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Full text of "The Nautilus"

FHE NAUTILUS 











Volume 107, Number 1 
March 24, 1993 
ISSN 0028-1344 




A quarterly devoted 
to malacology. 








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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 



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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 °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 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 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 


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 t urn i d 
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 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 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 


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^ 


^ 


^ 





km 

1 


' ^-— 


^^=^^8=- 


^^2 


\ 






^ ^ 


1^ ^^ 


% 

■^1 


DB Jy 


^"¥1 


i 


,54-70 




^) 


Ftl 



W/!?\ TerrigencHJS grainstcxie facies 
M i::::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; 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 




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: 








3 
I 






E. 



c ^ 1= _3 



■a 


^ 






^ 


V 




-b 


cc 


o 


<3 




t-. 


r^ 



VZ^^r77T77x y77777\^ 




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. 



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Abbott, R.T. 1958. The marine moliusks of (irand C'ayman 
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Abbott, R.T. 1974. American seashelis (2nd ed.). Van Nos- 
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Bengston, P. 1988. Open nomenclature. Palaeuntulogv 31: 
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Cerridwen, S.A. and B. Jones. 1991. Distribution of bivalves 
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de Jong, K.M and H.E. Coomans. 1988. Marine gastroputis 
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Hill, R.T. 1899. The geology and plusical geograph\ of Ja- 
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256 p. 

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Jung, P. 1972. Moliusks from the White Limestone Group of 
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Land, L.S. 1991. Some aspects of the late Cenozoic evolution 
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45-48. 

Land, L.S. and S. Epstein. 1970. Late Pleistocene diagenesis 
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Larson, DC 1983. Depositional facies and diagenetic fabrics 
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Liddell, W.D.,S.L. Ohihorstand AG. Coates. 1984. Modern 
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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 
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Parsons, KM. and C.E. Brett. 1991. Taphonomic processes 
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Petuch, E.J. 1988. Neogene histor\ of tropical .American mol- 
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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 
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Robinson, E 1963. Quaternary deposits. In. Zans, V A., L.J. 
Chubb, H R. Versey, J B Williams, E Robinson and D L. 
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Geological Survey Department of Jamaica 4: 50-54. 

Taylor, J.D. 1978. Faunal response to the instability of reef 
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Trechmann, C T. 1930. The Manchioneal Beds of Jamaica 
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N'alentine, J.W 1989 How good was the fossil record? Clues 
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Page 42 THE NAUTILUS, Vol. 107, No. 1 



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vii + 222p. 



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Volume 107, Number 2 
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A quarterly devoted 
to malacology. 



LIBRARY 



AUG 61983 



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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 










1 




1 




2. Calcitic layer 







1 


1 




1 




3. Number of aragonitic layers 







1 


2 




2 





-4 Position of opercular nucleus 







3 


1 




2 




5. Opercular shape 







1 







1 




6. Pigmentation pattern on head-foot re- 
















gion 










1 




1 




7. Duct(s) for accessory boring organ and 
















ventral pedal gland 















1 




8. Bursa copulatrix 







2 


1 




1 





9. Seminal receptacles at dorsal periphery 
















of albumen gland 







1 













10 Fenial shape 







2 


1 










11. Penial vas deferens 







1 










2 


12. Prostate 







1 













13. Accessory salivary gland(s) 







1 


1 




2 


2 


14. Straw-like membrane around glancl of 
















Leiblein 










1 




1 


1 


1.5, Posterior duct of gland of Leiblein 










1 




1 


1 


16. Ontral cusp of rachidian 















1 


1 


17 Margin of rachidian basal plate 










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 
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 



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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 



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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: 
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. 

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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 


49 


06 


0,64-0,80 


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 


24 


2,47-2,83 


2.65 


0.13 


NS 


.\perture length total Icn 


gth 


044-0,61 


0,50 


0,(.)5 


51-0,55 


0,52 


0.02 


NS 


Length last uhorl, total length 


0,65-0,87 


072 


0,06 


0,66-0,71 


69 


0.02 


NS 


,\pertiire length 'aperturt 


" width 


3,78-4,58 


4,27 


24 


3,55-4,87 


4,26 


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 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 


02 


0.57 


05 


7 


Posterior tooth 


6.66 


-0.002 


0.02 


0.76 


7 


Provinculum 


37.32 


0.01 


0.03 


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. 

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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 




95-1 10 


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. 



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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 
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Division of Mollusks 
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Natural History 
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Department of Invertebrates 
Field Museum of 
Natural History 
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Department of Biology 
College of Charleston 
Charleston, SC 29424 

Dr. William K. Emerson 

Department of Living Invertebrates 

The American Museum of Natural 

History 

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Division of Mollusks 
National Museum of 
Natural History 
Smithsonian Institution 
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Division of Mollusks 
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Museum of Comparative Zoology 
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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 42-0.50 mm long (ex- 
cluding immersed base), 39-0.42 mm wide, with 3.2- 
3 4 whorls; first whorl diameter 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. 

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Morch, O.A.L. 1875b, Ssnopsis molluscorum niarinorum lu- 
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Morch, O.A.L. 1876. A descriptive catalogue of the Scalidae 
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Nowell-l'stickc, G.W. 1959 A check list of the marine shells 
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.Sowerb>, GB, II. 1873-1874. Monograph of the genus Sca- 
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Souerby, GB. III. 1894. Descriptions of twelve new species, 
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Strong, ,AM. and L.G. Ilertlein. 1937. New species of Recent 
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Taki, I 1956 Anatomical study on Japanese P^pitoniidae (1 ). 
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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 



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 




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 







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, 



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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 



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Department of Biology 
College of Charleston 
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Dr. William K. Emerson 

Department of Living Invertebrates 

The American Museum of Natural 

History 

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Dr. Robert Hershler 
Division of Mollusks 
National Museum of 
Natural History 
Smithsonian Institution 
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Department of Mollusks 
Museum of Comparative Zoolog\ 
Harvard University 
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Department of Geology 
The Ohio State University 
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Department of Malacology 
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Natural Histor\ 
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% Department of Mollusks 
Museum of Comparative Zoology 
Harvard University 
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and Geophysics 

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Museum of Zoology 
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Dr. Ruth D. Turner 
Department of Mollusks 
Museum of Comparative Zoology 
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Dr. Geerat J. Vermeij 
Department of Geology 
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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 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 
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 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 




02 
oot 




1 5h 05 

1 iiT 07 



II 



L/W 



Sdit Ixittcmis 
SaiicK intcrtiilal 



1 17 
I 20 



04 

0.) 



I SS 



05 
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 
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Ituarte, i'. F. 1984b. El fen(Smeno de Ineubacion branquial 
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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 



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@ This paper meets ttie requirements of ANSi/NISO Z39. 48-1992 (Permanence of Paper). 



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