Prosobranchia: Muricoidea: Thaidinae), new com-
bination, with emphasis on the reproductive s\steni.
Malacologia 29i2);37.3-382.
Kool, S. P. 1989, Ph\ logeneticanal>sis of the suhlamii\ Thai-
dinae (Prosobranchia: Neogastropoda; Muritidae) Ph D
Dissertation, The George Washington I niversit\, Wash-
ington, DC. 3-42 pp.
Kool, S. P. and K. J. Boss. 1992. Sucella K.kling, 1798 (CJas-
tropoda: Muricidae): tvpe species. The Nautilus 106(1):
21-23.
Kool, S. P. 1993. Phylogenetic analysis of the Rapaninae (Neo-
gastropoda: Muricidae). Malacologia (in press).
Lamy, E. 1928. La ponte chez les gasteropodes prdsohraiiches.
Journal de Conch) liologie 72(2)80-148
Largen, M. J. 1967. The diet of (lie dog-wliclk, Succlla la-
pillua (Gastropoda Prosobranchia), Journal ot Zoology 151:
123-127.
Lebour, M. \'. 1937. Tlie eggs and larvae ol the lirilish pro-
sobranchs with special reference to those living in the
plankton. Journal of the Marine Biological .'\ssociation of
the United Kingdom 22:105-166,
Linnaeus, C. 175S. Systema Naturae, 10th ed, Stockhohn, 824
PP
Malatesta, .•\ 1960 Malacofauna Pleistocenica di Grammi-
chele Memorie per Servire alia Descrizione della Carta
Geologica d Italia 121-392, 19 pis, v\ith letter press,
Melvill, C. and R, Standen. 1898. Notes on a collection of
marine shells from Livelv Island, Falklands, v\ith list of
species. Journal of Conchology 9(4):97-105.
Montfort, D. de. 1810. Conchy liologie Svstematique, et Clas-
sification Methodique des Coquilles. Part 2, Paris, 676 pp.
Moore, H, B, 1936. The biology of Purpura lapilliis. I, Shell
variation in relation to environment Journal of the Marine
Biological Association, of the United Kingdom 21:61-89,
Moore, H,'^B, 1938, The biology of Nucclla lapillm. Ill Life
history and relation to environmental factors. Journal of
the Marine Biological Association of the L'nited Kingdom
23:67-74,
Murdoch, W, W, 1969. Switching in general predators: ex-
periments on predator specificity and stability of prey
populations. Ecological Monographs 39:335-354,
Nordsieck, F 1982 Die Europaischen Meeres-Gehiiuse-
schnecken (Prosobranchia), Fischer Verlag, Stuttgart, 539
PP
Oehlmann, J , E. Stroben and P, Fioroni, 1988, Zur .Anatomie
und Histologic des Fortpflanzungssystems von Nucella la-
pillus (L., 1758) (Prosobranchia, Stenoglossa). Zoologische
Anzeiger 221(3/4):101-116.
Pallas, P. S. 1774. Spicilegia Zoologica, Vol 1, Part 10 Lange,
Berolini, 41 pp., 4 pis.
Palmer, R I 1983. Growth rate as a measure ot food value
in thaidid gastropods: assumptions and implications for
prev morphology and distribution. Journal of Experimen-
tal and Marine Biology and Ecology 7395-124
Palmer, R. I. 1985. .Adaptive value of shell variation in Thais
lamctlosa. effect of thick shells on vulnerabilitv to and
preference bv' crabs. The X'eliger 27(4):349-356,
Pelseneer, P, 1911, Recherches sur I embryologie des gastro-
podes, Memoires de I'.^cademie ro\ ale de Belgique, Classe
des sciences 3(2):1-167,
Rafinesque, C. S. 1815. Analyses de la nature ou tableau du
univers et des corps organises. Barravecchia, Palermo, pp.
5-6. 136-149, 218-223.
Risbec, J 1937. Les irregularites et les anomalies du devel-
oppement emlirv oiuiaire chez Murex e'rinacetis L. et chez
Purpura lapillus L. Bulletin du Laboratoire Maritime de
Dinard 17:25-38,
Robertson, R, 1974, Marine prosobranch gastropods. Larval
studies and svstematics, Thalassia Jugoslavica 10(1/2):213-
238.
Roding, P. F 1798 Museum Boltenianum. Part 1, Hamburg,
156 pp.; Part 2. Hamburg, i-viii + 199 pp.
Sabelli, B. and S. Tommasini. 1987. Osservazioni sulla radula
e sulla protoconca di Ocenchra crinaceus (L.,1758) e Had-
riana orctca (Vokes, 1964), Bolletino Malacologico 23(9-
10):275-283,
Schumacher, C, F, 1817, Essay dun Nouveau Systeme des
Habitations des Vers Testaces, Copenhagen, 287 pp,
Sowerby, G. B. 1841. Conchological Illustrations, Part
II. Murex. A catalogue of recent species 9 pp. -I- Index,
pp. 1-22.
Spight, T. M. 1972 Patternsof change in adjacent populations
of an intertidal snail, Thais lamcUosa. PhD dissertation,
Universitv of Washington, Seattle, Washington, 325 pp.
Spight, T. M. 1976, Colors and patterns of an intertidal snail,
Thais lamellosa. Researches on Population Ecology 17:
176-190
Spight, T M, 1982 Population sizes of two marine snails with
a changing food suppl> Journal of Experimental Marine
Biology and Ecology 57:195-217,
Thorson, G 1941. Marine Gastropoda Prosobranchiata The
Zoology of Iceland 4(60): 1-150.
Thorson, G 1946 Reproduction and larval development of
Danish Marine Bottom Invertebrates Meddelelser
fra Kommissionen for Danmarks Fiskeri- og Havunder-
sogelser Serie: Plankton 4:1-523
Wenz, W. 1941. Prosobranchia. In: Handbuch der Palao-
zoologie. Volume 6. OH. Schindewolf, ed., Gebriider
Borntraeger, Berlin, part 5, pp. 961-1200.
THE NAUTILUS 107(2):58-62, 1993
Page 58
Thala esperanza, a new Costellariidae (Mollusca: Gastropoda) from
northern Puerto Rico
Jose H. Leal'
Donald R. Moore
Rosenstiel School of Marine am!
Atmospheric Science
Uni\ersity ol Miami
4600 Rickenbacker Causeway
Miami, Florida 33149-1098 USA
ABSTRACT
Thala esperanza n sp. is described from Piaya Esperanza,
municipality of Manati, on the northern coast of Puerto Rico.
Small size, slender and delicate shell, smaller spire angle, larger
number of spiral cords, and a distincti\e constriction below the
suture separates the new taxon from western Atlantic conge-
neric species.
Key words: Thala. Costellariidae, Puerto Rico, Atlantic Ocean,
Caribbean Sea, new species, systematics.
INTRODUCTION
The costellariid genus Thala H. & A. Adams, 1853 com-
prises about 12 Recent species, of which two are known
from the western Atlantic. Thala fovcata (Sowerby, 1834)
is known to occur at St. Vincent, Antilles and possibK at
the island of Sao Thome in the eastern Atlantic (Maes &
Raeihle, 1975). Thala floridana (Dall, 1884) is found
along the coasts of Florida, northern Cuba, the Gulf of
Mexico, Haiti, and the Yucatan Peninsula. Cernohorsky
(1970) and Abbott (1974) s\non\ mized T. floridana with
T. foveata, before Maes and Raeihle (1975) demonstrat-
ed that they are separate species. These latter authors,
following preliminary observations of Raeihle ( 1968), have
shown in a detailed anatomical study of T. floridana that
Thala belongs in the Costellariidae (as V'exillidae), and
provided support for the then still controversial proposal
that this latter taxon deserved full familial status apart
from Mitridae (e.g., Azuma, 1965; Ponder, 1972) due to
differences in gross anatomy and early life history.
In this work we describe a third western Atlantic spe-
cies of Thala collected in beach sediments from Manati,
' Current address: Department of Invertebrate Zoology, NHB
stop 118, National Museum of Natural History, Smithsonian
Institution, Washington, DC 20560, USA.
Puerto Rico. Soft parts and radula are unknown; none-
theless, conchological characters are distinctive enough
to justify the naming of a new species.
MATERIALS AND METHODS
Shells were measured using WILD M-5 and M-8 dis-
secting microscopes with ocular micrometers and pho-
tographed with a Nikon F2 camera with extension bel-
lows and MicroNikkor 55 mm objective. Scanning electron
micrographs were made using an ISI Double Stage DS-
130 scanning electron microscope at the Electron Mi-
croscope Laboratory, Rosenstiel School of Marine and
Atmospheric Science. Model I ANOVA with unequal
sample sizes (Table 1) was performed according to Sokal
and Rohlf (1981). Acronyms used throughout this paper
are: ANSP, Academy of Natural Sciences of Philadelphia;
FMNH, Field Museum of Natural History, Chicago;
MCZ, Museum of Comparative Zoology, Har\ard Uni-
versity, Cambridge; MNHN, Museum National d'His-
toire Naturelle, Paris; UF, Florida Museum of Natural
History, University of Florida, Gainesville; UMML, In-
vertebrate Museum, University of Miami Rosenstiel
School of Marine and Atmospheric Science; L'SNM, Na-
tional Museum of Natural History, Smithsonian Insti-
tution, Washington.
SYSTEMATICS
Family Costellariidae MacDonald, 1860
Genus Thala H. & A. Adams, 1853
Thala esperanza new species
(figures 1-6, Table 1)
Description: Shell fusiform (length width ~ 2.7), im-
perforate, reaching 5.0 mm in length, 1.9 mm width.
Spire angle about 27° Shell surface smooth, translucent,
golden brown to chestnut brown, usually with single
spiral baiul of lighter color on middle of last w horl. Peri-
ostracmn undetectable. Protoconcli usually decollated in
adult shells, when present transparent brown, smooth,
with 2 whorls, about 0.4 mm diameter, embryonic whorl
J. H. Leal and D. R. Moore, 1993
Page 59
Table 1. Linear shell measurements (mm) and counts for Thala esperanza new species, holotype and paratypes 1, 2, 5-12, and
T. foveata. Statistics for model I .\NO\'A with unequal sample sizes according to Sokal and Rohlf (1981); NS = not significantly
different.
T c.s/x
■ranza (n =
11)
7", fovea la (n = 6)
C'haracter
Range
.\
SD
Range
.\
SD
ANOVA
Total length
3.92-500
440
0,40
5-25-6,33
5,73
0.40
F = 24.14
p < 0.001
Shell uidth
1.36-1.88
1,62
0,16
2 04-2 24
2 15
0.07
F = 23.02
p < 0001
Length last whorl
2.80-3.48
3,17
022
3,72-4,28
3,97
0.19
F = 56.34
p < 0.001
.\perture length
1.92-2,44
2 18
0,15
2,84-3,32
3,02
0.20
F = 81.54
p < 0.001
.\perture wiilth
0.36-0,(50
0 49
0 06
0,64-0,80
0 71
0.05
F = 65.12
p < 0.001
Teleoconch whorls
3,25-5,00
4,18
0,45
4,00-4,75
4,42
0.26
NS
Spire angle (degrees)
26-29
26,9
1 14
35-37
36.0
0.89
F = 213.9
p <s 0.001
Spiral cords last u horl
23-31
27.4
2,20
16-18
16,8
0.75
F = 128.4
p «: 0.001
.\\ia\ ribs last whorl
24-32
28.3
2,65
21-27
23,0
2.28
F = 23.41
p < 0.001
Length wklth
2 17-3 05
2,73
0 24
2,47-2,83
2.65
0.13
NS
.\perture length total Icn
gth
044-0,61
0,50
0,(.)5
0 51-0,55
0,52
0.02
NS
Length last uhorl, total length
0,65-0,87
072
0,06
0,66-0,71
0 69
0.02
NS
,\pertiire length 'aperturt
" width
3,78-4,58
4,27
0 24
3,55-4,87
4,26
0 51
NS ■•■
tilted, about 0.3 mm diameter. Protoconch/teleoconch
transition orthocline. Teleoconch reaching 5 convex
whorls. Suture impressed. Constriction present abapi-
cally at suture, defining a spiral cord wider than re-
maining spiral cords. Combination of spiral and axial
elements producing clathrate aspect to entire teleoconch.
Axial sculpture of 24-32 flattened ribs on last whorl.
Interspaces as w ide as axial ribs on early w horls, '/4 rib
width on last whorl. Spiral sculpture of 23-31 cordlets
on last whorl. Cordlets much narrower than axial ribs on
early whorls, equal in width to axial ribs on last whorl.
Five or six adilitional abapical spiral ribs at ba.se slightK
wider than those on remainder of last whorl. Aperture
elongate (length/width ~ 4.5). Outer lip with 9-13 small
but well defined denticles that are absent in immature
shells. Columella arched, with 4 columellar folds that
follow orientation of spiral cords along base. Parietal
region excavated, thus columellar plaits not raised above
shell surface at base. Anterior siphonal canal wide, weak-
ly defined. Posterior siphonal canal delineated only along
inner surface of aperture, without notch.
Holotype : USNM 860280, 5.00 mm length. 1.64 mm
width.
Type locality: Playa Esperanza, about 40 km west of
San Juan, Municipality of Manati. northern Puerto Rico.
Paratypes: Paratypes 1-4, USNM 860281; Paratypes 5-
6, UF 193382-193383; Paratvpe 7, ASNP 391938; Para-
type 8, AMNH 232313; Paratype 9, MCZ 302588; Para-
tvpe 10, MNHN. unnumbered; Paratvpe 11, FMNH
223388; Paratvpe 12, I'MML 8375; Paratypes 13-16,
USNM 860282; All from tvpe locality, D. Piferer col.
05/1973. beach drift.
Remarks: Protoconchs are almost always decollated in
beach-collected shells of this new species. Only two of
17 shells examined in this study had intact protoconchs,
and those clearly were immature specimens. We do not
know whether protoconch decollation occurs while snails
are still alive. Decollation occurs frequently in unrelated
gastropods (e.g., Cadiicifer atlanticus Coelho, Matthews
& Cardoso, 1970, a buccinid from off Brazil, see Leal,
1991:151). The protoconch is ver\ thin and translucent
in T. esperanza. Decollation probably also occurs in T.
foveata. In examining one of the specimens illustrated
by Maes and Raeihle (1975) (UF 158165, St. Vincent,
Lesser Antilles), we found that the protoconch of that
specimen, present in Maes is. Raeihle s illustration, is now
missing (figures 7, 8).
Of the western Atlantic species of Thala, T. esperanza
can be more easily separated from T. floridana. We do
not provide statistical comparisons between these two
species, but such comparisons may be made using data
in Table 1 and in Maes and Raihle (1975). Thala flori-
dana has a much thicker and larger shell, with nearly
Page 60
THE NAUTILUS, Vol. 107, No. 2
Figures 1-6. Thala esperanza new species. 1. Holotype. Playa Esperanza, N4anati, Puerto Rico (SKM) 2. Juvenile shell of paratype
4 (SEM). 3. Protoconch of paratype 4 (SEM). 4. Paratype 1. 5. Parietal region sfiowing columellar plaits on holotype (SEM. 6.
Shell sculpture near adapical part of aperture on holotvpe (SEM). .\11 from type locality. Figures 7-9. Thahi foveata. UF 1.58165,
J. H. Leal and D. R. Moore, 1993
Page 61
twice the average width of T. cspcranza. Shells of T.
floridana are usuall\ dark brow n. hut some can be white
(Maes & Raeihle, 1975) (figure 1 1 ). Shells are never white
in r. esperanza. The 40° spire angle of T. floridana is
broader than that of T. esperanza (27°). Thala floridana
has a much coarser sculptural pattern (figures 10, 11),
with about 13 spiral cords and 20 axial ribs on the last
whorl. These numbers a\erage 27.4 and 28.3, respec-
tively, in T. esperanza (figures 1, 4, Table I) (See Ap-
pendix 1 for a list of specimens of Thala floridana ex-
amined).
The degree of overall shell similarit\ between T. es-
peranza and T. foveata is greater. Table 1 shows mea-
surements, proportions, and meristic counts for adult
shells of the two species. All known specimens of T.
foveata are thicker and significaiitK larger (Table 1 ) than
any shell in the t\pe series of T. esperanza. The base in
T. foveata is alw a) s more tapered. Thala esperanza dif-
fers in having flattened axial ribs with narrower inter-
spaces, significanth larger spire angle, and greater num-
bers of spiral cords and axial ribs on last w horl. There is
also a marked, distinctive constriction defining a wider
spiral cord just below the suture in the new species (fig-
ures 1, 4, 7, 8) (See .\ppendix 1 for specimens of T.
foveata examined).
Thala esperanza can be distinguished from its eastern
and western Pacific congeners by its much smaller size
and differences in shell shape and sculpture. Thala gra-
tiosa (Reeve, 1845) from tropical west America has a
somewhat similar shell contour, but is twice as large and
has a much coarser sculpture with relatively fewer ele-
ments (Keen, 1971; Sphon, 1969). The western Pacific
T. milium (Reeve, 1845) and T. todilla (Mighels, 1845)
also differ by their larger size, coarser sculpture, purple
color in the former and four-w horled protoconch in the
latter (Cernohorskv, 1970; Kav, 1979).
ACKNOWLEDGEMENTS
We thank D. Princz for making a\ ailable all specimens
of the type series. Loans of material lor comparison w ere
provided by E.G. Thompson and K. Auffenberg (UF),
S.P. Kool (MCZ), G. Rosenberg (ANSP), W.K. Emerson
and WE. Sage (AMNH), and M.G. Harasewych (USNM).
We are also indebted to M.G. Harasew\ch and to an
anonymous reviewer for criticisms that helped improve
the manuscript. N.A. Voss and P.L. Blackwelder allowed
for use respectively of the facilities at the Invertebrate
Museum and Electron Microscopy Laborator\-, RSMAS,
University of Miami.
LITERATURE CITED
.\bbolt. R T 1974 .\iiieric-aii seashells. Second edition \'an
Nostrand-Reinhold, New York. 66o p
.\zunia. M. 1965. On the raduiae of the faniiK \ e\illidae.
\ enus 24:53-.57
(>rnohorsk\, \N'. O 1970. S\stematics of the families Mitri-
dae and N'oiutoniitridae (Moilusca: Gastropoda). Bulletin
of the .Auckland Institute and Museum 8: i-ii + 1-190.
Ka\, E. \. 1979. Hawaiian marine shells. Reef and shore
fauna of Hawaii Section 4: Moilusca. Bernice P. Bishop
Museum Special Publication 64(4), Honolulu, 653 p.
Keen. .\ M 1971 Seasliells of tropical West .\merica. Marine
mollusks from Lower California to Colombia. Stanford
Universit\' Press, Stanford, 626 p.
Leal, J. H. 1991. Marine prosoliranch gastropods from oceanic
islands off Brazil: species composition and biogeography.
Uni\ersal Book Services, Oegstgeest, 419 p.
Maes. \' O and D Raeihle. 1975. Systematics and biology
of Thala floridana (Gastropoda: N'exillidae). Malacologia
15(l):43-67.
Ponder, W. F 1972. The morpholog) of some mitriform
gastropods with special reference to their alimentar) and
reproductive systems, Malacologia 1 1(2):295-342.
Raeihle. D 1968 Notes on capti\e Cerithium variabile
C.B .\dams and Milra floridana Dall .\muial Report of
the .\merican Malacological L nion 1968:35-36.
Sokal, R. R. and F. J. Rohlf. 1981. Biometry: the principles
and practice of statistics in biological research. Second
edition. W.H. Freeman, San Francisco, 859 p.
Sphon, G. G. 1969. Notes on the the Mitridae of the eastern
Pacific, 2. The genus Thala, with the description of a new
species. The N'eliger 12(11:84-88.
APPENDIX 1
Material of Thala foveata examined, all from Calliaqua,
St. \incent. British N'irgin Islands: LP 158165, 1 shell,
beach drift, H.H. Monroe, 01 1961, ex-McGinty Collec-
tion; UF 145855, 2 shells, H.H. Monroe, 01/1961, ex-
McGint\ Collection; "rockv co\e on windward side of
SE Point, Calliaqua Bav", 13°07'20"\, 6n 1'55"W, ANSP
324791, 1 shell, V.O. Maes 02 1972; 13°07'04"N,
61°12'20"W, ANSP 354555, 2 shells, H.H. Monroe 1956,
Ex-Einlay Collection.
Material of Thala floridana examined: East Coast of
Florida: St. Augustine, L'F 158167, 3 shells, Ted Yocius
10/1972, 225 ft, Ex-J.M. Bijur Collection; AMNH 246013,
2 shells, Ted Yocius, Jerome M. Bijur Collection; Fort
Pierce, AMNH 127640, 16 shells, Thomas Hughes leg.;
MCZ 298862, 3 shells, Bernadine Baker coll., Ex-Doris
Folsom 5 1976; Palm Beach, Bovnton Beach, UF 145856,
1 shell, McGinty 8 1944; UF 1(19469, 41 shells, McGinty
Collection, 02/1944; Bo\nton Beach, Ocean Ridge, UF
St. Vincent, West Indies, 7. Shell (SEM), 8. Shell, M. Shell sculpture near adapical part of aperture (SEM), Figures 10-1 I. Thala
floridana 10. ISNM S602S4, Bear Gut, Miami, Florida (SEM), 1 1. I'SNM 860283, off Miami Beach, Florida, Scale bars = 1 mm.
except for Figures 3, 5, 6, 9 = 0,:
Page 62
THE NAUTILUS, Vol. 107, No. 2
158164, 1 shell, T. McGinty 8/1944; Card Sound. UF
63207, 2 shells, M. Smith 1/1935, I'liiversity of Alabama
leg.; UF 13150, 1 shell, C.B. Lungren 1909; 1.6 km off
Miami Beach, USNM 860283, 1 shell, M. Almasi and D.
Marszalek 10/1978, 24 m depth; Bear Cut, Miami, USNM
860284, A. Emery 02/1956, beach drift; Bird Key, Bis-
cayne Bay, INSM 414387, 3 shells, Eolis Sta. 13, 2-10
ft., Henderson Coll. 1910; Florida Keys: Key Largo,
AMNH 242626, 4 shells, ex-Alice Denison Barlow Col-
lection; Little Molasses Reef, off Kev Largo, UF145851,
McGintv 4/3/1950; Bonefish Key, UF 192101, 4 shells,
Beal-Maltbie coll., Ex-McGintv Collection; UF 146530,
1 shell, McGinty Collection; UF 63202, 8 shells, M. Smith
coll.. University of Alabama leg.; UF 162451, 3 shells,
Blenn R. Bales, G.L. Warmke leg. 1989; UF 135603, 3
shells, V. Orr, H.G. Lee leg.; MCZ 118844, 10-H shells,
B.R. Bales 5/7/1940; MCZ 100714, 8 shells, B.R. Bales
leg.; AMNH 116412, 2 shells, 1940, Dr. and Mrs. Julius
Wisoff Collection; AMNH 114121, 1 shell, T. McGinty
coll.; AMNH 138995A, 1 shell, AS. Koto leg.; AMNH
189020, 4 shells, AS. Koto leg. 1949, J. M. Gate Collection;
AMNH 138995, 4 shells, A.S. Koto leg., M.K. Jacobson
Collection; AMNH 199169, 9 shells, AS. Koto leg.; Lower
Matecumbe Kev, USNM 53477, 3 shells, H. Hemphill,
in grass; Kev Vaca, UF 156266, 9 shells, McGinty 2/1939;
MCZ 153269, 25+ shells, B.R. Bales; Key Vaca, Boot
Key Harbor, UF 145853, 7 shells, McGinty coll. 3/1945;
MCZ 226804, 2 shells, grassy bottom, 1 fm. McGinty
coll.; No Name Key, UF 63203, 2 shells. University of
Alabama leg.; Little Torch Key, UF 123072, 33 shells,
M.C. Teskey 1968-1977, on sand patches; Grassy Key
(Florida Bay), MCZ 110198, 4 shells, under rocks, P.
McGinty leg.; Pumpkin Key, UF 63205, 2 shells. M
Smith 1/1935, University of Alabama leg.; Islamorada.
AMNH 121444, 3 shells, ex-E.C. Styles; Big Pine Key,
UF 63204, 4 shells, M. Smith coll.. University of Alabama
leg.; Key West, AMNH 8982, 3 shells. Constable, Jay
Collection; USNM 53541, 1 shell, H. Hemphill, on rocks;
Barraconta Key, 9 miles W of Key West, UF 13149, 2
shells, G.W. Van Hyning, 6/11/1958; Middle Sambo
Shoals, UF 145852, 6 shells, McGinty 6/1946; Dry Tor-
tugas: MCZ 258469, 1 shell, J.S. Schwengel, dredged;
Loggerhead Key, UF 158168, 1 shell, T. McGinty 8/1941,
0.5 fm; Garden Key, South Coaling Dock, UF 13151, 2
shells, G.W. Van Hyning 7/16/1938, on piling; Gulf
Coast of Florida: St. Martins Reef, near Aripeka Bird
Racks, 6 mi. W of Aripeka, Pasco-Hernando Counts line,
MC;Z 233672, 14 shells, W.A. Smith 5/25/1963; Ozone,
Crystal Beach, AMNH 246038, 3 shells, Dan Steger coll.,
Jerome Bijur Collection; AMNH 125612, 10 shells, S.
Levine leg. 1960; AMNH 189024, 5 shells, S. Levine leg.
i960, J.M. Gate Collection; AMNH 125928. 76 shells, S.
Levine leg. 1960; AMNH 194560, 66 shells, Gordon New-
ell-Usticke Collection; Ozone, Crystal Beach, St. Joseph
Sound, UF 193036, 27 shells, Dan Steger; UF 137573, 5
shells, D. Steger 1970. H.G. Lee leg.; AMNH 189023, 3
shells, mud and grass bottom, E. Marcott leg. 1963, J.
Gate Collection; Marco, UF 137649, 1 shell, H. Hemphill,
2 fms, H.G. Lee leg.; USNM 53542, 1 shell, H. Hemphill;
15-35 mi. off Ft. Walton (FL), MCZ 145877, 5 shells,
13-19 fms., LA. Burry; Featherbend Bank, UF 63206,
2 shells, M. Smith 1/1935. University of Alabama leg;
Okaloosa County, off Destin, UF 158166, 1 shell, UF
145857. 2 shells, McGinty 10/1941, 14 fms; Bahamas:
Andros. first island off Mintie Bar, SE end of South Bight,
USNM 271832, 7 shells, P. Bartsch; Abaco, Little Harbor,
USNM 180492, 5 shells, O. Bryant; Grand Bahama Is-
land: Dead Man's Reef, 26°34'45"N, 78°51'45"W, ANSP
371266, 1 shell. J. Worsfold, Ex-J. Worsfold; McLean's
Town. 26°38'45"N,77°57'30"W. ANSP 369060. 8 shells.
J. Worsfold, E.\-J. Worsfold; Eight Mile Rock, Hepburn
Town, "Garbage Hole", 26°31'30"N, 78°47'15"W, ANSP
370392. 4 shells, J. Worsfold, Ex-J. Worsfold; Wood Cay,
26°44'15"N, 79°5S'15"W, ANSP 369615, 3 shells. J. Wors-
fold, Ex-J. Worsfold; West End, Settlement Point,
26°42' 15"N, 78°59'50" W, ANSP 368626, 6 shells, J. Wors-
fold, Ex-J. Worsfold; West End, Hotel Jettv, 26°42'15"N,
078°59'50"W, ANSP 368711. 21 shells, J. Worsfold, Ex-
J. Worsfold; Nassau, New Providence Island, L'F 145849,
1 shell, McGinty 6/6/1947, 4-6 fms, Sta. 17; Cuba: Las
Villas, Caibarien, Cayo Salinas, AMNH 138924, 1 shell,
M.K. Jacobson leg., 07/1947; Esperanza (NW Cuba),
USNM 414396. 5+ shells, Barrera Expedition, Sta. 210,
2-3 fms; Santa Rosa (NW Cuba), USNM 414498, 1 shell,
Barrera Expedition, Sta. 209, 3-6 fms; Bermuda. AMNH
45068, 2 shells. Constable. Jav Collection; MCZ 24174,
2 shells, O. Brvant 1903; IISNM 223283, 6 shells, Havcock
coll.; Haiti, Dept. de lOuest, Saltrois, USNM 439975, 1
shell, Orcutt, Chamberlain Coll.
THE NAUTILUS 107(2);63-75, 1993
Page 63
Shell and Pallet Morphology of Early Developmental Stages of
Bankia goiildi (Bartsch, 1908) (Bivalvia: Teredinidae)
Antonieto S. Tan
Ya-ping Hu
Michael Castagna'
Richard A. Luiz
Michael J. Kennish
Alan S. Pooley
Rutgers University
Institute of Marine and Coastal
Sciences
P.O. Box 231
New Brunswick, NJ 0S9U3-0231 USA
' Virginia Institute of Marine and
Coastal Sciences
College of William and Mary
Wachapreague, VA 23480 USA
ABSTRACT
The margin of the lar\al shell ot Bankia gouldi (Bartsch) un-
dergoes a progressive change m shape during ontogen\ from
an initial D-shape, to nearly circular, and then to elliptical just
prior to metamorphosis. The length of the iiinge is 37.9 ± 2 3
lim, n = 30 (range; 34.0-43.0 iim). The D-stage larval shell is
inequilateral with the anterior end broader than the posterior
end. The provinculum, composed of two cardinal teeth and
three sockets in the left valve, and three cardinal teeth and two
sockets in the right valve, is well developed in shells as small
as 75 nm. The length of the central tooth in the right valve
ranges from 12 to IQ/iim, The other teeth are one-half to one-
third of the length of the central tooth. A lateral hinge s\ stem,
consisting of two lateral flanges on the left valve that interlock
with two lateral grooves on the right valve, is present in spec-
imens as small as 75 nm. This hinge system persists until meta-
morphosis. Metamorphosis, first seen in the development of a
ligament pit, occurred in larvae with shells as small as 197 ^m
in height; and leads to the development of the chondrophore,
apophysis, anterior and posterior slopes, disc, umbonal-ventral
ridge, dorsal cond\ le, ventral cond) le, denticulated ridges and
pallets. As\nimetrical growth results in the formation of the
characteristic form of the teredinid postlar\ al shell with a neu
axis of articulation oriented dorsoventrally along the condyles.
Each pallet consists of a proximal stalk and a distal blade witli
one to several segments. Pallets with one segment were first
observed in early postlarval shells at 0 4 nmi in length.
Key iL'ords: larvae, postlarvae, shell morphology, Teridinidae,
metamorphosis.
INTRODUCTION
Morphological classification schemes used in the iden-
tification of bivalve larval stages have been developed
from light microscopic examination of plankton (j0r-
gensen, 1946; Rees, 1950) or cultured larvae (Loosanoff
et al., 1966; Chanley & Andrews, 1971), More recently.
the scanning electron microscope (SEM) has enabled
workers to describe in detail the hinge and other minute
structures of bivalve larvae and postlarvae that are useful
in species identification (Turner & Boyle, 1975; Carriker
& Palmer, 1979; Chanley & Dinamani, 1980; Le Pennec,
1980; Lutz et al., 1982a, b; Lutz, 1985; Prezant, 1990;
Waller, 1991). A method of consistently orienting and
documenting the shapes and dimensions of larval shells
(Lutz et al., 1982b; Fuller et al., 1989b) has provided
an additional means of differentiating bivalve species.
The present manuscript is part of an ongoing effort to
document the gross morphology, hinge apparatus and
other morphometric features of larval and early postlar-
val shells of bivalve mollusks to facilitate ta.xonomic iden-
tification of individual specimens to the species level
(Fuller & Lutz, 1989a; Fuller et al., 1989c; Kennedy et
al.. 1991; Goodsell et al., 1992; Gustafson & Lutz, 1992).
Bankia gouldi (Bartsch, 1908) is an oviparous mollus-
can woodborer foiuid in tropical and temperate waters
of the western Atlantic Ocean (Turner, 1971). As an
adult, B. gouldi has a soft, vermiform body, a diminutive
shell that covers the anterior-most portion of the body,
and segmented pallets situated at the base of the siphons
beneath a muscular collar (Turner, 1966, 1971). The
pallets are not formed until after metamorphosis and,
therefore, are of no value in identification of larval shells.
In addition, previous descriptions of the larval shell, early
postlarval shell and pallet of B. gouldi are insufficient to
allow congeneric differentiation. Hence, a means of iden-
tifying younger specimens of B. gouldi is clearly needed.
This work presents both qualitative and quantitative data
on the morphology of the larval shell, early postlarval
shell and pallet of 6. gouldi that can aid in species iden-
tification of early ontogenetic stages. In particular, we
show that the following features are important in distin-
guishing the larvae of B. gouldi from T. navalis: slope
of the shoulders, length of provinculum/hinge-line, and
length of posterior tooth ot the left provinculum.
Page 64
THE NAUTILUS, Vol. 107, No. 2
Figure I. Inl.rnal surface vie-w of disart.culaU-cl larval sheik Antcru.r end of the left valve is oriented tc»vards Uie ■^''^^^-h';^;
the anter.or ernl of the nght valve .s or.enled towards the left. The runnber above each valve s.gn.tes the maximum Imear shell
dimension in ^in
Tan et al.. 1993
Page 65
3
X
o
X
CO
240
SHELL LENGTH (fim)
Fif»iir«- 2. Linear regression oi shell height plotted ;i>;aiiist shell
length ol larvae.
Table I . Dimensions in fim of the larval shell of Bankia gouldi
shown in Figures 1 and 2 1 SD = one standard deviation
Sam-
ple
Range
Mean ± 1 SD
size
Left Valve
Anterior tooth length
7.5-9.4
8.3 ± 0.8
7
Posterior tooth length
5.5-7.1
6.4 ± 0.6
7
Provincular length
36.0-40.5
38.1 ± 1.8
9
Right Valve
Anterior tooth length
3.9-4.9
4.5 ± 0.4
8
Central tooth length
12.5-15.5
14.5 ± 1.2
7
Posterior tooth length
4.5-5.9
5.0 ± 0.5
9
Provincular length
34.0-40.5
37.8 ± 1-9
9
MATERIALS AND METHODS
Specimens of Bankia gouldi were collected from wooden
frames of trays set out for a year in the York River at
Gloucester Point, VA by Dr. Mary Gibbons. Adults were
mass spawned, and the fertilized eggs w ere grown to the
early adult stage using standard techniques (Loosanoff
& Davis, 1963; Castagna & Kraeuter, 1984). The tem-
perature of the culture water ranged from 22 to 30°C,
and salinity, from 32 to 34 ppt salinity. Floating blocks
of wood were provided as substrata for metamorphosing
larvae. Larvae and postlarvae were sampled regularly
and preserved in 95% ethanoi. The preparation of spec-
imens for SEM examination was described previously by
Fuller et al. (1989b). The specimens were coated with
about 600 A gold-palladium using a Polaron Coating L'nit
E5100, and they were examined using a Hitachi-450
SEM. Calibration of orthogonality of the SEM was done
with the aid of metallic spheres or glass microspheres.
Dimensions in the SEM were calibrated using a cali-
brated ocular micrometer, a glass microsphere, or a grid
of known dimension. Specimens were prepared, consis-
tently oriented, and photographed as previously de-
scribed to facilitate comparison of shapes and measure-
ments (Lutz et al., 1982b; Fuller et al, 1989b, c). Shell
terminology is described in Rees (1950), Turner (1966),
Chaniey and Dinamani (1980), and Fuller et al., (1989c).
The maximum linear dimension, shell length, shell
height, provincular length, tooth length and the lateral
hinge length were measured from caiibarated scanning
electron micrographs of oriented shells. The shell height
and length of the shell, hinge line and pallet were also
measured using a compound microscope with the same
calibrated ocular micrometer used to calibrate the scale
of the SEM. These terms are defined in Fuller et al.
(1989c).
Whenever possible, we compared our results quanti-
tatively with previously published data. Variables that
changed with growth were compared using analysis of
covariance, whereas variables that were independent of
size were compared using the Student t-test, Tukey, or
Student-Neuman-Keuls multiple range test (Zar, 1984).
The larval shells in Figures 1 and 3 are referred to by
their maximum linear shell dimension, which is the
greatest distance between two points on a contour of a
valve oriented in a standard fashion as described pre-
viously. The early postlarval shells in Figures 5-7 are
referred to by their shell height. Numbers separated by
a slash (e.g. 57/62) indicate left and right valve dimen-
sions respectively.
RESULTS
Larval shell: One da> after fertilization the shell length
was 61.1 ± 4.5 Mill, n = 30 (range: 54-70 /um); shell
height was 49.3 ± 4.5 ^ni, n = 30 (range: 43-59 ^m);
hinge line/provincular length was 37.9 ± 2.3 fim, n =
30 (range: 34.0-43.0 fxm). Most valves of the straight-
hinge larvae are inequilateral with the anterior end
broader and protruding farther beyond the hinge line
than the posterior end (Fig. 1, 57/62).
The lar\al shell undergoes gradual allometric changes
during ontogeny from a D-shape, to a nearly circular
shape, and finally to an elliptical shape just prior to meta-
morphosis (Fig. 1). During larval growth the maximum
shell dimension shifts from an anteroposterior to a dor-
soventral axis. Larval shells are almost ecjuidimensional
(length = height) at approximately 143 ;um (Figs. 1, 2).
The relationship between the length and height of the
larval shell is described by the equation, Y = —29 92 +
1.26X (r^ = 0.97, n = 150), where Y = height and X =
length in ^m (Fig. 2). A second power polynomial equa-
tion (quadratic equation) fits the data points only slightly
better than the linear regression equation.
The provinculuin consists of two cardinal teeth and
three sockets on the left valve that interlock with three
cardinal teeth and two sockets on the right valve (Figs.
1, 3). The onset of provincular development is variable,
occurring at about 57 ^.m. An individual at 57 /um can
be in the prodissoconch I stage without any trace of a
provinculum, or in the early prodissoconch II stage with
a developing provinculum (Fig. 3, 57 ^ni). Development
of the provincular teeth in the right valve starts with the
formation ol the posterior tooth, followed by formation
Page 66
THE NAUTILUS, Vol. 107, No. 2
Figure 3. Scanning electron micrographs showing hinges of lar\'ai shells seen in Figure 1.
Tan et al.. 1993
Page 67
Figure i. Dursomedial \ ieu of the larval ihflls. Orientation as in Figure 1. The lateral hinge system consists of two lateral flanges
on the left valve and two lateral grooves on the right valve on each end of the provinculum. Horizontal valve width: left = 198
nm, right = 185 ^m.
of the anterior tooth, and cuhninating with the formation
of the central tooth (Fig. 3, 62 urn, 75 ^m). The prov-
inculum is well-developed in valves measuring 75 nm.
The longest provincuiar tooth is the centra! one, \\ hich
can reach 19 nm. The other teeth are either one-half or
one-third of the length of the central tooth (Table 1).
The left anterior tooth is consistentlv larger than the left
posterior tooth, but the right anterior and right posterior
teeth are not consistent!) different in size (Figs 1, 3). The
increase in tooth length relative to shell length is variable:
the total provinculum increases more sIovnIv on the right
valve than on the left valve. Oni\ the length of the left
anterior tooth regressed against shell length during grow th
was significant (r- = 0.57, Table 2), others were not. The
provincuiar length measured from scanning electron mi-
crographs agrees with the hinge-line length values of
articulated valves measured bv a light microscope.
Lateral flanges and grooves are present in valves as
small as 75 ^m (Figs. 1, 3), and persist until metamor-
phosis. Two lateral flanges on the left valve (Fig. 1, 81-
214 ^m. Fig. 4) interlock with the two lateral grooves
on the right valve (Fig. 1, 75-201 ^m. Fig. 4). Each
lateral flange or lateral groove extends from the ends of
the provinculum for about 1/10-1/15 of the perimeter
along the dorsal commissure of the larval shell (Figs. 1,
4). The length of the anterior and posterior lateral grooves
are equal in shells up to 109 nm (F"ig. 1), but in larger
valves the anterior lateral groove increases in size reach-
ing a length up to 12% greater than that of the posterior
lateral groove (Figs. 1,4).
Early postlarval shell: The first morphological evidence
of metamorphosis is the development of a ligament pit
beneath the posterior tooth and socket in valves mea-
suring 235/230 jum in height (Figs. 5, 6). After meta-
morphosis, the growth of the postlarval shell is very asym-
metrical (Pigs. 5, 7, 8). The postlarval shell is slightly
inequivalve, with the right valve longer than the left
valve (Fig. 8). The ventral margin of the right valve is
narrower than that of the left valve. The denticles on
the denticulated ridge are predominantly two pronged,
with the dorsal prong larger than tiie \ entral prong. The
ventral margin grov\'s faster medially than the anterior
shell margin, while the growth of the posterior margin
lags behind (Figs. 5, 7, 8). The length to height ratio of
Table 2. Linear regressions of the shell height, tooth length
or pro\incular length plotted against shell length of Banhia
gouldi shown in Figure 1. a = intercept, b = slope,
r- = coefficient of determination, P = probabilitv of the re-
gression coefficient (slope) of each regression, N = sample size.
Dependent
Variable
a
b
r-
P
N
Left Valve
Shell height
-31.42
1.27
0.98
<0.01
9
.\nterior tooth
5.88
0 02
0.57
0 05
7
Posterior tooth
6.66
-0.002
0.02
0.76
7
Provinculum
37.32
0.01
0.03
0 65
9
Right Valve
Shell height
-27.57
1.22
0.98
<0.01
9
Anterior tooth
4.89
-0.003
0.14
0.35
8
Central tooth
12.71
0.01
0.18
0.35
/
Posterior tooth
5.36
-0.003
0.09
0.43
9
Provinculum
42,13
-0 04
0.71
<0.01
9
Page 68
THE NAUTILUS, Vol. 107, No. 2
Figure 5. Internal .siirfac- view ol early posllarval shells. Orientation as ni Figure I. The number above eaeh valve signifies the
shell height measured in nm roughlv perpendieular to the hinge region. The number on the lower left of the valve is the she^l
length in fim measured roughly parallel to the hinge region. ,'\ = apophysis. UC: = dorsal eondyle, VC = ventral condvie, R -
nmbonal-ventral ridge.
Tan et al., 1993
Page 69
Figure 6. Scanning electron micrographs of the hinge region of earl\ postlarval shells seen in Figure 5. The number above each
valve signifies shell height in ^m, LP = ligament pit, C = chontlrophore.
Page 70
THE NAUTILUS, Vol. 107, No. 2
Figure 7. Scanniiif; i'l«lroii iiiicrograplis ot l\w cxU-riial surface (il t-arl\ pusllarval shells, OrieiilatHni and number representation
a.s in I'^igure 5. AS = anterior slope, FS = Posterior slope.
Tan et al, 1993
Page 71
Figure 8. Views from different directions of partially articulated early postlarval sfiells with two denticulated ridges. The anterior
(top left) and the posterior (top right) views show that the right valve (R) is slightly longer than the left valve. The ventral view
at the bottom shows that the ventral margin of the right valve is narrower than that of the left valve. Most teeth have two prongs.
The dorsal prong of the tooth is larger than the ventral prong. Horizontal valve widths are as follows top left valve = 235 ixm. top
right valve = 243 nm, bottom left valve = 225 m"'. bottom right valve = 239 ^m.
the postlarval shell continues to decrease with growth
(Figs. 9).
The chondrophore that forms at the base of the pos-
terior tooth and socket grows anteroventrally then me-
chally, assuming a recurved formation (Figs. 5, 6). A
saber-like apophysis develops beneath the anterior tooth
and socket, and elongates along the sagittal plane in a
posteroventral direction (Figs. 5. 6 height 255/235 Mm)
The dorsal portion of the posterior slope and the anterior
slope are reflected up and outward (Figs. 5, G, height
305-1140/290-1140 Mm). The base of the apophysis and
the adjacent reflected part of the anterior slope and umbo
are incorporated to form the dorsal condyle (Figs. 5, 6,
height 305-1140/290-1 140Mm). The enlarged chondro-
phore and dorsal condyle eventually conceal the pro-
vinculum. As growth proceeds, the ventral portion of the
disc and the anterior slope curves ventromedially (Figs.
5, 7, 8). The shell thickening at the medial side of the
ventral margin eventually develops into the ventral con-
d\le (Fig. 5, height 255/235Mm). An umbonal-ventral
ridge, e.xtending froin the larval shell to the ventral con-
dyle, forms on the inner shell surface (Fig. 5, height 335-
11 40/705- 11 40Mm). The umbonal-ventral ridge is visible
as an umbonal-ventral sulcus on the external shell surface
(Fig. 7, 415-970/355-1075 Mm). The posterior slope (au-
ricle) overlaps the disc forming the shelf. The asym-
metrical growth of the shell eventually results in the
formation of the characteristic form of the teredinid
Page 7S
THE NAUTILUS, Vol. 107, No. 2
B
J.
X
o
Ed
a
to
2.5
2.0
1.5
1.0
0.5
0.0
30 r
Y = 0.15 + 1.00 • X
r2 = 0.84. n = 41
0.0 0.5 1.0 1.5 2.0
SHELL LENGTH (mm)
Figure 9. Rflation.sliip between sliell length and .shell height
of early postlarval shells.
w
o
Q
K
El,
o
K
Cd
m
S
D
SHELL LENGTH (mm)
Figure 10. Relationship between shell length and number of
denticulated ridges on the exterior surface of early postlarval
shells.
postlarva, with the a.\is of articulation changing from an
anteroposterior to a dorsoventral orientation along the
condyles.
Early in the formation of the postlarval shell, dentic-
ulated ridges are added onto the anterior margin of the
shell, initially only on the anterior area of the anterior
slope (Fig. 7, height 250-280/240-250 urn). As the shell
grows larger, the denticulated ridges extend to the an-
terior median area of the anterior slope. The number of
denticulated ridges increases significantly with the in-
crease in shell length; regression equation, Y = 5.38 +
10.7X, (r- = 0.54, n = 41), where Y = number of ridges
and X = shell length in millimeters (Fig. 10).
Pallet: A one segment pallet was present at a shell length
of 0.4 mm; at a shell length of 1.2-1.3 mm, the pallet
had 13 segments. The pallet consists of a proximal stalk
and a distal blade with one to several segments (Fig. 11),
with the basal pallet segment being the largest. The angle
formed by the base of the blade and the stalk is larger
on the dorsal than the ventral side. The medial inner
wall of the blade is flat, while the shorter outer wall of
the blade is convex. The plane circumscribed by the distal
outer margin of the blade is tilted anteriorly such that
the distal border of the inner margin extends further
posteriorly. Each pallet appears hemispherical uhen
viewed from the distal end. The left and right pallets in
an individual are morphologically similar and have the
same number of segments (Fig. 11, height 250-505 jum),
v\ hereas pallets with the same number of segments from
different individuals can vary in form (Fig. 11, height
840-1415 Mm).
DISCUSSION
Larval shell: Teredo navalis Linnaeus, 1758 is the only
other teredinid species whose larval and postlarval shell
morphology has been oriented and photographed in a
consistent manner (Fuller et al.. 1989c). In lar\al shells
of similar size, the shoulders of T. navalis (Fig. 1, Fuller
et al., 1989c) are considerably steeper (less rounded) than
those of Bankia gouldi (Fig. 1). The shoulders refer to
the dorsal shell margin on the anterior and posterior ends
of the hinge (C'hanley & Dinamani, 1980). This differ-
ence in larval shell shape can be used to distinguish the
two species. On the other hand, the size of the specimens
at which the length approximates the height measure-
Table 3. Linear regression equations for the shell length and shell height of D-stage to recent postmetamorphic larvae, r- = coefficient
of determination, P = probability for the slope of each regression, N = sample size. Slopes with different superscripts are significantly
different (P < 0.0.5) by a Tuke\ multiple range test
Species
Intercept
Slope
r-
P
N
.\uthor
Teredo natxilis'
-49.03
1.40'^
0.99
<0.01
10
Fuller ('/ al.
(I9S9c)
Bankia sctacea
-22.50
1 14"
LOO
<0.01
8
Turner & Johnson
(1971)
liankiu f^onldi
-.38.72
L.34^
1.00
<0.0J
6
Culliney (1975)
liankia gouldi
-36.13
1.37^
0.99
<0.01
7
Mann & Gallager
(1985)
Rankia gouldi'
-.32.. 56
1.28^
0.98
<0.01
10
This stud\-
Data used to calculate the linear regression equations are derived measurements of the left \ alve of scaiuiing electron micrographs.
Analysis of covariance of the slopes is significant (F-statistic = 6.32, P < 0.001).
Tan et al., 1993
Page 73
Figure 1 1. Outer surface view of the pallets. The pallet length is the distance between the proximal (stalk) and distal end (hiade).
Numbers signify the average length of the left and right (H) pallets. Dorsal side of the pallet is oriented towards the top, I = inner
wall, O = outer margin of outer wall.
Page 74
THE NAUTILUS, Vol. 107, No. 2
nients can not bo used to differentiate B. gaiildi from T.
nat^alis. The length and height of the larva! shell of B.
goiildi are nearly equal at about 143 ^im, far beyond the
point where the umbo is first visible in a shell in our
standard orientation, at about 128 ;um (Fig. 1). Cailliney
(1975) as well as Mann and Gallager (1985) reported that
B. gouldi is equidimensional at 128 /xni ^nd 71.4 ^m,
respectively. This difference (144 nm vs 128 or 71.4 /um)
is probably due to foreshortening (shells tilted) in the
earlier studies. Fuller et al. (1989c) reported that T
navalis achieves «iuidinieiisionalit\ at 150 /im. The ini-
tial size of a B. goiiUIi larva, is much smaller than that
of the larviparus T. navalis due to differences in their
development. Despite the difference in initial larval shell
size, both species metamorphose at a shell height of about
230 ^m, as indicated b\ the appearance of the ligament
pit.
Table 3 shows that shell height increases faster than
shell length until just before metamorphosis in Bankia
goiildi (Culliney, 1975; Mann & Gallager, 1985), Teredo
navalis (Fuller et al., 1989c), and Bankia setacea (Tryon,
1863) (Turner & Johnson, 1971). The slopes are signifi-
cantly different (P = <0.001) as shown by an analysis
of covariance. Moreover, a Tukey multiple comparison
test reveals that the slope for B. setacea is significantly
lower than the other four species (P = <0.05, Table 3).
The data for B. gouldi (this study) and T. navalis (Fuller
et al., 1989c) are from measurements of SEM micro-
graphs of disarticulated left valves, whereas the data
from other studies are based on light microscope mea-
surements of articulated valves (Culliney, 1975; Turner
& Johnson, 1971; Mann & Gallager, 1985). We have
included the data from shells of recent postmetamorphic
larvae since the ligament pit, an earl) indicator of meta-
morphosis, is difficult to detect by the previous studies.
The larvae of Bankia gouldi and Teredo navalis can
be distinguished on the basis of the length of the prov-
inculum on the left valve. The average length of the
provinculum/hinge-line measured from micrographs of
the left valves of B. gouldi (38. 1 ± 1.8^ m, n = 9; Table
1) and T. navalis (46.7 ± 1.3 ^m, n = 9; Fuller et. al.,
1989c) are significantly different (Student t-test, P <
0.001). Culliney (1975) reported a similar difference be-
tween the hinge-line length of B. goiddi (40.4 ± 3.96
Mm, n = 35) and T. tiavalis (51.3 ± 2.19 Mm, n = 47).
The same conclusion is reached using a simple size of 21
for B. gouldi. The provincular length measurements of
the left valve are more useful than those of the right
valve, because the latter varies significantly with growth.
The larvae of Bankia gouldi and 6. inartensi (Stem-
pell, 1899) can be distinguished from Teredo navalis In
the length of the posterior provincular tooth of the left
valve. The left posterior tooth of B. gouldi (6.4 ± 0.6
Mm, n = 7) and B. martensi (6.4 ± 1.0 Mm, n = 10) are
significantK shorter that that of T. navalis (8.2 ± 0.3
Mm, n = 9).
Clo.se examination of the internal shell surface of the
valves of Teredo navalis (see F'ig I; Fuller et al., 1989c)
reveals a lateral hinge system similar to that in Bankia
gouldi The longer anterior component of the lateral
hinge system in larvae of B. gouldi longer than 109 Mm
appears to be due to slightly faster growth of the anterior
shell margin relative to the posterior shell margin. This
difference may be useful in differentiating the two spe-
cies.
Postlarval shell: Larvae of Bankia goiddi and Teredo
navalis can be distinguished by the number of ridges vs.
shell length in shells greater than 1 mm in length. Both
species develop a denticulated ridge at about 0.24 mm
in length. Denticulated ridges are added at about the
same rate in both species until the shell reaches 1 mm.
Subsequently, the number of denticulated ridges in T.
navalis exceeds that in 6. gouldi. For example, at a shell
length of 2mm, B. gouldi has 13 ridges, compared to
about 28 ridges in T. navalis. In the early larval stage
ot B. gouldi, the right valve is longer than the left valve
(Fig. 8), but the degree of asymmetry is less than that
in T. navalis. The shape of the later, postlarval shell
varies greatly with the type of wood used as a substratum
by the teredinids (Turner, 1966; Turner & Johnson, 1971 ),
and the environmental conditions at the collection site
(Hill & Kofoid, 1927).
Pallet: The morphology of the adult pallet is important
in identification of teredinid species (Turner, 1966; 1971).
In Bankia setacea, the pallet first appears at an early
postlarval shell length of 0..5-0.6 mm (Quayle, 1959). In
B. gouldi, the smallest individual with a pallet had an
early postlarval shell length of 0.4 mm. The pallet at this
early developmental stage is unsegmented and may be
confused with the nonsegmented pallet of other teredi-
nids. Caution is advised in using pallet morphology as a
taxonomic tool at this developmental stage.
ACKNOWLEDGEMENTS
We thank Mr. John Grazul for technical ad\ice related
to SEM, and Drs. R. Prezant, R. Tool, D. Marelli and B.
Kotrla for insightful discussions on several aspects of B.
gouldi biology. This work is supported partly by a post-
iloctoral fellowship grant to A. S. Tan from the Fisheries
and Aquaculture Technology Extension Center, Rutgers
L'niversity. Institute of Marine & Coastal Sciences con-
tibution no. 93-14; New Jersey Agricultural Experiment
Station contribution no D-324()2-l-93; Virginia Institute
ot Marine Science contribution no. 1778.
LITERATURE CITED
( :arrikfr. M. \\ and R K. Palmer. 1979. llltrastructural nior-
plHigciicsi,s of prodis.socoiich and carl) dissocoiicli \ai\es
of the oyster Crassostrea virginica. Proceediniis of the
National Shellfisheries Association (•)9: 10.5- 128
Castagiia, M and J N. Kraeuter. 1984. Maiivial for growing
the hard clam Merccnaria. Special Report in .Applied Ma-
rine .Science and Ocean lOngineering No, 249 Virginia
lii.slitiitc of Marine Science, Gloucester, X'irgiiiia, 1 Id p.
(:liaiilc\, 1' and J I) .Viuirews, 1971. Aids for identificalion
of l>i\al\t' lar\;ieof \'jigliiia Mahn'ologia 11( 1 ):4.5-l 19.
Tan et al., 1993
Page 75
Clianley, P, ami P Dinaniaiii 19S() CoinparatiNc ilescrip-
tions of some oyster larvae from New Zealand and (IhiJe,
and a description of a new genus of o\ster, Tio.slrca. New
Zealand Journal of Marine and Freshwater Research 14(2):
103-120
Culline\ , J L. 1975 (Aimparati\e lar\al de\elopmeiit ol the
shipworins Banhia gouldi and Teredo nuialis Marine Bi-
ology 29:245-25 L
Fuller, S. C. and R. A. Lutz. 1989a. Shell morpholog) ot
larval and post-larval mytilids from the North-Western
Atlantic. Journal of the Marine Biological Association of
the United Kingdom 69:181-218.
Fuller. S. C, R. A. Lutz and A. Poole\. 1989h Procedures
for accurate documentation of shapes and dimensions of
larval bivalve shells with scanning electron niicroscopv.
Transactions of the American Microscopical Societ\' I()8( 1 ):
58-63.
Fuller, S. C, Y. Hu, R. A. Lutz and M. Castagna. 1989c. Shell
and pallet morpholog\ in early developmental stages of
Teredo navalis Linne (Bivalvia: Tereiiinidae). The Nau-
tilus 103(1 ):24-35.
Goodsell, J. G., S. C. Fuller, A. G. Eversole, M C^astagna ami
R .A Lutz. 1992. Larval and early post!ar\al shell mor-
pholog\ of se\eral \enerid clams Journal of the Marine
Biological .Association of the United kingdom 72231-255
Gustafson, R. G. and R. A. Lutz 1992. Larval and earl) post-
larval development of the protobranch bivalve Solemija
velum (Mollusca: Bivalvia). Journal of the Marine Biolog-
ical Association of the United Kingdom 72:383-402.
Hill, C. L. and C. A Kofoid (eds.) 1927 Marine borers and
their relation to marine construction on the Pacific coast
Final Report of the San F'rancisco Ba\ Marine Piling Gom-
mittee. Universits of Galifornia Press, San f'^rancisco, 357 p
j0rgensen, C. B. 1946. Reproduction and larval development
of Danish marine bottom invertebrates with special ref-
erence to the planktonic larvae in the Sound (0resund),
Lamellibranchia. Meddelelser fra Kommissionen for Dan-
marks Fiskeri-og Ha\unders0gelser, Serie: Plankton 4:277-
311.
Kennedy, V. S., S. G Fuller and R A Lutz 1991. Shell and
hinge de\elopment of \oung Corbicula flumiiica (Miiller)
(Bivalvia: Gorbiculoidea). .American Malacological Bul-
letin 8(1991):107-111.
Le Pennec, M. 1980. The larval and post-larval hinge of some
families of bivalve molluscs. Journal of the Marine Bio-
logical Association of the United Kingdom (iO:601-617.
Loosanoff, \'. L. and H. C. Davis. 1963. Rearing of bivalve
molluscs. In Russell, F.S. (ed.). Advances in Marine Bi-
ology. Academic Press, New York, 410 p
Loosanoff, V. L., H. C. Davis and P. C^hanlcN 1966 Dimen-
sions and shapes of larvae of some marine bivalve molluscs.
Malacologia(4):351-435.
Lutz, R. A. 1985. Identilic.ilion ol bi\aKc lar\ae and postlar-
vae: a review of recent ad\ances. .American Malacological
Bulletin, Special Edition No. l(1985):59-78.
Lutz, R. .\ . J. Goodsell, M (,'astagna. S CJhapman, (.'. Newell,
H Hidu. R Mann, D Jablonski. \' Kennedy, S Sidall, R,
Cioldberg, H Beattie. G. Falmagne, A Ghestnut and A.
Partridge 1982a. Preliminary observations on the use-
lulness of hinge structures for identification of bivalve
lar\ae Journal of Shellfish Research 2(l);65-70.
Lutz. R A , R Mann. J G Goodsell and M. Castagna. 1982b.
Larval and earl\ post-larval development of Arctica is-
landica. Journal of the Marine Biological Association of
the United Kingdom 62:745-769.
Mann, R. and S. M. Clallager 1985. Physiological and bio-
chemical energetics ol larvae Teredo navalis L. ami Ran-
kia gotddi (Bartsch) (Bivalvia: Teredinidae). Journal ot
Experimental Marine Biology and Ecology 85:211-228.
Prezant, R S. 1990 Shell and hinge ontogeny of the Antarctic
bivaKe Lissarea notoreudensis. Journal of the Marine Bi-
ological .Association of the L'nited Kingdom 70:841-855.
Qua\ le, D. B 1959. The early de\elopment of Bankia setacea
(Trvon) In Ra\. D.C:. (ed.). Marine Boring and Fouling
Organisms. Uni\ersitv of Washington Press, Seattle, p. 157-
174.
Rees. C' 11 1950 The identilit'ation and classification of la-
mellibranch larvae Hull Bnlletin of Marine Ecology .3(9):
7.3-104.
Turner, R D. 1966 Sur\e\ and illustrated catalogue of the
Teredinidae (Mollusca: Bivalvia). Museum of Gompara-
ti\e Zoology. Harvard University. Gambridge, 265 p.
Turner, R D, 1971, 1, Identification of marine wood-boring
molluscs. In Jones, E B (;. and S.K. Eltringham (eds.).
Marine Borers. Fungi and P"ouling Organisms of Wood.
Organisation for Economic and Go-operation and Devel-
opment, Paris, p. 17-65.
Turner, R. D. and A. G. Johnson 1971. 13. Identification of
marine wood-boring molluscs. In. Jones, E.B.G. and S.K.
Eltringham (eds. ). Marine Borers. Fungi and Fouling Or-
ganisms of Wood. Organisation for Economic and Co-
operation and Development, Paris, p. 259-301.
Turner, R D and P. J. Boyle. 1975 Studies of bisalve using
the scanning electron microscope and critical point drying.
Bulletin of the .American Malacological L'nion for 1974,
p. 59-65.
Waller, T R. 1991. Evolutionar\ relationships among com-
mercial scallops (Mollusca: Bivalvia: Pectinidae). In.
Shumway, S.E. (ed.). Scallops: Biology. Ecology and Aqua-
culture Elsevier, New York, p. 1-132
Zar, J.H. 1984 Biostatistical Analysis. Prentice-Hall, Englewood
Cliffs, New Jersev, 718 p.
THE NAUTILUS 107(2):76-78, 1993
Page 76
A Simple Method for the In Vitro Culture of Perkinsiis marinus
S.J. Kleinschuster
S.L. Swink
Haskiii Slifllfisli Kfst'arcli Lalioralory
Department of Marine and Cxiastal Sciences
Instilnte of Maine and (Coastal Sciences
Ruti;ers UniversiU
Box' B-8
1 Miller A\ciiuf
Port Norris, NJ 08349 USA
INTRODUCTION
The oyster pathogen Perkinsun marinus Levine, 1978 is
an apiconiplexan protist tliat has caused sul)stantial mor-
tality among oyster populations along tiie mid-Atlantic
and Gulf coasts of the United States and may be capable
of cross-transmission among certain bivalve species (Gog-
gin et al., 1989). The life c\cle ot the organism has been
described previously (Mackin & Boswell, 1956; Perkins,
1966; Perkins & Menzel, 1966). Although many stages
of the life cycle have been addressed, little evidence
relative to mode of infection, infiltration and invasion
has been described under in vitro conditions. Such in-
formation would provide insight into many hitherto
unobservable aspects of the host/parasite relationship in
vivo. This study demonstrates that certain vegetative re-
productive aspects in the life cycle of the organism, nor-
mally occurring in the host, can be duplicated under
relatively simple if! vitro conditions.
MATERIALS AND METHODS
Parasitic cells were isolated from primary cultures of
tissue explants of visceral ganglia of Crassostrea virginica
Gmelin, 1791 and identified as Pcrkinstis ntarinns using
the technicjue of Ray (1952). Eoilowing isolation of the
protist, various mixtures of Leibowitz's (1952) medium
(L-15), oyster hemolymph, fetal bovine .serum (FBS),
various sugars, yeast extract and lactalbumin hydrolysate
were tested for cell growth potential. The osmolaritv of
each licjuid component was adjusted to 750 mOs/kg by
the addition of sea salts. The pH of each medium and/
or constituent was adjusted to 7.6 and filter sterilized.
(Cultures were grown at 28°C; in rilasks utider ambient
CO2/O2. Penicillin G (100 U/ml) and streptomycin (0.1
mg/ml) were routinely added to all cultures and the
medium (50%) exchanged weekly.
Mantle tissue lor challenge experiments w as from I'er-
hinsits-hee o\sters which were routinely biopsed and
maintained in isolated aciuaria. Sterile explants were ob-
tained by culturing the tissue for 4 days at 25°C in a 1:1
sterile sea water/hemolymph mixture supplemented with
penicillin G (100 U/ml), streptomycin (0.1 mg/ml) and
amphotericin B (0.25 ;ug/ml).
RESULTS
Healthy explant cultures of visceral ganglia of Crassos-
trea virginica are seen in figures 1 and 2. Similar explant
cultures being parasitized by Perkinsiis marinus are seen
in figures 3 and 4. Colonization of large and small groups
of oNster nervous tissue is readiK apparent as is the at-
tachment of parasitic cells to individual oyster cells. Var-
ious media preparations to be evaluated for optimal
growth were inoculated with parasitic cells from similar
cultures. Approximately two weeks after inoculation, it
w as evident that all media preparations evaluated would
support cell growth to varying degrees. Cultures con-
taining a high percentage of hemolymph displayed vig-
orous and rapid propagation (figures 5 and 6). Cells cul-
tured with a high percentage of L-15 and/or FBS (50%)
tended to differentiate into cells that resemble prezoospo-
rangia (figure 5, double arrows) (i.e., large cells greater
than ca. 20 um with the cellular \ olume consisting mostly
of a vacuole anil the c\ toplasin occup\ ing the perimeter
of the cell; as described by Perkins and Menzel, 1966).
Many non-flagellated daughter cells were formed in those
cells xielding the morula-like cell aggregates typical of
Pcrkinsus marinus. L'pon rupture of the mother cell wall
the daughter cells are released into the mediiun (figures
5 and 6, arrows). Shortly after release of the daughter
cells, until enlargement to form prezoosporangium-like
cells, thr\ had the ts pical Perkinsus Hiori/n/.s-like meront
structure consisting of a prominent, eccentric vacuole
(figure 6, double arrows) with a single vacuoplast (figure
6, V arrow) and a single nucleus located near the cell
wall.
The most successlul medium [ireparation, yielding rel-
ativt'K large numbers oi cells as well as a diversity of
S. J. Kleinsclnister and S. L. Swink, 1993
Page 77
>-
H
^\v
P
fX
N
-// v^
/
/.
S
V
^-^.:.
->
N
Figures 1-2. Photomicrographs of health\ visceral ganjihon cxplaiits from Crassostrea virginica. Neurite growth extending from
explants indicated by arrows, scale bar = (I 1 rmn
Figures 3-4. Photomicrographs of oyster ganglion explants parasitized by Perkinsus marinus (P). Notice individual neurons (N),
support cells (S), hemocytes (H) and attachment of parasitic cells to individual oyster cells (arrows), scale bar = 0.05 mm.
life cycle stages consisted of a 100 mi. solution of L-15
containing 10.0 ml. of FBS, 20.0 ml. of o\ster hemo-
lymph, 5.0 mg. taurine, 50.0 mg. glucose, 30.0 mg. ga-
lactose, 50.0 mg. fructose, 50.0 mg trehalose, 100.0 mg.
yeast extract, 300.0 mg. lactalbumin Inclrolysate, 1.0 ml
MEM N'itamin solution (100X)(Sigma C^hemical C^o.) and
0.1 ml lipid nnxture (1000X)(Sigma Chemical C:o.). Cul-
tures were routinely sub-cultured over a period of several
months Sterile mantle tissue was challenged with cul-
tured Perkinsiis marinus and became infected within
2-3 weeks. In fluid thioglycollate medium the parasite
cells from challenged tissue formed cells resembling the
prezoosporangia (or "hypnospores") normally seen in
oyster tissue infected with Pcrkin.stts marinus. The cells
also stained like Pcrkiustis marinus prezoosporangia in
Lugol's iochne solution (Rax, 1952).
Page 78
THE NAUTILUS, Vol. 107, No. 2
rj!^U\
-tx.
■pT-^S
%^
:-v^(S
^" ''<'^*
''^^
•*t'
il^\ <^'.
6 v-'at
Figures 5-6. Photomicrographs of groups of propagative cells of Perkinsus marinus in vitro. Larger groups represent several
generations. Figure 5. Notice presporangial cells (double arrows) and non-flagellated daughter cells (single arrows). Figure 6.
Perkinsus-\\ke cells with nieront structure (double arrows), vacuoplast (V arrow) and non-flagellated daughter cells (single arrows),
scale bar = 0.05 mm.
DISCUSSION
It is hoped that the technique described herein will con-
tribute to the understanding of the basic biology of this
parasite as well as to the host/parasite relationship. Re-
finement of culture conditions and further definition of
nutritional rec}uirements of the parasite in vitro would
be particularly helpful as cellular or molecular approach-
es to this understanding are attempted.
ACKNOWLEDGMENTS
The authors wish to thank Dr. F O. Perkins for his
invaluable assistance with this study . This work was fund-
ed by the New Jersey Agriculture E.xperiment Station,
Hatch Project number 32100 and is identified as paper
no. D-.32100-1-93 and as paper no. 93-09 for the Institute
of Marine and Coastal Sciences.
LITERATURE CITED
Goggin, C. L., K.B Sewell and R T.G. Lester. 1989. Cross-
infection experiments with Australian Perkiitsus species.
Diseases of Aquatic Organisms 7(l):55-59.
Liebowitz, A. 1963. The growth and maintenance of tissue
cell cultures in free gas exchange with the atmosphere.
American Journal of Hygiene 78:173-183.
Mackin, J. G. and T. L. B Boswell. 1956. The life cycle and
relationships of Dermocijstidiutn marinum. Proceedings
of the National Shellfish Association 46:112-115.
Perkins, F. O. 1966. Life History Studies of Dermocystidium
marinutu, an Oyster Pathogen, Dissertation, Florida State
University, Tallahassee, FL.
Perkins, F. O. and R. W. Menzel. 1966. Morphological and
cultural studies of a motile stage in the life cycle of Der-
mocystidium marinum. Proceedings of the National Shell-
fish Association 56:23-30
Ray, S. M. 1952. A culture technicjue for the diagnosis of
infection with Dermocystidium marinum Mackin, Owen
and CA)llier in Ovsters. Science 116:360.
THE NAUTILUS 107(2):79-80, 1993
Page 79
On the Identification of Fossil Terrestrial Gastropod Eggshells
Harold G. Pierce
Research Associate
University of Nebraska State Museum
Lincoln, NE 68588-0514, USA
Small (~1 mm), calcareous spheroids, often olilate, are
frequently recovered associated with iionmarine mol-
lusks from Quaternary localities on the High Plains. In
general, these eggshells of terrestrial gastropods have been
ignored by modern paleontologists. Tompa (1976a) cor-
rectly correlated fossil eggshells from two middle Pleis-
tocene localities in Kansas with the genus Vallonia. Tom-
pa followed with a benchmark paper (Tompa, 1976b) in
which he illustrated the eggshells of many ta.xa and pro-
vided a catalog of terrestrial species with partially or
wholly calcified eggshells. Improved, and gentler, prep-
aration techniques have resulted in encountering both a
greater abundance and frequency of these gastropod
eggshells, especially in material of mid-Pleistocene to
Recent age. Recently, I recovered a substantial number
of much older (21-28 Ma) gastropod eggshells from the
Oligocene-Miocene Cabbage Patch fauna of southwest-
ern Montana. Since these eggshells were associated with
a prolific and diverse terrestrial gastropod fauna (Pierce,
1992), an attempt was made to identify the taxa that
produced these eggshells.
In almost all cases, microscopic examination by trans-
mitted light, and careful dissection of some specimens,
has shown the fossil eggshells to be empty. Correlation
of the eggshells with the various taxa in the fauna can
be accomplished by comparison of hatchling size and
shape to the size and shape of the eggshells. The two
types of Cabbage Patch eggshells were both oblate sphe-
roids that differed primarily in size (Table 1 ). The smaller
eggshell matched very closely, in all dimensions, mea-
surements of the hatchlings of Vallonia berryi Pierce,
1992, from the same fauna. The larger eggshell type was
found to be marginally large enough, but too oblate, to
accommodate hatchling of Oreohelix brandi Pierce, 1992,
another component of this fauna. This match was also
rejected on the basis that modern Oreohelix are ovivi-
parous. Hatchlings of another species, Polygyroidea
montivaga Pierce, 1992, were a near perfect match, both
in size and shape. No other taxa of this fauna had hatch-
lings that corresponded to the measurements of these
eggshells (Pierce, 1992).
Current work involving the Skull Creek locality in
Butler County, Nebraska (late Pleistocene: Wisconsinan:
Peoria loess, ca. 20 Ka), provided a new collection of
gastropod eggshells. The Skull Creek material contains
at least two ciuite diilerent eggshell forms (Table 1). The
first, an oblate spheroid, H/W = 0.68, was very thin and
delicate, and is represented by only 8 specimens, half of
which are broken. In size and H/W, it compared closely
with the eggshells attributed to Oligo-Miocene Vallonia
berryi, and to mid-Pleistocene to Recent Vallonia pul-
chella (Mijller, 1774) (Tompa, 1976a). Measurements of
hatchling gastropods recovered as part of the Skull Creek
locality fauna showed that hatchlings of Vallonia gra-
cilicosta Reinhardt 1883, averaging 0.7 mm x 0.55 mm,
were an excellent fit, in both size and shape, for this
eggshell (Figures 1-3). The second eggshell type is a near
spheroid, H/W = 0.93 or greater, and occurs in two
slightly overlapping size ranges, each with distinct modes.
The larger eggshell of this type is quite abundant (>50)
(Figures 4-6). This eggshell size and shape was found to
be an approximate fit for hatchlings of three taxa found
in this fauna, all of which are known to have calcified
Figures 1-3. Small fossil eggshells and hatcliling of Vallonia
gracilicosta Reinhardt, 1883. \. Eggshell with crack and small
fragment missing, lot PM.SC 1002 2. Hatchling \'. gracilicosta.
lot P\ AL 1002 3. Broken eggshell, lot PMSC 1002.
Figures 4-6. Large eggshell with embryo and hatchling ot
Discus uhitneyi (Newcomb, 1864). 4. Hatchling D. whitneyi.
lot PDIS 1004. 5. Large eggshell, lot PMSC 1001 6. Embryonic
D whitneyi still retaining portions of its eggshell, PDIS 1016
,\ll figures 20 X, all from Skull Creek locality, Butler County,
Nebraska.
Page 80
THE NAUTILUS, Vol. 107, No. 2
Table 1. Mea-suremt'iits (in mm) of fossil gastropod eggshells
Eggshells
\Vidtli/SD
Range
Height/SD
Range
H/W
Oligo-Miocene taxa
Cabbage Patch mollustan fauna
Vallonia berryi
Polygyroidea montivaga
Late Pleistocene ta.xa
Skull Creek molluscan famia
Vallonia gracilicosta
Discus ichitneyi
Discus whitneyi? small
10
0,76 ± 0.03
0.57 ± 0.02
0.75
0.72-0.80
0.55-0.60
1(1
L60 ± 0.03
L24 ± 0.04
0.78
L55-I.65
1.15-L.30
4
0.80 ± 0.00
0.55 ± 0.00
0.68
0.80-0.80
0.55-0.55
10
1.20 ± 0.03
1.13 ± 0.01
0.94
1.15-1.25
1.12-1.15
5
1.03 ± 0.05
0,96 ± 0,04
0.93
0 95-1 10
0 90-1 00
eggshells (Tompa, 1976b, table 1): Nesovitrea hammonis
electrina (Gould 1841), averaging 1.15 mm x 0.75 mm,
Zonitoides arboreus (Say 1816), averaging 1.1 mm x 0.8
mm, and Discus whitneyi (Newcomb 1864), averaging
1.1 mm X 0.8 mm, with the latter two providing the
best fit. [Roth (1987:129) has shown D. whitneyi to be a
senior synonym of the well known D. cronkhitei (New-
comb, 1865)]. By chance, one eggshell contained a em-
bryo, identifiable as D. whitneiji (Figure 6). Roth V.
gracilicosta and D. whitneyi are very abundant (>200
specimens each) components of the Skull Creek mollus-
can fauna. Zonitoides arboreus and N. hammonis elec-
trina are less common (16-50 specimens each) compo-
nents. A second, smaller set (N=5) of the nearly spheroidal
eggshell type, microscopically indistinguishable from the
larger size, could not be matched precisely with any other
taxon, but would fit the extreme minimum hatchling size
of both D. whitneyi and N. hammonis electrina. Tompa
(1976b:867) has shown that, for some taxa, eggshell size
varies from individual to individual, and/or with size of
parent animal. It appears probable, therefore, that these
also represent D. whitneyi eggshells, but from small par-
ents.
Identification of eggshell types provides yet another
tool useful in reconstructing fossil faunas. As more egg-
shells are identified to genus and/or species, it may be-
come possible that additional taxa, represented only by
eggshells in a fauna, can be identified, thereby increasing
the recognized diversity of the fossil fauna and the ac-
curacy of paleoenvironmental reconstruction. However,
with rare exceptions rc()uiring SEM examination (Tompa
1976b), identification beyond generic level is, at this time,
often tenuous.
LITERATURE CITED
Gould A. A. 1841, Report on the Invertebrata of Massachu-
setts, comprising the Mollusca, Oustacea, Annelida, and
Radiata. Foisom, Wells, and Thurston, Cambridge. 373
p,, 213 figs,
Miiller, O. F. 1774. Vermium terrestrium et Duviatilum, seu
animalium Infusorium, Heiminthicorum et Testaceorum
non marinorum succinta historia. Volumen .Alterum Hei-
neck et Faber, Haviniae et Lipsiae. 214 p.
New comb, W 1864 Descriptions of nine new species of Helix
inhabiting California Proceedings of the California .Acad-
emy of Sciences (1) 3:1 1.5-119.
Newcomb, W. 1865. Descriptions of new species of land
shells. Proceedings of the California Academy of Sciences
(1)3:179-182.
Reinliardt, O 1883. Sitzungsberichte der Gesellschaft Na-
lurforschender Fruende, Berlin, 1883(2):42
Holh, B. 1987, Identifications of two (;alifornian land mollusks
tlescribed bv VVeslev Newcomb Malacological Review 20:
129-1.32,
Pierce, H, G, 1992. The nonmarine mollusks of the late Oli-
gocene-early Miocene Cabbage Patch Fauna of western
Montana. II. Terrestrial gastropod families other than Pup-
illidae (Pulmonata: Stylommatophora). Journal of Pale-
ontology 66(4):618-62S.
■Say, T. 1816. Conchology, Nicholson s British Encyclopedia
of Arts and Sciences, 1st (American) ed.
unpaged.
Tompa, A, 1976a Fossil eggs of the land snail genus Vallonia
(Pulmonata: Valloniidae). The Nautilus 90(l):5-7.
Tompa, A. 1976b. A comparative study of the ultrastructure
and mineralogy of calcified land snail eggs (Pulmonata:
Stylommatophora), Journal of Morphology 1.5()(4):861-888.
INSTRUCTIONS TO AUTHORS
THE NAUTILUS publishes papers on all aspects of the
biology and systematics of mollusks. Manuscripts de-
scribing original, unpublished research as well as review
articles will be considered. Brief articles, not exceeding
1000 words, will be published as notes and do not re-
quire an abstract. Notices of meetings and other items
of interest to malacologists will appear in a news and
notices section.
Manuscripts: Each original manuscript and accompa-
nying illustrations should be submitted in triplicate. Text
must be typed on one side of 8'/2 x 1 1 inch white paper,
double spaced throughout (including literature cited, ta-
bles and figure captions), with at least 1 inch of margin
on all sides. All pages must be numbered consecutively.
If printed on a word processor, the right margin should
be ragged rather than justified. Authors should follow
the recommendations of the Council of Biology Editors
Style Manual, which is available from the Council of
Biology Editors, Inc., 9650 Rockville Pike, Bethesda, MD
20814, U.S.A. The first mention of a scientific name in
the text should be accompanied by the taxonomic au-
thority, including year. Latin names and words to be
printed in italics must be underlined; leave other indi-
cations to the editor. Metric and Celsius units are to be
used.
The sequence of sections should be: title page, abstract
page, introduction, materials and methods, results, dis-
cussion, acknowledgements, literature cited, tables, fig-
ure captions, figures. The title page should include the
title, author's name(s) and address(es). The abstract page
should contain the title and abstract, which should sum-
marize in 250 words or less the scope, main results and
conclusions of the paper. The abstract may be followed
by a maximum of 8 key words. All references cited in
the text must appear in the literature cited section and
vice versa. In the literature cited section, all authors
must be fully identified and listed alphabetically. Follow
a recent issue of THE NAUTILUS for bibliographic style,
noting that journal titles must be unabbreviated. Infor-
mation on plates and figures should be cited only if not
included in the pagination. Tables must be numbered
and each placed on a separate sheet. A brief legend must
accompany each table. Captions for each group of illus-
trations should be typed on a separate sheet and include
a key to all lettered labeling appearing in that group of
illustrations.
All line drawings must be in black, high quality ink,
clearly detailed and completely labeled. Photographs
must be on glossy, high contrast paper. All figures are
to be consecutively numbered (figs. 1, 2, 3, ... , NOT
figs. la. lb, Ic, . . . NOR plate 1, fig. 1 . . .). Illustrations
must be arranged in proportions that will conform with
the width of a page (6% inches or 171 mm) or a column
(3V4 inches or 82 mm). The maximum size of a printed
figure is 6% by 9 inches or 171 by 228 mm. All illus-
trations must be fully cropped, mounted on a firm, white
backing, numbered, labeled and camera ready. The au-
thor's name, paper title and figure number(s) should ap-
pear on the back. Original illustrations must be between
one and two times the desired final size. It is the author's
responsibility that the line weight and lettering are ap-
propriate for the desired reduction. Original illustrations
will be returned to the author if requested. Color illus-
trations can be included at extra cost to the author.
Voucher Material: Deposition of type material in a
recognized public museum is a requirement for publi-
cation of papers in which new species are described.
Deposition of representative voucher specimens in such
institutions is strongly encouraged for all other types of
research papers.
Processing of Manuscripts: Upon receipt, every manu-
script is acknowledged and sent for critical review by at
least two referees. These reviews serve as the basis for
acceptance or rejection. Accepted manuscripts are re-
turned to the author for consideration of the reviewers'
comments. A finalized version of the manuscript is re-
turned to the editor and sent to press. Two sets of proofs
are sent to the author for correction. Changes other than
typesetting errors will be charged to the author at cost.
One set of corrected proofs should be sent to the editor
as soon as possible. Authors with institutional, grant or
other research support will be billed for page charges at
the rate of $60.00 per printed page.
An order form for reprints will accompany the proofs.
Reprints may be ordered through the editor.
Manuscripts, corrected proofs and correspondence re-
garding editorial matters should be sent to: Dr. M.G.
Harasewych, Editor, Division of Mollusks, NHB stop 118,
National Museum of Natural History, Smithsonian In-
stitution, Washington, DC 20560, USA.
THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.
rHE NAUTILUS
Volume 107, Number 3
February 2, 1994
ISSN 0028-1344
A quarterly devoted
to malacology.
FFR 1 A. iqP4
VUr.'-rv Hol6 M^,
EDITOR-IN-CHIEF
Dr. M. G. Harasewych
Division of Mollusks
National Museum of
Natural History
Smithsonian Institution
Washington, DC 20560
ASSOCIATE EDITOR
Dr. R. Tucker Abbott
.American Malacologists, Inc.
P.O. Box 2255
Melbourne, FL 32902
CONSULTING EDITORS
Dr. Riidiger Bieler
Department of Invertebrates
Field Museum of
Natural History
Chicago, IL 60605
Dr. Robert T. Dillon, Jr.
Department of Biology
College of Charleston
Charleston, SC 29424
Dr. William K. Emerson
Department of Living Invertebrates
The American Museum of Natural
History
New York, NY 10024
Dr. Robert Hershler
Division of Mollusks
National Museum of
Natural History
Smithsonian Institution
Washington, DC 20560
Dr. Richard S. Houbrick
Division of Mollusks
National Museum of
Natural History
Smithsonian Institution
Washington, DC 20560
Mr. Richard I. Johnson
Department of Mollusks
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138
Dr. Aurele La Rocque
Department of Geology
The Ohio State University
Columbus, OH 43210
Dr. James H. McLean
Department of Malacology
Los Angeles County Museum of
Natural History
900 Exposition Boulevard
Los Angeles, CA 90007
Dr. Arthur S. Merrill
% Department of Mollusks
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138
Ms. Paula M. Mikkelsen
Harbor Branch Oceanographic
Institution, Inc.
Ft. Pierce, FL 33450
Dr. Donald R. Moore
Division of Marine Geology
and Geophysics
Rosenstiel School of Marine and
Atmospheric Science
University of Miami
4600 Rickenbacker Causeway
Miami, FL 33149
Dr. Gustav Paulay
Marine Laboratory
University of Guam
Mangilao, Guam 96923
Mr. Richard E. Petit
P.O. Box 30
North Myrtle Beach, SC 29582
Dr. Edward J. Petuch
Department of Geology
Florida Atlantic University
Boca Raton, FL 33431
Dr. David H. Stansbery
Museum of Zoology
The Ohio State University
Columbus, OH 43210
Dr. Ruth D. Turner
Department of Mollusks
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138
Dr. Geerat J. Vermeij
Department of Geology
University of California at Davis
Davis, CA 95616
SUBSCRIPTION INFORMATION
The subscription rate per volume
is US $25.00 for individuals, US
$40.00 for institutions. Postage
outside the United States is an
additional US $5.00 for surface
and US $12.00 for air mail. All
orders should be accompanied by
payment and sent to: THE
NAUTILUS, P.O. Box 7279, Silver
Spring, MD 20907-7279, USA.
Change of address: Please inform
the publisher of your new address
at least 6 weeks in advance. All
communications should include
both old and new addresses (with
zip codes) and state the effective
date.
THE NAUTILUS (ISSN 0028-
1344) is published quarterly by
Trophon Corporation, 8911 Alton
Parkway, Silver Spring, MD 20910.
Second Class postage paid at Silver
Spring, MD and additional mailing
offices.
POSTMASTER: Send address
changes to: THE NAUTILUS
P.O. Box 7279
Silver Spring, MD 20907-7279
T H Ef7N AU T I L U S
CONTENTS
Volume 107, Number 3
February 2, 1994
ISSN 0028-1344
- ' -1 Jii
uorar/
Two New Tropical Western Atlantic Species of
Epitonium. with Notes on Similar Global Species and
Natural History 81
Wooft<! Hole. MA 02
Phalium (Semicassis) vector, a New Deep- Water Species
from the Central Indian Ocean 94
Ferussac and d Orbigny s " Histoire naturelle generate et
particuliere des Cephalopodes acetabidiferes". Dates of
Publication of Plates and Text 97
Alvania valeriae (Gastropoda: Rissoidae), a New Species
from off Southeastern Brazil 104
Robert Robertson
R. Tucker Abbot!
Annie Tiilier
Renata Boucher-Rodoni
Ricardo Silva Absalao
THE NAUTILUS 107(3):81-93, 1993
Page 81
Two New Tropical Western Atlantic Species of Epitonium, with
Notes on Similar Global Species and Natural History
Roberl Robertson
UepartmeiU of Malacology
The Acadenu ot Natural Sciences
1900 Benjamin Franklin Parkway
Philadelphia, PA 19103-1195, U.S.A.
ABSTRACT
Two large (up to 16.9 and 25, .'5 mm long), main -ribbed, shallow
water species of Epitonium are newly described; £, phymanlhi
from southeast Florida, the 15ahamas, and the West Indies, and
£. worsfoldi from the Bahamas and the West Indies. Where
possible, rib counts were made on each shell whorl to allow for
ontogenetic changes. Epitonium phijnmntlii lives with and feeds
on the sea anemone Phijmanthus crucifcr in Florida (where
E. lamellosum and another epitoniid Opalia crenata also live
with this anemone). Likewise, £. worsfoldi lives with and feeds
on a sand-d\\ elling anemone that is either Actinoporus clcgans
or Homostichanthus duerdeni in the Bahamas. There it is more
commonly parasitic on another anemone, Stichodactyla he-
lianlhus. .'\quarium observations on £. phymanlhi with £.
lamellosum revealed some major biological differences be-
tween these co-occurring species, e.g. fully everted proboscis
lengths and speed of movement. Epitonium phynianthi was
found with £ ivorsfoldi at only one localit> , as shells in the
Virgin Islands.
Key words: Epitoniidae. new species. Western .Atlantic
INTRODUCTION
The family Epitoniidae contains a large number of de-
scribed fossil and living species, and as with most groups
of mollusks there is as yet no world monograph. In the
interim, it could be argued that there should be a mor-
atorium on describing any new epitoniids. The literature
is very scattered and unsynthesized. There is not even a
published list of names. Nevertheless, it is here believed
that minimum requirements are met for describing two
new species, albeit v\ ithout anatomy.
This paper is a by-product of work on a book on the
marine prosobranchs of the Bahamas being prepared by
myself. Jack N. Worsfold, and Colin Redfern. New spe-
cies are not being described therein.
The conchological systematics of Recent Epitoniidae
(wentletraps) in the western Atlantic and eastern Pacific
(where cognate forms, subspecies or species can be ex-
pected) are relatively well know n thanks to Clench and
Turner (1950b, 1951, 1952, 1953) and DuShane (1974,
1979). Little systematic work has since been done on the
western Atlantic species. Even so, it was surprising to
conclude that two large, many-ribbed, shallow-water Ep-
itonium species had remained undescribed until now.
One ranges from southeastern Florida and the Bahamas
to the Lesser Antilles (Grenada). The other ranges from
the Bahamas to Cuba, Puerto Rico and the Virgin Islands.
Both may well have more extended ranges. As recently
as 1967, one of the species was first collected alive; it was
in less than 1 m of water within sight of a major marine
laboratory!
Clench and Turner had neither of these species avail-
able in the collections they studied [although a small,
badly broken shell of one of them was mi.xed in a vial
with one £. albidum (Orbigny, 1842) (MCZ 107820).
Some epitoniid species seem possibly or probabK to
be circumglobal (Robertson and Habe, 1965; Kilburn,
1972, 1985; DuShane, 1983; Paschall, 1987). Therefore,
an attempt has been made to consider the two new spe-
cies in a global context. Based on epitoniid collections at
MCZ, AMNH, ANSP and USNM and literature, shells
of 23 species from around the world (.\ppendi.x) were
assessed to be most similar to the shells described here.
I have tried to survey all the descriptions and illustra-
tions of Miocene to Recent species of Epitonium, sensu
lata, known from the Americas, and of most Recent
species from here and the rest of the world, especially
Europe and the Mediterranean. These are the fossils and
living animals most likely to be similar or identical to
the shells discussed here. European fossil epitoniids were
not surveyed because the literature is so extensive. Among
the most important references consulted on Epitonium,
sensu lata are:
1) world Recent species: Kiener (1838-1839); Sowerbv
(1844); Nyst (1871); Sowerby in Reeve (1873-1874);
Tryon (1887); Clessin (1896-1897); de Boury (1912-1913);
Kaicher (1980, 1981, 1983, the only really global con-
tribution in this century, albeit incomplete).
2) American Recent species: Morch (1875a, 1875b, 1876);
Dall (1889); Clench and Turner (1950a-1953); Nowell-
Page 82
THE NAUTILUS, Vol. 107, No. 3
Figures 1-4. Epitoniuiu phymaiithi liobt-rtsoii, new species. 1. .Apertural, 2. apical, and 3. basal \ie«sof liolot\pe. Shell 16.9
mm long, and 9.3 mm wide. Miami, Florida, ANSP 391939. 4. Unusually low-spired shell (not a paratype), 12,0 mm long, and
8,f) mm wide.
Usticke (1959); Keen (1971); Abbott (1974); DuShane
(1974, 1979, and references therein); Gundaker (1975);
de Jong and Coomans (1988); Espinosa and Fernandez
Garces (1990).
3) American late Cenozoic fossil species: Olsson (1916);
Maury (1910, 1917); Gardner and Aldrich (1919); Pilsbry
(1922); Woodring (1928, 1959); Mansfield (1930, 1935);
Pilsbry and Olsson (1941); Gardner (1947, 1948); Olsson
and Harbison (1953); Weisbord (1962); Jung (1969).
4) European Recent species: Parenzan (1970); Franchini
(1975-1976); Nordsieck (1982); Bouchet and War6n
(1986); Poppe and Goto (1971).
5) Indo-Pacific, Japanese, Australasian and South African
Recent species: Adams ( 1 861); McK ill and Standen (1903);
Jousseaunie (1911); Ircdale (1936); Kerslake (1958); Azu-
ma (1962); Macpliorson and Gabriel ( 1962); Kuroda, Habe
and Oyama (1971); Masahito and Habe (1973-1976);
Powell (1979); Kilburn (1985); DuShane (1988, 1990);
Nakayama (1991).
An attempt was made to count the ribs on each whorl
of each shell The need ior doing so was explained in a
previous paper (Robertson, 1983a). Counts on juveniles
can be different from those on adults. Indeed, a rib count
per whorl in species of Epitoniiim may increase, sta\'
the same, decrease, or decrease and then increase as the
shell grows in a species-specific maimer. These ta.\onomic
characters have hardly been used before.
The major study by Clench and Turner (1950b-1953)
was published before epitoniids were found to live with
or to forage for and feetl on benthic coelenterates, either
as parasites or predators (Thorson, 1957; Robert.son, 1963,
1981, 1983b; DuShane, 1988; Yamashiro, 1990; Nakay-
ama, 1991, etc.). Both species treated here have actinian
(sea anemone) hosts.
MATERIALS AND METHODS
The specimens studied were assembled during almost
two and a halt decades The two shallow -water species
are curiously uncommon, and I personally saw neither
of them alive. Only empty shells are available of E.
pliyiiumthi. but there are a few £. ivorsfoUli in alcohol
(see .ANSP "A" numbers in locality records).
Shells were studied with the aid of a Wild dissecting
R. Robertson, 1993
Page 83
Figures 5-6. Epitunium phymanllii. liicdnipletely grown
parat\ pe shells. 5. Only shell known from Bahamas, juvenile.
5 6 mm long, 3.1 mm wide. 6. Subaclull topotvpe. 10 8 mm
long, 6.3 mm wide.
microscope and camera lucida. First- whorl diameters
were measured and whorls counted as advocated by Rob-
ertson (1985). The camera kicida and a protractor were
used to measure spire angles to the nearest 5°. Specimen
numbers in the Specimens E.xamined sections relate to
Tables 1 and 2.
Rib counts were made on decollated shells b\' mea-
suring the mean width of each whorl on intact shells and
then extrapolating. .As landmarks, some ribs were lightK
marked with a pencil.
Repositories of examined specimens are indicated by
the following acronyms:
AMNH — American Museum of Natural History, Nev\'
York Cit>
AN'SP — The Academy of Natural Sciences of Philadel-
phia
MCZ — Museum of Comparative Zoology, Harvard Uni-
versity
USNM — National Museum of Natural History, Smith-
sonian Institution, Washington, D.C.
SYSTEMATICS AND NATURAL HISTORY
Superfamily Epitonioidea
Family Epitoniidae
Genus Epitoniiiin Roding, 1798
Figures 7-8. Epitunium phyinantlii. 7. Shell ape.\, .showing
a slightly decollated protoconch apex, the almost smooth pro-
toconch, the protoconch varix (shown as a line), some of the
early teleoconch axial ribs, and fine intervening spiral threads.
Topot\pe 8. Operculum (exterior). Topotype.
Epitoniurn phymanthi Robertson, new species
(figures 1-11, 13)
Jong & Coomans, 1988:54, fig.
•'Epitoniurn spec:
276,CAira5ao.
Shell morphology: Protoconch 0 42-0.50 mm long (ex-
cluding immersed base), 0 39-0.42 mm wide, with 3.2-
3 4 whorls; first whorl diameter 0 09-0.13 mm; whorls
slightK inflated, appearing smooth, shin), microsculp-
ture probably present; terminated with varix; cream-
white, tinged with amber at suture, terminal varix, and
columella (seen in transparenc\); some protoconchs
slightly tilted on teleoconchs. Entire shell reaching 16.9
2?
24
I
22
I
e
1 I I I
2 3 4 5
WHORL NUMBERS
Figure 9. Epitoniurn plojmanthi. Rib frequencies per whorl
on each w horl. The horizontal lines show means, the vertical
stippled boxes show ± one stanilard deviation, and the vertical
lines show ranges; n = sample sizes.
Page 84
THE NAUTILUS, Vol. 107, No. 3
Table 1. Epitonium phymanthi new species. Mensural and meristic shell characters. " + " indicates that a shell would lia\e been
larger had it not been slightly broken, or "+ + ' badly broken. Specimen 21 is the holotype All remaining specimens are paratypes
except specimen 28, which was not included in figure 9 because of the high rib counts.
Speci-
men
Length
(mm)
Width
(mm)
Spire
angle
Teleoconch
whorls
A,\ial
ribs per
whorl
no.
1
2
3
4
5
6
7
1
2.2+
1.5 +
60°
2.5 +
25
29
2
3.1 +
2.0+
55°
3.4 +
23
24
—
—
—
—
3
3.0+
1.6+
45°
3.2 +
21
23
22
—
—
—
4
2.3
1.5 +
55°
2.8
27
26
—
5
3,5
2.2
50°
3.8
23
24
28
—
—
6
4.2 +
2.7 +
50°
4.3 +
25
24
23
25
—
—
—
7
4.8
3.1
55°
4.4
28
29
29
29
—
—
—
8
5.5+
3.7 +
60°
4.7
27
30
29
33
—
—
—
9
5.3 +
3.3
55°
4.8
25
28
30
35
—
—
—
10
7.8
4.7
50°
5.6
27
29
28
36
45
—
—
11
7.8
4.6
50°
5.7
24
25
27
28
39
—
—
12
6.7
4.0+
50°
5.3
24
30
27
31
38
—
—
13
5.0+
2.3+
55°
3.7 +
24
27
26
—
—
—
—
14
8.7 +
4.4+
55°
5.3+
24
—
28
37
44
—
—
15
12.2+
3.9+
50°
5.2+
—
26
25
29
—
—
—
16
8.6+
5.2+
50°
5.8+
—
23
26
36
42
—
—
17
9.5 +
5.2+
50°
6.3+
24
24
21
26
32
42
—
18
11.2 +
6.7 +
50°
6.4 +
27
28
29
30
35
45
—
19
10.3 +
5.8+
45°
6.2
22
24
24
26
31
39
—
20
11.4
8.1
60°
6.2
23
21
23
26
33
34
—
21
16.9
9.3
55°
7.2
21
22
21
24
33
35
38
22
16.6
9.6
65°
7.7
23
24
22
23
26
34
40
23
15.3+
8.8
55°
6.2
—
21
25
26
40
36
—
24
5.6
3.1
45°
5.1
22
25
24
24
27
—
—
25
9.1 +
5.6
50°
5.7 +
—
21
25
28
—
—
—
26
16.9
9.5
50°
7.8
18
18
27
30
39
37
37
27
6.0+ +
4.5
—
2.6+ +
—
—
—
34
—
—
—
28
8.6+ +
4.6+ +
—
2.8+ +
—
—
39
45
—
—
—
29
5.9+ +
4.1+ +
—
1.6+ +
—
—
—
—
—
—
—
mm (Table 1). Teleoconch to 9.6 mm wide (including
ribs), with 7.8 whorls, initially thick, becoming relatively
thin, fragile w ith increasing shell size. Spire height vari-
able, usually low, spire angle 4.5°-65° (mean 53°; Table
1). Spire profile at first straight, convex on later whorls
of large shells. Whorls moderately to strongK inflated.
Axial ribs 18-45 per whorl, number increasing with shell
size (figure 9; table 1 ); ribs thick, low, with edges rounded
on upper whorls, thinner, higher, wav\' or crested at
shoulder on later whorls. Apical whorls fully attached to
preceding whorls, lower whorls slightly detached. Axial
ribs aligned, attached erectK from one w horl to the next
on upper whorls, rib alignment and attachment decreas-
ing with increasing shell size; ribs on later whorls rarely
in closely spaced pairs or bifurcating; ribs commonly
curled away from aperture. P'ine, regularK spaced spiral
threads between ribs on early whorls (about 12 above
suture), later whorls with irregular threads, smooth or
with fine incised spiral lines. Basal spiral cord lacking.
L'mbilicus narrow, or reduced to chink. Aperture
oblifjuely to roundK o\ al Teleoconch color cream white
or pure white. Operculum not available.
(Comparative remarks: This new species differs from E.
worsfoldi Robertson (see below) in that the latter species
has a thicker more highly spired shell, with coarser, un-
crested, less numerous ribs (especially on the last whorl
of large shells). Epitonium pht/nidiUhi most closeK re-
sembles species assignetl to de Bour\ 's (1912) "subgenus"
Foliaceiscala (Appendix). The most closeK- similar spe-
cies seem to be E. irregttlarc (Sowerby, 1844) from the
Philippines and Japan, and E. lyirgo (Masahito and Habe,
1976) from Japan (Appendix).
Etymology: Of Pliijriuiulhui, the genus of sea anemone
host.
Geographic range: SE Florida, the Bahamas, Virgin Is-
lands and Grenada (Lesser Antilles). Not known from
I^ermuda or the (Greater Antilles.
IVIaterial examined: (Table 1): SE Florida: Bear Cut,
Miami (25°43'N, S0°09' VV), 1 m, N, and E, Leeman leg,,
196.5, Holotype— ANSP. 391939, A NSP ,391940 & ANSP
391941, specimens 19-23, Leeman collection 1 specimen,
Bahamas: Dead Man's Reef beach, western Grand Ba-
hama (26°34'45"N, 78°51'45"W), S, Bowers leg,, Feb,
1981, via J,N, Worslold, ANSP 374362, specimen 24,
Virgin Islands: Lindbergh Bav, St, Thomas (18°20'N,
64°58'W), 2 m, MR, Hvett leg,, Oct, 30, 1968, ANSP,
R. Robertson, 1993
Page 85
Figures 10-1 1
Topotypes.
Epitoniuiu phymanlhi li\ing in aqi:
specimen 26. Hams Bay, St. Croi.x (17°47'N, 64°53'W),
G. Nowell-Usticke leg., Feb.-March 1957, AMNH 194365
(Specimens 1-18, 27-29)[co-occurred with 1 E. wors-
joldi]. Lesser Antilles: Saint George s Lagoon, Grenada
(12°03'N, 6r45'W), G. Nowell-Usticke Colin., AMNH
194429, Specimen 25. Specimen 21 is the holotype. All
remaining specimens except unnumbered specimen in
the Leeman collection and specimen 28 are paratypes.
A total of 29 specimens of £. phijinanthi was available
for this study. As with E. worsfoldi, many of the shells
are broken.
Natural history: In 1965, Neal and Eleanor Leeman col-
lected six living animals of £. phijmanthi slightly below
low tide line at a rocky area in Bear Cut, between Vir-
ginia Key and Key Biscayne, Miami, Florida, U.S.A. All
were associated with the actiniarian sea anemone Phij-
manthus crucifer (Lesueur, 1817). The anemones were
clinging to rock substrata, commonly in crevices, and
turtle grass (Thalassia festudiniini Banks and Solander
ex Konig) rhizomes, most of which were buried in sand.
The epitoniums were buried next to the Phymanthtis
Figure 12. t'liyiuanthus crucijer, the sea anemone liost ot
Epitonium pliijmantlii at Miami, Florida. Size not recorded,
but soniewlial less than life size. The anemone colors and pat-
terns vary considerably. Photo Neal Leeman.
columns under their radially extended oral discs Only
anemones buried in sand had E. phijmanthi with them.
A few clusters of sand-agglutinated Epitonium egg cap-
sules were observed but not studied.
On .'\pril 17, 1965, a pair of E. phijmanthi was found
with one Pliijmanthus. On May 12, Ma> 30, June 6, and
August 8 of the same year, single £. phijmanthi were
found w ith Phijmanthns. Unlike a predatory Epitonium,
w Inch swallows its coelenterate prey whole, £. phijman-
thi is a parasite, feeding on its large host poK p without
killing it. When fully extended, a large specimen of Phij-
1 1
Figure 13. Kpilunium phijntantln feeding on the upper col-
umn of its sliglitj) contractdl sea anemone host Plujmanlhus
crucifer. Note the cvtraordinariK long e\erted acremijojic pro-
boscis Miami, Florida Fpitoniums, unlike p\ ramidellids, lake
tissues for food rather than fluids. Drawing based on photo-
graphs by Neal Leeman.
Page 86
THE NAUTILUS, Vol. 107, No. 3
13
I
Figures 14-16. EpHonium worsfoldi Robertson, new species.
14. Apertural, 15. basal, and 16. apical (at lower magni-
fication) views of the holotype, 18.7 mm long, 9.1 mm wide.
Smith's Point, Grand Bahama Island. ANSP A17192.
inanthus crucifer is about 13 cm high and about 13 cm
wide, much larger than the wentletrap.
Epitoniums can vary their host preferences from place
to place. During the same year and months, at the same
locality, and associated with the same species of sea
anemone, the Leemans collected six living Epitonium
lamellosum (Lamarck, 1822)(ANSP 391948, A16850) and
one living Opalia crenata (Linnaeus, 1758)(ANSP
A16851). Thus Phijnianthus crucifer supports three par-
asitic epitoniid species at one locality. £, phijmanthi may
or may not be specific to Phymanthus elsewhere.
Colors and color patterns of P. crucifer vary consid-
erably. Usually, these anemones are variegated with
shades of green or brown. The white shell and body of
E. phijmanthi, and the white and reddish brown shell
and nearly white body of £. lamellosum, make neither
species cryptically colored with its host. At least during
the day, both species are buried in sand ne.\t to the
anemone.
The Leemans maintained £. phymanthi and E. la-
mellosum with Phymanthus crucifer in home aquaria
from mid-April to mid-October, 1965 (two E. lamellos-
um for as long as three months). Their observations are
abstracted below.
The external body coloration of E. phymanthi was
white, that of E. lamellosum was white except for light
\ellowish tentacles that taded to white in aquaria. The
fully everted proboscis of E. phymanthi was at least four
times the shell length, while the fully extended proboscis
of E. lamellosum was only slightly longer than the shell.
Roth species behaved as if they locate their host by che-
E . WORSFOLDI
I
_
T
T
T
(III
3 4 6 6
WHORL NUMBERS
Figure 17. Epitonium worsfoldi, new species. Rib frequencies
per whorl on each whorl. The horizontal lines show means, the
vertical stippled boxes show ± one standard deviation and the
vertical lines show ranges; n = sample sizes.
motropism, using their tentacles and slightK' everted pro-
boscises as do two Californian species (Smith, 1977; Saio,
1977).
Both E. phymanthi and E. lamellosum were seen to
feed on Phymanthus columns near the base and under
the oral disc, ingesting column tissues (with fluid? mu-
cus?). Anemone tentacles were not observed to be at-
tacked by either species. Duration of feeding in £. phy-
manthi was 2-10 minutes (animals not starved; mean of
5 observations ca. 4 min). Epitonium lamellosum (a
larger species, up to 45 mm shell length) starved for 3-
5 days fed for 45-97 minutes (mean of 4 observations:
64 min.). No purple secretion was seen to be released
from the pigmented mantle organ b\ either species dur-
ing feeding. One Phymanthus sur\ ived repeated attacks
by the two species for upwards of one month. The anem-
one w rithed near where it was attacked, and the column
also swelled. The anemone reacted least to E. phymanthi,
the smaller, slower species.
An £. phymanthi with a shell 7 mm long on August
11 grew to 1 1 mm by September 22, a mean rate of 0. 1 1
mm /day. Growth initially was faster (between August
11 and 21: 0.2 mm/day), but there was no growth be-
tween September 22 and October 5. Another £. phy-
manthi fed and survived from August 1 1 to September
2 without growing (22 days). Growth of £. lavwllosum
was rather faster and also erratic, but growth appeared
to be indeterminate (Leeman and Robertson unpub-
lished).
Epitonium phymanthi was observed to move very
slowly and remain motionless for long periods of time.
Epitonium lamellosum was more active. On several oc-
casions, £. phymanthi was observed following £. la-
mellosum mucous trails, and partially everting and in-
verting its proboscis, each time ending by touching an
R. Robertson, 1993
Page 87
Figure 18. Oral vie« of either Actinoponis elegans or Hom-
ostichanthus duerdeni. a sand-dwelling sea anemone, showing
beside it the holot\ pe and egg capsules of Epiioniiim worsfoldi
found with it A paratype shell also shows. The anemone is 7,5
cm in greatest diameter. Smith s Point, Grand liahama, I5a-
hamas. Photo Jack N. Worsfold.
.¥'-:C
Figure 19. Li\ ing Epitonitim uursfoldi near some of its sand-
agglutinated egg capsules Some of the shell ribs are abnormally
bent. .-WSP Ai7I94. Shell IS 8 mm long, 9.8 mm wide. Grand
I^ahama, Photo Jack N. Worsfold.
£. lamcllosum. On one occasion, an E. laincllosiun ap-
proached an £. phijmanthi in the same \va) . Once, in-
dividuals of the two species remained aperture to ap-
erture for about one hour, with the four tentacles curled
around each other.
Groups of egg capsules with £. phijmanthi were found
in early May, early June, and early October. After 36
days in isolation, one £. phijmanthi had two groups of
fresh egg capsules attached to it, indicating a capacit>
for sperm storage.
A small crab killed an £. phijmanthi with a shell 10
mm long. Three other, larger animals died from un-
known causes.
Epitonitim worsfoldi Robertson, new species
(figures 14-19)
?Turbo principalis Pallas. 1774: 33, pi 3, figs. 5-6. No localit\
Nonwn (hdritim. This species might not be large if the
figures are enlarged. The spire angle (if accurately drawn lis
30°-35°
?"Sca/a principa/is (Pallas)": Morch, 1875a. 1875b, 1876.Tortola.
British Virgin Islands. ProbabK not of Pallas.
Unidentified. Huber, 1982: 202, 2 figs. S of Balmoral Island,
Nassau, Bahamas. Reported to be I 5/8" long (=41 mm).
"Epitonium principaUs (Pallas, 1774)": Paschall, 1986:66, figs,
la and b. Next to a sea anemone, tureen Turtle 'Ke\"
[Ca> ], .-\f)aco, Bahamas. Not of Pallas Reported to be 1.75
inches long (= 44 mm).
:*£. principale "Roding, 1798": de Jong & Coomans, 1988:54,
fig. 275. "Rather common in Curai^ao and Arutia. Too
few axial ribs.
?"£. fricici (Dall): Espinosa anil Fernandez CJarces (1990:6),
probabl) not of Dall; three small shells from Bah!a Cien-
fuegos, C'uba, the longest 3.8 mm.
Shell morphology: Protoconch 0.52-0.54 mm long (ex-
eluding immersed base), 0.40-0.43 mm wide, with 4.1-
4.2 whorls; first whorl diameter 0.12 mm; whorls slightly
inflated; microsculpture as in £. alhidiim (Robertson,
19S3b); \\ hite, with pale brown at suture, terminal varix.
Entire shell attaining 25.3 mm (Table 2). Teleoconch to
11.5 mm wide (including ribs), with ca. 8.8 thin, fragile
whorls. Spire high, spire angle 35°-50° (mean 44°); spire
profile convex, rareK straight or initialK concave; whorls
moderately inflated. Axial ribs thick, low on upper w horls,
thinner, higher, upright or recurved without subsutural
or shoulder cresting (except remnants from breakage) on
later whorls; ninnbering 16-21 on all but first and last
whorls large shells (23 ribs on last whorl of 8.8+ whorl
shell), with ribs/ whorl initially decreasing then increas-
ing with shell size (figure 17); axial ribs on successive
whorls usually attached, attachments erect, rarely offset
away from aperture on lower w horls; rarely , all ril« bent
medialK after an injury (figure 19). Suture present in
Page 88
THE NAUTILUS, Vol. 107, No. 3
Table 2. Epitonium worsfoldi new species. Mensural and meristic shell characters. " + " indicates that a shell would have been
larger had it not been slightK broken, or "+ + bailK broken Specimen '57 is the liojotNpe
Speci-
men
Length
Width
S[)ire
Teleoconth
Axial ribs
per whorl
no.
(mm)
(mm)
angle
whorls
1
2
3
4
5
6
7
8
1
17.94-
8.9-1-
45°
7.34-
—
—
21
18
18
18
18
2
18.8
9.8
45°
8.0
20
20
19
18
18
19
19
20
3
9.4
5.3
40°
6.5
20
20
20
19
19
20
—
—
4
8.8
5.4
45°
6.6
22
21
20
20
21
21
—
—
5
—
4.7
—
—
19
19
19
18
19
20
—
—
6
—
—
—
3.3
23
21
21
20
—
—
—
—
7
—
—
—
—
20
bS
17
18
17
18
—
—
8
—
11.3-f
—
ca. 8,8
—
—
—
—
—
—
—
20
9
22.1
10.3
40°
8,3
20
20
19
19
18
18
18
18
10
15.6-1-
7.7
.35°
7.2
—
19
19
20
19
17
17
11
12.5
5.9-1-
40°
6.84-
19
19
19
18
19
20
—
12
17.2
8.0
40°
7.64-
—
—
21
20
19
19
19
13
14.3
7.2
40°
7.1
—
—
20
19
19
19
19
14
19.7-H
9.9
40°
8.2
—
—
19
19
19
19
19
20
15
19.6-t-
9.7
40°
8.0
—
21
20
20
20
20
21
19
16
19.7
9.6
40°
7.8
—
20
18
17
18
18
18
—
17
14.9
8.1
40°
7.3
—
20
19
18
18
18
19
—
18
18.3
9.5
45°
7,9
—
17
18
18
17
18
18
—
19
19.2
9.2
40°
8,1-H
—
—
21
19
19
19
19
23
20
4.4
2.4
4(J°
—
—
—
—
—
—
—
—
—
21
6.9
3.8
40°
5.4
—
20
18
19
19
—
—
—
22
25.3
11.5
45°
8.1
—
19
19
19
19
20
20
21
23
15.6
8.2
4.5°
8,0
—
—
—
—
—
—
—
—
24
21.5-1-
9.0-H
40°
7,9
—
—
19
17
17
17
17
—
25
18.9-1-
9.2-H
40°
7.8
—
—
—
19
19
19
—
—
26
17.7-1-
9.3 -H
45°
7.8
—
—
20
21
20
20
21
—
27
13.9-1-
6.4 4-
40°
6.94-
—
—
18
18
18
—
—
—
28
19.4
9.6
4.5°
7.7
—
—
20
20
20
20
20
—
29
15.0-F
7.54-
45°
7.1
—
20
18
18
17
17
17
—
30
18.6-h
10.0
40°
7.2
—
—
—
18
18
18
19
19
31
20.2
10.7
45°
8.2
—
18
18
18
18
18
18
19
32
18.8-1-
10.0
4.5°
8.0
—
—
18
18
18
18
18
19
33
■22A +
—
4.5°
8,3
—
—
20
20
20
20
19
21
34
17 1
8,4
45°
7.74-
—
—
18
17
18
18
18
19
35
12.9
7.2
45°
7.1
—
17
17
18
—
18
18
18
36
10.2-1-
4.74-
50°
5.54-
—
—
17
17
19
18
—
—
37
18.7
9.1
4.5°
7.8
—
19
19
20
20
19
20
—
38
e,.o+ +
2 94-4-
4.5°
5.64-4-
—
—
—
—
—
—
—
—
iiitcr-rib areas of uppermost whorls; middle, lower whorls
slightly detached. Areas between a.xial ribs with faint,
irregidarly-spaced, incised spiral lines; axial growth lines
present, Bod\ whorl v\ ithout basal spiral cord limbilical
chink narrovN (rarely almost closed). Colunieiiar callus
thin or thick, thinnest medially, not conforming to un-
derlying ribs. Columellar chinks present. Aperture
roundly oval. Teleoconch color pale tan, cream-white,
pure white (when bleached?). Operculum ])aucis|)iral,
growth wrinkles present, pale amber.
Comparative remarks: Epitonium worsjolili most close-
ly resembles species in the de Boury "subgenera" Hy-
aloscala [1889], Umiscala [\909], aiuj rapyriscala [\m)9]
(see Appendbx) as di.scu.ssed in Kilburn ( 1985). The shell
of £, worsfoldi most closely resembles /.', kraiissi (Nyst,
1871) of South Africa and E. me/ior (Mel vill andStanden,
1903) of the northern Arabian Sea (see Appendix).
Etymology: Named for Jack Nigel Worsfold, indefati-
gable naturalist and friend.
Geographic range: Bahamas, northern Cuba, and Puerto
Rico. Not known troni Bermuda, Florida, or the Lesser
Antilles.
Material examined: (Table 2): Bahamas: Fleming Road
iH'uch, Mosquito Point, 8 km SE of West End. (Irand
I5ahama (26°37'30"N, 78°54'00"W), J.N. Worsfold leg.,
ca. 1976, ANSP 370051, specimen 33- Smith's Point,
Grand Bahama (26°31'N, 78°37'W), 1 m, 1 living under
Stich()(lu(tyl(i ["Stoichactis"] heliatUhus, 1 large, with
egg capsules, li\ ing under either an :\ctinoporus clegans
R. Robertson, 1993
Page 89
or Homostichanlhns ducrdeni (Figure 18), J.N. VVors-
fold leg., August 1985, ANSP A17192, Holotype. ANSP
A17193, specimens 35, 37.— SW of Sharp Rocks Point,
near Peterson's Cay, S coast central Grand Bahama
(26°33'45"N, 78°33'30"\V), 1 in, 2 juveniles living with
egg capsules under Stichodactyla. J.N. Worstold leg.,
June 1984, ANSP A 17194.— Cold Rock, S. coast central
Crand Bahama (26°36'15"N, 78°22'15"W), 1 m, 2 ju-
veniles living under Stichodactyla. J.N. Worsfold, leg.,
June 1984, ANSP A17195. — Treasure Cay and vicinity,
Abaco, 12 shells, C. Redfern leg., Redfern collection,
specimens 7, 23-32. — W coast N end Elbow (Little
Guana) Cay, off NE Abaco (26°33'00"N, 76°56'45"W) K.
A. Robertson leg. ca. 1954, ANSP 359100, specimen 16. —
Wood Cay. Schooner ("ays, W of S Eleuthera, M
McNeilus leg., Dec. 1980, McNeilus collection, specimen
1.— North East Point, Arthurstown, Cat Island (24°38'N,
75°3S'W), 1 broken shell u ith an £. albidum. W.J. Clench
and H.D. Russell leg., 1936, MCZ 107820, specimen 36.—
Bahamas unlocalized, .ANSP uncatalogued, specimen 34.
Cuba: NW coast. Oasis Beach, Via Blanca Highway km
28, Matanzas (23°11'N, 82°04'W), C.J. Finlay leg. Feb.
1959, Finlay collection, specimen 15. — Same, km 27
(23°11'N, 82°05'W), C.J. Finlay, leg., 1951-1956, Finlay
collection, specimens 5,9,10,11, ANSP 359101, specimens
13,14. Puerto Rico; Punta Ostiones, 7 km S of Punta
Guanajibo, W coast, 1 shell, April 22, 1949, 3 shells. May
22, 1949, G.L. Warmke, leg., ANSP, specimens 3-4, 17-
18. — Ramey Air Force Base, NW coast, A. Phares, leg.,
ANSP, specimen 19. — Pinones Beach, 8-10 km E of San
Juan, NE coast (18°26.8'N, 65°55.7'W), Mrs. D. Hum-
phrey leg., 1970, Finlay collection, specimen 8. Virgin
Islands: Hams Bay, St. Croix (17°47'N, 64°53'W), with
21 £. plujmanthi shells, Feb. -March, 1957, G Nowell-
Usticke leg. AMNH 194365, specimen 38. Specimen 37
is the holotype. All remaining specimens except specimen
34 are paratypes.
A total of 38 specimens of £. worsfoldi was available
for study. Of these, 6 were collected alive: 2 large spec-
imens (the smaller the holotype), 2 juveniles, 1 with the
dried body deep in the shell, and 1 with the operculum
but no bod\ . The last 2 and most of the remaining shells
were collected from beach drift.
Many of the beach shells are in poor condition. The
fairly fragile shell, predators (crabs? fish?), and wave
action presumabK are responsible. Out of the 38 shells,
only 1 has an intact protoconch.
Natural history: At Grand Bahama, this species was found
alive three times with Stichodactyla hclianthus (Ellis and
Solander, 1786), the preferred host also of Epitonium
albidum (Orbigny, 1842)(Robertson, 1983b) and an oc-
casional host of Epitonium lamellosum. Like £. albi-
dum, E. worsfoldi was under the broad oral disc, next
to the column, in sand. The holotype of E. worsfoldi was
collected with an anemone that was either Actinoporus
elegans Duchassaing, 1850 or Homostichanthus ducr-
deni Carlgren, 1900, a sand-dweller (Figure 18). Feeding
(parasitism) on the anemones was not observed but must
occur. The sand-agglutinated egg capsules are like those
of £. albidum (Figure 19). The bathymetric range is
0-2 m.
CONCLUSIONS
The data in this paper are believed to warrant description
of the two new Recent western Atlantic species. Their
fossil ancestors seem not to be known in the Americas.
As documented in the Appendix, the most similar Recent
shells appear to occur in Ja|3an and South .Africa.
It will be noted in the .Appendix that the 23 species
have been assigned to a variety of genera and "subgen-
era" (many of the latter named by de Boury). Epitonium
worsfoldi has characteristics of three of these "subgen-
era combined. Epitoniid genera and subgenera need to
be much more broadly based.
It is surprising that two such large, shallow water spe-
cies in a well-collected area should have remained un-
de.scribed until now. The\ show the continuing role that
amateurs play in collecting, observing, photographing,
and providing material for systematic and biological
studies in museums and marine stations. Amateurs not
onl\ hnd undescribed species on rare occasions but, per-
haps more u.sefull\, they can also make aquarium ob-
servations such as those of the Leemans reported here.
ACKNOWLEDGMENTS
Most of the specimens were collected b\ amateurs, and
AMNH in the 1970's became the first museum to acces-
sion good specimens of one of the two species (from the
Nowell-Usticke Collection). Other amateurs who con-
tributed most to the discoveries by providing specimens,
data, and photographs are Neal and Eleanor Leeman
(formerly of Miami, Florida), C. John Finlay (formerly
of Cuba), and Jack N. Worsfold (formerly of the Ba-
hamas). Worsfold's Grand Bahama collection is now at
ANSP. Dr. Frederick M. Baver identified the Phyman-
thus, and Dr. Donald R. Moore helped the Leemans in
various other ways. Dr. Norman D. Paschall and Helen
DuShane were helpful in providing information. Dr.
Daphne Gail Fautin identified the Actinoporus or Hom-
ostichanthus from figure 18 (no specimen was kept). The
curators of the mollusk collections at MCZ, AMNH, and
USNM helped by providing access to their collections
and making loans. Other persons who helped with the
loan or gift of specimens are Dr. Marvin R. Hyett, Mrs.
Garwin (Marilee) McNeilus, Amy Phares, Colin Redfern,
and Germaine L. Warmke. Mary Fuges inked figures 7,
8, and 13. Harriet Robertson calculated the mensural
data and inked figures 9 and 17. Colin Redfern, Dr Gary
Rosenberg, Jack N. Worsfold, and two helpful reviewers
read various drafts of the manuscript.
LITERATURE CITED
Abbott, R.T. 1974. American seashells, 2nd ed \'an Nnstrand
Reinliold, New York, etc., 663 pp., 24 pLs
.■\danis, A. 1861. On the Scalidae or "wenlletraps of the Sea
Page 90
THE NAUTILUS, Vol. 107, No. 3
ot Japan; with descriptions of some new species .■\nnals
and Magazine of Natural Hi.story (3)8:479-484.
.\zunia. M 1962 Descriptions of five new species of Japanese
Kpitonlidae, N'eniis: The Japanese journal of
Malacology 22(2): 1. 50- 1. -35, (i figs.
Boucfiet, P. and A. Waren. 1986. Revision of the northeast
.Atlantic bathyal and abyssal .Aclididae, Eiilimi-
dae.Epitoniidae (Mollusca, Gastropoda). Bollettino Mala-
cologico Supplemento 2299-576
Boury, E. de. 1912-1913. Description de Scalidae nouveaux
ou pen conniis. Journal de (x)nchvliologie 60(2):87-107,
pi. 7 (1912). (3):169-196, pi. 8 (1913), (4):269-322, pis. 9-
11 (1913).
Clench, W.J. and H.D. Turner. 19.50a. The western Atlantic
marine mollusks described by C.B. Adams. Occasional Pa-
pers on Mollusks, Museum of Comparative Zoology, Har-
vard L'niversity l(15):233-403.
C;lench, VV.J. and R D Turner 19.50b. The genera Stlien-
orytis. Cirsotrcma, Acirsa, Opalia and Ainaea in the west-
ern Atlantic. Johnsonia 2(29):221-246.
Clench, W.J. and R.D. Turner. 1951. The genus EpUonium
in the western Atlantic. Part 1. Johnsonia 2(.30):249-288.
(;lench, W.J. and R D Turner 1952 The genera Epituniiirn
(Part II), Dt'prci^siscala. Cylinilrisccila, Nystiella and So-
Ititiscala'ui the western .Mlantic, Johnsonia 2(31 );289-356.
Clench, W.J. and R.D. Turner. 1953. The genera Epitonium.
Opalia and Cylindriscala in the western Atlantic [supple-
ment]. Johnsonia 2(32):36I-363.
Clessin, S. [189fi-]1897. Die Familie der Scalariidae. In:
Svstematisches C'onchvlien-Cabinet von Martini und
CheriHiitz [2nd. ed], Niirnberg, Rand 2, Alitheilung 13:1-
76, 18 pis.
Dall, W.H. 1889. Report on the Mollu.sca. Part II. Gastropoda
and Scaphopoda. Reports on the results of dredging in
the Gulf of Mexico . . . and in the Caribbean Sea . . . by
the U.S. coast survey steamer "Blake". . . . Bulletin of the
Museum of Comparative Zoology at Harvard College 18:
1-492, pis. 10-40,
DuShane. H. 1970. Two new Epiloniidae from the Galapagos
Islands (Mollusca: Gastropoda). The Veliger 12(3):330-
332, pi. 51.
DuShane, H. 1974. The Panamic-Galapagan Epitoniidae. The
\ eliger 16 Supplement: 1-84, 154 figs.
DuShane, H 1979. The family Epitoniidae (Mollusca: Gas-
tropoda)in the northeastern Pacific The X'eliger 22(2):91-
134, 68 figs.
DuShane, H, 1983. New developments in the famiU Epito-
niidae. The Western Society of Malacologists Annual Re-
port 15:10. Abstract.
DuShane, H. 1988. CJeographical distrihiUmii ol soiiu- Epi-
toniidae (Mollusca: Gastropoila) associated uilli lungiid
corals. The Nautilus 1()2( 1 ):.30-35.
DuShane, H 1990 Hawaiian i;[)itiinii(iac Ilauaiian Shell
News Supplement 1:1-17.
Espinosa, J. and R. Fernandez Garces 1990 La familia Ep-
itoniidae (Mollusca: Gastropoda) en Cuba. Poeyana, Ac-
ademia de Ciencias de C;uba No. 404:1-13, 8 figs.
Franchini, D.A. 1975-1976. FaiTiilia Epitoniidae in the Med-
iterranean Sea. LaConchiglia[Roine|[lll|7(79 80):10-I3
(1975), [IV] (81/82):6-7 (19751, [■'IN'" = V] 8(85/86):
3(1976).
Gardner, J 1947 The molhiscan fauna ol the ,\luin Bluff
Group of Florida. Part VIII Ctenobranchia (remainder),
Aspidobranchia, and Scaphopoda. United States . . . Geo-
logical Surve\ Professional Paper !42-H:ii + 493-656, pis.
52-62.
Gardner, J 1948 Mollusca from the Miocene and Lower
Pliocene of \irgiiiia and North Carolina, Part 2. Scapho-
potla and Gastropoda United States . . Geological Survex
Professional Paper I99-B;iii, 179-310, pis. 24-38.
Gardner, J. A. and T.H. Aldrich. 1919. Mollusca from the
Upper Miocene of South Carolina with descriptions of new
species. Proceedings of the Academv of .Natural Sciences
of Philadelphia 71:17-53, 4 pis.
Gundaker, G,G. 1975. Some Epitoniidae of the U.S. ea.st coast
and Gulf of Mexico. La C^onchiglia [Rome] 7 (73/74):I5-
17,(75/76): 10-11, 14-15, (77/78):14-19, (79/80):12.
(Huber, L] 1982, Who am I? [Epitonium sp J Of Sea and
Shore I2(4):202, 2 figs,
Iredale, T. 1936, Australian molhiscan notes. .No. 2. Records
of the Australian Museiuu 19(5):267-340, pis. 20-24.
Jong, KM, tie and HE. Coomans, 1988. Marine gastropods
from Curasao, .\ruba and Bonaire. Brill, Leiden, etc., 261
pp., 47pls.
Jf)usseaume, [F] 191 1 Fanne malacologitiue de la Mer Rouge.
Scalidae Memoires de la Societe Zoologiijue de France
24:180-246, pLs. 5-7.
Jung, P. 1969 Miocene and Pliocene mollusks from Trinidad.
Bulletins of American Paleontology 55(247):293-657, pis.
13-60.
Kaicher, S.D. 1980-1983. Card Catalogue of World-Wide
Shells. Pack # 23, Epitoniidae Part I (1980). Pack #
30,Epitoniidae Part II (1981). Pack # 35, Epitoniidae III
(1983).
Keen, A.M. 1971, Sea shells of tropical West .Xmerica. , . .
2nd ed, Stanford University Press, Stanford, California,
xiv + 1064 pp,, 22 pis,
Kerslake, J 19.58. A new Epitonium from eastern Australia.
Proceedings of the Royal Zoological Society of New South
Wales, ■T956-57":157-158.
Kiener, L.C. 1838-1839, Genre Scalaire. (Sea/aria, Lam.) In:
Species general et iconographie des cocpiilles vi-
vantes . Paris. 10:1-22. 7 pk
Kilburn, IVN, 1972, Taxonomic notes on South .\trican ma-
rine Mollusca (2), w ith the description ol new species and
subspecies of Conns, Nassarins. \'exillum and Dcmoulia.
Annals of the Natal Museum 21(2)391-437.
Kilburn. R.N, 1985, The family Epitoniidae (Mollu,sca: Gas-
tropoda) in southern .Africa and Mozambique Annals of
the Natal Museum 27(1): 239-337.
Kuroda, T , T Ilabe and K. Oyama. 1971. The sea shells ot
Sagami Bay collected by his Majesty the Emperor of Japan.
Maruzen.fokyo. xix -I- 7^1 + 489 ~l- 51 pp , 121 pis. 1
fold-out map.
Macpherson, J H and C J (Jabriel, 1962 Marine molluscs of
N'ictoria. Melbourne University Press; National Museum
ol X'ictoria Handbook No, 2, xv -I- 475 pp.
Mansfield, VV,(;, 1930 Miocene gastropods and scaphopods
of the (Choctaw hatchee Formation of Florida Florida State
Geological Survey Bulletin 3:1-189.
Mansfield, W.G 1935. New Miocene gastropods and scapho-
pods Irom .\laqua Oeek X'alley, Florida State of Florida
Department of Conservation Geological Bulletin 12:1-64.
Masahito ( I'rinte) and T Habe 1973, .V new epiloniid species
Ironi Japan, \ cniis The jap;inese journ;il of Malacology
31(4):1.).5-1.36,
Masahito (Prince) and I' H.ibc I H7.5 S\sU'malu- stiub ot
Japanese Epitoniidae II iMolluscal Bullcliii ol the Na-
R. Robertson, 1993
Page 91
tional Science Museum [Tokyo], ser. A (Zool) 1(4)191-
194.
Masahito (F'riiice) and T Hahe, 197(1 S\steniatic stud) ol
Japanese Epitoniidae (III), Bulletin of the National Science
Museum [Tokyo], ser. A (Zool.) 2(3):169-174.
Maur\, C: J 1910. New Oligocene shells from Florida Bul-
letins of American Paleontology 4(21): 121-168, 9 pis
Maury, C.J. 1917. Santo Domingo type sections and fossils
Bulletins of .American Paleontolog\ 5(29) 1-251, 39 pis,
Melvill, J C and R, Standen. 1903 The genus Scala (Klein)
Humphrey, as represented in the Persian Gulf, (Julf of
Oman, and North Arabian Sea, with descriptions of new
species. Journal of Conchology 10:340-351, pi. 7.
Miirch, O.A.L. 1875a. Synopsis familiae Scalaridarum Indi-
arum occidentalium. Oversigt over V'estindiens Scalarier
Videnskabelige Meddelelser fra den naturhistoriske For-
ening i Kjiihenluuii for Aaret 1874 (Tredie Aartis, sjette
Aargang):250-268,
Morch, O.A.L. 1875b, Ssnopsis molluscorum niarinorum lu-
diarum nccidentalivnn. Malakozoologische Blatter fiir 1874
und 1875 22:142-184.
Morch, O.A.L. 1876. A descriptive catalogue of the Scalidae
of the West India Islands. Journal of the Acadenn ol
Natural Sciences of Philadelphia (2)8:189-207, pi. 29.
Nakayama, T. 1991. A nev\ cpitoniitl species from the Pacific
coast of the Kii Peninsula, Japan, The Vdiger 34(1):88-
90.
Nordsieck, F. 1982. Die europiiischen Meeres-Gehause-
schnecken (Prosobranchia) vom Eismeer bis Kap\erden,
Mittelmeer und Schwarzes Meer. 2nd ed. F'ischer, Stutt-
gart, New York, .\ii +539 pp., 108 pis.
Nowell-l'stickc, G.W. 1959 A check list of the marine shells
of St. Croix U.S. Virgin Islands. . . . Private!) publisheil,
Christiansted, St. C'roi\, Virgin Islands, vi -I- 90 pp., 4pls,
Nyst, H. 1871. Tableau synoptique et synonymique des es-
peces vivantes et fossiles du genre Scalaria. . . . Memoires
de la Societe Malacologique de Belgique 6:77-147, pi 5
Olsson, A. [A] 1916. New Miocene fossils. Bulletins ol Amer-
ican Paleontology 5(27): 1-32, 3 pis.
Olsson, A. A and A. Harbison. 1953. Pliocene Mollusca of
southern Florida w ith special reference to those from North
Saint Petersburg. , . . The Academy of Natural .Sciences oi
Philadelphia Monograph 8:i-v, 1-457, 65 pis,
Pallas, PS 1774 Tcslacea quaedam rariora In Spicilegia
Zoologica, , , . Berlin, fasc. 10:32-34, pi, 3,
Parenzan, P. 1970 Carta d'identita delle eonchiglie del Med-
iterraneo. Vol. I. Gasteropodi. BiosTaras, Taranto, 283pp ,
53 pis.
Paschall, N.D. 1986. Lost and found; Atlantic "Precious Wen-
tletrap. Conchologists ol .America Bulletin 14(4) 6(S, figs
la and b
Paschall, N'D, 1987, Research results: one species worldwide :■
Epitonium hlainci/zelebnri. Conchologists of America
Bulletin 15(1): 13-14, 2 figs.
Pilsbry, HA. 1922. Revision of W.M. Gabb's Tertiary M<il-
lusca of Santo Domingo. Proceedings of the Academy ol
Natural Sciences of Philadelphia 73:305-435, pis. 16-47
Pilsbry, HA. and A. A. Olsson. 1941 A Pliocene fauna from
western Ecuador, Proceedings of the Academy of Natural
Sciences ol Philadelphia 93: 1-79, 19 pis, + 1 unnumbered,
Poppe, G T, and V. Goto 1991 European seashells \'ol I
Hemmen, Wiesbaden, Germany, 359 pp., 40 pis
P(.uell. A W' B 1979. New Zealand Mollusca . . C:ollins,
Auckland, etc., .\iv + 500 pp., 82 pis.
Robertson, R, 1963, Wentletraps (Epitoniidae) feeding on sea
anemones and corals. Proceedings of the Malacological
Society of London 35(2-3):51-(i3, pis, 5-7,
Robertson, R. 1970. Review of the predators anil parasites of
stony corals, with special reference to symbiotic proso-
branch gastropods. Pacific Science 24(l):43-54.
Robertson, R. 1981 ["1980"]. Epitonium millecostatum and
Coralliophilu clathrala. two prosobranch gastropods sym-
biotic with Indo-Pacific I'alythoa (Coelenterata:Zoanthi-
dae). Pacific Science 34(1): I -17.
Robertson, R. 1983a. A.xial shell rib counts as systematic char-
acters in Epitonium. The Nautilus 97(3):1 16-1 18,
Robertson, R, 1983b. Observations on the life histor\ of the
v\entletrap Epitonium alhidum in the West Indies. Amer-
ican Malacological Bulletin 1:1-12.
Robertson, R. 1985. Archaeogastropod biology and the sys-
tematics of the genus Triculia (Trochacea: Tricoliidae) in
the Indo- West-Pacific. Monographs of Marine Mollusca
No. 3:1-1()3,
Robertson, R, and T Ilabe 19(i5 Alcxania replaces Habea
(Epitoniidae). The Nautilus 78(4): 140-141.
Rc'iding, P F. 1798. Museum Boltenianum. . . . Pars Secun-
da. . . . Hamburg, viii -I- 199 pp.
Salo, S 1977 Observations on feeding, chemoreception and
toxins in two species of Epitonium. The Veliger 20(2):
168-172, 1 fig.
Smith, C'.R. 1977. (chemical recognition of pre\ by the gas-
tropod Epitonium tinctum (Gar[XTiler. 1864) The Veliger
1 9(3 ):33 1-340.
Smith, E..A 1910. On South African marine Mollusca, with
descriptions of new species. Annals of the Natal
Museum 2(2): 175-220, pis. 7-8.
So\\erb\. (;B II. 1844. Monograph of the genus .Saj/nrifl In:
Thesaurus Cxmchvliorum, London, 1:83 bis- 108 bis, pis,
32-35,
.Sowerb>, GB, II. 1873-1874. Monograph of the genus Sca-
luria In: Reeve, L.A., Conchologia Iconica. . . . London
19. 16 pis., 125 species [no pagination].
Souerby, GB. III. 1894. Descriptions of twelve new species,
chiefly from Mauritius. Proceedings of the Malacological
Societ) of London 1(2)41-44, pi 4
Strong, ,AM. and L.G. Ilertlein. 1937. New species of Recent
mollusks from the coast of western North America. The
Templeton Oocker Expedition of the C^alitornia .Academy
of Sciences, 1932. No. 35. Proceedings of the C^alifornia
Academy of Sciences (4)22(6): 159-178, pis. 34-35
Taki, I 1956 Anatomical study on Japanese P^pitoniidae (1 ).
Epitonium, Amaea and Papyriscala Bulletin of the Na-
tional Science Museum (Tokyo) 3(2)(39):71-79, pis. 13-
17
Thorson, (; 1957. Parasitism in the marine gaslropod-familv
Scalitlae. Vitlenskabelige Meddelelser fra Dansk naturhis-
torisk Forening i Kcibenhavn 119:55-58
TrNon, G.W. 1887. Family Scalariidae. In: Manual ol Con-
chology. . .. [ser. 1] 9:49-111, pis. 11-18.
Warmke, G L. and R.T .Abbott 1961. Caribbean sea-
shells. . , , Livingston, WyiHiewood, Pennsylvania, x + 346
pp., 44 pis
Weisbord, N.E. 1962. Late (;enozoic gastropods from north-
ern Venezuela. Bulletins of American Paleontologv 42(193):
1-672, 48 pis.
Wilson, B.R. and K. Gillett, 1972, Au.strahan shells, Tuttle,
Rutland. X'ermonl, and Tok\o, 168 pp,, Ut6 pis
Woodring, \V,P, 1928, Miocene mollusks from Bouden, Ja-
maica Part II Ciastropods and discussion of results. Car-
Page 92
THE NAUTILUS, Vol. 107, No. 3
iiegie Iiistitiiliiiii cil W'.isliint^lnii rublicalion .38,5:i-vii, 1-
56-4, JOpls.
VVoodring, W.P. 1959. Geology and paleontology of Clanal
Zone and adjoining parts of I'anania Description of Ter-
tiary molliisks (gastropods: \ errnflidac to Tliaididae).
[United StatesjCleological Siir\fv Prolessioiial Paper '>06-
B:iii, 147-2.39, pis. 24-;5S
Yamashiro, H. 1990. A wcntlelrap i'.pitoniiini hullatuiu as-
sociated with a coral Sundali/lithu robusta. \ enus: The
Japanese Journal of Malacology 49(4):299-305.
APPENDIX.
Conchological comparisons or notes on 23 species similar
to, or confused with Epitonhim phijmanthi and E. wors-
foldi, regardless of provenance.
Scala amathusia Melvill & Standen, 1903:341, pi. 7,
fig. 1. Kaicher, 1980: card 2300 [Epitonium]. Gulf of
Oman. Smaller than £. worafuldi (ft. 5 mm), with more
whorls (8-9) for size.
Scalaria costulata Kiener, 1838-1839:5-6, pi. 2, figs.
4. Tryon, 1887:pl. 12, fig. 59. Robertson, 1963:57-58, pi.
5, fig. 4. Robertson, 1970:45 [identification confirmed].
DuShane, 1988:30-32, fig. 1 (ribs curiously opisthocline).
ANSP 230639. Red Sea to Philippines. Thought to be a
synonym of S principalis (Pallas) by Sowerby (1844:88
bis), which he recorded from "Xipixappi, West Colum-
bia." Larger (35.3 mm); spire high (spire angle 30°);
suture slightly disjunct; up to 28 ribs on last whorl; um-
bilicus narrow, deep.
Scalaria crassa Sowerby, 1844:91 bis, pi. 33, fig. 55.
Kaicher, 1981: cards 3066, 3081 [Epitonium]. Philip-
pines. Unlike E. ivorsfoldi, ribs unequal, slightly fewer
(18) at largest size; may be spirally brown-banded.
Epitonium (Limiscala) cnjpticocorona Kilburn, 1985:
309-310, figs. 79, 126. South Africa. Smaller (7 mm);
weak "coronation" inside suture; 12-19 ribs; low spiral
lirae.
Scalaria dubia Sowerby, 1844:90 bis, pi. 33, fig. 41.
Not Epitonium duhium Roding, 1798:92. Sowerby in
Reeve, 1873: species 45, 45b wrongly as 75b. Tryon, 1887:
60, pi. 12, figs. 67-68. De Boury, 1912:95-97,' pi. 7, fig.
4. Kilburn, 1985:327. Type species of Foliaceiscala de
Boury, 1912. ANSP 19573 (broken); USNM G.177. Aus-
tralia, Samoa, etc. Larger (37 mm); spire angle 45°; like
£. phymanthi but lacks crests; ribs very low, thin, slightly
recurved, rarely enlarged, up to 42 on last whorl; suture
impressed but not disjunct; spiral threads crowded, vari-
able; last whorl inflated; umbilicus very narrow.
Scala emiliac Melvill & Standen, 1903:343-344, pi. 7,
fig. 6. Kaicher, 1980: card 2313 [Epitonium]. Kilburn,
1985:307, figs. 120-121. Pakistan to South Africa. Smaller
than E. phymanthi (12 mm); spire slightly higher (spire
angle 40°); 21-36 low, thread-like ribs; pale brown (if
not laded).
Scalaria Jriahilis Sowerby, 1844:95 bis, pi. 33, fig. 74.
Kaicher, 1980: card 2^29 [Epitonium]. Southern Austra-
lia. Higher spire than E phynianllii (spire angle 30°-
35°); spire profile evenly convex; no umbilicus.
Scala frielei Dall, 1889:313. Not illustrated. USNM
83727 (syntypes): AMNH 194388. North Carolina; Vir-
gin Islands. Clench & Turner (1952:300-301) and
Warmke & Abbott (1962:81, pi. 14a) misidentified this
species. Smaller (6.4 mm); 24-31 ribs on last whorl; no
shouldering; strong spiral threads; umbilicus narrow ; spire
fairly high (spire angle 45°). Epitonium phymanthi has
been misidentified as this species.
Epitonium (Nitidiscala) hancocki DuShane, 1970:332,
pi. 51, fig. 1 DuShane, 197432-33, fig. 73. Kaicher, 1983:
card 3612. Galapagos. Smaller (13 mm); spire higher
(spire angle 30°); fewer ribs (21); rib crests wavy.
Scalaria imperialis Sowerby, 1844:91-92 bis, pi. 33,
figs. 56-.57. Wilson & Gillett, 1972, pi. 13, figs. 5, .5a
[color]. [Epitonium inipcrialc]. Kaicher, 1980: card 2304.
AMNH 1.36625, 1.57310; ANSP 181726, 19,5661, 2.5.3841,
25.3842, uncat.; MCZ 294909, 294914; USNM 694170.
East Africa, Western Australia, Philippines and Queens-
land. Larger (40 mm); pale tan to dark reddish or pur-
plish brow !i, commonly in spiral bands (ribs white); more
ribs, increasing to 48 on last whorl; umbilicus wide and
deep; operculum grey-black.
Scalaria irregularis Sowerby, 1844:90 bis, pi. '33, figs.
40, 60. Philippines; Japan. USNM .343440. Up to 18 mm
long; spire angle 6.5°; ribs crowded, 37 on last whorl,
irregular in thickness but generally thin, with slight crest-
ing; strong spiral threads; slightly umbilicate; 1 pale brown
subsutural band present or absent on white. Resembles
£. phymanthi.
Scalaria kraussi Nyst, 1871:116. Kilburn, 1985:29.3-
295, figs. 97-100 [Epitonium (Hyaloscala)]. + Epiton-
ium shepstoncnsc E..^. Smith, 1910:204, pi. 7, fig. 15.
Kaicher, 1981: card 3113. .South Africa. Smaller (10-16
mm); spire averages higher (spire angle 25°-40°); 17-37
fine, low, erect or slightly reflexed ribs on later whorls;
umbilicus closed. Species believed to be most similar to
£. ivorsfoldi
Perlucidiscala lacrymula Jousseaume, 1911:198, pi. 5,
figs. 37-42. Kaicher, 1981: 3063 [Epitonium "lachry-
mida"]. Gulf of Aden. Smaller (5 mm); spiral threads
towards apex.
Scalaria latifasciata Sowerby in Reeve, 1874: species
117. Type species ot Papyriscala de Boury. Taki, 1956
[anatomy]. ANSP 70738, 219307, 234733, 243255; MCZ
294908 and 5 uncat. lots. Mozambique; Mauritius; Japan.
L'p to 18 mm long; spire low (spire angle 5.5°); trace of
whorl disjunction; thin, low ribs mostly not connected
from whorl to whorl; 32 ribs on last w horl, some irreg-
ularly spaced or thickened; weak to strong spiral threads;
no crests; 2 or 3 brown spiral bands; slightly imibilicate.
Scalaria lineolata "Kiener" Sowerby, 1844:91 bis, pi.
33, figs. 45, 46, 48. Confused with S. lineata Kiener not
Sav. Nvst, 1871:118, pi. 2, fig. 5, pi, 6, fig. 20. Kaicher,
1981: card 3125 [Epitonium]. ANSP 70742, 86246,
243289; MC:Z 294911 and 2 uncat. lots; USNM 198709.
Gulf of Aqaba; Mauritius; Philippines; Japan. Up to 19
mm long; spire angle 40°; ribs more numerous (16-33)
and more irregularly spaced and thickened ribs; no spiral
sculpture; umbilicate; 1-3 brown spiral band(s) per w horl;
operculum dark brown
Scalaria lyra Sowerby, 1844:89 bis, pi. 32, figs. 38-.39,
pi. 34, figs. 81-82; Sowerby in Reeve, 1873: species 23.
R. Robertson. 1993
Page 93
Kaicher, 1980: card 2341 [Epilimininl KillKirn, 1985:
308-309, figs. 122]. ANSP 195(iH, 19567, 119638; MCZ
187720; LlSNM 431817, 306339; 820891. Type species
of Limiscala de Boury. Fiji to Japan and the Red Sea to
Mozambique. Slightly smaller (20+ mm); spire lower
(spire angle 60°); 27-45 erect ribs; spiral threads fine and
dense; 0, 2 or 3 brown spiral bands.
Scala melior Melvill & Staiulen, 1903:345, pi. 7, fig.
9. Kaicher, 1980: card 2S33[Epitonium]. ANSP 164808
(syntype); USNM 424868. Pakistan; Gulf of Oman
Smaller (6.5 mm + ); 8 (teleoconch?) whorls; spire angle
30°; up to 23 ribs per whorl; spiral threads present or
absent. Resembles a small E. wor.sjoldi.
Scala micromphala Morch, 1875a:258 ("Vieques,"
Puerto Rico, one specimen collected by A.H. Riise) Not
illustrated. Photograph of holotype in Clench and Turner
(1951:258-260, pi. 112, fig. 3), wrongly synonymized by
them with E. occidentale Nyst, 1871, which according
to them has "12 to 15 costae on the body whorl. ' Ac-
cording to Morch there were 18 ribs on the 5 mm. -long
shell. An E. phymanthi that long would have had more
than 20 ribs. In addition, E. micromphala has subsutur-
ally crested ribs; in E. phymanthi the crests are on the
shoulders of the ribs. Morch wrongly likened S. microm-
phala to S. imperialis (see under that species)
Scala {Viciniscala'"') mimiticosia de Bour\', 1912:87-
90, pi. 7, fig. 1. Compared with Scala costulata and S.
principalis "West Columbia." DuShane ( 1974:20-22, figs.
20-26). Larger than £. worsfoldi (35 mm); spire high
(spire angle 35°); 21 mainly thin, low ribs on last whorl;
whorls all narrowly disjunct, but ribs attached; traces of
spiral striae; umbilicus narrow, deep; ribs slightly re-
flexed; crowded spiral threads.
Turbo principalis Pallas, 1774. Nomen dubium. Epi-
tonium principale (Pallas) Roding, 1798:91. De Bourv,
1912:89-90, 97. See above in synonymy of E. worsfoldi.
Scalaria rohillardi Sowerb\ , 1894:42-43, pi 4, fig. 5.
Kaicher, 1981: card 3037. Kilburn, 1985:305-307, figs.
118-119 [Epitonium (Papyriscala)]. Red Sea and India
to South Africa. Smaller (15 mm); more ribs (20-24 on
last whorl); umbilicate; pale tan with 2-3 brown spiral
bands (rarely absent).
Epitonium shcpstonense. See above under E. kraussi.
"Foracciscala" [=Foliaceiscala] virgo Masahito &
Habe, 1976:172, figs. 2-3. Compared with "Foraccis-
cala" duhia (Sowerb\, 1844). Japan. Paratype most re-
sembles £. phynuinthi, spire angle 40°; 50-60 weakly
lamellate growth riblets; suture slightly disjunct, whorls
connected by ribs; aperture constricted (holotype); oblique
spiral threads.
Epitonium (Nitidiscala) willetti Strong & Hertlein,
1937:171, pi. 35, fig. 5. DuShane, 1974:38-39, fig. 72.
Tropical eastern Pacific. Much smaller (3.2 mm); 18-
22 -h ribs.
THE NAUTILUS 107(3):94-96, 1993
Page 94
Phalium {Semicassis) vector, a New Deep-Water Species from
the Central Indian Ocean
R. Tucker Abbolt
Bailey-Matthews Shell Miimuiii
P.O. Box 1580
Saiiibel, FL 33957 USA
ABSTRACT
Phalium (Semicassis) vector, a new species of the family Cas-
sidae, is described from the Saya de Malha Bank, northeast of
Mauritius in the Indian Ocean It differs from its closest relative,
Phalium microstoma (von Martens, 1903) from Somalia, in
having a lighter, less elongate shell with stronger beads on the
spiral lirae.
Keii irords: Cassidae, Sa\a dc Malha liank. Indian Oceaii
INTRODUCTION
A number of new species of marine mollusks have been
collected over the past few years by Soviet research ves-
sels exploring the Saya de Malha submarine bank in the
central part of the Indian Ocean. Among these are species
of cassids not known at the time of my revision of this
family (Abbott, 1968).
In 1975, Kilburn described Phalium glahratum sub-
species fernandesi from deep water off Somalia. Two
years later it was dredged by the R/V Gordy on the
Saya de Malha Banks. With it were recovered at least
five specimens of a new Phalium referable to the sub-
genus Semicafisis Morch, 1852.
The bathymetry antl geologs of the Saya de Malha
Bank are discussed by Bouchet and Bail (1991:161), while
Bondarev and Rockel (1992) provided an excellent pop-
ular account of this bank, which is located along the
Mascarene Ridge north of the Mauritius and C^argados
Carajos Shoals (between 9°:3()'S and 12°2()'S, and 59°30'E
and 62°3()'E).
interstices of etjual width. CJolor opaque white with large,
faint, light-brow II blotch on dorsal surface of last whorl.
Protoconch (fig. 2) of 3'/2 whorls, obtuse-conical, glassy-
white, smooth. Teleoconch of 5'/2 whorls, the last bearing
23 flattish, spiral cords with fine beads at intersections
with fine axial threads. Suture simple, slightK impressed.
Spire convex-conic. Aperture % shell length. Outer lip
reflected, of variable thickness, enamel-white, crossed by
20-22 distinct, sharp white denticles. Both true and false
umbilici small, open, deep. Channel behind siphoual ca-
nal deep, narrow. Parietal wall slightly glazed over. An-
terior part of columella strongly and unevenly pustulose,
inner edge with 4-5 small denticles, outer edge with 3-
4 small, roimded projections. Soft parts and operculum
tmknown.
Type locality: Central Basin, Saya de Malha Bank, Mas-
carene Ridge, northeast of Mauritius, Indian Ocean,
dredged from sandy silt bottom at a depth of 80 meters,
R/V Gordy, 1989. More precise locality data did not
accoinpany the holotype.
Material examined: Holotype, USNM 860326, Paratypes
1-3, Donald Dan collection, all from the type locality.
The specimen illustrated by Bondarev and Rockel (1992;
fig. 5) is designated as paratype 4
Etymology: 1 take pleasure in naming this species after
the first recreational diving vessel to operate under the
Russian flag, the Vector.
Comparative remarks: This new species belongs to a
species group of Phalium (Semicassis) mainly limited to
SYSTEM ATICS
Phalium (Semicassis) vector Abbolt, new species
Figures 1-2
Synonymy:
Phalium sp 1. Bcindarev & Biicki
1992:28,32, fig 5
Description: Shell (fig. 1 ) reaching 60 mm in length,
light, strong, evenly ovate, with numerous, fine, flattish,
crowded, weakly-beaded spiral lirae separated b\ sunken
Table 1. Phalium (Semicassis) vector new spfc'\es l.inearshell
measurements (nun) and counts
llol(it\pc
I'aratype 1
Paratype 2
l'arat> pc 3
Length
53.6
56.8
53.4
(iO 0
Width
32,0
32. 1
33.4
33 5
Number ol
whorls
R. T. Abbott, 1993
Page 95
'--^
\.
Figures 1, 2. Phalitun {Seniicassis) vector new species. 1. A|Hiluidl, lalcral, and dorsal views of the holotype. 1.0 X. 2. Protoconch
of paratype 1. Botli from Saya de Maiha Bank, Mascarene Ridge, N E. of \4adagascar, dredged from sandy silt bottom at a depth
of 80 meters, R/V C;ordy, 1989. Scale bar = 500 nm.
the western Indian Ocean, although similar species, such
as P. sculptuni J. de C. Sowerby, 1840 (Miocene), P.
mehranictim (Vredenburg, 1925) (Upper Miocene) and
P. oligocalanticum (Vredenburg, 1925) (Oligocene) have
been reported from fossil beds on the mainland of India.
Phalium vector most closely resembles the eastern Af-
rican deep-water Phalium {SemicassU} microstoma (von
Martens, 1903), from which it differs in being niucli
lighter in weight, less elongate, and in having stronger
beads on the spiral lirae. Phalium vector lacks the four
or five spiral brown bands on the body whorl, but instead
has a faint tan blotch on the dorsal surface of the body
whorl. The very weak, rounded projections on the outer,
left edge of the columella of P. vector are reminiscent
of the fingertip projections found in the shallow-water
Indian Ocean Phalium Jaurotis (Jousseaume, 1888).
ACKNOWLEDGMENTS
I am indebted to Mr. Donald Dan of Ft Myers, Florida,
L SA, and to Mr. Valeri B. Darkin of Vladivostok, Russia,
for allowing me to examine and describe this new species.
\lr Darkin kindly presented the holotype of to the Na-
tional Museum of Natural History, Smithsonian Insti-
tution (USNM).
LITERATURE CITED
Abbott, R. T. 1968 The helmet shells of the world (Cassidae).
Part 1. Indo-Pacific Molliisca 2(9):15-202,
Bondarcs, I and D R5ckei, 1992 The shells of the Sa\a de
Malha Bank, La C:onchig!ia 23(262):21-34.
Biiuchet, P aiidP Bail, 1991. Volutes from the Sava de Malha
Page 96 THE NAUTILUS, Vol. 107, No. 3
Bank: The saga of Lijria surinanwn.si.s aiul a ni-u species. soiclae, Ceritliiidae, loniiidae, Cassidae, Buccinidae, Fas-
The Nautilus 105(4):159-164. ciolariiilae, Turhinellidae, urridae, Architectonicidae, Ep-
Kilburii, R. 1975. Taxonomic notes on South African marine ilotiiidae, l.inndae, and Thraciidae. Annals of the Natal
Mollnsca (5); including descriptii>Ms of new ta\a of Ris- Museum 22(2):.577-622.
THE NAUTILUS 107(3):97-103. 1993
Page 97
Ferussac and d'Orbigny's '^Histoire naturelle generale et
particuliere des Cephalopodes acetabidiferes": Dates of
Publication of Plates and Text
Annie Tillier
Renala Boucher-Rodoni
Biulogit-' dt'S lii\ fi'tebri'S niuriiis ft
Malacologie
LIRA 699 CNRS— Museum National
d Histoire Naturelle
55, rue Buifon
75005 Paris, FRANCE
ABSTRACT
Ferussac and d Orbigny s "Histoire naturelle generale et par-
ticuliere des Cephalopodes acclahuliferes is an essential con-
tribution to teuthology, because of the numerous new species
described, and because it represents the first important com-
pilation of cephalopods Issued between 1834 and 1848, in 21
jisraisons, the actual dates of publication of most ot its parts
were hitherto uncertain. A thorough bibliographic search has
allowed us to gather new information about the publication nl
this work. We can now identif > and date preciseU the '50 plates
issued in 1834 and the 72 plates issued in 1835; the remaining
42 plates are dated 1839-1841 or 1839-1842, D'Orbigny's text
was issued between 1839 and 1848 (livraison 12-21) The last
three livraisons (1845-1848) included only text
Key Words: Cephalopods; Publication dates; Ferussac and
d Orbigny.
INTRODUCTION
During the first half of the 19th Century , a vast amount
of material was collected Bn' the scientists participating
in French voyages around the world, the accounts of
which were published under the following titles: Voyage
aux terrea Anstrales (Ic Geographe, le Natiiraliste, la
Casiiarina) from 1800 to 1804 by Peron; Voyage de la
Coquille. from 1822 to 1825 by Lesson; Voyage en Arnc-
rique meridionalc , from 1826 to 1833 by d Orbigny;
Voyage autour du rnonde (Uranie et Physicienne), from
1817-1820and Voyage de I' Astrolabe, from 1826 to 1829
by Quoy and Gaimard; Voyage de la Bunite. from 1836
to 1837 by Eydou.x and Souleyet, The specimens collected
on all of these voyages were deposited in the Museum
National d'Histoire Naturelle, Paris, and described not
only by the collectors themselves, but also by other con-
temporary zoologists such as de Blainville, Lamarck and
Ferussac. These collections gave rise to numerous works
that constitute a considerable contribution to the devel-
opnient of natural history.
The publications issued during the first half of the 19th
century were delivered as a series of numbered parts, or
livraisons, and several years were often necessary to com-
plete a work. Such is the case for the "Histoire naturelle
generale et partictdiere des Cephalopodes acelabuli-
feres. . ." by Ferussac and d Orbigny, which was issued
in 21 livraisons, from 1834 to 1848, and whose publi-
cation was particularly lengthy and confused.
Ferussac intended to publish an important and general
work on Mollusca ("Histoire naturelle. generale et par-
ticuliere des Mollusques'). where all the monographs
written by him and other authors might be included. As
early as 1825, he had asked d'Orbigny to collaborate in
the publication of the "Histoire des Cephalopodes".
D'Orbigny started at once to produce plates and te.xt,
but in 1826 he left for an eight year long voyage to South
America. It was only in 1834-35, after his return, that
the first parts were distributed. Ferussac died in 1836,
and it was not until 1837 that d Orbigny could recover
his manuscript, and he then spent the follow ing two years
updating it (1839; issue of the first part of d'Orbigny's
text).
DESCRIPTION OF THE WORK
The "Histoire naturelle generale et particuliere des Ce-
phalopodes acetalndiferes. . ." comprises 2 volumes
bound according to the instructions of d Orbigny:
1 — A volume of text, starting with d Orbigny s Preface
(pp. I-LVI), followed by 361 pages of taxonomic de-
scriptions. The first livraison of text (livr. 12) is dated
1839 and largely postdates the Atlas.
An introduction to Mollusks and general remarks on
Cephalopods by Ferussac (96 pages), was issued in liv-
Page 98
THE NAUTILUS, Vol. 107, No. 3
raisoiis 1 to 6 (1834-1835). A copy of this text was coii-
.siilted at the "Bibliothecjue Nationale" in Paris, where
it is kept in the original wrapper, on the cover of which
appears: " Histoirc naturcllc ncncralc et part icxtUcre des
MoUusqiies — Monographic des C'cphalopodcs Cnjpto-
dibranches" . This part was originally intended to be
included in the work, but d'Orbigny considered it to be
obsolete. In his instructions to the liinilcr he specifies that
it is not part of the work on the Cephalopods and must
therefore be bound separately.
2 — An Atlas of 144 plates (90 plates named "Cryptodi-
branche", 52 named "Acetabulifere", one bearing both
names (Octopus pi. 23) "Cryptodibranche" and "Aceta-
buliiere ', and one having neither of them (Bellerophon
pi.T).
Winckworth (1942) stated that a large number of plates
were ready in 1826, when d'Orbigny left for South Amer-
ica; but were delivered only after his return in 1834.
Ferussac was at that time publishing the " Histoire na-
turcllc . . . des Mollusques terrestres et fluviatiles" in
collaboration with Deshayes. He also had to assume the
heavy costs of the "Bulletin Universel des Sciences et
de I Industrie' \ which he created and managed. How-
ever, Ferussac (1831:374) himself confirms that most of
the "Cryptodibranche ' plates were ready in 1831: "de-
puis longtemps tous les Cephalopodes de I'ordre des
Cnjptodihranches . . . composent cet ouvrage pret a etre
livre a I impression, et dont les planches au nombre de
pres de 80 sont tirees et enluminees".
HISTORICAL ACCOUNT OF PREVIOUS
RESEARCH
Sherborn and Woodward (1901:75) were the first to pub-
lish some dates, in a footnote to their paper on the pub-
lication dates of "Mollusques terrestres et fluviatiles"
by Ferussac and Deshayes. They are very concise and
do not mention their sources: "Of the dates of publication
of this work all that can at present be said is that the
first three out of 21 liv. appeared in Sept. 1834, nos.4-9
in 1835, no.l2 in 1839, nos.13-18 in 1840 taking the
work up to p. 240; and that it was finished in 1848".
The copy of Ferussac and d'Orbigny s work in The
Natural History Museum, London, gives some evidence
of how Sherborn deduced some of the dates. \ hand-
written note says:
"It is obvious, as the date 1837 occurs on pp. Ill -I- 27
& 1838 on pp. V + 73 & 1839 on p. 139— that no text
was published before 1837. 9 parts were published by
1835 (clearly these were all plates or part of the text to
volume II Atlas, which containing no species does not
concern us). Part 12 was published in 1839 & 13-18
(bringing the work up to p. 240) in 1840. \ fresh font
of type was used on p. 241 (compare the 444 's in the
pagination) & references to 1845 are frequent between
pp. 241-361.
I therefore regard \'ol. I, pp. I-LVI & 1-240 as 1840;
pp. 241-361 as 1848; Vol. II text, pp. 1-96 as 1835".
These considerations are dated October 1901. The last
part is repeated in the bibliograplu of the Index .\ni-
nialium (Sherborn, 1922: LlII), with an extra remark:
"[Dates] of plates unknown".
The references to 1835 most probabK come from the
Bibliographie de la France, which he thouroughly con-
sulted to date Ferussac and Deshav es' "Histoirc naturcllc
des Mollusques. . .". He thus probabK knew that the first
9 livraisons were issued in 1834 and 1835, and that thev
comprised only plates (although he did not know w hich
ones) and Ferussac's preface.
Thus Sherborn did not have sufficient information to
date the plates, but he attempted to date the text, rel\ ing
on the dates cited in the first pages of the text and in
the preface by d'Orbigny. His conclusion was that the
text could not have been issued before 1837 (he had no
data regarding livraisons 10 and 11).
As far as the dates of livraisons 12 (1839) and 13-18
("bringing the work up to p. 240 "; 1840) are concerned,
we could not determine his sources. He stated that a
fresh type font was used on p. 241, in the pagination,
but after a thorough examination of the different copies
we could consult, we noticed that such a change in the
pagination font of type actually occurs on page 273.
Winckworth (1942) had some additional data that al-
lowed him to propose dates for the plates issued in 1834-
1835. He possessed a bound copy of the 1834-1835 liv-
raisons, although he did not have enough information to
know precisely if this copy represented 9, 10 or 11 liv-
raisons, and w hich of the plates were issued in 1834 and
which were issued in 1835.
For the text, he used:
- the dates proposed b\ Sherborn and Woodward
(1901);
- d Orbign\ s "Mollusques vivants et fossilcs" (1845),
a thorough reading of which allowed him to point out
that all the plates and the text up to and including page
271 are mentioned. This led him to introduce an extra
break in the text, and to date as 1848 all the text following
page 27 1 ;
- the text of d Orbignv himself, where 1839 is the latest
bibliographic reference on page 210, and where the
"Mollusques vivants et fossiles" (1845) is first mentioned
on page 273;
- the indications on the back wrappers of the "His-
toirc. . . des mollusques terrestres et fluviatiles", with
which he, however, partly disagrees. This is discussed in
more detail below .
ORIGINAL RESULTS AND DISCUSSION
Three articles issued in the "Bulletin zoologique" and
in the "Annalcs des Sciences naturcllcs" enlightened us
as to the content of the first li\raisans, and encouraged
us to investigate further and to retrace the history of the
publication of the "Histoire generale et particuliere des
Cephalopodes Acctabidifcrcs". Successively, the follow-
ing data w ere gathered from 1 1 different sources:
A. Tillier and R. Boucher-Rodoni, 1993
Page 99
1 — "Bibliographic de la France, ou Journal general de
I'Imprimerie et de la Lihrairie' Paris, Pillet Aiiie:
23e annee, no. 39, 27 Sept. 1834, p. 615, no. 5229;
24e annee, no. 16, 18 Avril 1835, p. 246, no. 2099;
no. 23, 6 Juin 1835, p. 360, no. 3058.
The issue, nuini)er oi pages of text and plates of liv-
raisons 1-9, are announced fnit no details are given on
tfie content of tfie livraisons: Livraisons 1-3 (27 Septem-
ber 1834): 30 plates and 5 sheets of text; livraisons 4-6
(18 April 1835): 29 plates and 7 sheets of text; livraisons
7-9 (6 June 1835): 25 plates and 1 sheet with front page.
2 — "Bulletin Zoologiquc". 2e section, 1835:
- p. 14-16: some details on livraisons 1-3 are given;
- p. 63-65: all the species figured in the plates of
livraisons 4-6 are listed.
3 — "Annales des Sciences naturellcs", 2e serie, 3, Zool-
ogie. Paris, Crochard 1835, p. 192 (f.l2, March). As early
as March 1835, 12 livraisons of 9 to 10 plates each are
announced as read\ , 9 of which were on sale (whereas
La Bibliographie de la France announces their issue later
in the same year). A list of all the species issuetl in the
plates of the livraisons 1-9 is given.
4 — "Notice analytique sur les travaux de Zoologie de
Mr. Alcide d'Orbigny" (1856). Dumeril made two favor-
able reports at the Academic des Sciences, on the first
eleven livraisons, on the 15 December 1834 and 29 June
1835 (see also d Orbigny s Introduction, p. II). This allows
livraisons 10-11 to be dated 1835.
5 — "Registre des proces-verbaux et rapports des seances
de r Academic Royale des Sciences. Institut", vol.10,
1832-1835: p. 579, 22 September 1834: the delivery of
Ferussac's first three li\raisons is announced; p. 724, 29
June 1835: Dumeril indicates that the last five livraisons
are issued.
In the "pochette de la seance du 15 deeembre 1834"
we found Dumeril's report (two handwritten pages) de-
scribing the first three livraisons, i.e. an Introduction b\
Ferussac and 28 plates ("les figures de ranimal de I'Ar-
gonaute argo et de son anatomic, faites par Poli . . .
lithographiees par Chazal. Plusicurs especes nouvelles
de poulpes, et des dessins executes a Cadix d'apres les
dissections de deux tres habiles anatomistcs . . . repro-
duites par Jacob".
There is no trace in the "pochette of Dumeril's report
on the 29 June 1835, concerning the "last five livraisons".
Our sources are consistent with livraisons 1 to 1 1 being
delivered in 1834-1835, the first nine three by three. The
"last five livraisons" announced by Dumeril in 1835 are
thus livraisons 7-9 and 10-11.
6 — "La litterature jranqaise contemporaine, 1827-1844,
Dictionnaire Bibliographiqiie" b> MM. Ch.Louandre and
F. Bourquelot, Paris, Felix Daguin, t.3, 1848, p. 190-
191. The " Histoire . . . des Cephalopodes ' , is announced
as published in Paris, 1834-1842, in 20 livraisons, each
comprising 10 plates and corresponding text, 18 livrai-
sons of which were on sale. Thus we did deduce that 18
livraisons were issued from 1834 to 1842, livraison 18
being dated 1842; we know that the last livraison (21st)
was issued in 1848. From the e\ idence of the type font
change on p. 273 and of references to d'Orbigny (1845),
we consider that livraison 20 (pp. 273-320, see below)
also appeared in 1848. As for livraison 19, we date it
1845, since apparently it was not yet on sale in 1844, Init
the text up to p. 271 is quoted by d'Orbigny (1845).
7 — Two complete works (Text and Atlas) from the "Bi-
bliothcque Nationale": their peculiarity is that they have
been marked with different stamps (Bibliotheciue Im-
periale, Bibliotheque Royale or Bibliotheque Nationale).
.'Ml the plates are stamped, w hereas the text bears only
a few stamps. A thorough examination of these stamps
in the two volumes of text revealed that they were not
randomly distributed but seemed to correspond either to
the beginning or to the end of a set of livraison (Table
1). ^
In one of the copies of the Atlas, all the plates are
marked "Bibliotheque Royale", except 32 that are
stamped "Bibliotheque Imperiale". These 32 plates cor-
respond to most of the plates that we consider as be-
longing to livraisons 1-3. We conclude that the latter set
constitutes a homogenous lot, which can be dated 1834,
except for 4 plates of Argonauta, which we are sure
belong to livraisons 7-9 (Bull.lool. 1835 and Ann.Sci.nat.
1835).
All the plates in the second Atlas are stamped "Biblio-
theque Royale", except 46 that are marked "Biblio-
theque Nationale" (BN). Knowing what is included in
the first 9 livraisons, and from w hat we infer is included
in li\raisons 10-11 (confirmed b\ our source no. 8), we
deduce that the stamp "Bibliotheque Nationale" was
used for all the plates of livraisons 4-6, for almost all the
plates of livraisons 10-11 and for one plate of livraisons
7-9.
8 — An incomplete bound copy of the work present at
the National Museum of Wales, the title page of which
is "Histoire tiaturelle generale et particuliere des Mol-
lusques", contains the Introduction and general remarks
on Cephalopods by Ferussac, and 101 plates. It corre-
sponds perfectly to the first 11 livraisons, issued in 1834-
1835, as detailed in the preceeding paragraphs. Only one
discrepancy was observed: plate 25 of Octopus, printed
by Benard and stamped as BN, that we consider to be
part of the livraisons 10-11, is not included in this copy.
The National Museum of Wales also has a complete
copy with handwritten notes by Hovle, and a series of
77 plates, unbound, corresponding to an incomplete set
of livraisons 1 to 11. Plate 25 of Octopus is present in
this last series of plates.
9 — The complete copy present at The Natural History
Museum, London, bears two handw ritten notes. One, by
Sherborn on the first page of vol.1 (Text), dates the text.
Its content has been discussed above. The other, by
E.A.Smith on the first page of vol.11 (Atlas), says:
"Mr. Sherborn has been unable to discover anything def-
inite w ith regard to dates of publication of any of these
plates .
THE NAUTILUS, Vol. 107, No. 3
Table 1. Details of text livraisons, with all available information on the .sheets of each livraison, inclusion (P) or not in the BIMM
cop\, the stamped pages in the two complete works (I and 2) at the BN and the protiahle date of issue BR = Bibliotheque Royale;
BN = Bililii)tliei|ue Xationale.
Ni
umber
P = BIMM
Sheets
Pages stamps
of
sheets
cop\
Livraison
\ ear
1-6
1 BR 1-48
6
P
12
1839
7-12
49/BR1-96 BRl
5
13
1839-41
13-18
97/BRl + BR2-144/BR1
5
14
1840-41
a-c
I BRl-XXIV
19-20
145-160/BRl
5
P
15
1841
21-24
161/BR1-192'BR1
4
P
16
1841
25-29
193/BNl + BR2-232/BR1
5
P
17
1841
d-g
XXV/BRl-LVI
4
P
18
1842
30-34
233/BN1-272
5
19
1845
35-40
273/BN1-320
6
20
1848
41-45
321/8X1-361
5
21
1848
10 — An iiiconiplete bound copy of the work, from the
library ol the "Laboratoire de Biologie des Invertebres
marins et Malacologie" (BIMM) at the Museum National
d'Histoire Naturelle (MNHN) comprising: the introduc-
tion and preface by d'Orbigny (pp. I-LVI), the text p.
1-48 and 145-232, 30 plates, all coming from the same
printer (Imp. Lemercier, Benard et C).
We interpret this BIMM copy as corresponding to
livraison 12 (pp. 1-48) and livraisons 15-18 (pp. I-LVI;
145-232), i.e. five livraisons, each of 4 to 6 sheets of text
and 6 plates (livrai.son 17 is announced on back wrappers
as comprising 5 or 6 sheets of text and 6 plates). This
interpretation also takes into account the observations on
the other copies of the work.
There remain 12 plates of the Atlas that belong neither
to the first 11 livraisons nor to the BIMM copy. Printed
well before the text, they are thought to have been issued
in livraisons 13-14 (i.e. two livraisons of 6 plates each).
Thus all the plates were delivered before 1845 (cita-
tions in d'Orbigny, 1845). The livraisons 19, 20 and 21
include only text.
11 — "Notice analytique siir les travanx zoologiques el
paleontologiques de Mr.Alcide d'Orbigny" (1844).
D'Orbigny states that his manuscript was finished in 1839.
However, later, the death of the publisher interrupted
the printing of the text, and caused several trials. This
accounts for the gaps in the publication of livraisons from
1842 to 1845, and from 1845 to 1848.
TEXT
The data gathered from the incomplete BIMM copy,
together with the .stamps on the text in the two complete
copies of the BN have led us to propo.se that the text was
i.ssucd in 10 livraisons.
Livraison 12, pp. 1-48, 1839. This livraison is included
in the incomplete copy of the BIMM. The first page is
stamped "Bibliotheque Royale" in the BN. Its issue is
announced on the back wrappers of "Mollttsques tcr-
restres et fluviatiles" (VViiukworth, 1942). After three
years \\ itiiout aii\ liv raison being issued, it .seems obvious
that d'Orbigny, who had started to rewrite his manu-
script in 1837 (cf. Introduction p. Ill), had published the
initial part of it in this livraison. It also seems evident
that the Introduction, which cites works issued in 1840-
1841, cannot have been issued in 1839.
Livraison 13, pp. 49-96, 1839-1841. The stamp "Biblio-
theque Royale occurs on the first and last page of this
livraison, which constitutes 5 sheets. Further supporting
evidence is that this livraison is not included in the
incomplete BIMM copy. The announcements on the
back wrappers of "Molhisques terrestres et fluviatiles"
(Winckworth, 1942) state that 16 livraisons were issued
in 1839-1841. Lacking any further information, we date
livraison 13 as 1839-184L
Livraison 14, pp. 97-144, 1840-1841. The stamp "BiWiO-
theque Royale " is on p. 97 of both copies in the BN, and
also on p. 144 in one of these copies. The break at page
97 is thus confirmed twice. The incomplete BIMM copy
lacks pages 49-144, confirming the end of the livraison.
A reference to 1839, on p. 139, allows livraison 14 to be
dated 1 840- 184 1.
Livraison 15, Introduction, pp. I-XXI\', and text pp. 145-
160, 1841. This livraison is included in the BIMM copy,
and is stamped on p. I and p. 160 of the BN copy. Stamps
in the text corresponded to a livraison of only two sheets,
which seemed insufficient to us. When the first part of
the introduction is included, a livraison of 5 sheets is
obtained, stamped at the beginning and at the end. This
is consistent with the issue of the Introduction in two
separate livraisons (stamps on p. I and on p. XXV). Ref-
erences to publications issued in early 1841, as well as
indications on the abo\ e-quoted w rappers lead us to pro-
pose the date ot 1841 tor livraison 15.
Livraison 16, pp. 161-192, 1841. This livraison is in-
cluded in the BIMM copy. The BN copy is stamped at
the beginning (p. 161) and at the end (p 192). Livraison
16 was issued between two other livraisons ol 1841 and
is therelorc tlated 1841.
Livrai.son 17, pp 193-232, 1841 This livraison is
A. Tillier and R. Boucher-Rodoni, ]
[993
Page 101
Table 2. Pu
bhcation dates of the Atla
s pi;
ates, with identih-
Table 2. ('oiituiued
cation of the
= stamps on
correspond
the BibhotI
il1(J liv^'^icmi PI =
plate number; stp
ipies; BI = Biblio-
1
lilj^ liv
leque
Bihho
National
e cc
PI
sip
Li\ rai.son
Date
tlieqiie Impcnaie; ti.\ =
•tlieque .Nationaie
0
3
7-9
12/15-
18
March 1835
1839-42
PI
stp
Livraison
Date
4
Sepia 1
2
BN
12/15-
4-6
18
1839-42
Octopus
1
BI
1-3
(1834) 1826
March 1835
2
BI
1-3
22 Sept. 1834
BN
4-6
March 1835
3
BI
1-3
22 Sept. 1834
3
BI
1-3
22 Sept. 1834
3/2°
BI
1-3
22 Sept. 1834
3/2°
BN
4-6
March 1835
4
BI
1-3
(1834) 1826
3/3°
BN
4-6
March 1835
5
BI
1-3
22 Sept. 1834
4
7-9
March 1835
6
BI
1-3
22 Sept. 1834
4/2°
7-9
March 1835
6/2°
BI
1-3
22 Sept. 1834
5
BN
4-6
March 1835
6/3°
BI
1-3
22 Sept. 1834
5/2°
7-9
March 1835
7
BI
1-3
22 Sept. 1834
6
10-11
29 June 1835
8
BI
1-3
22 Sept. 1834
6/2°
10-11
29 June 1835
9
BI
1-3
22 Sept. 1834
7
7-9
March 1835
10
BI
1-3
22 Sept. 1834
8
7-9
March 1835
11
7-9
1835
9
10-11
29 June 1835
12
BI
1-3/4
-6
22 Sept. 1834
10
7-9
March 1835
13
BI
1-3/4
-6
22 Sept. 1834
11
BN
10-11
29 June 1835
14
BN
1-3/4
-6
22 Sept. 1834
12
BN
10-11
29 June 1835
15
BN
1-3/4
-6
22 Sept. 1834
13
BN
10-11
29 June 1835
16
BN
7-9
March 1835
14
12/15-
-18
1839-42
17
7-9
March 1835
15
13-14
1839-41
18
BN
10-11
29 June 1835
16
13-14
1839-41
19
BN
10-11
29 June 1835
17
12/15-
-18
1839-42
20
BN
10-11
29 June 1835
18
12/15-
■18
1839-42
21
BN
13-14
1839-41
19
12/15-
-18
1839-42
0 0
BN
10-11
29 June 1835
20
12/15-
-18
1839-42
23
13-14
1839-41
21
12/15-
■18
1839-42
24
BN
10-11
29 June 1835
22
12/15-
-18
1839-42
25
BN
10-11
29 June 1835
23
12/15-
-18
1839-42
26
12/15-
■18
1839-42
24
12/15-
■18
1839-42
27
13-14
1839-41
25
12/15-
-18
1839-42
28
13-14
1839-41
26
12/15-
-18
1839-42
29
13-14
1839-41
27
12/15-
-18
1839-42
Eledone
1
BI
1-3
22 Sept. 1834
Sepioteuthis 1
BN
4-6
March 1835
1/2°
BI
1-3
22 Sept. 1834
.">
BN
4-6
March 1835
2
BI
1-3
22 Sept. 1834
3
BN
4-6
March 1835
3
12/15-
18
1839-42
4
BN
4-6
March 1835
Argonauta
1
BI
1-3
22 Sept. 1834
5
BN
10-11
29 June 1835
1/2°
BI
1-3
22 Sept. 1834
6
12/15-
-18
1839-42
1/3°
BI
1-3
22 Sept. 1834
7
12/15-
-18
1839-42
1/4°
BI
1-3
22 Sept. 1834
Loligo IC
BI
1-3
22 Sept. 1834
1/5°
BI
1-3
22 Sept. 1834
lA
BN
10-11
29 June 1835
2
BI
7-9
March 1835
2
BN
4-6
March 1835
3
BI
7-9
March 1835
BN
4-6
March 1835
4
BI
7-9
March 1835
4
BN
4-6
March 1835
5
BI
7-9
March 1835
5
BN
4-6
March 1835
6
12/15-
18
1839-42
6
7-9
March 1835
Bellerophon
1
BI
1-3
22 Sept. 1834
7
7-9
March 1835
2
12/15-
■18
1839-42
8
7-9
March 1835
3
12/15-
18
1839-42
9
7-9
March 1835
4
13-14
1839-41
10
7-9
March 1835
5
12/15-
18
1839-42
11
BN
4-6
March 1835
6
12/15-
18
1839-42
12
BN
4-6
March 1835
7
12/15-
■18
1839-42
13
BN
7-9
March 1835
Cranchia
1
2
BN
4-6
7-9
March 1835
March 1835
14
15
16
BN
BN
4-6
7-9
4-6
March 1835
March 1835
March 1835
Sepiola
1
BI
1-3
22 Sept 1834
17
BN
4-6
March 1835
Page 102
THE NAUTILUS, Vol. 107, No. 3
Table 2. C^oiiliiiiicd
PI
stp
Livraison
Date
18
10-11
29 June 1835
19
7-9
March 1835
20
13-14
1839-41
21
BN
10-11
29 June 1835
22
12/15-
IS
1839-42
23
13-14
1839-41
24
13-14
1839-41
Loligopsis
1
BN
4-6
March 1835
2
7-9
March 1835
3
7-9
March 1835
4
1.3-14
1839-41
On>choteuthis
1
Bl
1-3
22 Sept. 1834
2
BN
4-6
March 1835
3
BN
4-6
March 1835
3/2°
BN
4-6
March 1835
4
BN
4-6
March 1835
5
BN
4-6
March 1835
6
BN
10-11
29 June 1835
7
BN
10-11
29 June 1835
8
BN
10-11
29 June 1835
9
BN
10-11
29 June 1835
10
BN
10-11
29 June 1835
11
BN
10-11
29 June 1835
12
12/15-
■18
1839-42
13
12/15-
■18
1839-42
14
12/15-
■18
1839-42
Ommastrephes
1
12/15-
•18
1839-42
r>
12/ IS-
IS
1839-42
eluded in the BIMM copy. Both BN copies are stamped
at the beginning (p. 193), and one of them also at the
end (p. 232). It is comprised of 5 sheets. The back wrap-
pers of "Molln.sqties terrestres et fluviatiles (1841), state:
"la 17eme livraison sera raise en vente d la fin d'aout"
(Winckworth, 1942:35). The date and content of this
Hvraison are given as 5 to 6 sheets of text and 6 plates.
Our conclusions concerning this livraison contradict
Winckworth, who disbelieved the announcement on the
back wrappers.
Livraison 18, Introduction pp. XXV-LVI, 1842. Page
XXV is stamped in the BN copy. This livraison is included
in the BIMM copy. This second part of the Introduction
comprises 4 sheets. We include it in livrai.son 18 since
we know that livraison 17 was made ol 5 or 6 sheets.
The date 1842 is given by Louandre and Bourquelot
(1848) and is supported by five of our sources.
Livraison 19, pp. 233-272, 1845. This livraison is not
included in the BIMM copy. It is stamped on the first
page (p. 233) in the BN copy. The date 1845 is based on
the fact that livraison 19 is not mentioned as issued in
the analyses of the "Utterature franqaise" of the period
from 1827 to 1844 (Louandre & Bourquelot, 1848), but
the pages of this livraison are (luoted in "Mollusqiics
vivants et fossiles" d'Orbigny (1845).
Livraison 20, pp 273-320, 1848. This livraison is not
included in the BIMM copy. It is stamped on the first
()age (p. 273) in the BN copy. A different type font was
used for the pagination, starting on p. 273 (especially the
5's), References to the "Mollusques Vivants et Fossiles"
(1845) appear in the te.xt, confirming that this livraison
was published after 1845. It was probably issued with
the 21st livraison in 1848.
Livraison 21, pp. 321-361, 1848. This livraison is not
included in the BIMM copy. The BN copy is stamped
on the first page (p. 321). "Mollusques Vivants et Fos-
siles" (1845) is cited in this livraison. It is the last livraison,
and is undoubtedly dated 1848.
ATLAS
The "Bulletin Zoologique" and the "Annales des Sci-
enees Naturelles ' list all the species figured in the plates
issued in livraisons 1-9. The incomplete copy at Cardiff
gathers the first eleven livraisons, as detailed in sources
2, 3, 8 (see above). Plates included in the BIMM copy
(source 10; livraisons 12 and 15-18) could not be dated
more precisely than 1839-1842.
Details concerning the dates of publication and the
corresponding livraison of each plate are summarized in
table 2. Our conclusions concerning the issue ot the Atlas
are as follows:
Livraisons 1-3, 22 September 1834: 30 plates.
Livraisons 4-6, March 1835: 29 plates (4 of which were
already issued in livraisons 1-3).
Livraisons 7-9, March 1835: 25 plates.
Livraisons 10-11: 29 June 1835: 22 plates.
Livraison 12, 1839: 6 plates.
Livraisons 13-14, 1839-1841: 12 plates.
Livraisons 15-18. 1841-1842: 24 plates.
Two plates issued in livraisons 1-3 (Octopus pl.l and
4) had been distributed as early as 1826 by d'Orbigny
hiuLself (footnote p. 18: "A la fin de 1825, nous avons
fait lithographier nos planches representant les figures
de I'Octopus Cuvierii et de VO.Lcchcnaultii. et elles out
ete distribuees a beaucoup de personnes ).
CONCLUSIONS
A comparative analysis of Ferussac and d'Orbigny s work
and that of d'Orbigny (1845) confirms that the dates
iiuoted by d Orbigny are not reliable. As an example,
Cranchia plate 1 includes three species: C. cardioptera,
C. minima and C. scahra. In d'Orbigny (1845), they are
cited as figured in plate 1, and dated respectively 1826
(p. 390), 1830 (p. 351 ) and 1839 (p. 240). Such inaccuracy
is common in d'Orbigny s dates, and was often the case
at that time.
Our bibliographic searches allowed us to acertain the
precise dates of i.ssue of the 102 plates of the .Mlas that
were i.ssued in 1834 and 1835, as well as their compo-
sition. The dates of issue of the remaining plates are
1839-1842 (livraisons 12-18), but we can find no details
on the actual content of each livraison.
A. Tillier and R. Boucher-Rodoni, 1993
Page 103
In the publication of the text, we recognize three main
stages:
1. 1834-1835. Livraisons 1-11: introduction and gen-
eral remarks by Ferussac. 102 plates of the Atlas. No
descriptions.
2. 1839-1842. Livraisons 12-18: Introduction and gen-
eral remarks by d'Orbigny. Text pp. 1 to 232. Atlas: the
remaining 42 plates.
3. 1845-1848. Livraisons 19-21: last part of the text,
pp. 233-361. No plates.
The above data and conclusions are summarized in
two tables. Table 1 provides information concerning the
livraisons of text and corresponding dates. Table 2 pro-
vides the publication date for each plate, and associates
the plate with a corresponding livraison.
ACKNOWLEDGMENTS
We gratefully acknowledge the help and advice of
Dr.PMordan, of The Natural Histor\ Museum, London,
throughout our bibliographic search, and for critical re-
view of the manuscript. Special thanks to Mr.J.R.Kenyon,
librarian of the National Museum of Wales, who pro-
vided most helpful documentation. We are grateful to
our colleague Alain Foubert for his help in preparing
the tables.
LITERATURE CITED
D Orbigny, A 1844. Notice analytique sur le travau.x zool-
ogiqueset paleontologiquesde M. Aicided Orbigny. Impr.
Cosson, Paris, 48p.
D Orbigin , .\. 184.5-1847. Mdilu.sqiit-s \i\ants et tossilt-s, ou
description dc toutes ies especes de coiiuiiles et de Moii-
usques .-K Deialiass Libr , Paris 605p , :35 pis.
D'Orbign\. .\ 1856 Notice analytique sur Ies travaux de
geologie. de paleontoiogie et de zooiogie de M. Alcide
d'Orbigny, 1823-1856. Impr. L.S. Crete, Corbeil. 60pp.
Ferussac, A. E. de. 1831. Bulletin des Sciences Naturelles et
de Geologie, 2e section, 24: 373-374
Ferussac, A. E, de and A. d'Orbign\. 1834-1848. Histoire
naturelle generate et particuliere des Cephalopodes ace-
tabuiiferes vivants et fossiles. J.-B Bailliere Libr., Paris.
Tome Premier, — Texte: Ivi -I- 361p Tome Second. — .At-
las: 144 pis,
Sherborn, C. D. 1905. Ou tlie dates ot publication of d Or-
bigny's "Moll. viv. et loss. ", "Paieont. univ." and "Paleont.
etrangere". Journal of ConchoJogy, 11 (6): 169-170,
Sherborn, C. D 1922-32 Index Animalium sive index nom-
inum quae ab AD MDC.'CLVIII generibus et speciebus
animalium imposita sunt Sectio Secunda A kaiendis la-
nuariis, MDCCX4 usque ad finem Decembris, MDCCCL.
British Museum (Natural Histor\ ), Loudon, cxlvii -I- 7,056
+ 114p
Sherborn, C. D. and B. B. Woodward, 1901 On the dates of
publication of the "Histoire naturelle generale et parti-
culiere des moilusques terrestres et fluviatiles and the
"Tableaux systematiques des Animaux moilusques", by the
Barons Ferussac and G, P, Deshayes Annals and Magazine
of Natural History, 7 (8): 74-76,
VVinckworth, R 1942 Notes on the publication of Ferussac
and Orbigny s Histoire des Cephalopodes. Proceedings of
the MaiacoJogical Society. 25 (1): 34-36.
THE NAUTILUS 107(3):104-106, 1993
Page 104
Alvania valeriae (Gastropoda: Rissoidae), a New Species
from off Southeastern Brazil
Ricardo Silva Absalau
Dcpailanu'nto de Zoologia
Institiilo df Biologia
Univcrsidade F"edt'ral do Km
Janeiro
lllia do FundSo, 21949
Kio de Janeiro, RJ. Brazil
ABSTRACT
Alvania (Pimctulum) valeriae new species, is found off the
coast of Rio de Janeiro and Espirito Santo States, soutlieastern
Brazil. It differs from other species of Alvania occurring off
the Brazilian coast in having more numerous axial ribs (about
•36 on the body whorl). Axial ribs are more pronounced than
the 12-14 spiral threads, and nodules are not formed at their
intersections.
Key words: Gastropoda, Rissoidae, Alvania. Brazil
INTRODUCTION
Alvania (Punctiilum) valeriae new species was collected
on the continental shelf off Rio de Janeiro and Espirito
Santo States, in ciredgings taken during the oceanograph-
ic operations Espirito Santo I (July to September, 1984),
and (>abo P'rio VII (March to July 1983), as part of routine
sampling by the Brazilian Navy to obtain basic ocean-
ograpliic data on the coastal and oceanic regions off Bra-
zil. The malacofauna found during these operations was
characterized by numerous small mollusks, among which
the Rissoidae were prominent (Absalao, 1989).
The first descriptions of rissoids that occur off Brazil,
or would later have their ranges extended to Brazilian
waters, are those of d'Orbigny (1842), Watson (1886),
who studied the material collected by the H.M.S. Chal-
lenger, and Dall (1889), who worked with material col-
lected by U.S.S. Blake (Romer & Moore, 1988). Since
1966, malacological material from off the Brazilian coast
has been obtained on a more regular basis, thanks to the
dredging operations of the Brazilian Navy (Absalao, 1986).
The ininiit<- specimens (< 5 mm) have received almost
no attention. The Rissoidae especially "have proved to
be very difficult to classify at all levels because of their
small size, diverse form, and confusion caused by con-
vergence, especially in shell characters" (Fonder, 1985).
Ahbcitt (1974) stressed that the entire group was in need
of revision before species could be assigned to subgenera.
The revision of Coan (1964) was mainly a bibliographic
compilation. Ponder (1985) made the last and most com-
plete revision of this family at the generic level.
ABBREVIATIONS
IBUFRJ = Instituto de Biologia da I niversidade
Federal do Rio de Janeiro, Brazil.
MORG = Museu Oceanografico da Funda^ao Uni-
versidade de Rio Grande, Brazil.
USNM = National Museum of Natural History,
Smithsonian Institution, USA.
MZUSP = Museu de Zoologia da Universidade de
Sao Paulo, State of Sao Paulo, Brazil.
MNUFRJ = Museu Nacional do Rio de Janeiro, State
of Rio de Janeiro, Brazil.
MNHN = Museum National d'Histoire Naturelle
(Paris), France.
SYSTEMATICS
Family Rissoidae H & A. Adams, 1854
Genus Alvania Risso, 1826
Type species: Turbo cimex Linnaeus, 1758.
There are three previous!) described species of Al-
vania that occur off the Brazilian coast: A. auheriana
(Orbigny, 1842), A. deliciosa Jeffreys, 1884 and A. xan-
thias (Watson, 1885). The first occurs in shallow waters
and the latter two at greater depths.
Alvania {Puncluhim) valeriae Absalao, new species
(figures 1-3)
Description: Shell minute, 1.6 to 2.4 mm long, white
ovate-conic, with rounded whorls. Suture well impressed,
slightly canalicuiated. Pri)toconch multispiral, smooth.
R. S. Absalao, 1993
Page 105
with 2'/2 whorls. Apical angle 52-65° Teleoconch with
about 3(i gently curved axial ribs and 12-14 spiral threads
on body whorl. Axial ribs override spiral threads without
forming nodules at intersections. Base rounded with 6-
8 spiral threads. Intritacalx present over entire teleo-
conch. Terminal \ari\ absent. L'mbilicus small, chink-
like, well defined. Operculum and ratlula unknown.
Type locality: Off southeastern Espirito Santo State,
southeastern Brazil (2U°3S'12"S, 40°16'12"W), 38 m depth,
22.5''C and 36.7 ppt salinity, Brazilian Naval Research
Vessel Almirante Saldanha, station 6449, August 23,
1984.
Type material: Holot\ pe IBUFRJ 1726, 1.74 mm height,
1.02 mm width. Parat>pe 1, MORG 26986 off north-
eastern Rio de Janeiro State, Brazil (23°08'18"S,
41°03'18"W), 85 m depth, 16.7°C and 36.2 ppt salinity,
Almirante Saldanha, station 6417, August 29, 1984,
1.80 mm height, 1.04 mm width. Paratype 2, USNM
860503, off northeastern Rio de Janeiro State, Brazil
(22°35'00"S, 40°50'00"W), depth 77 m, 19.2°C and 36.1
ppt salinity, Almirante Saldanha, station 6422, August
16, 1984, i.80 mm height, 1 04 mm width. Paratype 3,
MZUSP 27716, off Rio de Janeiro State, Brazil (23°16'50"S,
43°02'40"W), Almirante Saldanha station 6174, depth
92 m, 15.6°C and 35.5 ppt salinity, March 29, 1983, 1.80
mm height, 1.04 mm width. Paratvpe 4. MNUFRJ 5978,
off Rio de Janeiro State, Brazil (23°16'.50"S, 43°02'40" W),
Almirante Saldanha station 6174, depth 92 m, 15.6°C,
and 35.5 ppt salinity, March 29, 1983, 2.40 mm height,
1.28 mm width. Paratype 5, IBUFRJ 1727. off Rio de
Janeiro State, Brazil (23°16'50"S, 43°02'40"W), Al-
mirante Saldanha station 6174, depth 92 m, 15.6°C,
and 35.5 ppt salinity, March 29, 1983, 2.16 mm height,
1.20 mm width. Paratype 6. MORG 27681, off Rio de
Janeiro State, Brazil (23°16'50"S, 43°02'40"W), Al-
mirante Saldanha station 6174, depth 92 m, 15.6°C,
and 35.5 ppt salinity, March 29, 1983, 1.88 mm height,
1.04 mm width. Paratype 7, MORG 27682, off Rio de
Janeiro State, Brazil (23°16'50"S, 43°02'40"W), Al-
mirante Saldanha station 6174, depth 92 m, 15.6°C,
and 35.5 ppt salinity, March 29, 1983, 1.80 mm height,
1.04 mm width. Paratype 8, IBUFRJ 1728, off Rio de
Janeiro State, Brazil '(23°16'50"S, 43°02'40"W), Al-
mirante Saldanha station 6174, depth 92 m, 15.6°C,
and 35.5 ppt salinity, March 29, 1983, 2.00 mm height,
1.12 mm width. Paratype 9, MNHN, off Rio de Janeiro
State, Brazil (23°16'50"S, 43°02'40"W), Almirante Sal-
danha station 6174, March 29, depth 92 m, 15.6°C, and
35.5 ppt salinity, 1983, 1.84 mm height, 1.08 mm width.
Etymology: This species is dedicated to my wife, Mrs.
Valeria Gomes Veloso.
Range: Records of A. valeriae are restricted to the re-
gion between northern Rio de Janeiro State and southern
Espirito Santo State, southeastern Brazil.
Discussion: Alvania colomlriana is compared to A. val-
eriae because both species occur in the same biogeo-
graphic region. However, they are not sympatric. As
i
Figures 1-3. Alvania valeriae new species. 1. Holotvpe.
IliL'FRJ 1727; 120X, 2. Ornanu-ntatiDn details, 240x,' 3.
Protoconch, 240 x.
Alvania is largely a European group, comparisons with
those warm-water species were included
Alvania auheriana. A. deliciosa and A. xanthias have
respectively 14, 14 and 12-14 axial ribs, whereas A. vale-
riae has 32-40 ribs. This character alone is sufficient to
discern it from the other species occurring in Brazilian
Page 106
THE NAUTILUS, Vol. 107, No. 3
waters. In addition, /\. auberiana lias 5 spiral threads on
its teleoconch, A. deliciom has 10, A. xanthiaa lias only
4 spiral threads that are restricted to the base, whereas
A. valcriae has 12-14 spiral threads (figures 1 and 2) on
its teleoconch.
Alvania colombiana Romer and Moore, 1988 differs
from A. valeriae in having only 26 axial ribs, which
disappear on the base of the shell, and 7-9 spiral grooves.
Alvania valeriae has =36 (32-40) a.xial riblets that extend
onl\ slightly over the body whorl without reaching the
umbilical region of the shell, as well as 12-14 spiral
threads.
Alvania porcupinae (Jeffreys, 1884) has a smooth pro-
toconch very similar to that of A. valeriae, =32 axial
ribs that disappear smoothly towards the umbilical re-
gion, while A. valcriae differs in having =36 (32-40)
axial ribs that enter a little beyond the edge of the pen-
ultimate whorl. The diameter of the last two whorls
increases greatly in A. porcupinae, but little in A. valeri-
ae.
Alvania zylensis Gofas and Waren 1982, and A. im-
perspicna Palary, 1920 can be easily distinguished from
A. valeriae on the basis of protoconch sculpture. The
protoconch of A. valeriae is smooth, that of A. zylensis
has 12 zigzag spiral lines that intersect, forming a retic-
ulated pattern, while that of A. imperspicua has 7-8
spiral lines that also zigzag but do not intersect. In ad-
dition, A. imperspicua has protruding nodules on the
teleoconch, which are formed by the junction of axial
riblets and spiral lines. In contrast, A. valeriae lacks such
nodules.
Although A. valeriae is similar in shape to A. coni-
pacta, this latter species has fewer axial riblets (20-30)
and spiral threads (6-8) than the former, which has =36
(32-40) axial riblets and 12-14 spiral threads. Alvania
compacta also has nodules where axial and spiral orna-
mentation cross.
Alvania valeriae is unicjue among its southwestern At-
lantic congeners in having a large number of axial ribs
that clearly dominate the 12-14 spiral threads.
ACKNOWLEDGMENTS
1 am grateful to Prof. E. C. Rios from the Museu Ocean-
ografico da Funda^ao Universidade do Rio Grande, who
has provided bibliographic material. I am also much
obliged to the Laboratorio de I'ltraestrutura Olular e
Microscopia Eletronica do Instituto de Hiofisica Carlos
Chagas Filho, of the Universidade Federal do Rio de
Janeiro for the SEM micrograph, and to Dr. Jose Hen-
rique Leal, from the Rosenstiel Institute of Marine Sci-
ences, and to Dr. M. G. Harasewych from the Smith-
sonian Institution for their comments on the manuscript.
This work was partially supported by CNPq. grant no.
802240.87-9.
LITERATURE CITED
Abbott, H T 1974 ,\nierican Seasheiis, 2rnl cd \'aii .NostraiKl
Reinhoid, N.Y., 663 p.
.\bsalao, R. S. 1986 Moluscos da Comissao Oceanografica
Geocosta Rio I, RJ, Brasil Revista Brasileira de Biologia
46(1):27-31.
Absalao, R. S. 1989. Padroes distributives e zoogeografia dos
moluscos da plataforma continental brasileira. Parte III.
Comissao Oceanografica Espirito Santo I. Memorias do
Instituto Oswaldo C^ruz 84, Suplemento I\ '1-6
C^oan, E. 1964. A proposed revision of the Rissoacean families
Rissoidae, Rissoinidae, and Cingulopsidae (Mollusca:Gas-
tr(ipoda). The \'eliger 6(3)464-171.
Dall, W, H. 1889, Reports on the results of dredging in the
Gulf ol Mexico (1877-78) and in the Caribbean Sea (1879-
80) by the U.S. Coast Survey Steamer "Blake". 29 Report
on the MoUusca, part II. Gastropoda and Scaphopoda.
Bulletin of the Museum of Comparative Zoology IS 1-
492, pis. 10-40.
d'Orhigny, A. 1842. Mollusques, X'of 1 (part). In: R, de la
.Sagra (ed). Histoire Plusique, politique et naturelle de
rile de Cuba. Bertrand. Paris, p 209-26-1 + .\tlas. pis. 1-
28 4- II his.
Gofas. S. and A. Waren. 1982. Taxoriomie de yueiques Es-
peces du Genre Alvania (Mollusca, Gastropoda) des Cotes
Iberiques et Marocaines, Bolletino Malacologico 18(1-4):
1-16,
Ponder. W, F. 1985, A review of the Genera of the Rissoidae
(Mollusca: Mesogastropoda: Rissoacea). Records of the
.Australian Museum, Su|)plenieiit 4:1-221.
Rios, E. C, 1985. Seasheiis of Brazil. Museu Oceanografico
da Funda9ao I'niversidade de Rio Grande, 331 p. + 102
plates.
Romer, N. S. and D. R. Moore. 1988. A new species of A/t)ania
(Rissoidae) from the West Indian Region The Nautilus
102(4): 131-133.
Watson, R. B 1886, Report on the Scaphopoda and Gaster-
opoda collected In HM.S. Challenger during the \ears
1873-1876, Voyage of the H,M,S, Challenger. Zoolog> 15:
1-675; appendix A, p, 677-680; geographical distribution,
p 691-722; index, p, 723-75«; pis, 1-50,
INSTRUCTIONS TO AUTHORS
THE NAUTILUS publishes papers on all aspects of the
biology and systematics of mollusks. Manuscripts de-
scribing original, unpublished research as well as review
articles will be considered. Brief articles, not exceeding
1000 words, will be published as notes and do not re-
quire an abstract. Notices of meetings and other items
of interest to malacologists will appear in a news and
notices section.
Manuscripts: Each original manuscript and accompa-
nying illustrations should be submitted in triplicate. Text
must be typed on one side of 8'/2 x 11 inch white paper,
double spaced throughout (including literature cited, ta-
bles and figure captions), with at least 1 inch of margin
on all sides. All pages must be numbered consecutively.
If printed on a word processor, the right margin should
be ragged rather than justified. Authors should follow
the recommendations of the Council of Biology Editors
Style Manual, which is available from the Council of
Biology Editors, Inc., 9650 Rockville Pike, Bethesda, MD
20814, U.S.A. The first mention of a scientific name in
the text should be accompanied by the taxonomic au-
thority, including year. Latin names and words to be
printed in italics must be underlined; leave other indi-
cations to the editor. Metric and Celsius units are to be
used.
The sequence of sections should be: title page, abstract
page, introduction, materials and methods, results, dis-
cussion, acknowledgements, literature cited, tables, fig-
ure captions, figures. The title page should include the
title, author's name(s) and address(es). The abstract page
should contain the title and abstract, which should sum-
marize in 250 words or less the scope, main results and
conclusions of the paper. The abstract may be followed
by a maximum of 8 key words. All references cited in
the text must appear in the literature cited section and
vice versa. In the literature cited section, all authors
must be fully identified and listed alphabetically. Follow
a recent issue of THE NAUTILUS for bibliographic style,
noting that journal titles must be unabbreviated. Infor-
mation on plates and figures should be cited only if not
included in the pagination. Tables must be numbered
and each placed on a separate sheet. A brief legend must
accompany each table. Captions for each group of illus-
trations should be typed on a separate sheet and include
a key to all lettered labeling appearing in that group of
illustrations.
\\\ line drawings must be in black, high quality ink,
clearly detailed and completely labeled. Photographs
must be on glossy, high contrast paper. All figures are
to be consecutively numbered (figs. 1, 2, 3, ... , NOT
figs. la. lb, Ic, . . . NOR plate 1, fig. 1 . . .). Illustrations
must be arranged in proportions that will conform with
the width of a page (6% inches or 171 mm) or a column
(3'/4 inches or 82 mm). The maximum size of a printed
figure is 6% by 9 inches or 171 by 228 mm. All illus-
trations must be fully cropped, mounted on a firm, white
backing, numbered, labeled and camera ready. The au-
thor's name, paper title and figure number(s) should ap-
pear on the back. Original illustrations must be between
one and two times the desired final size. It is the author's
responsibility that the line weight and lettering are ap-
propriate for the desired reduction. Original illustrations
will be returned to the author if requested. Color illus-
trations can be included at extra cost to the author.
Voucher Material: Deposition of type material in a
recognized public museum is a requirement for publi-
cation of papers in which new species are described.
Deposition of representative voucher specimens in such
institutions is strongly encouraged for all other types of
research papers.
Processing of Manuscripts: Upon receipt, every manu-
script is acknowledged and sent for critical review by at
least two referees. These reviews serve as the basis for
acceptance or rejection. Accepted manuscripts are re-
turned to the author for consideration of the reviewers'
comments. A finalized version of the manuscript is re-
turned to the editor and sent to press. Two sets of proofs
are sent to the author for correction. Changes other than
typesetting errors will be charged to the author at cost.
One set of corrected proofs should be sent to the editor
as soon as possible. Authors with institutional, grant or
other research support will be billed for page charges at
the rate of $60.00 per printed page.
An order form for reprints will accompany the proofs.
Reprints may be ordered through the editor.
Manuscripts, corrected proofs and correspondence re-
garding editorial matters should be sent to: Dr. M.G.
Harasewych, Editor, Division of Mollusks, NHB stop 118,
National Museum of Natural History, Smithsonian In-
stitution, Washington, DC 20560, USA.
@ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).
rHE NALTILLS
Volume 107, Number 4
March 16, 1994
ISSN 0028-1344
A quarterly devoted
to malacology.
A*^w ----
EUITOR-IN-CHIEF
Dr. M. G. Harasewych
Division of Mollusks
National Museum of
Natural History
Smithsonian Institution
Washington, DC 20560
ASSOCIATE EDITOR
Dr. R. Tucker Abbott
American Malacologists, Inc.
P.O. Box 2255
Melbourne, FL 32902
CONSULTING EDITORS
Dr. Riidiger Bieler
Department of Invertebrates
Field Museum of
Natural History
Chicago, IL 60605
Dr. Robert T. Dillon, Jr.
Department of Biology
College of Charleston
C:harleston, SC 29424
Dr. William K. Emerson
Department of Living Invertebrates
The American Museum of Natural
History
New York, NY 10024
Dr. Robert Hershler
Division of Mollusks
National Museum of
Natural History
Smithsonian Institution
Washington, DC 20560
Mr. Richard I. Johnson
Department of Mollusks
Museum of Comparative Zoolog\
Harvard University
Cambridge, MA 02138
Dr. Aurele La Rocque
Department of Geology
The Ohio State University
Columbus, OH 43210
Dr. James H. McLean
Department of Malacology
Los Angeles County Museum of
Natural Histor\
900 Exposition Boulevard
Los Angeles, CA 90007
Dr. .Arthur S. Merrill
% Department of Mollusks
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138
Ms. Paula M. Mikkelsen
Harbor Brancli Oceanographic
Institution, Inc.
Ft. Pierce, FL 33450
Dr. Donald R. Moore
Division of Marine Geology
and Geophysics
Ro.senstiel School of Marine and
Atmospheric Science
University of Miami
4600 Rickenbacker Causeway
Miami, FL 33149
Dr. Gustav Paulay
Marine Laboratory
University of Guam
Mangilao, Guam 96923
Mr. Ridiard E. Petit
P.O. Box 30
North Myrtle Beach, SC: 29582
Dr. Edward J. Petuch
Department of Geology
Florida Atlantic University
Boca Raton, FL 33431
Dr. David H. Stansbery
Museum of Zoology
The Ohio State University
Columbus, OH 43210
Dr. Ruth D. Turner
Department of Mollusks
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138
Dr. Geerat J. Vermeij
Department of Geology
University of C^alifornia at Davis
Davis, CA 95616
SUBSCRIPTION INFORMATION
The subscription rate per volume
is US $25.00 for individuals, US
$40.00 for institutions. Postage
outside the llnited States is an
additional US $5.00 for surface
and US $12.00 for air mail. All
orders should be accompanied by
payment and sent to; THE
NAUTILUS, P.O. Box 7279, Silver
Spring, MD 20907-7279, USA.
Change of address: Please inform
the publisher of your new address
at lea.st 6 weeks in advance. All
communications should include
both old and new addresses (with
zip codes) and state the effective
date.
THE NAUTILUS (ISSN 0028-
1344) is published quarterly by
Trophon Corporation, 8911 Alton
Parkway, Silver Spring, MD 20910.
Second Class postage paid at Silver
Spring, MD and additional mailing
offices.
POSTMASTER: Send address
changes to; THE NAUTILUS
P.O. Box 7279
Silver Spring, MD 20907-7279
TH E€7NAUTI LUS
"Tine Bioloaica! Uboralofy/
,ole 0C3. ■.orraphic Institution y^/^^^ ^Qy J^umber 4
'""" March 16, 1994
IVIAR 2 5 1994 ISSN 0028-1344
CONTENTS
Woods Hois, MA Oi^t.
Robert Robertson Protoconch Size Variation Along Depth Gradients in a
Planktotrophic Epitonium 107
John I. Scheide Effect of Low Water Temperature on Ion Balance in the
Phyllis N. Bonaminio Zebra Mussel, Dreissena pohjjuorpha, and the Unionid
Mussel, Lampsilis radiata 113
Emily H. Yokes The Muricid T\ pes of Frank Collins Baker 118
J. P. Pointier Invasion of the Rivers of the Littoral Central Region of
R. N. Incani Venezuela by Thiara granifera and Melanoides
C. Balzan tuberculata (Mollusca; Prosobranchia: Thiaridae) and the
P. Chroseiechowski Absence of Biomphalaria glabrata, Snail Host of
S. Prypchan Schistosoma inansoni 124
Cristian F. Ituarte Corbicula and Neocorbicula (Bivalvia: Corbiculidae) in the
Parana, L'ruguay, and Rio de La Plata Basins 129
Book Review 136
THE NAUTILUS 107(4):107-112, 1994
Page 107
Protoconch Size Variation Along Depth Gradients in a
Planktotrophic Epitonium
Roberl Robertson
DfpartiiU'Hl ol Malacolog\
Tlif Acaiii-nn ol Natural Sciences
190(1 Benjamin Franklin Parkway
Philadclpliia, I'A 19103-1195, USA
ABSTRACT
The protoconch length of Epiluniiim t'chinaticuata ranges from
330 to 790 M'li '" tliP northern Bahamas. The length increases
along depth gradients from near 0 m to 40-52 m. Intermediates
are from intermediate depths. One planktotrophic species is
involved. A possible explanation is given. Svstematists should
be aware that the phenomenon ma\ occur in other plankto-
trophic epitoniids or other marine prosobranchs
Kill nurds: Epitiinuiiu. [imtoconch. size \ariation.
INTRODUCTION
". . . the protoconch may provide the best guide to rela-
tionships [within the Epitoniidae] yet. Of course ecological
factors . . nia> prove to influence protoconch form (as
has been established in other gastropod groups) but this
remains to be show n lor the Epitoniidae Kilburn. 19iS5:
There is scattered mentinii in the literature on marine
prosobrancli gastropods of one species having proto-
conchs that differ from shallow and "deep" water. None
of these cases seems to have been documented with mea-
surements, descriptions and illustrations. In some cases
different species perhaps were involved.
In this paper, I show that in a single planktotrophic
species of Epitonium much variation in protoconch size
occurs in the northern Bahamas. Size (length) increases
with increasing depth (and possibK also with exposure
to wave action). Populations from Grand i5aliama sam-
pled from 9, 10, 24, and 40 m, and from 1 I and 52 m
at Abaco all have larger protoconchs than do those from
nearby shallow sites (0-2 m).
The epitoniid studied is Epitonium echinaticosta [not
cchinaticostum] (d'Orbigny, 1842) [type locality: St.
Thomas, Virgin Islands; northern known limit: Bermuda
(.ANSP 100824); .southern know n limit: PYrnando de No-
ronlia, off Brazil (Leal, 1991)]. Because of its distinctive
shell morphology, especially its variably disjunct anil
commonly "open" teleoconch coiling (Rex & Boss, 1976),
and its 8-15 variably high and undulate axial ribs on the
last whorl (Robertson, 198.3b), Dall (1889) created for
this species a suligenus Cycloscala, which has sometimes
been ranked as a genus.
Species of Cycloscala. w hich occur in the Indo-Pacific
as well as the western Atlantic, have been separated by
degree of w horl disjunction and rib counts by de Boury
(1911), Woodring (1928), Kilburn (1985) and DuShane
(1990). Familiarity with a living population (Robertson,
1983a) clearly reveals that at least in the Bahamas there
is just one species, with intergrades between the extremes,
as concluded by C;lench and Turner (1951). They rec-
ognized only one western Atlantic species of Cycloscala,
and I concur.
In Bahamian £. echinaticosta, the frequencies of the
different teleoconch coiling types vary little at different
depths (Robertson, uniiublished) and thus do not affect
the conclusions here.
There is already some inlormation about the coelen-
terate host, feeiling and reproduction of £. echinaticosta
in the Bahamas (Robertson, 1983a). At least in shallow
water, the host is the actinian sea anemone Bunodeopsis
[or Viatrix] globulifera (Duchassaing, 1850), commonly
living on the leaves of turtle grass [Thalassia testudinum
(Banks and Solander ex Kiinig)] (It also occurs on Syr-
ingudium, Halimcda, Pcnicillus. and Acetahularia.) The
lower end of the bathymetric range of E. echinaticosta
is given by Abbott (1974) as 200 fathoms (366 m). Wheth-
er living aniinals occur at that depth, and whether the
Epitonium remains specific to B globulifera are un-
known. The Epitonium and anemone are most abundant
in the Freeport canals.
How is it known that E. echinaticosta is planktotroph-
icr* There is Leal s inulocumented statement that off Bra-
zil "planktotrophic ile\ elopment is indicated. More def-
initely, there is direct e\ idence of planktotrophv at least
in Bahamian shallow waters: eggs of this species have
Ijeen shown to be 98-106 ^m in diameter (mean = 102
^m; n = 10)(Robertson, 1983a). Two swimming veligers,
admittedK abnormal, were seen to hatch naturally from
egg capsules (Robertson, 1983a).
Page 108
THE NAUTILUS, Vol. 107, No. 4
Table I. l.()c-alit\ data, rt-positories of voui'lier.s, ualii {li-[)tli> and protocoiich leiigtlis of Epitonium echinaticosta from Grand
Haliaina, .Vbaco, Hiinini and the Berry Is., all northern Bahamas Data are arranged in order of inereasing depth .Abbreviations:
.WSP = The Acadenn of Natural Sciences of Philadelphia; CR = Clolin Kedfeni collection; GB = Grand Bahama; n = numbers of
specimens measurable (out of many unmpasurablei
Localities
Voucher
s
n
Depths
(m)
Lengths (^m)
(nos.)
Means
Ranges
ANSP .367 19.5 i
369'
■59
16
1
370
330-430
ANSP 370.521
2
0-2
380
370-380
ANSP .3672.34 &
367235
23
1
410
370-470
CR .3990
1
0-2
4.50
ANSP .32-429.3
2
0-2
450
440-460
ANSP .32556.3
0-2
460
CR 4.598
0-2
510
ANSP .372821
0-2
570
CR .398.3
0-2
590
ANSP .367 195
11
3-10':'
550
470-640
ANSP .370944 &
370.^
i21
9
640
ANSP 369074 &
369'i
•59
10
520
CR 76.38
11
650
610-740
ANSP .369476
24
670
ANSP 368803
40
660
{:R 529.5 & 5296
3
52
760
730-790
Running Moti Canal, (JB
26°29'45"N. 78°4r45"\V
North Hawksbill Creek, GB
26°32'00"N, 78°45'00"\\'
SiKer Coxp Canal. GB
26''.30'30"N, 78°39'30"\V
Treasure Cove, Abaco
26°42'00"N, 77°18'20"\V
(;hub Cav, Berr\ Is
25°25'N, 77°54'\\"
South Bimini
2,5°42'N, 79°17'W
Treasure Caw .\baco
26°40'00"N, 77°18'15"W
Settlement Point, West End,
GB 26°42'15"N, 78°.59'50"W
Sandy Point, .Abaco
26°0()'15"\, 77''24'00"W
Dead Man's Reef Beach, GB
26°34'45"N, 78°51'45"\V
Finders Point, GB
26°30'30"N, 78°46'30"W
Kings Bay. Dundee Ba\, GB
26°29T5"N, 7.S°43'.30"\V
Chub Rocks, Abaco
26°43'55"N, 77°13'05"W
Gold Rock, GB
26°.30'00"N, 78°22'0()"W
Tamarind, GB
26°30'45"N, 78°36'00"W
N, of N.VV. end (Great) Guana
Cav, Abaco 26°44'N, 77°09'W
"The hatching shell diameter is about 170 fim. i c much
larger than the egg, , . The protoconch is about 0.3 to
0.4 nun long [small ones], i.e it is much larger than the
hatching veliger shell - proof that the larva grows sub-
stantialK while in the plankton
(Robertson, iy8.'3a). There might be geographical vari-
aticni, but the data from Abaco as well as Grand Bahama
suggest that the larger proloconchs may grow in deeper
water every vviieri' the species occurs.
In the literature, there is already an indication of pro-
toconch size variation in another epitoniid. Two .SEM
photos of Epitonium dallianum (Verrili & Smith, 1880)
protoconchs (Bouchel & Waren, I98(r figs. 1180-1181)
show .some intrapopiilational shaix' and size variation. £.
(lallianiirn is deep-water and aniphi-.Mlantii- in distri-
bution.
Doubt has been cast on poecilogony (intraspecific vari-
ation in mode of reproduction) occurring in the Gas-
tropoda, especially by Hoagiand and Robertson (1988)
and Bouchet (1989). Poecilogony usually refers to plank-
totrophy versus lecithotrophy . By extension, lack of poe-
cilogony has been taken to mean that gastropod proto-
conchs are intraspecifically uniform in both form and
size. This paper calls in cjuestion size specificity.
MATERIALS AND METHODS
This study is based on 68 live-collected and empty shells
from shallow depths down to 40 m at southern Grand
Bahama island (mainly in the canals of Freeport), 3 emp-
ty shells from .52 m off northeastern .\baco, Bahamas, 9
maiiiK beachworn shells from shallower depths at the
same island, and a tew also from Bimini and the Berry
Islanils, all in the northern Bahamas, all but the last two
places on the Little Bahama Bank The .52 m station was
about .'3240 m horizontalK Irom the nearest shore.
R, Robertson, 1994
Page 109
Figures 1-2. SEM micrographs ot prolnKHK li and part nl tflfindiich (if: 1. Epitoniiini cchiiuilicDsta Irum less than 2 in (Ruiiiiiiig
Mini (laiial, Freeport, Grand Bahama), with ca 3,5 prntntonch whorls. Protoconch length = 410 ^in. Teleoconch whorls sutured.
2. £. cchinaticosta from .52 m (off Abaeo), with 5 protoconch whorls. Protoconch length = 770 /jm Note also the incised axial
grow th lines on both shells, an indication of planktotroph\ Teleoconch coiling of 2 was ojjen. increasing the measured protoconch
length onK slight!). Scale bar = 100 m"! for both micrograplis.
Protocoiielis w ere studied w ith light and scanning elec-
tron microscopy. Measurements were made to the near-
est 10 fim w ith the aid of a cahbrated ocular micrometer
in a Wild dissecting microscope at x,50.
The anterior end of the larval shell is obscured to a
var> ing degree b\ the first teleoconch whorl, which may
or may not have disjunct coiling begining before, at, or
after the protoconch vari.x. The length measuremenL
were made on the exposed part of the protoconch, i.e.
as far down as the protoconch varix (smaller than the
first teleoconch rib) can be seen. Where the teleoconch
becomes disjunct affects the length data slightly, but does
not account for the differences in protoconch size re-
ported here. Protoconch Is and first protoconch whorl
diameters were too corroded to be measurable accurateK
(but a size estimate is given below under Results!
The originalK '3 deep-water shells from Abaco are in
the private collection of Mr. Colin I^edfern (one was lost).
Nine other \()ucher specimens from .\baco also belong
to Redfern. Two other lots, from Bimini and the Berry
Islands, are at The Academy of Natural Sciences of Phil-
adelphia (ANSP). The rest, all from Grand Bahama and
the Jack N. Worsfold Collection, are also at ANSP.
Data on each population studied are gi\en in Table
1. No temperature or rearing data are available. The
data are graphed on figure 4.
RESULTS
First whorl diameters ol both the small and large pro-
toconchs are 100-140 /jm (the\ approximate the egg
sizes), and whorl numbers are 3.5 to 5. Whole protoconch
\\ idths of short and long larval shells are all 270-310 fiin.
Long and short protoconchs are similarly shaped (figures
1-2). All specimens have the usual planktotrophic Epi-
toniu/n protoconch microsculpture (incised axial lines).
Protoconch lengths var\ remarkabK ; from 330 to 990
^m. The smaller protoconchs have 3.5 to 4 whorls (figure
1) and the larger ones about 5 whorls (figure 2). Thus all
are multispiral, another indication of planktotrophy. On
the larger protoconchs, whorl ruimbers have increased,
not whorl sizes.
The data from the two Freeport canal populations
show that all protoconchs are very short but appear to
be slightly different from each other in size, with pop-
ulations from Running Mon having protoconchs no Ion-
Page 110
THE NAUTILUS, Vol. 107, No. 4
19^
77<
27<
26<
NORTH ATLANTIC OCEAN
LITTLE BAHAMA BANK
50 km
Settlement Point
West End
Dead Man's Reef Beach
North HawksbiU Creek ^ ^
"V 17 C
to
<
Q
M
«
o
►J
a.
A
\
■ South Bimini
GREAT BAHAMA BANK (NORTHERN END
27 =
26 =
Figure 3. Map ol llic iioitlicr ii li;iliaiiuis, sliouiiit; llic Little Ikiliani,! Bank. |)art (il the (in-at Baliaiiia Hank ami lli<' Idealities in
Talile I
ger than 430 ^ni and tlidSf ftoni Silver Cove no longer
than 470 juin.
The population sampled from Dead Man s Reef Reach
is puzzling because it ranges in protoconch length from
470 to 640 ixn\, o\('rla|)[)ing the largest shallow water
sliell Ironi elsew here and the smallest "deep" water shell.
\ possible explanation for the Reef anoinaK is that the
other two shallow localities are in sheltered canals, and
the Dead Man's Reel is on the exposed south coast —
where even near the surface, temperatures may be lower,
exposure to wave action max l)e greater, and hosts may
be more sparsely disper.sed. The beach shells may have
washed in from an estimated de|)th of 3-10 rn bexond
the near-beach reel
Judging by the Chub Rock and i'inders Point samples,
enlargement can occur in water as shallow as 9 and 1 1
m (Table 1). The Sandy Point shell has a moderatelx
large [irotoconch (.590 /jni long) which can be explained
by the presence of very deep w ater nearbx Un the North-
east Providence Channel, figure 3), where greater ex-
posure, colder shallow water and perhaps sparsely dis-
persed hosts can be expected.
The smallest .Abaco protoconch (430 /im) is larger tliaii
the smallest Crand Bahama protoconch (330 ^m). The
Freeport canals have esf)eciall\ dense populations ot
Bunoclcopsis (the anemone host).
DISCUSSION
Protoconch size bimodalitv in the .Vrchitectonicidae is
|)r()babl\ genetic in origin (Robertson, 1970), Since the
protoiiinch enlargement (il Rahamian V.pitonium echin-
R. Robertson, 1994
Page 1 1 1
2
4 3 4
• • • '
7 2 6 3 3 2
• ••••••
800
Epitonium echinaticosta PROTOCONCH LENGTH (jim)
Figure i. Epiliiniiim fcliiiialUDsln |iriil(KijrKli It-nullis vtrsus water (lepllis. Data troni Tahle 1. The spoli with iiuinbers .sliuvv
wliere two nr iimir data pniiits eciiiKule
olicosta at "depth occurs in two areas more than 100
km apart (figure 3), an ecological rather than a genetic
explanation seems necessary.
The factors most likely to account for the size \ ariation
of lar\al Epitoniinn cchinaticosla are:
1 Duration of the pelagic lar\a! stage. Mo\ement to
and fro by titlal and ocean currents.
2. Diurnal \ertical migration of larvae such as re-
ported in Mediterranean Epitonium spp. by Rich-
ter (1973, as Scalaria), who also recorded vertical
distances traversed i)\ other larval taxa in the order
of 100 m
3. Fast lar\ al grow th in the w arm w aters of the canals,
slower grov\ th near the cooler surface elsewhere,
and slowest growth at "depth" in the coolest water.
Scheltema (1967) and Pechenik (1984) showed ex-
perimentally that the larval growth rate in ////-
anassa obsoleta (Say, 1822) and Crepidula jurni-
cata (Linnaeus, 1758), two other prosobranchs.
responds to temperature in this manner.
4. In the absence ot metamori^hic cues, a long h\-
pothesized delay period and continuing grow th af-
ter the onset of competence. According to Pechenik
(1986), the lar\al prosobranch Bittiuin allcinatiint
(Say, 1822) can grow from 2.8-3.0 whorls to 3.7-
3.8 whorls during the delay periotl. The minimum
and maximum larval shell lengths at metamorpho-
sis for Crepidula jurnicata are 700 ^m and 2300
;um (Pechenik, 1984).
The great abundance of Bunodeopsis glohulijcra, the
sea anemone host of Epitonium echinaticosta, in the
shallow, warm Freeport canals (the artificial equivalents
of mangrove creeks) must be mentioned. Nearly all of
the live-collectetl epitoniin7is are from the canals There
there is a large reproducti\e population of the Epitonium
and it can be hypothesized that many ot the lar\ ae grow
last (in the warm water), swim near their parents, being
moved back and forth by tidal currents, and settle and
metamorphose promptK upon becoming competent (at
small size, 3.5 whorls).
Outside the canals, the abundance ot Buncodeopsis is
not known. It is even possible that £. echinaticosta has
one or more otlier coelenterate hosts at "depth." Some
ot the larvae in the sea could have come out of the canals
but it is also probable that many were spawned outside.
These larvae (from both sources) can be expected to drift
into "deep" water and to undergo diurnal vertical mi-
gration, v\ hich would involve intermittent slow growth
at "depth (cool water), lateral movement in currents,
and difficulty in finding host(s). The duration of the com-
petent period is believed long, and large (5-whorled)
larxae result.
Thus the duration of the pelagic larval stage is here
considered the most likeK factor bringing about the pro-
Page 112
THE NAUTILUS, Vol. 107, No. 4
toconch size variation of E. echinaticosta. But factors 2-
-4 listed above must also be iinoKod.
CONCLUSIONS
How widespread is this kind ot variation among other
mollusks? Partly allopatric and partK heterochronous
size \ariation in the "protoconth' ot the thecosome
pteropod Clio pyraniidata Linnaeus, 1767 has been re-
ported in central and eastern North Atlantic Recent
plankton and Late Pleistocene bottom deposits off north-
west .Africa by Diester-Haass and S. van der Spoel (1978).
They attributed the variation to genetic influences, tem-
perature differences, and "mi.xing of populations." Size
differences are smaller than those recorded here and
seem not to be comparable.
Long metamorphic dela> periods, like those reported
above in Bittiurn and C'repidula, seem a likely cause for
the phenomenon to be widespread (little being known
about prosobranch delay periods).
The prodissoconch sizes of some bivalves apparently
are controlled by temperature (Lutz & Jablonski, 1978).
As they pointed out, there could be paleoclimatological
(or paleoecological) implications.
This seems to be the best documented case of extreme
intraspecific protoconch size variation in one plankto-
trophic gastropod species, the variation probably induced
by differing times in the plankton plus a long delay
period (plus other factors). The variation is expressed
batlnnietrically. There is no indication that more than
one Epitonium species is involved, or that there is poe-
cilogony.
Ecophenotypic variation may be more frequent in
marine mollusks than currentK realized. Systematists
should be alert to the phenomenon treated here and
should beware of mere protoconch size differences for
distinguishing planktotrophic species.
ACKNOWLEDGMENTS
I thank Colin Redfern, of Boca Raton, Florida, lor mak-
ing the original discovery antl for loaning me all the
s[)ecimens from Abaco. I also acknow ledge extensive use
of Jack N. Worsfold's Grand Bahama collection (now at
ANSP). The detailed data in these two collections made
this study possible. Dr. K. Elaine Hoagland and partic-
ularK Dr Philippe Bouchet made penetrating and con-
structive criticisms Dr (Jarv Rosenberg and Helen
DuSliane also made liclplnl suggestions.
LITERATURE CITED
Bouchet, I' 1989 A review of poeciiogony in lii'strcipods.
Journal ol .Vlolluscan Studies 55:67-78.
Hdurhcl. I' .ukI \ W .in'-n l9S(i l^j'V isjoiMil lilt- iiDrthea.st
.VduiUic ballival and ai)v.ssai .Vciididae, F.uiiniidae, Epi-
toniidae (Moiiusca, Gastropoda) IJoiiettino Malacoiogico
Sn|>|)lement(i 2:299-.57(i
(blench, W J and R I) Turner 1951 The licnns I'.pitoniinu
ni ihe western ,\tlantic. jDlinsonia 2(:3()):2-49-2SS.
Dull W H ISWJ Heporls on (Iredijiiij; in the Cull ot
\lc\ii(i ,iihI in llic (iaribheun Sea. . .bv
the. . ."Ulake . .Heporl on the Moiiusca I'art II. -Gas-
tropoda and .Scaphopoda Bulletin ol the \hi.seum of Com-
parative Zooiogv at Harvard College 181— 192, 40 pis.
De Bourv, E. 19n. Diagnoses de Scalariidae nouveaii.\ ap-
partenant aux sous-genres Cijchncala et Sodiscald. Bul-
letin du Museum National d'Histoire Natnrelle[Paris] 17(51:
329-331.
Diester-Haass, L. and S. van der Spoel. 1978. Lati- Pleistocene
pteropod-rich sediment layer in the northeast .Atlantic and
protoconch variation of Cliu pyraniidata Liniie 1767. Pa-
laeogeographv, Palaeoclimatologv , Palaeoecologv 2-1:85-
109.
DuSliane. H 1990 liauaiian Epiloniidaf Hawaiian Shell
News Suppiement 1:1-17.
Hoagland. K, E, and R. Robertson. 1988. .An a.ssessnieiil ol
poecilogonv in marine invertebrates: phenomenon or fan-
tasy':' l^iological Bulletin 17-1:109-125.
Killiiini, R, N. 1985. The familv Epitoniidae (Moiiusca: Gas-
tropoda) in southern .Africa and Mozambique .Annals of
the Natal Museum 27(1): 239-337.
Leal, J H. 1991. Marine prosobranch gastropods I rom oceanic
islands off Brazil. . . . Universal fiook Services Dr W
liackhuys, Oegstgeest, Netherlands. \ + 419 pp.
laitz, R .A. and D. Jablonski 1978, Larval bivalve shell mor-
phometry: a new paleoclimatic tool':' Science 202: 51-53.
Pecheiuk, J. A. 1984. The relationship between temperature,
growth rate, and duration ol planktonic life for larvae of
the gastropod Crepidula fornicata (L.). Journal of Exper-
imental Marine l5iologv and Ecology 74:241-257.
l^eclienik, J ,A, 1986. Field evidence for delaved melamor-
pliosis of larval gastropods: Crepidula plana Say, C Jor-
nicata (L.), and Bittiuni ahcrnaium (Sav ). Journal of Ex-
perimental Marine Biology and Ecology 97:313-319.
Rex, M. .A. and K, J. Boss. 1976. Open coiliiii; in Recent
gastropods. Malacologia 15:289-297,
Richter, G. 1973. Field and laboratory observations on the
diurnal vertical migration of marine gastropod larvae.
Netherlands Journal of Sea Research 7:126-134, pi 1.
fioberLson. R. 1970. Svstemalics ot Indo-Pacific Philippia
(Psilaxis\ architectonicid gastropods with eggs and voung
in the umbilicus. Pacific Science 24:66-83.
Robertson, IS, 198.3a, (.)bser\ations on the lite liistorv ot the
wentletrap Epil(niiuni (.■cliindtiiosliini in the Bahamas.
The Nautilus 97:98-103,
lUibertson, R, 198.3b, .Axial shell rib counts as svstrnuitic char-
acters m E))ili)nium. The Nautilus 97:1 16-1 IS
Scliellema, l\. S, 1967, The relationship of temperature to the
larval devi-lopmcnl ol Sassarius ohsolctus (.(Gastropo-
da), Biological Bulletin 1.32:253-265.
W'oodring. \V . I' 1928. Mi<icene mollusks from Bowden. Ja-
maica Part II (Gastropods and discussion of results (Car-
negie liislitiilKin nl \\ .isliiiigliiii I'lililic.itioii ,38.5:i-v ii. 1-
564.
THE NAUTILUS 107(4):1 13-1 17, 1994
Page 113
Effect of Low Water Temperature on Ion Balance in the Zebra
Mussel, Dreissena polij morpha , dwd the Unionid Mussel, Lampsilis
radiata
John I. Scheide
l'livlli> N. Konaminio
l)<|i,irlinrnl nl i5iiiliii;\
(Central Miiliigaii Liii\fiMt\
ML I'lt-asant, Ml 488.59 I SA
ABSTRACT
Dreissrnil pnhiiniirpha. ai. i liiiiaird In hkiiii U'liipcialiirc (21
± 1°(]) l(ir 51 (.la\s 111 artilitial [Kiiiclualer, f\liil)ils liightT
hfni(il\ iiipli siHlium, chloride, potassiiiin and total osniolarit\
than mussels acclimated to 4-5°( : (cold), while hemoUmpli
calcium was increased with cold treatment. The room tem-
perature acclimated nuissel hemoKmph ion concentrations,
maintained lor 51 da\s m artiticial ponduater. were similar to
lliiisc meavuretl irom mussels sampled m the field, (xild-accli-
mation cau.scd Lampsilifi radiata hemoK mph sodium, chloride
and nsniolarity values to decrease, while calcium and the "oth-
er ions, presumably bicarbonate, increased. The positive net
sodunn and chloride flux and the calcium loss of zebra mu^vcK
acclimated to room temperature w as reduced upon direct trans-
ler into cold pondw ater Sodium and chloride net uptakes were
increased in cold-acclimated zebra mussels with an acute in-
crea.se in pontlwater temperature w hile calcium and potassium
net lo.ss increased. The effects of cold storage on zebra mussels
needs to be recognized due to possible changes in the ph\ si( ilog\
of the mussel with this treatment.
Kcij norda: Oild-acclimation. ion llu\, henioKiiipli kui cdii-
centrations. sodium, chloritle. cakium. pdtassiiiin
INTRODUCTION
The receiilK introduced zebra nuissel, Dreissena poly-
niurpha. has rapidly colonized the Great Lakes Region
(Ontario Ministry of Natural Resources, 1992). Zebra
mussels torni large colonies of a higii density, broadcast
spawn, and have a vehger dispersal stage that remains
in the water column for several weeks (Hebert, et al..
1991; Mackie. 1991; McMahon, 1991; Carton & Haag,
1993). Aside from impacting industries and utilities by
louling or clogging water intakes and associated piping
(Mackie, 1991; koCalak et ai. 1993; LePage, 1993), zebra
mussels also impact the ecology of their newly e.xploited
regions by decreasing the density and diversity of fresh-
water bivalves of tiie family L'nionidae (Hebert et al.
1991; Hunter & Bailey, 1992; Mackie 1993) and by de-
creasing phytoplankton availabilit\ (Leach, 1993).
.■\ninials in a freshwater habitat must regulate blood/
hemolymph and cell ion concentrations above the en-
vironmental concentrations. Freshwater bivalves, while
ionic hyperregulators for hemolymph and cell ionic com-
ponents, have reduced concentrations of hemolymph and
cell ions relative to other freshwater species. This results
in a reduced o\erall ionic/osmotic gradient (Murphy &
Dietz, 1976; Dietz, 1979; Kirschner, 1991). While he-
moly nipli ion concentrations are low, fresh water mussel
sodium ami chloride transport rates compare favorably
v\ ith those of other freshwater animals (Kirschner, 1983;
Scheide & Dietz, 19.Sfr Horohov et al. 1992). The zebra
nmssel is similar to other freshwater mussels in such
characters as relatively low hemolymph ion concentra-
tions (Horohov et al. 1992). Ion fliix \alues for sodium
and chloride in Dreissena polyniarpha show a higher
ion turnover with the net fluxes similar to those of other
fresh water bivalves (Horohov et al. 1992).
Current research protocols for long term storage of
zebra mussels in the laboratory recommend that zebra
mussels that are not used immediatelv be maintained at
temperatures below 10°C (Reid et al. 1993). The purpose
of this study was to measure the effect of cold storage
on ion balance in the zebra mussel and compare that
response to a representative of the l'nionidae. The net
ion flux of zebra mussels was measured at two pondwater
holding temperatures (room temperature and cold, 4°C).
The effect of direct (acute) transfer from one tempera-
ture extreme to the other was also determined.
MATERIALS AND METHODS
Freshwater mussels. Dreissena polynwrpha (Pallas, 1771)
antl Lampsilis radiata (Barnes, 1<S23), were collected
from Lakes Erie, Huron and Michigan with ambient
water temperatures ranging from 1 to 17°C. Zebra mussel
sizes ranged from 13 to 35 mm. Zebra mussels were
handled according to recommended zebra mussel con-
tainment procedures (Reid et al. 1993). .All laboratory
water was of Nanopure grade (Barnsteail). Large quan-
Page 114
THE NAUTILUS, Vol. 107, No. 4
Table 1. .'\ coinparisdii of field collcclcd, poridwaliT accli-
malfd (51 da\s In llic lahorator) al room or told IciiiptTatures)
zebra mussel lieinoK nipli ion comciil rations.
( omciilralion (niKq literl
Kooni temp,
pondwater
aeelimaletl
Cold temp,
pondwater
acclimateil
Field
collected
Sodium
C;liloriile
Potassium
("ali'inm
Other
()sniolarit\"
20.2 ± 0.4
24.3 ± 0.9
0.4 ± 0.1
3.4 ± 0.1
6.1 ± 1.6
."i4 ± I
13.5 ± 1.2*
18.0 ± 1.0*
0.2 ± 0.03*
6.0 ± 0.4*
7.1 ± 1.2
45 ± 2*
20.1 ± 1 ()
20.4 ± 0.9*
0.5 ± 0,1
3.4 ±0.2
3.6 ± 0.9
48 ± 2*
\alues represent tlie mean ± the standard error of the mean.
* SisnificantK ilifferent from the room temperature mussels
iP < 0.05), n = 8 for Imth field and room temperature mussels
and n = 16 for colli temperature acclimation mussels.
** I nils in mOsmole ki; HjO.
titles of zebra mussels (greater than 100) were placed in
at least 2 liters of artificial [jondw ater consisting of (niM/
L): 0.5 NaCl, 0.4 CaCU, 0.2 NaHC:03 ami 0.05 KCl (Dietz
& Branton, 1975). Mussels were divided into two groups.
One group was nialntained under lal)orator\ conditions
at room temperature u itii the water temperature ranging
between 20-25°C. The other group was maintained at
4-5°C for at least 10 days prior to u.se. I5olli groups were
maintained under constant light conditions. Mussels were
not fed.
Mussel hemoKniph was witlulrawn from individuals
by cardiac puncture (Fyhn & Costlow, 1975). Blood was
centrifuged at greater than 14,000 Xg for 1 minute (lEC
Ontra M). Total osmolarity was determined using a
\'apor Pressure Osmometer (Wescor, model 5500) with
a 0, 12.5, 25, 50 and 100 mOsm/kg H.O standard curve.
Sodium, potassium and calcium were determined using
flame photometr\ (Cloleman Model 51 Ca Flame Pho-
tometer, Bacharach). Clhloride was determined using a
chloride titrator (Labconco). All ion ileterminations were
referenced to commercially prepared standards and are
expressed as mEq/L. The measured ions were totalled
for each mu.ssel and subtracted from the total solute
osmolarity for each mus.sel. Tfie difference is noted as
"other" and assumed to be [jredominately bicarbonate
(Horohov et a/., 1992).
Zebra mussel net ion flux, an indicator of ion balance,
was determined using the methodolog\ of Dietz and
Branton (1975). Nhissels were rinsed in deionized water
for 30-60 minutes. The mussels were then transferred to
beakers containing 10 ml artificial pondwater. At time
zero, an arbitrary time cliosen when the majoritv had
opened and were siphoning following transfer, 4 ml of
bathing media were withdrawn. After 4 hours, anotiier
.sample of bathing media was withdrawn and the mussel
ti.ssue was removed from the shell and dried. The net
flux of each mussel was calculated as the change in the
bathing tnedia ion concentration corrected lor evapo-
rative loss divided b\ mussel dry weight and lime. The
net flux is expressed as fiEq/g'hr. A positive net flux
indicates that the influx occurring during the experi-
mental period was greater than the ion efflux. A negative
net flux indicated that mussel ion efflux exceeded the
influx for the experimental periotl.
All values are reported as the mean ± the standard
error of the mean with the sample size noted (n). The
Students 't' test was used to determine significant dif-
ferences.
RESULTS
Mussels maintained in both temperature conditions were
open and appeared to be siphoning, indicating normal
nmssel behavior patterns in each acclimation condition.
Zebra mussels maintained in pondwater at room tem-
perature had higher hemolymph sodiuiu and chloride
concentrations and therefore a higher hemolymph os-
molarity than those mussels maintained in pondwater at
4 to 5°C (Table 1). The acclimation time for each zebra
mussel group was 51 days. Hemolymph sodium con-
centrations decreased 26% in the cold treatment group
(P<0.05, n=8 for the room temperature group and n=l(j
for the cold group), while hemolymph chloride concen-
trations decreased 33%. Hemolymph potassium de-
creased 50% w ith the cold storage. HemoK mph calcimn
was increa.sed 76% in the cold-acclimatetl mussels over
room temperature-acclimated mussels. The observed os-
molarity was 13% less in the zel>ra nuissels maintained
at 4°C compared to mussels maintained at room tem-
perature, primarily reflecting the decrease in hemo-
Ivmph sodium and chloride. The cold treatment did not
significantly affect the "other" ions, presumabK indi-
cating no effect on bicarbonate.
Mussels acclimated 51 days at room teiuperature
maintained hemoK mph ion concentrations similar to the
hemolymph concentrations of mussels sampled in the
field (Table 1). Thus, Drcisscna poltjmorpha can main-
tain ion balance during room temperature acclimation
in artificial pondwater with \alues similar to animals
found in the natural envirt)nment This occurs in the
absence of feeding.
To in\estigate whether the cold induced change in
liemoK mph ion concentrations was a specific character-
istic of Drcissena fwlyniorpha or common to other fresh-
water bivalves, the effect of a 10 da\, cold acclimation
on Lampsilis radiata was determined (Figure 1). Most
of the trends observed with the zebra mussel were also
observed in Lampsilis. Hemolymph sodium in L. radiaia
was reduced 60%, while the chloride concentration was
reduced 75%. Hemolymph potassium remained similar
between the two treatment groups. Hemolymph calcium
was elev ated 56% in /.. radiata at the lower temperature.
Hemol\mi)h osmolarity was decrea.scd 2H'"i with the cold
treatment The remaining hemolymph ionic components
("other"), pri'sutnably bicarbonate, were elevated ap-
proximately 2 fold with cold treatment.
These data suggest that at least some freshwater bi-
valves mav reduce their osmoregulatory "set point in
J. I. Scheide and P. N. Bonaminio, 1994
Page 115
t-old It'iiipcratures as coiiipart'il to niiisscls in wariiuT
teiiipt'iatiirt's (laboratorN tf mpt-ratures). Tu test tlic cl-
teet (if an aeute temperature change on ion transpurt
capabilities, zebra mussels were acclimated to either 2 1
± 1°C water temperature or 4°C pondwater temperature
for at least 10 days then tested at the other temperature.
Mussels acclimateil to room temperatures \\ ere observed
to have a significant decrease in the sodium aiul chloride
net Duxes, in magnitude and sign, when mo\ed directK
to the cold water (P<0.05. n= 14 imissels, room anil
cold temperatures except for the sodium flux room tem-
perature nnissels n=13. Figure 2). The negative sodium
flu.x indicates a dominant efflux in the cokl temperature
testing condition whereas at room temperature the pos-
itive net sodium flux indicates a tlominant influx. The
negative sodium net flux was signiticantK dilfcrent from
zero (P<0.05, n= 14 mussels). The change in the net
chloride flux was similar to that observed for the sinliuni
net flux (P<().05, n= 14 mussels, for room and cold
temperatures) Interestingly, the tlirect transfer of tfic
room temperature mussels into the 4°(: pondwater re-
sulted in a decrease in the magnitude of the calcium loss
(P<0.05. n= 14, room and cold temperatures) indicating
a decrease in the calcium efflux. The potassium loss, while
being approximateK one order of magnitude lower than
the other ions, w as significantK less in mussels transferred
to the cold (P<0.05, n= 14, room antl cold temperatures).
The comparison between zebra mussel net fluxes ac-
climatetl to cold pondwater antl mussels acutely trans-
ferred to room temperature pondwater was similar to
those acclimated to room temperature (Figure '3). Mus-
sels, acclimated to the cold bathing media, had signiB-
cantK lower sodium and chloritle net fluxes than those
tlirectly transferred to room temperature (P<0.05 for
sodium and chloride, n=16 mussels for both temperature
groups). The sodium and chloride net Dux in the cold-
acclimated mussels, determined in the cold condition,
were not significantK different from zero(P>0.05, n=16).
Calcium and jjotassium losses were increased in mag-
nitude with the transfer of cold-acclimated zebra nnissels
to room temperature (P<0.01, n=16 each).
DISCUSSION
Zebra mussels in artificial pondwater maintained in lab-
oratory conditions without being fed had hemoKniph
ion values close to hemolymph values of mussels collected
from the field. However, mussels maintained in cold
pondwater, tor the same time period, lost sodium and
chloride with a resulting net reduction in hemolyni))li
osmolarity. The loss of sodium and chloride from Dreis-
scna pulyinorpha and Lampsilis radiata was similar to
the effect of salt depletion (Murphy & Dietz, 197(i; Scheide
& Dietz, 1982). Vet, the cold acclimated mussels were
in a medium that had sufficient ions for mussels to main-
tain sodium and chloride balance at a warmer tein|X'r-
ature.
Sodium and chloride net fluxes usually were positive
in room temperature acclimated zebra mussels. How-
vww Willi acute Iranslci nl rooiii temperature-accli-
mated zebra mussels to cold poiulvvater, mussels lost .so-
tliiim and chloriile The loss of sodium and chloride,
observed in the cold pondwater treatment, was reflected
in the reduced hemolymph sodium and chloride con-
centrations observed with cold pondwater acclimation.
Metabolic activit\, measureil as o\\gen consumption, of
zebra mussels is reduced w ith ctilder water temperatures
(McMahon & Alexander, 1991; Quigley, Gardner & Cor-
don, 1993). The apparent loss of sodium and chloride
observed in the room temperature-acclimated zebra
mussels w hen transferred to colli pondwater may be re-
lated to reduced metabolic acti\it\ . However, long term
acclimation of zebra mussels to cokl pondwater resulted
in a reduction of the net soilium and chloride loss to
values close to zero. The change in ion loss attribiiteil to
zebra mussel cold storage may be the result of a decreased
ionic gradient due to the animal loss of sodium and
chloride or may be due to changes in epithelial ion trans-
piirt and ion permeabilitv , thus the mussels become
"tighter to the loss of ions. Cold acclimated mussels,
transferred back tu the room temperature pondwater
exhibited a positive net Dux for sodium and chloride.
Thus, the ion transport mechanisms necessary for reg-
ulating sodium anil chloriile balance were present in the
cold acclimated zebra nmssels but their activity was re-
duceil in colli |ionilwatt'r
All net ion fluxes stuilieil varied with temperature re-
gime. Zebra nnissels not only exhibited decreased sodium
anil chloriile net fluxes, but decreases in magnitude of
calciinn and potassium net fluxes. At room temperature,
both calcium and potassium net fluxes were negative.
The artificial pondwater useil in this investigation, while
fulK compatible for maintaining unionid mussels (Dietz,
1985) and zebra mussel hemoK mph for extended periods
of time, may not be the best medium for zebra mussels.
The calcium loss indicates that this bathing medium is
negati\el\' influencing calcium balance. Even with the
calcium efflux, liemoK mph calcium values do not change
at room temperature, thus calcium is presumably being
mobilized from the shell to maintain a relativeh' constant
hemolymph calcium value
Even with the negative net calcium flux observeil in
the zebra nui.s.sel, hemolymph calcium concentrations
increased as the hemolymph osmolarity decreased. He-
molymph calcium also increased in the unionid, L. ra-
diata. during cold ponilwater conditions. The rise in
hemolvmph calcium concentrations in cold acclimated-
nuissels mav be assisted by the ilecrease in the net cal-
cium loss at the lower tem|)erature. The elevation of
hemolymph calcium has been noteil with the decrease
in hemoKniph soilium in imioniils (Scheide & Dietz,
1982). Only in L. radiata did the "other" ions increase
with cold acclimation. Similar changes have been ob-
served in the unionid, Ligumia subrostrata (Say) un-
ilergoing ihloriile de|iletion (Scheide & Dietz, 1982).
C^okl ponilwater storage of mussels causeil an increase in
calcium levels concomitant with the decrease of sodium
levels suggesting that compensatory mechanisms, al-
Page 116
THE NAUTILUS, Vol. 107, No. 4
Room
TemoefatLre
I I Cold
Tempefa'LTe
Rocn
Temperattre
I I Cold
TernoeraTure
CZ] c.
jj
Figures I -.'{. ('(iiiipari.sdiis 111 llic t'ftfit'- cil temperature im Ion
concentations in Larnpsilis radiata and Dreissena polijmorpha.
I. I leiiiiiK iTipli ion (.iiiicentratidns cii L(2nipsilis ratluita alter
a 10 (la\ aeeliniatidii period to room temperature (21 ± 1°(^
solid liar) and cold temperature (4 ± 1 °C:, liatehed bar) K'll)
refers to the potassium value multiplied l)y 10 for comparison
purposes. "Other relers to the difference between the mea-
sured total .solute and sum of the measured ion values, as noted
in the Materials and Methods section and indicates the he-
molymph bicarbonate concentration. The capped lines indicate
the standard error of the mean. Sigiiilicanl ilifferences are de-
noteil v\ilh a • representing P<0 ().5 with n = (S for room and
cold lemperalure-accliinated mussels 2. Kffectsof direct trans-
fer of 2I°(: ± \°C acclimated zebra mussels to 4°( : on the
sodium, chloride, calcium and potassium net fluxes The po-
tassium net flu.x is increased 10 fold lor comparatiM' purposes
(K*10). The room temperature acclimated nuissel net flu.x is
denoted by the solid bar {n= 14 mussels toi all i^mups except
tliougli not completely effective in maintaining room
temperature osmolarity, were acli\f in llie cold-aceli-
mated mtis.sels.
Freshwater animals iia\c set points lor liemoixnipli
blood sodimn and chloride (Kirschner. 1991 ). \lo\ement
of zebra mussels from the natural en\ironinenl to the
laboratory does not appear to negatively effect the mus-
sels for at least 51 days. The formulation of artificial
pondw ater appears to be suitable for tlie laborator\ niain-
tenante of zebra mussels. Zebra mus.sels, held at room
temperature, maintained hemolymph ion values close to
the collection hemolymph ion values, although the mus-
sels were in a negative calcium balance. The mussel
holding time in the laboratory may be increased by plac-
ing the mussels in cold pondwater and zebra mussel
storage at a cold temperature is recommended for lab-
oratory housing of zebra mussels not immediately being
used tor studies (Held, ei al. 1993). However, the ion
concentrations after holding may be depressed, partic-
ularly with a longer cold holding period and the lack of
feeding. Changes in mussel plu siology, due to cold stor-
age, need to be recognized as lurther research is per-
formed to understand the basic biology of the zebra
mussel.
ACKNOWLEDGEMENTS
This work was funded by a Research E.xcellence Fund
grant #42865 from the State of Michigan to JIS. The
cooperation of Consumers Power is greatly appreciated.
We thank Thomas Dietz, Stexe Donner, Ron Fobes and
Harokl SiKerman tor the critical re\ icw ol this manu-
script.
LITERATURE CITED
Diet/. T II 1979. Uptake of sodium aiKJ chloride In fresh-
water mussels. Canadian Journal of Zoology .37:1.56-160.
Dietz, T. H. 1985. Ionic regulation in freshwater rimssels: .\
brief review, .\merican Malacological Bulletin 3:233-242
Dietz, T. H. and W. D. Brantoii. 1975. Ionic regulation in
the freshwater mussel. Liguinia subroslruta (Sav). Jmirrial
ol (;oin|)arati\e PInsiiilogx 104: 19-26.
l\liii, II .iiiil I n t'ostliiw 1975 .Vnaerobic sampling of
l)iid\ nuids in l)i\al\e molluscs. Comparative Biochemistry
and Physiology 52A:265-268.
Carton, D W and W IV Haag. 1993 Sea.sonal reproductive
iNcles and settlement patterns of Drcissaia palyiuorpliu
lor the sodium group, n=13l and the coltl net llux is denoted
b\ the hatched bar (n = 14 mu.ssels for all groups). Signihcant
flux differences are denoted by • tP<0.05) 3. Effects of direct
transler of 4 ± 1°C^ acclimated zebra mussels to 21°C on the
sodiimi, chloride, calcium and potassium net fluxes The po-
tassiiini net flux is ini'ieased 10 lold for comparative purposes
(K- 10). The cold-acelimalcd net flux is denoted b\ the hatcheil
bar (11 = 16 mussels) and the room temperature nuis.sel net flux
is denoted In the solid bar (n=16 mus.sels). Significant lliix
differences arc denoted In • (P<0.05).
J. I Scheide and P. N. Bonaminio, 1994
Page 117
in \\ t'stcrn Lake Erif //r Nalcp.i T 1' amlD \\ S( lilocs-
ser it'ds. ). Zebra mussels: Biolog). UTii)aets ami nuitKil
t:RC: Press, Inc. Boca Raton, FL, p, 1 I 1-12.S
Hebert. P. D., C. C. Wilson. M. H Miirdoeh and b i.a/ar
1991. Deniograpli) and ecological impacts nl tlic m\ad-
ini; mollusc Drcissvna pdhjiiKiiiilid (:anadiati JoiuTiai ol
Zoolot;\ W). 405-409
Horohov. J . H Silverman. J W Lynn and T H. Diet/ 1992
Ion transport in the freshwater zebra mussel, Dreissena
pnhjmorpha. Biological Bulletin 1(S3:297— 'jOo
Hunter. H. D and J, F. Bailey. 1992. Dreissena ixilynuirphu
(Zebra nui.s.sel): Colonization of soft substrata and some
effects on I iiionid bixaKes. Tlie Nautilus lOtit-iO-fiT
Kirschner, L B I9S.'5, Sodium and chloritle absorption across
thelxHb surface: Frog skin and other cpilhrli.i \mcricaii
Journal of Physiologv"244:R429-H443
Kirschner, L. B 1991. \Vater and Ions //i; I'rosscr. ( : 1. led.)
Comparative .Animal Phvsiologv. l-'ourlh I'dition l"n\i-
ronmental and Metabc}|ic Animal l'h\si(il(ig\ \\dc\-
Liss.lnc. Neu York. p. 13-107
Kovalak. \V. P.. C, D. Longton and U H Simlhrr 1993
Infestation of power plant uatcr s\ stems b\ the zebra
nmssel [Dreissena polymurplia Pallus) In: Nalepa. T F
anil U.W Schloesser (eds. ), Zebra uurssels: Biologv . im-
pacts and control t"RC- Press. Inc. Boca Raton. FL ji 359-
3S0.
Leach. J. H. 1993. Impacts of the zebra uuissel [Dreissena
pohjmorpha) on water quality and fish spawning reefs in
western Lake Erie. In: Nalepa. T F and D W Schloesser
teds.). Zebra mussels: Biologv. unpads and control. CRC
Press. Inc. Boca Raton. FL. p. 381-397
LePage. W, L 1993. The impact of Dreissena pulyniurpha
on waterworks operations at Monroe, Michigan, .A case
histor\./;i. Nalepa, T. F. and D.W Schloesser (eds.) Zebra
mu.ssels Biolog\ , impacts anti i-oritrol CIU: Press. Inc.
Boca Raton. FL. p. 333-358.
Mackie. C; L, 1991. Biologv' of the exotic zel)ra unis.sel. nrci.s-
,sf/ia pohjmorpha. in relation to native bivaKes and its
potential impact in Lake St Clair Hvdrobiologit a 219
251-268.
Mackie. (.. 1. 199:) Biolog\ ol the zebra uuissel [Dreissena
polilinorplia) anil observations ot uuissel colonization on
unionid bixalves in Lake St Clair of tlie Great Lakes In:
Nalepa. T I" ami l.).\\. Schloesser (eds.). Zebra nuissels:
Biologv. iiupactsand i-onlrol (4U,' Press. Inc. Boca Raton,
FL. p. 153- Kio
McMahon, R. F. 1991 Mollusca: Bivalvia. /;;. Thorp. J H.
and .\. P. Covich (eds.) Ecology and classification of North
American freshwater invertebrates .\cademic Press, Inc.,
NY. p. 315-399.
McMahou. H F and J F Alexander 1991 Respirator) re-
sponses to temperature, h\po\ia and temperature accli-
mation in the zebra mussel, Dreissena pohimorpha i Pall I.
.\mericaii Zoologist 3174.^
Murpln. W A and 4' II Dietz 1976 The effects of salt
depletion on blood ami tissue ion concentrations in the
freshwater mussel. I.iiiuinia subrostrata (Sa\ ). Journal of
Comparative Plusiologv 108:233-242.
Ontario Ministrv ol Natural Resources. 1992. Zebra mus.sel
distribution Mmistrv ol Natural Resources. Ontario. Can-
ada, December. 1992
Quigley. M. A., W. S. (;ardner and W. M. Cordon 1993.
Metabolism of the zebra mussel [Dreissena pohjmorpha)
in Lake St. (Mair oi the (Jreat Lakes, h\: .Nalepa. T F and
D. W. Schloes.ser (eils. ). Zebra mussels: Biologv. impacts
ami control. C;RC: Pres.s, Inc, Boca Raton, FL, p. 295-306.
Reid. L^ F . J Biduell, J Carlton, L. Johnson, E, Mar.sden and
S.J Nichols 1993. Zebra mussel containment protocols.
NO.-\A Sea (Jraiit Office and L'S Eu\ ironmental Protection
.Agencv . 20 pages.
Scheide. J I and T II. Dietz. 1982. I4ie effects ot indepen-
dent S(idium and chloride depletion on ion balance in
freshwater mus.sels. (-an.iduin Journal of Zoologv 60 1676-
1682
Scheide. J I ami 4" 11 Dictz 19S6. Serotonin regulation of
gill c.WIP proiliictioii. ,Na, and uater uptake in freshwater
mussels Journal ol l-Aperimental Zoology 240:309-314.
THE NAUTILUS 107(4):118-123, 1994
Page 118
The Muricid Types of Frank Collins Baker
Emily II. Yokes
(^f(il(ii;\ l)c|);ir(iiicnt
riiluiif I iiiversit)
New Orleans, LA 70118-5698, USA
ABSTRACT
in llif lirsl leii scars ol lii.s scit'iililic caret-r V X', Baker iiaiiu-d
nine ta\a in the faiiiil) Muricklae, Of these only one, Haus-
tcllutn ruhidus (named as Murcx mcs.soriiis var. ruhidum). is
here eonsidered \ahd \ll others are synonyms ol older names.
Key uardf.: Miirieidae, (Gastropoda, VX'.. Haker
INTRODUCTION
Tlie name Frank C Baker is very familiar to workers
among land and fresli-water mollusks, i)iit for tfie first
ten years of his professional life lie thought ot himself
as an expert in marine mollusks, especialK the Muricidae,
Of his first 21 papers, published between 1888 and 1897,
only two (1895b, 1897b) are on non-marine Mollusca,
His first publication (1888) was a note describing the
marine mollusks collected along the coast of Massachus-
sets.
In the "(General Notes" of The Nautilus (3:46), it was
staled that "Mr. F (L Baker, formerly of Providence, R.I.,
is pursuing his studies at the Academy of Natural Sciences
ol I'hiladelpliia. He spent the months of January to
April, 1889, in Brevard County, Florida, and from this
experience came a diverse set of notes on the marine
mollusks (1889a), on the feeding habits of birds (1889b),
including the identification of a number of mollusks eat-
en by said birds, and on the habits of the muskrat (1889c).
The first molluseaii species he described was Ocinehra
jcnksii (Baker, I889d) and in 1890 when the American
Association ol (^oncliologists was initially annoimced (77k'
Nautilus, 3:140-143), among the charter members listed
was "F.C. Baker. Academy Natural Sciences. Philadel-
phia. Muricidae ." His second taxonomic paper (Baker,
lS9()a) was also on a muricid, with the redescriptidu ol
I'rosalpinx perrugatus ((Conrad, 184(j)
Among his most important contributions to the IkkK
of molluscan know ledge are his series ol seminal papers
on the i)rotoii)nchs of tiic Muricidae (Baker, 18901), IS9ld,
1894) and other marine gastropods (Baker, 1897c), to-
gether with several catalogues of the species ol Muricidae
(Baker, 1891e, 1895c, 1897a).
In 1894 he moved to (Chicago to beconu' the lirst v\\-
rator ol mollusks at the newly constructed Academv of
Sciences (see Baker, 1895a), a position he held for the
next 20 years (Franzen, 1958:30), and it is only then that
his interests turned to non-marine mollusks, beginning
with an important paper (1895b) on the transport of land
and Iresh-water mollusks by means of birds, fish, etc.
(possibly the first report of "ducks w ith dirty feet trans-
porting embryonic shells). Although he published three
more papers (1895c, 1897a, 1897c) on marine mollusks,
alter 1898 lie never again digressed from non-marine
lauiias.
In the time that he was study ing muricids Baker de-
scribetl nine new taxa, only four of which (those in the
1891d work) were figured. For the most part these species
have been ignored by subsecjuent workers. Keen (1971:
554) noted that Ricinula rugosoplicata Baker is a syn-
onym of Buccinum lugubre C.B. Adams, 1852, and Yokes
(1971) atteni])ted to assign the species to generic taxa.
w ith varying degrees ol success (see below ). Radw in and
D'Attilio (1976) did nothing with the species, e.xcept to
list two as "species of uncertain identity" (Intuherculatus
and jenksi [sic\ p. 217). Only two taxa named as "va-
rieties have ever been the subject of any discussion:
Murex haustclluin var. longicaudus and Murex mes-
sorius \ar. ruhidum.
In a monograph on the Indo-Pacific species of Chi-
coreus Iloiiart (1992, fig. 371) refigured the holotype of
Murex (Chicoreus) bitidjerculatus. from the collection
of the Chicago Academy of Sciences. This, combined
w ith an on-going study on the species of Ergalataxinae,
to which I susix'cted certain of Bakers species nright
belong, caused me to contact Mr. Ron \ asile. Curator of
Mollusks at the (Chicago Academy, and through him 1
arranged to borrow the t\ pe material ol these little known
taxa. To complete the study 1 also contacted the Academy
of Natural Sciences of Philadel])liia. where four aildi-
lional t\ pes were located, and the Wagner Free Institute
of Science. Philadelphia, w hich supposedly had one ty pe
lot that could not be located.
Because these taxa are so poorly known, it seems
worthw hile to present all ot them in one place, and they
are listed below in chronologital order as originally de-
.scribed. UnfortuiuiIcK , most \no\r to be synonyms of
presiousK described species. l''\cri though this is not a
E. H. Yokes, 1994
Page 119
"rt'visionarx work, lietause tlifre are multiple speci-
mens of certain ot tlie ta\a, it seems aiK isihie to ciesi^nate
lectot\pes. It is unlikely that most ot the taxa will ever
he treatetl in an\ strictK moiiot;raphic work,
SYSTEMATIC :S
Ocinchra jcuLsii
Figures 1, 2
Ocinchra jcnksii Baker, l.S89ci:S()
'■'I'riisdipinx jctiksii (Baker). \'okes:122.
Habilal: I nknown.
Lectolype: ANSP 36099 (here desiiinated) (fig. 1).
Paralectolypes: CAS 20696' (fig. 2); ?WFIS (ru.t toniul).
Discussion: Baker compared O. jcnk.'-iii lo the Cahlor-
nian species Occncbra circumtcxta Stearns, 1S72, i). gra-
cilliiita Stearns, 1S72, and O. inicliacli Ford, 1888, sug-
gesting that he perhaps thought it might also be from
California. However, examination of the twe) extant type
specimens indicates that it is an elongate form of the
xariable common southern .Australian Lcpsivlhi rinosa
(Lamarck, 1822) (see fig. 3),
In the original description. Baker (1889d:81) noted
that there were four specimens "obtained from the Wag-
ner Collection at the Wagner Free Institute of Science.
A search of the collections of the Institute did not unco\ er
any specimens but at the Academ\' of Natural Sciences
of Philadelphia, where Baker worked at the tii7ie, there
is a specimen (ANSP 36099; height 16.5 mm, diameter
9.5 mm) labeled as "syntvpe" that nearly matches the
cited dimensions ("Alt. 17, diam. 10 mm"). This speci-
men is designated as the lectotype. There is one para-
lectotype (CAS 20696) in the collections of the Chicago
Academy of Sciences, which is slightK smaller (height
15.1 mm, diameter 8.3 mm). The fate of the two addi-
tional paralectot\ pes is not known.
Murcx haiistclluin Linne var. lungicaudit^
Figure 4
Murcx htiuslcUum Linne var. longicaudus Baker, IS9li):5(i.
Ilau.slvlluni longicaudm (Baker), \'okes, 1971:6(1
Not Hau.stellum Umgicaudnni [.sic] (Baker). Kaiclier, 1973, 2 1 21
(= H. kurodai).
Not Hatistcllum longicaudus (Baker). Fair, I97():55, fig. 50 ( =
// kiirod(ii)
llaiislclliiiu haustclluni iHiiislclliini (ijnrie) PDiuier and \'llkt•^.
19,SS,S(i. tigs 4W:. 4(SD ('■h(ili.t\p<-"(lccl(.t\|)cjnl // !,„i-
gicdtidlist
Hauslelluiii huiislclluni longicaiidiis iBakcri llcmart. 1990;
332, figs. 12, 13, 29.
' Bcith till' Chicago At-adeni\ ot Sciences and the Caliturnia
At-adenu ot Sciences use the abl)re\ iation "CAS lor specinu-n
Itifiitilieation. In every case in this paper tiie reterenee (]AS is
til ihi' Cliicagd .\cadeni\ nt Sciences.
Habitat: Red Sea
Lectolype: ANSP 60965 (here designaled).
Paralectolype: CAS 2070! (fig. 4).
Discussion: Most authors (e.g., Radwin & D Attilio, 1976:
49; Ponder & Vokes, 1988:86) have considered this "va-
riety" to be a synonym of typical Murex hausteUiim.
Howe\er, Houart (1990:332) has expressed the opinion
that the differeuct's in the nature of the protocorich be-
tween Haustclluni Iwustcllurn s,s. (see Ponder & Vokes,
1988, fig. 78D; Houart, 1990, fig. 31) and forms from
the Indian Ocean named longicaudus (see Houart, 1990,
fig. 29) and Haustcllum sp. (Houart, 1990, fig. 32) in-
dicate three distinct species. Given that all three have
protoconchs of 2 to 2.5 whorls and the major difference
is in the whorl shape, ranging from rather conical in the
t\ pical Indo-West Pacific form, to cylindrical in the East
.African form {Haustclluni sp), to somewhat bulbous in
the Red Sea form {longicaudus), his argument is not
especially compelling.
The differences in teleoconch morphology are less be-
tween these three forms than between the typical //.
Iiaustclltini s.s. and populations named as //. kurodai
(Shikama, 1964) and H. vicdani Kosuge, 1980 (see Pon-
der & Vokes, 19S8, figs. 87A-C;: kurodai. vicda7ii, and
haustclluni s.s., respectively), thus it is probable that the
Indian (Jcean/Red Sea populations represent only mor-
phological variation as v\ell.
In the original description Baker (1891b:56) stated that
he had three specimens of this variety, one of which was
in tlie collections of the Academv of Natural Sciences of
Philadelphia and "two are in my collection." The spec-
mien in the collections of the Philadelphia Academy
(ANSP 60965) is here designated as the lectotype and
the one (fig. 4) in the Chicago Academ\ of Sciences (CAS
20701 ) is a paralectolype. I do not know w here the third
specimen is today,
Ocinchra pilshryana
Figures 5-7
Ocinchra rdshniana Baker, 1S911).57.
?Ur()salpinx piUliryaua (Baker), Wikes, 1971:125,
Habitat: ('e\lon
Lecioiype: ANSP 60307 (here designated) (fig. 5).
Paraleclotypcs: CAS 20699A and 20699B (figs. 6, 7).
Discussion: Raker stated that this species "has features
recalling the genera Troplion, Vrosalpinx. and Sistrum"
(1891b:58) and he is not the only person who has had
difficult)' placing it. Examination of the type specimens
(figs. 5-7) shows it to be a subjective synonvm of Er-
galatax contracta (Reeve, 1846). which also has been
attributetl to several different genera b\ other authors:
Buccinuni contractuui Reeve, 1846; Murcx calcareus
Dunker, I860; L'rosaliiinx innofahilis E.A. Smith, 1879;
Pcntadactylus cctjlonicuui Preston, 1909; and Morula
martcnsii Dall, 1923.
Page 120
THE NAUTILUS, \ol. 107. No. 4
13b '^M
Fifjurch 1,2. (hinclmi jcnk.sii Baker I. ANSP .■>W)99 (lectotxpe); height 16.5 mm. localit) unknown (X 3). 2. (^\S 20696 (para-
InldUpe); height 1.5.1 mm. locahty mikiiowii (X S). 3. LcpsicUa vinoaa (Lamarck), TAM C: 167197; height 8.2 mm. Hecherehe
Ba\, Tasmania. Australia (X 4). !•. Murcx haiislclliirii \ai longiniudns Baker, C'.\S 20701 (paralectotype); height 49,7 mm. Red
Sea (X 2,1. 5-7, Cktiichra pilshnjiina Baker. 5. ANSI' 60:507 (leef(it\ pe). height 15 5 nmi, Cieylon (X 3), 6. CAS 20699 (paraieetot) pe
A); lieight 16,0 mm. Ceylon (X .3,). 7. CAS 20699 (paraleel<)tv|)e B); heigiit 14.4 mm, Ceylon (X 3). 8, <^. Ricinula iSislrum)
rugusuplicala Baker. 8. A.N'SB 61306 (leelolype); height 1 1 .7 mm. Turtle Ba\ . Lower California (X 3). 9. C;AS 20703 (paraieetot) pe);
heiglit 11.0 mm. Turtle Bay. Lower California (.X 3). 10. Miircx (Chirorcus) hituhcrculatu.s Baker, C.\S 20702 (holotype); height
31.7 mm. Australia (X 2). 11. Ocinchra wanliana Baker, C.\S 20698 (holotype); height 12 5 mm, ".Australia" (X 3). 12. Ocincbrina
aciculata (Lamarck), Vokes C^oll.; height 10.6 imn, off Bota, Spain (X 3). LJ. Ocinebra rubra Baker, C.'\S 20697 (holotype); height
10.7 mm, locality unknown (.X 3). 14. Orcncbra clalhrata (Dall). Vokes Coll.; height 10.0 nmi, off Hollester Banch, California, 15
m (X 3) 1.1. Murex nicssorius var. rubidum 15aker, I SNM 87S003; height 30 5 mm, Anna Maria Ke\, Florida, 46 m (X 2) 16.
17. Purpura iThalfssa) problcrualica Baker. 16. CAS 2()704A (lectotx pel: height -30 9 mm, jafian (X 21 17. CAS 20704B (para-
ieetot) pe): height 22.8 mm, Japan (X 2)
E. H. Yokes, 1994
Page 121
Often eited as a another s\n(in\ni is Ergalalax rcciir-
rens Ireclale. 1931, named astNpeot tlie genus (ami hence
contracta is usualK cited as the t>pe species of Ergala-
tax): however, E. recurrens is not the same as £. con-
tracta but is a synonym of Miirex pauper Watson, 1883,
which is thus the correct name for the type of the genus
Ergalatax Iredale, 1931.
There was no comment on the nurnl)er of specimens
in the original type lot. There is one specimen (fig. 5) in
the collections of the Academy of Natural Sciences of
Philadelphia (ANSP 60307), which is here designated as
the lectot\ pe; two additional specimens (figs. (3, 7) at the
Chicago Academy of Sciences (C^.AS 20699) are desig-
nated as paraleetot\ pes.
RicinuUi (Sisfnnu) rugusuplicata
Figures 8, 9
Ruutula iSistriiiu) rugusoplkata liaki-r. l.S91b:58
Habitat: Turtle I5a\, Lower C'alilornia.
Lectolype: ANSP 61306 (here designated) (fig. 8).
Paraleclolype: CAS 20703 (fig. 9)
Uisrussion: Keen (1971:554) included R. rugoaoplicata
in the s> nonynn of Buccinutu lugnhrc (IB. Adams, 1852,
now placed in the Ocenebrine genus Trachypollia and
not Morula {Morunella), as Keen reported. There are
two specimens in the type lot, one at the Academy of
Natural Sciences of Philadelphia (fig. 8) and one at the
Chicago Academy of Sciences (fig. 9). leaker (1891b:58)
says "this pretty little shell was found in a collection
recently purchased by the Conchological Section of the
Academy of Natural Sciences from Mr. Henry Hem-
phill"; therefore, the Philadelphia specimen (ANSP
61306) is here designated as the lectotype.
Murex (Chicorcus) hituhcrculatus
Figure 10
Murex [Chicorcus) Itiluhcrculatu.^ Baker, l.S9Ul:133, |)l. II,
fig. 4.
Chicorcus hituhcrculatus (Baker). N'okes, 1971:24,
Murex hituhcrculatus Baker. Fair, 197(i:26; Houart, 1992 l()(i,
fii;. 371 (li(il(il\ pe).
Habitat: ,\ustralia.
Holotype: CAS 20702 (fig. 10).
Discussion: F"air (1976:26) was the first to suggest that
M. bitubercuUitus is a juvenile specimen of Chicoreus
capucinus (Lamarck, 1822). Examination of the holotype
shows that she was correct. .Although the localitv "Aus-
tralia at first seems misleading, C. capucinus does occur
in the tropical mangrove areas of northern Australia.
Ocinchra uardiana
Figure 11
Ocinchra uardiana Baker. 1S9U1 134. pi 11, fig. 5.
(Jcinchnna uanluma (Baker) NUkcs. 1971 127; Fair, 197(i:S(S,
tii; l)."> (,illrr B.iki-r).
Habitat: "Australia.
Holotype: CIAS 20698 (fig. 11 ).
Discussion: F]vidently the erroneous locality of "Austra-
lia convinced Baker (1891d:134) that this species, al-
though "having a superficial resemblance to Ocinebra
aciculata Lam., ' was to be distinguished by its larger
size, more ileveloped umbilical region, and its \ellow
color, differing from the "light rosy Mediterranean spe-
cies Ocinehrina aciculata (Lamarck, 1822). True, this
gerontic example is larger than most specimens of O.
aciculata liut l^adwin and D'Attilio (1976:125) cite 16
mm as maximum height for this species; they alsf) note
that the color ranges from orange-brown to red-brown
to purple-lirow II. Comparison of the type of O. wardiana
to a typical Mediterranean example of O. aciculata (fig.
12) fails to shf)\\ an\ significant differences. Nothing like
it has ever been recognized from .Vustralian waters.
Ocinchra ndmi
Figure 13
Ocinchra ruhra Baker, 1891d:134, pi 11, figs. 6, 7
Ocinehrina ruhra (Baker). Yokes, 1971. 12(x Fair, 197(i:73, fig.
52 (alter Baker).
Habitat: Lnknowii
Holotype: CAS 20697 (fig. 13),
Discussion: ,\lthough compared by Baker to Favartia
alvcata (Kiener, 1842) and F. pcasci (Tryon, 1880), the
only resemblance is the development of deep pits formed
by the intersection of the axial and spiral ribs. Baker
(1891d:135) iurther noted "it has some resemblances to
the figures of Ocinchra intcrfossa Cpr." and here he is
much closer to the truth. In fact, the same form (see fig.
14) was later named Tritonalia intcrfossa var. clathrata
by Dall (1919:324; holotype figured In Bormann, 1946,
pi. 4, fig. 2; Fair, 1976, fig. 315; Kaicher, 1978, no, 1615).
If one considers this a valid species, then Baker's name
is the correct one to use. Most w orkers, however, consider
the form to be a synonym of Ocenebra atropurpurea
Carpenter, 1865 (lectot>pe figured b>' Bormann, 1946,
pi. 4, fig. 8; Palmer, 1958, pi, 23, figs, 2, 3, 13; Fair, 1976,
fig. 316; Kaicher, 1978, no. 1627), which originally was
named as a variety of the common California species O.
interfossa Carpenter, 1864 (holotype figured by Bor-
mann, 1946, pi. 4, fig. 10; Palmer 1958, pi. 23, fig. 1;
Fair, 1976, fig. 318; Kaicher, 1978, no. 1614). Ocenchra
rubra /clathrata is no more than a shouldered form of
O. atropurpurea, and both are simply clathrate varieties
of the O. interfossa complex.
Purpura (Thalessa) prohlcmatica
Figures 16, 17
Purpura [Thalessa) ]nijhlcnuilu(i \\.iki-c lS91tl 135, pi 11. figs.
2. 3.
Page 122
THE NAUTILUS, \'ol. 107. No. 4
Habitat: Japan [Sfta coast, Japan, jiilc I-'ranze'ii, 1958:34,
f 11)111 lal)fl with .spciiiJifii.s].
Leclotype: CAS 207()4A (ilcsi^iialcd l)\ l-'raii/fii, 1958:
34) (fig. 17).
Paralec-totype: C:AS 2070 IB (fig. 10).
Dii^cu^sion: Iriasimicli as /'. pnihlcmulica is uiujiiestioii-
ahl\ a .sv'iioiiyin of the common Japant-sc specii'S Purpura
rlavigcra Kiister, 1800, one can only assume that Baker
was misled by Trvon (1880:163), who clismissed P. cla-
vigera as "a depauperate iorni" of P. tumulom Reeve,
1846, the species to which Baker compared prohlema-
lica. Baker (I891d:136) further added that "this species
is well figured on pi 46, f. 48 of the second volume of
Tryon's Manual, which is a reproduction of the shell
Lischke (1869, pi. 5, fig. 15) figured as P. tiitnulosa Reeve.
The latter is not Reeve's species l)ut the nodulose form
of P. clavigera, as Lischke demonstrates in his series of
illustrations (1869, pi. 5, figs. 12-l(i). Both morpiiotvpes
have been well figured In Kaicher (1980, no. 2481).
Reeve's species P. lunuilofia is a synonym of Thais
(Tenguella) marginalha (Blaiinilie, 1832), a widespread
Indo-Pacific species that does occur in eastern Africa, and
Reeve s locality ot "Zanzibar" is probabiv not erroneus,
as suggested by Tryon. Members of the P. clavigera
com[)lex have been placed in the subgenus Thais (Re-
ishia) Kuroila and Halx', 1971, type species: Purpura
hrunni Dunker, 1860.
There were said to be 12 specimens in the type lot,
two ol which were figured in the original description.
I-'raiizen (1958:34) selected the larger of tliese (Baker,
1891d, pi. 1 1, fig. 3) as the lectots pe. Both specimens are
figured liere (figs. 16, 17).
Murex messnrius var riilndmu
Figure 15
Manx mcssorins var nilnihiiii Hakcr, IS^^Hil l.iT iinulf iianic
list only); 1897a;377
Murcx (Murex) rertirviroslns nihuliis I5akfi , ( ^Iciiili .iiid Perez
Farfaiite, 1945:6, pi. .), figs. 1-7 (figs. 1. 2 iectol\pe).
Murex (Murex) ruhidus Baker. Yokes, 196;11(),S. |)1, 4. fig.s, .'j, 4.
Munx ruhidus Baki'r Nukes, 1971:92; Radwiii ami D'Mlilio,
1976:71, pi. 11, hg, 15; Fair. 1976:7:3, fig. 40.
Haustelhun ruhidus (Bakerl \ (ik<'s. 1990:17. pi. 1, tig 12
Murex ruhidus paniiiuicus I'rliKli. 199062, fig. 8.
Habitat: ( ledar Ke\s, Florida.
Lertotype: Harvard M(;Z 147272 (designated In Clench
and Perez Farfante, 1945:8).
Discussion: This is the most w idcK recognized ol leaker's
taxa, and was only inadvertantly named In Baker, wlm
credited the varietal name to ball (1889:196). In that
reference Dall describes the Florida specimens of Murex
/nessorius as 'often of a deep ro.se-color . . . less hooked
and spinose than the discolored specimen figured b\
Reeve [1845, pi. 22, fig. 90, which is a good example of
typical rnessoritis]" However, nowhere does Dall men-
tion the word "ruhiduni" and the name must date to
Baker s 1897 usage, w here he does gi\e a description of
sorts ("the shell being of a deep-pink color," 1897a:377),
which together with the Florida localitv' is sufficient to
identify the species
(Jeiich and Pc-rez Fartante (1945:8) recognized the
Florida form as a valid subspecies of the Pacific reeur-
virostris Broderip, 1833, changing "ruhiduni" to the or-
thographically correct ruhidus. and noting that the spe-
cies is not onl\ pink but comes in "cream, pink, pale
orange, salmon, or even red and some specimens possess
darker color bands at the shoulder and base of the body
w horl.
Althougli generally confined to the Florida and Ba-
hamian areas, specimens also occur in the Ba\ Islands of
Honduras, the northern coast of South America, and
Panama (see Vokes, 1990:17). The latter have been sep-
arated as a geographic subspecies by Petuch (1990:62,
fig. 8).
ACKNOWLEDGMENTS
I wish to thank Ron Vasile, Curator of Mollusks at the
Cyhicago Acadenn ol Sciences, for making a\ailable the
types in their collection. Gar\ Rosenburg, Earle E. Spa-
mer, and David S. Wiedner, all ol the Academv of Nat-
ural Sciences of Philadelpliia, located and loaned tvpe
material from their institution. Eugene .\. 13olt, Jr., of
the Wagner Free Institute of Science, Philadelphia, was
most helpful in attempting to track down the types of
O. jenksii, said to he in their collections. Ian Loch, the
Australian Museum, kindK provided the figured speci-
men of Lepsiella vinusa, as well as considerable help and
hospitalitv during in\ 1988 visit to the .\ustralian Mu-
seum.
ABBREVIATIONS OF REPOSITORY COLLECTIONS
ANSP - Academy of Natural Sciences. Philadelphia,
Pennsylvania, USA
CAS - Cliicago Acadenu oi Sciences. Chicago. Illinois,
L'SA
MCZ - Museum ot C^omparative Zoolog) . Harvard Uni-
versitN. (-ambridge. Ma.s.sachu.ssets, USA
TAM - The Australian Museum, Svclney. New South
Wales, Australia
USNM - National Museum of Natural History, Smith-
sonian Institution, Washington, D.(>., USA
\\ I'lS - Wagner Free Institute ol Science, Philadelphia,
I'ermsv Kania, I S.\
LITERATURE CITED
Baker. F, C, ISSS, |( )ii a icillcrlinu Irip Id ( )iisel Bay. Mass.].
rlir ( :(inilinl(igisl's Exclian.ge 2(9):! 19.
li.ikcr, I- C ISSOa, Notes dm Floridaii siiclls. The X.iutilus
.■5(5):5;5-54,
Baker. F (' IS.SOli Nolrs mi the lood ol hirtls rrorreiliiigs
ol .Vi'.ulciiiv ol X.iliiral Sciences of Pliilaclel|iliia 11:266-
270.
E. H Yokes, 1994
Page 123
Baker, F. C, 18S9c, Remarks upon tin- rdUiKl-Uiilcil iniiskial,
Neofiher alleni. True. Procedings nl Aciilcriu dl \,ihir.il
Sciences of Philadelphia 41:271-273
Baker, I"" (; IcS.SQd, De.scriplioii of a iiru -.[H'ck-sdl Ociitchra
The Nautilus .3(7):.S0-N1
Haker, P'. C;, 189()a. Reniarksou I rosalpiiix pcniifialiis (.'.mir
Proceedings of .-KcademN of Natural Sciences of Philadel-
phia 42;4(i-47; reprinted vcrhatuii ni The N.mlilu-- 4l,'5);
29-30.
Baker, F. C. f 890b. On the modibcation ol the apex in Munx
Proceeding of .Acadeni) of Natural Sciences ot Philadel-
phia 42:66-72.
Baker, F CI. 1890c. .\ catalogue of conchological alilire\ia-
tions The Nautilus 4l8):89-91
Baker, V C 1891a. Notes on a collection ot shells Ironi south-
ern Mexico. Proceedings of .\cadeni\ of Natural Sciences
of Philadelphia 4,3;45-.5o
Baker, F (.'. 18911). 1-leinarks on the Mmicitkie with descrip-
tions ol new species of shells. Proceedings of .Acadeiii\ ol
Natural Sciences of Philadephia 43:.56-61.
Baker, F. C;. 1891c. [continuation of] A catalogue of concho-
logical abbreviations. The Nautilus 4(10)T 1.5-1 17.
Baker, F (.'. 1891d. Descriptions of new species of Muricidae,
with remarks on the apices of certain forms Proceedings
ot Rochester .-Vcadem) of Science 1 129-137. i)l 1 I
Baker. F. C 1891e. Catalogue and s\iionym\ of the Recent
species of the famiK Muricidae. Proceedings of Rochester
Vtademy of Science 14.53-172.
Baker. V. C. 1892. Notes on a collection ol shells Ironi the
Mauritius, with a consideration of the genus Magilus of
Montfort. Proceedings of Rochester .Acadenn of Science
249-40, pi 9
Baker, F. V. 1.S94. Further notes on the eml)r\onie whorls of
the Muriciilae. Proceedings of the Acadein\ ot Natural
Sciences of Philadelphia 46:223-224
Baker, F. C. 1895a. A glance at the Chicago .\cadem\ of Sci-
ences. The Nautilus 8tl0):109-lll,
Baker, F. C. 1895b. The geographical distribution of the [land
and fresh-water] Mollusca. Science (New Series) 2(33): 179-
183.
Baker, F. C. 1895c. Prelniiinarx outline of a new classihcation
of the famiK Muricidae. Bulletin of the C^hicago Acadenn
of Sciences 2(2)169-189.
Baker, F. C: 1897a Critical notes on the Muricidae. Transac-
tions of the .\cadem\ of Science of St. Louis 7(16):371-
.391.
Baker, F C:. 1897b. On a collection of mollusks from Crand
Tower, Illinois. The Nautilus 1 1(3):28-.30.
Baker, F, C. 1897c. On the modification of the apex in gastropod
mollusks. .Annals of New York .Academx of Sciences 9:685-
704, pis. 18-20,
Bormann, M. 1946. .A survey of some West .American Oce-
nebras, with description of a new species. The Nautilus
60(2):.37-43, pi. 4.
Clench, W J and I Perez Farfante. 1945 Phe genus Murcx
in the western .-Vtlantic. Johnsonia I(17):l-58, pis, 1-29
Dall, \\ H 1889. Reports on the dredging, under the su-
pervision of Alexander .\gassiz, in the Gulf of Mexico
(1877-78) and in the Caribbean Sea (1.879-80), In the IS.
Coast Survey Steamer "Blake. Lieut. -C^ommander (^D
Sigsbee, L.S.N., and Commander JR. Bartlett, Ll.S.N.,
commanding X.\I.\ Report on the Mollusca; Part II, Gas-
tropoda and Scaphopoda. Bulletin ot Museum of C.om-
parative Zoology at Harvard C:ollege 18:1-492. pis. 111-
40.
Dall. \\ II 1919 Deveriplions of new species of Mollusca
Ironi the north PaciticOce.ui in the collection of the L'uited
States National Museum Proceedings of the II. S. National
Museum .56(2295) 29.3-37 I
Franzen, D, S. 1958 T\ pes ol molhisks described b\ 1" C
Baker - Part III. (Chicago .\cadein\ ol Sciences. The \.ui-
tilus 72(l):.3()-34
Fair. R. 1976. The Murcx book, an illustrated catalogue of
Recent Muricidae (Nhiricinae, Muricopsinae. Ocenebri-
nae). Priv.iteK published. Honolulu. Hawaii, 138 pp.. 23
pis.
Houart. R, 1990 New l,i\a .irid new recorils ot Indo-Pacilic
species ot .\/i/rc.v and lluiistclliuu (CJastropotla. Muricidae,
Miiricinae) Bulletin tin Museum .National d'Histoire Na-
turelle, Paris (Ser. 4) 12 (.sect. A) (2):329-347.
Houart, R 1992. The geiuis Chicoreus and relatetl genera
(Gastropoda: Muricidae) in the Indo-West Pacific. Me-
moires tlu Museum National d'Histoire Naturelle, Paris,
Zool. v.(A) 1.54:1-1.88,
Kaicher, S I^ 1973 Card catalogue of world-wide siiells,
Pack no 2 - Muricidae. Part I, nos. 99-195. Privately
published, St Petersburg, Floritla.
Kaiilier, S D 1978 (lard catalogue of workl-wiile shells.
Pack no 16 - Muricidae, Part HI, nos. 1540-1646 Pri-
\atel\ published, St Petersburg, P'torida
Kaicher, S D 1980 tlard catalogue of world-wide shells.
Pack no 24 - Thaididae. Part 1. nos. 2392-2497 Privately
published. St, Petersliurg. Florida
keen. .\ M 1971 Seashells of tropical west America. Marine
molhisks from Baja (."alifornia to Peru. Second Edition.
Stanford I niversitv Press. Stanford, California. \i\ + 106-1
pp., 22 color pis.
Lisclike, t;. F. 1869. Japanische Meeres-Concli\ hen. \olunie
1 Theodor F'ischer, C^assel, 192 pp , 14 pis
Palmer, K. \'. W. 1958. Type specimens of marine mollusks
described by P.P. Carpenter from the West Coast (San
Diego to British Columbia). Geological Society of .America,
Memoir 76. 376 pp.. 35 pis., frontispiece.
Petucli, E J 1990 .\ new molluscan faunule trom the Ca-
ribbean coast ol Panama. The Nautilus 104(2):57-71.
Ponder, W. J and E. H. X'okes. 1988. A revision of the Indo-
West Pacific fossil and Recent species of Miirex s.s. and
Haiislclluiu (Mollusca: Gastropoda:Mnricidae). Recordsof
the .Australian Museum. Supplement 81-160
Radwin, G.E and A. D'Attilio 1976. Murex shells of the
world; an illustrated guide to the Muricidae. Stanford llni-
versity Press, Stanford, California, 284 pp., .32 pis.
Reeve, L. \. 1845-1846. Conchologia Iconica, v. 3, Murex,
3« pis. (pis. 1-34 issued Apr to Oct., 1845; pis, .35, 36
issued Apr,, 1846)
Tryon, G. W 18.80 Manual ol (>'onchology, structural and
s\ stematic, w itli illustrations of the species, v. 2, Muricinae,
Purpnrinae Pliilatlelphia, Pennsv Kania, 289 pp., 70 pis.
\ likes E H 19(i3. Cenozoic Muricidae of the western .At-
lantic region. Part I - Murcx s s Tiilane Studies in Geologv
1(3): 9.3-12.3, pis. 1-4.
\ okes, E H 1971, Catalogue of the genus Murex Linne
(Mollusca: Clastropoda); Muricinae, Ocenebrinae Bulletins
of .American Paleontologv 61(268):l-l 41
\okes. E. H. 1990 Ceuo/oic Muricidae of the western .At-
lantic region Part \ 111 - Murcx ss, Hauftlcllum. Chi-
coreus, and Ilcxa))lcx, additions and corrections. Tulane
Studies in Geologv anti Paleontologv 23(l-3):l-96, pis. 1-
12. '
THE NAUTILUS 107(4);124-128. 1994
Page 124
Invasion of the Rivers of the Littoral Central Region of Venezuela
by Thiara granifera and Melanoides tiiberculata (Mollusea: Pros-
obranchia: Thiaridae) and the Absenee of Biomphalaria glabrata,
Snail Host of Schistosoma mansoni
J. p. Pointier
l„il)i>rjl(iirr dv Biiiliigif Marine ft
Malacoldgie
Centre ile Hiologie Tropicale
Ec'ole Prali<ine ties liante.s Etudes
.\\emie de \'illeneu\e
66860 Perpignan. France
R. N. Incani
lJt'|)artanientii de I'arasitulcigia
Facultad de Ciencias de la Salud
L'niversidad de C^araboho
\'aleneia, Venezuela
(". Balzan
P. Chrosoiechowski
S. Prypchan
Division de Parasitosis Intestinales y
Esquistosomiasis Direccion General
Sectorial de Malariologia y
Saneamiento Amhiental
Ministerid de Sanidad > Asistencia
Social
Maracav, Venezuela
ABSTRACT
Prior to 1975, some streams of the littoral central region of
Venezuela were active transmission sites for intestinal schisto-
somiasis. The snail intermediate host, BiomphuUnia glabrata.
was controlled but not eliminated b) repeated molluscicide
applications. Two thiarid snails, Thiara granifera and Melan-
oides tubereulata, began to invade these streams around 1975.
The rapid expansion of these two species was confirmed during
more recent surveys. Although other factors max be involved,
the exi.stence of stable populations ot tlie two thiarid species is
probabK linked to the absence of recolonization In B. glabrata.
the latter species has not been observed in the study region
during the last 15 \ears.
Key nords: Schistosomiasis, thiarid, recolonization
INTRODl'CTION
The littoral central region of N'enezuela iornis the north-
ern portion of an endemic area for intestinal schistoso-
miasis. Some of the rivers of tins region have previously
offeree!, and may still constitute, suitable habitats for the
snail intermediate host, Biumphalaria glabrata (Say,
1818); they may thus act as potential transmission sites
for tlie parasite (Incani, 1987). \ control program, based
cm I hemotherapy, sanitation, health education, environ-
mental improvements and api)lications of molluscicides
was iiiitiatet! in 194.). 'I'his control program resulted in
the near total inti'rruption ot transmission of Sehistosoma
mansoni (Sambon, 1907) during tfie 1970's (Incani, 1987).
Several malacological surveys carried out in the lower
course of the rivers before 1975 had revealed the pres-
ence of R. glalnata. More recent surveys, undertaken
between 1975 and 1983, and in 1990. demonstrated the
. ' \
10mm
Kipurc I. Thiara granijera [\fil; and Melanoides tubereulata
Uight).
J. p. Pointier et ai., 1994
Page 125
Figure 2. Map of the littoral ct-nlral region ol \ ene/.uela .showing the surveyed rivers
invasion of tliese rivers by two species of Thiaridae, Thiara
granifera (Lamarck. 1822) and Mclanoidcs tnlicrcnlata
(Miiller, 1774)(Figs. 1, 2).
Thiara granifera was introduced to tlie Caribbean
area around 1935 (Murray & Woopschal, 1965) and to
\ enezuela in 1970 (Chrosciechowski, 1973). It was first
observed in the littoral central region in 1975. The in-
troduction of M. tubcrculata to the Caribbean area prob-
ably occurred later. It was reported for the first time in
Te.xas in 1964, but according to Murrav (1964), it prob-
ably was introduced there several years earlier. M. tub-
crculata appeared in Venezuela in 1972 (Chrosciechows-
ki, 1973), and its presence in the rivers of the littoral,
central region coincided with that of T. granifera. This
paper presents the results of malacological surveys car-
ried out in all these rivers between 1975 and 1983, and
in 1990,
METHODS
Twent\-nine rivers were sampled along the littoral cen-
tral region of Venezuela (Fig. 2). All are permanent,
short and rocky streams descending rapidK' from the
mountains of the littoral cordillera; the U[)[)er stretches
constitute the typical habitat of the local thiarid snail
Pachycliilus lacvississimu.s (Sowerby, 1824). The lower
courses, often polluted in urbanized areas, provide suit-
able habitats for the snail B glahrata. OnK the lower
parts of the rivers were investigated during this study.
Qualitative data were collected on the presence or
apparent absence of the main species of snails before
1975, between 1975 and 1983, and in 1990 during the
control program monitored b\ the Ministrv of Health
(Chrosciechowski ct al.. 1987; Balzan, 1988). Samplings
using a quadrat method were carried out in si.x sites
located on the lower courses of the Cata, La Trilla, Ca-
muri Grande, .Anare, Los Caracas and El Botuco Rivers
in 1990. A\\ snails were manually collected and counted
in squares of 0.1 m" regularly spaced at one meter in-
tervals along four or five transects crossing the river.
RESULTS
Prior to 1975, seven of these rivers harbored B. glahrata
(Fig. 2 and Table 1) and were also active transmission
sites for schistosomiasis. These rivers were treated several
times with molluscicides (sodium pentachlorophenate)
during the 19(50's (.-Marcon de \o\a ct ai. 1987).
In 1975, T. granifera and M. tubcrculata were first
discovered in the Naiguata, .Anare and Los Caracas Riv-
ers, and in other rivers in subsequent years. General
survevs, carried out between 1975 and 1983, showed that
these two species hail become dominant in many rivers
(Table 1). In 1990, another survey demonstrated the in-
Page 126
THE NAUTILUS, Vol. 107, No. 4
Table I. Malai'ological i)b.scT\ati(iii,s made in t«ciit\ rivers of the littoral eeiilral region ol \'eiie/iiela B g Ruitttpluildrid filahrala,
I'.g. Tliiara graiiiffia. Ml Mclanoidcs ttihcrciilata. P.I. Parhyeliilufi Incvississimus.
Snail launa
Rivers
< 197,3
1975-198;5
1990
MdlluscieRle
application
(last \ear)
Sanclioii
Borlnirata
I'atanenio
San Esteban
l.a Trilla
Cat,.
(iiiarapitii
( lluironi
( 'liuao
El Liinon
Cliieliiriviche
L'rieao
Petaquire
Osorio
NEiciito
San Julian
Lria
N'aigiiata
Caniuri (irande
.\iiart'
Los (Caracas
El Botuco
La Seea
Osnia
Oritapci
Tixlasaiia
La Saljana
Carnao
( :liusp,i
B.g.
B.g.
B.g.
B.g.
B.g.
B.g.
B.g.
T.g./M.t. (198.3)
T.g./M.t. (1980)
T.g./M.t. (1977)
T.g./M.t. (1977)
r.g. (198.3)
T.g./M.t. (1976)
r.g. (1984)
T.g./M.t. (1980)
T.g./M.t
T.g/M.t
T.g/M.t
T.g/M.t
T.g. (1979)
T.g./M.t. (1978)
T.g./M.t. (1979)
T.g./M.t. (1979)
T.g/M.t. (1979)
T.g. (1979)
T.g. (1979)
T.g. (1979)
(1975)
(1979)
(197.5)
(1975)
T.g.
M.t.
T.g.
T.g./M.t.
T.g./M.t. /P.l.
M.t.
T.g.
T.g.
T.g.
T.g./M.t
P.l.
T.g/M.t.
T.g./M.t.
T.g.
T.g./M.t.
T.g./ M.t. /P.l.
T.g./M.t.
T.g./M.t.
T.g./M.t.
T.g.
T.g.
T.g./M.t.
T.g./M.t.
1967
1962
19S4
1962
1963
19(50
vasion of most of the rivers of this region by the two
thiarids and confirmed the absence of B. glahrota (Table
I).
Sampling carried out in 1990 by the ciuadrat method
determined the spatial colonization and densities of snails
in these six rivers. The two species of Thiaridae mainly
colonized the edges or protected areas in five of the six
sites (Figs. .3-14). These sites, with one exception, were
characterized by a strong current throughout the year
and the variable presence of rocks, stones, grt vel and
sand. Snail densities were not uniform, but showed an
aggregated distribution. In this ecological situation T
granifera was often more dense than M. tuhcrcuhita.
The observed maximum densities expressed as snails per
0 1 m- were of .50 T. granijcra and 67 M. tnherculata
lor ihcCata River; ,31 T. grant j era and 10 M. tuhcrculata
lor the La Trilla i^iver; 90 T. granifera and 1.5 M. tuh-
erculala for the Camuri (irande River; 23 T. granifera
and 5 M. tubereulata for the Anare River; and 46 T.
granifera and 9 T. tnherculala for the Los C-aracas River
(Figs. 3-14).
The sixth site, located on the El Botuco River, was
ecologically distinct and characterized by a sandy bottom
and very low velocities. Snail densities were low^ T. gran-
ifera reached a maximum dcMsil\ of 10 in<li\iduals per
0.1 m- and occupied mainK the center of the river,
w hereas M. tuhcrculata occurred at the edges at densities
as high as 9 individuals per 0.1 m- (Figs. 13-14).
DISCIISSION
Several biological control i^rograms using M. tuhcrculata
as a competitor ol Bioniphalaria .sp/J, were initiated in
the Claribbean area (Prentice, 1983; Pointier, 1989; Point-
ier ef al.. 1989). In four field trials on St. Lucia, B.
glahrata was apparenth eliminated from permanent
marshes and streams, 6 to 22 months after the introduc-
tion ot the competitor (Prentice, 1983). In Martinique,
M. tiiherciilata was introduced to groups of water-cress
beds and eliminated or strongly reduced densities of the
snail hosts, B. glahrata and B. straniinea. in less than
two years (Pointier ct al.. in press). In Xenezuela, the
introduction of thiarid snails to different types of habitats
in the Valencia lake region (Fig. 2) resulted in the elim-
ination of Bioniphalaria spp. from three artificial ponds.
However, results witc negative in several streams and
rivers (Pointier et al.. 1991).
The rivers of the littoral central region are verv short
and rocky, descending rapidly from the cordillera; the
water Nclocities ;ire often high. The thiarid snails mainly
J. p. Pointier et al., 1994
Page 127
Figures 3-8. Snail densities per 0.1 in- in tlirei' ri\ers (it tlie
littoral central regiiin (it N'enezuela, 3. C^ata River, Mclunuidcs
ttiberculala l. Cata Ri\er, Thiarn granijcra. 5. La Trilla Ri\-
er, M. tuherculata. 6. La Trilla River, T. granijera. 7. Canuni
Grande River, M. tubt'rculata. 8. (,'aniuri Grande River, T
grain j era
Figures 9-14. Snail densitit's per 0 I ni- in three rivers of the
littoral central region of Venezuela. 9. Anare River, Melanoides
tulicrculala. 10. Anare River, Thiara granijera. l\. Los Ca-
racas River, M. tuherculata. 12. Los Caracas River, T gran-
ijera L3. El Botuco River, M . tuherculata. 14. El Botuco River,
T. granijera.
occupv the edges and other protected areas (Figs. 3, 4).
Rocks and stones play an important role oBeriiig nianv
refuges against the strong current ami flooding. The riv-
ers of the N'alencia lake basin are ecologically different,
with substrates mainly of gravel and sand. Consequentlv ,
these latter rivers offer few suitable habitats for T gran-
ijera or especially for M. tuherculata (Pointier ct al.,
1991). The importance of water velocity as a limiting
factor for freshwater puimonate populations has been
described by several authors. Scorza et al., (1961) found
a negative hnear correlation between water velocities
and densities of B. glahrata in Venezuelan streams. In
Puerto Rico, B. glahrata populations did not survive in
portions of streams steeper than 20 meters per 1000 me-
ters of length (Harry and Cumbie, 1956). Jobin and Ippen
(1964) investigated the behavior of B. glahrata in irri-
gation canals and found that snails were dislodged when
the flow rate exceeded 65 cm/s. More recently, Dussart
(1987) studied the effects of water flow on the detach-
ment of several species of puimonate snails and found
significant differences between species. The resistance ol
thiarid snails to the current is not known, l)ut the distri-
bution patterns of the different species in the rivers of
littoral central region of N'enezuela indicate that T. gran-
ijera is better adapted than M. tuherculata. Experimen-
tal studies remain to be undertaken.
It is reasonably well established that T. granijera and
M. tuherculata began to invade the rivers of this region
around 1975. Before 1975 several streams were active
transmission sites for schistosomiasis and repeated mol-
luscicide applications allowed good control of the snail
host. However, in 19.S0, B. glahrata was again found in
the San Julian River and was responsible for the reap-
pearance of an important focus of S. mari.wni (Alarcon
de Noya, 1987). Several molluscicide treatments between
1980 and 1984 destroyed the B. glahrata colonies. Some
months after the last treatment, an important coloniza-
tion bv' T. granijera was noted. During the following
years, dense and stable populations of the thiarid snail
were established, and B. glahrata never reappeared. In
Puerto-Rico, a study of the distril^ution of freshwater
snails in a small stream showed that within five years,
T. granijera had displacetl B. glahrata from the central
portion of the stream, suggesting some form of compe-
tition (Butlers el al., 1980). Therefore, it seems probable
that the colonization of the San Julian River by dense
populations of the thiariil snail is now plaving a major
role in preventing the reappearance of the snail hosts.
The colonization of the rivers by the two species of
Thiaridae appears particularly efficient. When the cur-
rent is strong, T. granijera outnumbers M. tuherculata,
and the two snails are confined to the banks of the streams
Page 128
THE NAUTILUS, Vol. 107, No. 4
and to the protected areas. VVlieii the current is weak,
all the ri\er bed i.s colonized, l)ut 7". granifera occupies
the central portion of the habitat while M. tuherculala
occur.s at the edges.
The last nialacological sur\e\, carried out in 1990 in
the lower course of t\\ent\ three rivers demonstrated the
total dominance of the two thiarid species. However, a
few .v. punrtulata and /'. lacvissis.sinuis were collected
in the Oritapo, (]amuri Grande and Limon Rivers. The
current presence of stable and dense populations of the
two thiarids in nearly all the rivers of this region probabK
constitutes an imijortaiit deterrent to the reappearance
of B. glahrata, although other factors may be involved.
ACKNOWLEDGMENT
This work received financial sujiport from IINDP/World
Bank, WHO Special Progrannne for Research and Train-
ing in Tropical Disease. We should like to thank Dr.
Fergus McCullough, formerly Senior Scientist , WHO,
(ieneva. for his critical review of the manuscript. Vouch-
er specimens of snails are deposited at the Laboratoire
de Biologic des Invertebres Marins, Museum National
d'Histoire Naturelle de Paris, France.
LITERATURE CITED
Alarcon de Nova, B., O. Nova, l\ l rluuv, and J Hi.squez. 1987.
Reactivacion del foco l)i!liarziaiii)df Caraballeda en 1980-
83. Boletin de la Direccion ile Malarioli)i;ia \ Saiieamiento
Ambiental 27(l-4):86-9:5
15alzan, C 1988. Prograrna cic liiclia contra la t-squi.stosomiasis
eii Wnezuela. Ministerio de Sanidad y Asistencia Social:
l-ll
Butler. J \I , F. F. Ferguson. J \I Palmer and W H Johm
1980. Displacement of a colotu ol KuimphaUina glahrata
by an invading population of Tarclna granijcra in a small
stream in Puerto-Rico Caribbean journal of Science. 16
(l-4):7.3-79.
Chrosciechowski, P. 1973. In caiacol en busca dc niii-\a
re.sidencia. El Lago 30:813-814.
( ibrosciecliouski. P.. C. Ralzan, T. Camejo and B /Vlarcon de
Nova 1987 Recoleecion de caracoles, estimacion de su
poblacion en d habitat y traslado al laboratorio malacol-
ogico. In Cesari, N. & B. .\larc-on de Noya (eds.). Es-
ijuistiKumiasis mansoni diagnostic y control, p. 84-86.
lJu.ssart,(J. B.J. 1987 Effects of water How on the detachment
of some aquatic pulmonate gastropods .American Niala-
cological Bulletin 5( 1 ,):6.'5-72.
Harry, II W and B. G. Cmnbie 19.56 Stream ijradicnl as
a criterion of lotic habitats suitable for Auslralarhis fila-
hratiis in Puerto-Rico. .American Journal ol Tropital Med-
icine and Hygiene .5:921-928.
Incani, RN 1987 The Venezuelan experience in the control
of Schistosoma mansoni Memorias do Iiistituto Oswaldo
Cruz 82(,supp 41:89-93.
Jiibui. W, R, anil \ Ippen. 1964. Ecological design ol irri-
jiation channels. Science 14.5:1324-1326.
\1urra\, II I) 1964. Tarchia granijcra dwA Mclanindc\ tiih-
crcnlata in Texas. .American Malacofuical I iiion \rinnal
Report for 1964: 15-16. [Abstract}
\lurr.i\ 11 I) and L. J. Woopschall 1965 Ecology of Me-
lanoulcs tuherculala (Miiller) and Tarchia granijera in
South Texas. American Nialacological Union .Annual Re-
port for 1965:25-26. [Abstract],
I'onitier, J. P. 1989. Comparison between tuci biological con-
trol trials of Biomphalaria glahrata in a pond in (Guade-
loupe, French West Indies journal ol Medical and ,\pplietl
Malacology 1:83-95.
Poinlier. J P., A. Giiyard, and A. Mo.sscr. 1989. Biological
coiilml (il Biomphalaria glalnata and B. straminea by the
competitor snail Thiara tuhcrculata in a transmission site
of schistosomiasis in Martinique, French West Indies. .An-
nals of Tropical Medicine and Parasitology, 83(31:263-269.
Poinlier, J P., C Balzan, P. C^hrosciechowski, and R N Incani.
1991. Limiting factors in biological control ol the snail
\ectorsof scliistosomiasis in Wnezuela Journal of Medical
and .Applied Malacology 3:.53-67
I'rentice, M. .A. 1983. DisplncemenltA Biomphalaria glahrata
by the snail Thiara granijera in field habitats in St. Lucia,
West Indies. .Annals of Tropical Medicine and Parasitology
77:5 1 -.59.
Scorza, J. \'., J. Silva, L. Gonzalez, and R. Machado. 1961.
Stream velocity as a gradient in Aiislralorhis glahratiis
(Sav, 1818), Zeitschrift fiir Tropical Medi/in uiid Parasi-
tologic 12:191-196.
THE NAUTILUS 107(4): 129-135, 1994
Page 129
Corhicula and Neocorbiciila (Bivalvia: Corbiculidae) in the
Parana, Uruguay, and Rio de La Plata Basins
Cristian F. Ituartp
Musfii lie La I'lala
1900 La Plata
Buenos Aires, Argentina
ABSTRACT
CurlncuUl Megerle, 1811 is represented ni Arueiitniean waters
by Corhicula jlumuica (Mliller. 1774) and C'lnhu iila Idri^illicrli
(Pliiiippi, 1S44) The distrihution (il Corhicula ui the l'aran,'i,
L rnuua\ atui Hiii lie La Plata hasins is rexiewed liaved on
inihlished reports and eolleetions at the Mnseo de La Plata
iMLP) and Museo Argentino de t^iencias Naturales (M.A(A)
Sinee its intrmluetion about two deeades ago, Corhicula has
spread rapidK along the littoral regions of the Pio tie La I'lata
to Punto hullo on the western shore, and arroyo tAifre (.San
Jose n<'partTnent) on the eastern shore. The genus has reeentK
been reported to oeeur in the Uruguay River from Guale-
gna\ehn to Pio de La Plata .Additional reeords from the Parana
Piver at Paso de La Patria, Isia Barran(|uera and Bella Vista
are re|iorted In onler to elarif) the status and taxononn of the
eorbieulids of the area, a brief eomparative analysis ol the
morpholog\ of C'orhiiula and Scocorlniiila hischer, 1JS.S7, is
pro\ ided
Key i((;r(/\, Curhu ula. Scocorhicula. .\rgentina, (hslnbulmn,
introduced species
INTRODICTION
The family ('orliieuiidae is represented in the sciiitliern
neotropical region b\ the native genus Ncocorhiciila Fi-
scher, 1887, species of which were formerly attrihnted
to Corhicula Megerle, 1811 (e.g. Prime, 1865; Marshall,
1924, 1927). Parodiz and Hennings (1965) elevated the
rank and clarified the taxonomy of the genus Neocor-
bicula, earlier proposed as a section or subgenus of Cor-
hicula, and stated that South American taxa are referable
to Neocorhicula.
Species of the genus Corhicula ma\ have been intro-
duced into the Rio de La Plata basin l)etv\een 1965 and
1975. Specimens belonging to this genus were first col-
lected from the sandy banks at Punta Lara in February
1979 (Ituarte, 1981). In June 1978, Corhicula was col-
lected from the Jacui and Guaiba basins (Rio Grande do
Sul, Brazil), with the species identified by N'eitenheimer-
Mendes (1981) as Corhicula inanilcnsis (Philippi, 1844).
According to Morton (1979), tfiis species is a synonym
of C. fluminca (Miiller, 1774). Ituarte (1982) first re-
ported C. largillicrti (Philippi, 1844) at Colonia (Uru-
guay), on the eastern shore of the Rio de La Plata.
N'eitenheitner-Meiules and Olazarri (1983) provided
additional records along the L ruguay River from Las
Canas (south of Fray Bentos) to Pimta Gorda (Depart-
ment of Colonia) and from the eastern shore of the Rio
de La Plata to .\rroyo Cufre (Department of San Jose),
extending the known range of the genus. However, the
specimens collectetl in their sur\ey were not identified
precisely.
Ituarte (1984a) reported some biological aspects of a
population of C. largillicrti at Pinita Lara, Rio de La
Plata. Ituarte (1985) described the growth d\iiamics in
a natural population of C. fluminca at Pimta Atalaya,
Rio de La Plata, extending its known range along the
western shore of the estuary . In a brief communication,
Olazarri (1986) suggested the presence of two Asiatic
species of Corhicula . C. largillicrti antl C. fluminca. in
the L'ruguay Basin.
In the present paper, further data concerning the mor-
phological characterization ol the genera Corhicula and
\cocorhicula are proxided. The range of Corhicula spe-
cies in Argentina has been extended to the Parana River,
and the species inhabiting the Uruguay River are iden-
tified. The distribution of the genus Corhicula along the
Rio de La Plata is reviewed and variation in shell shape
due to environmental factors is discussed. N'oucher spec-
imens from this study are deposited in the invertebrate
collections of the Museo de La Plata and Museo Argen-
tino de Ciencias Naturales.
RESULTS
The .Asiatic Clam genus Corhicula is represented in Ar-
gentinean and L ruguayan waters by two species, C. flu-
minca and C. largillicrti. Present distribution of the
genus in the Rio de La Plata estuary (fig. 4) spans the
entire w estern shore from the delta of the Parana River
to Magdalena Cit\' (35°02'S, 58°28'\V) stretching ap-
proximately 75 km along the sandy intertidal zone. Bot-
tom deposits are composed of fine sands and Parana River
sediments. Beyond Magdalena City, only isolated indi-
Page 130
THE NAUTILUS, Vol. 107, No. 4
I'if^iiri's !-.'{. Scoii/rlni iild liiiKisii I. I'loiital scclioii .shovviiij;
spatial segregation ot male (ventral) and female (dorsal) tissues
within the visceral mass. 2. Sagital section oi the inner demi-
branch showing embryos brooded within the water tubes Co-
horts of different ages are visible .'5. Detail ol ligure 2, showing
the advaneeil stage ol development ot embryos. .All scale bars
= 0.5 mm
viduals of Corlncula were found. Darrigraii (personal
comrminication) collected both (.'. fluminca and C. lar-
gillierti at Piinta Indio.
The dislrihiition of Curhicula along the eastern shore
of the Rio lie la Plata estuary is diseontiiiiious. \eiteii-
heimer and Olazarri (1983) had reported several records
of Corlncula at San Jose and Colonia Departments with-
out iilentilieation to species. The only fully identified
record of L'ruguayan Corlncula in the Witt ile la Plata
River is a single lot of C. largillicrti from Ciolonia (MACN
30175) identified by Ituarte (1982). Along the southern
[jortions of the Rio de la Plata estuar\, Curhicula pop-
ulations seem to have been limited by substratum char-
acteristics (increased clay content) and a progressive in-
crease in salinity caused by inflow of sea water during
maximum high tides.
Three new records for both C. fluminca and C. lar-
gillicrti from the upper course of the Parana River are:
Paso de La Patria, Isla Barranquera and Bella Vista
(Province of Corrientes). Only C. fluminca was collected
at San Nicolas, on the lower course of the Parana River
(Ituarte, 1982).
The Parana River flow s into the Rio de La I^lata estuary
through a wide delta system, where both C. fluminea
and C. largillicrti have been collected The former spe-
cies seems to be more abimdaiit in the Sarmiento River,
the mouth of the Lujan River, and the .\rro\ <i Abra \ieja.
The genus Corhicula is present in the Uruguay River
from Nueva Palmira to Fray Bentos. Both C. fluminea
and C. largillicrti were found in nearly all of the 41
samples taken in 12 transects of the river in this area.
N'eitenheimer-Mendesand Olazarri (1983) reported Cor-
lncula from the following localities along the Uruguayan
coast without identifying the species: Department of Co-
lonia - Punta Gorda, Nueva Palmira; Department of
Soriano - Agraciada, CA)ncordia, Rio San Sakailor, Do-
lores; Department of Rio Negro - Las Canfias Rivulet
(southward to Fray Bentos). Of the many watercourses
that flow into the Uruguay River from the Argentine
shore, C largillicrti was found in oiiK one, the Parana-
cito Rivulet, This species was also collectetl at Santo Tome
(C^orrientes), the northernmost record Irom the Uruguay
River.
Shell shape and especially shell outline are aftected by
such environmental factors as substrate composition, hy-
drodynamic forces, and other hydrological characteris-
tics, inchKling tidal regime. A typical trigonal shell out-
line, inllateil beaks and unerodetl umbos (extended into
a rostrum posteriorly) are characteristic ol individuals of
bl, br.uicliial filament; bp. w ater tube expanded In Inrni a brood
pouch, e, embrso; 9, female tissues, 6, male tissues, I, inti'stine.
id, inner demibrancli, i] inlerlamellar junction, nic, mucous
cells; 1, iccliim, S, sloin.uli, wl walei lube
C. F. Ituarte, 1994
Page 131
-30-
San
BUENOS
-35
Dib. C.R.Tremouilles
Figure 4. Distriliiitioii of Corbicula species in the sniitliern Neotropical region, including new localities for C fluniinea and C.
lareillivrti. See text lor details.
C. fluminca that inhabit lentic and permanent en\ iron-
ments with a soft substratum composed of mud or silt
with \ariable sand content. Habitats composed of sand
bottoms, with flow iiig water and or that are tidali) af-
fected contain specimens with a rounded or oval shell
outline, and consequently higher Length/height (L/H)
and length/width (L/W) ratios.
E.xamples of tiie effects of environmental factors on
shell morphology can be found in the Rio de La Plata
at Olivos. Large populations of both C. largillierti and
C. fluminca were present along the extended sand flat
in the intertidal portion of the estuary. Hydrological
conditions are strongly affected by the reduced slope in
the intertidal. Tidal flows and waves increased in some
cases by southeastern winds produced shells with oval
outlines, eroded umbos and high values of L/H and L/
Page 132
THE NAUTILUS, Vol. 107, No. 4
4>
^
O
a^'
4fly 4^
^^
8
4r
■\
#
■;«■
##
Fij»ure> }{-•>. Si-rii". nl spriiiiifiis taken lidin Iklal pdiuls (left)
and MiiKU inlertiflal (rii;iit) luihilats in ()li\().s, Huenoi .Aires,
Rio de La Plata 8. Citrhicula largillicrli. '). Corhicula flumi-
iicd \ll scilc liais = Ml Tnni
l-'if;urc> .'>-7. ( ^>rl)itiiiiils liiini Hid ilc La I'lata. .S. Scacor-
Inciila hmosa (Matoii, 1809) froni llir aiT(i\o Migiielin 6. Ci>r-
hiciila /(n7;i7/uTh' (Pliilippi. ISlli ironi Piinta lara. Kii.scnada.
7. Corhicula fhimincii (Miillcr. 177 1), Ironi Olivos, I?iieiio.s
Aires ,\ll scale hars = 10 mm.
C. F. Ituarte, 1994
Page 133
'I'ahl.- I. Slirll IfTijitli slic-ll lifiiihl (LH) ami slicll lciii;lli shell ukUIi (L W I rutins lor (,'. larnillurti and (' jhinuma pcipulatioiis
licirii ililltTfiil fiivlroiimfnls at ()li\i)s. Win dr I, a I'lata (.v = nu-aii. S = staiidanl iIcn iaticjii )
(.'. ftuiuinca
C
. Iargilli(
■rli
1. H 1. w
n
A
S
X S
101
101
1 1:5
1 20
0 02
oot
1 5h 0 05
1 iiT 0 07
II
L/W
Sdit Ixittcmis
SaiicK intcrtiilal
1 17
I 20
0 04
0 0.)
I SS
0 05
0 10
W (figs. 8,9; Table 1 ). Populations that srttleii in substrate
depressions along the upper intertidal zone, vvhieh, like
tide pools of the rocky marine intertidal areas, constitute
small, lentic, micro-habitats with sand bottoms contain-
ing substantial amounts of mud or silt, luue a typical
trigonal shell outline, inflated beaks, thick shells, ami
lower L/W and L/H ratios. The posterior projection ot
the rostrum occurs only in Corhicula fliimiriea (Figs. S,9;
Table 1).
DISCUSSION
Parodiz and Hennings (1965) reported that the family
Corbiculidae is represented in the southern neotropical
region b\ the native genus Neocorhictila Fisher, 1887
In Argentina, Mcocorbicula liinu^a {Maton, 1809) is found
along the Llruguay and the Rio de La Plata Rivers, while
N. parancnsis (d'Orbigny, 1846) occurs in the Parana
River Basin. Although a considerable bod\ of evidence
has been presented to warrant the segregation ol South
American forms within the genus Ncocurhicula (Parodiz
& Hennings, 1965; Dall, 1902; Ituarte, 1984a, b, 1986)
many authors persist in referring autochtonous taxa to
the genus Corhicula. Conclusions based on this premise,
including biogeographic (Counts, 1980; McMahon, 1983)
and shell morphometric comparisons (Britton 6; Morton,
1979) are erroneous.
Corhicula and Neocorbicula differ substantially in shell
morphologv. The wide range of shell \ariability show n
by both genera in response to environmental factors re-
veals that the most relevant character is the presence in
Neocorhicula of a deepK marked pallial sinus (siphon
lengths 12-14 mm). In Corhicula the pallial sinus is ab-
sent or represented by a slightK curved pallial line, and
siphons as well as associated retractor muscles are scarce-
ly developed ( siphon lengths 3-4 mm) (figs, 5-7).
Branchial incubation of embryos, a characteristic com-
mon to both genera, is carried out in different ways.
Corhicula incubate embry os in large rnnnbers [ > 10,000,
as calculated for C. largillierti (Ituarte, 1984a)] within
the water tubes of the tw o inner demibranchs. The basic
structure of the inner demibranchs of Corhicula is un-
modihetl, and differs from the outer demibranchs only
in ha\ing fewer interlamellar junctions. Seocorhicula.
on the other hand, develops brood pouches by cellular
proliferation of the interlamellar junction epithelium of
the inner demibranchs. Each brood pouch contains a
single embryo. Gravid indi\iduals usually have 20-30
brood pouches, and occasionalK as many as 45 (Ituarte.
1984b) figs. 2,3). The incubation period of Corhicula is
\er\ short, with embryos released at a late veliger or
pediveliger stage (200-240 /um diameter) (Ituarte, 1984a;
Britton & Morton, 1979). In Ncororin'riWa, the incubation
period is longer and the offspring are released as fully
developed juveniles Birth size in Mcocorhicula is about
1.1 mm. Howe\er, some indi\iduals from each cohort
remain in the maternal gill until the\ reach 4-5 mm in
shell length (Ituarte, 1984a) (figs. 2,3). In Corhicula. all
embry OS w ithin a gra\ id indi\itlual show a similar degree
of ile\ elopment, while in Scocorhicula tw o or three gen-
erations of larvae may coexist w ithin each demibranch
(Ituarte, 1984b) (fig. 2).
Those species of both genera that occur in the Amer-
icas are simultaneous hermaphrodites. However, there
are differences in gonadal structure. During the early
stages of gonadal development of Neocorhicula the pre-
dominant elements in the acinar wall are large and vac-
uolated follicular cells, w ith germ cells scattered along
the base of the epithelimn (Ituarte, 1986). In Corhicula.
follicles are composed of minute follicular cells, with
gonia the dominant elements in the acinar wall (Ituarte,
1984a). In Corhicula there is no pronounced segregation
of male and female germinal cells within the gonad. If
any degree of localization exists (Kraemer, 1978), it dif-
fers from that shown in Neocorhicula, where the ovary
occupies the dorsal region of the gonad, while the testes
develop in the ventral half (Ituarte, 1986) (fig. 1). Fur-
thermore, in Neocorhicula. testes occupy more than 50'x'
of the gonad (Ituarte, 1986) while in Corhicula. they
occupy 15-30% (Kraemer & Lot, 1977; Britton & Morton,
1979).'
Corhicula largillierti was tiescribed from the Yangtse-
Kiang River in China, and its range includes northern
and Central China as well as the Korean Peninsula (Pras-
had, 1929). The taxonomy of the Asiatic species of Cor-
hicula was reviewed by Morton (1979), who reanalysed
Prashad's (1929) "affinities ' and regarded C largillierti
as belonging to a group of species that are synonymous
with C. fluminea (Morton, 1979: Table 11). However, a
re-examination of plots of affinities of Chinese, Korean
and southeastern Russian species reveals C. largillierti to
be distinct from an\ other species (Morton, 1979: Table
6). Moreover, there is no evidence in the text of Prashad
(1929) to support the s\iion\m\ of C largillierti with
C. fluminea.
The material studied and assigned to C. largillierti.
Page 134
THE NAUTILUS, Vol. 107, No. 4
which i.s now widoK (listril)utecl in Argentinean waters,
show.s morpliological ditierenee.s from C. jluminea that
are sufficient to to consider it a distinct species. Corbicula
largillierti is characterizetl In its inoch'rateiN thick shell,
with fine, clo.seK spaced concentric ridges and \ellowish
or greenish surface. Its shell is trigonal, nearly equilateral,
and not greatly inflated Its beaks are moderately inflated,
the posterior end slightK [irotrucled and obtuse at the
c\tremit\, e\en in larger indisiduals (fig. 6) Corbicula
fluniinca differs from C. lar^illicrti in having a more
solid shell, with coarser, more widely spaced surface sul-
cations. Its shell is more inflated and globous, with prom-
inent and inflated Ix-aks In C. fluminea. the posterior
margin is markedlv' protruded to form a characteristic
rostrum, especially in specimens larger than 15 mm (fig.
7).
In spite of the liighly variable shell morphology that
results from environmental factors, the differentiating
features listed above do not become masked, and the two
species are easily distinguished even when populations
of both species coexist in the same habitat.
ACKNOWLEDGMENTS
The author wishes to express his gratitude to Dr. Tad-
ashige Habe (National Science Museum, Tokyo) for his
assistance in the identification of specimens of Corbicula
largillierti. Thanks are also due to the following for pro-
\ iding specimens: Dr. F. Spinach Ross (Proyecto Eval-
uacion de Recursos Pesqueros del Rio Uruguay, CARU-
INIDEP-INAPE); Mr. R. Dillon, Mrs. A. Ibargoyen, Dr.
J. Garcia, Lie. F. Kaisin, Lie. N. Laiidoni, and Dr. A.
Rumi.
LITERATURE CITED
linttmi, J. C , D. R. Coldiron, L. P. Evans, Jr. C. GolightK, K
U. O'Kane, and J. R. Teneyck. 1979. Reevaluatioii of
the growth pattern in Corbicula fluminea (Miiller). [in]:
Brittoii, J. C. (ed.) Proceedings, First International Cor-
bicula Symposium. Texas Cliristiaii L'niversity Research
F"oundation, Fort Wortii, Texas, p. 15-3><.
Brittoii, J. C. and B. S. Morton. 1979. Corbicula in Nortii
America: The evidence revievvwl and evaluated [in]: Brit-
ton, J. C. (ed.) Proceedings, First liUeriiational Oirbicuia
Symposium. Texas Christian University Researcli Foun-
dation, Fort Worth, Texas, p. 2-19-287
Britton, J. C. and B. S. Morton. I9S6 Fob inorpiiism in Cor-
bicula fluminea (Bivalvia: Corbiculoiilea) from Ncirtii
.Vmerica Maiacologieal Beviev\ 19:l-4'5
Counts, C \. 1980. The •^vwn^ (.'■orbicula Mulilteld (Bivalvia:
Corbiciilidae) in .Africa and South America: Zoogeograpli-
ic and taxonomic prol)lems. Bulleliii nl llic Xnicncaii \lal-
acological Union for 1980:71-72
Dall. W. H. 1902. Noteon.\Vo™r/«ci//</ ImmIici- TIh' Nautilus
16:82-83.
Hayashi, Y. 1956. On the variation ol C.iirlii( iiln dui- t<i en-
vironmental factors. Venus l9(l):54-60.
Ingram, VV M 1918. The larger fresh water cl.iins ot Cali-
fornia, Oregon and Washington Journal ol I'ulouiology
and Zoology 40(-l):72-92.
Iluarte, C. F. 1981 Primera noticia acerca de la iiitroduccion
de pelecipodos asiaticos en el area rioplateusc NCotropica
27:79-82.
Ituarte, (;. F. 1982. C^oulribuciou al conocnniculo dc la bio-
logia de la tamilia Corbiculidae (Mollusca Pelecypoda) en
el Kio de La Plata. Doctoral Tlu-sis No 408 Facultad de
(aeiicias Naturales v Museo. I iiiversidad Naeional de La
Plata.
Ituarte, C". F. 1984a. .Aspectos bioioijico.s dc las poblaciones
de Corbicula largillierti iPliilippi. 1844) (Mollusca Pele-
cypoda) en el Kio dc La Plata Revista del Museo de La
Plata (nueva serie) 13(1 431:231-247.
Ituarte, i'. F. 1984b. El fen(Smeno de Ineubacion branquial
en Neocorbicula limosa (Maton, 1809) (Mollusca Pelecy-
poda). Neotropica 30(83):43-54.
Ituarte, C. F, 198.5. Growth dynamics in a natural i)opulatioii
ol Corbicula fluminea (Bisaivia Spliacriacea) at I'unta
Atalaya, Bio de La Plata, .\rgeutina. Studies on Neotrop-
ical Fauna and Environment 20(41:217-22.5.
Ituarte, C. F. 1986. Contribucion al conocimicMlo dc la Inol-
ogia reproducliva de Scocorbicula limosa (Maton, 1809)
(Mollusca Pelecypoda). .\nales de la Sociedad Caentifica
Argentina 214, I No. 47:1-27.
Kraemer, L 1978. Corbicula fluminea (Bi\aKia: Sphaeri-
acea): The functional morphology of its hermaphroditism.
Rullclin ol die .\merican Malacoloi;Jcal I niou. 197S40-
49
Kraemer, L. and S. Lott. 1977. Microscopic anatomy of the
\ isceral mass of Corbicula (Bivab ia: Sphaeriacea). Bulletin
ol the .American Malacological L niou. 197748-55.
Marshall, W B 1924. New Uruguayan Mollusks of the genus
Corhictilii Proceedings of the L'uited States National Mu-
seum ()(n2552):l-14.
Marshall, W. B. 1927. New species of mollusks ol the gciuis
Corbicula from Uruguay and Brazil. Proceedings ol the
United States National Museum 72(2699):l-7
McMahou, R F. 1983. Ecology of an invasive pest bi\al\e,
Corbicula In: K. M. Wilbur (ed 1 The Mollusca. .\cademic
Press, New York, 6:463-561.
Morion, n S 1979. Corbicula in Asia. In: Brillon, J. C. (ed.)
Proceedings, Urst International Corbicula Symposium.
Texas Christian University Research Foundation. Fort
Worth, Texas, p. 15-38.
Ola/arrl, J 1986. Las alinejas del genero Corbicula en el rio
Lruguay Hesinnenes de Comunicaciones, Seminario "El
rio Orugnay y sus Recursos" CAKl -INAPF-INIDFP, Entre
Bios, Argentina.
Parodiz, J. J. and L. Henniiigs. 1965 The Scocorhicula (Mol-
lusca Pelecypoda) of the Parana-L ruguay basin. South
America. Annals of the Carnegie Museum 38(3):69-96.
Prashad, B. 1929. Revision of the .Asiatic species of the genus
Corbicula III The species of the genus Corlncula from
China. Tibet, Formosa and Philippine Islands Memoirs,
Indian Museum 949-68
Prime, T. 18()5, Monograph of .\meriiau Corbiculidae (Re-
cent and fossil). Smithsonian Miscellaneous ( iolleclious 71-
80
SukI.ui, H M .nul B (;. Isom. 1963. Further studies on the
mtroihmd .\siatic clam Corbicula in Tennessee. Tennes-
.see Stream Pollution Control Board. Tennessee Depart-
ment of Pulilii- Health, N'aslnillc. Tennessee, 76 p.
Smith, M 11 . J C Button. P Buike, K K Chesser, M. W.
Suulh, ,iuil J ll.ii;cn. 1979. Cenetic variability in Cor-
buiila. ail iiiv.iding species. In: Britton, J. C. (ed.) Pro-
C. F. Ituarte, 1994
Page 135
ceedings. First Interiialiniial Cdrlncula S\ nipusium. Texas
Christian L'iiiversit\ Research Founilatioii. Kurt Wnrlh,
Texas, p. 243-248
N'eilenlieimer-Mendes, I L. 1981- C'orhicula inanili'u.'iis [Fhi-
Hppi, 1844), mohisco asiatico, ria liacia do Jaeui e do Guai-
ba, Rio Grande do Sul, Brasil iBi\al\ia Gorbieuhdae).
Iheringia (Zoologia) 60:63-74.
\ ciU-nlieimer-Meiides, I F anil J Olazarri. 1983. Primeros
registros de Curhicula Megerle, 1811 (Bivalvia Corbicu-
hdae) para el rio Iriiguav . Boletiii de la Sociedad Zoologica
ilel L rugua\ 1 50-53.
THE NAUTILUS 107(4):136, 1994
Page 136
Sea Slugs and their Relatives of Western Australia
by Fred E. Wells and (Clayton \V. Bryce. 1993. West-
ern Australian Museum, Perth, viii + 184 pp., 227 figs.,
1 map. Softcover, $30.00 U.S. ISBN 0-7309-5523-0.
The opisthobranch gastropods or "sea slugs" of Western
.Australia have been gloriousK introduced in this new-
book, a well-matched companion volume to Seashells of
Western Australia by the same authors (Western Aus-
tralian Museum, 1986) which treated only 16 species of
opisthobranchs. Produced in virtually the same size and
organization as the Seashells volume, it is also based on
the collections and research activities ot the Western
Australian Museum in Perth, undoubtedly including the
results of a series of international workshops, the last held
in .Albain in 1988. The authors extend this work as a
sample rather than a complete coverage of this diverse
fauna, but its a substantial snack, at any rate, of 224
species in 42 families and 6 orders (Cephalaspidea, The-
cosomata, .Anaspidea, Notaspidea, Sacoglossa, and 147
species of Nudibranchia). .\11 but a tew are beautifulK
illustrated as living, craw ling creatures in full (sometimes
gaudy) color; only a few bubbles and pteropods are pre-
sented as empty shells.
Each higher ta.xon is briefly diagnosed and most are
accompanied by a useful list of references from the pri-
mary literature (all but a few from the 1960's or later).
Species descriptions include size, rarity, common syn-
onyms, and overall geographical range plus specific oc-
currences in Western Australia. Again matching theSca-
shells \ olume, the geographical scope of this book includes
the entire coastline of the state of Western .-Australia,
from the tropical Kimberly region to the temperate Re-
cherche .\rcliipelago. The authors planned to (and in
fact do) include representatives of each of the si.x orders
from each of the three major zoogeographical regions,
w ith over 40 cited localities. Added to this volume are
records from late-1980's collecting on Christmas Island
and the Cocos (Keeling) Islands, Australian Territories
in the Indian Ocean. Although rather out-of-place, these
18 species records (7 of which are not found in Western
•Xustralia) do not seriously detract from the unity of this
\()lunie. Nevertheless, they might ha\(' been more ef-
fecliveK presented as a separate primar> publication.
Both the pages preceding and following the main tax-
onomic .section reflect tho.se presented by the Seashells
volume, with appropriate motlifications and uptlated in-
formation. The introduction covers various aspects of
opisthobranch biology (behavior, feeding, life histories),
and the chapter on zoogeography has been expanded.
C;hapters on collecting and cleaning methods, covered
f ulK in the Seashells \filume, are understandabK absent,
in deference to .Australia s recent collecting restrictions
as well as opisthobranchs' rare standing in the average
shell collection. Nothing is mentioned, however, on How-
to Photograph Sea Slugs, which would have been enor-
niousK welcome, since the photographs of li\ ing animals
(mam in obviousK underwater surroundings) are the
highlight of this book. A list of Western Australian shell
clubs, a brief glossary (but not nicely illustrated as in the
Seashells volume), and taxonomic index conclude the
volume.
By this book, the authors have sought to "stimulate
interest" and "demonstate both what is know n, and also
what is not known" about Western -Australian opistho-
branchs. They have succeeded on both counts through
their recognition of ongoing research and of the unre-
solved taxonomic problems in this group. Forty-five spe-
cies are identified to genus-level only (or with a tentative
species assignment), with some noted as new to science
(although, appropriately, none are originally described
in this book). Over half are stated as rare or uncommon;
many species are recorded from single localities. Ten
species were described in the 1980 s, another 10 in the
1990's (4 in 1993), as credit to the research efforts of a
number of active opisthobranch biologists. One photo-
graph of a living holotype (Chromodoris aurigera Rud-
man, 1990) is included I cannot detect any glaring iden-
tification errors (although the spelling of .Ajilustritlae is
incorrect), and I happily note that the overall SNStematic
treatment is up-to-date (perhaps due in part to a "who's
who " list of contributors): Akera is placed in .Anaspidea,
CijUndrolmUa in Sacoglossa, and Apltistrum anipluslre
in Ihjdatina, three recent changes often overlooked by
non-specialists. Some might contest the wording of a few-
evolutionary statements (e.g., "elysiids have lost their
shell"), but I think these are excusable in this context.
Wells and Bryce have elegantly met their intended
goal to illustrate the extraordinary diversitv of living
opisthobranchs from the long, winding coastline ot West-
ern Australia. Attendees at future Western Australian
Workshops (the next one in the Houtman Abrolhos in
1994) have a excellent guide to follow .
Paula M. Mikkelsen
21^1 .3U
INSTRUCTIONS TO AUTHORS
THE NAUTILUS publishes papers on all aspects of the
biology and systematics of mollusks. Manuscripts de-
scribing original, unpublished research as well as review
articles will be considered. Brief articles, not exceeding
1000 words, will be published as notes and do not re-
quire an abstract. Notices of meetings and other items
of interest to malacologists will appear in a news and
notices section.
Manuscripts: Each original manuscript and accompa-
nying illustrations should be submitted in triplicate. Text
must be typed on one side of 8'/2 x 11 inch white paper,
double spaced throughout (including literature cited, ta-
bles and figure captions), with at least 1 inch of margin
on all sides. All pages must be numbered consecutively.
If printed on a word processor, the right margin should
be ragged rather than justified. Authors should follow
the recommendations of the Council of Biology Editors
Style Manual, which is available from the Council of
Biolog> Editors, Inc., 9650 Rockville Pike, Bethesda, MD
20814, U.S.A. The first mention of a scientific name in
the text should be accompanied by the taxonomic au-
thority, including year. Latin names and words to be
printed in italics must be underlined; leave other indi-
cations to the editor. Metric and Celsius units are to be
used.
The sequence of sections should be: title page, abstract
page, introduction, materials and methods, results, dis-
cussion, acknowledgements, literature cited, tables, fig-
ure captions, figures. The title page should include the
title, author's name(s) and address(es). The abstract page
should contain the title and abstract, which should sum-
marize in 250 words or less the scope, main results and
conclusions of the paper. The abstract may be followed
by a maximum of 8 key words. All references cited in
the text must appear in the literature cited section and
vice versa. In the literature cited section, all authors
must be fully identified and listed alphabetically. Follow
a recent issue of THE NAUTILUS for bibliographic style,
noting that journal titles must be unabbreviated. Infor-
mation on plates and figures should be cited only if not
included in the pagination. Tables must be numbered
and each placed on a separate sheet. A brief legend must
accompany each table. Captions for each group of illus-
trations should be typed on a separate sheet and include
a key to all lettered labeling appearing in that group of
illustrations.
All line drawings must be in black, high quality ink,
clearly detailed and completely labeled. Photographs
must be on glossy, high contrast paper. All figures are
to be consecutively numbered (figs. 1, 2, 3, ... , NOT
figs. la. lb, Ic, . . . NOR plate 1, fig. 1 . . .). Illustrations
must be arranged in proportions that will conform with
the width of a page (6% inches or 171 mm) or a column
(3'/4 inches or 82 mm). Tlie maximum size of a printed
figure is 6% by 9 incites or 171 by 228 mm. All illus-
trations must be fully cropped, mounted on a firm, white
backing, numbered, labeled and camera ready. The au-
thor s name, paper title and figure number(s) should ap-
pear on the back. Original illustrations must be between
one and two times the desired final size. It is the author s
responsibility that the line weight and lettering are ap-
propriate for the desired reduction. Original illustrations
will be returned to the author if requested. Color illus-
trations can be included at extra cost to the author.
Voucher IVIaterial: Deposition of type material in a
recognized public museum is a requirement for publi-
cation of papers in which new species are described.
Deposition of representative voucher specimens in such
institutions is strongly encouraged for all other types of
research papers.
Processing of Manuscripts: Upon receipt, every manu-
script is acknowledged and sent for critical review by at
least two referees. These reviews serve as the basis for
acceptance or rejection. Accepted manuscripts are re-
turned to the author for consideration of the reviewers'
comments. A finalized version of the manuscript is re-
turned to the editor and sent to press. Two sets of proofs
are sent to the author for correction. Changes other than
typesetting errors will be charged to the author at cost.
One set of corrected proofs should be sent to the editor
as soon as possible. Authors with institutional, grant or
other research support will be billed for page charges at
the rate of $60.00 per printed page.
An order form for reprints will accompany the proofs.
Reprints ma\' be ordered through the editor.
Manuscripts, corrected proofs and correspondence re-
garding editorial matters should be sent to: Dr. M.G.
Harasewych, Editor, Di\ision of Mollusks, NHB stop 118,
National Museum of Natural History, Smithsonian In-
stitution, Washington, DC 20560, USA.
@ This paper meets ttie requirements of ANSi/NISO Z39. 48-1992 (Permanence of Paper).
MBL/WHOI LIBRARY
IJH 17Yb B
Live Music Archive
Librivox Free Audio
Metropolitan Museum
Cleveland Museum of Art
Internet Arcade
Console Living Room
Books to Borrow
Open Library
TV News
Understanding 9/11